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Vol. LXXXVIII MARCH 1980

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Devoted to Entomology in General

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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

The meetings of the Society are held on the 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; including subscription to the Journal, $15.00.

Members of the Society will please remit their annual dues, payable in January, to the
Treasurer.

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Officers for the Year 1980

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President , Dr. Betty Faber

American Museum of Natural History, New York 10024
Vice-president , Dr. Joseph Cerreta

Columbia University, New York 10032
Fordham University, New York 10458

: .

Secretary , Dr. Gerard Iwantsch
Assistant Secretary, Dr. Henry M. Knizeski, Jr

Mercy College, Cobs Ferry, New York 10522

Treasurer , Mr. Louis Sorkin

American Museum of Natural History, New York 10024
Assistant Treasurer , Mr. Kenneth Sherman

Rutgers University, New Brunswick, New Jersey 08903

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Trustees

Clas!s of 1980

Ms. Jane Harvey Cooper

Class of 1981
Dr. Ivan Huber
Ms. Laurene Ford

Publication Businesrs Manager
Irene Matejko

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Fordham University, New York 10458

Mailed June 27, 1980

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The Journal of the New York Entomological Society (ISSN 0028.-7199) is published quart,ierly for the Society by Allen Press-
Inc., 1041 New Hampshire, Lawrence, Kansa^ 66044. Second class postage paid dt New Brunswick, NeW Jersey and at
additional mailing office.

Known office of publication: W&lcsman Institute of Microbiology, 'New Brunswick, New Jersey 08903.

Journal of the N.Y. EntomologicaLSdciety, total No. copies printed 750. Paid circulation 490, mail subscription 470, free
distribution by mail 23, tdtal distribution 493, left-over 2^57 copie^ each quarter.

- A- r

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Journal of the

New York Entomological Society

VOLUME LXXXVIII MARCH 1980

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. Randall T. Schuh
American Museum of Natural History
Dr. Daniel Sullivan
Fordham University
Dr. Louis D. Trombetta
St. John’s University

CONTENTS

The seasonal occurrence of sexual brood and the pre- and post-nuptial behavior of the
honey ant, Myrmecocystus mexicanus Wesmael, in Colorado John R. Conway

Digestive system and associated organs in the adult and pupal male doryline ant Aenictus
gracilis Emery (Hymenoptera: Formicidae) S. Shyamalanath and James Forbes

Fecelia biorbis n. sp. (Heteroptera: Pentatomidae) , a new species from Haiti

Joseph E. Eger, Jr.

Abstracts of papers presented at the 51st Annual Meeting, Eastern Branch, Ento-
mological Society of America, Hershey, PA, September 26-28, 1979
Migration, distribution, and habitat preferences of Blattella germanica (L.) in urban,
low-income apartments (Dictyoptera: Blattellidae) R. C. Akers and W. H. Robinson
Comparative toxicity of etrimfos to gypsy moth larvae and house flies.

M. Akram and A. J. Forgash

Release of oviposition-deterring pheromone by apple maggot flies, Rhagoletis pomonella
(Walsh) (Diptera: Tephritidae) A. L. Averill and R. J. Prokopy

Oviposition response of the horn fly (Diptera: Muscidae) to environmental and manure char-
acteristics N. P. Bacon

The impact of the soil systemic, aldicarb and the foliar insecticide, pirimicarb on po-
tato plant growth B. A. Bajusz,

Z. Smilowitz, T. P. Mack, F. L. Petitt, C. A. Martinka and M. E. Whalon
Effect of electromagnetic devices on subterranean termites ( Reticulitermes ), dry wood ter-
mites ( Incisitermes minor), and German cockroaches ( Blattella germanica) R. Beal
The effects of mowing on the guild of sap-feeding insects associated with Kentucky blue-
grass Richard A. Bean and Robert F. Denno

Sweet corn insect pest management in New Hampshire

J. S. Bowman and A. T. Eaton
Bait sampling for white grubs (Coleoptera: Scarabaeidae) and wireworms (Coleoptera:
Elateridae) in Virginia corn land S. P. Briggs and W. A. Allen

Toxicity of the IGR L7063 to Choristoneura fumiferana (Clem.) (Lepidoptera: Tortricidae)
and two associated parasites J. R. Brush wein and J. Granett

Status of a biological control program against the cereal leaf beetle, Oulema melanopus
(L.) (Coleoptera: Chrysomelidae) T. L. Burger

Growth and survival of Hylotrupes bajulus (L.) larvae under favorable and unfavorable
conditions (Coleoptera: Cerambycidae) K. F. Cannon and W. H Robinson

Fungicidal inhibition of Beauveria bassiana, a pathogen of the Colorado potato beetle
( Leptinotarsa decemlineata Say.) (Coleoptera: Chrysomelidae) R. A. Clark

Insemination frequency and sperm precedence in the German cockroach (Dictyoptera:
Blattellidae) D. G. Cochran

Parahost behavior of adult Gasterophilus intestinalis (DeGeer) (Diptera: Gasterophilidae)

S. E. Cope and E. P. Catts

USDA Procedure for acting on new pest problems R. L. Cowden

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41

Herbivorous insects causing mortality of pine seedlings in an old field

W. J. Cromartie, M. A. Rivera, J. Gfrorer and D. Maser
Geographic variation in vagility in the milkweed beetle, Tetraopes tetraopthalmus
(Forster) (Coleoptera: Cerambycidae) M. A. Davis

Factors affecting honey bee Apis mellifera L. (Hymenoptera: Apidae) foraging on birds-
foot trefoil ( Lotus corniculatus ) G. L. DeGrandi and C. H. Collison

The mayfly fauna (Ephemeroptera) of two Connecticut streams P. J. Dodds

Impact of predators and parasites on the survival of Rhinocyllus conicus Froelich
(Coleoptera: Curculionidae) P. F. Dowd and L. T. Kok

Ovipositional preference of the grape colaspis, Colaspis brunnea (Fab.) (Coleoptera:
Chrysomelidae) in North Carolina A. T. Eaton, R. L. Rabb and J. W. Van Duyn
Presocial behavior in Cerceris watlingensis Elliott & Salbert (Hymenoptera: Sphecidae)

N. B. Elliott and T. Shlotzhauer
Effects of dietary nitrogen on 14C-glycine and 14C-formate incorporation into body urates
of Blattella germanica (Dictyoptera: Blattellidae)

J. A. Engebretson and D. E. Mullins
Niche relations of stinkbugs ( Podisus spp.) attacking eastern tent caterpillars E. W. Evans
Effect of temperature and light on aggregation behavior of Aedes stimulans larvae (Dip-
tera:Culicidae) in a woodland pool G. P. Fernald and J. F. Burger

A definitive method for determining pear seedling resistance against pear psylla, Psylla
pyricola Foerster (Hemiptera: Psyllidae) B. J. Fiori and R. C. Lamb

PTTH and ecdysone release in last instar larvae of the European corn borer, Ostrinia
nubilalis (Hiibner) (Lepidoptera: Pyralidae) D. B. Gelman and D. K. Hayes

Effects of decapitation on ecdysone and vitellogenin titers and egg maturation in Aedes
aegypti (Diptera: Culicidae) J. T. Greenplate

Control of neotenic development in a primitive termite (Isoptera: Hodotermitidae)

S. Greenberg and A. M. Stuart
Resmethrin sprays for relief from deer flies, Chrysops atlanticus (Diptera: Tabanidae)

E. J. Hansens

A field study of Aedes sollicitans (Walker) (Diptera: Culicidae) as a vector of filarial
worms W. M. Johnson and W. J. Crans

A systematic analysis of the genus Actici Robineau-Desvoidy in North America with de-
scriptions of two new species (Diptera: Tachinidae) M. A. Kamran

Removal method estimation of Blattella germanica (L.) (Dictyoptera: Blattellidae)
populations C. B. Keil

Assessment of insect damage to vegetation by remote sensing V. Klemas

Loss assessments and decision-making for the tobacco budworm, Heliothis virescens
(F.) (Lepidoptera: Noctuidae), on Maryland tobacco

D. M. Kolodny-Hirsch and F. P. Harrison
Effect of habitat characteristics on the succession of outlet-breeding black flies (Dip-
tera: Simuliidae) in New Hampshire D. J. Lake and J. F. Burger

Computer-assisted instruction for pest management: ORCHARD and ALFWEEV

P. A. Logan

Predicting the impact of Colorado potato beetle ( Leptinotarsa decemlineata Say) on
yield P. A. Logan and R. A. Casagrande

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45

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48

48

49

49

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51

51

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52

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55

Treatment of a nuclear polyhedrosis virus of Autographa californica Speyer with 5-
Bromodeoxyuridine and determination of changes in virulence in four lepidopteran
hosts J. T. McClintock and C. F. Reichelderfer

Consequences of host phenology for exotic scale insects M. S. McClure

Effect of Agromyza frontella Randani (Diptera: Agromyzidae) on alfalfa quality and
yield G. B. MacCollom, G. Baumann and J. G. Welch

Effect of pirimicarb on natural enemies of Myzus persicae (Sulzer) (Homoptera: Aphidi-
dae) T. P. Mack and Z. Smilowitz

Development of adult abdomen in Drosophila melanogaster Meigen (Diptera: Drosophili-
dae) M. M. Madhavan and K. Madhavan

Feeding behavior of Tabanidae (Diptera) on livestock L. A. Magnarelli

New approaches for screening passenger baggage at U.S. international airports

J. H. Mahaney

Mate-seeking strategies in male flower flies (Diptera: Syrphidae) C. T. Maier

A comparison of laboratory-reared and wild type gypsy moth Lymantria dispar (L.)

males (Lepidoptera:Lymantriidae) V. C. Mastro

Seasonal dispersal of pitcher plant mites (Acarina: Anoetidae)

T. N. Mather and E. P. Catts

Development of overwintered versus summer pupae of the alfalfa blotch leafminer
Agromyza frontella Randani (Diptera: Agromyzidae)

W. K. Mellors and R. G. Helgesen
A new Japanese beetle trap containing pheromone and floral lure as synergistic attract-
ants F. W. Michelotti and J. W. Seidenberger

Comparative effects of some insect growth regulators on development, morphogenesis,
and reproduction of the female rice weevil, Sitophilus oryzae (L.) (Coleoptera: Cur-
culionidae) J. Mkhize and A. P. Gupta

Effects of soybean plant populations and planting configurations on insect populations

J. W. Murphy and J. C. Smith
Biological control of the greenhouse whitefly, Trialeurodes vaporariorum (Westw.)
(Homoptera: Aleyrodidae) by Encarsia formosa Gahan (Hymenoptera: Aphelinidae)

J. R. Nechols

Attack ecology of the sugar maple borer, Glycobius speciosus (Say) (Coleoptera: Ceram-
bycidae) W. G. Newton

Biology of Leptothrips mali (Fitch) (Thysanoptera: Phlaeothripidae), a predator of orchard
mites M. P. Parrella and R. L. Horsburgh

Quantification of Myzus persicae (Sulzer) (Homoptera: Aphididae) damage to potato fol-
iage production F. L. Petitt, Z. Smilowitz and E. S, Nolan

Notal setae of first instar nymphs of five Periplaneta species (Dictyoptera: Blattidae)

P. K. Powell

Mating behavior of the bee Colletes thoracicus (Hymenoptera: Colletidae)

E. G. Rajotte and R. B. Roberts
Interactions among willow herbivores as a determinant of feeding and ovipositional pat-
terns of the imported willow leaf beetle, Plagiodera versicolora Laich. (Coleoptera:
Chrysomelidae) M. J. Raupp and R. F. Denno

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59

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60

60

61

62

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63

63

64

64

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Insects that deform black cherry C. O. Rexrode

Sexual behavior in Hawaiian Drosophila (Diptera: Drosophilidae) J. M. Ringo

Oviposition behaviour and egg distribution of the European apple sawfly Hoplocampa
testudinea Klug. (Hymenoptera:Tenthredinidae) B. D. Roitberg and R. J. Prokopy
Anomalous oviposition behavior by the gypsy moth (Lepidoptera: Lymantriidae)

M. C. Rossiter

Adaptations of the fall cankerworm Alsophila pometaria (Harris) (Lepidoptera: Geometri-
dae) to unapparency of host leaf flush J. C. Schneider

Delayed hatching of Japanese wax scale, Ceroplastes ceriferus (F.) (Homoptera: Coc-
cidae) in locations strongly affected by marine winds P. B. Schultz

Application of pheromone technology in the gypsy moth program

C. P. Schwalbe and E. C. Paszek
Life history strategies of spring feeding forest Lepidoptera D. F. Schweitzer

Effect of planting date on the abundance of insect pests on flue-cured tobacco

P. J. Semtner and T. R. Terrill
EDNH-stimulated ecdysone production by ovaries of Aedes aegypti (L.) (Diptera:
Culicidae) J. P. Shapiro and H. H. Hagedorn

Encapsulation of pollen attractants for honey bee, Apis mellifera L. diets (Hymenoptera:
Apidae) E. W. Herbert, Jr., H. Shimanuki and B. S. Shasha

Establishing economic parameters for green peach aphid, Myzus persicae (Sulz.) (Ho-
moptera: Aphididae) damage on commercial potatoes

Z. Smilowitz, M. E. Whalon, E. S. Nolan and C. A. Martinka
The bee louse, Braula coeca Nitzsch (Diptera: Braulidae), its distribution and preference
for queen honey bees I. B. Smith, Jr., and D. M. Caron

Rearing of Tabanus nigrovittatus Macquart (Diptera: Tabanidae) from egg to adult

R. K. Sofield and E. J. Hansens
Transmission of equine babesiosis and bovine anaplasmosis by Dermacentor albipictus
(Packard) (Acari: Ixodidae) D. Stiller, W. M. Frerichs, G. Leatch, and K. L. Kuttler
Soybean foliage consumption by the adult Mexican bean beetle, Epilachna varivestis
Mulsant (Coleoptera: Coccinellidae)

I. M. Sunzenauer, T. Elden, and A. L. Steinhauer
Humidity tolerances of 3 species of phytoseiid mites (Acarina: Phytoseiidae)

F. C. Swift and L. Blaustein

Hudson Porta-Pak ULV Sprayer R. Treichler

Integration of Rhinocyllus conicus Froelich larvae (Coleoptera: Curculionidae) and 2,4-D
for control of Carduus nutans L. J. T. Trumble and L. T. Kok

Characterization of the structural components of the granulosis virus of Plodia interpunc-
tella (Lepidoptera: Pyralidae) K. A. Tweeten

Biological characterization of AMBUSH insecticide M. Tysowsky

Gas-liquid chromatographic comparisons of the cuticular lipids from male and female
house crickets, Acheta domesticus (L.) (Orthoptera: Gryllidae)

E. C. Uebel and J. D. Warthen, Jr.

Life history and population dynamics of tree-dwelling Homoptera J. W. Webb

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Distribution of Phyllonorycter ( = Lithocolletis ) sp. in northeastern apple orchards

R. W. Weires and J. R. Leeper 81

Seasonal history of the leafhopper complex (Homoptera:Cicadellidae) on ornamental
honeylocust A. G. Wheeler, Jr. and K. Valley 81

Physiological age study of five species of mosquitoes (Diptera: Culicidae) in southeastern
New Hampshire J. J. Winegar 82

Book review 83

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(l), 1980, pp. 7-14

THE SEASONAL OCCURRENCE OF SEXUAL BROOD AND THE
PRE- AND POST-NUPTIAL BEHAVIOR OF THE HONEY ANT,
MYRMECOCYSTUS MEXICANUS WESMAEL, IN COLORADO

Dr. John R. Conway

Abstract. — Nocturnal observations and excavations of honey ant colonies
in the vicinity of Colorado Springs, Colorado provided information on pre-
and post-nuptial behavior and the seasonal appearance of the reproductive
brood. The mating flights of three different nests were observed on the
evenings of July 24, 25 and 28. The nuptial activity at one formicary lasted
an hour and a half and peaked between 7:26 and 7:36 PM when the sexuals
took wing. During this period 56 queens and 100-110 males were counted.
Dealated queens were observed after the nuptial flights. Queen larvae were
uncovered in nests between October and February and queen cocoons were
unearthed in May and June. Winged queens and males were taken from
nests approximately one month prior to swarming and were noted outside
colony entrances several days before and after the nuptial flights. A single
wingless queen was recovered from each of the three completely excavated
colonies.

Introduction

All ants are social insects, and most colonies contain three castes: queens,
males and workers. Typically, males and queens are produced in numbers
at certain seasons and emerge during a spectacular nuptial flight. After being
inseminated, the queen sheds her wings, excavates a small burrow and
nourishes the brood from her metabolized fat bodies and alary muscles
(Wilson, 1974).

This paper sheds some light on the seasonal appearance and duration of
the reproductive brood and the pre- and post-nuptial behavior of the honey
ant, Myrmecocystus mexicanus Wesmael in Colorado.

Materials and Methods

Information on the presence of M. mexicanus reproductives was collected
during the complete excavation of three formicaries and the partial un-
earthing of five others in the vicinity of Colorado Springs, Colorado in all
months except March, April, August and November. Workers and brood
from a December excavation were transferred to a vertical ant farm to study
the development of reproductives under laboratory conditions.

The nuptial flights of three separate colonies were observed on the eve-

8

NEW YORK ENTOMOLOGICAL SOCIETY

nings of July 24, July 25, and July 28. Dealated queens were located after
some mating flights by searching ridge and mesa tops near the active nests
with the aid of a double-mantle Coleman lantern.

Results and Discussion

Seasonal occurrence of sexual brood. — Queen larvae: Queen larvae were
uncovered in formicaries at depths ranging from 36 to 144 cm. The largest
numbexjincovered in one nest was 80 and they were found between 88 and
144 cm during the period from January 13 to February 3. The largest num-
ber, 38, was in a chamber 119 cm deep. They were probably deep in the
nest to protect them from the cold temperatures characteristic of that time
of the year.

Development of queen larvae begins as early as October 8 since a few
were unearthed in another colony at that time. They were at depths ranging
from 36 to 114 cm. From the foregoing, it is evident that the development
of queens begins at least 9.7 months prior to the nuptial flight in late July.
The queen larval stage is fairly long, extending from October 8 to February
3, a period of approximately 4 months.

Queen pupae: Queen pupae were uncovered in two colonies at depths

ranging from 6 to 178 cm during the period from May 31 to June 22. As a
result they were both closer to the surface and deeper in the nests than the
queen larvae.

Over 176 queen cocoons were removed from one nest in June, suggesting
the tremendous reproductive potential of a single colony. This figure is over
twice the maximum number of queen larvae uncovered in January and Feb-
ruary and leads to speculation that the number of queen larvae does not
peak until sometime between February and June.

The pupal stage appears to be considerably shorter than the larval stage.
Queen pupae were uncovered in formicaries between May 31 and June 22,
a period only 23 days long, and only 1.8 months prior to swarming. Queen
pupae appeared in the laboratory vertical nest much earlier, on January 14,
but their development was probably speeded by the warmer indoor tem-
peratures.

Queen cocoons were not observed during excavations between late July
and February. Apparently adults hatch in June and the pupae do not reap-
pear until the following spring.

Males and winged queens: Both males and alate queens were unearthed

in a colony as early as June 22, about a month before swarming, in chambers
at 36 cm, 48 cm, and 50 cm, but only males were present in two deeper
chambers at 64 cm and 83 cm. Winged queens were also taken from sub-
surface passages. Males and an alate queen were found below a rock at the
surface at another nest on July 16. This suggests that males and queens are

VOLUME LXXXVIII, NUMBER 1

9

at first deep in the nest, with males being deepest, but that they approach
the surface as the time of the nuptial flight approaches.

Others have reported adult reproductives present in the nests even earlier.
Gregg (1963) recorded males and females of M. mexicanus in colonies on
June 2 and Snelling (1976) noted reproductives of Myrmecocystus in nests
up to three months prior to their mating flight. Cazier and Mortenson (1965)
reported winged males and females of M. mimicus in subsurface burrows,
3 to 8 cm deep, as early as March 9, even though their nuptial flights occur
in late July and early August. This is a holding period of more than five
months.

Two newly-hatched winged queens appeared in my vertical laboratory
colony on February 28, approximately five months in advance of the nuptial
flight. But surprisingly, the first male did not appear in this nest until July
14, just two weeks prior to when the nuptial flight would ordinarily occur.
These unusual developmental times are attributed to the unnatural labora-
tory conditions.

Wingless queen: A single wingless queen was taken from each of the

three completely excavated nests. At one formicary she was in the second
deepest chamber at 138 cm along with 40 repletes, 3 depleted repletes, 2
semi-repletes, 25 queen larvae, many small larvae, and 1,819 workers. The
queen of another colony was in the 8 cm by 16 cm bottom chamber at a
depth of 178 cm, which also contained 2 flaccid repletes, 8 semi-repletes,
13 depleted repletes, 2 queen cocoons, many small larvae, about 700 work-
ers, and the remains of a wasp.

An exception was the third nest, where the queen was taken from a
chamber only 14 cm deep along with many small larvae and many workers.
She was in poor condition and could only move her legs and antennae
slightly, possibly due to injuries sustained when workers dragged her
through the passages to escape excavation.

McCook (1882), Creighton and Crandall (1954) and Snelling (1976) also
report but a single gravid female per colony. Thus, there appears to be but
one wingless queen, who is usually in the lowest part of the nest accom-
panied by repletes (in various stages of distension), brood, and a large con-
tingent of workers.

Appearance of reproductives before and after the nuptial flight. — Repro-
ductives appeared outside the entrances of the Colorado Springs nests sev-
eral days prior to swarming. This probably is a result of the fact that sexuals
display a circadian rhythm, with sharp increases in restlessness, at just the
hour each day at which the nuptial flights occur (McCluskey, 1967).

Males were observed outside the entrances as early as July 14, 10 to 14
days prior to swarming, and interacted with the workers. For example, on
one occasion a male left the entrance despite the efforts of a worker trying
to pull him back in. Another time a male went off the crater and a major

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NEW YORK ENTOMOLOGICAL SOCIETY

scurried about the cone as if searching for him. Males were even observed
being carried back to the nest by workers. But, on another occasion, it was
a worker who carried the male out of the entrance prematurely.

Winged queens were observed outside the entrances as early as July 20,
four to eight days before the nuptial flights. Some queens left the crater and
wandered away, while others went out and then returned. A dead winged
queen was found near the entrance of one nest four days before the nuptial
flight. Snelling (1976) reported a M. mexicanus queen outside a nest as early
as June 27.

At some nests winged queens and males were observed several days after
the nuptial flight. In fact, the last sighting of males and alate queens was on
July 29, one to five days after the observed nuptial flights. Snelling (1976)
reported M. mexicanus queens still in the nests as late as September 29.

Pre-nuptial activity. — The nuptial flights and pre-nuptial behavior of three
different colonies in the Colorado Springs region were observed on July 24,
July 25 and July 28. Slocumb (1966) reported mating flights at seven colonies
on the plains east of Colorado Springs as late as August 2. On the other
hand, Paul Nesbit, a local naturalist, noted a nuptial flight in the Garden of
the Gods on July 19, and recalled honey ants swarming as early as July 3
(personal communication). Thus, while this activity may occur anytime dur-
ing a one month period, it usually takes place in the last half of July.

Snelling (1976) reports that flights of M. mexicanus in eastern Arizona,
New Mexico and Colorado occur during the summer months, usually fol-
lowing an afternoon rain, and that in southern California, because of the dry
summers, these flights are postponed until the first fall rains.

There was some correlation with precipitation in the Colorado colonies.
It rained the night before swarming at two nests and at a third colony, it
rained both the day before and the afternoon preceding the flight.

Pre-nuptial activity began somewhat earlier in the evening than the usual
nocturnal exodus which commenced at 7:30 PM (Mountain Standard Time),
on the average. For example, on July 28, there were already 5 queens and
6 males in the entrance and surrounded by workers at one Colorado Springs
nest when observation began at 7:00 PM. Four queens and 21 males ap-
peared at 7:10 PM when the air temperature was 21°C. Males lined the
entrance, surrounded by workers on the crater, while queens periodically
came up and then receded from view. As more queens and males emerged,
they almost completely filled the entrance. Eight queens and 32 males ven-
tured farther from the opening at 7: 19 PM. By 7:23 PM there were 24 queens
and 40 males advancing onto the crater and a number reached the outer
edge a minute later. The overall worker activity increased greatly at 7:26
PM, when 42 queens and about 100-110 males were counted. Field obser-
vations on M. mimicus also indicated worker excitement around the nest
entrance just prior to and during emergence of the winged sexuals (Cazier

VOLUME LXXXVIII, NUMBER

1

and Mortenson, 1965). By 7:27 PM, reproductives were going off the crater
in all directions. Fifty-four queens and an estimated 70 males had advanced
about 0.6 m from the entrance at 7:29 PM. First a male took wing and rose
straight up, and then a few queens attempted to fly. Snelling (1976) reports
that males generally precede the females by several minutes. At 7:33 PM a
queen finally lifted off successfully. Some queens scaled grass tussocks, but
most tried to take off from the ground, usually after buzzing about the
surface futilely for a time. They then soared straight up and out of sight.
Queens took wing in all directions at 7:36 PM, but the flights were difficult
to follow due to impending darkness. Workers nipped at the queens on the
ground, apparently to prevent them from returning to the crater and to
induce them to fly. Thirty queens and 80 to 90 males were present at 7:42
PM. Nine queens and several males were seen around the crater at 7:50 PM
as the nuptial activity waned. At 8:04 PM there was but one queen and a
reduced number of workers on the crater. By 8:21 PM there was little ac-
tivity, and the number of ants on the crater was greatly diminished. A few
workers were seen a short distance from the entrance, and one was carrying
a dead insect. Cazier and Mortenson (1965) also reported workers foraging
during the nuptial activities. The last queen peered out the entrance at 8:21
PM, and then withdrew. As mentioned previously, unsuccessful winged
queens and males were observed several days after the nuptial flight, but
their ultimate fate in the colony is unknown.

The swarming of this nest lasted an hour and a half and peaked between
7:26 and 7:36 PM. Altogether 56 queens were collected (many escaped) and
100 to 110 males were counted during the pre-nuptial activity. Nesbit (per-
sonal communication) observed a nuptial flight of M. mexicanus in the Gar-
den of the Gods, during which he counted only half as many queens (28)
but approximately the same number of males (100).

Another colony was somewhat atypical in that flight attempts were ob-
served on three nights, July 22, 23, and 24, instead of on only one night.
This may have resulted from rain at the time of the nuptial flight on those
evenings. In any event, the queens actually took off only on July 24. Another
peculiarity of this nest was that the queens made a mass exodus twice on
July 23, once at 7:10 PM and later at 7:30 PM, but returned after each
departure.

In summary, queens and males first left the three observed colonies be-
tween 7:10 and 7:30 PM. Slocumb (1966) saw them exit from plains nests
even later, at 7:40 PM. Reproductives attempted to fly or took wing at times
ranging from 7:22 to 7:36 PM. Swarming ended when the unsuccessful
queens and males returned to the formicary, and this occurred between 7:40
and 8:04 PM. Slocumb (1966) reported that they came back later, at
8:30 PM.

Wingless queens: Although mating was not observed, it is presumed to

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occur during the crepuscular nuptial flight. It is possible that multiple in-
seminations take place since they have been observed in a number of for-
micine species (Wilson, 1974).

Pre-nuptial activity was observed at a colony on a mesa dotted with sim-
ilar nests on July 25, and a total of eight dealated queens were collected
later that evening. The first one was found at 8:05 PM on a nearby side
ridge, about a half hour after the nuptial flight. The others were captured
about 0.6 km away on the eastern end of the mesa, between 8:58 and 11:38
PM.

There is evidence that queens may not dig burrows until the night after
the nuptial flight. For example, on July 26, another dealated queen was
observed starting to dig a burrow at 6:20 PM, an hour before the normal
nuptial flight for that evening.

On July 27, at 8:50 PM, two more wingless queens were discovered dig-
ging burrows in the same region. Interestingly, one was digging in a clearing
around a Pogonomyrmex occidentalis colony. Mature M. mexicanus nests
have been observed in the immediate vicinity of this species on four other
occasions. Apparently they can tolerate the close proximity and avoid com-
petition by being active at different times during the day and night. The last
queen to excavate a burrow in this area was found on August 4, ten days
after swarming was first observed.

The queen-burrow entrances, which are 5 mm in diameter, were even-
tually closed with a plug of earth. The first closed entrance was noted on
July 27, two days after the nuptial flight. Snelling (1976) reports that Myr-
mecocystus queens dig to a depth of 15 and 46 cm before constructing a
brood chamber.

Four queen-burrows on the mesa were marked with rock cairns and left
for future observation. All four were closed by July 29, four days after
swarming, but unfortunately none reopened the following summer. This may
be indicative of the high mortality generally reported for colonizing queens.

Another wingless queen was collected the following summer on July 23
in the same area. Thus, dealated queens of M. mexicanus were found be-
tween July 23 and August 4, but no males were observed after the nuptial
flights.

Conclusions

Swarming in M. mexicanus in the Colorado Springs area typically occurs
about dusk on one evening in late July and may be associated with precip-
itation up to a day in advance. Specifically, it was observed at three different
colonies on July 24, 25 and 28.

Pre-nuptial activity begins somewhat earlier in the evening than the usual
stirrings, but interestingly, the peak activity, when the queens and males

VOLUME LXXXVIII, NUMBER 1

13

took wing, occurred at approximately the same time, 7:30 PM, as the normal
foraging exodus. In fact, after the flight the colony resumes its search for
food.

At one nest the pre- and post-nuptial activity lasted about an hour and a
half, during which time 56 queens were collected and 100 to 110 drones
were counted. Swarming ended when the unsuccessful queens and males
returned to the formicaries, and this occurred between 7:40 and 8:04 PM.

Although honey ants are reasonably abundant in the Colorado Springs
area there are several flaws in the nuptial process which probably restrict
their numbers:

1. Not all queens and males participate in the nuptial flight, since some
return to the colony afterwards.

2. Adverse weather may desynchronize the process and interfere with
aerial union. At one nest queens and males exited on three successive
nights but took wing on only one evening.

3. A few males and females leave nest entrances several days prior to the
flights and some are undoubtedly lost.

4. There appears to be a high mortality among the founding queens which
greatly decreases the reproductive potential.

5. The queens and colonies have rather specific habitat preferences, gen-
erally being found on the tops of ridges or mesas between 5,000 and
7,000 feet.

The metamorphosis of queens begins with the appearance of queen larvae
in October, approximately ten months prior to the nuptial flight. This de-
velopmental stage appears to be the longest, persisting at least four months
until February and perhaps longer.

Queen pupae were unearthed as early as May 31, about two months prior
to swarming, but were actually observed in the nests for only 23 days. Thus,
this stage may last less than one-fourth as long as the queen larval stage.
Over 176 queen cocoons were recovered from one nest, suggesting the large
numbers of queens which may be produced.

Adult males and females were removed from colonies as early as June 22,
about one month prior to swarming, and a period just slightly longer than
the pupal stage. This data needs to be carefully reexamined however, es-
pecially in light of reports of reproductives being present up to five months
in advance of the nuptial flight.

The discovery of but a single wingless mother queen in each nest is con-
sistent with the findings of most investigators.

Thirteen dealated queens were located after the mating flights between
July 23 and August 4. No males were found after the flights. Usually the
queens began to excavate their burrows within one-half to four hours after

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the flight, but one appeared to have delayed until the next evening, when
she was observed digging an hour in advance of the time for the flight.

The queen-burrow entrances were 5 mm in diameter, but began to be
closed only two days after the flight. Four closed entrances were marked
for future observation, but none reopened. These probably are indicative of
a high mortality rate among founding-queens.

Acknowledgments

I am indebted to Paul Nesbit who helped me locate M. mexicanus colo-
nies in the Garden of the Gods and allowed me to observe the nuptial
activities of a nest in his yard. Throughout the course of this investigation,
R. E. Gregg has made many valuable suggestions.

Financial assistance was provided by grants from the Walker Van Riper
Fund of the University of Colorado Museum and from the Kathy Lichty
Memorial Fund of the University of Colorado.

Literature Cited

Cazier, M. A. and M. A. Mortenson. 1965. Bionomical observations on myrmecophilous
beetles of the genus Cremastocheilus (Coleoptera: Scarabaeidae). Journal of the Kansas
Entomological Society 38:19-44.

Creighton, W. S. and R. H. Crandall. 1954. New data on the habits of Myrmecocystus melliger
Forel. The Biological Review, City College of New York 16:2-6.

Gregg, R. E. 1963. The Ants of Colorado. University of Colorado Press, Boulder. 792 pp.
McCluskey, E. S. 1967. Circadian rhythms in female ants, and loss after mating flight. Com-
parative Biochemistry and Physiology 23:665-77.

McCook, H. C. 1882. The honey ants of the Garden of the Gods, and the Occident ants of the
American Plains. J. B. Lippincott & Co., Philadelphia.

Slocumb, L. B. 1966. A report on the honey ants of the plains. M.A.T. Thesis, Colorado
College. 31 pp.

Snelling, R. R. 1976. A revision of the honey ants, genus Myrmecocystus. Natural History
Museum of Los Angeles County Science Bulletin 24:163 pp.

Wilson, E. O. 1974. The Insect Societies. Harvard University Press, Cambridge. 548 pp.

Department of Biology, Elmhurst College, Elmhurst, Illinois 60126
Received for publication August 20, 1979.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(l), 1980, pp. 15-28

DIGESTIVE SYSTEM AND ASSOCIATED ORGANS IN THE ADULT
AND PUPAL MALE DORYLINE ANT AENICTUS GRACILIS
EMERY (HYMENOPTERA: FORMICIDAE)1

S. Shyamalanath and James Forbes

Abstract. — This paper presents the first description of the anatomy and
histology of the alimentary tract, the postpharyngeal glands, the salivary
glands, the Malpighian tubules, and the mandibular glands in the male adult
and pupa of the Old World doryline ant, Aenictus gracilis. The alimentary
tract consists of the cibarium, the buccal tube, the infrabuccal chamber, the
pharynx, the oesophagus, the cardiac valve, the ventriculus, the pylorus,
the intestine, the rectal valve and the rectum. In the adult the infrabuccal
chamber lies posterior to its usual position in ants under the posterior floor
of the buccal tube and the anterior pharynx, while the pupa has a recess
where the infrabuccal chamber is usually present. The pharynx is divisible
into a broad anterior, an indented middle, and a tubular posterior region
into which the postpharyngeal glands open. The epithelium of the pharynx
has flattened cells in the adult and low columnar cells in the pupa. The
intima of the postpharyngeal glands and ducts bears long hair-like projec-
tions. Maxillary glands are absent. The epithelium of the oesophagus has
small folds in the pupa and the lumen is wider than in the adult. The con-
tinuation of the oesophagus into the ventriculus is the cardiac valve. A crop
and a proventiculus are absent. The ventriculus has a dorsal concavity to-
ward its posterior half. The ventricular epithelium of the adult consists of
digestive and regenerative cells. In the pupa the ventricular epithelium has
disintegrating and regenerating regions. The regenerating digestive cells are
of three distinct types that represent stages in the maturity of the epithelium.
The pylorus is internally divided into an anterior region where the 18-20
Malpighian tubules enter, and a posterior region. The posterior extension
of the intestine into the rectum is the rectal valve. The inner wall of the
rectum has 3 rectal pads. The very small salivary glands are of the simple
branched acinar type, and their ducts have a tortuous cuticular lined lumen.
The mandibular gland consists of about 12 pyriform cells and a reservoir.
Comparisons made with the other Old World and New World species pre-
viously described have revealed features of phylogenetic importance that
lend support to the concept of the triphyletic origin of the dorylines.

1 Part of a dissertation submitted to the Graduate School of Fordham University by the first
author in partial fulfillment of the requirements for the degree of Doctor of Philosophy.

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The study of the morphology, anatomy and histology of male ants is of
particular importance because as Janet (1902) recognized, they are the least
specialized of all of the nest forms and most wasp-like in appearance. This
wasp-like appearance offers evidence that ants evolved from wasp-like in-
sects. A review of the literature on the anatomy of ants indicates that the
males have been studied only in a very few of the numerous genera and
species. These studies have brought to light anatomical and histological
differences in the organs and systems at the subfamily, generic, and in some
even at the species level, that are of phylogenetic importance. As stated by
Wheeler (1910) the males are stable in structure and, if more readily ob-
tainable, would be valuable in the taxonomy of the group.

The digestive system of male ants was first described by Meinert (1861)
in a formicine, Formica rufa, and a myrmicine, Myrmica ruginodis. He
illustrated differences in the structure of the proventriculus in these two
species; this was the first indication of the importance of this organ as a
basis for determining phylogenetic relationships of various genera of ants.
Forel (1878) and Emery (1888) used the diverse features of the proventric-
ulus for taxonomic purposes. Descriptions of the anatomy of the digestive
system that followed were those of Janet (1902) in the gaster of a myrmicine,
Myrmica rubra, Mukerjee (1926) in an Old World doryline, Dorylus labia-
tus, Marcus (1953) in a dolichoderine, Dorymyrmex emmaericaellus. A
comprehensive and comparative study of the proventriculus in ants was
made by Eisner (1957), who formulated a phylogenetic dendrogram in which
he placed the dorylines parallel to the poneroid complex but with a different
origin than had been postulated for other groups of ants. More recent ob-
servations of this organ are those of a ponerine, Rhytidoponera me tallica,
(Hagopian 1963), and two species of myrmicines, Myrmica rubra, (Trachi-
mas 1967), and Solenopsis invicta (Buren 1972) = S. saevissima richteri
Forel (Tice 1967). The present investigation has revealed the absence of a
crop and a proventriculus in both the adult and pupa of the male of Aenictus
gracilis. Prior to Eisner’s paper (1957), Brown (1954) in a study on the phy-
logeny of ants stated that the precise affinities of the dorylines are unknown,
and it is possible that they are diphyletic in origin. Gotwald (1969) in his
exhaustive study of the mouthparts of 104 species of ants revealed distinct
differences between the New World and Old World dorylines, and in the
Old World doryline genera between Dorylus and Aenictus. Thus he advo-
cated a triphyletic origin for the dorylines separating the tribes Dorylini,
Aenictini, and Ecitonini from one another. Schneirla (1971) considered do-
rylines as monophyletic in origin on the basis of functional and behavioral
evidence.

In addition to the above mentioned papers on the digestive system of
ants, three other reports have been published on the internal anatomy of the

VOLUME LXXXVIII, NUMBER 1

17

New World doryline workers (Whelden 1963, Gotwald 1971, Gotwald and
Kupiec 1975).

The present work is the first comprehensive anatomical and histological
description of the digestive system of the male adult and pupa of the Old
World doryline Aenictus gracilis. Comparisons are made with those of the
other dorylines previously described.

Materials and Methods

The 12 male adults, and the 12 pupae at an advanced stage of development
used in this study were collected by the late Dr. T. C. Schneirla from a
bivouac on an eastern mountain slope above Dumaguete City, Negros Is-
land, Philippine Islands, in 1962. The specimens were preserved in 70%
ethyl alcohol. Each specimen was mounted on paraffin in a Syracuse watch
glass and dissected under 70% ethyl alcohol. Whole mounts of organs were
prepared by Lynch’s precipitated borax-carmine method (Galigher and Koz-
loff 1971), and specimens for histological study were processed by a double
infiltration technic (Trombetta and Forbes 1977). Frontal, sagittal, and trans-
verse serial sections were cut and stained with Harris’ haematoxylin, coun-
terstained with eosin and mounted in euparal. Drawings were made with the
aid of a camera lucida and a B & L trisimplex microprojector.

Anatomy of the Digestive System

The anteriormost part of the alimentary tract in the adult and pupa is the
cibarium that leads into a wide dorsoventrally compressed buccal tube. The
thin-walled buccal tube is broader anteriorly, and in the adult the posterior
floor opens into an elongated, flattened infrabuccal chamber that extends
beneath the anterior pharynx (Fig. 1). In the pupa an irregular recess is
located beneath the anterior floor of the buccal tube, where the infrabuccal
chamber is normally found in ants (Fig. 2). The pharynx that continues is
distinguishable into a broad anterior, an indented middle, and a tubular
posterior region. The lateral margins of the anterior region become pro-
gressively more sclerotized posteriorly into the indented region. Bundles of
muscle fibers extend from the dorsal surface of the anterior region to the
inner, anterior surface of the head capsule. Arising from the dorsolateral
wall on each side of the posterior region of the pharynx is a duct that divides
into numerous branching, and convoluted tubular postpharyngeal glands.
These glands extend anteriorly as far as the buccal tube and laterally and
posteriorly over the front part of the brain. The posterior end of the pharynx
continues as a narrow oesophagus passing between the brain dorsally and
the suboesophageal ganglion ventrally and through the neck, the thorax,
and the petiole into the gaster. In the gaster, about the beginning of the 4th

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Fig. 1. Diagram of a lateral dissection of the adult male of A. gracilis.

ABBREVIATIONS

A, anus; AB, aedeagal bladder; AG, accessory gland; AGD, accessory gland duct; Ao, aorta;

B, basal cell layer; Br, brain; BR, basal ring; BT, buccal tube; CV, cardiac valve; DC, DC1,
DC2, DC3, digestive cells; DM, dilator muscle; Dt, ductule; E, epithelium; ED, ejaculatory
duct; Ey, lower part of the eye; Gl, G2, G3, G4, gastric ganglia; GC, glandular cell; H, heart;
I, inner cell layer; IC, infrabuccal chamber; In, intestine; It, intima; IV, inner genitalic valve;
L, labium; Lu, lumen; M, mandible; MC, muscle coat; MG, mandibular gland; MT, Malpighian
tubule; Oc, ocellus; Oe, oesophagus; OL, optic lobe; OV, outer genitalic valve; PC, preoral
cavity; Pe, petiole; PeG, petiolar ganglion; PG, postpharyngeal gland; Ph, pharynx; R, rectum;
RC, regenerative cell; Re, recess; RP, rectal pad or gland; Rv, reservoir; RV, rectal valve;
SD, salivary duct; Se, secretion; SG, salivary gland; SI, salivarium; SoG, suboesophagal gan-
glion; T, testis; Tl, T2, T3, thoracic ganglia; Vn, ventriculus; VNC, ventral nerve cord; IX,
9th abdominal sternum; X, 10th abdominal tergum

abdominal segment, the oesophagus projects into the ventriculus and forms
a cardiac valve. The ventriculus is the largest organ of the alimentary tract
and extends to the middle of the 5th abdominal segment. Its anterior half
is broadly rounded, the posterior half is narrower and has a dorsal concav-
ity. The distal end of the ventriculus is constricted at the junction with the
intestine. The proximal region of the intestine is the slightly wider pylorus
where the Malpighian tubules enter. The rest of the intestine occupies the
5th through the 7th abdominal segments, and about halfway along it bends
dorsally, increases slightly in diameter, projects into the rectum, and forms
a rectal valve. The rectum is a thin-walled, pyriform sac in the 8th and the
9th abdominal segments and lies above the genital chamber. The inner wall
of the anterior region of the rectum has 3 rectal pads; two are lateral and
one is ventral in position. The rectum tapers to the anus that opens to the
outside on the ventral surface of the 10th tergum.

VOLUME LXXXVIII, NUMBER 1

19

Figs. 2-3. 2. Photomicrograph of a sagittal section of the head of the male pupa to show

the buccal tube and pharynx. x450; 3. Photomicrograph of a sagittal section of the cardiac
valve and anterior part of the ventriculus of the adult male. x400.

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Histology of the Digestive System

The epithelium of the buccal tube and the infrabuccal chamber is com-
posed of a single layer of small, squamous cells with very small, centrally
located nuclei and is lined by an intima. The intima on the roof of the buccal
tube is thinnest anteriorly, thickest, transversely ridged and weakly scler-
otized in the midregion, and thinner and more sclerotized with small hair-
like processes in the posterior region. The intima on the floor and in the
infrabuccal chamber is very thin. The epithelium is not surrounded by a
muscle coat in this region. In the pupa, the epithelium of the buccal tube is
composed of cuboidal to low columnar cells. The intima throughout this
region is uniformly thicker than that in the adult and has hair-like processes
along the floor pointing toward the preoral cavity.

The epithelium of the pharynx is composed of a single layer of flattened
cells with oval nuclei. The intima in the proximal part of the anterior region
on the roof and floor is thick, sclerotized and forms plate-like layers, and
sharp, hair-like processes on the floor project forward. Muscles are present
only on the roof of this region and consist of the oblique fibers that extend
to the head capsule and some longitudinal bundles that extend from front
to back. In the distal part of this anterior region, the lateral walls are curved
dorsally, and here the intima is more sclerotized than on the roof and floor.
There are three bundles of longitudinal muscle fibers separated by two bun-
dles of the oblique fibers. In addition, on the ventral side transverse muscle
fibers extend between the ventrolateral margins of the pharynx. The middle
indented region has heavily sclerotized, sharp, dorsal extensions of the lat-
eral margins and the lumen is U-shaped. The muscles here consist of lon-
gitudinal and the oblique fibers in the dorsal concavity and transverse fibers
that connect the dorsolateral arms. The distal region of the pharynx has an
intima that is thicker along the floor. The muscles in this region are a con-
tinuation of those of the indented region. In the pupa, low columnar cells
form the epithelium of the pharynx and the intima is thin and not sclerotized.

The postpharyngeal glands have a single-layered cuboidal epithelium. The
cytoplasm is acidophilic, granular, and vacuolated, and the small oval nuclei
are basally located. The glands and ducts are lined by a thin intima bearing
long, hair-like projections. The lumina are devoid of any secretion. In the
pupa the epithelium of these glands is folded and composed of low cuboidal
or squamous cells with spherical or flattened nuclei. The hair-like projec-
tions of the intima are less numerous than in the adult, and the lumen is
more spacious (Fig. 2).

The oesophagus has a low cuboidal epithelium, longitudinally folded in-
ward to form 4 or 5 ridges, and covered by a thin intima. Toward the distal
end the folds are reduced in height and the lumen is wider. The muscle coat
is composed of inner longitudinal fibers, some extending into the epithelial

VOLUME LXXXVIII, NUMBER 1

21

folds, and outer circular fibers. Toward the distal end a layer of 1 or 2
longitudinal fibers is present around the circular layer. In the pupa epithelial
folds are small and the lumen is wider.

In the adult and pupa the cardiac or stomodaeal valve has a reflected, low
cuboidal epithelium (Fig. 3). The intima terminates at the base of the outer
walls of the valve. No muscle fibers are present within the valve.

The epithelium of the ventriculus is composed of large columnar, digestive
or secretory cells, and small pyramidal, regenerative cells (Fig. 3). Some of
the digestive cells are tall and narrow and their free ends are swollen and
globular. In many of these cells the nucleus has migrated upward to the
globular region and is compact and irregular in shape. The cytoplasm is
neutral in its staining reaction and has basophilic granules throughout. In
some of these cells granules are concentrated toward the free margin. The
globular ends of some have ruptured and discharged the granular material
into the lumen of the ventriculus. The regenerative cells are widely scattered
clusters of a few cells each. They have a basophilic cytoplasm and oval
nuclei. The muscle coat surrounding the epithelium consists of an inner
circular and an outer longitudinal layer that are about equal in thickness;
the fibers of the outer layer are widely separated. Sections of the pupal
ventriculus show that the epithelium of the mid and posterior regions is
disintegrating, while that of the anterior and some of the midregions is re-
generating (Figs. 4a and b). The disintegrating cells are columnar in shape,
and their basally located nuclei are irregular in shape. The cytoplasm is
filled with fine, basophilic granules and small vacuoles. The free surfaces of
the cells are broken and the cytoplasmic material is released into the lumen.
The regenerating cells in the anterior ventral region are low, broad cells
with basally located, compact, spherical nuclei. The cytoplasm is finely
granular and homogeneous throughout with a thin compact layer of granules
just underneath the free surface. On the anterior dorsal region the cells are
tall, narrow columnar cells and the free surface is swollen and contains an
acidophilic secretion. In the midregion these cells on the ventral side are
also short and broad while those on the dorsal side are tall and narrow. The
cytoplasm of these cells in this region is uniformly granular, and the nuclei
are basally located and ellipsoidal in shape.

In the adult and pupa the pylorus is divided into anterior and posterior
portions by an inward fold of the epithelium (Fig. 5). Low columnar cells
in the anterior region, taller and wider cells over the fold, and small cuboidal
cells in the posterior region form the epithelium. The cytoplasm is compact,
finely granular and acidophilic. The intima, seen in some sections just behind
the entrance of the Malpighian tubules, is always more distinct over the
epithelial fold. Inner circular and outer longitudinal fibers form the muscle
coat.

The intestine in the adult and pupa has a folded, cuboidal epithelium

22

NEW YORK ENTOMOLOGICAL SOCIETY

Figs. 4-7. 4. Photomicrographs of sagittal sections of the ventriculus of the male pupa.

x325. a. Dorsal wall and b. Ventral wall; 5. Photomicrograph of a sagittal section of the
ventriculus and pylorus of the adult male. x215; 6. Photomicrograph of an oblique section of
the rectum of the adult male to show a rectal pad. x225; 7. Photomicrograph of a sagittal
section of the head of the adult male to show the mandibular gland. x300.

VOLUME LXXXVIII, NUMBER 1

23

covered by a distinct intima. The cells of the folds are taller, but the folds
diminish in height and disappear toward the distal region. The lumen con-
tains a finely granular basophilic secretion. An inner very thin layer of
longitudinal muscle, a middle compact layer of circular muscle, and an
outer layer of longitudinal fibers form the muscle coat. The epithelium and
the muscle coat project into the rectum and form a rectal valve. Here, the
circular muscle layer is thick and forms a sphincter.

In the adult and pupa the rectal epithelium is a single layer of flattened
squamous cells covered by an intima. The nuclei are spindle-shaped. The
muscles consist of a single layer of circular fibers and scattered longitudinal
fibers. A heavier band of circular muscle surrounds the anus and forms the
anal sphincter. Each rectal pad is composed of an inner tall, columnar layer
and a basal cuboidal layer separated by a lumen (Fig. 6). The inner columnar
layer of cells is a continuation of the epithelium of the rectum, and the
transition from the squamous to columnar cells is abrupt. The cytoplasm of
the inner and the basal cells is vacuolated and has basophilic granules. The
inner cells are faintly striated toward the lumen between the cell layers. The
intima covering the rectal pad is slightly thicker than that lining the rectum.

The adult and pupa have 18 to 20 long, slender Malpighian tubules that
open into the anterior end of the intestine at the pylorus (Fig. 5). They lie
coiled around the intestine and the accessory glands of the reproductive
system, some extending anteriorly as far as the proximal region of the ven-
triculus and some posteriorly as far as the rectum. The epithelium is com-
posed of a layer of 4 or 5 large cuboidal cells with a striated free surface.
The large nucleus has a prominent nucleolus and chromatin material. The
cytoplasm has coarse basophilic granules. The lumen is filled with a neutral,
granular secretion.

The salivary glands are microscopic in size and are located in the lateral
regions of the prothorax (Fig. 1). In the early stages of this work when the
organs and systems of the adults and pupae were dissected and studied as
whole mount preparations, the salivary glands and the gland ducts were not
recognizable. The salivary duct in the head and the lateral ducts in the neck
region were found. Examination of the serial sections did reveal the glands
and their ducts, and the following description is based on these observa-
tions. The glands are simple branched acinar glands and the acini are clus-
tered or distributed singly and interspersed in the fat cells. Each acinus
consists of a layer of pyramidal cells around a small lumen lined by a thin
intima and built on a distinct basement membrane. The cytoplasm of these
cells in the adult is homogeneous and basophilic with some vacuoles, while
in the pupa it is more vacuolated with large basophilic granules. The nuclei
are spherical or oval and centrally or basally located. The collecting ducts,
the lateral salivary ducts and the common salivary duct are composed of
cuboidal cells with ellipsoidal nuclei. The lumen of the salivary ducts, lined

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NEW YORK ENTOMOLOGICAL SOCIETY

with a heavy intima that has annular thickenings, is tortuous and forms
U-shaped loops within the cells of the ducts. The common salivary duct
proceeds below the brain and opens into the salivarium which is situated
between the hypopharynx and the labium. Where the salivary duct opens
into the salivarium a dilator muscle extends from the roof of the duct to the
ventral wall of the hypopharynx.

The adult and pupa have a pair of mandibular glands located at the bases
of the mandibles (Fig. 1). Each consists of a glandular portion and a res-
ervoir. The glandular portion is hemispherical with about 12 pyriform glan-
dular cells with intercellular spaces between them (Fig. 7). These intercel-
lular spaces are lined with a thin cuticle. The glandular portion is encased
in a thin membrane. The cells open by small ductules into the thin-walled
reservoir, and the intercellular spaces converge toward the reservoir. A
short duct leading from the reservoir opens laterally at the base of the
mandible. The cytoplasm of the glandular cells is homogeneous with fine
basophilic granules. The spherical or oval nucleus has a prominent nucleolus
and coarse chromatin granules. The membrane surrounding these cells is
composed of a layer of squamous cells covered by a thin cuticle. The wall
of the reservoir and the duct has a layer of squamous cells lined by a wrin-
kled intima.

Discussion

The adult has an infrabuccal chamber while the pupa has an anterior
recess in that region. The epithelium of the buccal tube and the pharynx
consists of squamous cells in the adult and cuboidal or low columnar cells
in the pupa. The pharynx of A. gracilis resembles in structure the Old World
doryline Anomma wilverthi (Bugnion, 1930), and in general histology the
New World Eciton (Whelden, 1963) and Cheliomyrmex morosus (Gotwald,
1971). However, variations are seen in the arrangement of spines along the
inner surface.

The epithelium of the postpharyngeal glands is folded in the pupa of A.
gracilis and not so in the adult.

The oesophageal epithelium of the pupa of A. gracilis has smaller folds
and the lumen is wider when compared to the adult. The long, acute spines
reported in the queens and workers of two species of Eciton (Whelden,
1963) are absent here.

A crop and proventriculus generally present in ants between the oespha-
gus and ventriculus are absent in A. gracilis and in the Old World D. la-
biatus. Eisner (1957) described the proventriculus in E. hamatum as degen-
erate. Whelden (1963) has reported a difference in the position of the small
crop in queens and workers of two species of Eciton ( hamatum and bur-
chelli), and the presence of a small proventriculus in both species. In the

VOLUME LXXXVIII, NUMBER 1

25

worker of C. morosus the crop and ventriculus lie juxtaposed and are con-
nected by a reduced membranous proventriculus (Gotwald and Kupiec,
1975). There is no mention of a cardiac valve at the posterior end of the
oesophagus in D. labiatus (Mukerjee, 1926). A simple cardiac or stomodaeal
valve is reported to be present in E. hamatum (Eisner, 1957) and in C.
morosus (Gotwald, 1971). In the adult and pupa of A. gracilis the posterior
end of the oesophagus continues into the ventriculus and forms a cardiac
valve; this valve is shorter in the pupa.

The ventriculus of the adult and pupa of A. gracilis resembles in general
form that of the ponerine male, Rhytidoponera me tallica, (Hagopian, 1963)
in having a dorsal concavity in its posterior half. The New World doryline

C. morosus worker has a tubular posterior portion that is referred to as the
pre-Malpighian tubule tract (Gotwald, 1971). This has not been reported in
any of the other species described (Mukerjee, 1926; Whelden, 1963). The
ventricular epithelium of the adult of A. gracilis is composed of large co-
lumnar digestive cells and small, scattered, pyramidal regenerative cells. In
the pupa, however, three types of digestive cells, each restricted to a region
of the ventriculus, were recognized, and these represent stages in the ma-
turity of the epithelium.

A pylorus that is divided into anterior and posterior portions by an inward
fold of the epithelium is present in the adult and pupa of A. gracilis. This
is indicated as a pyloric constriction in D. labiatus (Mukerjee, 1926). Such
a pylorus is not reported in the New World species described (Whelden,
1963; Gotwald, 1971).

The intestine is a tube of uniform diameter in all ants described. In D.
labiatus, however, it is reported to be divisible into an anterior narrow ileum
and a posterior broader colon (Mukerjee, 1926).

At the junction of the intestine and the rectum a rectal valve with a
sphincter is present in the adult and pupa of A. gracilis. A structure com-
parable to this has not been reported in any of the other dorylines described
(Gotwald, 1971; Mukerjee, 1926; Whelden, 1963). There seems to be no
consistency in the number of rectal pads or papillae; small rectal glands in

D. labiatus male (Mukerjee, 1926), usually 3 and rarely 6 in Eciton workers
and frequently 6 and infrequently 3 in Eciton queens (Whelden, 1963); 2
lateral papillae in the worker of C. morosus (Gotwald, 1971). The present
investigation revealed the presence of 3 rectal pads in the adult and pupa
of the male, two lateral and one ventral.

Maxillary glands situated on either side of the infrabuccal chamber and
opening into the posterior part of the buccal tube have been reported in the
male ponerine ant, R. metallica (Hagopian, 1963), in the queens and work-
ers of the two species of the New World doryline, Eciton (Whelden, 1963),
and in the workers of a New World doryline, C. morosus (Gotwald and

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NEW YORK ENTOMOLOGICAL SOCIETY

Kupiec, 1975). These glands are not present in adults and pupae of the male
A. gracilis.

There are 18-20 Malpighian tubules in the adults and pupae of the male
of A. gracilis. Mukerjee (1926) has reported the presence of numerous tu-
bules in the male of D. labiatus. Analysis of the Malpighian tubule count
made by Gotwald (1971) indicates that the range in number of tubules in
workers is less species specific. However, the mean numbers of tubules are
distinct for several species and he assumes that they could be correlated
with the body size; a higher mean for the larger Labidus species, and a
lower mean for the smaller Neivamyrmex species. The present count of 18-
20 for one of the smallest of the dorylines, Aenictus, does not support
Gotwald’ s assumption.

Salivary glands and gland ducts in the adult and pupa of A. gracilis are
very small and interspersed in fat cells. The glands are simple branched
acinar glands, and the ducts are typical in being straight with a tortuous
lumen. In the other dorylines described these glands are of different types;
short, thick, and branched tubules in Eciton workers and queens (Whelden,
1963), and small cylindrical lobes in workers of C. morosus (Gotwald, 1971).

Whelden (1963) reported much variation in shape and in size of the man-
dibular gland and the reservoir in the two species of Eciton. In the males
of these species he describes the posterior end of this gland as being fre-
quently bifurcate, the two branches passing one above, the other below the
optic nerve. The number of secretory cells in these glands varies from 100
to 1400. The mandibular gland in C. morosus (Gotwald and Kupiec, 1975)
is composed of a few irregularly-shaped cells with distinct nuclei; a reservior
was not reported. The glandular portion of A. gracilis resembles in all re-
spects that of the ponerine, R. metallica, (Hagopian, 1963). A. gracilis has
a reservoir into which the cellular ductules and the intercellular spaces of
the glandular portion open.

Gotwald and Kupiec (1975), analyzing the existing information on the mor-
phology, behavior and geographical distribution of the doryline tribes indi-
cate that the subfamily Dorylinae as presently constituted, is triphyletic.
The three lineages are the Ecitonini-Cheliomyrmecine, the Dorylini, and the
Aenictini. They advocate the retention of the subfamily Dorylinae to include
the tribe Dorylini and make a case for the creation of a subfamily Ecitoninae
already introduced by Brown (1973) to include the tribes Ecitonini and Che-
liomyrmecini. The status of Aenictini, they conclude, remains to be deter-
mined by further investigation.

The present investigation of the anatomy and histology of the digestive
system has brought to light several features of phylogenetic importance. In
the absence of a crop, a proventriculus, and maxillary glands, Aenictus is
distinctly different from the New World forms. The presence of a dorsal
concavity in the posterior half of the ventriculus, the pylorus internally

VOLUME LXXXVIII, NUMBER

27

divided into anterior and posterior portions, and the presence of a rectai
valve with a sphincter are the unique features exhibited in the digestive sys-
tem of Aenictus. These structures lend support to the elevation of the tribe
Aenictini to a subfamily rank, and add to the concept of the triphyletic
origin of the dorylines.

Literature Cited

Brown, W. L., Jr. 1954. Remarks on the internal phylogeny and subfamily classification of
the family Formicidae. Insectes Sociaux 1:21-32.

. 1973. A comparison of the Hylean and Congo-West African rain forest ant faunas, p.

161-185. In B. J. Meggers, E. S. Ayensu, and W. D. Duckworth, eds. Tropical forest
ecosystems in Africa and South America: a comparative review. Smithsonian Inst.
Press, Washington, D.C.

Bugnion, E. 1930. Les Pieces buccales, le sac infrabuccal et la pharynx des fourmis. Bull.
Soc. Royal Entomol. Epypte 40:85-210.

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

Eisner, T. 1957. A comparative morphological study of the proventriculus of ants. Bull. Mus.
Comp. Zool. 116:441-490.

Emery, C. 1888. Uber den sogenannten Kaumagen einiger Ameisen. Zeitsch. f. wissensch.
Zool. 46:378-412.

Forel, A. 1878. Etudes myrmecologiques en 1878 (premiere partie) avec l’anatomie du gesier
des fourmis. Bull. Soc. Vaudoise Sci. Natur. 15:337-392.

Galigher, A. E. and E. N. Kozloff. 1971. Essentials of practical microtechnique. Lea and
Febiger, Philadelphia, Pa. 531 pp.

Gotwald, W. H., Jr. 1969. Comparative morphological studies of the ants, with particular
reference to the mouth parts (Hymenoptera: Formicidae). Cornell Univ. Agr. Exp. Sta.
Mem. 408, 150 pp.

. 1971. Phylogenetic affinities of the ant genus Cheliomyrmexd Hymenoptera: Formi-
cidae). J. N.Y. Entomol. Soc. 79:161-173.

. and B. M. Kupiec. 1975. Taxonomic implications of doryline worker ant morphology:

Cheliomyrmex morosus (Hymenoptera: Formicidae}. Ann. Entomol. Soc. Amer.
68:961-971.

Hagopian, M. 1963. An anatomical and histological study of the male ponerine ant, Rhyti-
doponera metallica F. Smith (Formicidae, Hymenoptera). Ph.D. Dissertation, Fordham
University.

Janet, C. 1902. Anatomie du gaster de la Myrmica rubra. Carre et Naud, Paris 68 pp.
Marcus, H. 1953. Estudios mirmecologicos. Folia Universitaria, Cochambamba, Bolivia
6:17-68.

Meinert, F. 1861. Bitrag til de danske Myrers Naturhistorie. Kongl. Danske Vidensk. Selsk.
Skrift. Bind. 5:273-341.

Mukerjee, D. 1926. The digestive and reproductive systems of the male ant, Dorylus labiatus
Schuck. J. Asiatic Soc. Bengal, n.s. 22:87-92.

Schneirla, T. C. 1971. Army ants. A study in social organization. H. Topoff, ed. W. H.
Freeman and Co., San Francisco, CA. 349 pp.

Tice, M. 1967. The anatomy and histology of some of the systems of the male of the imported
fire ant, Solenopsis saevissima richteri Forel (Hymenoptera: Formicidae) Ph.D. Dis-
sertation, Fordham University.

28

NEW YORK ENTOMOLOGICAL SOCIETY

Trakimas, W. 1967. An anatomical and histological study of the male myrmicine ant, Myrmica
rubra L. (Hymenoptera: Formicidae). Ph.D. Dissertation, Fordham University.

Trombetta, L. D. and J. Forbes. 1977. A double infiltration technic for serial sectioning of
heavily sclerotized insects. J. Histotechnol. 1:25-26.

Wheeler, W. M. 1910. Ants, their structure, development, and behavior. Columbia Univ.
Press, New York. 663 pp.

Whelden, R. M. 1963. The anatomy of the adult queen and workers of the army ants Eciton
burchelli Westwood and Eciton hamatum Fabricius. J. N.Y. Entomol. Soc. 71:14-30
and 90-115.

Department of Biological Sciences, Fordham University, Bronx, NY
10458.

Received for publication October 16, 1979.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(l), 1980, pp. 29-32

FECELIA BIORBIS N. SP. (HETEROPTERA: PENTATOMIDAE),

A NEW SPECIES FROM HAITI

Joseph E. Eger, Jr.

Abstract. — A third species of Fecelia, F. biorbis n. sp. from Haiti, is
added to the genus. It is described, figured and compared to the other
species of Fecelia.

The genus Fecelia Stal, 1872, includes two previously described species,
F. minor (Vollenhoven, 1868) and F. nigridens (Walker, 1867). It was re-
cently revised by Grazia (1976); however, a single distinctive specimen in
the collection of the American Museum of Natural History was not included
in her work. This specimen represents a new species and is described here.

Fecelia biorbis n. sp.

Color above chestnut brown, scutellum somewhat lighter. Punctation
dense, irregularly distributed, concolorous or piceus (Fig. 1). Venter simi-
larly colored; punctation dense, piceus. Body length with membranes
18.6 mm.

Head 3.4 mm wide across eyes, 3.3 mm long at meson, evenly punctured,
transversely rugose. Juga strongly surpassing tylus, divergent anterior to
tylus; apices acute. Vertex flat, interocular width 2.2 mm. Antennal seg-
ments 1-4 nearly concolorous with dorsum, slightly red-tinged; last segment
lighter; first segment reaching apex of jugum; length of segments 1-5: 1.5;
2.4; 2.7; 2.9; 2.6 mm. Bucculae toothed anteriorly, evanescent posteriorly.
First rostral segment surpassing bucculae by approximately Vs length of
segment; second segment extending nearly to anterior margin of mesocoxae;
third segment reaching posterior margin of metacoxae; apex of rostrum
extending to middle of third visible abdominal sternite.

Pronotum 3.6 mm long at meson; width between apices of humeri 12.5
mm; disc with broad longitudinal band devoid of punctation, outlined by
punctation. Two roughly circular, lightly punctured maculae present poste-
rior to cicatrices, separated from latter by band of moderately dense punc-
tation, these maculae outlined by piceus punctation. Cicatrices with distinct
tubercle, lightly punctured near tubercle. Humeri strongly developed,
abruptly narrowing near apex into acute spines, densely punctured above.
Anterolateral margins strongly concave, denticulate.

Scutellum 5.0 mm wide at anterior margin, 6.4 mm long at meson. Punc-
tation large and irregularly distributed at base, becoming smaller and more

30

NEW YORK ENTOMOLOGICAL SOCIETY

Figs. 1-2. F . biorbis. 1. Dorsal aspect; 2. Female genital plates; first gonocoxae (GX1),
second gonocoxae (GX2), eighth paratergites (PT8), ninth paratergites (PT9), triungulum (TR).
Dimensional lines equal 1 mm.

VOLUME LXXXVIII, NUMBER 1

31

evenly distributed posteriorly, piceus on roughly anterior third of scutellum
and posteriorly near apex. Margin of basal angles and anterior V\ of lateral
margins piceus.

Each corium finely and densely puntured, punctation becoming somewhat
smaller and more dense posteriorly; with small, evenly distributed concol-
orous calli. Each corium with large, central piceus ring surrounding nearly
circular macula, the latter only lightly and shallowly black punctured. Poste-
rior margins convex, with finely broken submarginal piceus line.

Each connexivum strongly exposed; piceus punctation most dense ante-
riorly on each segment. Width of abdomen at widest point 9.8 mm.

Prosternum weakly sulcate; mesosternum carinate; metasternum flat. Os-
tiolar rugae spatulate, extending roughly lA distance from inner margin of
ostiole to outer metapleural margin. Legs with femora acutely toothed on
superior face at apex; superior face of tibiae flat or very shallowly sulcate.
Second to fifth visible abdominal sternites broadly and shallowly sulcate
mesially. Length of visible sternites 2-6 at meson (first segment not visible
mesially): 0.8; 1.1; 1.4; 1.4; 2.0 mm. Postero-lateral angles of segments 2-5
provided with small acute tooth; this tooth becoming more elongate and
spinose on segment 6.

Mesal margins of first gonocoxae with acute apices, narrowly separated
at base, diverging posteriorly, exposing triungulum and extending onto sec-
ond gonocoxae. Length of first gonocoxae 1.7 mm, width at widest point
1.6 mm. Eighth paratergites acutely spinose posteriorly; ninth paratergites
rounded posteriorly (Fig. 2).

Holotype.— Female, labelled (a) Furcy, 4000'; Haiti VII-10-56; B&B Val-
entine. (b) Sweeping in bushy ravine, (c) Haylini. Deposited in the American
Museum of Natural History, New York, N.Y.

Distribution. — Haiti.

Comments. — This species basically agrees with Grazia’s description of
the genus and resembles its congeners in the length and shape of the ostiolar
rugae, the presence of apical femoral spines, and the median furrow which
occurs on the basal abdominal sternites. It is most closely allied to F. ni-
gridens and agrees with this species in size, shape of the female genital
plates, and in the acute, divergent juga. The single representative of the
species described here is sufficiently distinct that dissection of the genitalia
was not deemed necessary. F. biorbis can be readily distinguished from the
two other species of this genus by the abruptly narrowed humeri and the
piceus ring on each corium.

Acknowledgments

I would like to thank Mr. C. W. Agnew of Texas A&M University for
the illustrations which appear in this paper, Dr. L. H. Rolston of Louisiana

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NEW YORK ENTOMOLOGICAL SOCIETY

State University for his advice, encouragement and criticism. Dr. J. C.
Schaffner of Texas A&M University for review of the manuscript, and Dr.
R. T. Schuh of the American Museum of Natural History for the loan of
specimens.

Literature Cited

Grazia, J. 1976. Revisao do genero Fecelia Stal, 1872 (Heteroptera, Pentatomidae, Pentatom-
ini). Rev. Brasil. Biol. 36(l):229-37.

Stal, C. 1872. Enumeratio hemipterorum II. Kongliga Svenska Vetenskaps-Akademiens Han-
dlingar 10(4): 1-159.

Vollenhoven, S. C. S. van. 1868. Diagnosen van eenige nieuwe soorten van Hemiptera Het-
eroptera. Versl. Ak. Amst. Nat. 2(2): 179.

Walker, F. 1867. Catalogue of the specimens of Hemiptera-Heteroptera in the collection of
the British Museum. Part 2:241-417.

Dept, of Entomology, Texas A&M University, College Station, Texas
77843.

Approved for publication as TA- 15694 by Director, Texas Agricultural
Experiment Station.

Received for publication December 11, 1979.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(l), 1980, pp. 33-82

ABSTRACTS OF PAPERS PRESENTED AT THE
51st ANNUAL MEETING
EASTERN BRANCH

ENTOMOLOGICAL SOCIETY OF AMERICA
HERSHEY, PA.

September 26, 27 and 28, 1979

Migration, distribution, and habitat preferences of Blattella germanica (L.)
in urban, low-income apartments (Dictyoptera: Blattellidae). R. C. Akers
and W. H. Robinson, VPI and State Univ., Blacksburg, VA 24061

This study monitored the dispersal behavior and determined density dis-
tribution and habitat preferences of B. germanica in 3 urban, low-income
apartment buildings. Three traps (1-qt. jars) were placed in the kitchen,
pantry, and bathroom of each apartment. Two hundred nymphal and virgin
adult phenotypic mutant German cockroaches were released into estab-
lished apartment building populations and trapped at weekly intervals. Trap-
ping proved that this insect could migrate between apartments and integrate
with the existing population. Trapping disclosed that the existing German
cockroach population was not distributed evenly throughout the buildings.
In each building at least one location ( = apartment) the number of captured
cockroaches was significantly greater than at any other location. In bldg. A,
80% of total cockroaches captured were from apts. 4 and 5; bldg. B, 75%
were from apt. 1; bldg. C, 55% were from apt. 6. This focal location cor-
responded to the apartment with the most severe sanitation problem. Hab-
itat preferences differed at the trapping sites in the 3 buildings: bldg. A,
kitchen — 22%, pantry — 39%, bathroom — 39%; bldg. B, kitchen — 21%, pan-
try— 37%, bathroom — 42%; bldg. C, kitchen — 41%, pantry — 21%, bath-
room— 38%. The German cockroach dominated other cockroach species in
the urban, low-income apartments studied. Laboratory-reared cockroaches
were able to disperse and integrate with the established population.

Comparative toxicity of etrimfos to gypsy moth larvae and house flies.
M. Akram and A. J. Forgash, Dept, of Entomol. and Econ. Zool., Rutgers
Univ., New Brunswick, NJ 08903

Etrimfos ( 0-0-dimethyl-0-(6-ethoxy-2-ethyl-4-pryimidinyl) phosphoro-
thioic acid ester), a new organophosphate insecticide was tested against
2nd, 3rd, 4th, and 5th instars of gypsy moth larvae ( Lymantria dispar L.)
and susceptible and multi-resistant strains of adult house flies ( Musca do-
mestica L.). For each dosage, 50 insects were tested in five replicates con-
taining 10 insects per replicate. Etrimfos was applied in acetone solution to
the thoracic dorsum with a microapplicator. Results showed an increased

34

NEW YORK ENTOMOLOGICAL SOCIETY

tolerance of gypsy moth larvae to this chemical with larval growth. Individ-
uals of the 3rd-, 4th-, and 5th-instars can tolerate up to 2.6, 5.2, and 28.8
times as much as 2nd-instar larvae. In gypsy moth an active microsomal
oxidative system has been reported, presumably, the increase in tolerance
to etrimfos of later instars is related to the changes in activity of the mixed-
function oxidases. Two susceptible strains (WHO/IN and Wilson) of house
fly showed a relatively high sensitivity to etrimfos. However, Es and A
strains, which were cultured under the pressure of resmethrin and diazinon,
respectively, were 4.5 and 14.2 times tolerant to this compound. Multire-
sistance to insecticides as the result of selection with a single material is well
recognized; presumably, diazinon selection was responsible for increased
tolerance to etrimfos. The lower tolerance of ES suggests a less developed
or different mechanism of resistance than A strain.

Release of oviposition-deterring pheromone by apple maggot flies, Rhago-
letis pomonella (Walsh) (Diptera: Tephritidae) A. L. Averill and R. J. Pro-
kopy, Dept, of Entomol., Univ. of Mass., Amherst, MA 01003

Following egglaying into a fruit, the female apple maggot deposits a mark-
ing pheromone on the fruit surface which deters subsequent oviposition.
Two techniques were developed to accurately quantify the amount of mark-
ing pheromone deposited. 1) Dusting a recently marked fruit with dry mag-
netic toner (Olivetti) used in paper copying machines renders the pheromone
trail readily apparent. 2) Feeding 9 9 amaranth results in a trail which is
conspicuously dyed. Pheromone trail length and area are then measured
with an ocular micrometer. For 14-day-old 9 9 ovipositing in 15 cm diameter
hawthorns, the average length of pheromone trail was 52.8 mm, the average
width .05 mm and the average area 2.7 mm2 (N = 99). Old flies produce
smaller trails than young flies and starved flies deposit much less material
than unstarved flies. There was a low, though significant, positive correla-
tion (r = +0.315) between time spent in ovipositor dragging and area of
pheromone trail. In individual flies, the degree of variability in pheromone
trail area was large among successive dragging bouts within single days.
The presence of amaranth dye (internally confined to the digestive tract) in
the pheromone trail suggests that a portion of the pheromonally active sub-
stances may be released from the digestive tract. Behavioral bioassays of
organs of the digestive and reproductive tracts strongly implicate the hindgut
(as well as the ovary) as a site of pheromone production or accumulation.

VOLUME LXXXVIII, NUMBER 1

35

Oviposition response of the horn fly (Diptera: Muscidae) to environmental
and manure characteristics, N. P. Bacon, West Va. Univ., Morgantown,
WV 26506

Horn fly control efforts have previously been directed primarily at chemical,
biological or sterile methods. Little work on behavioral control has been
carried out. The purpose of this study was to investigate horn fly oviposition
site selection behavior. Horn flies only oviposite on freshly defecated feces
(ca. 10 min), and then only on select pats. The responses of the flies to
selected environmental, manure and host-fly interaction characteristics were
studied on a rural Pennsylvania dairy farm. Fly number response, used to
assess oviposition preferences, indicated that manure pats which were def-
ecated in areas of full sunlight, and on grass or soil substrates attracted the
largest number of flies. Pats which were large in size (22. 8+ cm), tall in
height (4.3+ cm), and had features which gave topographic relief to the pat
(e.g. edge, cracks, depressions) also increased fly response. The expression
of these topographic features was found to be dependent on substrate, pat
moisture content, and pat height. Fly response was least when manure pats
were defecated under roofed areas, at night, on liquid substrates, on pats
with small height-size ratios, and on pats which lacked topographic relief
features.

These results offer another avenue of investigation for biological control
of the horn fly.

The impact of the soil systemic, aldicarb and the foliar insecticide, pirimi-
carb on potato plant growth. B. A. Bajusz, Z. Smilowitz, T. P. Mack, F.
L. Petitt, C. A. Martinka, and M. E. Whalon, Pesticide Res. Lab., Penn-
sylvania State Univ., University Park, PA 16802

In field experiments designed to evaluate the impact of an insect pest on
crop growth, different application rates of insecticides are often applied to
obtain insect populations of different levels. However, to accurately eval-
uate the insect component the impact of these insecticides on the growth of
the plant must be known. A field experiment designed to assess the effect
of different green peach aphid, Myzus persicae (Sulzer), densities on plant
growth was expanded to include tests on the impact of the foliar insecticide
pirimicarb and soil systemic aldicarb on plant growth and development. The
data indicated that both compounds appear to affect potato growth and
development. Pirimicarb applied on a weekly basis to fields containing five
aldicarb rates, 0-35 lb ai/A, reduced foliage biomass throughout the season
but did not appear to affect tuber biomass. Aldicarb appeared to prolong
the period of haulm growth and increasing soil dose delayed plant senes-
cence. The effects observed with increasing aldicarb concentrations closely

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paralleled those associated with decreasing aphid densities, and could have
erroneously been attributed solely to differing aphid numbers. Further ex-
perimentation is currently underway to increase our understanding of these
findings. It was concluded that the effects of chemical insecticides on plant
growth must be known before they are used in plant growth experiments to
manipulate insect population size.

Effect of electromagnetic devices on subterranean termites ( Reticulitermes ),
dry wood termites ( Incisitermes minor), and German cockroaches ( Blattella
germanica). R. Beal, U.S.F.S. Gulfport, MS 39501, M. Rust and D. Reier-
son, U. of Cal., Riverside, CA 92521, D. S. Dalton U.S.E.P.A., Wash. DC
20460 (proj. off.)

There has been a recent increase in the sale of “electromagnetic” pest
control devices for use against pest insects and rodents in private homes
and in commercial and industrial sites. EPA grants to 2 institutions tested
the efficacy of three devices. Field tests with subterranean termites exposed
to the devices in Mississippi for 6 months showed no significant effect on
tubing activity or frequency of stake attack. Field tests in roach infested
apartments in Los Angeles showed no measurable differences in control of
roach populations between exposed and unexposed buildings. Lab tests with
roaches showed no measurable differences on behavior of roaches in a re-
pellent/nonrepellent choice box situation. In addition, starting with either
adult or nymphal roaches there was no significant difference in the resultant
number of roaches after 12 weeks of exposure to the devices. Devices had
no effect on egg capsules of Periplanta americana. In lab tests with dry-
wood termites, no significant differences were found in mortality, amount
of wood consumed, or increase in termite biomass after a 3 month exposure
to the devices. As a result of these tests and tests against 3 rodent species,
EPA initiated enforcement action under the Federal Insecticide, Fungicide
and Rodenticide Act to stop the distribution of these devices because of
false and misleading claims.

The effects of mowing on the guild of sap-feeding insects associated with
Kentucky bluegrass. Richard A. Bean and Robert F. Denno, Department
of Entomology, Univ. of Maryland, College Park, MD 20742

Four species of Auchenorrhyncha dominate the sap-feeding guild of in-
sects associated with Kentucky bluegrass. They are in decreasing order of
abundance, Psammotettix lividellus (Zetterstedt), Graminellci nigrifrons
(Forbes), Doratora sty lata (Boheman) (Cicadellidae), and Delphacodes lu-
tulenta (Van Duzee). The densities of these sap-feeders were measured in
mowed (cut monthly) and uncut plots throughout the year to test the effects

VOLUME LXXXVIII, NUMBER 1

37

of disturbance and habitat modification on the structure of the guild. G.
nigrifrons is a fugitive species colonizing plots as adults in spring developing
a single generation and then emigrating in June. This species achieved equal
densities in both uncut and mowed plots. D. lutulenta and D. sty lata are
residents and overwinter as adults and nymphs respectively. Adults of these
species are mostly brachypterous and significantly lower densities occurred
in the mowed plots. The adults of the resident leafhopper P. lividellus are
macropterous and occurred at equal densities in cut and control plots. Fu-
gitive species were least affected by mowing followed by macropterous
residents. The densities of species that are closely tied to resources (bra-
chypterous) were most affected by habitat perturbation.

Sweet corn insect pest management in New Hampshire. J. S. Bowman and
A. T. Eaton, Univ. of New Hampshire, Durham, NH 03824

A modification of the New Jersey sweet corn IPM program has been of
economic value to participating growers for the past 3 seasons (1977-1979).
The program is based upon utilizing grower personnel trained to scout fields
for European corn borer and fall army worm feeding injury and operate black
light traps to detect corn earworm moth populations. The results of whorl
stage treatments of granular and spray treatments at 15% leaf feeding dam-
age demonstrated economic control of first brood European corn borer with
Dyfonate 10GK at 1.5 lb. ai/A or Furadan 10G at 1.0 lb. ai/A applied as
single treatments or two applications (5 days apart) of Penncap M 2F at 0.5
lb. ai/A. Promising results have also been obtained with Pounce 3.2EC at
0.1 lb. ai/A and Orthene 75S at 1.0 lb. ai/A. During 1977 and 1978 the late
arrival of the corn earworm moths along with low late season populations
resulted in savings of two to five applications of pesticides for an average
value of $18.38 per acre. The number of participating growers has increased
from three in 1977 to ten in 1979 representing 16 percent of the total sweet
corn acreage in New Hampshire. Problems associated with this grower-
operated program include difficulty in training personnel to identify light
trap catches as well as data gaps in scouting reports and light trap catches
when traps are inadvertently not operated. Also, an unreliable source of
black light traps has created problems in bringing new growers into the
program.

Bait sampling for white grubs (Coleoptera: Scarabaeidae) and wireworms
(Coleoptera: Elateridae) in Virginia corn land. S. P. Briggs and W. A. Allen,
Dept, of Entomol., VPI and State Univ., Blacksburg, VA 24061

Nineteen fields in two Virginia counties were used to confirm a solar
baiting technique for wireworms. In Westmoreland County, Virginia, eight

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fields in a corn-small grain-soybean rotation and five in a corn-corn agro-
nomic system were tested. In Montgomery County, Virginia, six fields in
a corn-corn system were used. The bait consisted of 30 cc of a corn-wheat
mixture (1:1 ratio) buried 15.0 cm deep and covered with black and clear
plastic. Baits were retrieved after remaining in the field for 14-15 days and
all wireworms and white grubs were collected. In addition to baiting, 40
cylindrical soil samples (20.3 cm diameter, 38.0 cm deep) per field were
taken to correlate relative sample (bait) populations with absolute sample
(core) populations.

A list of the white grub and wire worm species found was established. The
baiting technique is shown to be effective on both white grubs and wire-
worms in Virginia.

Toxicity of the IGR L7063 to Choristoneura fumiferana (Clem.) (Lepidop-
tera: Tortricidae) and two associated parasites. J. R. Brushwein and J. Gra-
nett, Univ. of Maine, Orono, ME 04469

Laboratory tests of Insect Growth Regulators (IGRs) have demonstrated
high levels of toxicity to the spruce budworm, Choristoneura fumiferana.
Field studies have indicated potential use of IGRs for spruce budworm
control programs. Our purpose was to determine comparative toxicities of
L7063 to spruce budworms and two associated hymenopteran parasites
Apantales fumiferanae Vier. (Braconidae) and Glypta fumiferanae (Vier.)
(Ichneumonidae). Overwintering budworm larvae on white spruce were col-
lected in the laboratory. Emerged second stage budworms were reared on
artificial diet until just prior to the instar tested. Newly molted, unfed 4th
and 5th stage budworms were fed artificial diet containing L7063 until larval
mortality or parasite emergence. Concentrations of L7063 in diet ranged
from 0.063-2.4 ppm. The LC50 for field collected, unparasitized budworms
was between 0.063 and 0.10 ppm. Larval A. fumiferanae and G. fumifer-
anae mortality was determined by dissection of L7063 killed hosts. Para-
sitism rates of A. fumiferanae and G. fumiferanae were determined by host
dissections and by numbers of emerged parasites. Parasitism rates for eight
experiments ranged from 4.6 to 27.9% and 2.9 to 12.9% for A. fumiferanae
and G. fumiferanae, respectively. For all L7063 concentrations tested, little
mortality occurred in stage 1 and 2 A. fumiferanae larvae or stage 1,2, and
3 G. fumiferanae larvae. Diet concentrations of L7063 between 0.42 and 2.4
ppm caused mortality of all 3rd stage A. fumiferanae with death occurring
before emergence from the host.

VOLUME LXXXVIII, NUMBER 1

39

Status of a biological control program against the cereal leaf beetle, Oulema
melanopus (L.) (Coleoptera: Chrysomelidae). T. L. Burger, USDA, APHIS,
Cereal Leaf Beetle Lab., Niles, MI 49120

The cereal leaf beetle, Oulema melanopus (L.), an Eurasian pest of small
grains, was first observed in the United States in 1959 in Berrien County,
MI. Aided by the prevailing winds, its range has expanded primarily to the
East to include 20 states and one Canadian province. Through a cooperative
Federal and State program five hymenopterous parasites: Tetrastichus julis
(Walker); Diaparsis carinifer (Thomson); Diaparsis n.sp.; Lemophagus
curtus (Townes); and Anaphes flavipes (Foerster) have been introduced as
an attempt at multiple species biological control of this pest. To date one
or more of these parasites have been released and/or recovered in 18 states
and one Canadian province. Results obtained from monitoring sites in states
where the program was first initiated, revealed high rates of parasitization
of eggs and/or larvae of the pest. Overall seasonal egg and larvae parasiti-
zation rates of 70 and 90 percent respectively recorded from low, medium,
and high host densities. States, such as MI, IN, OH, PA, and NY, which
experienced economic damage over moderately large areas in the past, no
longer have significant pest populations where high parasitization rates had
been encountered for one to two years. To date no single species of the
introduced parasites has exhibited dominance throughout its established
range in the United States. States are now in the process of assuming total
responsibility for the program.

Growth and survival of Hylotrupes bajulus (L.) larvae under favorable and
unfavorable conditions (Coleoptera: Cerambycidae). K. F. Cannon and W.
H. Robinson, Dept, of Entomol., VPI and State Univ., Blacksburg, VA
24061

Hylotrupes bajulus (L.), the old house borer, is unique among Ceram-
bycids in that it can survive and reproduce in seasoned softwoods. Larvae
bore just under the surface, hollowing out the outermost layer of wood
before continuing deeper into the wood. Larvae seldom break the surface,
thus preventing early disclosure. Detection in wood does not usually occur
until larvae are nearly full grown or adult insects emerge. Severity of damage
produced by this pest is linked to the developmental time of the larvae; it
can live and feed up to 10 years in structural wood. Previous investigations
and reports of H. bajulus in the U.S. have relied on European data and
field observations. The data presented here are based on a laboratory colony
and controlled field experiments of H. bajulus in Virginia. The experiment
was designed to study feeding, growth, and survival of H. bajulus under
favorable and unfavorable environmental conditions. Results of the feeding

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and growth studies reveal that under field and laboratory conditions small
(2.8-86.6 mg) larvae consume more wood per mg body weight than medium
(106.7-197 mg) and large (208.5-430.9 mg) larvae. Excrement (frass) is pro-
portional to consumption. Wood consumption and weight gain are influ-
enced by climatic conditions. Larval survival was not affected by climatic
conditions during the duration of the experiment.

Fungicidal inhibition of Beauveria bassiana, a pathogen of the Colorado
potato beetle ( Leptinotarsa decemlineata Say.) (Coleoptera: Chrysomeli-
dae). R. A. Clark, Dept, of Plant Path, and Entomology, Univ. of Rhode
Island, Kingston, RI 02881

Fungicides used to control Phytophthora infestans, the late blight path-
ogen, may limit the development of epizootics of B. bassiana in CPB pop-
ulations. Using agar-diffusion, 3 fungicides were bio-assayed for inhibition
of B. bassiana. Plates were inoculated with a spore suspension of 106 spores/
ml. Paper discs treated with fungicide were placed on the plates. Plates were
incubated 5 days at 25 C. Inhibition zones indicated B. bassiana was inhib-
ited by 3 fungicides. The insecticide azinphos methyl (Guthion) also had a
synergistic effect with at least one fungicide.

To refine quantification of the degree of inhibition, the fungus was grown
in Emerson’s Broth with doses of the pesticides added to the medium.
Flasks were incubated on a rotary-shake table at 25 C. For 7 days, at 24-hr
intervals, the contents of the culture flasks were filtered, the mycelial mat
dried and weighed. Although conventional fungicides (Manzecob, metiram,
chlorothalmil) retarded mycelial growth rate, the late blight specific exper-
imental CGA 48988 had little effect and thus holds the most promise for use
in integrated pest management for potatoes.

Insemination frequency and sperm precedence in the German cockroach
(Dictyoptera: Blattellidae). D. G. Cochran, VPI and State Univ., Blacks-
burg, VA 24061

A question of fundamental importance to insect reproductive biology is
how frequently a female can be effectively inseminated. The problem can
best be approached by use of genetic-mutant markers which allow positive
proof of parentage. Using the recessive marker rose eye ( ro ), it has been
shown that adult female German cockroaches, Blattella germanica (L.),
can mate effectively with more than one male. The most common period
for remating is between production of successive egg cases, but remating
also took place between the first mating and production of the first egg case.
In the first instance remating occurred in about 20% of the females studied
while in the second instance remating was rare.

VOLUME LXXXVIII, NUMBER 1

41

Because the German cockroach female normally produces only 3 or 4 egg
cases, the later ones with reduced numbers of progeny, the total impact of
second matings on all progeny produced tends to be small. That impact is
further diminished by the prevalence of incomplete sperm precedence which
allows sperm from the first mating to function even after a second successful
mating. These results show that monogamy is common in this species, but
that superimposed upon it is the willingness of the female to remate occa-
sionally. Presumably remating is opportunistic and occurs in response to
sperm depletion in the spermatheca.

Parahost behavior of adult Gasterophilus intestinalis (DeGeer)
(Diptera:Gasterophilidae). S. E. Cope and E. P. Catts, Dept, of Entomol.
and Appl. Ecol., Univ. of Delaware, Newark, DE 19711

In Delaware adult horse bot flies occur from mid June to mid October.
Adult activity was closely observed in the vicinity of grazing equines. In-
dividual flies were followed by a mark and release method. The onset of
daily parahost activity was correlated to temperature. Males hover inter-
mittently around hosts during the mating period and are rarely seen after
mid day. Male flies also defend single or clustered horses from other males
through the use of audible encounters and aggressive pursuit. Female fe-
cundity approximated 1000 eggs. The majority of oviposition was completed
by mid day and the egg-laying pattern varied in location and concentration.
Females are capable of ovipositing the full egg complement in less than 1
hour and exhibit a broad range of tolerance to environmental conditions.
Flight characteristics of females are indicative of physiological age. Lon-
gevity of adults in their natural habitat is 1-2 days. Adults which were
captured in the field and held under laboratory conditions lived for a slightly
longer period of time. These studies represent an advance in the knowledge
of the biology of Gasterophilus intestinalis and help to establish a basis for
future experimentation with regard to the development of trapping devices
as a method of augmenting host treatment as a control measure.

USDA Procedure for acting on new pest problems. R. L. Cowden, USDA,
APHIS, PPQ, Hyattsville, MD 20782

Of the economically important pests that occur in this country, most have
come in from foreign sources. With the current widespread interest and
concern in pesticides, environmental pollution, and energy conservation,
early detection and decision as to what should be done has become increas-
ingly important. It is the function of the New Pest Work Group to review
available data and make recommendations to the Deputy Administrator of
the Animal and Plant Health Inspection Service as to the best courses of

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action concerning newly reported plant pests. Upon detection of a new
insect species of pest potential, a determination is made of its probable
economic importance in this country. Experts from various Federal and
State agencies, universities, industry, and others are called together to dis-
cuss and provide advice on future courses of action. Most new species
detected are likely to be of little economic importance. A second group may
cause concern because of the uncertainty of their pest potential in this coun-
try. Thirdly, there are others which are known economic pests on crops or
in situations similar to those in the area where they are detected. In the
latter two categories, the temporary quarantines, control or eradication,
parasite introduction, crop destruction, and other options are considered
when recommending action.

Herbivorous insects causing mortality of pine seedlings in an old field. W.
J. Cromartie, M. A. Rivera, J. Gfrorer and D. Maser, ENVL, Stockton
State College, Pomona, NJ 08240

As part of a study of establishment of Pitch Pine ( Pinus rigida ), seedlings
1-8 yr old in an abandoned agricultural field were censused, tagged and
sampled during two years. Percentage defoliation was estimated in spring,
summer and fall, 1978, and measured in spring, 1979, on a sample of 30
trees. The principal defoliator was the Red-headed Pine Sawfly ( Neodiprion
lecontei Fitch). Damage by shoot borers (mainly Rhyacionia sp.) and suck-
ing insects, including a scale (Tourney ella sp.) and aphids (Cinara sp.) was
also estimated. Intensity of herbivore attack was variable from tree to tree.
Thirty-five percent were killed or severely stunted by defoliation or loss of
numerous shoots to tip moths. Some trees, however, were virtually unaf-
fected, even when they were adjacent to trees that were killed. Defoliation
levels increased from 1978 to 79, except on trees that were 90% defoliated
prior to spring, 1978; these showed signs of recovery. A correlation was
found between foliar concentrations of potassium and level of defoliation.
It is hypothesized that a positive feedback mechanism is responsible for the
pattern of herbivore attack on young Pitch Pine.

Geographic variation in vagility in the milkweed beetle, Tetraopes tetraop-
thalmus (Forster) (Coleoptera: Cerambycidae). M. A. Davis, Dartmouth
College, Hanover, NH 03755

This study shows that extensive variation in flight and migratory behavior
exists within and between Tetraopes populations. The vagility of milkweed
beetles in different populations in New England was compared by flight
testing the beetles with a still air tethering device.

The vagility of populations varied seasonally, with peak vagility coincid-

VOLUME LXXXVIII, NUMBER 1

43

ing with the peak flowering period of its host plant. Most beetles tested were
relatively sedentary, although each population possessed a certain propor-
tion of beetles that were highly vagile. Populations differed significantly in
the proportion of highly vagile beetles they contained. It is hypothesized
that this geographic variation in vagility is due to differences in the age and
remoteness of milkweed patches throughout the range of Tetraopes.

Factors affecting honey bee Apis mellifera L. (Hymenoptera: Apidae). For-
aging on birdsfoot trefoil ( Lotus corniculatus). G. L. DeGrandi and C. H.
Collison, Penn. State Univ., University Park, PA 16802

Honey bees are essential for commercial seed production of birdsfoot
trefoil because of self-sterility and incompatibility systems that make cross-
pollination necessary. Six commercial varieties, Dawn, Empire, Mackinaw,
Missouri-20, Kentucky-Ecotype, and Viking were evaluated for flower pro-
duction, nectar secretion, and attractiveness to honey bees. Nectar was
removed from one-day-old florets with a micropipette and sugar concentra-
tion determined with a Baush and Lomb refractometer. Both greenhouse
and field studies of first year growth were conducted simultaneously. The
total volume of nectar and sugar concentration between varieties did not
differ significantly at the 0.05 level of probability in either study. Flowers
contained between 0.21-0.25 p\ of nectar, and had a sugar concentration of
14.6%-17.5%. Significant differences were found in flower production, with
Empire producing the most florets (mean of 497/10 plants) throughout the
season. Viking and Kentucky-Ecotype were the first to bloom, and averaged
the least, 21 and 48 florets, respectively. No significant differences were
found between Missouri-20, Dawn, or Mackinaw, with each averaging be-
tween 88-99 florets. Flowering and attractiveness to honey bees was sig-
nificantly correlated, with Empire plots having the greatest foraging activity
and Viking plots the least. Missouri-20, Dawn, and Mackinaw were equally
attractive, but were not foraged as heavily as Empire.

The mayfly fauna (Ephemeroptera) of two Connecticut streams. P. J.
Dodds, Univ. of Connecticut, Storrs, CT 06268

The life histories and ecological relationships of mayflies inhabiting two
Connecticut streams were studied. Forty-eight species were collected from
each stream. Ten were selected for life history analysis utilizing head length
frequency classes and arbitrary age groups based on wing pad development.
This was correlated with adult flight periods to determine temporal parti-
tioning of the habitat. Two basic life history patterns were found. In the
first, nymphs appeared in late summer and autumn and grew throughout the
winter, emerging in the spring. Nymphs and adults of the second type were

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both present in the summer with eggs diapausing for most of the year. Most
species showed a seasonal succession with narrow temporal overlap, al-
though species of the genus Ephemerella exhibited more significant overlap.
The mayfly community as a whole illustrated seasonal changes with greatest
differences occurring between the two rivers in early summer and fall. Num-
bers increased in early spring and decreased thereafter until early summer,
when hatching of summer species induced another increase; numbers de-
clined again through the summer months. Species diversity also decreased
from spring to fall in both rivers. Such changes in the mayfly fauna were
associated with life history phenomena, including delayed hatching, matu-
ration, and adult emergence.

Impact of predators and parasites on the survival of Rhinocyllus conicus
Froelich (Coleoptera: Curculionidae). P. F. Dowd and L. T. Kok, Dept, of
Entomol., VPI and State Univ., Blacksburg, VA 24061

Several predators and parasites frequently observed in the vicinity of
thistle plants infested by the thistle head weevil, Rhinocyllus conicus Froe-
lich, were studied during 1978 to update their impact on R. conicus. Poten-
tial predators were collected from the flower heads of musk thistle ( Carduus
nutans) in Pulaski Co., VA during the oviposition period of R. conicus in
the spring. Exposure tests using insects in the families: Cleridae, Canthar-
idae, Lampyridae, Elateridae, Nabidae, and Formicidae indicated no pre-
dation on the eggs and adults of R. conicus. However, similar studies with
spiders in the families Salticidae and Thomisidae showed predation on the
adult weevils and third instars. Salticid spiders appeared to prefer larvae
(6.5/wk), while the thomisid spiders were more important predators of adult
weevils (3/wk). These spiders, however, were not abundant and thus were
not important mortality factors. Parasitism was studied by collecting 50
terminal heads of plumeless thistle ( Carduus acanthoides) from each of 2
selected sites, and 100 terminal heads and peduncles of musk thistle from
each of 3 sites during early summer. Twenty percent of the heads were
dissected soon after collection and those remaining were held for parasite
emergence. Four species of parasites were recovered, 2 of which were pre-
viously reported on R. conicus. Rates of parasitization of R. conicus in the
2 plumeless thistle sites were 0 and 14.8 ± 8.7%, as compared to 5.0 ±
1.8%, 1.2 ± 1.3% and 0.27 ± 0.55% in the 3 musk thistle sites. The rate of
parasitization generally was not sufficiently large to be a major mortality
factor of R. conicus.

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45

Ovipositional preference of the grape colaspis, Colaspis brunnea (Fab.)
(Coleoptera: Chrysomelidae) in North Carolina. A. T. Eaton, R. L. Rabb
and J. W. Van Duyn, Univ. of New Hampshire, Durham, NH 03824, and
North Carolina State Univ., Raleigh, NC 27650

The effects of four different soils on ovipositional preference of the grape
colaspis were investigated in field cage and laboratory tests. A 1976 field
cage test (RCB design) utilized soybeans growing in caged areas measuring
2.74 x 1.83 m. A thin layer of test soil occupied the top 3 cm of the soil
profile; the caged areas were divided into quadrants for the four soils. Adults
of Colaspis brunnea were swept from surrounding soybean fields and re-
leased inside the cages. After allowing six weeks for oviposition and growth
of the larvae, soil samples were taken inside the cages. The number of larvae
recovered from the soil demonstrated a preference for clay loam followed
by loam, muck, and sand. The former were highly creviced soils; the latter
had few surface crevices. The following year, repetition of the test in the
field and laboratory gave similar results. The lab test utilized cylindrical
plastic containers, 15 cm in diameter x 4 cm deep, into which 10 male and
10 female Colaspis were placed. After several days, eggs were carefully
recovered from the soil, and the number of eggs recovered in each quadrant
(soil type) were compared. Additional testing with the same containers dem-
onstrated a preference for creviced soil over smooth soil. Further laboratory
observations suggested that adults preferred to oviposit in areas of low light
intensity.

Presocial behavior in Cerceris watlingensis Elliott & Salbert (Hymenoptera:
Sphecidae). N. B. Elliott and T. Shlotzhauer, Hartwick College, Oneonta,
NY 13820

Earlier studies had shown that females of C. watlingensis shared nests.
Marked females were observed on San Salvador Island, Bahamas, during
November and December, 1978 to determine the extent of nest sharing and
switching. Newly emerging females remained in the parental nest for several
days. In most cases several females provisioned these nests during this
period. After a few days the young females switched to new nests. Nest
switching was preceded by a day’s searching activity during which females
flew restlessly about, often biting the sand’s surface. In some cases the new
nests may have been freshly dug, but we observed females taking over
already established nests. Females always chose new nests at least one
meter away from their original nests, although entrances in this species were
often only a centimeter apart.

Activities of the individuals in shared nests were compared, and one was
always a more effective provisioner. The major provisioner always left the

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nest first in the mornings, spent less time in the nest between provisioning
trips, and provided more prey per day. The other females spent more time
within the nest, often with the head in the entry. This behavior served to
guard the nest against enemies, especially ants, the major predators of the
species. Thus there is the beginning of division of labor in this species.

Effects of dietary nitrogen on 14C-glycine and 14C-formate incorporation into
body urates of Blattella germanica (Dictyoptera: Blattellidae). J. A. En-
gebretson and D. E. Mullins, VPI and State Univ., Blacksburg, VA 24061

The relationship of dietary nitrogen to urate metabolism and the incor-
poration of two radiolabeled precursors has been examined in Blattella ger-
manica. Male and female cockroaches maintained on diets ranging from 0
to 42% protein and injected with either 14C-glycine or 14C-formate were
examined for retention of total radioactivity in whole insects and their urate
fractions.

Urate synthesis/storage increased in cockroaches maintained on the
higher protein diets, whereas urate levels decreased in those insects on low
nitrogen diets. Greater radiolabel retention was found in insects maintained
on the high protein diet when injected with either 14C-glycine or 14C-formate;
however, cockroaches injected with 14C-formate retained almost twice as
much radioactivity as those injected with 14C-glycine.

Incorporation of 14C-glycine into the urate pool varied significantly among
insects maintained on different dietary regimes. Insects on higher dietary
nitrogen levels incorporated more radiolabel into urates. The pattern of re-
formate incorporation into body urates differed from that observed for 14C-
glycine. More radioactivity was found in the urates from insects maintained
on the intermediate nitrogen diets. Insects fed a 42% protein diet incorpo-
rated significantly less 14C-formate into their body urates. These levels were
comparable to those obtained from insects maintained on 0 and 5% protein
diets.

Niche relations of stinkbugs ( Podisus spp.) attacking eastern tent caterpil-
lars. E. W. Evans, Cornell Univ., Ithaca, NY 14853

The importance of interspecific competition in shaping the life history char-
acteristics of insects is presently unclear. The present study addresses the
issue by considering patterns of prey exploitation of three species of pred-
atory stinkbugs, Podisus maculiventris, P. placidus, and P. modestus (He-
miptera: Pentatomidae), which were found together, attacking eastern tent
caterpillar larvae (Lasiocampidae: Malacosoma americanum). Tents of M.
americanum and more than 500 associated adults of the three predator
species were censused in 1977-79 near Ithaca, N.Y. While only Podisus

VOLUME LXXXVIII, NUMBER 1

47

maculiventris adults were found about the tents during the first half of the
larval period of M. americanum, adults of all three predators were found
together about the tents thereafter. During this period of overlap, the three
predators did not differ in the sizes of prey they consumed, despite differ-
ences in the sizes of these predators themselves. The three predators did
differ, however, in their locations: P. placidus was most often found inside
tents, P. maculiventris on the surface of tents or within 30 cm of the tents,
and P. modestus more than 30 cm from the tents. These results suggest that
the three species have diverged in where they hunt for prey. As individual
predators readily attack prey already subdued by other individuals, this
divergence in site of hunting may reduce interference in consumption of
prey, and this may reflect the importance of interspecific competition in
determining how these predators exploit tent caterpillars.

Effect of temperature and light on aggregation behavior of Aedes stimulans
larvae (Diptera: Culicidae) in a woodland pool. G. P. Fernald and J. F.
Burger, Dept, of Entomol., Univ. of New Hampshire, Durham, NH 03824

Behavior of Aedes stimulans larvae in southeastern N.H. was studied
relative to daily changes in water temperature. Observations were made on
a 41 x 2.5 m woodland pool divided into 14 sections. Temperature readings
and larval counts were taken in each section from initiation of hatching at
2.7°C (March 6, 1979) until adult emergence was complete (May 12, 1979).
Maximum vertical and horizontal temperature gradients occurred on bright,
sunny days. Maximum surface temperatures occurred over 5 cm deep mar-
gins in the shallower sections. Aggregating behavior of A. stimulans was a
complex response to temperature and incident solar radiation. These were
major factors initiating and directing horizontal movement of larvae. Diurnal
warming and nocturnal cooling induced the daily formation and dispersal,
respectively, of larval aggregations. Vertical and horizontal distribution was
more uniform when temperature gradients were low. Larvae were mostly
inactive below 5°C except for vertical movement associated with feeding
and respiration. Horizontal movement in response to light was greatest at
7-10°C. Above 10°C horizontal movement was a thermotactic response to
increasing temperature. Larvae did not cross thermal barriers more than
2°C below temperatures in the aggregation site. Larvae tended to move
toward areas of higher temperature until water temperature in shallow sec-
tions reached the optimum (18-19°C). Thereafter, larvae tended to avoid
areas of higher temperature (>20°C) and seek areas of lower temperature
in deeper sections of the pool.

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A definitive method for determining pear seedling resistance against pear
psylla, Psylla pyricola Foerster (Hemiptera: Psyllidae). B. J. Fiori1 and R.
C. Lamb2, SEA, USDA, Geneva, NY1. New York State Agric. Expt. Stn.,
Geneva, NY 144562

Reduction of egg deposition and nymphal mortality have been shown to
be responsible for resistance against the pear psylla in Pyrus ussuriensis
and crosses of P. ussuriensis with P. communis. Exposure of candidate
seedlings to unconfined adults and counting the number of nymphs per seed-
ling after appearance of 5th instars (25-30 days) has been suggested and
employed to determine resistance. The method does not permit seedling
replication in a single test, does not exclude preference and permits some
seedlings to escape infestation and remain unclassified. To avoid these se-
rious disadvantages the following method was devised using known resistant
and susceptible pear stock grafted to potted rootstock. Dialysis tubing was
used to confine 5S and 5 9 psylla adults on two leaves of each plant for a
10-day period; tubing and adults were removed, eggs were counted and
reduced to 15 per leaf by rubbing. After an additional 15-20 days (appear-
ance of 5th instars) nymphal counts were taken. In 4 tests using 2 plants
each of 1 resistant and 2 susceptible stocks, average egg deposition and
number of nymphs was reduced 57-83% and 87-100% respectively on re-
sistant stock. Results indicate the number of eggs deposited within 10 days
can be used to determine resistance of candidate seedlings, thus reducing
testing time from 25-30 to 10 days, and that known resistant and susceptible
standards should be included in each test. The method provides for at least
2 replications per seedling, excludes preference and does not allow escapes.

PTTH and ecdysone release in last instar larvae of the European corn borer,
Ostrinia nubilalis (Hiibner) (Lepidoptera: Pyralidae). D. B. Gelman and D.
K. Hayes, Insect Reproduction and Livestock Insects Laboratories, USDA,
Beltsville, MD 20705

European corn borer larvae were reared under a photoperiodic regimen
of 16 hours of light and 8 hours of dark and a temperature of 30°C, conditions
which promote pupation. Based on a comparison between % pupation in
controls (unligatured larvae) and % pupation or formation of intermediates
in ligatured larvae, it was concluded that PTTH release, under the above
conditions, begins approximately 24 hours before the formation of the phar-
ate pupa, and that ecdysone release by the prothoracic glands occurs at
least 6-9 hours after PTTH release. Both PTTH and ecdysone release as
well as the formation of the pharate pupa appear to be rhythmic in that
PTTH release and the formation of the pharate pupa are associated with the
onset of darkness, while ecdysone release is associated with the onset of

VOLUME LXXXVIII, NUMBER 1

49

dawn. Ligatured larvae which did not show signs of pupation by 96 hours
post ligation never did show such signs, although most remained alive for
30-40 days. This indicated that effective concentrations of PTTH are not
released before the 24 hours preceding pharate pupal formation. Since many
headless pupae did show adult development, including completely formed
wings and legs with scales, it can be concluded that neurohormones and
hormones needed for this development and produced by structures in the
head have been released before the formation of the pharate pupa.

Effects of decapitation on ecdysone and vitellogenin titers and egg matu-
ration in Aedes aegypti (Diptera: Culicidae). J. T. Greenplate, Cornell
Univ., Ithaca, NY 14853

Egg development in Aedes aegypti is governed by an intricate control
system involving several hormones. Early studies were limited to measuring
oocyte growth alone. However, recent discoveries have allowed the study
of a sequence of intervening steps and products. Early decapitation exper-
iments associated egg maturation with the release of a factor from the head
at 4-8 hr after blood-feeding. Later in vitro research suggested that a brain
hormone (released after blood-feeding) prompts the ovary to secrete ecdy-
sone which stimulates the fat body to synthesize vitellogenin, the yolk pro-
tein precursor. In vivo support for this scheme has been lacking, but recent
studies by this author add such support. A. aegypti females were decapi-
tated at various times after blood-feeding. At 20 and 48 hr post blood meal
(PBM), the animals were dissected and divided into two groups: females
with maturing oocytes (yolk length > 100 ptm) and females with arrested
oocytes (yolk length ^ 50 pm). Females maturing oocytes at 20 and 48 hr
contained much higher titers of ecdysone and vitellogenin respectively (mea-
sured via radioimmunoassay and rocket-immunoelectrophoresis respective-
ly) than did females in oogenic arrest. These experiments agree with early
work in which egg maturation in A. aegypti was linked to the initial release
of a brain hormone. In addition, the association of brain hormone release
and egg maturation with the production of large amounts of ecdysone and
vitellogenin lends in vivo support to the latest in vitro studies.

Control of neotenic development in a primitive termite (Isoptera: Hodoter-
mitidae). S. Greenberg and A. M. Stuart, Dept, of Zool., Univ. of Mass.,
Amherst, MA 01003

Functional reproductives in colonies of the lower termites have been
shown to inhibit the precocious sexual development (neoteny) of competent
larvae. The mechanism of this inhibition has not been conclusively dem-

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onstrated. It has been postulated that this inhibitory influence is exerted
through sex-specific non-volatile pheromones that are given off together
with the feces of functional reproductives: larvae obtain the postulated in-
hibitory pheromones by proctodeal feeding. However, no pheromones
which inhibit neotenic development in larvae have ever been isolated from
any termite species. The hypothesis that the inhibitory pheromones are
contained in feces from functional reproductives was tested using the west
coast damp-wood termite Zootermopsis angusticollis (Hagen) (Isoptera:
Hodotermitidae). As expected neotenic reproductive development was in-
hibited in colonies of larvae that contained a pair of functional reproduc-
tives. When filter paper impregnated with feces and hind gut contents from
male and female reproductives and larvae were fed to groups of larvae that
did not contain a pair of functional reproductives, neotenic development
was actually stimulated compared to control colonies fed filter paper only.
These results suggest that in the termite Z. angusticollis, the postulated
inhibitory pheromones are not contained in feces from functional reproduc-
tives nor circulated in colonies by proctodeal feeding. Instead, proctodeal
feeding may have a primarily nutritive role, in this termite species, as well
as being essential for the transfer of symbiotic protozoa.

Resmethrin sprays for relief from deer flies, Chrysops atlanticus (Diptera:
Tabanidae). E. J. Hansens, Rutgers Univ., New Brunswick, NJ 08903

Resmethrin appeared promising as a safe insecticide which would reduce
deer fly annoyance to harvesting crews and other agricultural workers in
fields near large salt marsh breeding areas. During two seasons resmethrin
EC was applied at 0.025, 0.05, and 0.1 lb/A using conventional air-blast
sprayers. Spray was directed from the edge of the fields into the vegetative
barrier between these fields and the salt marsh. The sprays were applied
between 7:00 and 8:45 a.m. when wind was less than 5 mph. Observations
were made before treatment and approx. 1, 4, 8, 24, and 48 hr after treatment
both in treated and nearby untreated areas. Effectiveness was measured by
averaging 5 counts of flies taken in 10 figure-8 sweeps with an insect net
around the observor’s head. Pre-treatment counts varied from 15 to more
than 200 deer flies/10 sweeps. Dosages of 0.1 and 0.05 lb/A reduced flies
visiting a human host by more than 95% at 1 hr post-treatment, by 50-95%
after 8 hr and 0 to 85% after 24 hr. Sprays with 0.025 lb/A were nearly as
good. For New Jersey only label approval has been obtained for sprays for
deer fly control using resmethrin EC applied at 0.025 to 0.05 lb ai/A in
agricultural and recreational areas. Such applications can be expected to
give greatly reduced deer fly annoyance for 1-2 days.

VOLUME LXXXVIII, NUMBER

51

A field study of Aedes sollicitans (Walker) (DipteraiCulicidae) as a vector
of filarial worms. W. M. Johnson and W. J. Crans, Mosquito Research and
Control, Rutgers Univ., New Brunswick, NJ 08903

Laboratory studies have shown Aedes sollicitans to be a susceptible host
for the larvae of Dirofilaria immitis (Leidy). This mosquito has long been
suspected of being a major vector of dog heartworm along the New Jersey
coast. To date there has been no direct field evidence to support these
findings and infective stage larvae have never been recovered from wild
populations. Ae. sollicitans were collected from C02-baited CDC light traps
during the summer of 1978 to determine if filarial worms could be recovered
at a site where dog heartworm was known to be endemic. The traps were
operated once weekly from June to October at 3 sites along a 3 km transect
from a breeding marsh into the upland. Ae. sollicitans were anesthetized,
sorted and pooled into groups of 150, and infective stage filarial worms were
extracted by a modified Baermann technique. Approximately 100 mosqui-
toes from each collection were individually dissected to detect developing
stages of the parasite. Dissections revealed developing filariae as early as
June, but infective stage larvae were not recovered by extraction until late
August and September. Results showed that natural populations of Ae.
sollicitans were capable of harboring filarial parasites but suggest that there
may be a seasonal distribution in the transmission to vertebrate hosts.

A systematic analysis of the genus Actia Robineau-Desvoidy in North
America with descriptions of two new species (Diptera: Tachinidae). M. A.
Kamran, Dowling College, Oakdale, NY 11769

The genus Actia R.-D. consists of small flies which are parasitic on mi-
crolepidopterous larvae. They are an abundant group and have a worldwide
distribution comprising ca 60 species.

The genus Actia was erected by Robineau-Desvoidy in 1830. Over the
years its generic limits were invariably given a broad interpretation with the
result that hundreds of specimens in various museum collections, loosely
ascribed to Actia R.-D., in fact did not belong there. Recent authorities
have looked at this assemblage much more critically and sharply defined the
generic limits of the many taxa included therein.

This paper presents the important diagnostic characteristics of the tribe
Siphonini, and all points of discussion relevant to generic limits in the sub-
tribe Siphonina. A key is provided for the separation of all the known North
American genera, viz., Actia R.-D., Aphantorhapha Towns., Ceranthia R.-
D., Ceromya R.-D., Chaetostigmoptera Towns., Peribaea R.-D., Pseu-
dosiphona Towns., Siphona Meig., etc.

The genus Actia R.-D. is reviewed. Among the recorded North American

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NEW YORK ENTOMOLOGICAL SOCIETY

species of Actia, the following 4 are recognized as valid: A. autumnalis
(Towns.), A. diffidens Curr., A. interrupta Curr., and A. rufescens
(Greene). Two new species, A. labellata and A. pauciseta, are described.
A key for the separation of the North American species is provided.

Removal method estimation of Blcittella germanica (L.) (Dictyoptera: Blat-
tellidae) populations. C. B. Keil, VPI and State Univ., Blacksburg, VA
24061

The estimation of cockroach population numbers is a continuing problem.
Mark and recapture techniques have met with variable success. In this
study, a removal method estimation was used in preparation for a sterile-
male experiment on two Navy ships. The objectives were (1) to document
reduction of population numbers prior to releases by estimating pre- and
post-removal population size and (2) to make inferences concerning popu-
lation age structures.

In cooperation with Navy entomologists, collections were made on 2
ships, one 62.5 m and the other 159.2 m in length. A synergized aerosol
pyrethrin spray was used to flush cockroaches from harborages. Known or
suspected harborages were then sprayed with 1% baygon in oil. All cock-
roaches appearing within 15 minutes of cessation of spraying were collected
with a vacuum cleaner.

Two captures brought the population of the smaller ship to manageable
levels. Capture probability was .6851 ± .0063. Pre- and post-removal num-
bers were calculated with ca. 9% coefficient of variation. The large ship
required 4 collections. Capture probability was lower (.5077 ± .0021), as
was the coefficient of variation (ca. 5%). All captures contained a prepon-
derance of adults. Treatment caused a shift in the sex ratio toward a majority
of adult females. Nymphal populations showed a 1:1 sex ratio. Male nymphs
appeared to have 5 instars while females had six.

Assessment of insect damage to vegetation by remote sensing. V. Klemas,
Univ. of Delaware, Newark, DE 19711

Insects cannot be detected directly from satellites or aircraft. However,
changes in the spectral reflectance of plants can be used to assess insect
damage to vegetation over large wooded and agricultural areas. Examples
of successful remote sensing of insect damage include gypsy moth (Por-
thetria dispar (L.)) defoliation of mixed woodlands, citrus blackfly ( Aleuro -
canthus woglumi ) infestations in citrus groves, bark-beetle impacted coni-
fers, and various crop diseases. Remote sensing techniques can significantly
enhance the capability of ground-survey teams to assess insect damage be-
cause remote sensors map large areas at relatively low cost; use infrared

VOLUME LXXXVIII, NUMBER 1

53

wavelengths which penetrate deeply into the leaf tissue to detect early loss
of plant vigor; and employ multispectral analysis techniques which can de-
tect smaller spectral reflectance changes than the human eye. While a lim-
ited amount of ‘‘ground truth” is still required, remote sensing techniques
are being adopted by many groups and agencies as a cost-effective tool for
assessing insect and disease damage to vegetation.

Loss assessments and decision-making for the tobacco bud worm, Heliothis
virescens (F.) (LepidopterarNoctuidae), on Maryland tobacco. D. M. Ko-
lodny-Hirsch and F. P. Harrison, Dept, of Entomol., Univ. of Maryland,
College Park, MD 20742

Despite the economic importance of tobacco, very little information con-
cerning tobacco injury by the tobacco bud worm is available. Therefore,
during the growing season of 1977 field studies were conducted to investigate
this pest on air-cured tobacco in southern Maryland. Specific objectives of
our research were (1) to determine the damage-density relationships result-
ing from artificial infestations of the budworm at several growth stages of
tobacco (2) to determine the theoretical economic-injury levels (EILs) of
the budworm throughout several growth stages of tobacco. Percent yield
and value reduction of tobacco was correlated with larval density at several
growth stages of tobacco by multiple and linear regression procedures. Re-
sults of this study show that sensitivity to defoliation varies markedly with
tobacco growth. Throughout most growth stages plants were able to com-
pensate for some degree of foliar loss. Compensation was highest during
growth periods showing rapid meristematic activity. Sensitivity was most
evident during root formation and production of floral parts, although mark-
edly less during the latter. Based on the above data, EILs for the budworm
were determined by integrating several economic parameters with value-
density relationships. EILs indicated that throughout most developmental
stages of Md. 609 tobacco, large populations of bud worms can be tolerated
without sufficient damage to warrant suppression. Average EILs of 25%
approach optimal benefit-cost relations.

Effect of habitat characteristics on the succession of outlet-breeding black
flies (DipteraiSimuliidae) in New Hampshire. D. J. Lake and J. F. Burger,
Dept, of Entomol., Univ. of New Hampshire, Durham, NH 03824

Hatching patterns, adult emergence sequence and temporal succession of
outlet-breeding black flies, including habitat selection in sympatric sibling
species of Simulium venustum and S. verecundum, were studied in N.H.
Larval populations were sampled weekly at 5 pond outlet sites in a 10 km

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radius around Waterville Valley. Habitat characteristics were determined
by measuring the hydrological parameters, including pH, conductivity, dis-
solved gases, temperature, principal ions, benthic substrates and current
velocities. Seventeen species were collected at study sites in 1978-79, 6
were isomorphic sibling species of S. v enustum/v erecundum . S. venustum
sibling species begin hatching from late March to mid-May. The CC sibling
hatches first, followed by A/C, AC (gB) and CC2. All siblings appear to be
univoltine except CC, which persists at some sites until late July. Both
verecundum (ACD and A/C) species hatch from mid-May to mid- June, de-
pending on water temperature at specific sites. Hatching of other outlet-
breeding species after mid-May occurred simultaneously at several sites and
was strongly correlated with water temperature. Adult emergence closely
paralleled hatching sequence in all species. Succession patterns were strong-
ly dependent on rate of increase in water temperature. Differences in hy-
drological parameters between sites were too small to explain differences
in species occurrence at the study sites.

Computer-assisted instruction for pest management: ORCHARD and ALF-
WEEV. P. A. Logan, Univ. of Rhode Island, Kingston, RI 02881

Two interactive computer programs were written to help students learn
to manage pests in dynamic agro-ecosystems. ORCHARD, written at Mich-
igan State Univ. in 1977, considers principles of managing apple mites. Its
3 levels of difficulty require a student to 1) become familiar with simple
predator-prey population dynamics (Lotka-Voltera), 2) to be aware of how
changes in either the predator or the pest populations can affect the overall
population stability, and 3) to make a profit in an orchard through an inte-
grated program using predatory mites and contemporary miticides. ALF-
WEEV, originating at Michigan State and completed at the Univ. of Rhode
Island in 1978, is a computer simulation of an alfalfa field and some of the
insects found there, including alfalfa weevil (primary pest), aphids (second-
ary pest), weevil larval parasitoids and honey bees (non-target beneficials).
Students manage pests and optimize yield (both quantity and quality of hay)
by making critical day-to-day decisions on cutting or spraying strategy. Re-
alistic weather forecasts, dynamic plant growth processes (changes in
height, biomass, protein, fiber, and digestible nutrients), legal restrictions,
etc. add to the game environment. Both programs are fully tested and are
standardized to be easily adaptable to any computer. Both are supplemented
with student guides and programmer manuals. ALFWEEV also has a sep-
arate teacher guide.

VOLUME LXXXVIII, NUMBER 1

55

Predicting the impact of Colorado potato beetle ( Leptinotarsa decemlinea-
ta Say) on yield. P. A. Logan and R. A. Casagrande, Univ. of Rhode Island,
Kingston, RI 02881

The economic significance of CPB density as it changes in time has never
been accurately quantified. We have developed an expedient dynamic pre-
dictor of CPB impact on early maturing potato varieties to serve until a
more accurate multidimensional simulation can be built. To do this, we
established a gradient of densities as follows: All adults in each of several
4 x 4 m plots were collected twice weekly and redistributed at 3, 1, 0.5,
0.1, and 0.05 times lx lot density, taking advantage of the CPB’s relatively
sedentary behavior. Twice weekly, for 8-10 weeks, 30 counts per plot were
made of egg masses per stem and larvae per stem. Of several alternatives,
3rd order exponential models (over degree-day base 10, adjusted for crop
emergence date) fit these count data best. Comparison of plot yield with
area under the incidence curve provided a useful, linear yield estimator.
Finally, we developed a generalized incidence curve algorithm, capable of
density-dependent shaping adjustment, suitable for projecting a first gen-
eration larval incidence curve (and subsequent yield loss estimate) from 1
or more early season larval samples.

Treatment of a nuclear polyhedrosis virus of Autographa californica Speyer
with 5-Bromodeoxyuridine and determination of changes in virulence in four
lepidopteran hosts. J. T. McClintock and C. F. Reichelderfer, Univ. of
Maryland, College Park, MD 20742

The use of chemical mutagens to produce mutations in vertebrate viruses
has been reported by several investigators. Chemical mutagenesis might
provide a mechanism to increase virulence in insect pathogens, and ulti-
mately enhance their potential for utilization in an integrated pest manage-
ment program. The purpose of this study was to treat an insect virus that
displays a relatively high degree of virulence for one insect and assess
changes in virulence for several alternate hosts. Increased virulence with
insect viruses has been achieved by serial passage and one published report
exists concerning treatment with chemical mutagens which suggests that
virulence might be under polygenic control.

This presentation concerns the effect of treatment of a nuclear polyhe-
drosis virus of Autographa californica, the alfalfa looper, with the chemical
mutagen 5-Bromodeoxyuridine. Treatment was initially carried out in Tri-
choplusia ni, the cabbage looper, and the effect of treatment assessed in
this host along with subsequent effects on virulence in Heliothis virescens,
the tobacco bud worm; Heliothis zea, the corn earworm; and Spodoptera
frugiperda, the fall army worm.

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The consequences of using alternate hosts as a starting point for treatment
with mutagens was found to be significantly important, as well as the overall
effects of treatment on the LC50 and LT50.

Consequences of host phenology for exotic scale insects. M. S. McClure,
Connecticut Agric. Exp. Stn., New Haven, CT 06504

Two armored scales, Fiorinia externa Ferris and Tsugaspidiotus tsugae
(Marlatt) (Homoptera: Diaspididae), native to Japan coexist on Eastern
hemlock and on numerous other native and exotic conifers in Connecticut.
Studies were conducted from 1976-1979 to examine the phenological rela-
tionships between these exotic scales and their hosts and related effects on
scale success. Differences in the phenology of 14 host species and related
differences in concentrations of nitrogen and water in the newest foliage in
June during peak colonization by nymphs resulted in differential scale suc-
cess. Nymphs which colonized hosts whose foliage was relatively immature
and contained high concentrations of nitrogen and water suffered less mor-
tality, developed faster, and ultimately laid more eggs than nymphs which
colonized hosts with relatively mature foliage containing low concentrations
of nitrogen and water. The phenological relationships between Eastern hem-
lock and its exotic scales favored the success of F. externa whose nymphs
colonized 2 weeks earlier than nymphs of T. tsugae. Greenhouse experi-
ments demonstrated that earlier colonization by F. externa had adverse
effects on T. tsugae success by reducing the amount of foliar nitrogen avail-
able to T. tsugae nymphs and by causing a greater portion of them to
colonize older, low nitrogen foliage where fewer survived. At 18 of 20 sites
where these scales coexisted on hemlock during 1976, F. externa demon-
strated superior competitive ability during three years by excluding T. tsu-
gae (14 sites) or by significantly reducing its population densities. The com-
petitive superiority of F. externa is likely due to its greater compatibility
with hemlock phenology.

Effect of Agromyza frontella Randani (Diptera: Agromyzidae) on alfalfa
quality and yield. G. B. MacCollom, G. Baumann and J. G. Welch, Vt.
Agric. Exp. Stn., Univ. of Vt., Burlington, VT 05405

The alfalfa blotch leafminer, Agromyza frontella, a newly introduced al-
falfa insect, was first reported in the northeast U.S. during 1968. Conflicting
views on economic damage to alfalfa have in some states resulted in chem-
ical control recommendations. Five years of field experimentation on the
control of the alfalfa blotch leafminer, and its subsequent effect on yield
and quality have shown no benefit from insecticidal treatment. Quality as
measured by amounts of protein and percentage digestibility showed no

VOLUME LXXXVIII, NUMBER 1

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significant differences between carbofuran and Pydrin treated plots having
less than 1 mine/stem and control plots having over 30 mines/stem. No
differences in yield as measured by dry weight were detected between the
treated and untreated plots. Hand-separated leaflets, into blotched and un-
blotched samples from 1st cut alfalfa, showed no significant difference in
crude protein. The average weight of a blotched leaflet was 0.9 mg less than
an unblotched, indicating that one might expect yield differences, but this
was not reflected in 5 years of evaluation. It is suggested that the alfalfa
plant may compensate for the mining activity by putting out more leaflets,
but this aspect has not yet been investigated. On the basis of these studies,
the alfalfa blotch leafminer is not considered a serious alfalfa pest in Ver-
mont, and as a result growers are advised not to initiate insecticidal treat-
ment.

Effect of pirimicarb on natural enemies of Myzus per sic ae (Sulzer) (Homop-
tera: Aphididae). T. P. Mack and Z. Smilowitz, Pesticide Res. Lab.,
Dept, of Entomol., Pennsylvania State Univ., University Park, PA 16802

The effects of pirimicarb on the predaceous natural enemies of M. per-
sicae, the green peach aphid (GPA) were determined in a potato ( Solarium
tuberosum L. var Katahdin) field as a step in the development of an inte-
grated pest management system for the GPA on tablestock potatoes. These
effects were determined by visually counting predators occurring on 24 ran-
domly selected potato stems on several prespray and 4 postspray (20, 41,
116 and 164 hr) sample dates. The plots were treated with encapsulated
methyl parathion 18 days prior to the test to facilitate GPA population
growth and thus attract natural enemies. Pirimicarb was applied at 4 oz
active ingredient/acre.

Coccinellid larval populations in treated and untreated plots were similar,
with population peaks occurring 20, 41 and 116 hr postspray. Coccinellid
pupal populations in both treatments were similar until 41 hr postspray,
when the untreated population exponentially increased at a much fast-
er rate than the treated population. Hippodamia convergens Guerin
(Coleoptera:Coccinellidae) adult populations in treated and untreated plots
were markedly different. The H. convergens adult population in the treated
plot decreased exponentially until the end of the study, while the untreated
population increased. Coleomegilla maculata (DeGeer) (Coleoptera:Coc-
cinellidae) adult populations were also higher in untreated than treated
plots. The adult Coccinella transver so guttata richardsoni Brown (Coleop-
teraiCoccinellidae) populations were variable in both treatments, with dif-
ferent peaks.

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Development of adult abdomen in Drosophila melanogaster Meigen (Dip-
teral Drosophilidae). M. M. Madhavan and K. Madhavan, Biol. Dept., Holy
Cross College, Worcester, MA 01610

The cell lineage of the histoblasts during metamorphosis of Drosophila
has been studied by histological methods. The development of adult abdo-
men is characterized by three morphogenetic processes. (1) Mitosis and
consequent increase in the number of cells in the histoblast nests. This
mitotic activity parallels the increase in the titre of ecdysone. (2) Pro-
grammed cell death among the larval epidermal cells and the coordinated
spreading of the histoblasts to their final positions replacing the epidermal
cells. By 24 hr after pupariation, fusion of the anterior and posterior dorsal
nests and that of ventral nest with spiracular anlage is complete. The dorsal
nest fuses with the spiracular anlage by 28 hr and by 36-40 hr after pupar-
iation the histoblasts have completely covered the abdomen. From the pat-
tern of cell death, and spreading of histoblasts it is concluded that the larval
cells provide a cue for the advancing histoblasts to what extent they can
spread in the formation of the definitive segment border. (3) Differentiation
of the imaginal epidermal cells resulting in the secretion of the cuticle and
cuticular outgrowths characteristic of adult abdomen. The anterior dorsal
histoblast nest gives rise to the region of the tergite bearing cuticular out-
growths, the posterior dorsal nest, possibly, to the intersegmental mem-
brane and to the acrotergite of the following segment, the spiracular nest to
the spiracle and the area surrounding it, and the ventral nest to the sternum
and the pleura of the corresponding hemisegment.

Feeding behavior of Tabanidae (Diptera) on livestock. L. A. Magnarelli,
Conn. Agric. Exp. Stn., New Haven, CT 06504

The feeding behavior of Chrysops, Tabanus, and Hybomitra species was
observed to determine preferences in feeding sites on dairy cattle and pat-
terns of blood ingestion. Females of 7 Chrysops species fed more frequently
from the head areas than any other body region, whereas those of 4 Tabanus
and 4 Hybomitra species commonly chose the sides, belly region, or legs.
Tabanus lineola and Tabanus quiquevittatus fed avidly from the legs, and
because of their abundance, these insects were of particular annoyance to
livestock in pastures. Deer flies typically fed to repletion after initial feeding
attempts, while horse flies (which attempted to feed from the sides and legs
of animals) were frequently interrupted by host body movements and, con-
sequently, had to pierce their mouthparts repeatedly into the animal’s skin
before they imbibed blood. It was not uncommon to observe H. lasioph-
thalma and Tabanus atratus females flying from one animal to another after
unsuccessful feeding attempts. Internal examinations of host-seeking horse

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59

flies, captured in pastures by dry-ice baited canopy traps, revealed small
amounts of fresh blood in midguts and only partially developed (Stage II)
ovarian follicles. Capillary precipitin tests sometimes detected mixed blood
from bovines and equines. Feeding success (acquisition of full blood meals)
varies with the insect species, site chosen for blood ingestion, and host
response to biting individuals. Repetitive feeding attempts by tabanids on
different host animals reinforce the importance of these insects as potential
vectors of certain pathogens by mechanical transmission.

New approaches for screening passenger baggage at U.S. international air-
ports. J. H. Mahaney, USDA, APHIS, PPQ, Hyattsville, MD 20782

Deregulation of airlines, increasing number of wide-bodied aircraft and
low fares have caused a crisis at American international airports. Since 1960
air traffic has increased nearly 600%. Federal inspectional forces at airports
have increased by only 100%. Further staffing increases are not likely. In-
adequate airport facilities along with inadequate staffing have caused trav-
elers to sit for over an hr in aircraft parked outside of congested terminals.
Government review agencies believe there are too many Customs inspectors
at airports and that baggage spot checks would suffice. Due to intense pres-
sure from Congress, airlines and traveling public, Customs has devised in-
spectional systems to expedite travelers. Newer systems as Citizen By-Pass
and One Stop require only 90 seconds processing. There is even a goal of
60 seconds. As passenger traffic increases (2 million additional seats will be
available for the European run this summer and there is a projection of 25%
increase of travel) it is felt that restrictions will be made on Customs in-
spections at certain saturated airports. Recommendations may be made for
“spot checks” or for the European honor system called “Red door-green
door” whereby the traveler decides whether or not to go through Customs.
Any such departures from present inspectional systems could be most dan-
gerous and extremely destructive to American agriculture which is valued
annually at nearly $500 billion.

Mate-seeking strategies in male flower flies (Diptera: Syrphidae). C. T.
Maier, Dept, of Entomol., Conn. Agric. Exp. Stn., New Haven, CT
06504

Although male syrphid flies exhibit many different types of mate-seeking
behavior, most search only near flowers and other resources utilized by
females. I observed the mate-seeking behavior of males of more than 40
species in the subfamily Milesiinae and extensively studied that of Mallota
posticata (Fabr.) and Somula decora Macq. in a sandy area in Illinois.
Males of both M. posticata and S. decora utilized two different site-depen-

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dent searching strategies. In the morning, they patrolled flowering plants at
the forest edge and in a nearby field to find feeding females. In the afternoon,
they waited near treeholes, potential egg-laying sites, to intercept females
arriving to oviposit. Both mate-seeking areas were aggressively defended
against conspecific and some nonconspecific intruders. Resident males of
M. posticata won 95% of the encounters with conspecific males entering
their territories near treeholes. Active territorial defense (i.e. flying in re-
sponse to intruders), probably the most energetically costly activity per-
formed by males near treeholes, accounted for only 2.4% and 2.9% of the
total activity budget of M. posticata and S. decora, respectively. The mat-
ing success of males at each resource is probably related to the abundance
of males and females and to the degree of sperm precedence.

A comparison of laboratory-reared and wild type gypsy moth Lymantria
dispar (L.) males (Lepidoptera:Lymantriidae) V. C. Mastro, Gypsy Moth
Methods Development Center, APHIS, PPQ, USDA, Otis AFB, MA 02542

Studies carried out between 1957 and 1973 explored the effects of gamma
irradiation and chemical sterilants on male gypsy moth sterility. These and
other studies indicated that laboratory reared insects were inferior to wild
type males. Recent improvements in laboratory culturing of the gypsy moth
have made possible the mass culturing of insects for sterile male release and
virus production. However, little was known about the “quality” of insects
produced. The experiments described summarize two years of comparison
of wild type and laboratory-reared gypsy moths. Specifically these studies
compared: the periodicities of eclosion dispersal and attraction to phero-
mone sources; the spacial distribution of dispersing males; the ability of
males to locate natural and artificial pheromone sources; and the longevity
of males in the field.

Results of these trials indicate that male gypsy moths reared in the lab-
oratory for seventeen generations are competitive with several wild strains
of males. Laboratory reared males are able to locate virgin females and
(+) disparlure baited traps as well as feral males. In addition periodicities
of the three activities monitored are similar. Furthermore, laboratory reared
males were captured in nearly the same age classes as wild males. Dispersal
patterns of the strains tested were not identical; but differences did not
indicate that laboratory-reared males are at competitive disadvantage.

Seasonal dispersal of pitcher plant mites (Acarina: Anoetidae). T. N. Mather
and E. P. Catts, Univ. of Delaware, Newark, DE 19711

The structure of the symbiotic community of aquatic arthropods associ-
ated with pitcher plants ( Sarracenia purpurea ) was studied from sites in

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61

New Jersey and Delaware. Comparisons were made between these com-
munities, and similarities and differences in species abundance and richness
were noted. Anoetid mites, believed to be Anoetus gibsoni, were most
abundant from New Jersey plants, but were found from Delaware plants as
well. Little more than a species description is found in the literature con-
cerning this mite. A. gibsoni commonly uses a non-feeding hypopal stage
for dispersal from its overwintering site inside old pitcher plant leaves. Mites
overwinter as hypopi, and dispersal of mites begins in the spring after new
leaves are open. Techniques were investigated to determine mechanisms of
mite dispersal to new leaves. Emergence traps placed over individual over-
wintering leaves were serviced twice weekly and freshly emerged insects
were collected and examined for paratenic mites. Newly formed leaves were
marked and collected at two day intervals, and their contents were examined
for mite dispersants. Insect visitors were selectively blocked from entering
newly opened leaves with screens of different mesh size. Parasaggital sec-
tions of leaves glued to glass were used to observe pitcher plant biota in
situ.

Development of overwintered versus summer pupae of the alfalfa blotch
leafminer Agromyza frontella Randani (Diptera: Agromyzidae). W. K. Mel-
lors and R. G. Helgesen, Dept, of Entomol., Cornell University, Ithaca,
NY 14853

The overwintered versus summer pupae of the alfalfa blotch leafminer
(ABL), Agromyza frontella, differed consistently in their temperature-de-
pendent developmental rates. Overwintered pupae developed about 25%
faster than summer pupae. This difference was significant in attempts to
computer simulate the timing of spring emergence of the ABL. The transi-
tion from summer to overwintered developmental rate characteristics in
ABL pupae was analyzed by comparing simulations using each set of rates
to observed adult emergence from field samples incubated at a constant
20°C temperature in the laboratory. In August, emergence was adequately
simulated using the observed pupation dates and summer pupal rates. In
November, emergence was adequately simulated using the overwintered
pupal rates and assuming that the pupae were at the same stage of devel-
opment after completion of a presumed period of diapause development. By
November, the variation in developmental state among pupae in the field
had decreased compared to the time interval during which these individuals
had pupated in the late summer. These developmental differences have con-
sequences in terms of both the ecology and management of the ABL.

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A new Japanese beetle trap containing pheromone and floral lure as syn-
ergistic attractants. F. W. Michelotti and J. W. Seidenberger, J. T. Baker
Chemical Company, Phillipsburg, NJ 08865

A simple, effective all-plastic Japanese beetle trap has been developed
consisting of a set of interlocking vanes and an hourglass-shaped disposable
plastic bag that is attached through four slits and hooks. A controlled-release
Hereon™ strip containing minute quantities of Japanese beetle pheromone
(R,Z-5-(l-decenyl)dihydro-2(3H)-furanone) is affixed to one of the vanes
through an adhesive backing. A floral lure pack, containing a 69%/31% mix-
ture of 2-phenethyl proprionate and eugenol, respectively, is inserted
through slots in the vanes. The trap is then suitably suspended by a hang-
tie and the replaceable plastic bag can be conveniently disposed of when
filled with beetles along with household refuse. Trap performance tests were
carried out in Wooster, OH, during August 1978 in selected residential
areas. The trapping devices caught large numbers of beetles (ca. 20,000)
over a two-day period. Average catch per trap was over 1600 beetles. These
catches were achieved without having to spray with chemicals. It was es-
tablished that use of the natural sex attractant in combination with the floral
lure, markedly increases the number of beetles caught when compared to
captures using the floral lure alone.

Comparative effects of some insect growth regulators on development, mor-
phogenesis, and reproduction of the female rice weevil, Sitophilus oryzae
(L.) (Coleoptera: Curculionidae). J. Mkhize and A. P. Gupta, Dept, of Ento-
mol. and Econ. Zool., Rutgers Univ., New Brunswick, NJ 08903

S. oryzae is an important stored products’ pest throughout the world.
Previous studies of the effects of insect growth regulators (IGRs) have
shown that this insect pest is difficult to control, because it develops within
the kernels. The objective of this study is to compare the effects of R-20458,
Ro 20-3600, MV 678, methroprene and hydroprene on the developmental
stages, morphogenesis, F2 progeny, productivity index, percent reduction
in adult emergence, oviposition, and egg hatchability of S. oryzae. These
effects were studied in insects reared on IGR-treated wheat as well as in
those treated topically. Both methods produced similar effects but at dif-
ferent dosages. All developmental stages are susceptible to the IGRs, the
4th larval and the pupal stages being the most susceptible. The number of
larval-pupal and pupal-adult intermediates increased at higher dosages.
Sclerotization was not uniform in pupal-adult intermediate, and in most the
elytra and the hindwings were malformed. Internally, the ovarioles were
attenuated. Ft progeny decreased at higher dosages. Productivity index was
lowest (2.06) and percent reduction in adult emergence highest (97.94) at a

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topical dosage of 0.3 p\ of 30 ppm of R-2045 and Ro 20-3600, for example.
None of the IGRs significantly affected oviposition and hatchability. A com-
parative account of the dosages used and their effects will be presented.

Effects of soybean plant populations and planting configurations on insect
populations. J. W. Murphy and J. C. Smith, VPI and State Univ., Blacks-
burg, VA 24061

Essex variety soybeans were planted on July 5, 1978 in 12, 24 and 36-inch
row spacings. Within each row spacing were populations of 1, 2 or 4 plants
per square foot corresponding to 43,560, 87,120 and 174,240 plants per acre,
respectively. In linear row foot comparisons, weekly shake cloth observa-
tions revealed that 36-inch row spacings had the highest numbers of the
green cloverworm, Plathypena scabra (F.) and total numbers of phytoph-
agous insects and predaceous insect totals. The 24- and 12-inch row spacings
contained the second highest and least number of insects, respectively.
There were no differences in the numbers of total phytophagous insects
present in plant populations of 1, 2 or 4 plants per square foot. Significantly
higher numbers of green cloverworms were found in plant populations of 4
and 2 plants per square foot compared to those found in populations of 1
plant per square foot. The greatest number of predaceous insects were found
in plant populations of 4 and 2 plants per square foot though the numbers
found in 2 plants per square foot were not significantly different from those
found in populations of 1 plant per square foot. Generally, highest numbers
of phytophagous insect totals, green cloverworms and predaceous insects
were found in those planting configurations with the widest row spacings
and densest plant populations within the rows. These results indicate a
preference of insects in selection of soybean phytohabitat based on plant
spacing.

Biological control of the greenhouse whitefly, Trialeurodes vaporariorum
(Westw.) (Homoptera: Aleyrodidae) by Encarsia formosa Gahan (Hyme-
noptera: Aphelinidae). J. R. Nechols, Dept, of Entomol., Cornell Univ.,
Ithaca, NY 14853

The greenhouse whitefly, Trialeurodes vaporariorum (Westw.) is a seri-
ous pest of many commercially-grown greenhouse crops. Control of this
pest by conventional chemical methods is difficult for a number of reasons.
Biological control by its parasite, Encarsia formosa Gahan, on the other
hand, is a practical method for controlling the whitefly — both on short- as
well as long-term crops. However, to use this natural enemy effectively,
growers and extension agents need to know: (1) what greenhouse temper-
atures to maintain; (2) how many adult parasites to release (in relation to a

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given host density); and (3) when these parasites should be released. From
previous studies, both the optimal temperatures for the parasite (21-27°C),
and the parasiteihost release ratio (about 1:30) have been established. How-
ever, the problem of when to release adult parasites had not been well
established. Therefore, I exposed each nymphal whitefly stage to E. for-
mosa females. The results showed that both 3rd and 4th instar whiteflies
are: (a) the most highly accepted stages by ovipositing Encarsia females,
and (b) the most suitable stages for the parasite’s development and survival.
Thus, timing the release of E. formosa adults to coincide with these host
stages results in efficient mass-rearing and successful colonization of the
parasite. The application of this information to biological control trials on
poinsettia and Clematis has contributed to the successful suppression of
whitefly populations on these crops in commercial greenhouses.

Attack ecology of the sugar maple borer, Glycobius speciosus (Say) (Co-
leoptera: Cerambycidae). W. G. Newton, Dept, of Environ, and For. Biol.,
State Univ. of N.Y., Syracuse, NY 13210

This study was initiated to describe the status of sugar maples at the time
of attack by the sugar maple borer, Glycobius speciosus (Say). A ten acre
plot in a northern hardwood stand near Wanakena, NY, was chosen for
examination. In 1978, all trees with evidence of sugar maple borer damage
were felled. Uninfested stems were taken from the study plot as controls.
Growth ring analysis was performed on three to five cross-sections from
each tree. Parameters of the trees which were assessed for the time of attack
include: growth rate, diameter at base, diameter at breast height, diameter
at attack, height of attack, total height, aspect of attack, and aspect of scar.
Trees were also described as they were at the time of study. Scars were
aged, and this information was used to characterize the pattern of attack
within the plot. Projections were made in order to describe the stand at each
year during the infestation’s history. The parameters of the trees from the
time of attack were compared with the parameters at the time of study to
illustrate the changes that take place after infestation. The most important
factor describing trees prior to attack seems to be growth rate.

Biology of Leptothrips mali (Fitch) (Thysanoptera: Phlaeothripidae), a pred-
ator of orchard mites. M. P. Parrella and R. L. Horsburgh, Shenandoah
Valley Res. Sta., VPI and State Univ., Steeles Tavern, VA 24476

A two-year sampling program in orchards on integrated pest control pro-
grams revealed that Leptothrips mali (Fitch) was one of the most abundant
mite predators during 1977-78. A study of its biology was undertaken as
part of an evaluation of this thrips’ potential for inclusion in an integrated

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mite control program. Laboratory rearing conditions were 23.9° ± 1°C, 14-
hr photoperiod, and 80-100% RH. A total of 1521 eggs were removed from
apple leaves; 92% were on the underside of the leaves along the midvein.
The majority of eggs were laid singly; however, they were occasionally
found in pairs. The duration (x ± S.D. days) of the developmental stages
were: egg — 7.9 ± 1.3 (N == 145), first instar — 6.0 ± 0.7 (N = 34), second
instar — 6.3 ± 0.9 (N = 18), prepupa — 1.3 ± 0.4 (N = 29), pupa I — 1.7 ±
0.5 (N = 19), and pupa II — 4.0 ± 0.5 (N = 27). Total development time (x
± S.D.) from egg-adult was 27.0 ±1.2 days (N = 21). Longevity of adult
females (x ± 5.0 days) was 37.7 ± 25.9 and the total number of eggs ovi-
posited per female (x ± S.D.) was 37.9 ± 30.4. The presence or absence of
males did not affect female longevity or fecundity. However, L. mali is
arrhenotokous; unfertilized eggs become males. No direct feeding on apple
leaves was observed and the thrips survived (x ± S.D.) for 2.8 ± 0.83 (N =
16) days on apple leaves alone. Normal development occurred on a strict
diet of V erbascnm thapsis pollen.

Quantification of Myzus persicae (Sulzer) (HomopterarAphididae) damage
to potato foliage production. F. L. Petitt, Z. Smilowitz and E. S. Nolan,
Pesticide Research Lab., Pennsylvania State Univ., University Park, PA
16802

The dynamic nature of plant growth processes necessitates that when
assessing damage the developmental stage of the plant must be considered
as well as the abundance of the pest. Five aldicarb levels were used in field
plots to foster different green peach aphid, Myzus persicae (Sulzer), pop-
ulation densities. Plant foliage and tubers were harvested weekly during the
growing season and wet and dry weights were recorded. Green peach aphids
were sampled weekly in each aldicarb level. Foliage production models were
developed for each treatment by regression of accumulated foliage dry
weight on degree-days. These models were used to estimate foliage pro-
duction for each treatment. Determination of the impact of pest density on
foliage production for a specific time during the season was effected by
regression of accumulated foliage dry weight on green peach aphid numbers.
Quantification of the damage, resulting in a relative damage index, was
accomplished by regression of the percentage foliage losses on green peach
aphid numbers. The relative damage index, which was expressed as per-
centage foliage loss per green peach aphid, varied as the growing season
progressed. The seasonal damage index was the relative damage index re-
gressed on accumulated degree-days for foliage. Creation of the seasonal
damage index facilitated examination of seasonal changes in the effects of
aphid feeding on foliage dry weight accumulation. Early season infestations

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resulted in greater percentage foliage reduction than did late season infes-
tations in 1977.

Notal setae of first instar nymphs of five Periplaneta species (Dictyoptera:
Blattidae). P. K. Powell, Dept, of Entomol., VPI and State Univ., Blacks-
burg, VA 24061

The cockroach species Periplaneta americana, P. australasiae, P. brun-
nea, P. fuliginosa, and P. japonica are all well known pests of man. How-
ever the morphology of the first instar nymphs of these species has not
previously been well studied. Close observations of the pro-, meso-, and
metanota with a scanning electron microscope at magnifications of 2000 x
and above revealed the presence of at least six types of setae. These can be
divided into distinct groups on the basis of length, shape, and location. The
types include 1) long setae, found on all five species and located primarily
along the notal margins, 2) moderately-long setae, the most common setal
type, found on all five species both along and within notal margins, 3) short
setae, located along and within notal margins on all five species and the
most dominant setal type on P. americana and P. japonica, 4) proclinate
setae, which lie directly on the surface of the cuticle of P. australasiae and
P. fuliginosa, 5) chalazae, located on the underside of the posterior margins
of the nota of all five species, and 6) small setae, located on the postero-
dorsal margins of the nota of P. australasiae and P. fuliginosa.

The chalazae and the proclinate setae are the most unusual. Owing to the
position of the chalazae and their proximity to the next posterior segment,
they probably function as mechanoreceptors. The proclinate setae lie di-
rectly on the surface of the cuticle. Their sockets seem to be sunken into
the cuticle.

Mating behavior of the bee Collet es thoracicus (Hymenoptera: Colletidae).
E. G. Rajotte and R. B. Roberts, Dept, of Entomol. and Econ. Zool., Rut-
gers Univ., New Brunswick, NJ 08903

A nesting aggregation of Colletes thoracicus was found in the sand traps
of a golf course near Trenton, N.J. in May and June, 1979. Protandrous
males fly 3-8 cm above the sand searching for females. Such males will land
and briefly inspect any small dark object. An attractive female (probably
virgin) emerging from the ground is covered by a tumbling mass of 5-15
males. Such males also attempt mating with each other. When one effects
genital union with a female, other males disperse immediately. The copu-
latory pair then flies in tandem to nearby trees or shrubs to complete mating.
One pair remained in copula for approximately 3.5 minutes. The male clasps
the dorsum of the female abdomen. Both face the same direction. Attractive

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females seem to release a sex pheromone that excites males. Males attempt
copulation with freshly killed attractive females. The severed abdomen of
a female seems at least as attractive as the head and thorax. Males were
also attracted to the nozzle of a syringe that expelled air from a container
filled with attractive females. Males attracted to the nozzle attempt to cop-
ulate with each other. In a previous study males have been observed to dig
into the sand a few seconds before an emerging female had breached the
surface. Behavior of these bees suggests the presence of a female sex pher-
omone. The rapidity with which the unsuccessful males lose interest in a
copulating female suggests the presence of a pheromone which negates the
effects of a female sex pheromone.

Interactions among willow herbivores as a determinant of feeding and ovi-
positional patterns of the imported willow leaf beetle, Plagiodera versico-
lora Laich. (ColeopteraiChrysomelidae). M. J. Raupp and R. F. Denno,
Dept, of Entomol., Univ. of Maryland, College Park, MD 20742

Current plant-herbivore theory suggests that young compared with mature
foliage of long-lived plants, such as trees, is preferred by specialist herbi-
vores. Analyses of leaves of the weeping willow, Salix babylonica, indicate
that young leaves are less physically tough and have greater moisture and
nitrogen content compared with mature leaves. Thus, young leaves appear
to be a better resource than mature ones. Comparisons of the feeding be-
haviors of adults and larvae of Plagiodera versicolora indicate that adults
preferentially feed on young leaves but larvae show no significant preference
for either young or old leaves. While older leaves may be less nutritious,
the survivorship of eggs and feeding success of young larvae may be greater
on mature compared to young leaves. This was confirmed when willow
leaves of different ages were surveyed and young leaves were found to be
missing more often or more severely damaged by adult beetles and other
herbivorous insects. Consequently, eggs and/or larvae of P. versicolora are
probably much more likely to be disturbed or destroyed on young compared
to old leaves. If egg or juvenile mortality is greater on young leaves, then
selection should favor oviposition on mature foliage. Laboratory and field
surveys of female egg-laying patterns confirm this prediction. It appears that
the cost of laying eggs on leaves of poor nutrient quality is outweighed by
a gain in juvenile survivorship there.

Insects that deform black cherry. C. O. Rexrode, Northeastern Forest Exp.
Stn., Delaware, OH 43015

Many black cherry, Prunus serotina Ehrh. seedlings and saplings have
forked or crooked main stems. This results in pole stands with a high pro-

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portion of deformed trees, and consequent reduced timber value. Results
of a 2-year study of stem deformity in black cherry in Pennsylvania and
West Virginia revealed that insects, diseases, forest, and browsing by deer
were the major causes of injury to the terminal shoots of seedlings and
saplings. Twenty-seven species of insects from 19 families and 5 orders were
associated with young black cherry trees in six study areas. Of these
species, Archips (Lepidoptera: Tortricida) accounted for up to 36 percent
of the deformed stems and Cecidomyia serotinae O.S. (Diptera: Cecido-
myiidae) accounted for 0 to 43 percent of the deformity. Reducing stem de-
formity in black cherry caused by these two species of insects is a complex
problem because the degree of infestation varies from year to year. Although
insect damage may be found in less than 20 percent of the trees in a plan-
tation in any one year, the damage to shoots is compounded over a period
of years. Also, the insects that cause the deformity have different habits
and life cycles, which makes control difficult.

Sexual behavior in Hawaiian Drosophila (Diptera: Drosophilidae). J. M.
Ringo, Univ. of Maine, Orono, ME 04469

Endemic Hawaiian species of Drosophila are distinguished by their di-
verse and spectacular morphology and mating behavior, and by their ex-
traordinary rates of speciation. Many, perhaps hundreds, of these species
have a dramatic sexual habit: mating at lek or arenas, a sexual habit once
thought to be confined to birds.

The frequencies and sequencing of sexual displays in D. grimshawi, a
“model” Hawaiian species, and in two closely related, morphologically
similar species have been measured and found quite distinct. Individuals of
these species can be separated with little error by discriminant functions.

The frequencies of some male displays in D. grimshawi are density-de-
pendent. Increasing density causes courtship to increase. When females are
present the ratio of contact to noncontact aggression increases linearly with
density (fighting becomes more “intense”). The frequency of communal
displays and aggression are positively correlated only when females are
present, supporting the idea that the tendencies to disperse are balanced in
a lek. These experiments show that whereas males initiate courtship towards
male and female conspecifics at random, they respond to the presence of
females in some of their other sexual behavior (aggression and communal
displays).

The sequential organization of male sexual behavior is nonrandom but
loosely organized. Information theoretical analysis shows differences be-
tween this behavior and the courtship rituals of continental Drosophila and
other insect species.

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Oviposition behaviour and egg distribution of the European apple sawfly
Hoplocampa testudinea Klug. (HymenopteraiTenthredinidae). B. D. Roit-
berg and R. J. Prokopy, Univ. of Mass., Amherst, MA 01003

When presented with apple blossoms in laboratory cages, 15 of 18 (83%)
female sawflies visited previously searched blossoms and 15 of 16 (94%) un-
searched blossoms. Following feeding on pollen and nectar, females crawled
to the receptacle where they displayed little (duration x = 5.7s ± SE1.2,
N = 14) overt searching behaviour. Females pierced the receptacle with
their ovipositors and deposited a single egg (duration of successful ovipo-
sition x = 121.0s ± SE9.9, N = 13). Following oviposition attempts which
exceeded 45s, females fed on exudate from the oviposition puncture. Of
females which visited previously un-searched blossoms, 16 of 19 (84%) at-
tempted oviposition. Of females which visited blossoms which had just re-
ceived an oviposition, 18 of 23 (77%) attempted oviposition. These data
strongly suggest that, unlike another apple pest, Rhagoletis pomonella, fe-
male sawflies do not deposit an oviposition deterring substance. Sawfly
larval infestations were normally distributed among the fruit of three sam-
pled apple trees (N = 208 infestations in 838 fruit examined.) Selection pres-
sure for development of an oviposition deterring substance in this sawfly
may be weak because (1) there is normally a profusion of host blossoms,
(2) larvae are mobile and move between fruits in a cluster, and (3) sawflies
usually (87% of 110 cases examined) deposit an egg in only 1 or 2 of the ca.
4-5 blossoms per cluster, thus leaving the other 2-4 fruits for subsequent
exploitation by larvae.

Anomalous oviposition behavior by the gypsy moth (Lepidoptera: Lyman-
triidae). M. C. Rossiter, State Univ. of New York, Stony Brook, NY 11794

Choice of oviposition site has been correlated with subsequent larval fit-
ness. Previous research on the gypsy moth, Porthetria dispar, indicates that
the female oviposits on the tree which serves as the food source (except
under conditions of very high density). Because the female moth does not
fly, her choice of pupation site becomes her oviposition site.

This study of oviposition preference was done in oak-pitch pine wood-
lands in New York, New Jersey, Pennsylvania and Massachusetts. Study
sites contained gypsy moth populations with densities ranging from low to
high. Oak, the predominant tree in all localities, is considered to be the most
preferred food in the northeastern U.S. Trees were sampled along a transect
and were scored according to tree type and presence or absence of egg
masses. Analysis of the data showed that preference for oviposition on pitch
pine was significant for all localities except New Jersey.

Pitch pine is not a suitable food for early larval development as instars I-

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III cannot or will not eat the needles, yet females prefer pitch pine as an
oviposition site. During the first 48 hr after hatch, the larvae will disperse,
a passive ballooning process dependent upon the wind, if appropriate food
is not encountered or if conditions are crowded. Thus, hatchlings oviposited
on pitch pine must disperse to avoid starvation. This data provides a foun-
dation for further study on the advantages and disadvantages resulting from
preferential use of pitch pine as an oviposition site.

Adaptations of the fall cankerworm Alsophila pometaria (Harris) (Lepidop-
tera: Geometridae) to unapparency of host leaf flush. J. C. Schneider,
Princeton Univ., Princeton, NJ 08540

The fall cankerworm is subject to selection to increase synchrony of time
of egg hatch with host foliation. I have asked the question of how the fall
cankerworm has reacted evolutionarily to variation in time of host foliation
on two spatial scales: 1) within stand^among individual trees (1 m) and 2)
among stand — among species of hosts (100 m). Hatchlings can survive for
only about three days without food. Late hatch by a few days on red maple
resulted in a 50% decline in potential fecundity. Red maple leaves toughen
at an age of 28 ± 2 days. Thus the larval stage must coincide with an
approximately 30-day long window. Individual trees tend to foliate consis-
tently early or late. The average weight of prepupae dropping out of a given
tree is positively correlated with the average weight of the apterous females
climbing the tree the next fall. Thus female aptery results in oviposition on
the same tree on which the female fed as a larva and presumably increases
synchrony of hatch with foliation. The average date of foliation of black
oaks can be six days later than that of red maples. The egg masses in a
stand of black oak hatched three days later on average than those in a stand
of red maple 150 m away. This is due to differences in the hatching char-
acteristics and times of oviposition of the clones specializing on the two
species. Thus differences in hatching time of host races match differences
in host species’ average times of foliation.

Delayed hatching of Japanese wax scale Ceroplastes ceriferus (F.) (Ho-
moptera: Coccidae) in locations strongly affected by marine winds. P. B.
Schultz, Virginia Truck and Ornamentals Res. Stn., Virginia Beach, VA
23455

The Japanese wax scale, Ceroplastes ceriferus is generally distributed
throughout the southeastern states. It attacks a wide variety of ornamental
plants and is very prolific. The scale overwinters as an adult female and
oviposition begins in eastern Virginia in early May. Scale populations on
Ilex cornuta var. bufordii at five sites in the Norfolk metropolitan area were
examined weekly beginning June 12, 1978 and the percentage of hatched

VOLUME LXXXVIII, NUMBER

71

eggs was determined weekly at each site. All weather stations in the study
area provided daily maximum and minimum temperatures which were com-
pared to the hatching rate.

On June 12, 1978 the hatching at the southern exposure of the most urban
location was 80%, while hatching had not begun at locations near Chesa-
peake Bay or the Atlantic Ocean. One week later all sites, except near the
Atlantic Ocean, had hatching percentages of 70% or higher; only 15% were
hatched at Cape Henry along the ocean. The delay in hatching of the wax
scale at Cape Henry is attributed to the strong eastern airflow along the
coastline that results in lower maximum temperatures in spring months. In
the urban location the buildings protect the scale-infested plants from the
marine breezes and produce the heat-island effect, with reduced evening
cooling due to reradiation.

Application of pheromone technology in the gypsy moth program. C. P.
Schwalbe and E. C. Paszek, USD A, PPQ, Otis Methods Development Cen-
ter, Otis AB, MA 02542

The optically active (+) enantiomer of disparlure is a potent attractant of
male gypsy moths. Traps baited with this material are effective devices for
determining the occurrence and distribution of this pest. A series of field
tests was conducted to define probabilities of capture of moths within var-
ious arrays of traps and to describe adult dispersal behavior in areas trapped.
As trap density decreases, the probability of capturing moths declines. Trap
arrays with intertrap distances (ITD) of 800 m recover ca. 1% of the released
population; 22% was recovered when traps within the array were spaced
175 m apart. It appears that dispersal behavior of males is similar in trapped
and untrapped areas. A grid of traps (ITD = 175 m) established on an area
where moths had been released 48 hr earlier. The distribution of captured
insects was similar to that found when moths were released into the trap
grid. Most insects (ca. 70%) were captured in traps 275 m from the release
point. These data were used to develop a system for estimating probable
boundaries of infestations based upon adult distribution.

The techniques have been applied in two pilot tests in natural infestations.
Both surveys yielded capture patterns which indicated the centers of the
infestations. In one test area, ca. 9 sq. mi. had been presumed to be infested
(based upon earlier surveys). Using the new techniques, it was found that
only ca. 1 sq. mi. was actually infested, thus minimizing the area to be
subsequently involved in control efforts.

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NEW YORK ENTOMOLOGICAL SOCIETY

Life history strategies of spring feeding forest Lepidoptera. D. F. Schweitz-
er, Peabody Museum, Yale Univ. New Haven, CT 06520

Data obtained from outdoor observations in CT, primarily with the noc-
tuid tribe Lithophanini, indicate that overwintering as non-diapausing eggs
results in very early, highly staggered hatching. Early spring oviposition
results in similarly staggered, but later, hatching commencing near the time
of spring foliation. A winter egg diapause usually results in synchronous
hatching shortly after foliation of the host. However, some Catocala have
staggered hatching despite such a diapause. Spring ovipositing Lithophanini
attained closer synchrony (measured in degree-days base 5°C) of egg hatch-
ing with foliation in 1975 and 1978, years with rather cool, late springs, than
in 1976 and 1977, when temperatures over 30°C caused early foliation. Such
adjustable synchrony apparently reduces the risk from unfavorable weather
which is especially likely to follow an early foliation, but results in less
favorable food quality. Staggered hatching dates appear to maximize mini-
mum larval fitness in an environment characterized by erratic weather and
unpredictable foliation dates.

Effect of planting date on the abundance of insect pests on flue-cured to-
bacco. P. J. Semtner and T. R. Terrill, Southern Piedmont Res. and Con-
tinuing Education Center, Blackstone, VA 23824

Research was conducted to determine the effect of planting date on the
occurrence of tobacco hornworms, Manduca sexta (Linneaus) (Lepidop-
tera: Spingidae), tobacco bud worms, Heliothis virescens (Fabricius) (Lep-
idoptera: Noctuidae), and tobacco flea beetles, Epitrix hirtipennis (Mel-
sheimer) (Coleoptera: Chrysomelidae) on flue-cured tobacco. Tobacco was
transplanted at 10-day intervals for 3 dates beginning May 6. The experiment
was established in a split-split plot design replicated 3 times.

At 2 wk after the last transplanting, tobacco flea beetles caused signifi-
cantly (P < 0.05) more damage to tobacco transplanted on the middle date
than on tobacco transplanted early or late. During 1977, the late planted
tobacco had significantly fewer flea beetle feeding holes than the early or
medium planted tobacco. The smaller leaf size of the late tobacco and a
decline in flea beetle populations during late May and early June were re-
sponsible for reduced tobacco flea beetle feeding. When compared to early
and medium planted tobacco, flea beetle populations were significantly low-
er on late tobacco during June. However, during August the late tobacco
had the highest flea beetle populations.

Tobacco budworms were most prevalent (P < 0.01) on early planted to-
bacco, averaging V/i times more budworms/acre than tobacco planted 10
days later and 8 times more budworms than tobacco planted 20 days later.
Tobacco hornworms were most abundant on late planted tobacco.

VOLUME LXXXVIII, NUMBER 1

73

EDNH-stimulated ecdysone production by ovaries of Aedes aegypti (L.)
(DipteraiCulicidae). J. P. Shapiro and H. H. Hagedorn, Dept, of Entomol.,
Cornell Univ., Ithaca, NY 14853

In vitro evidence supports the hypothesis that the egg development neu-
rosecretory hormone (EDNH) of the yellow fever mosquito, Aedes aegypti,
acts by stimulating ecdysone synthesis and secretion by the ovary. Ecdy-
sone may subsequently stimulate vitellogenin synthesis by the fat body. We
studied specific aspects of the ovarian response to EDNH in vitro.

Incubations of ovaries with mid-brain of adult females stimulate ecdysone
secretion into medium. Ovaries and incubation medium contain negligible
amounts of ecdysone prior to incubation, as determined by radioimmu-
noassay (RIA). Upon incubation of ovaries with a saline extract of heads,
nanogram quantities of ecdysone appear in the medium, with little remaining
in ovaries. Actual secretion in vivo is indicated by distribution of ecdysone
in tissues from blood-fed animals.

Ecdysone secretion by ovaries upon exposure to EDNH extracted from
whole heads occurs in a time-dependent manner, beginning within one hour
and continuing at least 10 hours. Secretion is dose-responsive to whole head
extract.

The ovary does not develop the capability to respond to head extract until
36 hours after eclosion; maximal responsiveness is reached by 60 hours at
27°C. Development of ovarian responsiveness to EDNH may be due to the
influence of juvenile hormone during the first 36 hours after emergence.

Encapsulation of pollen attractants for honey bee, Apis mellifera L. diets
(Hymenoptera: Apidae). E. W. Herbert, Jr., H. Shimanuki and B. S.
Shasha, USDA, Beltsville, MD 20705

Honey bees usually prefer to collect fresh pollen when given a free choice
between pollen and pollen substitutes. The addition of attractive fractions
from pollen to increase consumption of bee diets was studied. Pollen sub-
stitutes containing whey and yeast as protein sources were supplemented
with polymers (2, 4, 6 or 8% dry weight) produced by encapsulation of the
chloroform fractions from fresh bee collected pollen in a starch matrix. Diets
containing each level of starch-encapsulated lipid was fed to four colonies
of caged bees and the rate of brood rearing and diet consumption was mea-
sured for 12 weeks. The bees fed whey-yeast diet containing starch encap-
sulated pollen lipids were able to rear significantly more brood to the sealed
stage than bees fed the whey-yeast diets without attractants. Bees fed the
diets containing either 2 or 4% encapsulated lipids reared 2 x/i times as much
brood to the sealed stage as control bees. Bees fed both levels (2 and 4%)
equalled the levels by bees fed pollen. Bees fed the poorest diet (containing
8% lipid) were able to rear twice as much brood as control bees. Consump-

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NEW YORK ENTOMOLOGICAL SOCIETY

tion by bees of all diets containing pollen extracts was significantly better
than for bees fed the attractant free control. The addition of attractive frac-
tions from pollen to substitute diets not only increased consumption of the
diet but also eliminated the possibility of spreading chalkbrood disease,
Ascosphaera apis by the addition of whole pollen to diets.

Establishing economic parameters for green peach aphid, Myzus persicae
(Sulz.) (Homoptera:Aphididae) damage on commercial potatoes. Z. Smilow-
itz, M. E. Whalon, E. S. Nolan and C. A. Martinka. Pesticide Res. Lab.,
Dept, of Entomol., Pennsylvania State Univ., University Park, PA 16802

Following establishment of green peach aphid, Myzus persicae (Sulzer)
(GPA), impact on potatoes the economic consequences of the damage must
be assessed. The procedure used for GPA-CAST, a computerized pest man-
agement system for GPA on commercial potatoes incorporated a market
model, management cost and pest damage index. Pest impact data were
obtained from plant production models and pest densities. A matrix model
was developed to estimate market values which generated 12 distinct crop
values. The percent loss/GPA/leaf throughout the growing season, seasonal
damage index (SDI), was interfaced with the market model to determine
monetary losses. The monetary damage index (MDI), dollar loss/GPA/leaf,
the monetary counterpart of SDI, was obtained as follows: MDI = SDI/FT
x crop value (FT = foliage tuber ratio). Dynamic economic injury levels
(EIL) were obtained from these values and insect management costs per
acre. EIL = management cost/MDI. The economic threshold levels (ETL)
were calculated from the EIL and the projected GPA population growth
rate. The latter was based on the expected accumulated degree days (°D)
for GPA at a specific interval during the growing season. A 3-4 day period
between recommendation and time of application permitted growers ample
time to take appropriate action. The time period averaged out to be 100
accumulated °D. Therefore, the ETL for any interval during the growing
season became the EIL that occurred 100 accumulated °D prior to the cur-
rent accumulated °D for GPA.

The bee louse, Braula coeca Nitzsch (Diptera: Braulidae), its distribution
and preference for queen honey bees. I. B. Smith, Jr., and D. M. Caron,
Maryland Dept, of Agric., Annapolis, MD, 21401 and Univ. of Maryland,
College Park, MD 20742

A survey was conducted in Maryland to determine the infestation level
of bee lice in honey bee colonies. 1881 colonies in 272 apiaries were ex-
amined during inspections by the Maryland Department of Agriculture. Bee
lice were observed in 28% of the apiaries and 18% of the colonies. In apiaries
with lice 50% of the colonies contained lice.

VOLUME LXXXVIII, NUMBER 1

75

Four frame honey bee nucleus colonies were established and stocked with
50 Braula coeca each. One or more lice were present on 24% of the mated
queen honey bees observed between August and December. Only 2% of the
virgin queens observed harbored lice during the same period. Almost no
lice were present on queens during preceding months. In an established
apiary known to be infested with Braula , no lice were observed on queens
April through June 15. 62% of the queens examined from June 16 through
the rest of the season harbored lice and 58% of these lice were pale in color,
indicating Braula were less than one-day old. It appears that newly emerged
bee lice are attracted to mated queens.

Rearing of Tabanus nigrovittatus Macquart (Diptera: Tabanidae) from egg
to adult. R. K. Sofield and E. J. Hansens, Dept, of Entomol. and Econ. Zool.,
Rutgers Univ., New Brunswick, NJ 08903

Host-seeking female salt marsh greenheads, Tabanus nigrovittatus, were
collected from box traps on a salt marsh near Cedarville, NJ. Flies were
allowed to blood feed on human forearm or restrained guinea pig. Ten egg
masses were oviposited by these flies. The egg masses turned grey-brown
with a chalky covering several hours after oviposition. The egg masses were
loosely cemented together. Each egg mass was removed from the cage and
placed on moist filter paper in a petri dish until hatching occurred. Seven
masses hatched after 5 days yielding 508 larvae. Larvae were reared at 27
C in individual 8 dram vials with wet filter paper. Every 2-5 days the filter
paper was changed and fresh food (house fly maggot or earth worm) pro-
vided. Larval mortality was highest in the first 40 days. Only 12.5% of the
larvae survived this period. Three to five months after hatching, half the
larvae were subjected to 5 C for 40 days and the remainder for 80 days. The
rearing temperature was then returned to 27 C. Pupation of 26 larvae from
four egg masses occurred 6-9 months after hatching. A single larva pupated
60 days after hatching. The pupal stage lasted 7-12 days. Of the 14 females
reared, 7 oviposited a mass of infertile eggs, 3 took a blood meal after
oviposition, and one fly oviposited a second mass of infertile eggs.

Transmission of equine babesiosis and bovine anaplasmosis by Dermacen-
tor albipictus (Packard) (Acari: Ixodidae). D. Stiller, W. M. Frerichs, G.
Leatch*, and K. L. Kuttler, Animal Parasitol. Inst., SEA, USDA, Belts-
ville, MD 20705, and Commonwealth Sci. and Indust. Res. Org., Indoo-
roopilly, QLD 4068, Australia

Although the 1-host winter tick, Dermacentor albipictus, is a common
parasite of horses and cattle and is widely distributed in the U.S., its role
as a potential vector of the hemoparasitic diseases equine babesiosis and
bovine anaplasmosis has been little studied. Experiments to assess the vec-

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NEW YORK ENTOMOLOGICAL SOCIETY

tor competence of the inornate form of D. albipictus for the horse hemo-
parasite Babesia caballi revealed that (1) 70% of female ticks fed on an
infected pony became infected, as demonstrated by Giemsa-stained smears
of tick hemolymph; (2) the Fj progeny of these females were infected trans-
ovarially and transmitted the infection when test-fed on susceptible ponies;
and (3) male ticks transmitted the infection after being removed from an
infected pony, held for 1 hr, and transferred to a susceptible pony. Similar
experiments with this tick and the cattle hemoparasite Anaplasma margin-
ale indicated that (1) nymphal ticks, fed as larvae on an infected calf, trans-
mitted anaplasmosis when test-fed on a susceptible calf and (2) male ticks
transmitted the infection after being removed from an infected calf, held for
1 or 4 hr, and transferred to susceptible calves. Tests of transovarial trans-
mission are in progress. This is the first unequivocal evidence incriminating
D. albipictus as an experimental vector of these pathogens and suggests
that this tick may be a potential vector of these agents in nature.

Soybean foliage consumption by the adult Mexican bean beetle, Epilachna
varivestis Mulsant (Coleoptera: Coccinellidae). I. M. Sunzenauer1, T. El-
den2, and A. L. Steinhauer1, Dept, of Entomol., Univ. of Maryland, College
Park, MD 207821 and USDA, SEA, Beltsville, MD 207052

Foliage feeding studies have been conducted on a variety of soybean
insects, but are lacking for the adult Mexican bean beetle. Such information
is needed by researchers developing decision-making models for the Mex-
ican bean beetle in soybeans. In this study the daily feeding rate of the adult
Mexican bean beetle was determined by measuring leaf area consumption.
Daily oviposition rates were also obtained. Three different populations of
adults were used. Population I consisted of 2nd generation adults, Popula-
tion II consisted of adults which had overwintered in the field, and Popu-
lation III consisted of adults which had overwintered in the laboratory. Pairs
of beetles in each study were set up in petri dishes in an environmental
chamber with fresh foliage, which was changed after specific time intervals,
and the number of eggs was recorded. Consumed leaf area was measured
by a LI-COR Leaf Area Meter from the Lambda Instruments Corporation.
The mean daily leaf area consumption rates for Populations I, II, and III
were 4.65 cm2/day, 4.87 cm2/day, and 3.80 cm2/day respectively. The mean
daily oviposition rates for Populations I, II, and III were 11.10 eggs/day,
6.77 eggs/day and 5.37 eggs/day. The lower feeding and oviposition rates for
Population III resulted in part from the 7 months in cold storage.

VOLUME LXXXVIII, NUMBER 1

77

Humidity tolerances of 3 species of phytoseiid mites (Acarina: Phytosei-
idae). F. C. Swift and L. Blaustein, Rutgers-The State Univ., New Bruns-
wick, NJ 08903

The effects of humidity on hatching success, survival of adult females,
and rates of feeding and oviposition were determined for Neoseiulus fallacis
(Garman), Neoseiulus mckenzei (Schuster and Pritchard), and Amblyseius
andersoni Chant. Each factor was measured at relative humidities of 43, 51,
64, 75, 85, 90, and 97%. All experiments were conducted at 25°C and a 16:8
light-dark cycle. Eggs of A. andersoni showed the greatest resistance to
desiccation. Also, A. andersoni females deprived of food survived longer
than either N. fallacis or N. mckenzei at lower humidities, but the latter
species survived longest at the highest humidity. Survival time over the
range of humidities was exponential for all 3 species. The feeding rate of N.
fallacis was most affected by humidity, showing approximately a 3-fold
increase from the highest to the lowest humidity. Oviposition rates of N.
fallacis and A. andersoni were affected only slightly by humidity, but the
oviposition rate of N. mckenzei declined rapidly at humidities below 75%.
A ratio of oviposition rate/feeding rate was used as a measure of humidity
niche breadth, using Levin’s formula (1968), divided by the number of re-
source states. According to this formula, A. andersoni had the widest niche
breadth (0.96), followed by N. fallacis (0.82) and N. mckenzei (0.50). How-
ever, N. fallacis compensated by certain behavioral adaptations; namely,
higher feeding rates at lower humidities and the placement of eggs in mi-
crohabitats of higher humidity.

Hudson Porta-Pak ULV Sprayer. R. Treichler, H. D. Hudson Manufactur-
ing Company, 500 North Michigan Ave., Chicago, IL 60611

The Hudson Porta-Pak Sprayer is the only back-carried motorized unit
that is designed specifically for use in ultra-low-volume pesticide applica-
tion. The air volume output at full throttle is 400 CFM; the air speed at the
nozzle is 230 MPH. The pesticide tank holds 106 ozs; the fuel tank 68 ozs.
With a nominal discharge rate of 1.6 ozs. p.m. a full tank supplies 1 hr of
application or Vi hr at a full discharge rate. Walking at 1. 1 MPH, over 1 mile
will be covered discharging 1 tank full at 1.6 ozs. p.m. If the application is
horizontal, the 50 foot swath and 1 pesticide tank contents will cover a IV2
acre area. Hudson Porta-Pak is ideal for fruit and ornamentals. Solutions,
emulsions and suspensions of wettable powders can be used to apply pes-
ticide to trees, shrubbery and truck crops. Porta-Pak is especially useful in
cattle feed lots and swine confinement areas to control mosquitoes, flies and
other pests. The turbulent air stream and spray swath of 40 ft. vertical and
over 50 ft. horizontal makes it especially suited for both fruit and ornamental

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trees. The Hudson Porta-Pak Sprayer is ideal for mosquito and fly control
in parks, stadiums, playgrounds and picnic areas.

Integration of Rhinocyllus conicus Froelich larvae (Coleoptera: Curculion-
idae) and 2,4- D for control of Carduus nutans L. J. T. Trumble and L. T.
Kok, Dept, of Entomol., VPI & State Univ., Blacksburg, VA 24061

Herbicidal effect on Rhinocyllus conicus Froel., a thistle head weevil,
was studied by examining the mortality, emergence rates and weights of
weevils developing from plants treated with 2,4-D (LVA). Infested heads,
obtained by caging ovipositing R. conicus on primary heads of musk thistle
( Carduus nutans L.), were treated with 2,4-D at 1.68 kg/ha 0 to 3 weeks
after oviposition. Mortalities of larvae developing from untreated plants and
those plants sprayed 1 to 3 weeks were significantly lower than mortality
from plants sprayed within 48 hr of oviposition. The latter failed to support
larval development beyond the second instar. Developmental times and
weights of weevils that emerged from blooms sprayed at 1,2, and 3 weeks
were not significantly different from controls. Plants sprayed up to 2 weeks
after oviposition (late-bud to early-bloom) did not produce viable seeds, but
treatments at 3 weeks after oviposition (full-bloom) allowed 10% germina-
tion of seeds not damaged by R. conicus in primary heads, and plants
survived to produce additional heads. Treatment of musk thistles with 2,4-
D at late-bud to early-bloom stage of the primary heads prevented formation
of viable seeds without adversely affecting R. conicus development.

Characterization of the structural components of the granulosis virus of
Plodia interpunctella (Lepidoptera: Pyralidae). K. A. Tweeten, 2810 Fawkes
Dr., Wilmington, DE 19808

Procedures were developed for isolation of the following structural com-
ponents of Plodia interpunctella granulosis virus: granulin, enveloped nu-
cleocapsids, nucleocapsids, capsids, and DNA. Biophysical and biochemi-
cal techniques were utilized to extensively characterize these components.
Two molecular forms of DNA were extracted from the virus. Twenty-five
percent of the DNA was isolated as covalently closed circles; the remaining
seventy-five percent consisted of relaxed circular molecules. The molecular
weight of the DNA was estimated to be 80 x 106 daltons from sedimentation
in neutral or alkaline sucrose gradients and from electron micrographic
length measurements. The polypeptide composition of each viral component
was determined by sodium dodecyl sulfate discontinuous polyacrylamide
slab gel electrophoresis. The enveloped nucleocapsids consisted of fifteen
structural proteins ranging in molecular weight from 12,600 to 97,300. Of
these proteins, eight were present in nucleocapsids. The major nucleocapsid

VOLUME LXXXVIII, NUMBER

79

proteins had molecular weights of 12,500 (VP 12) and 31,000 (VP31). Gel
electrophoresis provided evidence that VP31 was the major capsid poly-
peptide while VP 12 was an extremely basic protein which was located inside
the capsid, probably in association with the viral DNA.

Biological characterization of AMBUSH insecticide. M. Tysowsky, ICI
Americas, Biological Research Station, Goldsboro, NC 27530

AMBUSH, a synthetic pyrethroid insecticide being developed by ICI
Americas has been used successfully under grower conditions over the past
two years for control of Lepidoptera and a number of selected Coleoptera.
Rates of 0.05-0.2 lb ai/A have been evaluated on pests which attack apples,
celery cole crops, corn, cotton, lettuce, potatoes, tomatoes and soybeans.
However, very little has been reported concerning the biological properties
of this new chemical.

AMBUSH works by contact and it is extremely effective when ingested.
The material acts as disruptant. Lab and field tests have demonstrated that
although insects may not die quickly, feeding stops within minutes after
contact. Death may take from 24-72 hours. AMBUSH provides a high level
of activity against various instars of lepidopterous larvae with some control
of adults and suppression of egg hatch. Bioassays with Lepidoptera have
shown that this material possesses ovicidal activity comparable to that
achieved by chlordimeform. Tank mixes with other insecticides are not
recommended as they can result in delayed crop maturity.

The material is environmentally sound and has a wide margin of safety
to humans. However, like the natural pyrethrins, it is toxic to bees and fish.
AMBUSH is very stable under sunlight conditions and it is tightly bound
to soil particles.

Gas-liquid chromatographic comparisons of the cuticular lipids from male
and female house crickets, Acheta domesticus (L.) (Orthoptera: Gryllidae).
E. C. Uebel and J. D. Warthen, Jr., Biol. Act. Nat. Prod. Lab., AEQI,
USD A, SEA, Beltsville, MD 20705

The main objective in the investigation of the cuticular lipids of the cricket
was to check for compounds that were more abundant on the adults of one
sex than on the other and which might serve as recognition pheromones for
members of the opposite sex. R. Paul (Nature, 1976. Vol. 263:404-405) has
demonstrated that the males of several species of ground crickets are able
to detect chemical stimuli from conspecific females and respond with calling
songs. The hydrocarbons and cuticular lipids from mixed sexes of house
crickets have been identified by Hutchins and Martin (Lipids. 1968. Vol.
3:250-255) and Blomquist et al. (Comp. Biochem. Physiol. 1976. Vol.

80

NEW YORK ENTOMOLOGICAL SOCIETY

54B:38 1-386). Gas-liquid chromatography on 2% OV 101 showed both a
qualitative and quantitative similarity between the cuticular lipids present
on adult females, 8th-instar males, and 8th-instar female nymphs. However,
the adult males produced much more material chromatographing between
36- and 37-carbon n -alkane standards. This large peak of material consisted
of ca 13% saturated and 87% unsaturated hydrocarbon. Approximately 33%
of the 37-carbon unsaturates were monoenes and 66% were dienes. At the
present time we are trying to characterize these male 37-carbon mono- and
dienes by locating the positions of the double bonds and the methyl branch-
ing. Whether these male-produced monoolefins and diolefins have a com-
municative function between the males and females or between other males
remains to be investigated.

Life history and population dynamics of tree-dwelling Homoptera. J. W.
Webb, Oak Ridge Nat. Lab., Oak Ridge, TN 37830

There are few instances in which the role of the host plant in the popu-
lation dynamics of herbivorous insects has been assessed in the field. Two
studies of leaf feeding Homoptera were undertaken to help elucidate this
role. Estimates of field population numbers and biomass were analyzed
along with data on host plant physiological condition and phenology. Spring
population numbers of the South African psyllid Acizzia russellae were
correlated with nitrogen levels in the leaves of its host, Acacia karroo. Also,
populations were almost 10 times greater on the stump sprouts of cut trees
than on leaves of uncut trees. In another study, aphid populations on 8
species of North American hardwoods showed similarly dramatic growth
on stump sprouts following spring cutting. Homoptera which feed on cellular
contents did not increase on cut plants, probably because leaf nitrogen con-
centrations were no higher than in uncut plants. Phloem feeding species,
however, apparently responded mainly to the increased flow rate of nu-
trients in growing tissues of cut trees. Estimates of consumption using aphid
densities from cut plants and literature values for ingestion were about 25%-
50% of leaf standing crop. The uptake and cycling of such quantities, re-
sulting from changes in plant condition, could have significant effects on
plant function. These results clearly demonstrate that the condition of the
host plant is a major factor influencing population densities in phloem feed-
ing Homoptera, and suggest that feedback effects on plants may also be
important.

VOLUME LXXXVIII, NUMBER 1

81

Distribution of Phyllonorycter (= Lithocolletis ) sp. in northeastern apple
orchards. R. W. Weires and J. R. Leeper, Hudson Valley Lab., Highland,
NY 12528

Surveys were conducted during 1976-1978 throughout Northeastern apple
orchards to determine which leafminer species were responsible for recent
outbreaks. Samples of leaves were collected from beneath the apple trees
during the winter months. Apple branches and leaves were collected during
the summer months from several New York, Connecticut, Vermont, and
Massachusetts commercial orchards. All stages were either collected or
reared from the leaf samples for specialists to examine. In addition, phero-
mone traps containing the Phyllonorycter blancardella sex attractant were
deployed in several New York orchards and the adult males captured were
saved for identification. Phyllonorycter crataegella was the predominant
species found in Massachusetts, Connecticut, and the area East of the Hud-
son River in New York state. Phyllonorycter blancardella was the predom-
inant species found in Western New York, the Champlain Valley, and Ver-
mont. Both species were found in several orchards in Ulster, Greene,
Saratoga and Washington counties of New York. The pheromone trap catch-
es were predominately restricted to detecting the Phyllonorycter blancar-
della species with only very low numbers of crataegella found in the traps.
Both species have strains which are not controlled by the present organo-
phosphate insecticide programs.

Seasonal history of the leafhopper complex (Homoptera:Cicadellidae) on
ornamental honeylocust. A. G. Wheeler, Jr. and K. Valley, Bureau of Plant
Industry, PA Dept. Agric., Harrisburg, PA 17110

Leafhoppers have been included among the arthropods known to injure
ornamental honeylocust. Since little biological data were available to im-
plicate leafhoppers as pests, we studied the complex on ‘Sunburst’ honey-
locust by sampling weekly the terminal 36 cm of 4 branches on 2 trees at
Harrisburg, PA, during 1975-77. Four common species, exhibiting spatio-
temporal differences were found: Macropsis fumipennis, Stragania apical-
is, Orientus ishidae, and Empoasca sp. Eggs of M. fumipennis hatched
shortly after leaf flush; nymphs fed on petiolules and rachises and were
present from mid-April to late May. Adults matured in mid-May and per-
sisted until late July. Large populations near the sample site, in the virtual
absence of honeylocust plant bud ( Diaphnocoris chlorionis ), failed to pro-
duce the leaflet distortion characteristic of mirid injury. Nymphs of S. ap-
icalis, present from late April to mid-May, fed mainly on leaflets. Although
adults were found until September, we were unable to determine whether
this species is bivoltine. Nymphs of O. ishidae were found from early June

82

NEW YORK ENTOMOLOGICAL SOCIETY

to early July; adults, until late July. This introduced species fed on leaflets
of water sprouts, yellowing the foliage. Empoasca adults, which appeared
during June, may migrate each season into Pennsylvania. This leafhopper
feeds mainly on the second flush of new growth and produces 2 or 3 gen-
erations by September. Our observations indicate that distortion of honey-
locust foliage should be blamed on honeylocust plant bug, not on the pe-
tiolule-feeding M. fumipennis or other leafhoppers.

Physiological age study of five species of mosquitoes (Diptera: Culicidae)
in southeastern New Hampshire. J. J. Winegar, Dept, of Entomol., Univ.
of New Hampshire, Durham, NH 03824

Adult females of Aedes excrucians , A. canadensis, A. cantator, A. vex-
ans, and Coquillettidia perturbans were collected by human-bait capture,
net sweeping, and C02-baited CDC light traps in Exeter, New Hampshire,
from June to September, during 1978. Mosquito specimens were stored at
— 12°C for later dissection. Physiological age for 1,818 females was deter-
mined by the Polovodova method. The data revealed that females of each
species could complete a minimum of two gonotrophic cycles. A. excrucians
and A. canadensis had the greatest longevity, with 12.8% of A. excrucians
and 10.9% of A. canadensis surviving to become 3-parous, one A. cana-
densis female was 4-parous. Females of A. cantator, A. vexans, and C.
perturbans determined to be 2-parous were uncommon, comprising 0.6% to
3.0% of total number dissected. All species studied are anautogenous. Parity
data showed that A. excrucians and A. canadensis were univoltine and
emerged synchronously. A. cantator and A. vexans appear to have three
distinct broods, while C. perturbans had a single brood, but emerged rela-
tively asynchronously. A. excrucians and A. canadensis may be epidemi-
ologically important in southern New Hampshire, particularly as potential
vectors of dog heartworm, Dirofilaria immitis. Absence of multiple feeding
in most A. cantator, A. vexans, and C. perturbans indicates these species
probably played a very minor role as disease vectors in New Hampshire,
in 1978, possibly due to extremely dry weather during the summer.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(l), 1980, p. 83

BOOK REVIEW

Comprehensive Virology 12. Newly characterized protist and invertebrate
viruses. Heinz Fraenkel-Conrat and Robert R. Wagner, ed. Plenum Pub-
lishing Co., New York. 1978. 344 p. $29.50.

This volume contains 5 chapters and entomologists will be particularly
interested in the first and longest one, by T. W. Tinsley and K. A. Harrap,
which provides a summary of viruses of invertebrates. The baculoviruses
which comprise the nuclear polyhedrosis and granulosis viruses are already
being used as “living insecticides” in the United States and in several other
countries, because they differ from all viruses that infect higher animals and
plants. They have normally a very narrow host range and their morpholog-
ical characteristics are well illustrated by electron micrographs provided in
this volume. Other groups of insect viruses, such as the entomopoxviruses,
iridoviruses, the isometric RNA viruses to which the densonucleosis virus
belongs and the rhabdoviruses are described in some detail. Main emphasis
is on morphology because other characteristics have been studied to a lesser
degree. An excellent review of the ecology and epizootiology is included in
the first chapter and the literature citations comprise 20 pages. Unfortu-
nately the chapter has been prepared several years ago and its publication
delayed so that only a few references are as recent as 1975. This is especially
regrettable since the authors, working at the Unit of Virology at Oxford,
England, have made significant contributions in recent years and have char-
acterized baculoviruses by modern techniques.

The remaining chapters describe viruses of fungi, cyanobacteria and pseu-
domonads, as well as the intriguing bacteriophages found in antibiotic-pro-
ducing strains of Penicillium. The chapter on this subject is by T. I. Tik-
chonenko of the Ivanovsky Institute of Virology in Moscow, where most
of this work has been done in recent years. A good subject index is provided.

Karl Maramorosch,

Waksman Institute of Microbiology ,

Rutgers — The State University ,

New Brunswick, N.J.

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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

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Founded in 1872 — Incorporated in 1885
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Officers for the Year 1980

President , Dr. Betty Faber

American Museum of Natural History, New York 10024
Vice-president , Dr. Joseph Cerreta

Columbia University, New York 10032

Fordham University, New York 10458
Assistant Secretary , Dr. Henry M. Knizeski, Jr.

Mercy College, Cobs Ferry, New York 10522

Secretary , Dr. Gerard Iwantsch

Treasurer , Mr. Louis Sorkin

4

American Museum of Natural History, New York 10024

Assistant Treasurer , Dr. Sally B. Padhi

Rutgers University, New Brunswick, New Jersey 08903

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Class of 1980

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Ms. Jane Harvey Cooper

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Trustees

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Class of 1981

Dr. Ivan Huber

•

Ms. Laurene Ford

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Dr. Irene Matejko

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Drexel University, Philadelphia, Pennsylvania 19104

Fordham University, New York 10458

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Mailed July 16, 1980

The Journal of the New York Entomological Society (ISSN 0028-7199) 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.

Journal of the N.Y. Entomological Society, total No. copies printed 750. Paid circulation 490, mail subscription 470, free
distribution by mail 23, total distribution 493, left-over 257 copies each quarter.

/; v; ,::YY Yii '

Journal of the

New York Entomological Society

VOLUME LXXXVIII JUNE 1980 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. Streu

Publication Committee
Dr. Randall T. Schuh
American Museum of Natural History
Dr. Daniel Sullivan
Fordham University
Dr. Felix J. Bocchino
The College of Mt. Saint Vincent

CONTENTS

Environmental partitioning in lowland tropical rain forest cicadas

Allen M. Young 86-101

Simultaneous use of a foraging trail by two leafcutter ant species in the Sonoran
desert Alex Mintzer 102-105

Causes of life history differences between the morphs of Pemphigus populitrans-
versus Lorenz Rhomberg 106-112

Epistylus cambari (Ciliata: Peritrichida) and dragonfly nymphs, an epizoic asso-
ciation Reid Jilek 113-114

Total concentration of free amino acids and the corresponding morphogenetic
changes during the embryogenesis of Chilomenese sexmaculata Fabr. (Coleop-
tera: Coccinellidae)

Devinder Mukesh, Dalbinder Singh Sidhu and Nirmal Kumar 115-119

A conspectus of Pentatomini genera of the Western hemisphere. Part I (Hemiptera:
Pentatomidae) L. H. Rolston, F. J. D. McDonald, and Donald B. Thomas, Jr. 120-132

Anatomy of the male reproductive systems of the adults and pupae of two dory-
line ants, Dorylus (. Anomma ) wilverthi Emery and D. (A.) nigricans Illiger

Francis C. Ford and James Forbes 133-142

Apparent nest site competition between the paper wasp Polistes fuscatus (Hyme-
noptera: Vespidae) and the house wren David L. Gibo 143-145

Overwintering of Polistes fuscatus in Canada: use of abandoned nests of Do-
lichovespula arenaria David L. Gibo 146-150

Population trends of the alfalfa weevil (Coleoptera: Curculionidae) and its asso-
ciated parasites in Maryland and New Jersey, 1966-1970

Robert F. W. Schroder and William W. Metterhouse 151-163

Book review

164-165

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(2), 1980, pp. 86-101

ENVIRONMENTAL PARTITIONING IN LOWLAND TROPICAL
RAIN FOREST CICADAS

Allen M. Young

Abstract. — The peak emergence periods and habitats of cicadas were ex-
amined discontinuously over several years in lowland tropical rain forest of
northeastern Costa Rica. Two distinctive ecological features of this region
which might affect cicadas are (1) the existence of primary and secondary
forest habitats, and (2) the occurrence of a short dry season. Although the
data on adults concern six genera and eleven species, emphasis is placed
on quantitative sampling of nymphal casts from ground plots for Zammara
smaragdina, Fidicina s eric arts, Fidicina mannifera, Fidicina pronoe, Fidi-
cina spinocosta, and Quesada gigas. If selection pressures favor emergence
adaptations such as allochrony or habitat non-overlap among some species,
it was expected that the census data would show these patterns. Generally
the data support this hypothesis. For example, of the two most abundant
cicadas, Z. smaragdina and F. sericans, the former emerges primarily dur-
ing the long wet season while the latter emerges during the dry season. Both
species occur in primary forest but Z. smaragdina also occurs in advanced
secondary forest. Another, F. mannifera , is generally not very abundant,
has low emergence almost continuously throughout the year, and sings at
times of the day when others are less active. Others such as F. spinocosta
and F. pronoe and Q. gigas emerge in young secondary forests but in
different seasons. Owing to the high selective value placed upon the cicada
song in breeding, temporal and spatial partitioning of the environment may
be an adaptive measure that reduces interspecific competitive conflicts that
could arise by adults of abundant species being active at the same times of
the year or in the same habitats.

Introduction

This paper reports the peak emergence periods and habitats for several
genera and species of cicadas (Homoptera: Cicadidae) in the lowland trop-
ical rain forest of northeastern Costa Rica. While six genera and eleven
species of cicadas occur in this region (Young 1972; pers. obs.), quantitative
data on emergences were obtained for three genera and six species: Zam-
mara smaragdina Walker, Fidicina mannifera (Fabricius), Fidicina seri-
cans Stal, Fidicina spinocosta (Walker), Fidicina pronoe (Walker), and
Quesada gigas (Olivier) (Figs. 1-4). These are large-bodied cicadas that
were previously determined (pers. obs.) to occupy different habitats (Table

VOLUME LXXXVIII, NUMBER 2

87

Fig. 1. Zammara smaragdina \ female, male, male (lateral view). This is a loud and large-
bodied green-and-brown cicada of the Caribbean lowland tropical rain forest.

1). Owing to the availability of several distinct habitats and a pronounced
dry season (e.g., Slud 1974), it was predicted that different species may
exhibit different habitat preferences and annual periods of peak emergences
if they were in competition for some resources. While this study did not
determine these resources, the data indicate considerable allochrony and
non-overlap in habitat for these cicadas.

Region and Habitats

The cicadas were studied at "Finca La near La Virgen de

Sarapiqui (220 m elev.), Heredia Province. The site is a 1,000-acre farm
complex with primary forest (mostly fragmented into remnants) and a va-
riety of secondary forest habitats. The dominant tree in the primary forest
remnants is Pentaclethra macroloba (Willd.) Ktze. (Leguminosae) whose
life history has been described by Hartshorn (1975). Advanced secondary
forest (over 25 years old) is dominated by trees of Goethalsia meiantha

88

NEW YORK ENTOMOLOGICAL SOCIETY

Fig. 2. Majeorona bovilla and Fidicina mannifera (left to right) are large, black-and-brown-
bodied cicadas of primary forest; their songs are loud and F. mannifera sings primarily at
dawn and dusk.

(Donn. Sm.) Burret (Tiliaceae) forming a canopy of about 25 m (DBH = 9
cm) and with a dense understory of Pentaclethra seedlings and saplings.
Young (2-10 years old) secondary forest contain a large number of woody
shrubs and treelets (canopy less than 4 m) such as Vernonici patens H.B.K.
(Compositae) and Vismia guianensis (Aubl.) Pers. (Guttiferae). A short dry
season falls between January and April (Frankie et al. 1974). Generally the
rainfall is less than 225 mm per month during the dry season, especially in
March and April (Fig. 5). The pattern shown in Figure 5 is typical for several
years (e.g., 1971-78) (pers. obs.).

Census Methods

Population size, density, and sex ratio were obtained for each cicada
species by collecting nymphal casts from plots of known size. The majority
of cicada censuses were taken between June 1972 and July 1974, although
a few additional ones were made in 1976 and 1978. These censuses were

VOLUME LXXXVIII, NUMBER 2

89

Fig. 3. Left to right: Fidicina sericans, F. pronoe, F. amoena, and F. spinocosta. With
the exception of F. sericans, these cicadas occupy young secondary forest. Fidicina sericans
and F. pronoe are black-bodied and they are active primarily in the dry season; F. amoena
and F. spinocosta are green-bodied wet season species and the latter sings primarily at dusk.

discontinuous and were timed to sample habitats at different times of the
year. Zcunmcira smaragdinci was censused in 23 nested quadrats (see “quad-
rat area” in Fig. 6), each 6 by 5 meters, located on a hill covered with
Goethalsia (advanced secondary forest — Fig. 7) for a total of 30 days over
the three years. Eleven additional scattered quadrats of the same size, each
one around a large Pentaclethra in forest remnants (Fig. 8), were used to
census Z. smaragdinci , F. mannifera, and F. sericans for a total of 28 days
in the same period. Several other isolated plots of varying size (“road-edge
plot” — 6 by 5 m; “hill-top plot” — 9 by 6 m; “river-edge plot” — 12 by 8 m)
were established in different types of secondary forest (Fig. 6) to census F.
spinocosta and others. A total of 34 days were spent censusing nymphal
casts from these three plots. Fidicina pronoe was censused (March 1976)
in a pasture covered with Vernonia shrubs and following a brief dry season
census of Z. smciragdina in the “quadrat area”; Quesada gigas and F.

90

NEW YORK ENTOMOLOGICAL SOCIETY

Fig. 4. Quesada gigas and Fidicina pronoe (inset) feeding at branches of Vernonia patens
(Compositae) in young secondary forest during the dry season, a time of peak emergence of
these species.

pronoe were censused in February 1978 in a plot (20 by 10 m) in a grove of
nutmeg trees, Myristica fragans Houttuyn (Myristicaceae). During most
censuses, two or three trained workers collected nymphal casts together
from each quadrat or plot. The nymphal casts of different cicadas were

Table 1 . Habitat associations of adult cicadas in northeastern Costa Rica, below 250 m elevation.

VOLUME LXXXVIII, NUMBER 2

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92

NEW YORK ENTOMOLOGICAL SOCIETY

Fig. 5. The rainfall pattern at “Finca La Tirimbina" during a one and one-half year period
(data courtesy of Dr. J. Robert Hunter). A dry season occurs between January and April, with
monthly rainfall during these months usually falling below 225 mm. The patterns shown are
typical for several years in this region.

readily distinguishable from one another (see also Young 1972), facilitating
field censuses.

Results

Habitat Separation by Adult Cicadas

Of eleven species that occur in this region, adults of seven were seen and
heard in primary forest remnants, and these species comprise the greatest
range in body length (Table 1). Only one cicada, Z. smaragdina , overlaps
with other habitats, namely advanced secondary forest and the highest num-
ber of congeneric species occurs in primary forest (Table 1). In primary
forest, adult Z. smaragdina feeds on, and choruses from Pentaclethra trees,
as does F. sericans. In advanced secondary forest, the former species is
similarly associated with Goethalsia trees. Where this habitat is contiguous
with primary forest, sometimes dense aggregations of F. sericans adults are

VOLUME LXXXVIII, NUMBER 2

93

Fig. 6. The areas used to examine cicada emergences: (a) "quadrat area” on a Goethalsia-
covered slope along the Rio Tirimbina (positions of quadrats shown by the numbered squares);
(b) a series of large Pentaclethra trees in primary forest, shown here by numbered or lettered
circles; (c) three large "plots” as indicated here. The areas designated as "lote A” and "lote
B” indicate sites of cultivation for crops such as cocoa, black pepper, nutmeg, etc. Modified
version of a map provided by Dr. J. Robert Hunter.

seen in Goethalsia trees. In some patches of advanced secondary forest
there are feeding aggregations of F. mannifera , F. sericans, and Z. sma-
ragdina on Zanthoxylum procerum Donn. Sm. (Rutaceae). Both F. sericans
and Z. smaragdina oviposit in seedlings of Pentaclethra in primary forest,

94

NEW YORK ENTOMOLOGICAL SOCIETY

Fig. 7. The Goethalsia -covered "quadrat area” where Zammara smaragdina was cen-
sused.

while the former also oviposits in a variety of understory woody shrubs in
advanced secondary forests. In young secondary forests, Q. gigas and F.
pronoe feed and oviposit in shrubs of Vernonia and occasionally in Inga
sp. (Leguminosae). Fidicina spinocosta adults are associated with a wide
range of woody shrubs including Vismia. Carineta indecora is a small cicada
associated with the primary forest understory (Young 1972), especially near
light gaps, while F. amoena is most frequently heard in large trees along
streams in primary forest. Majeorona bovilla adults occur in patches along
borders of primary forest and Proarna sallei choruses in tall Cordia allio-
dora (R. & P.) Oken. (Boraginaceae) along the borders of primary forest
(see also Young 1972).

Some cicadas exhibit pronounced seasonality in adult activity as indicated
primarily by choruing activity. The greatest number of chorusing Z. sma-
ragdina are heard during the mid- wet season, especially in July-August
while intense chorusing in F. sericans takes place in the late dry season,
especially March-April. Others, such as F. pronoe and Q. gigas are most

VOLUME LXXXVIII, NUMBER 2

95

Fig. 8. Zammara smaragdina (female). Below: primary forest remnants dominated by Pen-
taclethra are habitats for Zammara smaragdina, Fidicina sericans, Fidicina mannifera, and
Majeorona bovilla.

96

NEW YORK ENTOMOLOGICAL SOCIETY

Fig. 9. Peak emergence periods for Zammara smaragdina in the "quadrat area” for three
successive years. Peak numbers of nymphal casts appear in mid-wet season, July and August,
each year.

active near the end of the dry season (April); F. spinocosta, F. amoena, C.
indecora and M. bovilla are active during the wet season. Two cicadas, F.
mannifera and P. sallei , are generally heard throughout the year but with
some diminishing activity in November-December. In summary, of the
eleven species seven are active during the wet season (including two which
are active throughout the year); the wet season generally lasts eight months.
Although F. mannifera adults are active in the same habitat with F. sericans
and Z. smaragdina and at the same times of the year, chorusing intensity
is greatest in the early evening (5:00-6:00 P.M.) in the former cicada. Most
of the other cicadas chorus throughout the day, although F. spinocosta and
Q. gigas often exhibit pronounced bursts of activity near dawn and dusk.
In fact, while F. pronoe generally sings throughout most of the day and
occasionally near dusk, Q. gigas in the same habitat exhibits a large burst
of singing at dusk and sings only sporadically during the day.

VOLUME LXXXVIII, NUMBER 2

97

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Patterns of Cicada Emergences

The highest number of nymphal casts of Z. smaragdina were found during
mid-wet season (July-August) and very few during the late dry season
(March-April) and the emergences declined over the three years (Fig. 9).
Although these data are discontinuous (Fig. 9), visits to the area following
periods when no censuses were taken revealed the same abundance. The
number of cicadas emerging from each quadrat varied considerably on each
census date (as much as 100-fold) and the sexes occurred equally (Table 2).
A few quadrats consistently yielded the greatest numbers of cicadas; of the
2,418 cicadas emerging from the area over three years, about 15% came
from quadrat no. 23. Other quadrats produced less than ten cicadas each
year, and overall the area produced about 3.5 cicadas per square meter for
the study period or about one cicada per square meter each year. All nymph-
al casts censused in the area were Z. smaragdina.

The mean number of nymphal casts of this cicada obtained from the
Pentaclethra trees was 19.37 ± 31.18 (SE; N = 11) for the same period,
and this figure is within the range of mean values for the “ quadrat area”
during periods of peak emergence (Table 2). However, Fidicina sericans
was the most abundant cicada emerging at Pentaclethra, producing 28.46 ±
48.99 nymphal casts per tree during the late dry season, while F. mannifera
was far less abundant producing 4. 1 ± 6.5 nymphal casts for the same trees
and period. Although the sample size of Pentaclethra is small, F. sericans
emerged at all eleven trees, while Z. smaragdina at eight, and F. mannifera
at four. Subsequent anecdotal observations (e.g., January-February 1976)
indicated that although Z. smaragdina peaks in emergence in the wet season
and F. sericans during the late dry season, a few fresh nymphal casts of
the latter have been collected during brief dry periods amid an otherwise
wet season. Likewise, a few Z. smaragdina emerge during brief rainy pe-
riods during the dry season.

In some areas of primary forest, dense emergences of F. sericans were
observed during the dry season. For example, during a five-week period in
the 1975 dry season, 903 nymphal casts of this cicada (9.3 cicadas per square
meter) were collected from the “river-edge plot” (Fig. 6) while only 1,800
nymphal casts were collected from the Pentaclethra over three years (5.4
cicadas per square meter). The river-edge emergence of this cicada repre-
sented a ten-fold increase when compared to the emergence of Z. sma-
ragdina in the “quadrat area.”

Fidicina spinocosta emerges during the wet season in young secondary
forest areas such as the “road-edge plot” (Fig. 7) where 120 nymphal casts
(1.5 cicadas per square meter) were collected in 1973 and 1974 (seven col-
lections); most of these were clumped around two Vismia trees. Another
148 nymphal casts (two collections) were collected in the 1974 wet season

VOLUME LXXXVIII, NUMBER 2

99

from the “hill-top plot” (Fig. 7) (2.7 cicadas per square meter). Seventy-
five nymphal casts of Z. smaragdina and one each of F. sericans and F.
mannifera were also collected from this plot during the same period. Two
other cicadas, F. pronoe and Q. gigas, emerge in young secondary forest
habitats, but in ones that are abandoned pastures overgrown with the shrub
Vernonia. Fidicina spinocosta and the other cicadas mentioned above
emerge in areas containing a great variety of woody shrubs and treelets.
The census of 15 Vernonia in a field produced 53 nymphal casts of F.
pronoe (4.5 ±3.7 per tree), while eleven nymphal casts of Q. gigas were
also found. The census in the nutmeg grove produced 24 F. pronoe and 14
Q. gigas.

Discussion

The data indicate that different species of cicadas occurring in a lowland
tropical rain forest region occupy different habitats and have different peak
emergence periods each year. The primary forest contained the largest num-
ber of cicada species. Furthermore, the body size range of these species is
greater than for species in secondary habitats. The range in body sizes in
insect species might be an indicator of a larger number of ecological niches
available (Schoener and Janzen 1968). Secondary habitats tend to support
fewer cicada species and this was also the case for advanced secondary
forest regions in the mountains of central Costa Rica (Young 1975).

Other studies have shown that insects and plants exhibit seasonal re-
sponses to changes in rainfall, timing various activities with a particular
season (e.g., Janzen 1967; Schoener and Janzen 1968; Frankie et al. 1974).
This study and a previous one (Young 1972) show that different species of
cicadas may have allochronic emergences that result in breeding activities
(courtship, mating, oviposition) also being allochronic among the species.
The emergence of a greater number of cicada species in the wet season may
be a result of this period being longer than the dry season in this region.
The factors responsible for triggering cicada emergences in the tropics have
not been determined. Perhaps a drying out of the soil acts as a proximal cue
for the emergence of F. sericans, F. pronoe, and Q. gigas late in the dry
season. Increased rainfall and soil moisture content in the wet season may
be a cue for the emergence of Z. smaragdina, F. spinocosta, and others.
The two most abundant cicadas, Z. smaragdina and F. sericans emerged
from the same patches of primary forest and the former also emerged in
advanced secondary forest. The discovery that peak emergences in these
species are allochronic suggests a temporal partitioning of the environment
that results in breeding activities taking place at different times each year.
A similar pattern exists for cicadas such as F. spinocosta and F. pronoe in
young secondary forest habitats.

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NEW YORK ENTOMOLOGICAL SOCIETY

The emergence data also suggest that when two cicadas have similar
abundance levels in the same habitat, they tend to have highly allochronic
or seasonal emergences. But a species such as F. mannifera, with relatively
low emergence in a given month, although emerging in the same forest
patches as F. sericans and Z. smaragdina, is essentially active throughout
most of the year. Even though F. mannifera is active with two or more
congeneric species at the same time, it sings primarily at times of the day
when the others do not. The data also show that a similar daily partitioning
of daylight hours for singing takes place between two late dry season cicadas
in the same habitat, F. pronoe and Q. gigas.

Certain types of microhabitats within the primary and secondary forest
habitats may provide very suitable conditions for cicada development and
survival. Pentaclethra is one of the most abundant trees in lowland tropical
rain forests of Costa Rica (Hartshorn 1975) and the present study and a
previous one (Young 1972) have shown that cicadas often emerge near them.
Other suitable microhabitats for cicadas in this region are patches of Goe-
thalsia in advanced secondary forests and patches of Vernonia and Vismia
in young secondary forests. North American periodical cicadas have been
shown to have definite habitat and microhabitat associations (e.g., Dybas
and Lloyd 1962, 1974) as do some non-periodical species in the same region
(Moore 1966). The types of trees and shrubs present in an area may deter-
mine the availability of suitable oviposition sites and nymph development
sites for cicadas in tropical habitats. The observed large differences in num-
ber of nymphal casts collected in different quadrats in the same area also
suggest environmental heterogeneity affecting cicada oviposition and/or de-
velopment within the habitat.

Although the proximal factors determining the observed patterns of en-
vironmental partitioning by cicadas were not investigated, such patterns
could be explained by selection pressures resulting from a limited avail-
ability of resources. Such selection pressures would involve the determi-
nation of physiological and behavioral responses of mature cicada nymphs
to changes in environmental factors indicating season and act to time the
peak emergence with either the wet or dry season. Allee et al. (1949), among
others, emphasize the role of limited resources in resulting in selection pres-
sures expressed through interspecies competition. If, for example, in the
present study, suitable singing, feeding, and/or oviposition sites are limited
resources for cicadas in tropical habitats, selection may result in allochronic
emergence patterns among some species occupying the same habitats, or
cause divergence in habitat occupancy among species active during the same
time of the year. An additional factor might be the continued cutting down
of primary forest, favoring those cicadas that occupy secondary forests.

VOLUME LXXXVIII, NUMBER 2

101

Acknowledgments

This research was funded by National Science Foundation grant GB-
33060. Field assistance was given by W. Albright, D. Ewart, R. Kimber,
K. Kinsell, K. Richter, J. Thomason, and M. J. Young. Dr. J. Robert Hunter
gave me permission to work at “Finca La Tirimbina.” Dr. Dieter C. Wass-
hausen (Smithsonian) identified the tree species and Dr. Thomas E. Moore
(University of Michigan) gave generous assistance with identification of ci-
cadas. I thank Greg Raab of the Milwaukee Public Museum for the prepa-
ration of figures.

Literature Cited

Allee, W. C., A. E. Emerson, O. Park, T. Park and K. P. Schmidt. 1949. Principles of Animal
Ecology. Philadelphia: W. B. Saunders Co., 837 pp.

Dybas, H. S. and M. Lloyd. 1962. Isolation by habitat in two synchronized species of peri-
odical cicadas (Homoptera: Cicadidae: Magicicada). Ecology 43:444-459.

and . 1974. The habitats of 17-year periodical cicadas (Homoptera: Cicadidae:

Magicicada spp.). Ecol. Monogr. 44:279-324.

Frankie, G. W., H. G. Baker and P. A. Opler. 1974. Comparative phenological studies of
trees in tropical wet and dry forests in the lowlands of Costa Rica. J. Ecol. 62:881-919.

Hartshorn, G. S. 1975. A matrix model of tree population dynamics. In: Tropical Ecological
Systems, F. B. Golley and E. Medina, eds., pp. 41-51, New York: Springer Verlag.

Janzen, D. H. 1967. Synchronization of sexual reproduction of trees within the dry season in
Central America. Evolution 21:620-637.

Kato, M. 1956. The biology of cicadas. Bull. Cicada Mus. Tokyo, 319 pp.

Moore, T. E. 1966. The cicadas of Michigan (Homoptera: Cicadidae). Papers Mich. Acad.
Sci., Arts, and Letters 60:75-96.

Schoener, T. W. and D. H. Janzen. 1968. Notes on environmental determinants of tropical
versus temperate insect size patterns. Amer. Natur. 102:207-224.

Slud, P. 1964. The birds of Costa Rica: distribution and ecology. Bull. Amer. Mus. Nat. Hist.
128.

Young, A. M. 1972. Cicada ecology in a Costa Rican tropical rain forest. Biotropica 4:152-
159.

. 1975. The population biology of neotropical cicadas. I. Emergences of Procollina and

Carineta in a mountain forest. Biotropica 7:248-258.

Invertebrate Division, Milwaukee Public Museum, Milwaukee, Wisconsin
53233.

Received for publication April 26, 1979.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(2), 1980, pp. 102-105

SIMULTANEOUS USE OF A FORAGING TRAIL BY TWO
LEAFCUTTER ANT SPECIES IN THE
SONORAN DESERT

Alex Mintzer

Abstract. — An episode of simultaneous utilization of a foraging trail by
two attine ants, Acromyrmex versicolor (Pergande) and Atta mexicana (F.
Smith), is described. No aggressive behavior between the two species was
noted, except near an Acromyrmex nest entrance. This observation of si-
multaneous trail use is of particular interest because both species have sim-
ilar habits and they overlap in selection of food plant species in the Sonoran
Desert.

Acromyrmex versicolor (Pergande) is a common leafcutting ant in the
desert regions of Arizona and Sonora. The natural history and foraging
behavior of this ant has been described by Wheeler (1907) and Gamboa
(1975, 1976). The leafcutting ant Atta mexicana (F. Smith) ranges from
extreme southern Arizona through most of Mexico (Smith 1963). Both
species culture fungi and they occur at a study site 46 km south of Sonoita,
Sonora, Mexico. The foraging behavior of Atta mexicana at this site has
been described (Mintzer 1979).

Both ant species cut material from annual and perennial plants and collect
dry vegetation. Atta mexicana and Acromyrmex versicolor exhibit a broad
overlap in their selection of target plant species. They cut pieces from the
winter annual Plantago insularis Eastwood when it is available, and utilize
fresh and dry material from the common large desert perennials Larrea
tridentata Coville, Olneya tesota Gray, and Prosopis velutina Woot. Their
temporal periods of foraging activity are very similar. Both species forage
during the day in winter and spring (1974-77; 24 days total observation
period).

These mycophagous ants use trails as a regular component of foraging
activity. On March 25, 1977, simultaneous use of a 6-8 m trail segment by
these two species was noted by the author and field assistant Barry Pullen.
Both ant species were collecting leaf pieces from creosote bush, Larrea
tridentata , and Plantago insularis. The trail extended along the side of an
arroyo channel. The slope was rough but generally free of debris, and the
ants made no effort to clear or level the trail surface. The trail was 6-10 cm
wide; trails of Atta mexicana are usually 3-5 cm across (unpub. observa-
tion), while those of Acromyrmex versicolor are up to 14 cm in width (Gam-
boa 1975). One end of the trail terminated at a nest opening of the Aero-

VOLUME LXXXVIII, NUMBER 2

103

Figs. 1, 2. 1. Section of trail used by Atta mexicana and Acromyrmex versicolor. Lower

left: Acromyrmex versicolor forager with cut item from Plantago insularis. Upper right: Atta
mexicana forager with cut leaf from Larrea tridentata. 2. Another section of the trail. From
lower left to upper right: Acromyrmex versicolor ; Atta mexicana ; A. mexicana (submajor with
forage item); Acromyrmex versicolor ; A. versicolor (with forage item). Atta mexicana foragers
have longer legs than the Acromyrmex versicolor foragers of comparable size.

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NEW YORK ENTOMOLOGICAL SOCIETY

myrmex colony, which was surrounded by a pile of cut pieces from L.
trident ata. Acromyrmex workers were returning to this nest entrance with
forage items during the period of observation, while Atta workers with
forage items traveled in the opposite direction, towards their own tunnel
opening (see figures). Acromyrmex workers attacked and chased Atta for-
agers they encountered on this pile or near the nest entrance. B. Pullen
observed six dead or injured Acromyrmex and one dead Atta worker near
this entrance. Most Atta workers left the trail about 60 cm from the Acro-
myrmex nest entrance and foraged individually, returning to the trail after
securing leaf pieces. Foragers of both species were well represented when
the trail was first noted around 1300 hr local time. Seventy Acromyrmex
and 66 Atta workers passed a fixed point on the trail during a five minute
period. No aggressive interactions were noted on the trail, except within 25
cm of the Acromyrmex nest opening. However, Atta became less abundant
on the trail later in the afternoon.

The Atta colony involved in this episode produced up to 30 trails simul-
taneously, and at least two Acromyrmex versicolor colonies were located
near the margins of its foraging area. The one episode of simultaneous use
of a trail occurred shortly after a period of rain, when Acromyrmex was
very active on the surface. The Atta foragers may have initially been at-
tracted to the accumulation of L. trident at a leaf pieces around the Acro-
myrmex nest entrance. From my observations, it was impossible to deter-
mine whether both species are capable of following each other’s trails, or
which ant produced the initial trail.

In the laboratory, some attine species follow trails produced using ab-
dominal gland extracts from other attine species and genera. In Atta texana
(Buckley), the poison gland in the abdomen is the source of the primary
trail pheromone (Blum et al. 1964; Moser 1967). Moser notes: Tn the field,
however, although the trails of various species cross, the workers generally
find their own trails easily.” For a more typical account of relations between
foragers of two Atta species, check Weber (1969). At the study site in
Mexico, Acromyrmex was active on only a few observation days, and trails
of the two species were not observed near each other on other occasions
when simultaneous foraging activity occurred.

Wilson (1965) describes a case of simultaneous trail use by two ants on
Trinidad, and reviews two cases discovered by W. M. Wheeler in neotrop-
ical forest. In two of these cases involving Crematogaster limata parabi-
otica Forel [with Camponotus femoratus (Fabricius) and Monads debilis
(Emery)], some of the shared trails led to food resources such as membra-
cids used by both ants involved, and no aggressive interactions were noted.
In the third case, Camponotus beebei Wheeler utilized trails of Azteca
chartifex Forel, and some aggressive interactions were noted on the trails.

VOLUME LXXXVIII, NUMBER 2

105

In all of these reports, the two ant species involved belong to different
subfamilies (Myrmicinae, Dolichoderinae, or Formicinae). In contrast, Atta
and Acromyrmex are closely related myrmicine genera (Weber 1972). Si-
multaneous use of a foraging trail is particularly interesting in this case
because the species involved are members of a smaller and more recently
evolved phyletic group with similar and unique dietary habits, and they may
be expected to compete for resources where they occur together.

Acknowledgments

I thank Barry Pullen for his assistance at the study site, and for reviewing
the manuscript. I also thank T. E. Moore, D. Bay and W. D. Hamilton for
assistance and comments during preparation of this manuscript. Field study
of Atta mexicana was supported by a travel grant from the Rackham School
of Graduate Studies of the University of Michigan.

Literature Cited

Blum, M. S., J. C. Moser and A. D. Cordero. 1964. Chemical releasers of social behavior II.

Source and specificity of the odor trail substances in four attine genera. Psyche 71:1-7.
Gamboa, G. 1975. Foraging and leaf-cutting of the desert gardening ant Acromyrmex versi-
color (Pergande). Oecologia 20:103-110.

. 1976. Effects of temperature on the surface activity of the desert leafcutter ant Ac-
romyrmex versicolor. Amer. Midi. Nat. 95:485-491.

Mintzer, A. 1979. Foraging activity of the Mexican leaf-cutting ant Atta mexicana (F. Smith)
in a Sonoran Desert habitat. Insectes Sociaux 26:364-372.

Moser, J. C. 1967. Trails of the leafcutters. Natural History 76, no. 1.

Smith, M. R. 1963. Notes on the leafcutting ants, Atta spp., of the United States and Mexico.
Proc. Ent. Soc. Wash. 65:299-302.

Weber, N. A. 1969. Ecological relations of three Atta species in Panama. Ecology 50:141-
147.

. 1972. Gardening Ants, the Attines, Mem. Amer. Philo. Soc. 92.

Wheeler, W. M. 1907. The fungus-growing ants of North America. Bull. Amer. Mus. Nat.
His. 23.

Wilson, E. O. 1965. Trail sharing in ants. Psyche 72:2-7.

Museum of Zoology and Division of Biological Sciences, The University
of Michigan, Ann Arbor, Michigan 48109.

Received for publication November 6, 1979.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(2), 1980, pp. 106-112

CAUSES OF LIFE HISTORY DIFFERENCES BETWEEN THE
MORPHS OF PEMPHIGUS POPULITRANSVERSUS 1

Lorenz Rhomberg

Abstract. — The aphid Pemphigus populitransversus produces galls of two
distinct morphs on cottonwood petioles. There are pronounced between-
morph differences in both the number and maturity distribution of aphids
within the galls. The progress of gall maturation was followed over the
summer to determine whether these life history differences are explainable
solely by the fact that galls of the “globular” morph are initiated some seven
weeks later than those of the “elongate” morph or whether qualitative dif-
ferences exist in the pattern of gall maturation. It was found that the pattern
of maturation for the two morphs was similar, with globulars being always
seven weeks behind. However, the number of nymphs and mature alates
was about three times greater in globular galls.

Introduction

Pemphigus populitransversus Riley is a gall-forming aphid attacking
leaves of cottonwoods, principally Populus deltoides Marshall and P. sar-
gentii Dode. A nymphal stem mother hatches from a sexually produced egg
in the spring and initiates a gall on a cottonwood petiole. The growing petiole
tissue quickly encloses her in a capsule within which she parthenogeneti-
cally and viviparously produces nymphal fundatrigeniae which, as winged
adults, leave the gall and fly to secondary hosts of the family Cruciferae.

In the study of a dimorphism of galls in this aphid Senner and Sokal (1974)
found large differences in the numerical distribution of developmental stages
in the two morphs. In fact, they noted that “the two samples can be differ-
entiated on life history characteristics more clearly than on gall shape, the
initial criterion for separation [into two morphs]” (p. 368). Galls of the
“globular” morph contained fewer nymphs, most (98.1%) of which were in
the smallest of three size classes. Galls of the “elongate” morph had about
half their nymphs in the two largest size classes and had more alates (adult
fundatrigeniae). Senner and Sokal hypothesized that “nymphs in the ‘glob-
ular’ galls . . . , although necessarily born sequentially, develop synchro-
nously and ... the bulk of alate production must be in the late fall. In the

1 This paper is publication number 347 from the Graduate Program in Ecology and Evolution
at the State University of New York at Stony Brook, Stony Brook, N.Y. 11794.

VOLUME LXXXVIII, NUMBER 2

107

‘elongate’ sample . . . the nymphs seem to develop sequentially throughout
the summer with alates presumably being produced continuously” (p. 368).

Subsequently, Faith (1979a) discovered that the globular-elongate dimor-
phism was based on different times of gall initiation. Elongate galls are
initiated in early May by stem mothers hatching from overwintering eggs.
They are formed on the early flush of leaves (which are pre-formed in the
bud) soon after bud-break. Globular galls are founded by stem mothers
hatching some six weeks later from eggs that are laid in the spring. These
galls are on the second, morphologically distinct, flush of leaves (Faith
1979a). The ranges of the two morphs seem to overlap essentially completely
(Bird et al. 1979).

Faith (1979b) proposed that this difference in time of gall initiation pro-
vides a simple alternative to Senner and Sokal’s explanation of the life
history differences between the morphs. He showed that globular galls pro-
duced their first alates during the first days of August — again six weeks later
than elongate galls attain this milestone. Perhaps globular galls are simply
less mature, and at the time of collection of Senner and Sokal’s original
dimorphic sample (August 5, 1966 in West Point, Ga.) even the oldest
nymphs were still small. The differences in the mean and variance of the
distribution of life history stages should then disappear later on in the sea-
son.

It is the purpose of this paper to show that differences in gall age between
the morphs can indeed account for the life history dimorphism discovered
by Senner and Sokal. There is some evidence, however, that some devel-
opmental synchrony may exist in globular galls, but on a scale much smaller
than Senner and Sokal originally envisaged.

Materials and Methods

Galls of Pemphigus populitransversus were collected at roughly weekly
intervals from a midsized (approximately 15 cm dbh) eastern cottonwood
tree {Populus deltoides ) in Port Jefferson, L.I., N.Y. Five elongate galls
were collected each time beginning on 13 June 1975 and ending on 7 Sep-
tember 1975. Five (occasionally six) globular galls were collected each time
beginning on 31 July 1975, and ending on 4 October 1975. Galls could be
assigned to one or the other morph unambiguously by noting whether they
were on early or late leaves. Galls were immediately preserved in 70%
ethanol. Later, each gall was opened and censused for the number of small,
medium, and large sized nymphs, as well as the number of alates. Proce-
dures were those of Senner and Sokal (1974).

Abbreviations and definitions of variables are listed in Table 1. Several
empty or parasitized galls were eliminated from further study. Since Senner
and Sokal coded the numbers of nymphs in each size category into 10 abun-

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NEW YORK ENTOMOLOGICAL SOCIETY

Table 1. Variables used in this study.

Abbreviation

Definition

# SM NYM

The number of “small” nymphs in the gall, roughly those in the first
or second instar.

# MD NYM

The number of “medium” nymphs in the gall, roughly those in the
third, or penultimate, instar.

# LG NYM

The number of “large” nymphs in the gall, roughly those in the
fourth, or ultimate, instar.

# ALATES

The number of adult fundatrigeniae in the gall.

G LH MN

By coding small, medium and large nymphs as 1, 2 and 3 respectively,
the mean nymphal stage attained by all nymphs in the gall. Coded
abundances of each stage were used.1

G LH VAR

By coding small, medium and large nymphs as 1, 2 and 3 respectively,
the variance in nymphal stage attained by all nymphs in the gall.
Coded abundances of each stage were used.1

1 Abundances were coded into 10 classes following Senner and Sokal (1974): 0: 0 nymphs,
1: 1-10 nymphs, 2: 11-20, 3: 21-30, 4: 31-40, 5: 41-50, 6: 51-60, 7: 61-70, 8: 71-80, 9: 81 and
greater.

dance classes, the mean developmental stage and its variance (G LH MN
and G LH VAR respectively) were calculated for each gall using their coding
scheme. I have reported the nymph abundances themselves as uncoded
counts, however.

The changes in variance in developmental stage attained (G LH VAR)
over the season are plotted in Figure 1. During June the variance for elon-
gate galls shows a sharp rise as the first nymphs mature. By July 1 a max-
imum variance (and mean) of the distribution is attained, and remains at
that level until September (at which time the supply of elongate galls was
exhausted). Alates are continually leaving the gall during this time, so the
stable mean and variance represent a steady state distribution of develop-
mental stages.

Globular galls progress along an essentially identical curve, save that it
is displaced some 50 days later in the season. The initial rise is about as
rapid as for elongates, and the same steady state is maintained from then
on, at just the same level, until leaves drop in the autumn.

Figure 2 shows the progress of gall population size over the season, with
each bar divided according to the numbers present for each developmental
stage. The maximum population size of globulars is much greater than that
of elongates. This maximum is attained more quickly (and lost more quickly)

Results

VOLUME LXXXVIII, NUMBER 2

109

.8 T

\ 1 b

JUNE

» ‘ * —

JULY

1 1 h

AUG

SEPT

OCT

160

200

240

DAY

280

Fig. 1. Changes in life history variance (the variance of the distribution of maturities) in
elongate galls (solid points) and globular galls (open points). Vertical bars indicate ± one
standard error of the mean. Sample sizes may be obtained from Figure 2. Abscissa is number
of days since January 1, 1975.

than in elongates. It should be noted that at the end of the initial, essentially
“all small nymph” stage, globular galls have already attained about one
third of their eventual peak population, whereas elongates have only
reached about one seventh of their maximum at the analogous time. Fur-
thermore, the initial phase of increase in life history variance is accompanied
by rapid population size increase in elongates, but not so in globulars. In-
stead, the rapid increase in population just follows the attainment of the
stable distribution of life history stages.

Discussion

At any given time until mid-August galls belonging to the globular morph
will indeed have a very different distribution of developmental stages than
those of the elongate morph, as noted by Senner and Sokal. These differ-
ences seem to be a consequence of the different ages of galls of the two
morphs at any one time. They vanish in mid- August as globular galls “catch

MEAN GALL POPULATION

no

NEW YORK ENTOMOLOGICAL SOCIETY

400 T

200 ■’

Elongate

4

3 a

JUNE

JULY

AUG

SEPT

oct]

800

600

400 •

200 •

160 2 00

Globular

240

5

I

280

KEY

alates

large nymphs
medium nymphs
small nymphs

4

i

JUNE

JULY

AUG

SEPT

OCT

160

200

240

280

DAY

Fig. 2. Changes in mean gall population size for elongate (top) and globular (bottom) galls.
Each bar is divided into segments proportional to the numbers of small, medium, and large
nymphs, and alates. The sample sizes are shown above each bar. Abscissa is the same as
Figure 1 .

VOLUME LXXXVIII, NUMBER 2

111

up” in maturity. The changes in the variance of the developmental stage
distribution are identical in pattern. The approximately 50 day displacement
of the curve for globular galls almost exactly matches the 47 day difference
in time of gall initiation on Long Island (average of 1976, 1977 and 1978
[Faith 1979b]). The duration of the phase of low life history variance (when
almost all nymphs are in the small size class) for globular galls is brief, but
it is just around this time that numerous samples considered by Senner and
Sokal were collected.

A more extensive investigation of the full data base of which their col-
lections were a subset shows a seasonal pattern of life history changes very
much like the one here described (Bird et al. 1979). In fact, knowledge of
this pattern has enabled these authors to allocate galls of unknown status
to one or the other morph.

In contrast to the pattern of development, the number of nymphs pro-
duced is very different between morphs. Peak nymph population sizes differ
(t. = 3.23 , P < 0.02) with globulars reaching levels over three times as high
as elongates. The seasonal progression in globular gall population size is not
simply displaced from that of elongate galls. Because of the higher popu-
lation size attained by globular galls, the numbers of small nymphs at the
time of Senner and Sokal’ s collections are about equal to the total popu-
lations of the more mature elongate galls, giving the impression that small
globular nymphs have accumulated without developing further.

A closer look at Figures 1 and 2 reveals some support for a much smaller
scaled version of the “developmental synchrony” hypothesis in globular
galls, spanning some two weeks in August instead of the whole season. In
these galls the phase of rapid increase in life history variance happens with-
out an appreciable increase in the number of nymphs in the gall, as though
a large initial cohort begins to mature before many new small nymphs are
produced. Elongate galls, in contrast, show a gradual rise in number of
nymphs during their phase of life history variance increase.

In conclusion, an investigation of the progress of maturation of globular
and elongate galls of Pemphigus populitransversus shows that the between-
morph differences in maturity distribution of the nymphs can be ascribed
largely to their different ages at any point in time. A large scale qualitative
difference in the progress of maturation, such as that hypothesized by Sen-
ner and Sokal, is not found. However, some differences independent of gall
age are found; these are peak population size and its rate of attainment.

Whether the life history differences between morphs can be considered
to be adaptations to different circumstances is difficult to say. Certainly the
existence of the globular morph enables exploitation of a new resource,
namely the late flush of leaves, which seems to be unexploited by other
Pemphigus species. Stem mothers seem to have no limit to the numbers of

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NEW YORK ENTOMOLOGICAL SOCIETY

nymphs they can produce. Unborn nymphs may still be found in stem moth-
ers taken from galls on abscised leaves in the autumn. However, the fecun-
dity of stem mothers should be influenced by genetic and environmental
factors. The greater fecundity and consequent higher population size of
globular galls may be a phenotypic response to maturation in the warmer
climate of mid-summer or to nutritive or anatomical differences between
the early and late flushes of leaves. Faith (1979a) has shown strong asso-
ciation of population size with measurements of the leaves both within and
between morphs.

Acknowledgments

This study was supported by grant number BMS 720221 1A04 from the
National Science Foundation to Robert R. Sokal. The manuscript was read
by D. P. Faith, B. Riska, and R. R. Sokal.

Literature Cited

Bird, J., D. P. Faith, L. Rhomberg, B. Riska and R. R. Sokal. 1979. The morphs of Pemphigus
populitransversus : Allocation methods, morphometries, and distribution patterns. Ann.
Entomol. Soc. Am. 72:767-774.

Faith, D. P. 1979a. Strategies of gall formation in Pemphigus aphids. J. New York Entomol.
Soc. 87( 1):2 1—37.

. 1979b. A study of a dimorphism in the life history of the gall-forming aphid Pemphigus

populitransversus. Ph.D. Thesis. State University of New York at Stony Brook, Stony
Brook, New York.

Senner, J. W. and R. R. Sokal. 1974. Analysis of dimorphism in natural populations with
consideration of methodological and epistemological problems. Syst. Zool. 23:363-386.

Department of Ecology and Evolution, S.U.N.Y. at Stony Brook, Stony
Brook, New York 11794.

Received for publication October 8, 1979.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(2), 1980, pp. 113-114

EPISTYLUS CAMBARI (CILIATA: PERITRICHIDA) AND
DRAGONFLY NYMPHS, AN EPIZOIC ASSOCIATION

Reid Jilek

Abstract. — Epistylis cambari was found attached to the dorsal abdominal
surface of 30% of the dragonfly nymphs examined. Colonies ranged from
20-30 individuals with but one colony attached to each dragonfly nymph.

Introduction

Members of the genus Epistylus characteristically form epizoic associa-
tions (Sleigh 1975). Associations involving Epistylus have been recorded
from crayfish (Kudo 1954), chironomid larvae (Jahn 1949), fishes (Crites
1977; Rogers 1971), and turtles (Bishop and Jahn 1941). This study has
revealed the presence of Epistylus cambari from dragonfly nymphs.

Materials and Methods

A total of 30 dragonfly nymphs was collected from ponds 47, 48, 49, and
50 on the Delaware Reservoir Wildlife Area, Delaware County, Ohio, during
April 1979. The nymphs were identified as Gomphus quadricolor Walsh
(family Gomphidae) and Pachydiplax longipermis Burmeister (family Li-
bellulidae).

The nymphs were maintained for one week in individual petri dishes filled
with distilled water. After one week the nymphs were examined.

Identification of Epistylis cambari to species was performed following
removal from the dragonfly nymphs. E. cambari were stained with Harris
haematoxylin and Rose Bengal and examined by light microscopy.

Results

Nine of the 30 dragonfly nymphs (6 of 12 G. quadricolor and 3 of 18 P.
longipennis) were found to have single colonies of Epistylis cambari at-
tached to their dorsal abdominal surfaces. Each colony was comprised of
20-30 individuals. No evidence of pathology was observed at the site of
attachment of E. cambari.

Discussion

Dragonfly nymphs provide a suitable environment for Epistylis. The rath-
er sedentary existence of the nymphs allows for attachment of telotrochs
and subsequent development into a colonial form. This epizoic association

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NEW YORK ENTOMOLOGICAL SOCIETY

is further enhanced by the hardened sclerotized abdominal plates of the
dragonfly nymphs.

The presence of E. cambari attached to dragonfly nymphs represents a
new host record. The presence is, however, not inexplicable since dragonfly
nymphs commonly co-inhabit areas with crayfish, a known host of E. cam-
bari and the ponds are inhabited by numerous crayfish.

Literature Cited

Bishop, E. L. and T. L. Jahn. 1941. Observations on colonial peritrichs (Ciliata: Protozoa)
of the Okoboji Region. Proc. Iowa Acad. Sci. 48:417-421.

Crites, J. L. 1977. The occurrence of Epistylis niagarae (Ciliophora: Peritrichida) on fishes
from the island region of western Lake Erie. Ohio J. Sci. 77:193-194.

Jahn, T. L. 1949. How to Know the Protozoa. Wm. C. Brown Co. Dubuque, Iowa. 234 pp.
Kudo, R. R. 1954. Protozoology. 4th Ed. Charles C Thomas, Springfield, 111. 966 pp.

Rogers, W. A. 1971. Disease in fish due to the protozoan Epistylis (Ciliata: Peritricha) in the
Southeastern U.S. Proc. 25th Ann. Conf. Southeastern Assoc. Game and Fish Comm.
25:493-496.

Sleigh, M. 1975. The Biology of Protozoa. Edward Arnold Limited, 25 Hill Street, London,
London W1X 8LL. 315 pp.

Department of Zoology and The Center for Lake Erie Area Research,
The Ohio State University, Columbus, Ohio 43210.

Received for publication November 19, 1979.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(2), 1980, pp. 115-119

TOTAL CONCENTRATION OF FREE AMINO ACIDS AND THE
CORRESPONDING MORPHOGENETIC CHANGES DURING
THE EMBRYOGENESIS OF CHILOMENESE SEXMACULATA
FABR. (COLEOPTERA: COCCINELLIDAE)

Devinder Mukesh, Dalbinder Singh Sidhu1 and Nirmal Kumar

Abstract. — The concentration of free amino acid (FAA) pool in relation
to the various morphogenetic events occurring during embryogenesis of
Chilomenes sexmaculata Fabr. was determined at different incubation pe-
riods. During the first 12 hours of embryogenesis, when the zygote under-
goes an active cleavage, the concentration declines gradually. However, it
rises sharply at 15 hours of development when the cleavage energids
undergo an active migratory activity. The amount of FAA decreases at 24
hours and remains almost steady up to 35 hours of incubation when the
embryo starts elongation and segmentation. From then on there is a contin-
uous increase in FAA up to 75 hours followed by a declining trend towards
hatching as the yolk gets depleted.

Introduction

Extensive reviews on the biochemical investigations pertaining to the ni-
trogen metabolism during embryogenesis of insects have earlier been made
by Chen (1962, 1966) and Corrigan (1970). To get a greater insight into the
development of an embryo it is necessary to elucidate the relationship of
biochemical changes to the corresponding morphogenetic events occurring
during this period. Since the major processes underlying differentiation at
the embryonic level aim at protein synthesis which in turn depends on the
supply of free amino acids (FAA) from the yolk reserves, FAA thus have
a distinct relationship to morphogenesis.

The present attempt unveils a relationship between the total concentration
of FAA and the corresponding morphogenetic events occurring during the
different incubation periods in Chilomenes sexmaculata Fabr.

Materials and Methods

The adults of C. sexmaculata were collected from the Calotrips procera
plants during the months of November to February around Patiala (India)
and were reared in the laboratory according to Sarswat (1976) on Aphis
neri. Numerous batches of eggs were obtained, incubated at 28 ± 2°C for

All correspondence to this author.

116

NEW YORK ENTOMOLOGICAL SOCIETY

Table 1. Concentration of FAA pool and various morphogenetic events occurring during
embryogenesis of Chilomenes sexmaculata Fabr.

Incubation
period (hr)

/x M FAA/ 100 eggs
(±SD)*

Morphogenetic event

0.25

1.51 ± 0.030

Maturation of the male as well as female pronucleus
(Fig. 1).

1.00

1.48 ± 0.029

Fusion of male and female nucleus (Fig. 2).

3.00

1.34 ± 0.027

Cleavage of the zygote, giving rise to 4-energids stage
(Fig. 3).

5.00

1.46 ± 0.029

8-cleavage energids stage (Fig. 4).

7.50

1.16 ± 0.023

The process of cleavage continues.

12.00

1.11 ± 0.022

240 ± 16 energids stage (Fig. 5).

15.00

1.77 ± 0.035

Migration of the cleavage energids towards the
periplasm starts (Fig. 6).

19.00

1.68 ± 0.035

A syncytial primary epithelium is formed (Fig. 7).

24.00

1.37 ± 0.028

A regular layer of primary epithelial cells observed (Fig.
8).

31.00

1.39 ± 0.028

The ventro-posterior portion of the primary epithelium
becomes thickened, the cells being columnar, so as to
give rise to the definitive germ primordium (ventral
plate) (Fig. 9).

35.00

1.32 ± 0.026

Elongation of the germ band or ventral plate takes place
and the germ band becoming faintly distinguishable
into an anterior protocephalon and the posterior
protocorm (Fig. 10).

45.00

1.42 ± 0.028

Further elongation of the germ band resulting in the
formation of three protocormic regions (Fig. 11).

60.00

1.45 ± 0.029

The germ band becomes segmented with delineated
appendages (Fig. 12).

72.00

1.48 ± 0.029

75.00

1.55 ± 0.030

An active growth period, the appendages further
develop and the process of cephalization starts,
secondary yolk cleavage, yolk poly-hedra formed.

90.00

1.35 ± 0.026

The appendages become fully formed and the head also
becomes distinct with the usual appendages (Fig. 13).

100.00

1.30 ± 0.026

Fully formed embryo.

110.00

1 .22 ± 0.024

Hatching takes place.

* Standard deviation. Each reading is an average of three determinations.

Fig. 1. Abbreviations: CN, cleavage energid; FN, female pronucleus; G, germ band; GA,
gnathal appendages; LR, labral appendages; MN, male pronucleus; PC, pole cell; PE, primary
epithelium; PM, protocorm; 1PM, 2PM, 3PM, 1st, 2nd and 3rd protocormic regions; PMB,
premandibular appendages; PN, protocephalon; TA, thoracic appendages; YC, yolk cell.

definite periods and subsequently fixed in Carl’s fixative at 60°C for 2 hours.
The permanent preparations were made according to the method of Rempel
and Church (1971) to observe various developmental changes in the embryo.

The FAA extracts from the various egg samples at specific incubation
periods were prepared according to Sidhu and Kang (1979). The quantitation
of FAA pool was made by the method of Troll and Cannon (1953).

Results and Discussion

The concentration of FAA pool and the various morphogenetic events
occurring during the embryogenesis of Chilomenes sexmaculata Fabr. are
recorded in Table 1.

At 0 hour, i.e., immediately after oviposition, the concentration was
1.51 ± 0.030 /xmoles/100 eggs, which decreased three hours later to 1.34 ±
0.027 /amoles/100 eggs. This decrease in FAA concentration corresponds to
the period when the exochorion of the newly laid egg becomes hard and

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NEW YORK ENTOMOLOGICAL SOCIETY

dark in color, apparently indicating the utilization of FAA in this process.
It has further been noted that there is a continuous decrease in FAA con-
centration up to 12 hours of incubation, i.e., during the active cleavage
activity when the zygote nucleus undergoes multiple synchronous divisions.
This decrease suggests that the breakdown of yolk proteins has not yet
started and the need of amino acids for protein synthesis is met from the
already existing FAA pool accumulated during maturation of egg. The ini-
tiation of protein synthesis immediately after fertilization has earlier been
reported by Lockshin (1966) in some coleopterous insects.

At about 15 hours of incubation, when energids start migrating, the con-
centration of FAA suddenly increases to a fairly high value (1.77 ± 0.035
/amoles/100 egg). This increase is due perhaps to the lysis of yolk proteins,
with the result that the mixing-up of the yolk contents facilitates the pe-
ripheral migration of the cleavage energids. This type of increase in FAA
concentration has been observed by Chen and Briegel (1965) in Culex pipe ns
morestus at a stage when blastoderm formation and elongation of germ band
(Idris 1960) takes place.

The FAA concentration sharply decreases (1.37 ± 0.28 /amoles/lOO eggs)
at 24 hours of incubation when a regular primary epithelial layer gets estab-
lished, revealing a higher rate of protein synthesis in comparison with the
breakdown of yolk proteins. Thereafter the concentration remains almost
steady up to 34 hours of development. During this period the differentiation
of the ventral plate or the germ band is completed.

At 45 hours of development the amount of FAA again rises (1.42 ± 0.028
/nmoles/ 100 eggs), when the germ band undergoes further elongation and
primary segmentation. This trend in FAA concentration increase was ob-
served in the following 30 hours of development. The peak is reached at 75
hours of incubation when the embryo shows an active development. This
increase in concentration can be attributed to an active lysis of the yolk
proteins, corresponding to an active growth period when cephalization as
well as the phenomenon of secondary yolk cleavage takes place.

After this period, the FAA concentration shows a continuous decline
when the appendages become fully formed and the head becomes quite
distinct. This decline in concentration prior to hatching is due perhaps to
the fact that the yolk reserves become increasingly exhausted at this stage
and the supply of amino acids can no longer keep pace with the demand for
more amino acids for protein synthesis. Chen (1966) and Indira (1963) have
also observed a similar type of decrease in FAA concentration towards
hatching.

VOLUME LXXXVIII, NUMBER 2

119

Literature Cited

Chen, P. S. 1962. Free amino acids in insects; in Holden: Amino Acids Pools, pp. 115-138.
Elsevier, Amsterdam.

. 1966. Amino acid and protein metabolism in insect development. Adv. Insect Physiol.

3:53-132.

and H. Briegel. 1965. Studies on protein metabolism of Culex pipiens L. V. Changes

in free amino acids and peptides during embryonic development. Comp. Biochem. Phys-
iol. 14:463-473.

Corrigan, J. J. 1970. Nitrogen metabolism in insects; in Campbell: Comparative biochemistry
of nitrogen metabolism. I. The invertebrates, pp. 387—488. Academic Press, London.

Indira, T. 1963. Biochemical and cytological studies during development and ovarian growth
in Sphaeroderma malestum (Duf.). Ph.D. Thesis, Annamalai University, South India.

Idris, B. E. M. 1960. Die Entwicklung im novmalen Ei von Culex pipiens (Diptera). Z. Morph.
Okol. Tiere 49:387-429.

Lockshin, R. A. 1966. Insect embryogenesis: Macromolecular synthesis during early devel-
opment. Science 154:775-776.

Rempel, J. G. and N. S. Church. 1971. Embryology of Lytta viridana LeConte (Coleoptera:
Meloidae). Ill Appendiculate 72h embryo. Can. J. Zool. 49:1563-1571.

Sarswat, G. G. 1976. Studies on the trends in consolidation of nervous system in Coleoptera.
Ph.D. thesis, Agra University, Agra (India).

Sidhu, D. S. and H. K. Kang. 1979. Metabolic reserves and pool size of free amino acids
during metamorphosis of Callosobruchus maculatus (F.). Entomon. 4(l):57-59.

Troll, W. and R. K. Cannon. 1953. A modified photometric ninhydrin method for the analysis
of amino and imino acids. J. Biol. Chem. 200:803.

Department of Zoology, Punjabi University, Patiala- 147002, India.
Received for publication December 13, 1979.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(2), 1980, pp. 120-132

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

L. H. Rolston, F. J. D. McDonald and Donald B. Thomas, Jr.

Abstract. — The Pentatomini of the Western Hemisphere is divided into
3 sections. Keys are provided to separate these sections and the genera of
one section, viz. those genera that have a median tubercle at the base of
the abdominal venter and a metasternal production apposing this tubercle.
Four new monotypic genera are diagnosed, each based on a new species:
Elanela hevera Rolston, n. sp., Glaucioides englemani Thomas, n. sp., No-
cheta adda Rolston n. sp. and Vidada dollingi Rolston n. sp. The genus
Phineus Stal is removed from this group and placed in Discocephalinae.

There is no conspectus of Western Hemisphere genera of Pentatomini
more recent than the work of Stal (1867). Stabs conspectus is now inade-
quate because of the many genera subsequently added to the tribe.

As a foundation for this conspectus we have previously provided keys
and diagnoses for the Western Hemisphere families of pentatomoids,
subfamilies of pentatomids and tribes of the nominate subfamily (Rolston
and McDonald 1979).

The present work divides the Pentatomini into three sections, which sep-
arate rather clearly with the exception of sexually dimorphic species of the
genus Dendrocoris, and provides a key to the genera of section 3. Genera
of this section have on the third abdominal sternite (second visible) a median
tubercle that is apposed by the posterior margin of the metasternum. The
metasternum is produced ventrad, at least posteriorly, and often interlocks
with the abdominal spine.

The genus Bathycoelia, most species of which are African, is included in
this section since 2 species of the genus were described from specimens
supposedly collected in South America (Jensen-Haarup 1931 and 1937).
These specimens were examined and they do indeed belong in Bathycoelia.
Although the presence of this genus in South America is not unthinkable,
its existence there needs confirmation.

The genus Phineus Stal, which has customarily been placed among the
genera of section 3, is transferred to Discocephalinae on the basis of rostral
and trichobothrial characters (Rolston and McDonald 1979).

Key to Sections of Pentatomini

1 . Abdominal venter bearing median tubercle or spine at base 2

VOLUME LXXXVIII, NUMBER 2

121

- Base of abdominal venter smooth medially, not clearly produced

Section 1

2. Metasternum projecting ventrad (at least between metacoxae) with
posterior margin in apposition to median tubercle on base of abdom-
inal venter (Fig. 23) Section 3

- Median tubercle or spine at base of abdominal venter free distally,
not in apposition to posterior margin of metasternum (Fig. 24)

Section 2

Key to the Western Hemisphere Genera of Pentatomini
Key to Genera of Pentatomini, Section 3

1. At least hind femora with two rows of spines on inferior surface

(Fig. 1); body depressed Placocoris Mayr

- Inferior surface of femora unarmed; body normally convex 2

2. Rostrum lying in deep mesial groove extending almost or entirely

length of abdominal venter Bathycoelia Amyot & Serville

- Abdominal venter not grooved mesially or only shallowly so ba-

sally 3

3. Tarsi 2-segmented Phcilaecus Stal

- Tarsi 3-segmented 4

4. Superior surface of at least mesothoracic femora prolonged distally

as small spine (Fig. 2) 5

- Superior surface of femora unarmed distally 7

5. First rostral segment lying entirely between bucculae; ostiolar ru-

gae reaching more than halfway from inner margin of ostioles to
lateral margin of metapleura (Fig. 3) Pallantia Stal

- First rostral segment projecting beyond bucculae; ostiolar rugae

reaching less than halfway from inner margin of ostioles to lateral
margin of metapleura (Fig. 4) 6

6. Juga surpassing tylus, acute apically; superior surface of tibiae sul-

cate Arvelius Spinola

- Juga not surpassing tylus; superior surface of tibiae rounded except

distally Taurocerus Amyot & Serville

7. Metasternum bifurcate anteriorly and extending onto swollen meso-

sternum (Fig. 5) 8

- Metasternum usually entire anteriorly, occasionally with pair of

carinae 10

8. Anterolateral margins of pronotum fimbriate; ostiolar rugae ex-

tending halfway from mesial margin of ostioles to lateral margin of
metapleura Neopharnus Van Duzee

- Anterolateral margins of pronotum entire; ostiolar rugae extending

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NEW YORK ENTOMOLOGICAL SOCIETY

about 3 fourths of distance from inner margin of ostioles to lateral
margin of metapleura 9

9. Rostrum extending well beyond abdominal sternite bearing tu-
bercle Pharnus Stal

- Rostrum extending little or not at all past metacoxae

Praepharnus Barber & Bruner

10. Mesosternal carina projecting cephalad between procoxae and be-
yond anterior limits of procoxae (Fig. 6) 11

- Mesosternal carina if projecting cephalad terminating between pro-
coxae 12

1 1 . Apex of scutellum emarginate; ostiolar rugae nearly reaching lateral

margin of metapleura Evoplitus Amyot & Serville

- Apex of scutellum entire; ostiolar rugae extending about 2 thirds

of distance from inner margin of ostiole to lateral margin of meta-
pleura Pseudevoplitus Ruckes

12. Ostiolar rugae reaching more than halfway from inner margin of

ostioles to lateral margin of metapleura 20

- Ostiolar rugae reaching less than halfway from inner margin of

ostioles to lateral margin of metapleura 13

13. Distal end of first rostral segment clearly projecting beyond buc-

culae (Figs. 7, 8) 14

- Distal end of first rostral segment not or scarcely projecting beyond

bucculae 18

14. Ratio of head width across eyes to length of head 10:8-10:10 15

- Ratio of head width across eyes to length of head 10:7 or less 16

15. Humeral angles conspicuously produced; rostrum reaching meso-

coxae Myota Spinola

- Humera angles little produced; rostrum reaching metacoxae

Vidada Rolston, new genus

16. Metasternum saddle-shaped, transversely depressed between meta-
coxae and mesocoxae (in part) Brachystethus Laporte

- Metasternal production nearly flat or obtusely carinate 17

17. Rostrum reaching mesocoxae; antennal segment 2 much shorter

than 3 Lopadusa Stal

- Rostrum reaching metacoxae; antennal segment 2 much longer

than 3 Elanela Rolston, new genus

18. Metasternal production nearly flat Serdia Stal

- Metasternal production sloping or arched 19

19. Metasternal production greatest posteriorly, sloping dorsad ante-
riorly, weakly carinate anteriorly Marghita Ruckes

- Metasternal production longitudinally arched, carinate for entire

length Stictochilus Bergroth

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123

20. Distal end of first antennal segment clearly surpassing apex of

head 21

- Distal end of first antennal segment not or scarcely surpassing apex

of head 24

21. Metasternum much more produced than mesosternum, often bul-
bous (Fig. 9) (in part) Brachystethus Laporte

- Metasternal production flattened or obtusely carinate, together

with abdominal spine and mesosternal carina forming nearly con-
tinuous profile (Fig. 10) 22

22. Metasternal production flattened; distal end of first rostral segment

surpassing bucculae 23

- Metasternal production obtusely carinate; distal end of first rostral

segment not or scarcely surpassing bucculae Paratibilis Ruckes

23. Mesosternal carina forming thin wedge between procoxae, pro-
jecting slightly onto prosternum (Fig. 1 1) Tibilis Stal

- Mesosternal carina little elevated, not projecting beyond mesoster-
num (in part) Janeirona Distant

24. Interocular distance more than twice width of one eye; juga well

separated apically 25

- Interocular distance less than twice width of one eye; juga contig-
uous or nearly so apically Nocheta Rolston, new genus

25. Distal end of first rostral segment clearly exceeding bucculae 26

- First rostral segment lying entirely between bucculae 27

26. Apex of rostrum extending to or beyond posterior margin of third

sternite; juga no longer than tylus Pharypia Stal

- Apex of rostrum not projecting beyond metacoxae; juga surpassing

tylus (in part) Janeirona Distant

27. Posterior margin of metasternum excavated medially and apex of
abdominal spine fitting into this excavation (Fig. 14)

Glaucioides Thomas, new genus

- Posterior margin entire medially or with shallow subvertical

sulcus Banasa Stal

Elanela Rolston, new genus
Type species. — Elanela hevera Rolston, n. sp.

Diagnosis. — Median tubercle of abdominal sternite 3 (second visible) fit-
ting into notch in posterior margin of metasternum. Metasternum produced,
flat and bifurcate posteriorly, becoming obtusely carinate anteriorly. Meso-
sternum obtusely carinate mesially, forming continuous profile along meson
with metasternum, anteriorly apposing two low carinae on prosternum
which diverge and continue as anterior prosternal margin. Ostiolar ruga on

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NEW YORK ENTOMOLOGICAL SOCIETY

Figs. 1-8. 1 . . Placocoris viridis — right hind femur. 2, 3. Pallantia macula — 2. Left mid
femur, apex. 3. Right metathoracic scent gland. 4. Arvelius latus — right metathoracic scent
gland. 5. Pharnus inconspicuus — metasternum. 6. Pseudevoplitus longicornis — mesosternal
carina, lateral. 7. Lopadusa augur — right buccula, lateral. 8. Elanela hevera — left buccula,
lateral. (SI, S2), sternites one, two.

each metapleuron extending between 1 fourth and 1 half of distance from
mesial margin of ostiole to lateral margin of metapleuron. Femora unarmed;
tibiae sulcate; tarsi 3-segmented. Bucculae posteriorly curving to surface of
head. Basal segment of rostrum projecting past bucculae (Fig. 8); apex of
rostrum reaching metacoxae. Length of head about 6 tenths of width across
eyes. Juga and tylus subequal in length; lateral margins of juga deeply con-

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125

Figs. 9-17. 9. Brachystethus rubromaculatus — meso-, metasterna, lateral. 10. Janeirona

bergi — meso-, metasterna, lateral. 11. Tibilis parva — pro-, mesosterna, lateral. 12. Elanela
hevera — head, dorsal. 13. Elanela hevera — female genitalia. 14-17. Glaucioides englemani.
14. Metasternum, ventral. 15. Female genitalia. 16. Spermatheca. 17. Left paramere. (SI, S2),
sternites one, two.

cave before eyes (Fig. 12). Basal segment of antennae stout, reaching to or
slightly beyond apex of head.

Elanela hevera Rolston, n. sp.

Stramineous with castaneous to black punctures and markings. Length
7.5 mm, width at humeri 4.5 mm.

Head sparsely punctate excepting line of contiguous punctures on each
side touching ocellus mesially and curving anteriorly then laterally toward
middle of eye. Lateral margin of juga parallel between anterior convexity
and concavity before eyes, narrowly bordered in brown. Width of head

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across eyes 1.9 mm, length 1.2 mm; interocular width about 1.05 mm; width
across ocelli about 0.65 mm. Antennal segments 0.4, 1.0, 0.8, 1.5, 1.7 mm
long; distal half of segments 4 and 5 black. Rostral segments 2 through 4
about 1.0, 0.7, 0.5 mm long; distal end of segment 2 at middle of mesoster-
num, of segment 3 at mesocoxae, of segment 4 at metacoxae.

Pronotal punctures arranged in line on anterior and anterolateral submar-
gins, along posterior boundary of cicatrices and posterior boundary of nar-
row transverse callus lying immediately caudad of cicatrices; much smaller
punctures border posterior pronotal margin; a few small punctures inter-
spersed with many larger punctures scattered over pronotal disk. Antero-
lateral margins straight, narrowly ridged at edge. Humeri not produced.
Pronotal width at humeri 4.4 mm, length at meson 1.7 mm.

Scutellar punctations sparse on basal disk, dense elsewhere excepting
impunctate apex; large stramineous callus present in each basal angle; fovea
small, inconspicuous; basal width and length each 3.0 mm. Large stramin-
eous spot on disk of each corium located in an elongate impunctate area;
punctation elsewhere similar in density and strength to that on lateral por-
tions of scutellum. Connexivum broadly exposed, black to castaneous with
2 pale macules on each segment, one arching mesad from lateral margin,
one arching cephalad from posterior margin, these much reduced on first
visible segment, confluent on last segment; dark areas densely punctate,
light areas nearly impunctate.

Venter sparsely irregularly punctate, most strongly so on propleura.
Evaporative area matte, minutely granular, without fine rugae, castaneous.
Spiracles on sternite 2 exposed; peritremes dark. Posterolateral angles of
sternites (except second) produced as small black spines.

Genital plates as in Figure 13.

Holotype. — Female, labeled (a) "Peru: Iquitos, 100 mi. n.e., on Napo
R.” (b) "Mar. 20, 1969. B. K. Dozier.” Deposited in Florida State Collec-
tion of Arthropods (Gainesville, Florida). No paratypes.

Glaucioides Thomas, new genus

Type species. — Glaucioides englemani Thomas, n. sp.

Diagnosis. — Abdominal sternite 3 (second visible) bearing anteriorly di-
rected spinose tubercle, this tubercle received by elongated socket on pos-
terior surface of metasternum. Socket slanting anteroventrad, closed ven-
trally (open in related genera) (Fig. 14). Metasternum pubescent, mildly
convex, elevated posteriorly, sloping and subelevated anteriorly. Low flat
mesial carina on mesosternum broadened anteriorly, obsolescent on pos-
terior !4.

Apex of scutellum angulate, not bluntly rounded as in related genera.

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127

Ostiolar rugae sinuate, elongate, extending nearly to the edge of metapleura.
Femora unarmed; tibia terete; tarsi 3-segmented.

Basal segment of rostrum lying entirely within bucculae, latter evanescent
posteriorly. Jugae and tylus subequal in length. Basal segment of antennae
not surpassing apex of head.

Etymology . — Named to reflect morphological similarity to the Indo-Ma-
laysian genus Glaucias Kirkaldy 1908.

Glaucioides englemani Thomas, n. sp.

Stramineous, devoid of darkened punctures; abdominal tergites and some-
times jugae, pronotum and connexivum thinly margined with red; acuminate
tips of connexival angles black. Weak punctures densest on pronotum and
apical Vs of scutellum; head without or only feebly punctate; dorsal surface
strigose at bases of jugae.

Head width across the eyes 2.1 mm, length 2.2 mm; interocular width 1.3
mm. Antennal segments: 0.51, 0.75, 1.25, 1.50, 1.51 mm long; distal Vi of
segments 3 and 5, and nearly all of segment 4 dark brown. Rostral segments
2 through 4: 1.1, 1.0, 0.8 mm long, tip of rostrum just attaining abdominal
tubercle.

Anterolateral pronotal margins straight, obtuse; humeri not produced;
cicatrices inconspicuous. Pronotal width 5.6 mm, length at meson 2.5 mm.

Basal disk of scutellum mildly convex and more sparsely punctate than
apex. Apex of scutellum angulate, pointed; length 4.0 mm. Hemelytra with
weak, dense punctures, nearly translucent; membrane hyaline.

Venter of abdomen broadly V-shaped in cross section, sloping sides flat,
impunctate. Peritremes of spiracles concolorous with venter.

Tergite 6 of female bearing a pair of triangulate, posteriorly directed teeth
(Fig. 15). Spermathecal bulb with a lateral process (Fig. 16). Pygophore of
male open, ventral border broadly V-shaped from caudal view; inferior mar-
gin broadly, shallowly emarginate, subtended by a broad, shallow sulcus
which becomes obsolescent laterally. Pygophore otherwise unelaborated.
Proctiger broad, dorsal aspect trapezoidal, with a shallow, transverse, me-
sial depression. Parameres squat, goblet shaped, with a broad spatulate
apophysis extending from rim of its bowl (Figs. 17, 18). Aedeagus with a
pair of elongate thecal processes (Fig. 19).

Length. — 8 10.1 mm, 9 11.3 mm. Width: 8 5.6 mm, 9 6.3 mm.

Etymology. — This species is named for H. Dodge Engleman, M.D., of
Coco Solo, Panama, who has presented us with numerous specimens of
Pentatomidae for study, including one of the types upon which this species
is based.

Holotype. — 8 Sinop, Brazil, (Mato Grosso): Lat. 12°3T, Long. 55°37'.

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Figs. 18-24. 18, 19. Glaucioides englemani. 18. Left paramere, lateral. 19. Aedeagus. 20.

Nocheta adda — genital cup. Figs. 21, 22. Vidada dollingi. 21. Head, dorsal. 22. Female genitalia.
23. Serdia concolor — metasternum. 24. Ramosiana insignis — metasternum. (Co.l.), conjunc-
tival lobe, (m.p.l.), median penial lobe, (Th.), theca, (Th. ap.), thecal appendage.

Oct. 1974. M. Alvarenga Collr. Deposited in the American Museum of Nat-
ural History.

Paratypes. — 36 6 same locality, date and collr; 1 6 same locality and
collr., Oct. 1975; 1 6 Barro Colorado Island, Panama, 5 May 1937, S. W.
Frost collr; 1 9 Barro Colorado Island, Panama, 1-9 May 1964, W. D. and
S. S. Duckworth collrs; 16 Barro Colorado Islan, Panama, 1 May 1973,
D. Engleman collr; 19 Mepane Kamp, Surinam, Aug. 1961, J. P. Shulz

VOLUME LXXXVIII, NUMBER 2

129

collr; 1 9 Mt. Duida, Venezuela, 25 Feb. 1924, unk. collr; 1 9 Kuyuwini
River, British Guiana, 22 Nov. 1937, W. G. Hassler collr. Deposited in the
collections of the American Museum of Natural History, U.S. National
Museum, California Academy of Sciences and the authors.

Nocheta Rolston, new genus

Type species. — Nocheta adda Rolston, n. sp.

Diagnosis. — Median tubercle on abdominal sternite 3 (second visible) fit-
ting into notch in posterior margin of metasternum. Metasternum produced,
surface flat with lateral ramus on each side between meso- and metacoxae,
apposed anteriorly by mesosternal carina. Mesosternum tumescent; mesial
carina broad basally and continuing profile of metasternum when viewed
laterally, narrowing anteriorly to acuminate termination between procoxae.
Arms of chevron shaped production on prosternum diverging cephalad of
procoxae, continuing along anterior submargin of proplura, terminating be-
hind eyes. Ostiolar sulcus and ruga on each side reaching about 8 tenths of
distance from mesial margin of ostiole to lateral margin of metapleuron.
Femora unarmed; tibiae sulcate; tarsi 3-segmented. Bucculae evanescent
posteriorly. First rostral segment lying entirely between bucculae. Juga con-
vergent before tylus; lateral margin of juga deeply concave before eyes.
Basal segment of antennae scarcely projecting past apex of head.

Nocheta adda Rolston, n. sp.

Stramineous with dark castaneous to black punctation and markings.
Length about 9.1 mm, width at humeri 5.2 mm.

Line of dense punctation along both margins of tylus continuing over
vertex, converging at base of head; mesial margin and longitudinal band on
disk of juga densely punctate; punctures about eye irregularly disposed.
Lateral jugal margins sigmoid, nowhere parallel, narrowly bordered in
black. Width of head across eyes 2.5 mm, length 1.5 mm; interocular dis-
tance 1.2 mm; distance across ocelli about 0.85 mm. Ocelli rather larger,
about 0.2 mm in diameter. Antennal segments 0.5, 1.5, 1.6, 2.2, 2.1 mm
long; color stramineous to fulvous, first 3 segments darkly dotted, third
faintly so, fifth fuscous on distal half. Rostral segments 2 through 4 about
0.8, 0.9, 0.5 mm long; segment 2 reaching procoxae; apex of segment 4
nearly attaining mesocoxae.

Narrow pale callus traversing pronotum along posterior margins of cica-
trices. Row of closely spaced punctures surrounding cicatrices, a few punc-
tures scattered on cicatrices. Anterolateral margins straight with single row
of punctures between narrowly reflexed margin and submarginal impunctate
band. Punctation elsewhere on pronotum mostly in transverse lines, these

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longer and somewhat vermiform basally. Humeri not produced. Pronotal
width 5.2 mm, length at meson 2.2 mm.

Scutellar punctations in transverse vermiform castaneous bands; basal
angles without callus or fovea; width at base 3.5 mm, length 4.0 mm; frena
extending along basal 6 tenths; apex acute. Coria blotched with castaneous;
punctation stronger than on scutellum; membrane fumose with about a doz-
en simple veins. Connexiva moderately exposed, alternately banded where
exposed with dark border on each side of intersegmental sutures.

Venter sparsely and weakly punctate excepting moderately dense punc-
tation on propleura. Evaporative areas matte, not rugulose, concolorous
with remainder of meso- and metapleura. Spiracular peritremes concolorous
with surrounding area of sternites.

Posterior surface of pygophore with deep mesial triangular impression
below emargination of dorsal border; subacute tooth present on each side
of emargination. Parameres acicular, partially obscuring subvertical ridge
on anterior wall of genital cup (Fig. 20).

Holotype. — Male labeled (a) “Hyutanahan Rio Purus Brazil. S. M.
Klages.” (b) “March 1922.” Deposited in U.S. National Museum. Type no.
72138. No paratypes.

Vidada Rolston, new genus

Type species. — Vidada dollingi Rolston, n. sp.

Diagnosis. — Metasternal production almost flat, posterior margin lightly
notched mesially, anterior margin truncate and slightly less produced than
adjacent part of mesosternum. Mesosternal carina broad, flattened between
mesocoxae and procoxae, compressed between procoxae, not extending
onto prosternum. Prostemum tumescent with low carina on each side curv-
ing anterolaterad from between procoxae. Ostiolar ruga on each metapleu-
ron extending about 3 tenths of distance from mesial margin of ostiole to
lateral margin of metapleuron. Femora unarmed; tibiae sulcate; tarsi 3-seg-
mented. Bucculae sloping caudad to surface of head. Basal segment of ros-
trum projecting beyond bucculae; apex of rostrum reaching mesial tubercle
on abdominal venter. Length of head about 8.5 tenths of width across eyes.
Juga contiguous before tylus, their lateral margins moderately concave be-
fore small obtuse anteocular process (Fig. 21). Basal segment of antennae
projecting little beyond apex of head.

Vidada dollingi Rolston, n. sp.

Dorsum castaneous becoming fuscous on pronotum anteriorly and disk
of head; venter dark castaneous; appendages light castaneous excepting
antennal segment 3 distally and 4 fuscous. Length about 12.3 mm, width
across humeri 5.9 mm.

VOLUME LXXXVIII, NUMBER 2

131

Head flat above, jugal margins reflexed; punctation dense, somewhat ru-
gose. Eyes appearing small relative to head width; width of head across
eyes 2.35 mm, length 1.9 mm; interocular width 1.5 mm; distance across
ocelli 1.1 mm. Length of antennal segments 0.8, 0.5, 1.1, 1.4 mm (last miss-
ing). Rostral segments 2 through 4 about 1.6, 1.5, 1.0 mm long.

Anterolateral margins of pronotum reflexed, slightly sinuous. Punctation
dense, all of disk behind cicatrices somewhat rugose. Width at humeri 5.9
mm, length at meson 2.6 mm.

Scutellum less densely punctate but basally more rugose than pronotum;
fovea in basal angles deep; basal width 3.6 mm, length 4.7 mm; frena ex-
tending along basal 6 tenths of scutellum. Coria punctate similarly to prono-
tum, immaculate; membrane darkly fumose with about 7 weak simple veins.
Connexiva almost unicolorous, modestly exposed.

Thoracic venter strongly and densely punctate ; abdominal venter weakly
but densely punctate laterally, becoming almost impunctate mesially. Evap-
orative area strongly rugose, punctate. Spiracles small, oval, with black
peritreme.

Genital plates as in Figure 22; surface of 9th paratergite convex, strongly
so basally.

Holotype. — Female, labeled (a) “Machu-Picchu, Cuzco. 8-V-65” (b) “ar-
busto” (c) “1067” (d) “CUZCO.” Deposited in Museu Nacional, Rio de
Janeiro, Brazil.

No paratypes.

Comments. — This species is dedicated to W. R. Dolling, of the British
Museum (Natural History), whose unstinting assistance to fellow hemipter-
ists is indispensable.

Acknowledgments

We wish to express our gratitude to Drs. N. Mpller Andersen (Univer-
sitetets Zoologiske Museum), W. R. Dolling (British Museum (Natural His-
tory)), H. Dodge Engleman, Richard C. Froeschner (U.S. National Mu-
seum), Jocelia Grazia (Universidade Estadual de Campinas), Frank W.
Mead (Florida State Collection of Arthropods) and Randall T. Schuh
(American Museum of Natural History) for loans of specimens.

Literature Cited

Jensen-Haarup, A. C. 1931. Hemipterological notes and descriptions VI. Entomol. Medd.
Copenhagen 17:319-336.

. 1937. Einige neue Pentatomidenarten aus der Sammlung des Zoologischen Museums

in Hamberg (Hem. het.). Entomol. Rundschau 54:321-324.

Rolston, L. H. and F. J. D. McDonald. 1979. Keys and diagnoses for the families of Western
Hemisphere Pentatomoidea, subfamilies of Pentatomidae and tribes of Pentatominae
(Hemiptera). J. N.Y. Entomol. Soc. 87(3): 189-207.

132

NEW YORK ENTOMOLOGICAL SOCIETY

(LHR) Department of Entomology, Louisiana Agricultural Experiment
Station, Louisiana State University, Baton Rouge, Louisiana 70803; (FJDM)
Department of Plant Pathology and Agricultural Entomology, University of
Sydney, Sydney, N.S.W., Australia 2006 and (DBT) Department of Ento-
mology, University of Missouri, Columbia, Missouri 65211.

Received for publication October 9, 1979.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(2), 1980, pp. 133-142

ANATOMY OF THE MALE REPRODUCTIVE SYSTEMS OF THE
ADULTS AND PUPAE OF TWO DORYLINE ANTS,
DORYLUS ( ANOMMA ) WILVERTHI EMERY AND
D. (A.) NIGRICANS ILLIGER12

Francis C. Ford1 2 3 and James Forbes

Abstract. — This is the first study of the anatomy of the male reproductive
systems of the adults and pupae of the doryline ants, Dorylus ( Anomma )
wilverthi and D. (A.) nigricans Illiger. The external genitalia and the ter-
minal gastric sterna are included. The reproductive systems consist of the
testes and vasa efferentia, the vasa deferentia, the seminal vesicles, the
accessory glands, the bound accessory gland duct, the ejaculatory duct and
wedge, the aedeagal bladder, and the external genitalia. The external geni-
talia of these ants are composed of a basal ring and three pairs of valves,
the outer, the middle, and the inner, typical of the formicid pattern. The
male systems in these two species are similar in their shapes and the ar-
rangement of the organs. Although the testes in the nigricans adult are
absent, in the pupa there are 50-55 follicles in each testis. Wilverthi has 35-
40 testicular follicles in both the adult and pupa. The valves of the external
genitalia and terminal gastric sterna of these two species show no individual
variations, other than size, in their shapes. Male systems have been de-
scribed for only three other doryline ants, the Old World Dorylus labiatus,
the New World Eciton hamatum and Neivamyrmex harrisi. This system is
compared in these dorylines. Important differences exist between the male
reproductive systems of the two African Dorylus species herein studied and
the two New World dorylines previously reported. The Dorylus species
have a larger number of testicular follicles, the shapes of their accessory
glands are different, the basal ring of the genitalia is fused to the outer
valves, a membrane joins the ventral margins of the inner genitalic valves,
and the shapes of the genitalic valves and the subgenital plate are different.

This paper presents the first description of the anatomy of the male re-
productive systems of the adults and pupae of two Old World dorylines,
Dorylus ( Anomma ) wilverthi Emery and Dorylus ( Anomma ) nigricans II-

1 Hymenoptera: Formicidae.

2 Part of the Ph.D. dissertation of the first author at the Department of Biological Sciences,
Fordham University.

3 Present address: Department of Biology, Bronx Community College, CUNY, Bronx, N.Y.
10453.

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NEW YORK ENTOMOLOGICAL SOCIETY

liger. Descriptions of the genitalic valves and the terminal gastric sterna are
also included. Male systems have been described for only three other dor-
yline ants, the Old World Dorylus labiatus (Mukeijee 1926), the New World
Eciton hamatum (Forbes 1958) and Neivamyrmex harrisi (Forbes and Do-
Van Quy 1965). This system will be compared in these dorylines.

Seven to nine specimens each of the adults and the pupae were kindly
furnished by Albert Raignier, S.J. of Belgium, who collected these ants in
the Republic of the Congo, Africa, now designated Zaire, during June of
1956 and November of 1957. The pupae were advanced in development.
The entire reproductive systems were dissected from the gasters of the
adults and pupae, stained with borax carmine, and prepared as whole
mounts for study. The external genitalia and terminal segments were also
removed and mounted for study. Illustrations were prepared with the aid of
a B and L trisimplex microprojector.

The reproductive systems consist of the testes and vasa efferentia, the
vasa deferentia, the seminal vesicles, the accessory glands, the bound ac-
cessory gland duct, the ejaculatory duct and wedge, the aedeagal bladder,
and the external genitalia (Figs. 1-3). Throughout the following descriptions,
the adult system of wilverthi will be described first, followed by differences
in the pupal system. Comparisons will then be made with the adult and pupa
of nigricans.

The testes of wilverthi are located in the dorsal median region of the
gaster and extend through the 2nd and 3rd gastric segments above the ven-
triculus (Fig. 1). The posterior portion of each testis overlies most of the
seminal vesicles. Each testis is composed of about 35-40 slender, thin-
walled testicular tubules that are about 8 mm in length. Each tubule ends
in a narrow duct, a vas efferens. The number and the arrangement of the
testicular tubules in the pupa of wilverthi are similar to those of the adult.
In the adult of nigricans the testes and vasa efferentia are absent. However,
in the pupa each testis consists of 50-55 tubules, and it is similar in its shape
and position to that of wilverthi. The testes of these two species are not
covered with connective tissue sheaths or capsules.

The vasa efferentia join to form a short duct, the vas deferens. Where the
vasa efferentia unite, the vas deferens is slightly more dilated than it is
distally. The distal portion enters the expanded seminal vesicle just below
its anterior end. The vas deferens is absent in the adult of nigricans but
present in the pupae of wilverthi and nigricans.

The seminal vesicles are large, prominent, thick-walled, U-shaped tubes
that lie in the 3rd and 4th gastric segments (Fig. 1). The first part of the
proximal arm lies at right angles to the direction of the testicular tubules.
It continues forward and ventrolaterally, then bends sharply posteromediad
toward the posterior region of the ventriculus. The proximal arm is slightly
shorter than the distal, but both arms are nearly equal in diameter. The

VOLUME LXXXVIII, NUMBER 2

135

Fig. 1. Diagram of a lateral dissection of the posterior portion of the gaster of the adult
male ant Dorylus ( Anomma ) wilverthi. Abbreviations: Ag, accessory gland; Ed, ejaculatory
duct; InV, inner valve; M, Malpighian tubule; MV, middle valve; OV, outer valve; R, rectum;
SV, seminal vesicle; T, testis; V, ventriculus; Vd, vas deferens; VIII-IX, Roman numerals
designate abdominal segments.

position and arrangement of the seminal vesicles in the pupa of wilverthi
and in the adult and pupa of nigricans are similar to that in the adult of
wilverthi. Differences in the diameters of these organs between the adults
of wilverthi, the larger species, and nigricans are about 25-30 percent.
Likewise in both species, the diameters of the seminal vesicles of the pupae
are about Vx to V3 smaller than those of the adults. The seminal vesicles of
the nigricans adults were packed with sperm, while those of wilverthi had
sperm scattered throughout. No sperm is present in these organs in the
pupae. Toward the posterior margin of the 4th gastric segment each seminal
vesicle tapers gradually, bends dorsomedially, and joins the accessory gland
on its ventro median surface.

Each accessory gland is a prominent, thick-walled, S-shaped tube lying
above the seminal vesicles, and located on either side of the posterior region
of the ventriculus (Fig. 1). Their free ends are close to the midline and point
forward. The gland bends sharply laterally and anteriorly a short distance,
then dips ventroposteriorly, turns toward the midline at the posterior margin
of the 4th gastric segment, where the seminal vesicle joins it on its ventro-
median surface. Beyond this junction each accessory gland tapers slightly,
joins the gland from the other side, and forms a single tube. The first portion
of this tube is the bound accessory gland duct, and the end portion where

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VOLUME LXXXVIII, NUMBER 2

137

the wedge is located is the ejaculatory duct. The bound accessory gland
duct bends dorsally at the anterior edge of the 5th gastric segment.

The ejaculatory duct passes through the basal ring and between the an-
terior borders of the large, prominent, outer genitalic valves and enters the
middorsal surface of the aedeagal bladder (Figs. 1 and 4). The posterior end
of the ejaculatory duct is depressed into the middorsal wall of the aedeagal
bladder.

The aedeagal bladder is a muscular- walled, ovoid sac that lies beneath
the basal portions of the inner genitalic valves, and it opens posteriorly
between the inner valves. Muscle fibers on the outer wall of the aedeagal
bladder are attached to the anteroventral surfaces of the inner valves.

The accessory glands, the bound accessory gland ducts, the ejaculatory
ducts and the aedeagal bladders in the wilverthi pupa and the nigricans
adult and pupa are similar to those of the wilverthi adult.

Genitalia and Terminal Gastric Segments

The external genitalia of wilverthi and nigricans (Figs. 1 through 6) are
retracted into a cavity within the last few gastric segments beneath the
rectum and the anus, and only the posterior tip of the ninth sternum or
subgenital plate projects from the end of the gaster. This arrangement is
characteristic of the dorylines (Borgmeier 1955). The external genitalia of
these ants are composed of a basal ring and three pairs of valves, the outer,
the middle, and the inner, typical of the formicid pattern (Clausen 1938;
Snodgrass 1941; Krafchick 1959).

The basal ring or lamina annularis is a narrow, ring-like segment that is
moderately sclerotized throughout (Figs. 2 and 3). The dorsal part is wider
than the lateral and ventral portions, and it is fused to the dorsal, anterior
borders of outer valves; its middorsal region forms a bridge between the
outer valves. On the ventral part there is a small middorsal apodeme.

The outer valves or parameres are the largest and most heavily sclerotized
of the three pairs (Figs. 2 and 3). They are laterally convex. From a side
view the overall shape of the valve is somewhat C-shaped with the open
part of the C directed downward. There is no division into an anterior lamina
parameralis and a posterior paramere. The anterior portion of the valve is
large and cup-shaped, and it covers the anterior tips of the middle valves
and the bases of the inner valves. The posterior portion of the valve, when

Figs. 2, 3. 2. Diagram of a lateral view of the male genitalia of wilverthi. 3. Diagram of a

dorsal view of the male genitalia of D. wilverthi. Both diagrams are drawn to the same scale.
Abbreviations: BR, basal ring; InV, inner valve; MV, middle valve; OV, outer valve; Sp,
spathe.

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NEW YORK ENTOMOLOGICAL SOCIETY

Figs. 4-6. 4. Diagram of a right lateral view of the ejaculatory duct, the inner genitalic

valve, and the aedeagal bladder of the adult male of D. wilverthi. Abbreviations: AB, aedeagal
bladder; Ap, lateral apodeme; Ed, ejaculatory duct; InV, inner valve; Sp, spathe. 5. Diagram
of the dorsal view of the ninth sternum or subgenital plate of the male of D. wilverthi. 6.
Diagram of a ventral view of the eighth sternum of the male of D. wilverthi.

viewed laterally enlarges slightly. The ventral margin of this posterior por-
tion is covered with numerous, long, slender hairs. Dorsally and ventrally
the outer valves are separated from each other.

The middle valves or volsellares are the shortest of the three pairs, and
they are finger-shaped (Figs. 1, 2, and 3). The basal portion is the tallest,
and it tapers posteriorly. The ventroanterior part of the basal portion is
joined to the ventroposterior tip of the base of the outer valve by bands of
muscle. These middle valves lie within the posterior portions of the outer

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139

valves, and they extend as far as or slightly beyond the posterior margins
of the outer valves. The outer surface of these middle valves is convex.
Fine, sensory hairs extend along the middle of the outer surface of this
valve.

The inner valves or laminae aedeagales constitute the aedeagus or male
intromittant organ (Figs. 1 through 4). The basal portion of the valve is
laterally convex and has a ridge-like apodeme on its outer surface. Numer-
ous bands of muscle extend from the lateral apodemes to the inner surface
of the base of the outer valves. The remaining two-thirds of these valves
are bent downward in the middle, they are scimitar-like in shape, and they
are heavily sclerotized except at their tips. A heavy, non-sclerotized mem-
brane, the spathe, joins these valves dorsally almost to their tips. A small
triangular sclerite is present on its anterior end. These valves are also united
ventrally by a membrane that extends almost to their tips. These valves
extend slightly beyond the posterior margin of the outer valves.

The eighth sternum is roughly rectangular in shape and moderately scler-
otized throughout (Fig. 6). Its anterior border is thickened and deeply in-
dented, while the posterior margin is weakly sclerotized and rounded in
shape. Two patches of dark, sensory hairs are present on its ventral surface.

The ninth sternum, or subgenital plate, is narrow in shape, bifid poste-
riorly (Figs. 1 and 5), and is heavily sclerotized throughout. The anterior
margin is thick; in the central region of this segment there is a depression
on the dorsal surface into which many muscle fibers are inserted. Here,
also, there are two small, lateral, triangular apodemes. The ventral surface
of the posterior end is covered with many sensory hairs. Posteriorly the tips
of the segment extend beyond the end of the gaster and the posterior margins
of the external genitalia.

In the wilverthi pupa the arrangement, shapes, and position of the valves
and terminal gastric sterna are similar to those of the adult, but they are not
as heavily sclerotized; in the adult they are dark brown in color. In the
nigricans adult and pupa the arrangements, shapes, and positions of the
valves and terminal gastric sterna are similar to those of wilverthi , and these
structures in the pupa are less sclerotized. The genitalic valves and the
terminal gastric sterna in nigricans are smaller in size, it is a smaller ant,
and they are reddish-yellow in color.

Discussion

The testes in the adults and pupae of wilverthi and the pupae of nigricans
are large and extend through the second and third gastric segments. The
testes in the adults of nigricans are absent. In N. harrisi (Forbes and Do-
Van Quy 1965) the testes are small and lie in the posterior region of the
third gastric segment, in E. hamatum (Forbes 1958) they are large and

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extend through the first three gastric segments, and in D. labiatus (Mukeijee
1926) no specific location was designated. The testicular follicles in both
wilverthi adults and pupae, and nigricans pupae are long, slender tubules,
35-40 in each testis of wilverthi and approximately 50-55 in each testis of
nigricans. The testicular follicles in N. harrisi and E. hamatum are long,
slender tubules, 22-25 in each testis of N. harrisi and 20 in each testis of
E. hamatum. In D. labiatus each testis contains “a fair number” of tubular
follicles. The testicular follicles of all of the ants end in short narrow ducts,
the vasa efferentia.

No connective tissue sheath covers the testes of the two Dorylus species
herein reported. A single capsule covers the testes in N. harrisi, while each
testis is covered by a capsule in E. hamatum. No capsule is reported by
Mukeijee (1926) in D. labiatus. In wilverthi and nigricans the vasa effer-
entia join to form a short duct, the vas deferens. This arrangement is similar
in N. harrisi and E. hamatum. Structures comparable to vasa efferentia are
shown, but not labelled, by Mukeijee in his illustrations of the reproductive
system of D. labiatus.

The seminal vesicles in wilverthi and nigricans adults and pupae are
prominent, thick-walled, U-shaped tubes lying in the third and first half of
the fourth gastric segments. In N. harrisi (Forbes and Do-Van Quy 1965)
and E. hamatum (Forbes 1958) the position, shape, and arrangement of
these organs are similar to that in wilverthi and nigricans. The terminology
of this organ in male ants has been clarified by Hung and Vinson (1975).
Mukeijee (1926) reports an organ in D. labiatus that he calls the seminal
duct and describes a swelling at the anterior end, the collecting sac, and a
larger dilation at its posterior end, the vesicula seminalis. He reports that
this organ is usually U-shaped and also notes variations in the position and
size of the vesicula seminalis. In this study nothing resembling the descrip-
tion and diagram of the seminal duct of D. labiatus was found.

The accessory glands in the adults and pupae of wilverthi and nigricans
are prominent, thick-walled, S-shaped tubes lying in the fourth gastric seg-
ment. In N. harrisi and E. hamatum the accessory glands are tightly coiled
tubes situated on either side of the intestine in the fourth gastric segment.
The accessory glands of D. labiatus are large, slightly curved, thick-walled
tubes, which in some cases have an appendix. Mukerjee indicates variations
and anomalies in the reproductive system of D. labiatus. No such anomalies
or variations were observed in the small number of specimens available for
this investigation.

The bound accessory gland duct in wilverthi and nigricans is very short.
In N. harrisi this organ is approximately equal to the accessory gland in
length. In E. hamatum this duct encircles the ventriculus five or six times
and is 28 to 31 mm in length. In D. labiatus no bound accessory gland duct

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141

is reported, although a portion of what is shown as the ejaculatory duct may
be the bound accessory gland duct.

The ejaculatory duct in wilverthi and nigricans is similar to that of N.
harrisi and D. hamatum, and in all these ants a cuticular wedge is present
in this organ. The ejaculatory duct, illustrated by Mukeijee (1926) in D.
labiatus appears to be long, and he shows a wedge-shaped structure in the
posterior part of this duct which he designates “penes.” He found in a
single specimen a blind diverticulum on the dorsal side of this duct. No such
structure has been reported in any other doryline.

An aedeagal bladder is present in wilverthi , and nigricans as well as N.
harrisi and E. hamatum. Mukerjee (1926) made no mention of this organ in
D. labiatus.

The valves of the external genitalia and the terminal gastric sterna of these
two Dorylus species show no individual variations, other than size, in their
shapes. This is contrary to what Borgmeier (1955) reported in his study of
the Neotropical dory lines where even in what may be related species dis-
tinctive differences are present in some genitalic valves and the subgenitalic
plate.

Important differences, therefore, exist between the male reproductive
systems of the two African Dorylus species herein studied and the two New
World dorylines previously reported. The Dorylus species have a larger
number of testicular follicles, the shapes of their accessory glands are dif-
ferent, the basal ring of the genitalia is fused to the outer valves, a membrane
joins the ventral margins of the inner genitalic valves, and the shapes of the
genitalic valves and the subgenital plate are different.

Literature Cited

Borgmeier, G. 1955. Die Wanderameisen der Neotropischen Region (Hymenoptera, Formic-
idae). Studia Entomologia, Nr. 3, Editora Vozes Limitada, Petropolis, R. J. Brasil, 3:9—
716.

Clausen, R. 1938. Untersuchungen iiber den mannlichen Copulatinsapparat der Ameisen,
speciell der Formicinae. Mitteil. Schweizer. Entomol. Gesells. 17:233-346.

Forbes, J. 1958. The male reproductive system of the army ant, Eciton hamatum Fabricius.
Proc. Xth 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 (Haldman) (Hymenoptera: Formicidae). J.
N.Y. Entomol. Soc. 73:95-111.

Hung, S. C. F. and S. B. Vinson. 1975. Notes on the male reproductive system in ants
(Hymenoptera, Formicidae). J. N.Y. Entomol. Soc. 83:192-197.

Krafchick, B. 1959. A comparative study of the male genitalia of North American Ants (For-
micidae) with emphasis on generic differences. Dissertation, University of Maryland.

Mukerjee, D. 1926. The digestive and reproductive systems of the male ant, Dorylus labiatus
Schuck. J. and Proc. Asiatic Soc. Bengal, n. s. 22:87-92.

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NEW YORK ENTOMOLOGICAL SOCIETY

Raignier, A. and J. Van Boven. 1955. Etude taxonomique, biologique et biometrique des
Dorylus du sous-genre Anomma. Ann. Mus. R. du Congo Beige, Sc. Zool. 2:1-359.
Snodgrass, R. 1941. The male genitalia of Hymenoptera. Smithson. Misc. Col. 99, No. 14.
86 pp., 33 pis.

Department of Biological Sciences, Fordham University, Bronx, New
York 10458.

Received for publication December 17, 1979.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(2), 1980, pp. 143-145

APPARENT NEST SITE COMPETITION BETWEEN THE PAPER
WASP POLISTES FUSCATUS (HYMENOPTERA: VESPIDAE)
AND THE HOUSE WREN

David L. Gibo

Abstract. — Small colonies of Polistes fuscatus can suffer significant mor-
tality due to attacks by house wrens. The wrens apparently compete with
the wasps for nest sites.

The house wren Troglodytes aedon uses a variety of cavities as nest sites,
including nest boxes, discarded tin cans, holes in banks, natural cavities in
trees or stumps and bleached skulls of livestock (Bent 1948). Colonies of
social wasps, particularly of the genus Polistes, also make use of similar
nest sites and have been recorded in nest boxes (McAtee 1929, 1931), dis-
carded tin cans and similar items (Rau 1942), holes in banks (Hungerford
and Williams 1912) and cavities in trees and skulls of livestock (Rau 1931,
1942). Consequently, it is not surprising that there are occasional records
of wrens and other small birds being displaced by social wasps (Bent 1948;
Hart 1941; McAtee 1929, 1931). However, because the male house wren is
exceedingly aggressive, he attempts to occupy all available nest sites in his
territory (Bent 1948), T. aedon is potentially capable of competing with
social wasps.

This note reports on a nesting pair of T. aedon that systematically exter-
minated approximately 15-17 colonies of the social wasp, Polistes fuscatus
(Fabricius) during apparent competition for nest sites. The study area was
located at the Wasp Studies facility on the Erindale Campus of the Univer-
sity of Toronto, Ontario, Canada. The particular location, which has been
described previously (Gibo 1978), consists of two sets of nest boxes (sites
A and B) and an abandoned shed located in a field adjacent to a woodlot.
At site A 40 nest boxes were arranged in a 5 x 8 grid, with each nest box
approximately 0.75 m from any other nest box. At site B 8 nest boxes were
arranged in a 2 x 4 grid. The two sets of nest boxes were separated by
approximately 20 m. The shed was approximately 20 m from nest box site
A and 10 m from site B. The facilities had been utilized by a resident pop-
ulation of P. fuscatus for the past 6 seasons.

Poultry screen, with openings of 2.5 cm in diameter, was placed over the
openings of the nest boxes and the windows of the shed. The screen was
installed to protect the P. fuscatus colonies from attacks from birds, chiefly
blue jays, red- winged blackbirds, Baltimore orioles, and other medium size
species that are known to prey on P. fuscatus colonies in early summer
(Gibo 1978). House wrens, which had not been observed at the facility in
previous seasons, were able to slip through the poultry screen.

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NEW YORK ENTOMOLOGICAL SOCIETY

The pattern of colony initiation for P. fuscatus was typical. Foundresses
initiated colonies in April or May and often were joined by other foundress-
es. By mid-June approximately 20-45 colonies were established in the study
area and these remained active for the rest of the season. The colonies
increased in size steadily and the first workers appeared by late June to
early July. Although small nests with only a few cells were often abandoned
early in the season, they were not knocked down and remained in place,
empty and deserted, throughout the season. Predation on the colony by
birds normally does not occur in this area until late June or early July, and
in past seasons was prevented at the nest boxes with the screen (Gibo 1978).

In the spring of 1979 the normal pattern of colony development was ob-
served at grid A until May 22 when a colony, with two foundresses and 13
cells, was found knocked down and all eggs and adults were missing. By
this date at least 13 and possibly 15 colonies already were established. On
June 5 three more colonies, one a two foundress colony with 12 cells, and
the rest single foundress colonies, were found destroyed. All had construct-
ed at least 10 cells and, judging from records of previous years, normally
would have survived. Piles of twigs were found on the bottoms of two of
the nest boxes that had been occupied by the P. fuscatus colonies. By June
12 all of the initial colonies were either destroyed or deserted and piles of
twigs (nest building efforts of the T. aedon male) were found in all but two
nest boxes. The pattern of colony development was similar at grid B. Two
colonies were initiated and both were destroyed by early June. The nest
boxes received deposits of twigs. A female joined the male on approximately
the first of June and one of the nest boxes was subsequently utilized to rear
a brood.

The colonies at the shed had a normal season. Nineteen colonies were
established by mid-May, 18 of which were still surviving by mid- June. The
single failure was not associated with destruction of the nest, but with loss
(or desertion) of the single foundress. Colonies in the shed were completely
destroyed by birds on two previous occasions, both in early July. The first
attack occurred early in the study, prior to installation of the poultry screen
on the shed windows. The second attack occurred two years later when
birds entered through a hole near the door. This year the wrens apparently
avoided the colonies in the shed.

The complete extermination of the box nesting colonies at the study sites
differs from reports of other bird -Polistes interactions. It has been docu-
mented that birds prey on the adult wasps and brood, and usually attack
prior to or shortly after the emergence date of the first few workers. At this
time the colonies represent a large and relatively defenseless food supply
(Gibo 1978; Gibo and Metcalf 1979). Although the house wrens may have
been feeding on the wasps, the behavioral patterns observed, including (1)
removal of all Polistes nests in boxes, regardless of size (most nests only
contained a few eggs when removed), (2) filling the nest boxes with sticks,

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145

and (3) ignoring the P. fuscatus colonies in the shed, indicate that compe-
tition for nest sites was the main Polistes- wren interaction. Consequently,
territorial behavior and nest site competition by house wrens appears to
represent a major threat to P. fuscatus colonies during the early period of
colony initiation and may occasionally be responsible for local extinction of
a concentration of colonies. Despite the tendency of the P. fuscatus foun-
dresses to initiate colonies in the immediate area of the parental nest site
(Gibo 1972; Rau 1940; West-Eberhard 1969), the total destruction of local
concentrations of colonies as a result of predatory or territorial activities of
birds, suggests that dispersal must play a major role in maintaining the
population.

Acknowledgments

This research was supported by National Research Council of Canada
grant number A5991. I am indebted to Brigette Kemper for assistance with
fieldwork.

Literature Cited

Bent, A. C. 1948. Life Histories of North American Nuthatchers, Wrens, Thrashers, and their
allies. Dover Pub., New York.

Gibo, D. L. 1978. The selective advantage of foundress associations in Polistes fuscatus
(Hymenoptera: Vespidae): a field study on the effects of predation on productivity. Can.
Ent. 110:519-540.

and R. A. Metcalf. 1978. Early survival of Polistes apachus (Hymenoptera: Vespidae)

colonies in California: a field study of an introduced species. Can. Ent. 110:1339-1343.

. 1972. An introduced population of social wasps, Polistes apachus , that has persisted

for 10 years (Hymenoptera: Vespidae). Bulletin S. Calif. Acad. Sci. 71:53.

Hart, R. E. 1941. Blitzed birdhouse. Nat. His. 48:256.

Hungerford, H. B. and F. X. Williams. 1912. Biological notes on some Kansas Hymenoptera.
Ent. News 23(6):241-260.

McAtee, W. L. 1931. Paper wasps (. Polistes ) in bird houses. Proc. Ent. Soc. Wash. 33(7): 186.

. 1929. Paper wasps ( Polistes ) as pests in bird houses. Proc. Ent. Soc. Wash. 31(7): 136.

Rau, P. 1942. The nesting habits of Polistes as a factor in taxonomy. Ann. Ent. Soc. Amer.
35:336-338.

. 1940. Co-operative nest-founding by the wasp Polistes annularis Linn. Ann. Ent.

Soc. Amer. 33:617-620.

. 1931. The nests and nesting sites of four species of Polistes wasps. Bull. Brooklyn

Ent. Soc. 26(3): 11 1-1 19.

. 1929. The habitat and dissemination of four species of Polistes wasps. Ecology 10: 191—

200.

West-Eberhard, M. J. 1969. The social biology of polistine wasps. Misc. Publ. Univ. Mich.
(Mus. Zool.) 140.

Department of Zoology, Erindale College, University of Toronto, Missis-
sauga, Ontario L5L 1C6, Canada.

Received for publication December 20, 1979.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(2), 1980, pp. 146-150

OVERWINTERING OF POLISTES FUSCATUS IN CANADA: USE OF
ABANDONED NESTS OF DOLICHOVESPULA AREN ARIA

David L. Gibo

Abstract. — Over 350 females of the social wasp P. fuscatus were found
overwintering in southern Ontario in two old D. arenaria nests. Most of the
wasps were in a damaged nest that was attached to the southern side of a
building and fully exposed to the sun. Mid- winter mortality was 2.8% or
less. A test nest fitted with thermistors and placed in the original location
of nest A showed that although the overwintering wasps can occasionally
gain a moderate amount of protection (2.5°C) from low ambient tempera-
tures in the intact nest, there was essentially no protection in a damaged
nest. This study presents further evidence that adult P. fuscatus can survive
nearly complete exposure to the midwinter temperature regime of southern
Ontario.

Young queens of the social wasp genus Polistes are very opportunistic
in their choice of overwintering sites and often appear to require only mod-
est protection from the weather. Various species have been found overwin-
tering in a variety of natural and man-made sites, including cracks and
crevices in buildings (Gibo 1972; Snelling 1954; West-Eberhard 1969), under
shingles and loose tarpaper (West-Eberhard 1969; Gibo unpublished data),
in crevices in rocky bluffs (Rau 1930), under bark (Bohart 1942) and in
stumps (Hermann et al. 1974). The wasps were often found in clusters, and
several species may be present (Bohart 1942; Hermann et al. 1974; Snelling
1954). Old abandoned Dolichovespula nests are also utilized by Polistes as
hibernation sites. Green et al. (1976) found 39 P. fuscatus aurifer queens in
a nest in southeastern Washington and Zabriskie (1894) collected two Dol-
ichovespula nests from buildings during winter in New York and found
approximately 160 overwintering Polistes in each.

The occasional discovery of large numbers of Polistes in old Dolicho-
vespula nests suggests that the wasps may select the nests, when available,
in preference to other sites. However, the advantages in utilizing the aban-
doned nests are not readily apparent. Although these aerial nests are usually
inaccessible to some potential predators active during winter, such as
shrews and ground dwelling rodents, they are vulnerable to other predators
such as squirrels, raccoons and birds. Furthermore, the old nests would be
expected to offer little protection from low winter temperatures. The insu-
lating properties of Dolichovespula nests are known to be relatively modest,
and nest temperatures significantly higher than ambient temperatures are

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only achieved through large energy expenditures by a population of active
adult wasps (Gibo et al. 1974a, 1974b, 1976). An abandoned nest occupied
by inactive Polistes should, with some lag, simply track the air temperature.
The extent of the lag depends upon the effectiveness of the insulation, the
rate of change in the air temperature, and the weather (particularly wind
velocity). When air temperatures drop rapidly and then rebound the insu-
lating properties of the nest material could, under favorable conditions
(primarily little or no wind), have a moderating effect on the nest temper-
ature. Of course, this moderating effect will also apply during brief periods
of high ambient temperatures. A slightly more moderate temperature regime
and a reduced amplitude in the daily temperature fluctuations, could, on
occasion, be important for P. fuscatus survival. However, the actual ther-
mal environment of old Dolichovespula nests during winter is unknown.
The purpose of this note is to (1) report the use of old Dolichovespula nests
by P. fuscatus in Canada, (2) report the midwinter survival rate of these
wasps, and (3) examine the effectiveness of an old Dolichovespula nest in
moderating the winter temperature regime.

Two old nests of D. arenaria were collected in midwinter in southern
Ontario and examined for P. fuscatus. Nest A was collected on January 2,
1979. This nest had been partially destroyed by winter storms in 1978 and
was missing part of its envelopes, partially exposing the combs on one side.
It contained 354 P. fuscatus. Nest B was collected on January 11, 1979 and
contained only 7 P. fuscatus. Midwinter mortality was low for both groups:
2.8% (10 wasps) for nest A and 0% for nest B. All of the overwintering P.
fuscatus were females and, presumably, young queens. (West-Eberhard
(1969) reported that all overwintering and overwintered P. fuscatus females
that she examined (N = 28) during her studies in Michigan were found to
be inseminated, and thus were potential queens.) In nest A a few wasps
were found overwintering between envelope layers but most were either
clustered on the combs or wedged head first into the cells. This latter posture
is often observed when P. fuscatus are collected from their own nests in
the fall. The wasps in nest B were clustered between the combs. Approxi-
mately 20 overwintering Diptera (Tachinidae) were also found in nest A.

The sites of the two old nests were similar. Both were attached to the
overhang of the roof of a four-storey concrete building of the Erindale Col-
lege campus of the University of Toronto. The nests were attached to the
outer edge of the overhang, approximately 2 meters from the walls and 20
meters above the ground. Nest A was located on the southern side of the
building and nest B was on the northern side. The difference in occupancy
of the nests may have been due to the late season activity of the wasps.
Every sunny day during the late summer and fall large numbers of P. fus-
catus males and females were observed to gather on south facing walls of
the building in the afternoon. The male wasps attempted to mate with pass-

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ing females while the females spent much of their time investigating cracks
and crevices in the masonry, apparently searching for overwintering sites.
During the same period only an occasional wasp was observed on the shaded
north facing walls.

In order to investigate the thermal environment of an old D. arenaria nest
an intact nest that had been attached to a branch was obtained in the fall of
1979 and fitted with 2 thermistor probes. The probes were placed in the
sections of the nest occupied by the overwintering P. fuscatus in nest A:
between the central combs and in space between the roof of the nest and
the top comb. Two additional probes were placed near the nest (5 cm and
1 m) to monitor air temperatures. The entire apparatus was clamped to a
pole and, in early January of 1980, was securely attached to the overhang
of the roof at the original location of nest A. Data were collected for 10
days. Records of the ambient temperature, nest temperatures and wind ve-
locity were recorded in the early morning, midday, and late afternoon or
whenever the ambient temperature was unusually high or low.

The results showed that an intact nest did provide some protection. Dur-
ing the first three days the ambient temperature ranged between -8° and
+ 1°C and the nest temperature between -7.5° and +3°C. When there was
little wind (velocity < 2 km/hr) and the ambient temperature was changing
rapidly, nest temperatures 2.5°C warmer than the ambient temperature were
recorded. These differences occurred during the late afternoon and in the
midmorning. When the nest was shaded and the ambient temperature was
stable or changing slowly, and when wind velocities were approximately 5
km/hr or greater, nest temperatures were equivalent to the air temperature.
However, on clear days exposure to the sun could warm the interior of the
nest by approximately +2°. The space between the top comb and the roof
of the nest proved to be the most insulated area (i.e., experienced the least
change).

After the third day of observations a winter storm removed part of the
nest envelopes and partially exposed the combs on one side. The damaged
test nest now was in approximately the same condition as nest A when it
was occupied by overwintering P. fuscatus in January of 1979. As would
be expected, the thermal regime of the test nest changed. During the next
six days the weather was unsettled and measurements were made under a
variety of combinations of wind, cloud cover, and temperature. The ambient
temperature ranged between — 9.5°C to +2°C, and the nest temperature
ranged from -9°C to +7°C. In cloudy periods and at night the nest tem-
perature was always within 0.5°C of the ambient temperature. During the
midday when the nest was exposed to full sun it apparently trapped heat, and
nest temperatures 5°C higher than the ambient were recorded, even on days
when the wind velocity was approximately 15 km/hr.

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149

The nest temperature records indicate that under certain weather condi-
tions an intact D. arenaria nest can moderate the ambient temperatures by
2.5°C. This difference may occasionally be significant for P.fuscatus sur-
vival, particularly when the ambient temperature drops a few degrees below
lethal levels for a relatively short period (e.g., before dawn). Although many
overwintering P. fuscatus are very tolerant of low temperatures, and can
survive at least 48 hours at -20°C, they are killed by exposure to 48 hours
at -25°C (Gibo 1972, 1976). Temperatures approaching or falling within the
lethal range usually occur in southern Ontario each year (Gibo 1972, 1976).
Since nightly lows of - 17°C had already been recorded at the College prior
to the time the wasps in nests A and B were collected in January of 1979,
the wasps in nest A had probably experienced these temperatures with little
or no moderation.

A reduction of the amplitude of the daily temperature fluctuations only
occurred within the intact nest and was restricted to overcast days with
little or no wind. On sunny days the test nest had approximately the same
or a greater daily temperature fluctuation than the ambient temperature.
Exposure to the sun caused even greater fluctuations when the test nest
was damaged. Because survival of P. fuscatus was high in a damaged nest
exposed to the sun, the amplitude of these fluctuations does not appear very
important to the wasps. This assertion is supported by other studies that
show that overwintering P. fuscatus are capable of withstanding frequent
cycles of freezing and thawing (Gibo et al. 1976).

In summary, a relatively modest moderation of the winter thermal regime
can be obtained by Polistes that overwinter in old D. arenaria nests. This
moderation occurs under specific conditions: an undamaged nest, little or
no wind, no exposure to sun, and a rapidly changing ambient temperature.
When any of these conditions are not met the nest temperature equals, or
in some cases exceeds the ambient temperatures. Since the lowest temper-
atures usually occur at night, and often persist for several hours, little or no
thermal moderation should be experienced by P. fuscatus overwintering in
old D. arenaria nests during these critical periods. Possibly the abandoned
nests offer their advantages to overwintering wasps. In addition to shelter
from rain, snow and ice storms, the nests provide partial protection from
the drying effects of the wind. The temperature data indicated that even
damaged nests can provide this protection. Humidity is another consider-
ation. When ambient temperatures are above 0°C, nests which have been
recently soaked by rain would maintain high humidities. Whether this effect
would persist at temperatures below 0°C remains to be determined. In any
case, the study offers further evidence that overwintering P. fuscatus are
sufficiently cold hardy to survive nearly complete exposure to the midwinter
temperature regime of southern Ontario.

150

NEW YORK ENTOMOLOGICAL SOCIETY

Acknowledgments

I would like to thank Joan Simmons for help in collecting nests A and B
and Shelly Chaiken for help in setting the test nest in place on the roof. This
research was supported in part by National Research Council Grant number
A5991.

Literature Cited

Bohart, G. E. 1942. Notes on some feeding and hibernation habits of California Polistes
(Hymenoptera: Vespidae). Pan-Pac. Ent. 43(1):30.

Gibo, D. L. 1972. Hibernation sites and temperature tolerance of two species of Vespula and
one species of Polistes (Hymenoptera: Vespidae). J. N.Y. Ent. Soc. 80(2): 105-108.

. 1976. Cold-hardiness in fall and winter adults of the social wasp Polistes fuscatus

(Hymenoptera: Vespidae) in southern Ontario. Can. Ent. 108:801-806.

, R. M. Yaraschavitch and H. E. Dew. 1974a. Thermoregulation of colonies of Vespula

arenaria and Vespula maculata (Hymenoptera: Vespidae) under normal conditions and
under cold stress. Can. Ent. 106:503-507.

, H. E. Dew and A. S. Hajduk. 1974b. Thermoregulation in colonies of Vespula ar-
enaria and Vespula maculata (Hymenoptera: Vespidae). II. The relation between col-
ony biomass and colony production. Can. Ent. 106:873-879.

, A. Temporale, T. P. Lamarre, B. M. Soutar and H. E. Dew. 1976. Thermoregulation

in colonies of Vespula arenaria and Vespula maculata (Hymenoptera: Vespidae). III.
Heat production in queen nests. Can. Ent. 109:615-620. (1977).

Green, A., R. D. Akre and P. Landolt. 1976. The aerial yellowjacket, Dolichovespula arenaria
(Fab.): Nesting biology, reproductive production and behavior (Hymenoptera: Vespi-
dae). Melanderia 26:1034.

Hermann, H. R., D. Gerling and T. F. Dirks. 1974. The cohibemation and mating activity of
five polistine wasp species. J. Ga. Entomol. Soc. 9:203-204.

Rau, P. 1930. The behavior of hibernating Polistes wasps. Ann. Ent. Soc. Amer. 23(3):441-
446.

Snelling, R. C. 1954. Notes on nesting and hibernation of Polistes (Hymenoptera: Vespidae).
Pan-Pac. Ent. 28(4): 177.

West-Eberhard, M. J. 1969. The social biology of polistine wasps. Misc. Publ. Mus. Zool.
Univ. Mich. 140:1-101.

Zabriskie, J. L. 1894. Notes on some parasites of Vespa. J. N.Y. Ent. Soc. 2:81-86.

Department of Zoology, Erindale College, University of Toronto, Missis-
sauga, Ontario L5L 1C6, Canada

Received for publication February 7, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(2), 1980, pp. 151-163

POPULATION TRENDS OF THE ALFALFA WEEVIL
(COLEOPTERA: CURCULIONIDAE) AND ITS ASSOCIATED
PARASITES IN MARYLAND AND NEW JERSEY, 1966-1970

Robert F. W. Schroder and William W. Metterhouse

Abstract. — Populations of the alfalfa weevil, Hypera postica (Gyllenhal),
in Maryland and New Jersey and of associated parasites changed greatly in
the period 1966-70. The alfalfa weevil declined rapidly during the period
under the increasing pressures of the introduced parasites Bathyplectes cur-
culionis (Thomson), Tetrastichus incertus (Ratzeburg), Microctonus cole si
Drea, Microctonus aethiopoides Loan and Bathyplectes anurus (Thomson).

The biology and ecology of the alfalfa weevil, Hypera postica (Gyllenhal),
and of the associated parasites in Maryland and New Jersey were reported
for the period 1961 to 1967 by Blickenstaff et al. (1972). This study included
details concerning the life history, habits, and movement of the weevil, the
importance of fall-laid eggs, the number of generations/year, the levels of
populations, the establishment, spread and percent parasitism of introduced
parasites, and the results of surveys of fall and winter populations of adults
done so investigators could predict larval populations and damage. Much
other work done on the biology and life history of the alfalfa weevil was
listed by Cothran (1966).

The population surveys of the alfalfa weevil begun by Blickenstaff and
Huggans (Blickenstaff et al. 1972) in 10 fields in Maryland and New Jersey
in 1964 and 1965 were continued and are reported here for the period from
fall 1966 to spring 1970. The fields in Maryland and New Jersey that were
surveyed from fall 1966 through March 1968 were the same as fields sur-
veyed by Blickenstaff et al. (1972). Thus, Maryland fields 1 and 2 were in
Howard County, near Clarksville; fields 3, 4, 5, and 6 were at Belts ville,
Prince George’s County; field 7 was in Howard County near Jessup; fields
8 and 9 were at Crowns ville, Anne Arundel County; and field 10 was in
Montgomery County near Rockville. In New Jersey, the survey fields es-
tablished by Blickenstaff in cooperation with the New Jersey Department
of Agriculture were: fields 1 and 2 in Sussex and Warren Counties in the
upper Appalachian Valley; field 3 in Warren County in the Lower Appala-
chian Highlands; fields 4 and 5 in Hunterdon and Mercer Counties in the
Lower Appalachian Piedmont area; and fields 6-10 in Monmouth, Burling-
ton, Gloucester, and Cumberland Counties in the Inner Coastal Plain area.
The single major change in field locations was made in March 1968 when 8
out of the 10 Maryland fields were dropped and 8 new fields were chosen.

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NEW YORK ENTOMOLOGICAL SOCIETY

This change was made to provide a more complete evaluation of weevil
populations in the major alfalfa producing areas of central and western
Maryland. Two of the original 10 fields (field 9 at Clarksville and field 10 at
Belts ville) were retained for comparison. The new locations were: fields 1
and 2 at Goshen and Boyds, Montgomery County; fields 3 and 6 at Frederick
and Thurmont, Frederick County; fields 4 and 5 at Boonesboro and Smith-
burg, Washington County; and fields 7 and 8 at Westminster and Sykesville,
Carroll County.

To maintain uniformity in sampling and analysis of the data collected, we
determined populations of larvae from April through July by examining plant
material (foliage) from 1- or (4-ft2 samples or removed adults from ground
samples during the winter season with a suction device. However, in March
1968 the number of samples (Vi ft2) for adult alfalfa weevil collections was
increased from 10 to 20 per field. The sampling of insects in the field and
the processing of samples in the laboratory were generally the same as those
reported by Blickenstaff et al. (1972), and where modifications were intro-
duced, they are noted.

Until 1968, adult alfalfa weevils were collected October through March
from each field with a D-Vac suction machine designed and developed by
Dietrick (1961). However, when more extensive field surveys were under-
taken in 1968, the machine was mounted on wheels to reduce the time, labor
and fatigue involved in sampling (Schroder 1970). The adults that were col-
lected with the vacuum were separated from debris by sieving and then
collected by forceps and counted.

Collections of larvae were made by cutting ten !^>-ft2 foliage samples at
ground level and placing the foliage with the insects in paper bags. The
samples from the 10 New Jersey alfalfa fields were collected by personnel
of the New Jersey Department of Agriculture and then processed at Belts-
ville, Maryland. The samples from the 10 Maryland alfalfa fields were col-
lected by personnel of USDA and processed at Beltsville. A maximum of
50 3rd-4th instar larvae were removed from each of the foliage samples and
placed with a fresh bouquet of alfalfa in a container. The container consisted
of a 6 in. glass tube (2" diam.) sealed at one end with parachute cloth and
a cork with a vial inserted in the center at the other end. The alfalfa was
placed in the vial and sealed with a sponge. This provided a cage for the
larvae to develop to adults. The cages were checked daily for the emergence
of parasites from the larvae. Additional collections of larvae were made by
sweeping the alfalfa at times when low numbers of larvae were present.
Foliage samples were then processed by using a high-speed blender-water
extraction technique (Schroder 1974) instead of the alcohol-kerosene meth-
od of Blickenstaff and Huggans (1969). By the new procedure, the 1st to
4th instar larvae were dislodged from the alfalfa in a high speed blender and
then separated from the debris by washing through a series of sieves. The

VOLUME LXXXVIII, NUMBER 2

153

number of larva e/Vi ft2 was then determined by totaling the number obtained
for recovery of parasites and the number recovered by extraction. Then the
number of larvae/ft2 for each field could be calculated and recorded.

Microctonus cole si Drea (Braconidae) and M. aethiopoides Loan (Bra-
conidae), parasites of adult alfalfa weevils, are difficult to recover because
of the diapause of the host and the diapause of the parasite larvae in the
host. Neal et al. (1971) demonstrated that topical application of the juvenile
hormone cis-trans and trans-trans 10,11 epoxyfarnesenic acid methyl ester
brought forth larvae of both Microctonus parasites. Once the parasite larvae
emerge from the host and if adult Microctonus did not emerge, species
identification was a problem. There were no dependable external body char-
acters available for use in distinguishing the larvae of these parasites until
Fuester (1970) separated them by the size of their mandibles.

There were three methods used to determine parasitism of adults: 1) hold
the weevil adults on fresh bouquets of alfalfa in small cages until the para-
sites emerge, 2) dissect adults, and 3) treat with synthetic juvenile hormone.
Depending on the availability of the hormone, it was applied topically on
the venter of the abdomen with 100 fx g of the hormone in 0.5 ^liter acetone
with a microinjector. The weevils were then held for emergence of parasites
for 30 days and then dissected to determine presence of parasite larvae.

Alfalfa Weevil Adult Populations

1966- 1967

Surveys for alfalfa weevil adults begun in September 1966 in Belts ville,
Maryland signaled that the adults did not start to return to alfalfa fields until
early October and reached a peak, a high of 6.7 adults/ft2, in mid-November
(Table 1). Numbers then declined by 67% to 2.3/ft2 in mid-March 1967. A
similar decline was apparent in New Jersey fields, although only 2 collec-
tions were made. The fewer collections made in New Jersey make it difficult
to pinpoint the peak population for this area as accurately as for Maryland,
but we feel that the figures in Table 1 provide a valid estimate of the levels.
Thus, populations in New Jersey were approximately Vi those in Maryland,
but the declines (63%) from mid-November to mid-March were very similar.
The same approximate 2 to 1 difference in adult numbers in Maryland vs.
New Jersey and the % decline in adults was reported by Blickenstaff et al.
(1972) for the winters of 1964-66.

1967- 1968

Regular collections were put off until October 30, 1967, in Maryland be-
cause earlier collections in Beltsville were not successful. The totals showed
fewer alfalfa weevils than in 1966 and a later return of the adults to the fields

154

NEW YORK ENTOMOLOGICAL SOCIETY

Table 1. Alfalfa weevil adults/ft2 in 10 Maryland and 10 New Jersey survey fields. 1966-
19701.

Date

Maryland

1966-1967

New Jersey

9/23

.02

10/20

2.86

—

11/14

6.66

—

11/17

—

3.54

12/12

4.74

—

1/17

2.92

—

2/27

1.30

—

3/16

2.26

—

4/4

1967-19681 2

1.32

10/30

1.24

11/21

—

1.36

11/20

1.68

—

12/19

2.26

—

3/25

1.63

—

3/27

1968-1969

.88

11/22

.51

—

12/20

.29

—

1/16

—

.18

1/27

.16

—

2/19

.15

—

3/11

.14

—

4/8

—

.26

4/11

.76

—

4/19

.45

—

4/24

.41

1969-1970

10/14

.43

—

10/22

.51

—

10/30

.51

—

11/24

.96

—

12/3

.48

—

12/11

.77

—

2/27/70

.11

—

3/10

.23

—

3/26

.21

1.4

4/29

.66

—

1 Average for 10 fields, 10 Vi ft2 samples/fields. — = no samples taken that date.

2 Sample size for adult collections was increased from 10 to 20 Vi ft2 sample s/field. Change
effective from 3/25/68^1/29/70.

VOLUME LXXXVIII, NUMBER 2

155

(Table 1). Populations peaked at 2.3/ft2 in mid-December and declined to
1.6/ft2 by late March, a reduction of only 30%. In New Jersey, populations
were again about Vi those in Maryland and the overwinter reduction was
35%. This was the first year since the surveys were begun in 1964 that the
decline in numbers of adults from fall to spring was less than 60%. This
sharp reduction in measurable overwintering mortality plus the late return
of adults in the fall of 1967 suggests that fewer adults returned to the fields
in the fall, but that larger numbers than normal returned in the spring. Such
spring migrations of adults had not previously been considered important
in the Maryland-New Jersey area, but are a common occurrence in New
York (Poinar and Gyrisco 1962).

1968- 1969

Collections of adults in both states in the fall of 1968 showed greatly
reduced populations and much later appearance of adults, confirmation of
the trend reported for 1967 (Table 1). As in 1967, this late return was signaled
by failure to collect them in numerous alfalfa fields in the Beltsville area.
For this reason, regular collections were not begun until November. Al-
though populations were low throughout the 1968-69 season, numbers of
adults were actually higher in the spring of 1969 than the previous fall,
another indication of change in the migration of the weevil. The trend was
obvious in both states though populations remained lower in New Jersey
than in Maryland.

1969- 1970

D-Vac sampling was begun October 14, 1969, when the first adults were
recovered by sweeping. Once again, populations in Maryland were rela-
tively low (Table 1), and the decline in overwintering Maryland adults was
lower than in the 1966-67 season. In New Jersey, sampling was not done
during the fall season; however, the samples taken in March 1970 showed
an average 1.4 adults/ft2, an increase over the average for the spring of 1968
and 1969.

Alfalfa Weevil Larval Populations

The mean numbers of alfalfa weevil larvae/ft2 for the 10 Maryland and 10
New Jersey alfalfa fields (calculated from the combined means of the larvae/
ft2 in individual fields) are shown in Figures 1 and 2. In 1967, larval popu-
lations peaked earlier than anticipated in both states and reached the highest
levels seen since the surveys were begun in 1964, 370 and 265 for Maryland
and New Jersey, respectively (Figs. 1 and 2). Populations in both states
remained above the 100 larvae/ft2 level for over 6 weeks. The New Jersey
peak occurred approximately 3 weeks later than the Maryland peak.

156

NEW YORK ENTOMOLOGICAL SOCIETY

APRIL

MAY

JUKE

JULY

Fig. 1. Maryland: Mean number alfalfa weevil larvae/ft2 per 10 fields ( ) and mean

percent parasitism of the larvae by B. curculionis ( ) and T. incertus (. . .), 1967-70. The

figures ABC represent the pooled standard deviations of the means.

In 1968 the larval buildup peaked almost one month later than in previous
years in Maryland and 2 weeks later in New Jersey. In addition, the average
number (over 10 fields) for both states was 75% of that for 1967 (100% in
Maryland and 65% in New Jersey). In Maryland, populations remained at

VOLUME LXXXVIII, NUMBER 2

157

Fig. 2. New Jersey: Mean number alfalfa weevil larvae/ft2 per 10 fields ( ) and mean

percent parasitism of the larvae by B. curculionis ( ) and T. incertus (. . .), 1967-69. The

figures ABC represent the pooled standard deviations of the means.

the peak level for about 2 weeks, which created a plateau rather than a
sharp peak as in previous years (Figs. 1 and 2).

Alfalfa weevil larval populations in 1969 were lower in both states than
in 1968. The peak in Maryland occurred about 1 week later than in 1968;
the peak in New Jersey occurred about 1 week later than in Maryland. A
comparison of the mean peak larval populations of the 10 fields in both
states from 1967 to 1970 showed a 86 and 92% reduction in numbers of
larvae/ft2 in Maryland and New Jersey, respectively (Table 2).

In 1970 the larval populations declined to the lowest level since the sur-
veys were begun (Figs. 1 and 2).

Thus, since 1967, the number of larvae collected in field 10 in Maryland,
one of the two fields sampled all 4 years, has declined from a peak of 980

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NEW YORK ENTOMOLOGICAL SOCIETY

Table 2. Peak number of alfalfa weevil larvae for each of the 10 fields in Maryland and
New Jersey, 1967-1970.

Peak number larvae/ft2

New Jersey Maryland

Field

no.

1967

1968

1969

1970

1967

19681

1969

1970

1

157

62

21

45

316

41

77

27

2

168

76

18

10

531

231

46

71

3

133

27

35

35

557

45

39

12

4

401

93

21

18

585

129

140

38

5

351

45

30

9

544

334

161

58

6

194

55

15

17

147

97

61

28

7

124

103

12

7

541

65

14

63

8

532

211

32

16

169

96

49

31

9

813

216

7

17

320

84

42

26

10

593

67

81

20

980

86

49

29

X

346.6

95.5

27.2

19.4

469.0

120.8

67.8

38.3

1 Fields 2 and 10 in

Maryland sampled again in

1969; all others

in 1969

were new

locations.

to 29 larvae/ft2. The overall decline was 92% for all 10 fields (Table 2).
Nevertheless, there were some areas that had severe larval damage. In New
Jersey, larval populations also declined by 92% and so have remained at
about Vi the Maryland levels.

Correlations Between Adult and Larval Populations

Blickenstaff et al. (1972) indicated that correlation of adult alfalfa weevil
populations collected in November and March with the peaks of larval pop-
ulations that followed were helpful in predicting potential damage to alfalfa.
It was one of the objectives of continuing the two-state surveys to predict
potential damage from such correlations. The relationship between these
adult numbers and the peak larval populations in individual fields is shown
in Table 3. Significant correlations were obtained for Maryland fields in
1966-67 except for adults collected in February. Also, significant correla-
tions were obtained for New Jersey fields for November 1966 and March
1968. Likewise, in 1968-69, significant correlations were obtained between
Maryland adults collected in January and March and peak spring larval
numbers and between New Jersey adults collected during April 1969 and
the peak in spring larvae. Correlations were not obtained between Maryland
adults collected in 1969 or 1970, and the 1970 peak larval populations and
correlations were not available for the 1969-70 season in New Jersey be-
cause samples were not collected. The change in behavior of the overwin-
tering adults in Maryland identified from the sampling is evident in the lack
of the uniform correlation between adults and peak larval populations dem-

VOLUME LXXXVIII, NUMBER 2

159

Table 3. Correlations between winter collected adult alfalfa weevil populations and peak
larval populations the following spring.

Adults

collected

1966-67

1967-68

1968-69

1969-70

Md.

N.J.

Md.

N.J.

Md.

N.J.

Md.

Oct.

.19

Nov.

85**1

.73*

.35

.05

.57

.21

Dec.

.73*

.18

.14

Jan.

.66*

.72*

Feb.

.59

.32

March

.76**

.52

.37

.67*

98**

.31

April

.51

.88**

.20

1 ** Indicates significance at 1%. * Indicates significance at 5%.

onstrated by Blickenstaff et al. (1972). However, the significance of the
correlations (Table 3), despite this change, support the idea that the number
of overwintering adults can be helpful in predicting potential larval popu-
lations.

Parasites of the Alfalfa Weevil

The history of releases and recoveries of the alfalfa weevil parasites in
Maryland and New Jersey was reviewed by Blickenstaff et al. (1972) and
Dysart and Day (1976). It was the main objective of the continuing surveys
to determine the incidence of and percent parasitism by alfalfa weevil par-
asites in the 10 alfalfa fields of Maryland and New Jersey.

Larval Parasites

As indicated, the alfalfa foliage in Vi ft2 at 10 or 20 sites in each field on
each sampling date was shaken to dislodge larvae and those in the 3rd and
4th stadia were held for parasite recovery. The supplemental surveys were
processed in the same manner so we would have additional larvae, if nec-
essary, to observe for parasite recovery. A maximum of 50 such larvae from
each sample were placed in containers with fresh alfalfa and reared to adults
in the laboratory at room temperature. The containers were examined week-
ly for emerging parasites.

Bathyplectes curculionis (Thomson) (Ichneumonidae), an introduced lar-
val parasite, was the predominant parasite of the alfalfa weevil collected in
the 10 Maryland and 10 New Jersey alfalfa fields from 1967 to 1970. In
Maryland, the highest level of parasitism for a single field was 64% on June
20, 1968, in New Jersey, the highest level recorded was 61% on June 19,
1968. The increased synchronization between populations of B. curculionis
and the peak population of alfalfa weevil larvae in Maryland and New Jersey

160

NEW YORK ENTOMOLOGICAL SOCIETY

through the 1969 season that is shown on Figures 1 and 2 suggests an in-
creased effectiveness of the parasite, especially in New Jersey. The mean
percent parasitism of alfalfa weevil larvae by B. curculionis shown in these
figures was calculated by using the means for percentage parasitism for the
individual fields.

Bathyplectes cinurus (Thomson) is another introduced larval parasite. It
was recovered in 1970, for the first time in these surveys, in New Jersey.
There it was parasitizing 30% of the larvae collected in field 7. The parasite
was not recovered from larvae collected in any of the 10 Maryland alfalfa
fields during the time of these surveys. This is not surprising since the first
record for recovery of B. anurus in Maryland was in Prince George’s Coun-
ty in 1972. The first recora for recovery in New Jersey was in Monmouth
County in 1967 (Dysart and Day 1976).

Tetrastichus incertus (Ratzeburg) (Eulophidae) is an introduced lar-
val parasite that becomes abundant in Maryland and New Jersey al-
falfa in late summer, generally after the first cutting when host density is
low; it therefore appears later than the Bathyplectes species. This parasite
has been established in Maryland since 1964; its spread and distribution in
the eastern United States, especially in Maryland and New Jersey, was
reported by Schroder et al. (1969). The incidence and mean percent para-
sitism of the alfalfa weevil by T. incertus in the 10 Maryland and 10 New
Jersey alfalfa fields for the period 1967-70 was calculated by using the means
for percentage parasitism for the individual fields and is shown in Figures
1 and 2. Plainly T. incertus parasitism increased very rapidly in the Mary-
land survey fields: it was 14% in 1967 and 73% a year later. Thus, it seems
to be an effective late season parasite of alfalfa weevil larvae in Maryland.
However, it was not recovered until September in New Jersey in 1967 and
then only in the southern 4 fields; in 1968 and 1969, it was found in only 2-
3 fields, and parasitism was less than 9%; and in 1970, it was recovered
from only 2 fields, and there levels of parasitism were less than 2%.

Adult Parasites

According to Brunson and Coles (1968) the adult parasite Microctonus
aethiopoides was firmly established by 1968 in about 100 square miles of
area in central New Jersey. It was introduced into Maryland in 1965 at
Beltsville but was not recovered through 1967 (Blickenstaff et al. 1972). The
first record for recovery of M. aethiopoides in Maryland was in Frederick
County in 1969 (Dysart and Day 1976). Microctonus colesi, another parasite
of adult alfalfa weevils, was recovered in low numbers from both Maryland
and New Jersey by the time of the reported surveys (Drea 1968).

In 1967, M. colesi adults were recovered from adult weevils collected in
5 of 10 Maryland fields where Microctonus parasitism ranged from 2 to

VOLUME LXXXVIII, NUMBER 2

161

50%. In New Jersey, in 1967, Microctonus spp. parasites were collected
from 4 fields, M. cole si from 2 (1 of the 6 adults and 1 of the 16 adults
collected, respectively), and M. aethiopoides from 2 (1 of the 6 adults and
1 of the 8 adults collected, respectively). Also, collections of larvae made
in spring of 1968 and held for adult eclosion showed M. cole si present in 3
of 10 New Jersey fields with 16.7% parasitized in field 10, 33.3% in field 7,
and 66.7% in field 6. One Microctonus sp. emerged from 10 adults collected
in field 3. In Maryland, collection of adults in the spring of 1968 showed
Microctonus parasitism in 9 of 10 fields, always at rates less than 5.95%.
However, when overwintering adult alfalfa weevils collected in Maryland
fields during the winter of 1968-69 were dissected to determine percent
parasitism by Microctonus spp., 5 of 10 fields contained parasitized adults,
and parasitism ranged from 18 to 66%. Also, 1, 3 and 3 Microctonus im-
mature parasites were recovered from adult weevils collected January 1969
from New Jersey fields 2, 8 and 9, respectively. In addition, M. colesi
emerged from adult weevils collected April 1969 in 3 fields. Parasitism
ranged from 20 to 33%.

In 1969, in Maryland, larvae were collected from fields in the spring
and reared to adults. Then 50 of these adults from each collection taken in
fields 3-8, and 10 were treated with the synthetic juvenile hormone to re-
cover M. colesi. All living adults were then dissected 30 days after treatment
to determine whether any more parasites were present in the host. M. colesi
emerged from alfalfa weevil adults collected from 6 of the 7 fields and par-
asitism ranged from 2 to 22% (including the 3 recovered by dissection); the
mean was 9.4%.

Microctonus aethiopoides emerged from adults collected in 1969 from
Maryland fields 6, 8 and 9. No parasites emerged from adults collected in
fields 1-5, and 10. The highest level of parasitism was 19.2% in field 6. In 1970,
collections made in the 10 Maryland survey fields in spring showed M. colesi
present in 8 fields and parasitism ranged from 12.5% in field 3 to 89.3% in
field 4. Parasites were not recovered from alfalfa weevil adults collected in
fields 7 and 10.

In New Jersey, alfalfa weevil adults were reared from larvae collected in
spring 1969. These weevils were held for recovery of parasites until August
19, 1969. On this date 50 adults from fields 3-8, and 10 were each treated
with 100 pg of the same synthetic hormone used previously. As a result,
M. colesi emerged from 2 to 16% of the adults collected from the 7 fields.

Summary and Conclusions

1. Adult alfalfa weevil population levels in New Jersey and Maryland de-
clined steadily from 1967 until spring 1969, but collections from Maryland
fields in fall 1969 showed a slight increase over the previous year. This

162

NEW YORK ENTOMOLOGICAL SOCIETY

increase was not reflected by an increase of larvae the following spring.
Adult population levels in New Jersey remained about lA to Vi lower
than population levels in Maryland.

2. Populations of alfalfa weevil larvae declined from a peak of nearly 1,000/
ft2 in 1967 to the lowest level recorded, less than 100/ft2, in 1970. Despite
this near 10-fold decline, there were isolated areas reporting serious dam-
age. Larval populations in New Jersey for the 4 year period remained
about Vi to V3 lower than larval populations in Maryland.

3. Previous to 1968, Maryland and New Jersey alfalfa fields had shown a
characteristic decline in adult weevils from fall to spring of approximately
66%; however, the decline was not evident from 1968 to 1970. This
change in behavior of the adult weevil was reflected in the lack of cor-
relation between overwintering adults and peak larval populations for
the three seasons, 1968-70. Nevertheless, correlations were obtained
between spring adult populations and the subsequent peak larval popu-
lations for the period 1966-69, except for the March 1967 collections in
Maryland. The conclusion of Blickenstaff et al. (1972) that the number
of adults per given time is helpful in predicting potential larval popula-
tions is therefore supported.

4. Bathyplectes curculionis was the predominant parasite of the alfalfa
weevils collected in all 10 Maryland and New Jersey alfalfa fields in 1967-
70. The increased synchronization of peak populations of B. curculionis
and the peak population of larvae in Maryland and New Jersey suggests an
increased effectiveness of the parasite during that period, especially in
New Jersey. The high levels of parasitism, from 64% in Maryland to 61%
in New Jersey, and this increased synchronization made this a very ef-
fective parasite.

5. Bathyplectes anurus was recovered in 1970, the first recovery in these
surveys, from one of the 10 alfalfa fields in New Jersey.

6. Parasitization by T. incertus built up very rapidly in Maryland fields from
a low of 14% parasitism in 1967 to a high of 73% a year later. The parasite
appears late in the season, generally after the first cutting and at a time
of low host density. T. incertus give indications of being an effective late
season parasite of alfalfa weevil larvae in Maryland. However, in New
Jersey, T. incertus practically disappeared in 1968-70. The low parasit-
ism levels may be a result of the much lower host density in New Jersey
alfalfa fields.

7. Microctonus colesi and M. aethiopoides, parasites of adult alfalfa weevils,
were difficult to recover because of the diapause of the host and the dia-
pause of the parasite larvae in the host. Topical application of synthetic
juvenile hormone to adult weevils induced early emergence of Microc-
tonus parasites. Species identification was a problem if adult Microctonus
did not emerge. However, based on the available data, both species are

VOLUME LXXXVIII, NUMBER 2

163

established and increasing in the survey areas of Maryland and New
Jersey.

Acknowledgments

Grateful thanks are extended to Wm. P. Dodson, Jr., Agricultural Research
Technician of the Beltsville laboratory, who rendered valuable aid through-
out the study in all sampling and rearing, and to members of the New Jersey
Department of Agriculture, who made sample collections for larval popu-
lations and assisted in taking samples for adults in New Jersey.

Literature Cited

Blickenstaff, C. C. and J. L. Huggans. 1969. Four methods of sampling to measure populations
of alfalfa weevil larvae. J. Econ. Entomol. 62(3):556-557.

, and R. F. W. Schroder. 1972. Biology and ecology of the alfalfa weevil,

Hypera postica, in Maryland and New Jersey, 1961 to 1967. Ann. Entomol. Soc. Amer.
65(2):336— 349.

Brunson, M. H. and L. W. Coles. 1968. The introduction, release, and recovery of parasites
of the alfalfa weevil in eastern United States. USDA Prod. Res. Rep. 101, 12 pp.
Cothran, W. P. 1966. A bibliography of the alfalfa weevil, Hypera postica. Bull. Entomol.
Soc. Amer. 12(2): 151-160.

Dietrick, E. J. 1961. An improved backpack motor fan for suction sampling of insect popu-
lations. J. Econ. Entomol. 54(2): 394-395.

Drea, J. J. 1968. A new species of Microctonus (Hymenoptera: Braconidae) parasitizing the
alfalfa weevil. Entomol. News 79(4):97-102.

Dysart, R. J. and W. H. Day. 1976. Release and recovery of introduced parasites of the alfalfa
weevil in eastern North America. USDA, Prod. Res. Rep. 167. 61 pp.

Fuester, R. W. 1970. Separation of first-stage larvae of 2 species of Microctonus (Hymenop-
tera: Braconidae) which attack the alfalfa weevil. Ann. Entomol. Soc. Amer. 63(6): 1777-
1778.

Neal, J. W., W. T. Hollaway and W. E. Bickley. 1971. Response of Microctonus aethiops
and M. colesi, parasites of the alfalfa weevil, to a mixture of cis-trans- and trans-trans-
10,1 1-epoxyfarnesenic acid methyl ester. J. Econ. Entomol. 64( 1):338— 339.

Poinar, G. O. and G. G. Gyrisco. 1962. Flight habits of the alfalfa weevil in New York. J.
Econ. Entomol. 55(2):265-266.

Schroder, R. F. W. 1970. A modified suction machine for sampling populations of alfalfa
weevils on alfalfa. J. Econ. Entomol. 63(4): 1329-1330.

. 1974. Separating alfalfa weevil larvae from alfalfa foliage samples. Proc. Entomol.

Soc. Wash. 76(3):312-314.

— , J. L. Huggans, D. S. Horn and G. T. York. 1969. Distribution and establishment of

Tetrastichus incertus in the eastern United States. Ann. Entomol. Soc. Amer. 62(4):812-
815.

Beneficial Insect Introduction Laboratory, Agricultural Research, Sci. &
Educ. Admin., USDA, Beltsville, Maryland 20705 and Division of Plant
Industry, New Jersey Department of Agriculture, Trenton, New Jersey
08025.

Received for publication March 28, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(2), 1980, pp. 164-165

BOOK REVIEW

Herbivores: Their Interaction with Secondary Plant Metabolites. Edited by

Gerald A. Rosenthal and Daniel H. Janzen. 1979. Academic Press, New

York. 718 pp. $59.50.

“Chemical ecology” is a maddening field. Almost devoid of theory (and
utterly dependent on the coarse verbal “ploy-counterploy” model, which
has changed little since proposed by G. Fraenkel in his classic paper “The
raison d’etre of secondary plant substances” 21 years ago), it consists large-
ly of a collection of more or less pretty anecdotes. Yet the prettiest anec-
dotes are always the new ones; whenever a story is developed in detail,
carried beyond the stage of descriptive natural history, its prettiness goes
away and the evolutionary and functional scenarios become clouded. A
good example is the phytoecdysones. There is indisputable appeal in the
idea of plants outfoxing insects by confounding their development with large
doses of hormone analogues. Yet most phytoecdysones that are ingested by
insects are degraded by the midgut epithelium and cannot “function” in this
way. In the “insect hormones” chapter of this new collection, Karel Slama,
who started this story going, sidesteps the degradation issue on page 693.
Reese tackles it head-on on pp. 323-324.

“Herbivores” — the title is misleading and may help sales for the wrong
reasons — is the latest attempt to integrate “chemical ecology.” It brings
together a great many pieces of the puzzle, but puzzle it remains. The field
clearly still suffers from confusion over the concept of “function” in an
evolutionary-biological context. The ecological and evolutionary chapters
still suffer lapses into the unwarranted assumption that “compound X
evolved in order to deter (poison) species Y” just because it is seen to do
so now. This is tantamount to saying the function of DBCP is to sterilize
workers in chemical plants!

Yet, full of hot air as they are — perhaps because they are — the ecological
and evolutionary chapters are where the fun is. For most biologists the
superb reviews of the major classes of secondary compounds are “chicken-
wire chemistry” potentially useful for reference. Janzen’ s chapter, “New
horizons in the biology of plant defenses,” is outrageous as usual — and
much to the point. “Herbivores do not eat Latin binomials,” he says, re-
minding that Gertrude Stein did not mean a rose is a rose is a rose bio-
chemically. Then he says “plants are anachronisms,” and tells about things
superbly adapted for seed dispersal by creatures which are extinct. (Phy-
logenetic inertia, the last refuge of scoundrels, is indeed untestable — and
probably true.) His last two subheadings are “pitfalls” and “one-liners.”
Perhaps his next overview will include “pratfalls” and “howlers,” both of
which certainly apply in this field.

VOLUME LXXXVIII, NUMBER 2

165

Among the various chapters, Chew and Rodman’s stands out as a cou-
rageous (and ultimately unsuccessful, but that isn’t important) attempt to
calculate the energetic costs to a plant of defending itself chemically. The
costs of defense have generated a lot of hot air, and this is the first attempt
to do something concrete with the idea. Chew and Rodman don’t succeed
because it just isn’t possible now to isolate that segment of the system from
the plant as a whole, just the same as ‘‘optimization theory” in ecology has
relied on extremely naive calculations which compartmentalize the time and
energy budgets of organisms in questionable ways. A great many evolu-
tionists act as if evolution can do anything. But the process of adaptation
is non-Markovian; it does matter where you have been; and the mere fact
that doing something is energy-inefficient means nothing about the oppor-
tunity of stopping. Perhaps lots of things are anachronisms, sensu Janzen.

Past work in this field has had a strong taxonomic bias; many ‘‘chemical
ecologists” acted as if all herbivores had six legs. This book has less of an
insect slant, but it still belongs on the shelf next to Keeler, VanKampen,
and James’ “Effects of Poisonous Plants on Livestock” (Academic Press,
1978) for the sake of balance. There are signs that phytopathologists, ento-
mologists, vertebrate biologists, and vegetation scientists are finally con-
verging on a common realization that the same compounds may have
multiple “raisons d’etre.” This is all to the good. Anyone who considers
him- or herself a “chemical ecologist” or “coevolutionary biologist” should
have and read “Herbivores”; Janzen’ s cautionary chapter should be re-
quired reading for anyone considering doing this for a living. The cost of
the book is ridiculous on its face, but decomposes to 8.30/page, which now-
adays ain’t all that bad.

The chemistry in the book is safe. Take the biology with a grain of salt
(which, of course, may be dangerous to your health).

Arthur M. Shapiro, Department of Zoology, University of California,
Davis.

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Journal of the

New York Entomological Society

VOLUME LXXXVIII SEPTEMBER 1980

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. Randall T. Schuh
American Museum of Natural History
Dr. Daniel Sullivan
Fordham University
Dr. Felix J. Bocchino
The College of Mt. Saint Vincent

CONTENTS

j Su Zan Noguchi Swain (Mrs. William Kenneth Firmage) Asher E. Treat 168-169

Observations on the nesting behavior of Tachytes tricinctus (F.) on San Salvador
Island, Bahamas (Hymenoptera: Sphecidae, Larrinae)

Nancy B. Elliott and Paul Salbert 170-173

Life history, morphology, and taxonomy of Acutalis tartarea (Say) (Homoptera:
Membracidae) James H. Tsai and Dennis D. Kopp 174-185

Ecology of necrophilous and carpophilous Coleoptera in a southern New York wood-
land. Part I — Overview Dominick J. Pirone and Daniel J. Sullivan, S.J. 186-196

\Platanthera cristata (Michx.) Lindl., a new host for the red-banded leaf roller

T. L. McCabe and C. J. Sheviak 197-198

! Evolutionary origin of bees (Hymenoptera: Apoidea) U. N. Lanham 199-209

Transmission of vibrations along plant stems: implications for insect communication

Paul D. Bell 210-216

Over-exploitation of larval host plants by the butterflies Heliconius cydno and Heli-
conius sapho (Lepidoptera: Nymphalidae: Heliconiinae: Heliconiini) in Costa
Rica? Allen M. Young 217-227

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(3), 1980, pp. 168-169

SU ZAN NOGUCHI SWAIN
(Mrs. William Kenneth Firmage)

Su Zan Noguchi, third of the five gifted daughters of Minosuke and Tomi
Ogawa Noguchi, was born in Colorado. She attended the University of
Colorado at Boulder, being graduated in 1938 with the degree of Bachelor
of Fine Arts. Even before her graduation she had illustrated various biolog-
ical publications for members of the university faculty.

While serving as Staff Artist for the School Nature League at the Amer-
ican Museum of Natural History she was married to Dr. Ralph B. Swain,
a well known entomologist in the United States Department of Agriculture,
by whom she later had two sons, Tom Alfred and Ralph Adrien. With Dr.
Swain she prepared and illustrated The Insect Guide (Doubleday 1948), one
of the finest and most successful of such popular works. Together, also,
they wrote and illustrated the section on insects in Compton s Pictured
Encyclopedia (now published by Encyclopedia Britannica).

In 1953, while returning with his family to duty with the U.S.D.A. in
Nicaragua, Dr. Swain was ambushed en route and killed by bandits. With
heroic courage and fortitude Su Zan with her two boys, then only ten and
twelve years old, made her way back to the United States “to put together
their shattered life.” In Chatham, New Jersey she resumed her career as
author and illustrator, producing in succession Insects in their World (1955),
Plants of Woodland and Wayside (1958), and First Guide to Insects (1964)
all published by Doubleday. She illustrated nature books and articles for
many other authors and publishers, and has had her work exhibited in mu-
seums and galleries throughout the country, most recently in The Five No-
guchi Sisters Exhibit in honor of Japanese- Americans, sponsored by the
Arts Council of Sterling, Colorado. Her two sons have both attained dis-
tinction, Tom as an herpetologist and Ralph in the field of mass communi-
cations.

Elected in 1953 to Honorary Life Membership, Su Zan has contributed
to the New York Entomological Society the cover design of its Journal as
well as a memorable seminar on the history of entomological illustration,
with examples from the rare book room of the American Museum of Natural
History Library, and spiced with refreshments in the form of various edible
insects. In the letter informing her of her election (four years earlier!) Dr.
E. S. Hodgson, then Secretary of the Society wrote: “We know of no other
person who has produced such consistently excellent drawings and paintings
of insects and other natural history subjects. We greatly admire the com-
bination of realism and scientific accuracy with sensitive artistry in your
work.”

In 1969 Su Zan was married to William Kenneth Firmage. Though nom-
inally retired, both she and her husband continue active work, Su Zan in

VOLUME LXXXVIII, NUMBER 3

169

Su Zan Swain Firmage at home in her study in Great Barrington, MA.

oils and wood carving. They travel widely, always returning to their summer
home in Massachusetts or to their winter home in Florida.

Asher E. Treat
Tyringham, Massachusetts

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(3), 1980, pp. 170-173

OBSERVATIONS ON THE NESTING BEHAVIOR OF TACHYTES
TRICINCTUS (F.) ON SAN SALVADOR ISLAND, BAHAMAS
(HYMENOPTERA: SPHECIDAE, LARRINAE)

Nancy B. Elliott and Paul Salbert

Abstract. — Nesting females of Tachytes tricinctus (F.) were observed on
San Salvador Island, Bahamas, during June, 1978. This paper reports the
first published data on nest construction and predatory behavior in this
species. Females captured acridid grasshoppers, Ophullela pelidna pelidna
Burmeister and Delia sp., which were carried to the nest in flight. The nests
were multicelled with one to five cells per nest; cells were located below
the level of the main burrow, and contained two to nine prey. Evidence is
presented that females of this species capture all the prey for a particular
cell before ovipositing on one of the prey.

Introduction

Individuals of the genus Tachytes are large active solitary wasps. Typi-
cally females of this genus dig multicellular nests surrounded by distinctive
circular mounds of soil (Evans and Kurczewski 1966). They provision these
nests with Orthoptera from various families.

Bohart and Menke (1976) included Tachytes tricinctus (F.) in the T. dis-
tinctus species group, and reported its distribution to be the West Indies.
In June, 1978, we observed a nesting aggregation of females on San Salvador
Island, the Bahamas. Our observations, reported here, represent the first
published information about nesting behavior in this species.

Results

We observed females nesting in a sparsely-vegetated area of hard-packed
sand at the edge of an abandoned field adjacent to the old naval base now
operated by the College Center of the Finger Lakes. While we did collect
males of the species on San Salvador during the period of the study, none
were seen at the study site. Twelve nests were observed in the area which
represented a narrow strip, approximately 10 by 50 meters. Other species
of digger wasps including Cerceris zonata Cresson and Prionyx thomae (F.)
also nested in the area as did several ground-nesting bees. Our observations
were made between 6 and 19 June 1978.

During collecting trips made to the island yearly in November through
December since 1975, we have collected many sphecids, but never T. tri-

VOLUME LXXXVIII, NUMBER 3

171

cine t us (Elliott et al. 1979). Therefore, we conclude this is a summer nesting
species on the island.

Nest construction. — We found several burrows in the process of construc-
tion, but did not observe their actual beginning. These burrows were closed
and were marked by large spoil heaps, indicating extensive digging within.
One nest, first observed at 1415 hours on June 15, was noted open the next
day at 828 hours.

Orientation. — A female about to leave her nest to hunt for prey, walked
out of the entry onto the spoil heap. She then took flight, hovering above
the nest, facing the entry and flying in successively wider and higher circles
around the nest. Orientation lasted a total of ten seconds. The female then
flew away to hunt, leaving the burrow open.

Predatory behavior. — Females were often away from the nests for several
hours hunting for prey. The prey, always acridid grasshoppers, were carried
to the nest in flight. Typically the burrow was open, and the female entered
directly carrying her prey. One female, returning to a nest that had acci-
dentally been covered in her absence, was seen holding her prey with all
three pairs of legs as she approached. Then, supporting herself with the
middle legs, and holding the prey with the hind legs, she used the mandibles
and forelegs to dig at the closed entrance. Eventually she released the prey
to dig, but soon retrieved it.

Another female, encountering a grasshopper inside an insect net, stung
it and prepared it for carriage. She grasped the antennae with her mandibles
and turned it sideways under her body, preparing it for transport. This
grasshopper, however, was larger than any we saw females carry to the
nest. It weighed 240 mg, and all prey weighed averaged only 93.8 mg. Hence
the mounting behavior observed may have been atypical.

Two species of acridids were used as prey. They were Ophulella pelidna
pelidna (Burmeister) and Delia sp. They ranged in weight from 36.3 mg to
214 mg (n = 13; x = 93.8 mg). Females capturing the prey ranged in weight
from 110 mg to 133 mg (n = 5;;t = 126.5). The mean ratio of weight of prey
to weight of wasp carrying it was .75 (n = 12; range = .33-1.61). We noted
a slight tendency for females to take larger grasshoppers later in the period
of our study, and this seemed to indicate that females simply captured those
individuals of suitable species available, thus taking larger grasshoppers as
the prey species matured. A similar tendency was observed in T. crass us
Patton by Evans and Kurczewski (1966).

Nest characteristics. — Eleven nests were excavated and reconstructed
using the methods described by Salbert and Elliott (1979). Table 1 sum-
marizes the important nest and cell characteristics we quantified. The entries
of relatively new nests were surrounded by circular mounds of soil. These
were abraded or completely absent at older nests. The shape of each entry
burrow depended on the substrate and might deviate from the vertical if

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NEW YORK ENTOMOLOGICAL SOCIETY

Table 1. Nest and cell characteristics of Tachytes tricinctus (F.).

Characteristic

Mean

Range

Number

Diameter of spoil heap

7.5 cm

6.5-10.5 cm

9

Height of spoil heap

3.3 cm

2. 7-3. 5 cm

4

Maximum depth of burrow

14.5 cm

2.5-19.0 cm

11

Depth of cells

20.3 cm

7-29.0 cm

42

Number cells/nest*

2.6

1-5

8

Length of cell

2.9 cm

1. 7-4.0 cm

8

Width of cell

1 .2 cm

0. 6-2.0 cm

7

Height of cell

1.2 cm

1 .0-1.5 cm

5

Prey/cell (when whole)

4.2

2-9

4

Length of cocoon (when present)

1.5 cm

1 .2—1 .7 cm

6

* An older nest which contained 21 cells, many of them already empty has not been included
in this measurement.

bypassing a rock. While many nests had nearly vertical burrows, one ran
vertically for about 1 1 cm; then it became nearly horizontal. The soil below
the level of this particular burrow was very rocky. The cells were located
at or below the level of the main burrow. With a single exception, all cells
had been closed off from the main burrow. One old nest, associated with
three entrance holes, contained 21 cells, many of them bearing evidence
that their occupants had already emerged. Newer nests contained from one
to five cells. The cells with whole prey contained from two to nine grass-
hoppers. All prey, for which placement could be determined, were placed
in the cell with the head inward, but varied as to whether they had been
placed venter up or on their sides.

Nest closure. — The females closed their nests at night and sometimes
during midday. A female preparing to leave the nest in the morning, re-
moved the sand closure with her mandibles. Possibly sand is pushed into
the entry to form the closure as a result of additional digging going on within
the nest.

Discussion

In general our observations on the nesting behavior of Tachytes tricinctus
agree with those made by Williams (1913) on its North American relative,
T. distinctus Smith, and those summarized by Bohart and Menke (1976) for
all species of the Tachytes distinctus species group. The females of these
species all prey on acridids, which are carried to the nest in flight, and
construct multicellular nests with one to several prey per cell.

A few reports indicate that females of Tachytes validus must capture all
the prey to be used in a particular cell before ovipositing (Evans and Kur-
czewski 1966; Kurczewski and Ginsburg 1971). Our observations indicate

VOLUME LXXXVIII, NUMBER 3

173

that this may also be the case for T. tricinctus. In one excavation we found
a single acridid in the burrow about 2.5 cm from the open cell. The cell
itself contained two more fresh grasshoppers, but no egg. We had captured
the female returning with prey late in the afternoon. Presumably the grass-
hopper in the burrow and the one she was carrying would also have been
placed in the open cell. Evans and Kurczewski (1966) reported that females
of T. validus stored prey for a particular cell in a separate pocket of the nest
prior to oviposition. Perhaps females of T. tricinctus leave prey in the bur-
row for the same reason.

Acknowledgments

We thank Dr. Donald Gerace, Director of the College Center of the Finger
Lakes Bahamian Campus and Mrs. Gerace for their assistance during our
study. This study was financed in part by a grant from the Hartwick College
Board of Trustees.

Dr. Ashley B. Gurney, U.S.D.A. Systematic Entomology Laboratory,
determined the prey.

Literature Cited

Bohart, R. M. and A. S. Menke. 1976. Sphecid Wasps of the World: a Generic Revision.
Univ. Calif. Press, Berkeley, pp. 695.

Elliott, N. B., F. E. Kurczewski, S. Claflin and P. Salbert. 1979. Preliminary annotated list
of wasps of San Salvador Island, the Bahamas with a new species of Cerceris (Hyme-
noptera: Tiphiidae, Scoliidae, Vespidae, Pompilidae, Sphecidae). Proc. Entomol. Soc.
Wash. 81:352-365.

Evans, H. E. and F. E. Kurczewski. 1966. Observations on the nesting behavior of some
species of Tachytes (Hymenoptera: Sphecidae, Larrinae). Jour. Kans. Entomol. Soc.
39:323-332.

Kurczewski, F. E. and S. E. Ginsburg. 1971. Nesting behavior of Tachytes ( Tachyplena )
validus. Jour. Kans. Entomol. Soc. 44:113-131.

Salbert, P. and N. Elliott. 1979. Observations on the nesting behavior of Cerceris watlingensis
(Hymenoptera: Sphecidae, Philanthinae). Ann. Entomol. Soc. Amer. 72:591-5.
Williams, F. X. 1913. The Larridae of Kansas. Univ. Kans. Sci. Bull. 8:121-213.

Department of Biology, Hartwick College, Oneonta, New York 13820.
Received for publication January 8, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(3), 1980, pp. 174-185

LIFE HISTORY, MORPHOLOGY, AND TAXONOMY OF ACUTALIS
TARTAREA (SAY) (HOMOPTERA: MEMBRACIDAE)1

James H. Tsai and Dennis D. Kopp

Abstract. — Acutalis tart are a (Say) is a membracid species found through-
out the northeastern 2A of North America. We present information on its
biology and field life history in Florida and draw comparisons with the
biology of Micrutalis malleifera (Fowler).

A discussion of the polymorphic forms of A. tartarea is presented along
with the reasoning for suppression of the names Acutalis semicrema (Say)
and Acutalis inornata (Ball), both of which are color polymorphs. The im-
mature stages of A. tartarea are described and illustrated.

Acutalis tartarea (Say) and closely related Micrutalis spp. were found in
rotary flight traps near and on the undergrowths of coconut plantings during
a study of homopterous insects associated with lethal yellowing (LY) dis-
ease of coconut palms in Florida (Tsai 1980). Simons and Coe (1958) found
the treehopper Micrutalis malleifera (Fowler) to be the vector of pseudo-
curly top virus (PCTV) in tomatoes grown in Florida. Their work provided
the first documented record of a treehopper vector of a virus disease.

In light of Simons and Coe’s work, A. tartarea was considered a possible
vector of LY due to its proximity to diseased trees. Since little was known
of its biology and life history the following study ensued. The purpose of
this paper is to report on the life history, morphology, and taxonomic status
of A. tartarea.

Materials and Methods

Since common ragweed ( Ambrosia artemisiifolia L.) was found to be a
natural host of A. tartarea in the field, it was the principal host plant in
laboratory rearings. Seedlings were transplanted to styrofoam pots ca. 5 cm
in diameter and held in a greenhouse equipped with an evaporative cooling
system that maintained a temperature at ca. 27°C. Relative humidity was
not controlled. China aster ( Callistephus chinensis (L.) Nees), eggplant
{Solarium melongena L.) and periwinkle {Catharanthus roseus (L.) G. Don)
also were used for life history studies in the laboratory, under the same
conditions as common ragweeds.

A. tartarea adults were collected in the Fort Lauderdale area to establish

Florida Agricultural Experiment Station Journal Series No. 2346.

VOLUME LXXXVIII, NUMBER 3

175

laboratory colonies. The stock colony of A. tart are a was reared on common
ragweed in a rearing room at 24 ± 1°C and R.H. 75 ± 5%. Light intensities
of 7,000 lux were provided by a bank of six fluorescent lights with a 12 hour
photophase and 12 hour scotophase. The same conditions were provided
for life cycle study.

In order to study nymphal development, host plants were modified by
trimming the excess foliage from the terminal tips and placing these trimmed
tips in rearing cages. Rearing cages consisted of 4.5 cm plastic petri dishes
with their sides notched to accept the terminal stem of the host plant. The
portion of the stems which passed through the notch in the petri dish was
wrapped with sponge rubber to form a tight seal and prevent escapes. Dark
colored paper was placed on the bottom of the dish to facilitate the obser-
vation of molted skins. Ventilation was provided by cutting a small hole in
the petri dish lid and glueing a fine screen over it. Daily observations were
made on each rearing cage throughout the duration of the study.

The morphological characters of each life stage were described and illus-
trated with the aid of a camera lucida mounted on a Wild® 5M dissecting
microscope at 25 x magnification. Measurements of nymphs in each instar
were made with a Bausch and Lomb Zoom 7 Stereoscope® and a calibrated
ocular micrometer.2

Results

Laboratory Life History Studies and Field Observations. — A. tart area
eggs hatch and the nymphs pass through 5 instars before moulting into the
adult. Adult females generally lay eggs in clusters of 12-15 eggs each. These
eggs are inserted into the epidermal tissue of the host plant which has been
slit by the female ovipositor. Eggs are inserted to a depth that leaves ca.
V3 of the egg exposed. Of the 82 plants examined, the favored oviposition
sites were in the axis area of the leaf (70%) and the stem terminus (27%).
Approximately 3% of the time, eggs were found completely exposed and
held by the dense epidermal hairs of the host plant. On common ragweed,
the number of eggs laid by a single female ranged from 12 to 62 and incu-
bation time ranged from 10-23 days with a mean of 15.7 ±3.2 (S.D.) days.
On aster incubation time was significantly longer (t = 2.72, df = 38, P =
0.01) (see Table 1).

Early instar nymphs exhibit gregarious, sedentary behavior near the ter-
minal portion of the ragweed plant. Early instars remain in the area of egg
hatch and, as nymphal development progresses, later instar nymphs dis-

2 ® The use of a trademark name is not an endorsement of one product over another that is
equally effective.

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Table 1. Developmental time (in days) of Acutalis tartarea (Say) reared on common rag-
weed ( Ambrosia artemisiifolia L.) and China aster ( Callistephus chinensis (L.) Nees at
24 ± rc.

Common ragweed China Aster

Life stage

Sample

size

Mean

Range

S.D.

Sample

size

Mean

Range

S.D.

Egg

24

15.7

10-23

±3.2

16

19.5

15-26

±3.1

1st Instar

22

4.5

3-8

±1.9

12

9.2

8-11

±0.9

2nd Instar

22

5.5

3-10

±1.5

11

9.6

7-12

±1.9

3rd Instar

22

4.5

3-7

±1.7

9

10.4

8-13

±1.6

4th Instar

17

6.6

4-11

±2.1

14*

12.0

10-15

±1.5

5th Instar

15

12.1

7-30

±3.1

13

13.0

10-16

±2.1

Total mean
development time

49.9

73.7

Adult longevity

Female

14

12.1

3-45

±4.1

9

32.0

8-55

±6.4

Male

21

7.9

1-30

±3.1

7

20.0

7-31

±5.6

* Due to the increasing mortality, insects were supplemented by individuals from the colony
reared on China aster. Their ages were determined on the basis of wing-pad development.

perse on the plant. Nymphal feeding occurs on the stems and petioles with
individuals maintaining a head down stance. No feeding was observed on
the leaves.

When reared on common ragweed, the mean nymphal development time
for the 5 nymphal instars was 34.2 days at 24 ± 1°C (see Table 1). The mean
time of nymphal development for instars 1 through 4 was between 4-6 days
each and that for the 5th instar was about twice that of the earlier instars
on common ragweed. Nymphs reared on aster have a longer developmental
time (t = 2.62, df = 155, P = 0.01) (see Table 1).

Laboratory reared male and female adults live significantly longer (t =
2.73, df = 33, P = 0.01; t = 2.98, df = 14, P = 0.01 respectively) on China
aster host than on common ragweed. On both hosts, female longevity is
greater (t = 2.83, df = 21, P = 0.01; t = 2.78, df = 26, P = 0.01 respec-
tively) than that of the male (see Table 1). Rearings were also attempted
using eggplant (Solatium melongena L.) and periwinkle ( Catharanthus ro-
seus (L.) G. Don) but A. tartarea could not survive on either of these hosts.

Field observations substantiated egg cluster size, oviposition site selec-
tion, nymphal behavior and nymphal feeding orientation. Adult flight activ-
ity in the field occurred after 9:00 a.m., this seemed to be related to tem-
perature and possibly light intensity. Greatest flight activity occurred during
the warmest portions of the day. The most common field host of A. tartarea
in Florida was common ragweed but the species was occasionally found on
Bidens bipinnata L. (Spanish needles).

VOLUME LXXXVIII, NUMBER 3

177

1

Fig. 1. Lateral view of adult male Acutalis tartarea (Say).

A monthly population survey was made from May 1978 to April 1979 by
visually counting all A. tartarea on 35 ragweed plants randomly selected in
the field (Fig. 8). The population was at very low levels from August through
November. During this period the growth of ragweed in the field was greatly
affected by environmental factors such as high temperature, dry months,
and long day length.

Morphology

Adult. — The adult A. tartarea (Fig. 1) possesses the typical characteris-
tics of Subfamily Smiliinae with its prothoracic tibiae not foliaceous, its
metathoracic tarsi equal in size to the prothoracic and mesothoracic tarsi,
three longitudinal veins in the tegmen arising near the base of the forewing
and five apical cells present in the forewing. The head is twice as broad as
long and the pronotum extends beyond the abdomen, but not covering the
tegmen. The colors of the head, pronotum tegmen and legs vary from green
to black. The adult female is 3.7 ± 0.07 mm (range: 3. 5-3. 9 mm) in length
and 1.9 ± 0.00 mm (range: 1.8- 1.9 mm) in width. The male is 3.3 ± 0.09
mm (range: 2. 8-3. 4 mm) long and 1.7 ± 0.03 mm (range: 1.5-1. 9 mm) wide.

Since the immature stages of this species have not been previously de-

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2

5

Figs. 2-5. 2. Side view of egg of Acutalis tartarea (Say). 3. Lateral view of 1st instar

nymph of A. tartarea. 4. Lateral view of 2nd instar nymph of A. tartarea. 5. Lateral view of
3rd instar nymph of A. tartarea.

VOLUME LXXXVIII, NUMBER 3

179

scribed or illustrated in the literature, their description and measurements
are presented below.

Egg (Fig. 2). — Mean length 1.0 ± 0.02 mm (range: 0.9-1. 1 mm), width
0.3 ± 0.00 mm (range: 0.2-0. 3 mm). Color white and translucent with one
end blunt and the other end pointed.

First Instar (Fig. 3). — Mean length 1.4 ± 0.03 mm (range: 1. 1-1.6 mm),
width 0.3 ± 0.00 mm (range: 0. 3-0.4 mm). Color pale green. Dorsum bearing
well developed paired spines, 1 pair anterior projecting from vertex of head,
2 pair anterior projecting from dorsum of prothorax, 1 pair anterior pro-
jecting from dorsum of mesothorax. One posterior projecting pair from dor-
sum of metathorax and posterior projecting pairs from dorsum of abdominal
segments 3, 4, 5, 6, 7, and 8.

Head: Clypeus triangular and hirsute; eyes prominent, projecting lat-

erally, pigmented red. Vertex bearing 1 pair of well developed anterior pro-
jecting spines; rostrum projected caudad between prothoracic legs to me-
tacoxae. Antenna located in front of eye, basal 3 segments enlarged and
terminal segments filamentous.

Thorax: Dorsum of prothorax bearing 2 pairs of anterior projecting

spines, one on either side of median line; 1 pair of anterior projecting spines
on dorsum of mesothorax; 1 pair of posterior projecting spines on dorsum
of metathorax. Pro, meso and metacoxae well developed; femora sparcely
hirsute; pro and mesotibiae laterally compressed and expanded with setae
on lateral edges; metatibiae Vs longer than pro and mesotibiae, only slightly
compressed; tarsi 2 segmented, tarsal claws swollen at base.

Abdomen: Dorsum of abdominal segments 1 and 2 unarmed, com-

pressed against mesothorax, segments 3-8 each armed with a pair of pos-
terior projecting dorsal spines, one on either side of median line; venter of
segments 1-8 convex and with prominent setae; segment 9 elongate, taper-
ing to apex, subequal in length to combined length of segments 5-8; segment
9 bearing a dorsal pair of posterior projecting spines, entire segment seta-
ceous.

Second Instar (Fig. 4). — Outline and shape similar to previous instar,
except in size. Mean length 1.8 ± 0.05 mm (range: 1. 6-2.1 mm), width 0.5
± 0.01 mm (range: 0.4-0. 7 mm). Color pale green. Paired dorsal spines on
either side of median line bearing secondary lateral setae.

Head: Similar to previous instar in outline and shape. Dense setae on

clypeus, frons and vertex.

Thorax: Paired dorsal spines with secondary setae on dorsal edge, tips

fuscous. All paired dorsal thoracic spines projecting anteriorly. Pro and
mesothoracic legs similar in form and size, tibiae laterally compressed,
dense setae on margins. Tibiae of metathoracic legs triangular in cross sec-
tion, IV2 times length of pro and mesothoracic tibiae.

Abdomen: Dorsum of abdominal segments 1 and 2 with short, prominent

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7

Figs. 6, 7. 6. Lateral view of 4th instar nymph of Acutalis tartarea (Say). 7. Lateral view

of 5th instar of A. tartarea.

VOLUME LXXXVIII, NUMBER 3

181

setae. Dorsum of segments 3, 4, 5, 6, 7 and 8 with prominent posteriorly
projecting spines, spines with secondary setae on dorsal margin. Abdomen
triangular in cross section, folding along line of junction of the lateral and
ventral plates. Segment 9 elongate, tapering to apex.

Third Instar (Fig. 5). — Outline and shape similar to previous instar except
in size and caudal edge of the meso and metathoracic lateral plates devel-
oped into wing buds. Mean length 2.6 ± 0.10 mm (range: 2.0-2. 8 mm),
width 0.7 ± 0.05 mm (range: 0.5-0. 8 mm). Color pale green. Paired dorsal
spines on head, thorax and abdomen with tips black. Venter of abdomen
concave.

Head: Frons densely covered with setae, caudal edge of clypeus forming

a ridge between eyes.

Thorax: Similar to previous instar except wing buds apparent on meso

and metathoracic segments. Mesothoracic wing buds covering V2 of meta-
thoracic wing buds. Metathoracic wing buds covering Vi of 3rd abdominal
segment.

Abdomen: Abdominal segments 1 and 2 covered by caudal edge of meta-

thoracic plate. Venter of abdominal segments concave. Segment 9 elongate,
tapering to apex.

Fourth Instar (Fig. 6). — Outline and shape similar to previous instar ex-
cept in size. Mean length 3.3 ± 0.15 mm (range: 2. 5-3. 9 mm), width 0.9 ±
0.04 mm (range: 0. 7-1.1 mm). Color pale green. Mesothoracic wing buds
extending onto 3rd abdominal segment. Metathoracic wing buds well de-
veloped and covered by mesothoracic wing buds. Paired dorsal spines prom-
inent, with black tips. Venter of abdomen strongly concave.

Head: Setae dense on clypeus and frons. Paired dorsal spines on vertex

smaller than thoracic or abdominal dorsal spines.

Thorax: Prothoracic dorsal sclerite extending caudo-mesad over V2 of

metathoracic dorsum between paired dorsal spines. Mesothoracic wing buds
covering metathoracic wing buds and extending onto 3rd abdominal seg-
ment.

Abdomen: Venter of abdomen strongly concave. Segment 9 elongate,

tapering to apex and subequal to the combined length of segments 2-8.

Fifth Instar (Fig. 7). — Outline and shape similar to previous instar except
in size. Mean length 5.7 ± 0.21 mm (range: 3. 3-7. 7 mm), width 1.3 ± 0.10
mm (range 0.8-1. 8 mm). Color pale green. Body setae dense. Paired dorsal
spines on head, thorax and abdomen with prominent black tips. Venter of
abdomen strongly concave.

Head: Setae dense on clypeus and frons. Paired dorsal spines on vertex

Vs to V2 length of thoracic and abdominal dorsal spines.

Thorax: Prothoracic dorsal sclerite extending between paired dorsal

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spines of mesothorax. Mesothoracic wing buds covering metathoracic wing
buds and extending caudally onto 3rd abdominal segment.

Abdomen: Venter strongly concave.

Taxonomy

In America, north of Mexico, the genus Acutalis historically has had
three described species: A. tart are a (Say), Acutalis semicrema (Say) and
Acutalis inornata (Ball). Differentiation of these three species was based
upon coloration of the pronotum and pigmentation of the venation of the
tegmen (Van Duzee 1908). A. tart area was the darkest in coloration and
this traditionally has been applied to specimens collected throughout the
Northeast, North Central and West Central portions of North America. A.
semicrema, originally described from Florida material, was not as darkly
pigmented as A. tartarea. More recent distribution records show the range
of A. semicrema to be very similar to that of A. tartarea. A. inornata,
described by Ball (1905) from Florida, is entirely green in color with no
black markings. Most distribution records of A. inornata have been from
Florida. However, intensive collecting in more northern states has produced
specimens that would be classified as A. inornata (Kopp and Yonke 1973).

As early as Matausch (1912) field observations were casting doubt upon
the species status of A. tartarea and A. semicrema. His observations in-
dicated that many of the females in his New Jersey collecting site were of
the A. semicrema form while the males were of the A. tartarea form. This
also was observed by the authors, who view this coloration pattern as a
form of sexual dimorphism.

Caldwell (1949) studied the genitalia of the three described species of
Acutalis and could not find distinctive differences. He treated these three
forms as trinomials, Acutalis tartarea tartarea (Say), Acutalis tartarea sem-
icrema (Say) and Acutalis tartarea inornata (Ball). From Caldwell’s treat-
ment it is apparent he used the trinomial to designate color varieties of the
same species, not, as today, to denote subspecies.

We feel that Caldwell’s treatment of these three species as one is entirely
justified. We view A. tartarea as a single species that in the adult stage
presents several different polymorphic color forms to the environment. In-
tensive collecting of local populations in Missouri and Florida has produced
representatives of all three forms.

Field observations of populations in Florida offer another line of evidence
for the acceptance of the single species with several color morphs. In Flor-
ida, where the three morphs can be most frequently encountered, mating
between the forms has been observed from February to June. These matings
in the natural habitat are strong evidence indicating a lack of reproductive
isolation necessary for species consideration of each form. No infertility

VOLUME LXXXVIII, NUMBER 3

183

was observed in the laboratory studies in which 9 pairs of green and black
forms were crossed. Of the 152 individuals produced, 40 of the green forms
were found to be female, and 86 of the black- headed forms were males.

The following listed synonymy is presented for the species Acutalis tar-
tare a (Say).

Micrutalis tartarea Say, 1830:242:1.

Micrutalis semicrea Say, 1830:242:2.

Acutalis anticonigra Fairmaire, 1846:498.

Tragopa brunnea Provancher, 1872:320.

Acutalis inornata Ball, 1905:119.

Acutalis tartarea semicrema Van Duzee, 1917:529.

Acutalis tartarea tartarea Severin, 1927:33.

Acutalis tartarea inornata Caldwell, 1949:498.

Acutalis tartarea semicrema Caldwell, 1949:498.

Discussion

A. tartarea has a broad range and presents a variety of color polymorphs
within populations. Three of these polymorphs have been described as
species, traditionally separated from each other by their colors and patterns
of coloration. Matausch (1912) doubted the validity of species status for
both A. tartarea and A. semicrema based on field observations in New
Jersey. Caldwell’s (1949) studies on the insect’s genitalia indicated that there
was a single species of Acutalis in America, north of Mexico, and he listed
the polymorphs as varieties of A. tartarea.

Field observations by the present authors indicate that many populations
had 2 or more polymorphs present which freely interbreed in nature, thus
supporting the observations of Matausch (1912). Laboratory rearings were
accomplished with several polymorphs and no abnormal infertility was ob-
served.

We view A. tartarea as a single species with color polymorphs throughout
its range. These morphs show different proportions in different geographic
areas. The biological mechanism of expression of these polymorphs is not
understood but may be a complex interrelationship of ecotypical color
forms, temperature related color expression, sexual dimorphism, and/or a
nymphal host plant relationship. Further autecological studies will be re-
quired to fully understand these forms.

Our observations on the life history of A. tartarea shows similarities to
the life history of M. malleifera as observed by Simons (1962). These sim-
ilarities are: mean incubation time of the eggs, mean development time of
instars 1 to 3, presence of adults in the field year round in Florida, gregarious
behavior of early instar nymphs, head down feeding stance of the nymphs

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8

Fig. 8. Monthly total numbers of adults and nymphs of Acutalis tartarea (Say) found on
common ragweed.

and highest reproductive activity during the active growing seasons of their
respective hosts.

A. tartarea has 5 nymphal instars with a total mean development time
from egg to adult of about 50 days. This is about 25% longer than M.
malleifera. Simons (1962) indicated M. malleifera had 4 nymphal instars,
which is remarkably unusual since all other treehoppers, whose biology
have been studies, have 5 nymphal instars. The difference may have been
due to rearing hosts (Simons, personal communication).

In Florida populations of both A. tartarea and M. malleifera show fluc-
tuation trends through the season. Both species are most active and reach
highest populations during the warmest months. Lowest field populations
of M. malleifera occur from October through April (Simons 1962). Popu-
lations of A. tartarea show rapid decreases during July, reaching a low
during the months of August through November (Fig. 8). Seasonal popu-
lation fluctuations of both species are related to the seasonal growth cycle
of their host plants.

The favored field host of A. tartarea in Florida is common ragweed.
Laboratory rearings on this host were quite successful. China aster was also
used as a laboratory rearing host for A. tartarea. The species could be
continuously reared on this host but individuals reared on it showed longer
development time than those reared on common ragweed (see Table 1). It
was also noted that adults of A. tartarea which had been reared form the

VOLUME LXXXVIII, NUMBER 3

185

egg stage on China aster, had a greater adult longevity than those reared on
its native host plant.

Acknowledgment

We thank Ms. Grace Johns for conducting portions of this research.

Literature Cited

Ball, E. D. 1905. Some new Homoptera from the South and Southwest. Proc. Biol. Soc.
Wash. 18:117-120.

Caldwell, J. S. 1949. A generic revision of the treehoppers of the Tribe Ceresini in America
north of Mexico, based on a study of the male genitalia. Proc. U.S. Natl. Mus. 98:49 1—
521.

Fairmaire, L. M. H. 1846. Revue de la tribu des membracides. Ann. Soc. Entomol. de France
4:479-531.

Kopp, D. D. and T. R. Yonke. 1973. The Treehoppers of Missouri: Part 2. Subfamily Smi-
liinae; Tribes Acutalini, Ceresini, and Polyglyptini (Homoptera: Membracidae). J. Kan-
sas Entomol. Soc. 46:233-276.

Matausch, I. 1912. Observations on some North American Membracidae in their last nymphal
stages. Bull. Amer. Mus. Nat. Hist. 31:331-336.

Provancher, L. L. 1872. Description de plusierurs Hemipteres nouveaus. le Nat. Canadien
4:350-352.

Say, T. 1830. Descriptions of new North American hemipterous insects, belonging to the first
family of the section Homoptera of Latreille. J. Acad. Nat. Sci. Philadelphia 6:235-244.

Severin, H. C. 1927. A third report upon the Membracidae (treehoppers) of South Dakota.
Proc. South Dakota Acad. Sci. 11:33-49.

Simons, J. N. 1962. Life-history and behavioral studies on Micrutalis malleifera, a vector of
pseudo-curly top virus. J. Econ. Entomol. 55:363-365.

and D. M. Coe. 1958. Transmission of pseudo-curly top virus in Florida by a tree-

hopper. Virology 6:43-48.

Tsai, J. H. 1980. Lethal yellowing of coconut palm: search for a vector. In: Vectors of Plant
Pathogens, K. F. Harris and K. Maramorosch, eds., pp. 177-200 (Chapter 11), New
York, N.Y.: Academic Press.

Van Duzee, E. P. 1908. Studies in North American Membracidae. Bull. Buffalo Soc. Nat.
Sci. 9:29-127.

. 1917. Catalog of the Hemiptera of America north of Mexico excepting the Aphididae,

Coccidae and Aleurodidae. California Agric. Expt. Sta. Entomol. Tech. Bull. 2:1-902.

(JHT) Agricultural Research Center, University of Florida, 3205 S. W.
70th Avenue, Fort Lauderdale, Florida 33314 and (DDK) Cooperative Ex-
tension Service Agriculture and Applied Science, North Dakota State Uni-
versity, Fargo, North Dakota 58105.

Received for publication March 8, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(3), 1980, pp. 186-196

ECOLOGY OF NECROPHILOUS AND CARPOPHILOUS
COLEOPTERA IN A SOUTHERN NEW YORK
WOODLAND. PART I— OVERVIEW1

Dominick J. Pirone2 and Daniel J. Sullivan, S.J.3

Abstract. — Almost 20,000 individual Coleoptera were collected, pinned
and labeled from 6 pitfall traps baited with decomposing smelt fish to attract
necrophilous beetles, and 6 traps baited with decaying cantaloupe melon for
carpophilous species. Six of these traps (3 pairs with each type of bait) were
located on a dry slope, and 6 others in an adjacent wet valley. This field
sampling was done over an 8-month period (April thru November) at Ford-
ham University’s Calder Ecology Center, Armonk, Westchester County,
New York. Approximately equal numbers of beetles were taken from each
bait (fish and melon), and from each habitat (slope and valley). The trapped
Coleoptera were concentrated in 10 major Families (99.4%), plus 5 minor
Families (0.4%) and 18 adventitious Families (0.2%). These 10 major Fam-
ilies also comprised 76.5% of the total 217 species trapped. The 3 most
numerous Families were the Staphylinidae (37.1%), Silphidae (21.5%) and
Nitidulidae (17.9%). These were followed by the Hydrophilidae (7. 1%), Lep-
todiridae (4.9%), Histeridae (3.4%), Carabidae (2.3%), Ptiliidae (2.0%),
Scarabaeidae (1.9%), Scolytidae (1.3%). Certain species were consistently
attracted to fish (necrophilous) and others to melon (carpophilous). How-
ever, in both cases, many of the beetles are actually predatory, and some
are general scavengers. Closely related species are usually separated tem-
porally, most often by fortnightly periods. The geography, geology and bot-
any of the research area are discussed as representing a southern New York
woodland situated less than 50 k north of midtown Manhattan.

Introduction

The purpose of this research was to identify and enumerate the families,
species and individuals of necrophilous and carpophilous Coleoptera col-
lected at traps baited with decaying fish and melon. The field study was
done in 2 ecologically distinct but adjacent areas: a dry slope and a moist

1 This research was supported in part by a National Science Foundation Grant GU-3554 to
the second author.

2 Assistant Professor in the Department of Biology, Manhattan College, Riverdale, New
York 10471.

3 Associate Professor in the Department of Biological Sciences, Fordham University, Bronx,
New York 10458.

VOLUME LXXXVIII, NUMBER 3

187

valley in an open woodland in southern New York State. The sampling
collections were made over an 8-month period from April thru November.

Decaying carrion and fruit are an important food source for many species
of insect larvae and adults, especially Diptera and Coleoptera. Beetles are
always of particular interest because of their sheer numbers (perhaps
300,000 species) which comprise about Vs of the world’s known insect
species. In addition, their well-known niche-specialization provides oppor-
tunities for remarkable diversity. For these reasons, Coleoptera can be stud-
ied to record food preferences between such energy sources as decaying
fish and melon. Further comparisons can then be made in relation to habitat
differences between a dry slope and a moist valley.

Therefore, these data were used to tabulate: (a) both the numerical and
temporal relations between the species; (b) the food preferences for fish or
melon; (c) the habitat association of the slope and valley. Such quantitative
information concerning the relative proportions and preferences of ecolog-
ically related species should be continually added to the insect ecological
literature (Matthews and Matthews 1970, Wallace 1974). Other authors such
as Arnett (1970) caution further that it is not enough to know only the orders
and families that occur in such habitats and have certain food preferences —
but also that the species themselves, their population fluctuations and phe-
nology must be studied. It is hoped, therefore, that the research presented
here will provide basic information for use in taxonomic catalogs and future
ecological studies. This Part I contains a summary overview of the entire
project at the family level.

Materials and Methods

Field study. — The field work was done at Fordham University’s Calder
Ecology Center, Whippoorwill Road, Armonk, Westchester County, New
York 10504. This sampling aspect of the research was conducted over an
8-month period from April thru November 1970. The field study area was
2.5 hectares, and was located in the northwest part of the 45 hectare Ford-
ham Ecology Center. It consisted of 2 distinct but adjacent sections: a wide,
western-exposed slope; a flat, narrower, stream-laced valley at the base of
the slope and 27.4 m lower in altitude.

Geography, geology and pedology. — The Fordham Ecology Center is lo-
cated 48.3 k north of midtown Manhattan in the County of Westchester,
and ranges from 145-200 m above sea level. It is underlain by the Manhattan
Prong of the igneous, highly metamorphosed Fordham Gneiss of the New
England Upland (Muller 1965, Schuberth 1968), and may be of proterozoic
(Precambrian) age, ca. 1 billion years old. It is about 56 k north of the
southernmost terminus of the last Wisconsin glaciation which began reced-
ing about 15,000 years ago. This Fordham property is composed of the

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NEW YORK ENTOMOLOGICAL SOCIETY

Hollis association of soils (laid down on glacial till). It is a shallow, sandy
loam 25-50 cm to bedrock, and slightly acidic, low in nitrogen and calcium,
marginal in phosphorous, but adequate in potassium for tree growth. The
slope is dry with shallow soil, while the valley is wet.

Botany. — The flora of the slope in this open woodland setting is dominated
by typical beech (Fagus grandifolia Ehrhart), birch ( Betula lenta L.) and
red maple ( Acer rubrum L.). The main associated trees are rock-chestnut
oak ( Quercus prinus L.), black oak ( Q . velutina Lamarck) and red oak ( Q .
rubra L.). The shrub understory is composed of maple-leaved viburnum
(Viburnum acerifolium L.), blueberries (Vaccinium spp.), pinxter-flower
(Rhododendron nudiflorum Torrey), and mountain laurel (Kalmia latifolia
L.). Some common vines, herbs, ferns, club-mosses and true mosses com-
plete the list of the slope macrophytes.

In the valley, however, only infrequently are beech and birch found (lim-
ited to high spots), while the red maple, so common on the slope, is com-
pletely replaced by sugar maple (Acer saccharum Marshall). The oaks are
entirely absent, perhaps because of the moist, deeper soil of the lower val-
ley. Hence, the characteristic trees of the valley are the white ash (Fraxinus
americana L.), black ash (F. nigra Marshall), bitternut hickory (Cary a cor-
diformis Koch), mockernut hickory (C. tomentosa Nuttall), and tulip-poplar
(Liriodendron tulipifera L.). Witch-hazel (Hamamelis Virginia na L.) is
usually a somewhat isolated species elsewhere, but in the valley it is com-
mon and attains a large size and extensive coverage. Spicebush (Lindera
benzoin Blume) dominates the high shrub layer. It is in the herbaceous
strata, however, that the well-watered valley differs very strongly from the
drier slope. The darker, deeper valley soil supports a nearly continuous
carpet of lush herbaceous growth, ranging from just above ground level to
over 1 m in height.

In summary, the total number of macrophyte species for each of the 2
study areas was: slope 29 and valley 78, with many species common to
both. By combining the total species for both slope and valley, a breakdown
of the above data reveals that there was a total of 22 species of trees; 17
shrubs and woody vines; 47 herbs, woodrush, grasses; 13 species of ferns,
clubmosses and true mosses.

Meteorology. — The Fordham Ecology Center is located just inside the
southern boundary of the climatic division of New York State known as the
"Hudson Valley.” The annual normal temperature for the study area is
about 1 1°C. The temperatures on the slope ranged from a high of 32°C on
the early date of May 10th to a low of — 21°C on January 22nd. In the valley,
a similar pattern was recorded, but usually 2.24°C lower in each case. The
highest temperatures each day were reached between 1-3 PM, while the
lowest were between 4-7 AM.

On days without precipitation, the relative humidity in the valley was

VOLUME LXXXVIII, NUMBER 3

189

usually 5-10% higher than that of the slope. The precipitation is about 112
cm annually, and well-distributed throughout the year, with a fairly regular
monthly average of slightly over 8.9 cm. During 1970, however, only 75 cm
fell in the study area.

Traps. — Each of the 12 pitfall bait traps consisted of a 3.78 1 cylindrical
tin can (size no. 10). In order to allow for rain and runoff water drainage,
25 tiny nail holes were hammered into the bottom of each trap in a regular
pattern. A hole was dug for each trap so that the rim of the can was exactly
flush with the surrounding woodland floor substrate. The outer circumfer-
ence of each trap was packed with soil so that no intervening gaps remained.
A .47 1 uncoated cardboard container was placed inside each trap to hold
the bait. A 23 x 23 cm square of galvanized 1.3 cm hardware screen was
placed on top of each trap. Four 2.0 cm thick small stones were placed at
each of the 4 corners of the screen to permit a crawl-space for the anticipated
beetles to enter the trap. This in turn was covered by a 23 x 23 cm square
piece of 0.6 cm masonry hardboard which acted as a rain-shield. Finally, in
order to protect the pitfall trap from racoons and other bait-raiders, a large,
heavy rock was placed over each rain-shield.

Bait. — Six traps were baited with fresh cantaloupe melon ( Cucumis melo
L.), sliced into sixths and the seeds removed. On the 1st day of baiting, 2
of the slices were placed in the cardboard container. On the 2nd day of
collecting and replacement of the bait, 1 of the 2 slices of melon was re-
moved from the trap and replaced by a fresh piece. Hence, a cyclic regimen
was maintained so that 1 slice of melon was usually 0-3 days old, and the
other slice 4-7 days old in each of the 6 melon traps.

The 2nd set of 6 traps was baited with 2 frozen, whole (18-23 cm) Atlantic
smelt ( Osmerus mordax Mitchell) with viscera and all body parts left intact.
The same regimen was followed in baiting with fish as with melon, so that
2 fish in different stages of decomposition were always in each bait con-
tainer.

Three melon and 3 fish pitfall traps were located on the slope, and an
equal number in the valley area. These 12 traps were placed in pairs (1
melon- 1 fish) at approximately 12 m distance from each other. These pairs
in turn were located 24 m apart.

Collections were made from the traps twice weekly (3-4 days apart) be-
tween 1-7 PM, during the 8-month study period. The removed bait was
carefully dissected in an aluminum baking pan in the field in order to insure
collection of all specimens which had bored into the bait (usually nitidulids
and scolytids). Other beetles were removed directly from the bait con-
tainers, and put together with the aforementioned Coleoptera into 57 g
screw-top labeled jars containing 80% ethyl alcohol mixed with 2% glycerine
by volume. Each area was sampled 46 times for a total of 552 separate
collections.

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NEW YORK ENTOMOLOGICAL SOCIETY

Table 1 . Families of Coleoptera collected from traps baited with fish for necrophilous bee-
tles and melon for carpophilous species over an 8-month period (April thru November) at
Fordham University’s Calder Ecology Center, Armonk, Westchester Co., N.Y.

Family

Individuals

Species

No.

%

No.

%

1. Staphylinidae

7,407

37.1

62

28.6

2. Silphidae

4,300

21.5

7

3.2

3. Nitidulidae

3,575

17.9

24

11.1

4. Hydrophilidae

1,427

7.1

a

—

5. Leptodiridae

979

4.9

6

2.8

6. Histeridae

684

3.4

17

7.8

7. Carabidae

463

2.3

23

10.6

8. Ptiliidae

390

2.0

4

1.8

9. Scarabaeidae

385

1.9

17

7.8

10. Scolytidae

266

1.3

6

2.8

Minor Families (5)

73

0.4

16

7.4

Adventitious families (18)

43

0.2

35

16.1

Totals

19,992

100.0

217

100.0

a All hydrophilids were in the genus Cercyon, but current taxonomic literature could not
identify specimens to species.

Curating. — The field-collected beetles were curated in the Entomological
Laboratories of Fordham University, Bronx, New York 10458. Each of the
approximately 20,000 adult beetles was individually pinned, labeled, and
then stored in 40 of Fordham’ s “California Academy of Sciences” style
insect drawers.

Results and Discussion

Necrophilous and carpophilous families. — Almost 20,000 beetles were
collected during the 8-month period (April thru November) from 6 traps
baited with fish and 6 with melon. Six traps (3 pairs with each type of bait)
were located on the dry slope and the other 6 in the wet valley. The trapped
Coleoptera were concentrated in 10 major families (19,876 beetles or 99.4%
of total) as shown in Table 1. In addition, there were also 5 minor families
which included only 73 individuals or 0.4% of the total and were associated
with fungi or decaying organic material. Finally, 18 other families (43 beetles
or 0.2%) were categorized as “adventitious,” i.e., phytophagous forms or
wanderers on the forest floor which were probably accidentally caught in
the traps. From the point of view of numbers of species, 217 species were
collected and identified. Of these, 76.5% belonged to the 10 major families,
7.4% to the 5 minor families, and 16.1% to the adventitious families.

10 major families. — Considering that almost 20,000 individual Coleoptera
were collected in the combination fish-melon and slope-valley traps, there

VOLUME LXXXVIII, NUMBER 3

191

Table 2. Seasonal summary over an 8-month period of the 10 major families of Coleoptera
collected from traps baited with fish for necrophilous beetles and melon for carpophilous
species in a dry slope habitat and an adjacent wet valley.

Month

Slope

Valley

Total

Monthly

Fish

Melon

Total

Fish

Melon

Total

Fish

Melon

Total

%

Apr

227

114

341

337

95

432

564

209

773

3.9

May

978

247

1,225

547

255

802

1,525

502

2,027

10.2

Jun

509

765

1,274

356

639

995

865

1,404

2,269

11.4

Jul

1,309

574

1,883

1,371

747

2,118

2,680

1,321

4,001

20.2

Aug

793

1,536

2,329

975

1,148

2,123

1,768

2,684

4,452

22.4

Sep

960

858

1,818

791

914

1,705

1,751

1,772

3,523

17.7

Oct

260

672

932

217

1,018

1,235

477

1,690

2,167

10.9

Nov

159

172

331

138

195

333

297

367

664

3.3

Totals

5,195

4,938

10,133

4,732

5,011

9,743

9,927

9,949

19,876

100.0

Percent

51.3

48.7

51.0

48.6

51.4

49.0

49.9

50.1

100.0

were some interesting, if not surprising, similarities in the overall totals
(Table 2). For instance, 9,927 necrophilous beetles were collected in the fish
traps, and almost an equal number (9,949) of carpophilous beetles were on
melon. Similarly, the slope traps (fish-melon) accounted for 10,133 beetles,
and the valley bait (also fish-melon) trapped 9,743. There are no significant
statistical differences between these pairs of numbers. However, when the
families and especially the species are examined more closely, then differ-
ences are seen in the preferences for the bait (fish or melon), and to a lesser
degree for the habitat (slope or valley). A more complete picture of this
Part l overview will be given in subsequent papers which will analyze in
detail the trophic and geographic differences not only between the 10 major
families, but also among the species of each.

For the purpose of this introductory presentation, however, a brief sum-
mary of the 3 most numerous families (Staphylinidae, Silphidae, Nitidulidae)
will be given below (Table 3). These 3 families comprised 76.5% of the
individual beetles collected in this study, and 42.9% of the total species.

1) Staphylinidae. — The staphylinids contained more individuals (7,407 or
37.1%) and more species (62 or 28.6%) than any other family. They were
attracted to melon over fish by a 2:1 ratio (4,704:2,703), and there was a
10% difference between the number in the valley (4,103 or 55.4%) and the
slope (3,304 or 44.6%).

2) Silphidae. — Although this family ranked 2nd in the number of speci-
mens (4,300 or 21.5%), it dropped significantly to 6th place in the number
of species, having only 7 species represented or 3.2% of the total. In addi-
tion, unlike the Staphylinidae, these beetles were almost absent from the
melon traps (only 0.8%). The slope also reflected a similar reversal in niche

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NEW YORK ENTOMOLOGICAL SOCIETY

Table 3. Comparison of the 3 most numerous Families of Coleoptera (Staphylinidae, Sil-
phidae, Nitidulidae) collected from traps baited with fish for necrophilous beetles and melon
for carpophilous species in a dry slope habitat and an adjacent wet valley over an 8-month
period (April thru November).

Family

Slope

Valley

Total

Total

Fish

Melon

Total

Fish

Melon

Total

Fish

Melon

Staphylinidae

No.

1,171

2,133

3,304

1,532

2,571

4,103

2,703

4,704

7,407

%

35.4

64.6

44.6

37.3

62.7

55.4

36.5

63.5

100.0

Silphidae

No.

2,457

24

2,481

1,807

12

1,819

4,264

36

4,300

%

99.0

0.1

57.7

99.3

0.7

42.3

99.2

0.8

100.0

Nitidulidae

No.

262

1,888

2,150

144

1,281

1,425

406

3,169

3,575

%

12.2

87.8

60.1

10.1

89.9

39.9

1 1.4

88.6

100.0

Totals

No.

3,890

4,045

7,935

3,483

3,864

7,347

7,373

7,909

15,282

%

49.0

51.0

51.9

47.4

52.6

48.1

48.2

51.8

100.0

preference, accounting for 57.7% of the trapped silphids, while only 42.3%
were collected in the valley traps.

3) Nitidulidae. — This was the 3rd most common family with 3,575 or
17.9% of the total individuals. Because of the few number of silphid species
noted above, however, the Nitidulidae moved up to become the 2nd largest
family as to number of species (24 or 1 1.1%). Of these, strong preferences
were shown for melon (88.6%), while 60.1% were trapped on the slope.
Hence, the nitidulids seemed to combine the preference of the Staphylinidae
for melon with that of the Silphidae for the slope. Another way of looking
at these data would be that the Nitidulidae occupied niches which were less
competitive when considered in relation to the other two families.

Phenology. — During the 8-month period of this study (Table 2), only 773
beetles (3.9%) were collected in April. This increased to 2,027 (10.2%) dur-
ing May, and 2,269 (11.4%) in June, so that both months were quite similar.
However, the peak months were July (4,001 or 20.2%), August (4,452 or
22.4%) and September (3,523 or 17.7%). The October collecting (2,167 or
10.9%) was similar to that of May and June, while November (664 or 3.3%)
returned to the low of April. This can be summarized by saying that 60.3%
of the beetles were trapped during the peak months of July, August and
September. But only about half as many (32.5%) were collected in May,
June and October. As a result, April and November being the beginning and
the end of the season, accounted for only 7.2% together.

In reference to the 3 most numerous families, the Staphylinidae peaked
from August through October. On the other hand, May, July and August
were the months in which the Silphidae were best represented, while the
Nitidulidae were trapped most frequently in June, August and September.

VOLUME LXXXVIII, NUMBER 3

193

These data indicate that August was the high month for collecting all 3
families. This was apparently due to phenological overlap, although the
other months show some temporal niche-splitting.

Many species of beetles were represented by a very small 1st generation
early in the spring or summer. After this initial appearance, they were not
trapped again for a long period during the remainder of the summer, but
reappeared with a large 2nd generation in the fall. Species with such a
pattern are usually designated as bivoltine, but it should be pointed out that
many spring specimens may have been overwintering adults (especially
among the Carabidae). In this case, such species are actually univoltine.
Winter diapause and summer aestivation are factors which should be studied
in this regard. Hence, the continuing need in insect ecology for observation
on the bionomics of each species in the field.

The results show that certain species are attracted to decomposing fish
(Eecrophilous) and others to melon (carpophilous). However, in both cases,
many of the beetles are actually predatory, and some are general scaven-
gers. Closely related species are usually separated temporally, most often
by fortnightly periods. Some of the seemingly weaker species emerge earlier
than the stronger ones, and so gain a definite competitive advantage in time.

Fuller (1934), working for 4 Vi years in Australia, recorded far fewer
species of beetles on carrion than is reported in this study, although the
families represented were remarkably similar. Howden (1950) recorded 98
species of Coleoptera on carrion. The Familes Staphylinidae, Histeridae and
Carabidae made up 50% of the beetles. She concluded that the effect of the
surrounding flora on a carcass was expressed in moisture relationships, and
our data on the dry slope and wet valley differences support this interpre-
tation. She found that more beetle species and individuals participated in
the succession in the spring than in the summer. In autumn, the carrion
regained some of the lively and abundant qualities of the spring, but the
fauna was not as rich.

Walker (1957) also showed that the succession of organisms associated
with decaying material is influenced by variation in vegetation and micro-
climate. In addition, in an area of only a few hectares, the animals associated
with similar decaying materials in adjacent habitats may be quite different.
He used cantaloupe, fish and cornmeal as bait. The major groups of the
trapped beetles included 15 species of staphylinids, 7 nitidulids and scarabs
each, 3 histerids, 2 carabids, and 1 species each of hydrophilid, leiodid,
ptiliid, mycetophagid — all taken on cantaloupe. In the fish baited traps, the
following families of Coleoptera were recorded: 18 species of staphylinids,
8 species each of silphids and histerids, 6 scarabs, 2 hydrophilids and tro-
gids, plus 1 nitidulid, carabid, dermestid, ptiliid, monotomid. Walker cor-
rectly noted that most of the field research on how attractive a bait is at
certain times in its decay cycle fails to take into account whether the species

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NEW YORK ENTOMOLOGICAL SOCIETY

are naturally available as adults at the time of the study. Our work at Ford-
ham attempts to elucidate this neglected aspect of insect ecology.

Reed (1958) reported an analysis of dog-carcass microcommunities with
particular reference to serai development, seasonal variation and environ-
mental factors. He found that the summer months had peak populations
with 217 species of insects of which only 96 were coleopteran species.

Payne (1965) used as carrion baby pigs ( Sus scrofa L.). Carrion free of
insects decomposed very slowly, merely drying and retaining its form for
many months. However, 90% of the carrion open to insects was removed
in 6 days. Only 2 coleopterous families, the Staphylinidae and Histeridae,
were reported during the 2-3 weeks of his study. He determined that by the
5th day of exposure of pig carrion, beetles were the dominant adult insects
present. Payne found that the activities of insects were more influenced by
temperature than by any other environmental factor, while moisture ranked
second. Payne agrees with Fuller (1934) in stating that insects hasten carrion
disintegration by dissemination of bacteria, secretion of digestive juices, and
by the mechanical processes of burrowing and tunneling through the car-
casses. The Coleoptera and Diptera comprised 60% of the total fauna taken
on carrion. Of 522 organisms collected, 422 species were insects, with 217
being beetles. Once again, the staphylinids and histerids were the dominant
Coleoptera (92 species or 42.4%).

Extensive field studies on carrion beetle responses to poikilothermic (fish)
and homoiothermic (chicken) bait have been done by Shubeck (1968, 1969,
1971, 1975a,b, 1976a, b) over many years using specially designed pitfall
traps as well as specially designed ground and air traps. As in our study, he
agreed with Walker (1957) that the differences in the species collected were
due to the degree of carrion decay within the habitats, which in turn were
caused by differences in microclimate. At the Family level, the overwhelm-
ing majority of beetles taken were Silphidae, Staphylinidae, Histeridae,
Leiodidae, Scarabaeidae and Nitidulidae. Similar supporting studies have
been done using lizards and toads (Cornaby 1974), and mammals (Johnson
1975).

The data obtained in our own field collections at the Fordham Ecology
Center also fit very well with the population generalizations of Preston
(1948). He noted that in any association there are just as many very rare
species as there are common ones, but species of moderate abundance are
vastly more numerous than either. Wilhm (1968) stated that the widespread
similarity in numerical distribution patterns of individuals and species is one
of the most interesting and still unexplained properties of ecological sys-
tems. Our results concur with these conclusions that a relatively small per-
centage of the species contain a large number of individuals, and a large
percentage contain only a small number of individuals. Hence, the necrophi-

VOLUME LXXXVIII, NUMBER 3

195

lous and carpophilous Coleoptera can be used as ecological indicators of
the numerical and temporal relations between species.

Acknowledgments

Special appreciation is expressed to Dr. Samuel S. Ristich of the Boyce
Thompson Institute for Plant Research, Cornell University — for his advice
on the ecological methodology used in this field study.

Literature Cited

Arnett, R. H. 1970. Objectives of a taxonomic catalog of Coleoptera. Coleopt. Bull. 24:76-
84.

Cornaby, B. W. 1974. Carrion reduction by animals in contrasting tropical habitats. Biotropica
6:51-63.

Fuller, M. E. 1934. The insect inhabitants of carrion: a study in animal ecology. Austral.
Council for Sci. Indus. Res. Bull. 82:1-62.

Howden, A. T. 1950. The Succession of Beetles on Carrion. Unpublished M.S. Thesis, North
Carolina State College, Raleigh, North Carolina. 83 p.

Johnson, M. D. 1975. Seasonal and microserai variations in the insect populations on carrion.
Am. Midi. Nat. 93:79-90.

Matthews, R. W. and J. R. Matthews. 1970. Malaise trap studies of flying insects in a New
York mesic forest. I. Ordinal composition and seasonal abundance. Jour. New York
Entomol. Soc. 78:52-59.

Muller, E. H. 1965. Bibliography of New York Quaternary Geology. New York State Mus.

and Sci. Service Bull. no. 398, Albany, N.Y. 116 p.

Payne, J. A. 1965. A summer carrion study of the baby pig Sus scrofa Linnaeus. Ecology
46:592-602.

Preston, F. W. 1948. The commonness, and rarity of species. Ecology 29:254-283.

Reed, H. B. 1958. A study of dog carcass communities in Tennessee, with special reference
to the insects. Am. Midi. Nat. 59:213-245.

Schuberth, C. J. 1968. The Geology of New York City and Environs. The Natural History
Press, Garden City, N.Y. 304 p.

Shubeck, P. P. 1968. Orientation of carrion beetles to carrion: random or non-random? Jour.
New York Entomol. Soc. 76:253-265.

. 1969. Ecological studies of carrion beetles in Hutcheson Memorial Forest. Jour. New

York Entomol. Soc. 77:138-151.

. 1971. Diel periodicities of certain carrion beetles (Coleoptera: Silphidae). Coleopt.

Bull. 25:41-46.

. 1975a. Do diurnal carrion beetles use sight, as an aid to olfaction, in locating carrion?

The William L. Hutcheson Memorial Forest Bull. 3:36-39.

. 1975b. Flight activity of certain carrion beetles; Silpha noveboracensis, Staphylinidae,

Histeridae. Ibid. 3:40^13.

. 1976a. An alternative to pitfall traps in carrion beetle studies. Ent. News. 87:176-178.

. 1976b. Carrion beetle responses to poikilotherm and homoiotherm carrion (Coleop-
tera: Silphidae). Ibid. 87:265-269.

Walker, T. J. 1957. Ecological studies of the arthropods associated with certain decaying
materials in four habitats. Ecology 38:262-276.

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NEW YORK ENTOMOLOGICAL SOCIETY

Wallace, B. 1974. Radioactive wastes and damage to marine communities. BioScience 24: 1 64—
167.

Wilhm, J. L. 1968. Biomass units versus numbers of individuals in species diversity indices.
Ecology 49:153-156.

Department of Biological Sciences, Fordham University, Bronx, New
York 10458.

Received for publication March 25, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(3), 1980, pp. 197-198

PLATANTHERA CRISTATA (MICHX.) LINDL., A NEW HOST FOR
THE RED-BANDED LEAF ROLLER1

T. L. McCabe and C. J. Sheviak

Abstract. — Larvae of the tortricid, Argyrotaenia velutinana Wlk., have
been recorded feeding on the seeds of the orchid, Platanthera cristata
(Michx.) Lindl., on Long Island, New York.

Four plants of the orchid, Platanthera cristata (Michx.) Lindl., were
collected August 15, 1979 from pitch pine barrens on Long Island, New
York, and were subsequently cultivated at Albany. Within ten days two
plants showed webbing and frass resulting from the activity of a caterpillar
in each inflorescence. By October 28th, a mature larva of the red-banded
leaf roller, Argyrotaenia velutinana Wlk., was found feeding within a seed
capsule. Normal larval behavior is leaf rolling, i.e., it feeds and rests from
within a curled leaf (Forbes 1923).

Chapman and Lienk (1971) gave host records for a great many deciduous
and coniferous trees. They stated that the moth is of low incidence in nature
making it difficult to identify native primary hosts and that virtually all
reports of secondary hosts have been from the immediate vicinity of apple
orchards, where it is a major pest. The present collection was made in an
undisturbed community. The orchid is a coastal plain species that reaches
its northern limits on Long Island. The moth is found throughout the East.

The larvae were first noticed toward the end of the orchid’s period of
bloom, indicating that the eggs were laid in the mature inflorescence at about
the peak of flowering. This suggests that the moths were attracted to the
flowers. The moth proboscis is less than two millimeters long, and, as the
orchid’s nectariferous spur is about five millimeters long, the moth probably
does not serve as a pollinator. The larvae constructed a loose silken tube
running nearly the length of the inflorescence, and ate the contents of six
or seven seed capsules, entering each at the middle, before pupating. One
larva pupated within a seed capsule on 8 October and was refrigerated on
28 November. The pupa was reintroduced to room temperature on 16 Feb-
ruary and a male emerged on 2 March 1980. The pupal exuvia protruded
from the capsule.

Pupal exuvia, mature larval head capsule, reared adult, and specimens of

1 Published by permission of the Director, New York State Museum, State Education De-
partment, Journal Series No. 302.

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the orchid are coded tlm 79-78 and CJS 1606, respectively, and are deposited
in the New York State Museum.

Literature Cited

Chapman, P. J. and S. E. Lienk. 1971. Tortricid fauna of apple in New York. N.Y. State
Agric. Exp. Sta. Special Public. Ithaca, 122 pp.

Forbes, W. T. M. 1923. The Lepidoptera of New York and neighboring states. Cornell Uni-
versity Agric. Exp. Sta. Mem. 68:1-729.

New York State Museum, State Education Department, Cultural Edu-
cation Center 3132, Albany, New York 12230.

Received for publication March 26, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(3), 1980, pp. 199-209

EVOLUTIONARY ORIGIN OF BEES
(HYMENOPTERA: APOIDEA)

U. N. Lanham

Abstract. — Bees are generally supposed to be descendants of sphecid
wasps, largely on the basis of similarities in thoracic structure. If it be
assumed that these similarities are convergent, resulting from independent
evolution for high-precision flight, then an argument can be made for a close
relationship between bees, ants, and scolioid wasps.

In 1951 I wrote that “the pollen and nectar-feeding bees arose from the
primarily predaceous wasps . . . ,” even then a well-worn phrase, and went
on to say that “. . . but acceptable theories as to the transition between the
two modes of life have not been worked out.” This last also was true then
and is now, in spite of the publication of a book on the evolution of the
Hymenoptera (Malyshev, 1968). Nor does the present paper remedy the
situation, in the sense that “at last the mystery is solved.” Nevertheless
aculeate phylogeny, although an inherently frustrating subject, in the man-
ner of all phylogenetic studies, is worthwhile insofar as it organizes obser-
vation of the immense complexity of morphology and behavior to be found
in the aculeate Hymenoptera.

It is the modern consensus that the group of wasps that gave rise to the
bees is the sphecids. Two recent works supporting this theory are those of
Bohart and Menke (1976:31) and Brothers (1975). Among the sphecids,
Malyshev (279ff) selects the pemphredonines as the group offering a tran-
sition from sphecid to bee. Some of these are small wasps nesting in twig
cavities that they fill with small, soft-bodied insects such as aphids. Indi-
vidual cells along the length of the twig cavity are lined with a tough trans-
parent membrane produced by mandibular glands of the female. The most
wasp-like of the bees, according to Malyshev, are the hylaeines, some of
which also nest in twig cavities with individual cells lined by thin transparent
membranes. Aphid-feeding pemphredonines, Malyshev reasons, could have
shifted from aphids, themselves a mixture of protein and honeydew, to a
pollen-nectar food store for the young. This would mark the genesis of bees.

Hylaeine bees and pemphredonine wasps resemble each other both su-
perficially and in those characteristics, mainly involving the thorax, which
in Brothers’ opinion tie all of the bees closely to the sphecids. A discussion
of these common characters will be deferred for the moment to concentrate
on the differences. If a Hylaeus is examined carefully at high magnification,
it will be found that at least a few of the hairs are compound (branched or

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feathery). Such hairs are especially abundant in the short filter fringe on the
prothoracic lobe, which covers a large spiracle. Hylaeine bees are somewhat
atypical in being relatively hairless; most bees are abundantly long-hairy,
with many branched hairs. In fact, the presence of branched hairs is abso-
lutely diagnostic in comparing bees with sphecids: none of the sphecids
have such hairs; all bees do have them.

It is unreasonable, in my opinion, to think that the few compound hairs
of the hylaeines represent a basis for the selective evolution of compound
hairs in the bees. Rather, they represent the situation found in the many
species of parasitic bees which do not have to transport pollen in external
baskets or brushes; these are also nearly hairless, and also have a few
compound hairs. Hylaeines transport pollen to the brood cells in the crop,
gathering the pollen with their front legs into the mouth. It seems more
reasonable to think that they display a specialized reduction of hairs such
as occurs in the parasitic bees, made possible by the crop-transporting
mode. Better representatives than the hylaeines of primitive bees are per-
haps the euryglossines, limited to Australia, which are not superficially
wasp-like. Although these, like hylaeines carry pollen in the crop (presum-
ably the primitive mode) rather than externally, they are abundantly long-
hairy, with many compound hairs. The hylaeines could then be regarded as
specialized descendants of the euryglossines.

It is difficult to imagine that the presence of a few compound hairs in the
manner of the hylaeines would provide the broad and stable base necessary
for the evolution of compound hairs which, judging from their absolute
absence in the many thousand species of sphecid, vespid, pompilid, tiphiid,
and scoliid wasps, are not evolved except in highly unusual circumstances
(Lanham 1979). If we take the compound hairs as representing a fundamen-
tal rather than trivial character of the bees, then we are led to consider the
possibility that the pemphredonine-hylaeine transition is not the best hy-
pothesis for the evolutionary origin of bees.

Another difference that leads to the same conclusion is one which for
some reason escaped general notice until rather recently (Lanham 1960 and
Bohart and Menke 1976:27). This is the presence on the hind leg of the
sphecid of a strigil composed of a hind tibial spur (sometimes modified)
facing a brush-lined concavity at the proximal end of the basitarsis. When
the wasp disposes of particulate debris, it pulls the opposite hind leg, which
is stocked with the particles by a variety of grooming movements, through
the strigil in the manner that the antenna moves through the strigil of the
front leg. In all bees the hind strigil is absent, and debris is disposed of by
shuffling together the two brushes on the inner surfaces of the widened hind
basitarsi. It has been suggested that the evolution of the basitibial brush in
the bees obliterated the ancestral sphecid strigil, but it seems to me that the
brush is a character which has been inherited directly from primitive astrig-

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201

APID

Fig. 1. Phylogenetic tree of the aculeate Hymenoptera. Letters refer to strategies or major
adaptive shifts, numbers to the one or more major taxa that result.

ilate aculeates without the intervention of a strigilate phase. One sees on
the slender basitarsis of some primitive ants the apically directed hairs which
make of it a rather bee-like, one-way disposal apparatus when shuffled
against the opposing brush. Wilson (1962:413) has described this cleaning
movement in certain primitive ants, and I have observed it in the non-acu-
leate Pelecinus.

If the compound hairs and absence of the hind strigil are taken so seriously
that it is thought best to shift the origin of bees to an area of the aculeates
where both characters are present, then the only site available in the living
fauna is one near the ants. Ants are so specialized that they could not serve
as ancestors for the bees, but both could be considered to be derived from
a stock of astrigilate wasps with compound hairs at some stage in the life
cycle. In addition, as will be brought out subsequently, the ancestral stock
of the bees and ants would be social and nest in cavities, most likely in
wood.

It is obvious that if the origin of bees is shifted away from the sphecids,

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then the similarity in thoracic structure must be regarded as the result of
convergent evolution. Brothers, who has made a detailed study of the acu-
leate thorax in his phylogenetic studies, finds three characters of the thorax
which in his numerical procedure tip the balance heavily in favor of a close
bee-sphecid relationship.

1 . The antero-lateral corner of the mesoscutum is enlarged so that it extends
far anterior to the tegula. This is Brothers’ character state 21.2, which
“arose uniquely on internode 4-5 and thus is a very good indicator of
the holophyly of the sphecid and apid groups.” The position adopted
here is that it arose on two occasions, once within the strigilate wasps,
once within the astrigilate wasps.

2. The ventral angles of the propodeum are slender and so elongate that
they almost or quite encircle the thorax, meeting (or nearly so) at the
ventral midline. This is character state 23.2, which “is uniquely present
in the sphecid and apid groups and apparently arose on internode 4-5,
thus forming a strong indicator of the holophyly of this grouping.” The
position taken is that this structure is the inevitable consequence of the
enlargement of the antero-lateral corners of the mesoscutum. There is no
other way in which a broad support to brace the prothorax (which carries
the industrially important front legs) against the mesothorax can be man-
aged, when the slanting abutment against the anteriorly narrowed meso-
scutum of other aculeates is abolished. Thus character state 23.2 is a
mechanical correlate of 21.2 and cannot be weighted as a separate evo-
lutionary event.

3. The origin of the postphragmal muscle shifts posteriorly to a position on
the dorsal surface of the apparent propodeum. Brothers interprets the
origin of this muscle as always being the metapostnotum, so that the
large “triangle” of the propodeum is the metapostnotum. This is char-
acter state 35.3, which he says is “unique in the Aculeata and apparently
in the Hymenoptera as a whole. This remarkable modification of the
metapostnotum which is present in the sphecid and apid groups provides
extremely strong evidence of the holophyletic association of these
groups, having arisen on internode 4-5.” The small postphragmal muscle
inserts on the posterior surface of the 2nd phragma, a curved sclerotized
band whose anterior face provides the area for the origin of the longi-
tudinal indirect flight muscles. In character state 35.3 the phragma be-
comes vertical instead of slanting far back of and below the insertion of
the postphragmal muscle (Brothers 1976, figs. 1-12). When the phragma
became vertical, the bundles of the longitudinal muscles become more
nearly equal in length, and the functional center of the phragma has
retreated, making necessary a posterior shift in the origin of the post-
phragmal muscle. I am unable to explain the functional significance of

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203

this, except to say that it would seem reasonable that equal-length lon-
gitudinal fibers would deliver more smoothly the power that arches the
mesoscutum and depresses the wings. At any rate, the net result of all
three character states is the consolidation of the indirect flight muscles
(which furnish nearly all of the energy of flight) into a compact subspher-
ical or elliptical mass. It may be that character state 35.3 is also part of
the only solution to the problem of enhancing flight ability (a matter to
be discussed in the next section) and thus, like state 23.2, not to be
weighted as independent of the other two character states.

Enlargement of the antero-lateral corners of the mesoscutum is here taken
to be the basic morphological character from which other major thoracic
characters follow. Its primary function is that of increasing the area of in-
sertion, hence the size of the vertical (elevator) indirect flight muscles. Judg-
ing from the figures of the thoracic musculature of Apis in Snodgrass (1942)
the longitudinal and vertical muscles are of equivalent size. This would
mean that the upstroke would deliver approximately the same power as the
downstroke. This aspect of the functional morphology of the aculeate thorax
is a complex subject that has not been investigated. However, nearly every
field entomologist has seen the extreme example: the large anthophorine
bees or the bembecine wasps, which hover perfectly still, then accelerate
swiftly to a speed where they become invisible (an object this size becomes
invisible at about 35 miles per hour) and with such accuracy that they easily
avoid the net. No other aculeates approach the powerful and high-precision
flight of such bees and sphecid wasps. Intuitively it would seem that con-
tinuous delivery of power at a uniform level would give an aerodynamic
fine-tuning not attainable in the presumably more ragged flight of a wasp
with a considerable difference in delivery of power in upstroke and down-
stroke. The effect on flight musculature of the anteriorly narrowed meso-
scutum (characteristic of all aculeates above the bethyloid level except sphe-
cids and bees) can be seen in the figures by Duncan (1939) of the thoracic
musculature of the vespid wasp Vespula pennsylvanica (Sauss.). Here the
longitudinal indirect flight muscles are, says Duncan, enormous, being much
the largest muscles of the body. The size of the vertical muscles, which
insert laterally on the mesoscutum, is limited by the shortened area for
insertion as compared with the longitudinals. Some tiphiid and pompilid
wasps seem to have the anterior portion of the mesoscutum squared off
rather than narrowed, but this is illusory. If the long, overlapping pronotal
collar is pried off, underneath can be seen the sclerotized, narrowed end of
the mesoscutum.

In brief, it is proposed that a major adaptive characteristic was arrived at
independently in the bees and sphecids. This adaption is the high-precision
flight made possible by the enlargement of the elevator flight muscles. In

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the bees, this is useful in flower visiting and return and entry to a nest. In
the sphecid wasps it is useful in visiting flowers for fuel, searching for and
catching prey, and returning to and entering a nest. In both it is, especially
in the spacious, competitive desert environment, helpful in eluding birds or
such swift insect predators as asilid flies or, indeed, some of the sphecid
wasps themselves.

With the bees now in theory detached from the sphecids, a possible home
for them is provided in the accompanying phylogenetic tree. Letters indicate
major strategies or adaptive advances that separate the tree into a number
of sectors, each of which contains one or more higher taxa.

Bethyloids (sector 1) are for the most part small to minute parasitic wasps.
These probably could not be directly ancestral to any of the aculeate groups
with large-sized individuals, since wing venation is much reduced in the
forewings and absent in the hind wings. The genus Scleroderma of the bethy-
lids has long figured in speculations on the origin of ants (see Malyshev p.
217ff). A female lays many eggs on a large host larva and stays with it,
providing rudimentary care for the young. Eventually a few generations of
wasps are present on a single larva, forming a pre-social aggregation in-
cluding many wingless, ant-like females. The phylogenetic position of the
bethyloids is not well understood, but they have a mixture of characters
belonging to other superfamilies, so could possibly be specialized descen-
dants from a primitive stock of aculeates.

Strategy A is a modification of the parasitic mode in which relatively large
and powerful wasps dig through loose soil or rotting wood in a more pred-
atory fashion after beetle larvae which they paralyze with the sting, and on
which they lay a single egg. Primitively they leave the prey in place, but
some dig a cell which shelters the larva and the subsequent wasp larva, or
even move the host larva to another shelter. Sector 2 includes two major
families, the scoliids and tiphiids, which are particularly abundant in the
warmer parts of the world and with many species in the Southern Hemi-
sphere. These families, and two or more smaller families, are sometimes
put in a superfamily, the Scolioidea. They can be termed the non-nesting
astrigilate wasps, although as already pointed out, the behavior of some of
the species grades into the better defined nesting behavior of the more ad-
vanced wasps.

Strategy B is an adaptive shift in which some of the astrigilate wasps use
arboreal nesting sites where they store prey in cavities in wood. These clean
and dry sites in the tropical forest might have proved a major new nesting
environment which made possible a train of events leading to the ants and
bees. These nesting cavities, which would be enlarged, using the mandibles,
were from the beginning, and remained so during the early stages of bee
evolution, communal nest chambers. This communal mode leads easily into
progressive feeding, then through pre-social communities to true societies.
One of the consequences of moderately large societies, it seems to me, must

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have been the opportunity to widen the food base. Primitive hunting wasps,
for example, might be transformed from fairly specialized hunters into gen-
eral hunters and scavengers, bringing all kinds of edibles into the nest. In
some instances, the at first tentative use of a new food source might open
up an area for specialization. While most modern ants, for example, are
generalists, a few are specialists on such prey as other ant species or ter-
restrial isopods. Social aculeates living in communal nests produce a sig-
nificant evolutionary event — larvae with long hairs (Lanham 1979). Long-
haired larvae, otherwise unknown in the aculeates, occur in about 90% of
the genera of ants and in many of the genera of allodapine bees. Further-
more, at least one species of allodapines and about half the genera of ants
have compound larval hairs. The modified hairs of ant larvae have a great-
er variety of form than any other feature of ant morphology. Some 30 types
have been described and appear to be adaptations for various characteristics
of life in communal cells (Wheeler and Wheeler 1976). Communal nests
(with only a handful of apparently irrelevant exceptions) are known among
the aculeates only in the ants (essentially all) and in some bees (some al-
lodapines and Bombus). Thus, the bees have been placed in a phylogenetic
milieu that could explain the two features separating them from the sphe-
cids: absence of the hind strigil and presence of compound hairs.

Nowhere have there been found scolioid wasps that suggest the kind of
transition made in strategy B. There is only one extensive, unexplored group
remaining among the larger scolioid wasps. This is the subfamily Thynninae,
with about 500 species in Australia and some hundreds in South America.
All the known species have winged males and wingless females. Males are
larger than the females and carry them about during mating. For most
species, the females are observed only when found on the conspicuous,
flower- visiting males. Some spend most of their time burrowing about in the
ground in search of beetle larvae. Probably the biology of fewer than 5% of
the species is known in Australia (Burrell 1935). The females of these sting
the host larva, oviposit, and leave it in place. In only one species, which is
taxonomically isolated from the rest of the thynnines ( Diamma bicolor
Westwood) is the female known to drag its prey to its own burrow. In
species other than Diamma the males, unlike nearly all other aculeates,
play a role in the economy of the wasp other than fertilizing eggs. Some
carry the females, attached to the tip of the male abdomen, to feeding sites,
either flowers or honeydew-secreting scale insects, where they position
themselves so that the female can drink. Others carry nectar or honeydew
to the female, carrying a droplet in a basket of hairs on the underside of the
head (Given 1954), or feeding by regurgitation. Perhaps somewhere in this
ancient and abundant group there may be found small societies of females
(even attended by solicitous males) which would bear witness to the former
existence of sector 3. A few species live in wood.

This is the place to point out that the delineation of a clean, arboreal

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communal nesting site as the site for the evolution of the ancestral stock of
bees may have fashioned a two-edged sword, which may provide evidence
that it was indeed the sphecid wasps that gave rise to the bees. It has been
rather recently discovered (see Matthews 1968 and Bohart & Menke
1976:172) that a genus ( Microstigmus ) of small neotropical pemphredonine
wasps live in diminutive (18 x 12 mm) bag-like nests suspended from the
underside of the leaves of forest trees. The species M. comes Krombein,
measuring about 3 mm in length, constructs the nest from fibers scraped
from the undersurface of the leaf. It would seem that this cavity would offer
the ideal opportunity for the establishment of a communal nest, yet this
species remains determinedly sphecid, constructing in the loose fibers filling
the lower half of the bag up to 15 separate cells, with one naked larva reared
in each. Nevertheless, one can imagine that in the yet unstudied species of
the genus there may be found an evolutionary sequence in which cell making
is abandoned and the larvae have become hairy, in the manner of other
aculeates living communally. Only 17 species of Microstigmus are listed in
the Bohart and Menke catalogue, but apparently more than a hundred more
have since been discovered. These wasps represent a most interesting de-
parture from the biologies of other sphecids, but it is probably a specializa-
tion too recent to have been involved in the evolution of the bees or ants
or both.

Strategy C was the commitment to on-foot foraging by a wingless female
worker caste. Reproductive males and females are generally winged, but
flight ability is degenerate, serving to form mating swarms and to aid in
dispersal. This social, on-foot foraging mode of existence laid the foundation
for the evolution of the most successful group of aculeates, the ants, with
a number of species exceeded only by the bees and in numbers exceeding
all other aculeates. Although they are dominant in the arboreal fauna of the
tropics below 2,500 meters of altitude, the key to their success has been the
invasion of the nutrient rich ground-air interface, where they rule in all the
sun warmed land environments on the earth.

Sector 4 is the living family Formicidae, the ants. In the clean arboreal
environment there was no selection of the hind strigil and the ants remained
almost entirely astrigilate. A few of the ground-dwelling genera, such as
Neoponera, have a fairly well developed hind strigil, but these occur in
only few and scattered instances, and are taken to be independent origins
of this structure. The other thousands of species of ground dwelling astrigi-
late ants are able to take care of hygiene by mutual grooming and the fact
that the legs are so long that the hind legs can be pulled through the strigil
of the front legs.

As has been said, probably about half the genera of ants have larvae with
compound hairs. A few species also have compound hairs in the adult stage.
Two are known in North America. Triglyphothrix striatidens (Emery), an

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introduced species, has trifid hairs. The larva has hairs with “multifid tips”
(Wheeler & Wheeler 1973). The native species Acanthomyops plumipilosus
(Buren) has plumose hairs, the genus itself generally has denticulate hairs
as larvae. The similarity between larval and adult hairs, particularly in the
peculiar first example, tends to support my contention (1979) that the genetic
expression of compound hairs can be shifted from the larval to the adult
stage. However, the numerous examples of exotic ants with odd shaped
hairs in the adult need to be checked against larval morphology to learn
more of this phenomenon.

Strategy D is a rather minor development in which the females usually
become wingless and solitary parasites of the nests of other Hymenoptera.
Some extra-North American species are said to parasitize larvae of other
insect orders.

Sector 5. The great majority of species are in the family Mutillidae. The
families Eotillidae, Typhoctidae, and Anthoboscidae (females winged) are
included in this sector because at least some of the species of each have
compound hairs, thought to be sporadic, genetically based reminiscences of
a cavity-nesting past.

Strategy E is the specialization for pollen feeding that marked the origin
of bees. One other group of aculeates, the masarine vespid wasps, also
provides its young with a food store of pollen, but this was a relatively small
scale venture, involving less than a thousand species. Bees total about
20,000 species, and have more than any other group cooperated in the evo-
lution of the flowering plants. As said in the discussion of Sector 3, the
social mode makes possible a widening of the nutritional base. Among the
edibles brought in to these early societies there would certainly be pollen,
which was available in the Cretaceous, when these early phases in hyme-
nopteran evolution were taking place (Evans 1973). Instances in which pol-
len becomes an important food source would produce overwhelming selec-
tive advantage for the changes in control genes that would transfer
expression of compound hairs to the adult stage. The pollen-gathering mas-
arine wasps, mentioned above, did not have a communal-nesting ancestry,
never had compound hairs in their evolutionary background and the modern
species do not have compound hairs.

Sector 6 comprises the bees, usually put in a superfamily divided into
several families. Thus use of the term “apids” here is merely a convenience,
following the terminology used by Brothers, and does not represent a state-
ment as to the rank given the taxon. It is supposed here that the bees were
primitively social arboreal cavity nesters. Today only a minority of bees are
social, either primarily or secondarily, being represented by several
hundreds of species, most of them dwelling in arboreal nests constructed of
wax and resins, but the allodapines resembling at least superficially the
supposedly primitive mode in which there are small communal nests in twig

208

NEW YORK ENTOMOLOGICAL SOCIETY

cavities. Among the wax-nesting bees are the usually nonarboreal Bombus,
of which a few species nest communally during at least part of the seasonal
cycle. None of these have long-hairy larvae. In the more primitive bee
societies, reversion of social bees to the solitary mode, or even the reverse, j
is quite feasible (Wilson 1971:102). The vast majority of living bees are
solitary, are especially well represented in arid regions, and usually nest in
the ground in the manner of many sphecid species, providing each larva
with its own earthen cell.

Strategy F represents the transition between the primitive astrigilate non-
nested to the strigilate ground nesters. Here the burrow and the cells are
carefully constructed, with a brood cell of compacted earth. The nest is
often multicellular, thus being visited often between foraging trips, and even
when unicellular, with several burrows being dug, the cell is often provided
with several prey insects or spiders. It is here postulated that the develop-
ment of the hind strigil is of adaptive value in keeping clean in this mode of
life, and in helping maintain hygiene in the brood cells.

Sector 7 includes the pompilids (spider wasps) and vespid wasps. Pom-
pilids are either ground nesters (sometimes using a preformed cavity) or
construct above-ground nests of mud cells. All are solitary. They are rather
poor fliers, and have a tendency to collide with rather than gracefully alight
upon flowers. They tend to do much of their hunting on foot. The primitive
subfamilies of the vespids are solitary, constructing their cells in the ground,
or in twigs building mud cells above ground. The advanced vespine wasps
build arboreal nests of papery materials (sometimes the same sorts of nests
are made in large underground cavities). These wasps are always social.
Each larva has a separate paper cell.

Strategy G involves the evolution of high-precision flight, the morpholog-
ical expression being the expansion of the antero-lateral corners of the meso-
scutum. It may be that the failure of this character to develop in the vespids
was due to the evolution of the yet unexplained peculiarity of the wings,
which in all but a few primitive forms are kept folded lengthwise when not
in use. This might have some advantage in the nest life, but might well be
a liability in flight. Its advantage in the sphecids is probably a reflection of
their generally more intense foraging and nesting activity: the frequent visits
to flowers for nectar for fuel, alternating flight and on-foot pursuit of prey,
sometimes the prey captured in flight, and the wasp hovering alertly, ready
to dash away at any disturbance.

Sector 8 includes the sphecid wasps. The biology of the exceptional social
forms was discussed earlier. The thousands of species of solitary forms,
mostly ground nesting, prey on insects or, more rarely spiders. Unlike the
bees and scoliids, there are few endemic or aberrant groups in Australia.

VOLUME LXXXVIII, NUMBER 3

209

Literature Cited

Bohart, R. M. and A. S. Menke. 1976. Sphecid wasps of the world. A generic revision.
University of California Press, Berkeley. 685 p.

Brothers, D. J. 1975. Phylogeny and classification of the aculeate Hymenoptera, with special
reference to Mutillidae. U. Kansas Sci. Bull. 50:483-648.

. 1976. Modifications of the metapostnotum and origin of the 'propodeal triangle’ in

Hymenoptera Aculeata. Systematic Entomol. 1:177-182.

Burrell, R. W. 1935. Notes on the habits of certain Australian Thynnidae. J. New York Ent.
Soc. 43:19-28.

Duncan, C. D. 1939. A contribution to the biology of North American Vespine wasps. Stanford
Univ. Publ., Biol. Sci., 8(1), 272 p.

Evans, H. E. 1973. Cretaceous aculeate wasps from Taimyr, Siberia. Psyche 80:166-178.

Given, B. B. 1954. Evolutionary trends in the Thynninae (Hymenoptera:Tiphiidae) with
special reference to feeding habits of Australian species. Trans. R. Ent. Soc. London
105:1-10.

Lanham, U. N. 1951. Review of the wing venation of the higher Hymenoptera (Suborder
Clistogastra) and speculations on the phylogeny of the Hymenoptera. Ann. Ent. Soc.
America 44:614-628.

. 1960. A neglected diagnostic character of the Apoidea. Ent. News 71:85-86.

. 1979. Possible phylogenetic significance of complex hairs in bees and ants. J. New

York Ent. Soc. 87:91-94.

Malyshev, S. I. 1968. Genesis of the Hymenoptera and the phases of their evolution. (First
English translation.) Methuen & Co., London. 319 p.

Matthews, R. W. 1968. Nesting biology of the social wasp Microstigmus comes. Psyche
75:23-45.

Snodgrass, R. E. 1942. The skeleto-muscular mechanisms of the honey bee. Smithsonian
Misc. Coll. 103(2), 120 p.

Wheeler, G. C. and J. Wheeler. 1973. Ant larvae of four tribes: second supplement. Psyche
80:70-82.

. 1976. Ant larvae: review and synthesis. Memoir Ent. Soc. Washington, (7), 108 p.

Wilson, E. O. 1962. Behavior of Daceton armigerum (Latreille), with a classification of self-
grooming movements in ants. Bull. Mus. Comp. Zool. Harvard 127:403^422.

. 1971. The insect societies. Belknap Press of Harvard U. Press, Cambridge, Mass.

548 p.

University of Colorado Museum, Box 218, Boulder, Colorado 80309.
Received for publication April 10, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(3), 1980, pp. 210-216

TRANSMISSION OF VIBRATIONS ALONG PLANT STEMS:
IMPLICATIONS FOR INSECT COMMUNICATION

Paul D. Bell

Abstract. — Many insects are known to communicate with percussive vi-
bration and vibrations transduced with acoustic songs via plant tissue. Ar-
tificial percussive vibrations degrade in acceleration and velocity away from
a vibration source along plant stems; the free end distorting and loosing
energy faster than the fixed end. Woody stemmed plants vibrate within a
narrow frequency band and thus transmit signals further than fleshy ones.

Nutritional imperatives, oviposition sites and predator avoidance consti-
tute the major reasons many insects frequent certain species of plants.
Nevertheless, another use of plants to insects may be to act as a commu-
nication channel. Representatives from many insect orders vibrate their
plant substrate during mating and aggressive behavior (Table 1).

It is well documented that insects detect substrate vibrations with sensory
neurons inervating the tympana and subgenual organs (Cokl et al. 1977;
Fudalewicz-Niemezyk et al. 1978; Huber 1978; Kalmring et al. 1978).

Here are investigated the vibration transmission characteristics of various
plants that tree crickets, Oecanthus spp. (Orthoptera: Gryllidae), are found
on north of Huttonville, Ontario, Canada. The acceleration, velocity, wave
form and duration of induced artificial vibrations that approximated O. ni-
gricornis (Walker) percussive vibratory signals were measured from plants
that tree crickets do and do not mate on. The transmission characteristics
and vibrational physics of these plants below (fixed end) and above (free
end) a vibrational source were determined.

Materials and Methods

An Exact 126 signal generator was set to deliver a 5 ms, 100 Hz sinusoidal
pulse every 5 sec. A Marsland woofer converted these pulses into vibrations
via a plastic probe. These signals as modified by the plant were measured
with a Bruel and Kjaer 4344 accelerometer and a B.&K. 2304 impulse pre-
cision sound level meter for acceleration and velocity. The accelerometer
and probe were mounted to the plant with a thin layer of bees’ wax as
outlined by Brock (1972). The vibrations were recorded with a Uher 4000
Report IC tape recorder and displayed on a Tektronix 455 oscilloscope (Fig.
1).

Ten each of orchard grass ( Dactylis glomerata L.), raspberry canes ( Ru -

VOLUME LXXXVIII, NUMBER 3

211

Table 1. Insects employing plant tissue to transduce vibratory signals.

Insect

Plant

Context

Reference

Orthoptera

Neconema thalassinum

leaf?

courtship

Ragge (1965)

Copiphora rhinocerous

stem ?

courtship

Morris (1980)

Neoconocephalus spp.

stem ?

courtship

Whitesell (1969)

Neoconocephalus ensiger

stem ?

aggression

Gwynne (1977)

Conocephalus nigropleurum

stem ?

aggression

Morris (1971)

Mogoplistinae

leaf, stem ?

courtship,

post-

copulation

Love & Walker (1978)

Oecanthus nigricornis

Rubus spp. etc.

courtship,

post-

copulation

Bell (1979a)

Oecanthus burmeisteri

sunflower leaf

calling

Prozesky-Schulze
et al. (1975).

Plecoptera

Perlidae etc.

Phalaris spp.

courtship,

calling

Rupprecht (1968)

Hemiptera

Oncopeltus fasciatus

Asclepias spp.

mating

Walker (1979)

Homoptera

Dictophora europea

leaf?

calling

Strubing (1977)

Euscelis incisus

leaf?

calling

Traue (1978)

Coleoptera

Golofa ported

palm leaf,

stalk aggression

Eberhard (1977)

Eusattus spp.

bark ?

?

Tschinkel & Doyen
(1976)

Brentus anchorago

bark ?

?

Johnson, L. K.
(pers. comm.)

Nothorrhina muricata

bark ?

courtship

Faber (1953)

Diptera

Helius flavipjes

Pile a spp.

male as-
semblages

Zalom (1979)

Euaresta spp.

Ambrosia spp.

calling

Batra (1979)

Liparia spp.

Phragmites spp.

calling

Mook & Bruggemann
(1968)

Tephritis spp.

Senecio spp.

calling

Tauber & Toschi
(1965)

bus spp.), Canada thistle stems ( Cirsium arvense L.), common cattail leaves
( Typha latifolia L.), toadflax (Linaria vulgaris Hill), and goldenrod ( Soli -
dago spp.) were individually tested for their vibration transmission char-
acteristics. The parts of all plants ranged from 30-35 cm in height with a

212

NEW YORK ENTOMOLOGICAL SOCIETY

Fig. 1 . Vibration generation and reception apparatus, gen., signal generator; spkr., speaker;
acc., accelerometer; vm., vibration meter; rec., tape recorder; osc., oscilloscope.

base diameter not varying by more than 0.5 mm within species. The acce-
lerometer was affixed to each plant at 180° to the probe at 2.5, 5, 7.5, 10,
12.5, and 20 cm from the top of the plant base clamp. Canada thistle stems
required shaving for probe and transducer attachment. The probe attached
to each plant 10 cm above the base clamp (Fig. 1).

Results and Discussion

Intraspecific plant vibration transmission variation as indicated by the
standard deviation for each of the sample groups of 10 was small. Variation
was usually greater further away from the probe, especially above (20 cm
above base) (Fig. 2).

Orchard grass, raspberry cane, Canada thistle and goldenrod had nearly
flat vibration transmission curves; while cattail and toadflax had rapid signal
decay with greater distortion away from the probe (Fig. 2, 3). Tree crickets
have never been observed by the author to mate on the latter two species.
Tree crickets mate on all others (Bell, unpublished data). Cattail and toad-
flax also displayed a 'whip’ effect on either side of the probe, i.e., the
intensity of the signal increased momentarily before diminishing. In all
plants excepting raspberry cane the deterioration of the vibrations was faster
at the distal end.

Generally the vibrations were more distorted above the probe. Orchard
grass, goldenrod and to some extent raspberry cane had a wave form re-
sembling a sine wave with a slow decay, that is a vibration which is slowly
dissipating energy over a narrow frequency band, i.e., Tinging.’ This was
especially evident 2.5 cm above the base, thereby allowing these stiff
stemmed plants to transmit vibrations further with more energy than more
flexible stalked plants (Fig. 3).

Substrate vibrational communication in insects has certain advantages
over acoustic, chemical and visual modalities. Predators are known or ex-

VOLUME LXXXVIII, NUMBER 3

213

— 1~- — u -T 1 r— — 1 • 1

Orchard grass

1 1 r 1 1 1 r-

Raspberry cane

“ — l 1 1 1 1 1 1 —

Canada thistle

-2 * * + 4

0 ■ 4

i 4 * * *

+ 4 * 4

4

t

03-

04-

* f

I +

I'

o d>

-5 „ 0 s 0 ,

■J . 0 o

4 o <> 0 $

! C 43 0 0 - if

' f

0

I

Cattail leaf

Toadflax

Goldenrod

?. * • * * *

4 + 4

4 4 4

* 4

* 1

* ^ 1

3

1 ‘

4 .

*

t -

•

. « 4 4 I

t " ‘ * |

o * o 0 o

0

* I

1 1 1 1 1 1 1 — -

I □

I

10

10

10'

2.5 5 75 10 12.5 15 20

5 5 75 10 12.5 15 20

5 5 75 10 125 15 20

Cm. above plant base

Fig. 2. Acceleration (A), and velocity (□) of artificial vibrations. Each point represents the
mean value of 10 plant specimens; bars equal 1 standard deviation, arrows indicate levels
below meter sensitivity.

pected to use acoustic cues to locate insect prey (Bell 1979b; Walker 1964;
Rentz 1975). Certain parasites display positive phonotaxis to singing insects
(Cade 1975; Soper et al. 1975). Thus it may be advantageous for some insects
to reduce the occurrence of acoustic displays. However, selection against
acoustic displays will eliminate some opportunities for mate attraction. Sub-
strate vibration signals have the ability of signalling only conspecific fe-
males, while avoiding detection by predators and other males at longer
distances (Morris 1980; Walker 1979). Acoustic signalling is attenuated, ab-
sorbed, refracted, reflected and diffracted by variations in foliage mass,
height, species composition, humus layer, soil, and atmospheric conditions
(Linskens et al. 1976; Martin and Marler 1977; Wiley and Richards 1978).
In addition the airborne sounds of some insects are highly directional (Leroy
1976; Paul and Walker 1979).

214

NEW YORK ENTOMOLOGICAL SOCIETY

2o*vy^v^

viyvv-

Ifcr-

15 lj\l\fwvv*

i/\/\AA/\fr^^ y^vw-

12.5

J\JVwv^^ — - — -

L- L

probe

7.5

■/yw v\a — i/yv\/\A/v\/^ — — T^y\ /Vvw — - —

ytyw— VlAv^^—

2 . 5 pjm^

^Y\/\Aaa/w' — ~ -j^Vl/vwww

60 ms

Orchard grass Raspberry cane Canada thistle Cattail leaf Toadflax Goldenrod

Fig. 3. Oscillograms of plant transduced artificial vibrations.

Just as in acoustic signalling, the directionality of pheromones will limit
mate attraction to those within the broadcast field. Olfactory signalling has
been shown to function in close range communication, e.g. (Paul 1976), but
would be ineffective in unfavorable air currents.

Visual signalling is limited in many nocturnal insects and hampered diur-
nally in some by dense vegetation (Bell 1979a).

Vibrations on the other hand proceed along plant tissue in front of and
behind the signaller in a predictable manner. In the absence of interfering
(in contact) foliage there is less potential degradation of vibrations over
short (<1 m) distances than of visual and chemical signals. Traue (1978)
reported that vibrations from leafhoppers were perceived 90 cm away
through plant stems. A vibratotactic function for O. nigricornis percussive
vibration signals has been indicated by preliminary experiments. Also, the
temporal integrity of the male tree cricket calling song is preserved as sub-
strate transduced vibrations through the insects legs, and may assist ori-
enting females (Bell, in progress).

The present study has demonstrated that the quality of vibration signals
is dependent on the species of plant involved in transduction. Tree crickets
and many other insects may exploit the vibration transmitting characteristics

VOLUME LXXXVIII, NUMBER 3

215

of plants to signal conspecific mates. It is likely that as plants age, develop
new structure, and succumb to changes in turgor pressure, they begin or
cease to have vibrational characteristics that are advantageous for insect
signalling. Selection would favor insects which could successfully select
a substrate that could transmit their messages over greater distances without
distortion and degradation. These individuals would increase their likelihood
of attracting potential mates only. It is probable that certain plants such as
orchard grass and raspberry cane are frequented by tree crickets because
of their high kQ’ resonance qualities, which allow vibrational energy to be
released over a narrow frequency band, thereby travelling further.

Acknowledgments

I thank Dr. J. H. Fullard and two anonymous reviewers for their sugges-
tions to improve the manuscript. Financial support through the N.R.C.
(Canada) grant #4946 to Dr. G. K. Morris, and O.G.S. and N.S.E.R.C.
postgraduate scholarships to P.B. was appreciated.

Literature Cited

Batra, S. W. T. 1979. Reproductive behavior of Euresta bella and E. festiva (Diptera: Te-
phritidae) potential agents for the biological control of adventive North American rag-
weeds ( Ambrosia spp.) in Eurasia. J. N.Y. Ent. Soc. 87:118-125.

Bell, P. D. 1979a. Mate choice and mating behavior in the black-horned tree cricket, Oecan-
thus nigricornis (Walker). M.Sc. Thesis, University of Toronto, iv, 81 pp.

. 1979b. Acoustic attraction of herons by crickets. J. N.Y. Ent. Soc. 87:126-127.

Broch, J. T. 1972. The Application of the Bruel and Kjaer Measuring Systems to Mechanical
Vibration and Shock Measurements. Larson and Son, Denmark, 311 pp.

Cade, W. H. 1975. Acoustically orienting parasitoids: fly phonotaxis to cricket song. Science.
190:1312-1313.

Cokl, A., K. Kalmring and H. Wittig. 1977. The responses of auditory ventral-cord neurons
of Locusta migratoria to vibration stimuli. J. Comp. Physiol. 120:161-172.

Eberhard, W. G. 1977. Fighting behavior of male Golofa poteri beetles (Scarabeidae: Dynas-
tinae). Psyche 83:292-298.

Faber, A. 1953. Eine Unbekannte art der Lauterzeugung bei Kafern: das Trommeln von
Nothorrhina muricata. Jh. Ver. vaterl. Naturk. Wurttemburg. 108:71-83.
Fudalewicz-Niemczyk, W, A. Petryszak, M. Rosciszewka and M. Olesky. 1978. Sense organs
in some more primitive orders of insects. Prseglad. Zool. 22:123-128.

Gwynne, D. T. 1977. Mating behavior of N eoconocephalus ensiger (Orthoptera: Tettigoni-
idae) with notes on the calling song. Can. Ent. 109:237-242.

Huber, F. 1978. The insect nervous system and insect behavior. Anim. Behav. 26:976-981.
Kalmring, K., B. Lewis and A. Eichendorf. 1978. The physiological characteristics of the
primary sensory neurons of the complex tibial organ of Decticus verrucirorus L. (Or-
thoptera: Tettigoniidae). J. Comp. Physiol., 127:109-121.

Leroy, Y. 1976. Etho-ecologie des communications acoustiques des animaux. C. R. Soc.
Biogeogr. Seance. 457:45-69.

Linskens, H. F., M. J. M. Martens, H. J. G. M. Hendricksen, A. M. Roestenberg-Sinnige, W.
A. J. M. Browvers, A. L. H. C. van der Staak and A. M. J. Strik-Jansen. 1976. The
acoustic climate of plant communities. Oecologia 23:165-177.

216

NEW YORK ENTOMOLOGICAL SOCIETY

Love, R. E. and T. J. Walker. 1979. Systematics and acoustic behavior of scaly crickets
(Orthoptera: Gryllidae: Mogoplistinae) of Eastern United States. Trans. Amer. Ent. i
Soc. 105:1-66.

Martin, K. and P. Marler. 1977. Sound transmission and its significance for animal vocaliza-
tion. I. Temperate habitats. Behav. Ecol. Sociobiol. 2:271-290.

Mook, J. H. and C. G. Bruggemann. 1968. Acoustic communication by Lipara lucens (Dip-
tera, Chloropidae). Ent. Exp. & Appl. 11:397-402.

Morris, G. K. 1971. Aggression in male conocephaline grasshoppers (Tettigoniidae). Anim.
Behav. 19:132-137.

. 1980. Calling display and mating behavior of Copophora rhinoceros Pictet (Orthop-
tera: Tettigoniidae). Anim. Behav. 28:42-51.

Paul, R. 1976. Acoustic response to chemical stimuli in ground crickets. Nature 263:404-405. I

and T. J. Walker. 1979. Arboreal singing in a burrowing cricket, Anurogryllus arbo- i

reus. J. Comp. Physiol. 132:217-223.

Prozesky-Schulze, L. O., P. M. Prozesky, F. Anderson and G. J. J. Van der Merroe. 1975.
Use of a self-made leaf baffle by a tree cricket. Nature 255:142-143.

Ragge, D. R. 1965. Grasshoppers, Crickets and Cockroaches of the British Isles. F. Wayne,
London, xii, 299 pp.

Rentz, D. C. 1975. Two new katydids of the genus Melanotus from Costa Rica with comments
on their life history strategies (Tettigoniidae: Pseudophyllinae). Ent. News 86:129-140.

Rupprecht, R. 1968. The drumming of Plecoptera (Insecta). Z. vergl. Physiol. 59:38-71.

Soper, R. E., G. E. Shewell and D. Tyrell. 1975. Colcondamyia auditrix nov sp. (Diptera:
Sarcophagidae), a parasite which is attracted by the mating call of its host, Okanagana
rimosa (Homoptera: Cicadidae). Can. Ent. 108:61-68.

Strubing, H. 1977. Sound production or substrate vibration as a means of communication in
small Cicadidae. Discussed using as an example Dictyophora europaea (Homoptera-
Cicadina: Fuloroidea). Zool. Beitrage 23:223-232.

Tauber, M. J. and C. A. Toschi. 1965. Life history and mating behavior of Tephritis stigmatica
(Coquillet). Pan. Pac. Entomol. 41:73-79.

Traue, D. 1978. Zur Biophysik der Schallabstrahlung bei Kleinzikaden Jassidae. Zool. Bei-
trage. 24:155-164.

Tschinkel, W. R. and J. T. Doyen. 1976. Sound production by substratal tapping in beetles
of the genus Eusattus (Tentyriidae: Coniontini). Coleop. Bull. 30:331-335.

Walker, W. F. 1979. Mating behavior in Oncopeltus fasciatus : circadian rhythms of coupling,
copulation duration and ‘rocking’ behavior. Physiol. Entomol. 4:275-283.

Walker, T. J. 1964. Experimental demonstration of a cat locating orthopteran prey by the
prey’s calling song. Fla. Entomol. 47:163-165.

Whitesell, J. J. 1969. Biology of the United States cone-headed katydids of the genus Neo- I
conocephalus. M.Sc. Thesis, University of Florida, vi, 73 pp.

Wiley, R. H. and D. G. Richards. 1978. Physical constraints on acoustic communication in
the atmosphere: implications for the evolution of animal vocalizations. Behav. Ecol.
Sociobiol. 3:69-94.

Zalom, F. G. 1979. Notes on male assemblages of Helius flavipes (Macq) with reference to
mating habits (Diptera: Tipulidae). J. Kan. Ent. Soc. 52:553-555.

Department of Zoology, Erindale College, University of Toronto, Missis-
sauga, Ontario, L5L 1C6 Canada.

Received for publication April 30, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(3), 1980, pp. 217-227

OVER-EXPLOITATION OF LARVAL HOST PLANTS BY THE
BUTTERFLIES HELICONIUS CYDNO AND HELICONIUS SAPHO
(LEPIDOPTERA: NYMPHALIDAE: HELICONIINAE:
HELICONIINI) IN COSTA RICA?

Allen M. Young

Abstract. — I examined two small individuals of a larval host plant species
(Granadilla sp.) of Heliconius cydno and H. sapho (Lepidoptera: Nym-
phalidae: Heliconiinae: Heliconiini) in northeastern Costa Rica to determine
approximately the degree to which each one was utilized repeatedly. From
the rather detailed conceptual framework and ecological studies of Heli-
conius butterflies I expected to find low clutch size and exploitation of fresh
meristem for H. cydno and H. sapho as members of the “Granadilla- feed-
ing” species group. The observations on one of the two plants agreed well
with these expectations. On the other host plant individual, however, fe-
males of these species placed unusually high numbers of eggs on available
fresh meristem. In subsequent observations when no apical meristem was
present, no immature stages were found. The subtle interplay of many en-
vironmental factors, including shifting scarcities of fresh meristem on host
plants, determines the intensity of oviposition on a particular plant.

Introduction

In some butterflies females searching for oviposition sites do not always
accurately assess the suitability of a host plant individual in terms of larval
growth and survival (Chew 1977). The pattern of egg placement on individ-
ual host plants by species which oviposit singly is often determined by the
degree of patchiness of the host plant species (Chew 1977; Benson 1978;
Young 1980). In many tropical butterflies such as Parides, females often
place eggs on tiny seedlings of the host plant, clearly too small to support
larval growth (Young, pers. obs., 1969-80, Costa Rica). But in some groups
such as the Heliconiinae exploiting the Passifloraceae as larval host plants
(Brower and Brower 1964; Gilbert 1975; Benson et al. 1976, and several
other references) there appears to be careful assessment of host plant in-
dividuals by female Heliconius (Benson et al. 1976; Benson 1978). It is
therefore of considerable interest to report here patterns of repeated ex-
ploitations of individuals of a host plant of Heliconius cydno in Costa Rica.
An incidence in which a related species, Heliconius sapho, overexploited
one of the host plants used by H. cydno is also reported.

218

NEW YORK ENTOMOLOGICAL SOCIETY

Materials and Methods

I observed female H. cydno deposit eggs on two immature individuals of t
Granadilla in northeastern Costa Rica. I also discovered a large number of
eggs of H. sapho on one of these plants. The general locality is a 4,000-acre
farm complex about 8 km north of La Virgen, Heredia Province, a region
within the premontane tropical wet forest zone (220 m elev.) (Holdridge
1967). Information on the biology of H. cydno and H. sapho in Costa Rica
has been summarized elsewhere (Smiley 1978; Young 1973, 1978). Both
species are abundant along borders of forest.

The initial observations of H. cydno at one host plant were made on
February 14, 1977. I marked the location of the plant, took notes on its size j
and appearance (number of fresh and old leaves, herbivore damage, etc.),
and returned many times until March 6, 1977 to make observations on the
eggs and larvae present. I examined the patterns of larval feeding and any
interactions among larvae.

The specific locality of the first plant is “Finca La Tigra,” mixed cacao
( Theobroma cacao L.) plantations and various stages of forest succession.
On August 3, 1978 I discovered H. cydno ovipositing on another individual
of the same plant species but this time at “Finca El Uno,” an area of cacao,
Hevea rubber, and strips of young secondary succession about 5 km from
the “La Tigra” site. I recorded the immature stages present and made
observations until August 8. The location of this plant was marked and I
returned to examine for the presence of H. cydno or other Heliconius in
five additional periods: December 1-4, 1978; March 12-20, 1979; June 30,
1979; September 11, 1979; and February 21-22, 1980. Each time I recorded
the appearance of the plant, noting presence of fresh leaves and other fea-
tures. Upon returning to the site of the “El Uno” plant on June 13, 1980 I
noted the presence of new meristem and many eggs scattered on it. Obser-
vations were made through July 2 during which it was determined that the
species was H. sapho. I observed the abundance of larvae and their distri-
bution on the plant.

Results

The host plant is an immature Granadilla , a subgenus in the passiflora-
ceous subfamily Laurifoliae, which consists chiefly of small, erect plants
without tendrils (Dr. K. S. Brown, Jr., pers. comm.). The plant is possibly
Passiflora guazumaefolia. At this locality H. cydno oviposits on at least
one other Passiflora, this time a vine, P. vitifolia with different taxonomic
affinities within the family (Young 1978). Owing to the tentative nature of
host plant identifications I refer to them as “ Granadilla A” (La Tigra) and
“ Granadilla B” (El Uno).

“ Granadilla A”. — This individual was found in a light gap along a well-

219

VOLUME LXXXVIII, NUMBER 3

shaded footpath (Fig. 1A) in disturbed primary forest and at the time of its
discovery it had four fresh leaves (new meristem) (Fig. IB). The plant was
about 0.25 m tall (Fig. 2A) with few signs of herbivore attack to both older
leaves or fresh meristem. At 1130 hours on February 14, 1977 one "fresh”
female H. cydno was noticed inspecting the plant and two eggs were placed
on it within a 20-minute period. The first egg was deposited at 1140 hours
next to the midrib on the ventral side of one of the apical fresh leaves; a
second egg was already present on the same leaf. Another egg was placed
on the petiole of a fresh leaf (Fig. 2B) ten minutes later. No additional eggs
were found. Judging from the white color of the egg already present, this
egg was probably a few days old and possibly not from the same female
witnessed placing two eggs. Fresh eggs of H. cydno are yellow (Young
1973). This plant had a total of ten leaves, including the fresh ones. No
other Heliconius eggs or larvae were found on the plant.

The following day the older egg hatched and matched the general descrip-
tion for the first instar of H. cydno (Young 1973). This larva rested on the
tip of the fresh leaf bearing the two eggs. Later that day one of the other
eggs disappeared, possibly the result of predation by ants or cannibalism by
the larva present. At 1530 hours that day, an ant approached the larva and
it dropped on a silken thread from the leaf. Several small reddish-brown
(unidentified) ants were seen at the conspicuous extra-floral nectaries on the
petiole of one of the lower yet fresh leaves. On the following day (February
16) the larva had eaten away a strip of fresh leaf tissue on the leaf bearing
the surviving egg. Later that day a new egg was found on the petiole of
another fresh leaf.

The first instar begins feeding from the edge of a fresh leaf, cutting deep
notches into the tissue and eventually reaching the midrib area. This larva
stayed on the same fresh leaf for all five instars, completing molt cycles on
it and eventually developing a feeding pattern in which entire sections of
the leaf are chewed away (Fig. 3 A, B). Prior to pupating it eventually moved
onto two other fresh leaves at the top of the plant and attaining a body
length of about 35 mm. The two additional eggs present disappeared and
this larva was the only one on this plant for the study period.

I noticed that the larva fed both day and night, but intermittently and with
no consistent pattern. The larva eventually pupated on an older leaf of the
host plant. It is estimated that the larvae ingested approximately 140 mm2
of fresh leaf tissue by the time of pupation. Older leaves were not consumed.
The portion of fresh leaf tissue consumed represents about 60% of the total
amount of fresh leaf tissue available on the plant during the study period.
No additional fresh leaves were produced during this period. Other than the
total of four eggs counted in the first few days, no additional eggs were
found thereafter.

“ Granadilla B”. — This plant was discovered again as the result of an H.

220

NEW YORK ENTOMOLOGICAL SOCIETY

Fig. 1. (A) The forest habitat understory where “ Granadilla A” was discovered. (B)

“ Granadilla A” showing the fresh meristem (stem and leaves) contrasting in shade from older
leaves.

VOLUME LXXXVIII, NUMBER 3

221

Fig. 2. (A) The fresh leaves and other parts of meristem on “ Granadilla A”; note few

signs of any appreciable herbivore damage. (B) Positions of two eggs of Heliconius cydno on
the fresh meristem of “ Granadilla A”; note that one egg is near the midrib of a young leaf
while the second egg is at the base of the petiole; note the conspicuous extrafloral nectaries
on a petiole to the right.

222

NEW YORK ENTOMOLOGICAL SOCIETY

Fig. 3. (A) Note the damage from feeding by the larva of H. cydno on the fresh leaf in the

lower left. (B) Fouth instar larvae of H. cydno with typical pattern of damage to a fresh leaf
on 'Granadilla A”.

cydno female inspecting it (August 3, 1978, 0930 hours). It was found in a
strip of dense herbaceous vegetation along a shaded trail within a cacao
plantation. The plant was about 0.30 m tall when discovered and of a total
of 12 leaves present six were fresh meristem. As with the first plant, the
fresh leaves are easily distinguished from the older ones by their light green
color and soft texture. The plant was actually a sucker shooting up from a
thick woody stem.

A tattered female of H. cydno deposited a total of 15 yellow eggs on the
apical stem of this plant (Fig. 4A) within a ten-minute period on the morning
of discovery. The butterfly made four different visits to the plant, after flying
off and fluttering through nearby vegetation before returning to deposit more
eggs. The eggs were placed singly (Fig. 4B). Within five days after ovipo-
sition the folded apical region of the host plant had “leafed out” (Fig. 4C),
making a spectacular display of bright yellow eggs packed into a relatively
small area of fresh leaflets and stem. The result is a “loose” clustering of
many eggs on the apical and fresh portion of the plant (Fig. 4C). An ex-
amination of the plant following the oviposition revealed no additional eggs
or larvae, although an empty pupal shell of Heliconius was found on a
seedling adjacent to the host plant. At the time of oviposition a raiding
column of army ants ( Eciton sp.) passed within a few centimeters of the
host plant but did not affect oviposition.

VOLUME LXXXVIII, NUMBER 3

223

Fig. 4. (A) Heliconius cydno in the act of placing several eggs on the fresh meristem of

“ Granadilla B”. (B) Egg of H. cydno on the fresh petiole of “ Granadilla B”. (C) The distri-
bution of 15 eggs of H. cydno on the unfolded fresh leaflets and stem of “ Granadilla B”.

Egg number remained the same for the first four days following the burst
of oviposition activity. By the fifth day, however, there were only eleven
eggs and no first instar larvae to account for the remaining four. One of the
remaining eggs was shriveled up. The same plant was re-examined in De-
cember and two larvae (second instar and third instars) of H. cydno were
present on the plant (December 1, 1978). No eggs were present. At this time
the plant was about 30 cm tall with roughly 50% of the biomass being fresh.
The two larvae occupied different leaves. Several ants were seen patrolling
the apical area of the plant on two days following the discovery. For the
period March 12-20, 1979 fresh leaves were entirely absent from the plant
and no Heliconius early stages were found. The plant was about the same
size as it was in the previous December although no new growth was pres-
ent. I noticed three fresh adults of Philaethria dido (Heliconiinae) flying in

224

NEW YORK ENTOMOLOGICAL SOCIETY

the vicinity of the plant on one day. The general condition of the herbaceous
vegetation was lush at this time. The plant was in the same general condition
on June 30, 1979 and no early stages of H. cydno were found on it. The
plant was briefly observed in September 1979 and February 1980 and again
there was no new growth and no signs of H. cydno on it.

The plant was re-examined on June 13, 1980 at which time it had fresh
meristem about 40 cm high and bearing eight full-sized and folded leaflets.
The bottom woody part of the plant had ten old leaves. A total of 20 Hel-
iconius egg shells were found scattered on the fresh leaflets and there were
14 second instar larvae in a tight aggregation on one leaflet. No unhatched
eggs were present. By June 16 only eight larvae remained and the leaflets
were now completely unfolded and the egg shells gone. No observations
were made until June 29 at which time it was discovered that all of the fresh
meristem had disappeared and that only one chrysalis was present on an
older leaf. A thorough search of the surrounding vegetation within a four-
meter square area failed to turn up additional chrysalids. The chrysalis
hatched July 2 and proved to be H. sapho.

Discussion

Heliconius cydno and H. sapho are species in the “ melpomene ” group,
being closely allied with other species such as H. pachinus, melpomene ,
and heurippa (Brown and Mielke 1972; Benson 1978). The species are clas-
sified as laying medium to large eggs, solitary meristem feeders as a larvae,
and generally requiring large host plant individuals (Benson 1978). The
clutch size is generally small (Benson 1978) and a species such as H. mel-
pomene places eggs singly on subterminal leaflets and young tendrils (Alex-
ander 1961). The usage of meristem tissues by Heliconius species comprises
a major aspect of the radiation of the group (Benson et al. 1976). Heliconius
cydno is oligophagous while the allied H. melpomene in Costa Rica is mon-
ophagous (Smiley 1978). The species oviposits on at least one non -Grana-
dilla host plant at the site of the present study (Young 1978) as it does in a
Costa Rican mountain wet forest site (Young 1973). Passiflora vitifolia is
a common and widespread host plant of H. cydno and several other Heli-
coniinae at the study site (Young 1978) as well as at nearby k Tinea La
Selva” (Smiley 1978). Twenty-seven species of Granadilla are used by at
least eight species in the “transitional” and melpomene groups (Benson et
al. 1976).

Many species in the “Granadilla- feeding” group (Smiley 1978) place eggs
singly on fresh meristem and the larvae feed primarily on these tissues
(Benson et al. 1976). The availability of fresh meristem is a limiting factor
in the populations of some Heliconius species (Benson 1978; Smiley 1978).
The butterflies are adapted for searching for and placing eggs on this portion
of the individual host plant (Gilbert 1975).

VOLUME LXXXVIII, NUMBER 3

225

Assuming that the observed host plant was indeed P. guazumaefolia and
the fact that many species in the Granadilla -lineage of the Passifloraceae
have conspicuous extrafloral nectaries that attract ants (Gilbert 1975; Ben-
son et al. 1976) the observed loss of eggs, apparently from ant predation,
is not unexpected. It is also known that some species of Granadilla possess
stipules resembling Heliconius eggs and larvae (Benson et al. 1976) thus
discouraging repeated oviposition in Heliconius which place eggs singly on
meristem. Such factors reduce the likelihood that a female Heliconius will
oviposit repeatedly on the meristem of a particular individual of the host
plant species available in the habitat. The species of Heliconius exploiting
such host plants generally have aggressive larvae so that there is usually no
more than one larva per meristem (Benson et al. 1976).

Yet “Granadilla B” presents a different picture. The observed placement
of fifteen eggs on a single fresh meristem by H. cydno is a notable departure
from the expected clutch size. The usual clutch size of this species is small
(1-3 eggs per plant) (Benson 1978; Young 1978). Presumably the plant was
too small to support all of the larvae. Perhaps ant predation and aggressive
interactions among the larvae would have taken a heavy toll thus reducing
effective clutch size considerably. The apparent placement of 20 eggs on
fresh meristem of the same plant by H. sapho and the subsequent dwindling
of larvae in this gregarious species is a further indication of the interaction
of reduced food supply and possibly predation as reducing larval numbers
in this species. Presumably the eggs were placed in a relatively loose cluster
! at an even earlier stage in the unfolding of the fresh meristem. The data
suggest that the larvae defoliated the plant of the fresh meristem and that
only a portion of them survived.

Why should some Heliconius “invest” about five times the expected
number of eggs in a small individual of Granadilla ? The answer is clearly
beyond the scope of this paper, yet it is interesting to suggest one expla-
nation based upon the concept of limiting resources affecting the butterfly
at the study site. The consecutive choices by a female butterfly searching
for oviposition sites is influenced greatly by the spatial distribution of suit-
ably host plant individuals (Chew 1977; Benson 1978; Young 1980). If host
plant individuals are very patchy and concealed, one might expect exploi-
tation of alternative host plant species or more intense exploitation of the
individual host plant when encountered. Although several species of host
plant might be available in the habitat, their suitability for oviposition by
meristem specialists such as H. cydno and H. sapho is determined largely
by the availability of fresh meristem tissues. My general impression of the
relative abundance of P. vitifolia to P. guazumaefolia at the locality of the
present study is that the former species is far more abundant. There seems
to be a greater number of patches per unit of habitat and the biomass of
each patch is considerably larger. Whether or not H. cydno and H. sapho
exhibit a physiological preference for one species over the other is not

226

NEW YORK ENTOMOLOGICAL SOCIETY

known, although one study showed that the H. cydno exhibits approxi-
mately equal oviposition frequency on several host plant species in different
subgenera (Smiley 1978). Therefore the differential in terms of actual host
plant usage in the wild is probably largely determined by relative patch
structures among host plant species and the availability of fresh meristem
among the species.

The scarcity of Granadilla ( P . guazumae folia) coupled with perhaps re-
duced availability of fresh meristems on other host plants such as P. vitifolia
may induce repeated oviposition on small patches of those host plants with
fresh meristem. Chew (1977) noted that there is sometimes considerable
discrepancy between oviposition behavior and the suitability of the chosen
host plant individual for larval growth. I have noted similar situations in
Parides (Papilionidae) ovipositing on Aristolochia seedlings and Morpho
peleides oviposit on tiny seedlings of Machaerium seemanii in Costa Rica.

Acknowledgments

This research was funded by grants from the Friends of the Museum of
the Milwaukee Public Museum. I thank Dr. Keith S. Brown, Jr. for making
an attempt to identify the host plant, although he bears no responsibility for
the tentative determination as discussed in this paper should it prove to be
erroneous.

Literature Cited

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

Benson, W. W. 1978. Resource partitioning in passion vine butterflies. Evolution 32:493-518.

, K. S. Brown, Jr. and L. E. Gilbert. 1976. Coevolution of plants and herbivores:

Passion flower butterflies. Evolution 29:659-680.

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

Brown, K. S., Jr. and O. H. H. Mielke. 1972. The heliconians of Brazil (Lepidoptera: Nym-
phalidae). Part II. Introduction and general comments, with a supplementary revision
of the tribe. Zoologica 47:1-40.

Chew, F. S. 1977. Coevolution of pierid butterflies and their cruciferous foodplants. II. The
distribution of eggs on potential foodplants. Evolution 31:568-579.

Gilbert, L. E. 1975. Ecological consequences of a coevolved mutualism between butterflies
and plants, p. 210-240. In: L. E. Gilbert and P. H. Raven, eds., Coevolution of Animals
and Plants. University of Texas Press, Austin. 246 p.

Holdridge, L. R. 1967. Life Zone Ecology. Tropical Science Center, San Jose, Costa Rica.
104 p.

Smiley, J. 1978. Plant chemistry and the evolution of host specificity: new evidence from
Heliconius and Passiflora. Science 201:745-747.

Young, A. M. 1973. Notes on the biology of the butterfly Heliconius cydno (Lepidoptera:
Heliconiinae) in Costa Rica. Wasmann J. Biol. 31:337-350.

VOLUME LXXXVIII, NUMBER 3

227

1978. “Disappearances” of eggs and larvae of Heliconius butterflies (Nymphalidae:
Heliconiinae) in northeastern Costa Rica. Entomol. News 89:81-87.

-. 1980. Evolutionary responses by butterflies to patchy spatial distributions of resources
in tropical environments. Acta Biotheoretica 29:37-64.

Section of Invertebrate Zoology, Milwaukee Public Museum, Milwaukee,
Wisconsin 53233.

Received for publication June 10, 1980.

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INDEX OF AUTHORS

BELL, PAUL D. Transmission of vibrations along plant stems: implications for insect

communication 210

CONWAY, JOHN R. The seasonal occurrence of sexual brood and the pre- and post-
nuptial behavior of the honey ant, Myrmecocystus mexicanus Wesmael, in Colorado 7

EGER, JOSEPH E., JR. Fecelia biorbis n. sp. (Heteroptera: Pentatomidae), a new

species from Haiti 29

ELLIOT, NANCY B. and PAUL SALBERT. Observations on the nesting behavior
of Tachytes tricintus (F.) on San Salvador Island, Bahamas (Hymenoptera: Sphecidae,

Larrinae) 170

FORD, FRANCIS C. and JAMES FORBES. Anatomy of the male reproductive systems
of the adults and pupae of two doryline ants, Dorylus ( Anomma ) wilverthi Emery and

D. (A.) nigricans Illiger 133

GIBO, DAVID L. Apparent nest site competition between the paper wasp Polistes

fuscatus (Hymenoptera: Vespidae) and the house wren 143

GIBO, DAVID L. Overwintering of Polistes fuscatus in Canada: use of aban-
doned nests of Dolichovespula arenaria 146

HENDRICKSON, R. M., JR. Continuous production of predacious mites in the green-
house 252

JILEK, REID. Epistylus cambari (Ciliata: Peritrichida) and dragonfly nymphs, an epi-

zoic association 113

KRAMER, JOHN PAUL. The house-fly mycosis caused by Entomophthora muscae:

influence of relative humidity on infectivity and conidial germination 236

LANHAM, U. N. Evolutionary origin of bees (Hymenoptera: Apoidea) 199

MATOUSCHEK, M., A. DUBENDORFER, and E. KUBLI. Studies on the incorpo-
ration of [32P]-phosphate into low molecular weight RNA of Drosophila melanogaster

cell lines 241

McCABE, T. L. and C. J. SHEVIAK. Platanthera cristata (Michx.) Lindl., a new

host for the red-banded leaf roller 197

MINTZER, ALEX. Simultaneous use of a foraging trail by two leafcutter ant species

in the Sonoran desert 102

MUKESH, DEVINDER, DALDINDER SINGH SIDHU and NIRMAL KUMAR. Total
concentration of free amino acids and the corresponding morphogenetic changes
during the embryogenesis of Chilomense sexmaculata Fabr. (Coleoptera: Coccinellidae) 1 1 5

l

PIRONE, DOMINICK, J. and DANIEL J. SULLIVAN, A. J. Ecology of Necrophilous

and Carpophilous Coleoptera in a southern New York woodland. Part 1 — overview 1 86

RHOMBURG, LORENZ. Causes of life history differences between the morphs of

Pemphigus populitransversus 106

ROLSTON, L. H., F. J. D. McDONALD, and DONALD B. THOMAS, JR. A con-
spectus of Pentatomini genera of the western hemisphere. Part 1 (Hemiptera: Penta-

tomidae) 120

ROLSTON, L. H. and F. J. D. McDONALD. Conspectus of Pentatomini genera of

the Western Hemisphere — Part 2 (Hemiptera: Pentatomidae) 257

SCHAEFER, CARL M. The sound-producing structures of some primitive Penta-

tomoidea (Hemiptera: Heteroptera) 230

SCHRODER, ROBERT F. W. and WILLIAM W. METTERHOUSE. Population trends
of the alfalfa weevil (Coleoptera: Curculionidae) and its associated parasites in Mary-
land and New Jersey, 1966-1970 151

SHYAMALANATH, S. and JAMES FORBES. Digestive system and associated organs
in the adult and pupae male doryline ant Aenictus gracilis Emery (Hymenoptera:

Formicidae) 15

TREAT, ASHER E. SuZan Noguchi Swain (Mrs. William Kenneth Firmage) 168

TSAI, JAMES H. and DENNIS D. KOPP. Life history, morphology, and taxonomy

of Acutalis tart area (Say) (Homoptera: Membracidae) 174

YOUNG, ALLEN M. Environmental partitioning in lowland tropical rain forest cicadas 86
YOUNG, ALLEN M. Over-exploitation of larval host plants by the butterflies Heli-
conius cydno and Heliconius sapho (Lepidoptera: Nymphalidae: Heliconiinae: Heli-

coniini) in Costa Rica? 217

Book Reviews

MARAMOROSCH, KARL. Comprehensive virology 12. Newly characterized protist

and invertebrate viruses. Heinz Fraenkel-Conrat and Robert R. Wagner, eds. 83

SHAPIRO, ARTHUR M. Herbivores: Their interaction with secondary plant metab-
olites. G. A. Rosenthal and D. H. Janzen, eds. 164

ii

Journal of the

New York Entomological Society

VOLUME LXXXVIII DECEMBER 1980

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. Streu

Publication Committee
Dr. Randall T. Schuh
American Museum of Natural History
Dr. Daniel Sullivan
Fordham University
Dr. Felix J. Bocchino
The College of Mt. Saint Vincent

CONTENTS

The sound producing structures of some primitive Pentatomoidea (Hemiptera: Het-

eroptera) Carl W. Schaefer 230-235

The house-fly mycosis caused by Entomophthora muscae : influence of relative hu-
midity on infectivity and conidial germination John Paul Kramer 236-240

Studies on the incorporation of [32P]-phosphate into low molecular weight RNA of
Drosophila melanogaster cell lines

M. Matouschek, A. Diibendorfer and E. Kubli 241-251

Continuous production of predacious mites in the greenhouse

R. M. Hendrickson, Jr. 252-256

Conspectus of Pentatomini genera of the Western Hemisphere — Part 2 (Hemiptera:
Pentatomidae) L. H. Rolston and F. J. D. McDonald 257-272

Acknowledgment, List of Honorary, Life, and Sustaining Members 273

Index to Scientific Names of Animals and Plants for Volume LXXXVIII 275-278

i-ii

Index of Authors for Volume LXXXVIII

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(4), 1980, pp. 230-235

THE SOUND-PRODUCING STRUCTURES OF SOME PRIMITIVE
PENTATOMOIDEA (HEMIPTERA: HETEROPTERA)

Carl W. Schaefer

Abstract. — The Cydnidae and Thyreocoridae have a small stridulitrum on
the under side of the hindwing’s postcubitus. The plectrum is a striated lima
on the first tergite. Possession of the stridulitrum and plectrum is probably
plesiomorphic in a complex of primitive Pentatomoidea.

The Thaumastellidae (a member of the complex) also have a stridulitrum,
near the postcubitus of the hindwing, and a similar plectrum. Some Scutel-
leridae and Tessaratomidae have a similar stridulitrum. Other morphological
features ally these two families with — but are insufficient to warrant includ-
ing them in — this complex of primitive pentatomoids.

Members of the families Cydnidae and Thyreocoridae, both primitive in
the Pentatomoidea, possess a stridulitrum on the lower surface of the post-
cubital vein of the hindwing (Dupuis 1953; Leston 1954, 1957; Lawson and
Chu 1971; Draslar and Gogala 1976). The stridulitrum is small (less than a
millimeter long) and composed of cuticular ridges. The part of the vein that
bears it is raised, darkened, and more heavily sclerotized than the rest of the
vein. The plectrum across which presumably the stridulitrum strums, is
a striated lima on the anterior edge of the first abdominal tergite (Draslar and
Gogala 1976). Muir (1907) and Leston (1954) discuss the functioning of the
stridulitrum and plectrum, and Gogala (several references: see 1978) de-
scribes and characterizes the sounds produced and discusses their functions.

The possession of the stridulitrum by these families assumes greater in-
terest now that they appear to be part of a larger complex of primitive
Pentatomoidea (Thaumastellidae, Plataspidae, Cyrtocoridae, Megarididae,
Canopidae, and perhaps Lestoniidae [Schaefer, in preparation]), and now
that Stys (1964) has described a similar stridulitrum in the Thaumastellidae.
Moreover, some (but not all) members of two other families, not members
of the complex, also have a postcubital stridulitrum. Some, perhaps most,
Tessaratomidae have such a stridulitrum (Muir 1907; Dupuis 1953, Au 1969),
and so do certain Scutelleridae (Au 1969). Spermathecal features ally parts
of these families to the group of primitive pentatomoids (Schaefer, in prep-
aration).

The distribution of the postcubital stridulitrum is clearly important. Un-
fortunately, when Leston (1957, p. 372) wrote that he had found it in “a
further twenty-five genera” of Cydnidae, he did not name them. His concept
(1961) of the Cydnidae differs from the more generally accepted one of
Froeschner (1960), and one cannot be sure if Leston found such a striduli-

VOLUME LXXXVIII, NUMBER 4

231

trum in the Amnestinae, the Scaptocorinae, or the Garsauriinae. Moreover,
his description (1954) of the plectrum in several of these bugs requires some
amplification.

Here I describe briefly the stridulitrum in the Cydnidae (except Garsau-
riinae, of which I had no representatives) and Thyreocoridae, and discuss
the cladistic significance of its presence in these families and the Thaumas-
tellidae.

Results

The presence and some characteristics of the postcubital stridulitrum are
given in Table 1.

The size and number of teeth of the stridulitra of all families vary consid-
erably, as does their “packing”; but the ridged form of the teeth and — more
importantly — the overall shape of the stridulitrum itself, are constant; al-
though, in Tessaratoma, the teeth are more chisel-like and less ridge-
shaped. Variation probably occurs between sexes and among species, as
Draslar and Gogala (1976) have shown for two species (of different subgen-
era) of Sehirus .

As Au (1969) remarks, the possessors of the stridulitrum can be placed
into either of two groups: those with many teeth (Scaptocorinae, Amnesti-
nae, Thyreocoridae, and Scutellerini) and those with few teeth (Cydninae,
Sehirinae, Thaumastellidae, and Tessaratomidae) (see Table 1).

The plectra of all species studied, except those in the Thaumastellidae
and Scutelleridae, is a very small striated lima located sublaterally on the
anterior border of the first tergite. Its position does not change from species
to species, although its development does (see scanning electron micro-
graphs in Draslar and Gogala 1976); the striations in Amnestus are so fine
as to be invisible except at high (200 x) magnification.

Leston (1954) places the plectrum in the Sehirinae on the third metatho-
racic phragma (scutum 3), but admits there is some confusion about the
naming of the sclerites in this region. In his figure 4, he placed the plectrum,
correctly, on the anterior border of the first tergite, and not on phragma 3.

As in the others, the plectrum of Tessaratoma papillosa is on the first
abdominal tergite, not on a “sclerite . . . between the metathorax and first
abdominal segment” (Muir 1907, who misnumbered the segments; an error
repeated by Au 1969). The plectrum is paired and resembles those of the
Cydnidae and Thyreocoridae closely, although it is more medially situated
than are theirs.

The plectrum of the Thaumastellidae closely resembles those above, but
occurs on the second abdominal tergite, not the first.

The scutellerid plectrum (in C allidea duodecimpunctata ) is as Au (1969)
describes and figures it for Scutellera nobilis Distant: “a heavily sclerotized

232

NEW YORK ENTOMOLOGICAL SOCIETY

Table 1 . Characteristics of the stridulitrum of some primitive Pentatomoidea.

Species

Total teeth

No. teeth/mm

Jugal

hole

Reference

Cydnidae: Cydninae
Aethus indicus (Westwood) (9)

18

65

no

this paper

A. indicus (9, #')

17-20

' —

—

Leston, 1954

Pangaeus bilineatus (Say) (9, 6)

30-40

130*

no

this paper

Microporus obliquus Uhler

30-35

(45/mm
distally,
190/mm
proximally)
90/ mm

no

Lawson &

Cydnidae: Sehirinae

Sehirus cinctus (Palisot de

12-15 or 32*

distally;

250/mm

proximally

72-75

no

Chu, 1971
this paper

Beauvois ( 9)
Sehirus bicolor (L.)

17-19

Leston, 1954

Dismegistus fimbriatus Thunberg

16-18

50

Leston, 1956

Cydnidae: Scaptocorinae
Scaptocoris divergens

84

(estimate)

150

no

this paper

Froeschner (d)
Atarsocoris gisellae

65

150

no

this paper

(Carvalho) (<3)

Cydnidae: Amnestinae
Amnestus spinifrons (Say) (6)

70

380

no

this paper

Thyreocoridae

Thyreocoris scarabaeoides (L.) (<3)

60

250

9

this paper

Corimelaena pulicaria (Germar) (<3)

ca. 200

ca. 700

9

this paper

Galgupha ovalis (Hussey)

ca. 100

ca. 200

yes

Lawson &

Thaumastellidae

Thaumastella aradoides Horvath

44

Chu, 1971
Stys, 1964

Scutelleridae

Callidea duodecimpunctata F. (9 , c?)

300

250

no

this paper

Tessaratomidae

Tessaratoma javanica Thunberg (<3)

12-15

10

no

this paper

Cyrtocoridae
Cyrtocoris laceratus
Herrich-Schaeffer
C. sp.

Plataspidae

Coptosoma cribraria (F.)
Brachyplatys vahlii (F.)

Megarididae
Megaris sp.

no

stridulitrum

no

stridulitrum

no

stridulitrum

no

stridulitrum

no

. stridulitrum

this paper
Au, 1969

this paper
Au, 1969

this paper

* Two specimens; possibly sexually dimorphic.

VOLUME LXXXVIII, NUMBER 4

233

file-like sclerite”; however, it is not on the “anterolateral area of the first
abdominal tergum” but (as indeed she figures it), single and medial on the
tergum. The plectrum in both Scutellera and Call idea is much larger than
it is in the Thaumastellidae, Cydnidae, Thyreocoridae, and Tessaratomidae;
in those scutellerids it occupies the medial fifth of the first tergite’s anterior
border. Its striations run front- to-back, not side-to-side, as do those of the
other four families.

Leston (1957) remarks that the jugal wing area of the Cydnidae is smaller
than that of the Thyreocoridae and, when folded under, does not overlap
the stridulitrum and intervene between it and the plectrum. Therefore, the
cydnid jugal area lacks the hole McAtee and Malloch (1933), and Leston
(1954), believe to characterize the Thyreocoridae, and Lawson and Chu
(1971) describe for Galgupha. I too have not seen such holes in the Cydnidae
(Table 1), but Draslar and Gogala (1976) show that they accommodate indi-
vidual stridulitral teeth in this family, and are so small as to be undetectable
at the magnifications usually used in dissecting. Muir (1907) mentioned
abrasions in this region of the Tessaratoma papillosa Thunberg wing. Au
(1969) did not report holes in the Scutelleridae nor did I find them in
Callidea duodecimpunctata .

Discussion

There are three groups of pentatomoids: those without the postcubital
stridulitrum, those with it and a first-tergal plectrum, and one family (Thau-
mastellidae) the homology of whose stridulitrum and plectrum is problem-
atic.

Those families lacking the PCu stridulitrum include the Pentatomidae,
Urostylidae (Urostylinae and Saileriolinae), Acanthosomatidae, some Scu-
telleridae, and some Tessaratomidae (unpublished observations); other fam-
ilies probably lack it, but have not been studied.

The location and structure of both the stridulitrum and the plectrum are
so similar in the Cydnidae and Thyreocoridae, that it seems unlikely to have
evolved more than once: the character unites the two families, as Dupuis
(1953) and Leston (1954 and elsewhere) have emphasized. However, the
presence of both structures in some Tessaratomidae — a group in other re-
spects not close to those two families — weakens the argument that the com-
mon possession of these structures requires close common ancestry of their
possessors.

The stridulitrum of Thaumastella aradoides Horvath (Thaumastellidae)
is just anterior to, not on, the postcubitus (Stys 1964); nevertheless, the
general position of this stridulitrum, and its structure, suggest strongly it is
homologous with those of the Cydnidae and Thyreocoridae. The striduli-
trum of the strongly brachypterous T. namaquensis Schaefer and Wilcox
may be at the second-third tergal border, but this is uncertain (Schaefer and

234

NEW YORK ENTOMOLOGICAL SOCIETY

Wilcox 1971). However, the plectra of both species are alike, a small striated
area not on the first tergite but on the second.

Despite these differences in position, I believe the thaumastellid stridu-
litrum (of T. aradoides ) and plectra to be homologous with those of the
Cydnidae and Thyreocoridae, because the structures are so similar in all
three families. Accepting this position argues for the inclusion of the Thau-
mastellidae in the unit containing the other two families. However, penta-
tomoid families phylogenetically closer to the Thaumastellidae than these
two lack the stridulitrum and the plectrum. Therefore, I consider possession
of a postcubital stridulitrum and abdominal plectrum to be an autapomorphy
helping to define a complex of primitive Pentatomoidea, a complex defined
also on other characters (Schaefer, in preparation). Within the complex the
character is symplesiomorphic, and the stridulitrum and plectrum have been
lost in several members (Table 1); but loss has not occurred so frequently
(only three times) as to vitiate my argument that stridulitrum and plectrum
have indeed evolved only once.

The postcubital stridulitrum occurs in some 12 genera of the Scutelleridae
(Au 1969), and so presumably does the first-tergal plectrum, although Au
did not check for its presence in all genera. Ten of the 12 genera belong to
the Scutellerini, as defined by Schouteden (1904), who treats the scutellerids
as a subfamily, not a family. One exception is Steganocerus, considered a
member of the Scutellerinae: Sphaerocorini until Leston (1952) transferred
the genus to his more restricted Scutellerini (a fact apparently overlooked
by Kumar [1965]). The other exception, Chiastosternum, is listed next to
Steganocerus by Schouteden in his list of Sphaerocorini.

The stridulitrum appears not to occur in the other scutellerid subfamilies,
the other scutellerine tribes, and certain members of the Scutellerini ( sensu
Leston) itself.

In spite of the differences between the plectrum of the Scutellerini and
those of the Cydnidae, Thyreocoridae, and Tessaratomidae, the placement
of all plectra on the first abdominal tergite indicates, tentatively, they are
homologous. That they are in fact homologous must await a cladistic study
of the Scutelleridae vis a vis these other pentatomoids, a study based on
other characters.

I do not believe either the Scutelleridae or the Tessaratomidae belong to
the group of primitive Pentatomoidea of which the Cydnidae, Thyreocori-
dae, and Thaumastellidae are members, although some scutellerids (and
possibly some tessaratomids) share some spermathecal characteristics with
some members of the group (Schaefer, in preparation). It seems to be pos-
sible that the two families evolved from members of the primitive group,
and that the stridulitrum and plectrum are plesiomorphic in the Scutelleridae
and Tessaratomidae. In this regard it is significant that Kumar (1965) con-
siders the Scutellerinae (= Scutellerini of Schouteden) to be primitive with

VOLUME LXXXVIII, NUMBER 4

235

respect to female genitalia, and Au (1969) considers their hindwing ventation
primitive; for it is in this subfamily that stridulitrum and plectrum are found.

Acknowledgments

I am grateful to Dr. John D. Lattin (Oregon State University), who sent
me a copy of Eunice Chizuru Au’s unpublished thesis. I also thank the
National Science Foundation, which supported part of this work.

Literature Cited

Au, E. C. 1969. The taxonomic value of the metathoracic wing in the Scutelleridae (Hemiptera:
Heteroptera). MA thesis, Ore. State Univ. Pp. 129.

Draslar, K. and M. Gogala. 1976. Struktura stridulacijskih organov pri zuzelkah iz druzine
Cydnidae (Heteroptera). Biol. Vestn. (Ljubljana) 24:175-200.

Dupuis, C. 1953. Appareil stridulatoire et stridulation des Cydnidae et Tessaratomidae (Het-
eroptera, Pentatomoidea). Cah. Nat. (Bull. Natur. Parisiens) (N.S.) 8:25-27.

Froeschner, R. C. 1960. Cydnidae of the Western Hemisphere. Proc. U.S. Nat. Mus. 111:337-
680.

Gogala, M. 1978. Ecosensory functions in insects (with remarks on Arachnida). Pp. 123-153
in Sensory Ecology, M. A. Ali, ed., Plenum Publ. Corp.

Kumar, R. 1965. Contributions to the morphology and relationships of Pentatomoidea (Hem-
iptera: Heteroptera) Part I. Scutelleridae. J. Entomol. Soc. Qd. 4:41-55.

Lawson, F. A. and J. Chu. 1971. A scanning electron microscope study of stridulating organs
in two Hemiptera. J. Kan. Entomol. Soc. 44:245-253.

Leston, D. 1952. Notes on the Ethiopean Pentatomoidea (Hemiptera): VIII, Scutellerinae
Leach of Angola, with remarks upon the male genitalia and classification of the subfam-
ily. Publ. Cult. Co. Diam. Angola 8:11-25.

. 1954. Strigils and stridulation in Pentatomoidea (Hem.): some new data and a review.

Entomol. Mon. Mag. 90:49-56.

. 1956. The Ethiopean Pentatomoidea (Hemiptera): XXII, on Dismegistus Amyot and

Serville (Cydnidae). Proc. Roy. Entomol. Soc. London (A) 31:87-94.

. 1957. The stridulatory mechanisms in terrestrial species of Hemiptera Heteroptera.

Proc. Zool. Soc. London 128:369-386.

. 1961. (Review of Froeschner, 1960). The Entomologist 94: 102-104.

McAtee, W. L. and J. R. Malloch. 1933. Revision of the subfamily Thyreocorinae of the
Pentatomidae (Hemiptera-Heteroptera). Ann. Carnegie Mus. 21:191-411.

Muir, F. 1907. Notes on the stridulating organ and stink-glands of Tessaratoma papillosa
Thunb. Trans. Entomol. Soc. London 1907:256-258.

Schaefer, C. W. and D. B. Wilcox. 1971. A new species of Thaumastellidae (Hemiptera:
Pentatomoidea) from southern Africa. J. Entomol. Soc. S. Afr. 34:207-214.

Schouteden, H. 1904. Heteroptera Fam. Pentatomidae Subfam. Scutellerinae. Genera Insec-
torum Fasc. 24, pp. 98.

Stys, P. 1964. Thaumastellidae — a new family of pentatomoid Hemiptera. Acta Soc. Entomol.
Cechoslov. 61:236-253.

Systematic and Evolutionary Biology Section, Biological Sciences Group,
University of Connecticut, Storrs, Connecticut 06268.

Received for publication June 12, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(4), 1980, pp. 236-240

THE HOUSE-FLY MYCOSIS CAUSED BY ENTOMOPHTHORA
MUSCAE : INFLUENCE OF RELATIVE HUMIDITY ON
INFECTIVITY AND CONIDIAL GERMINATION

John Paul Kramer

Abstract. — Studies on the effects of various relative humidities on the
infectivity of Entomophthora muscae conidia showed that Musca domes-
tica exposed to conidial showers falling through atmospheres with relative
humidities ranging from zero to 100% acquired fatal infections. Studies on
the germination of conidia held at various humidities showed that the fun-
gus produces long germ tubes only from secondary conidia at 100% R.H.
These results suggest that E. muscae can be transmitted within fly popu-
lations during periods of dry-cool weather, and that the relative humidity
within the boundary layer surrounding the fly’s body is at or near the sat-
uration point. Temperature maintained during the studies was 21 ± 3°C.

Introduction

The mycosis of the house fly caused by Entomophthora muscae has been
recognized for more than a century. One factor that undoubtedly influences
the occurrence of this disease in populations of Musca domestica and other
species is atmospheric humidity. Wilding and Lauckner (1974), in a review
of the literature pertaining to this subject, noted that epizootics occur mainly
in wet and cool periods but are known from dry and warm ones as well. In
their own studies these authors found little correlation between atmospheric
conditions and the incidence of the disease in the field. Berisford and Tsao
(1974) also found no striking relationship between the incidence of E. mus-
cae infections and weather conditions. Baird (1957) noticed accidental trans-
mission of E. muscae among caged flies held in the laboratory at about 23°C
and a relative humidity of 50%; for reasons unknown the fungus was not
transmissible after the cultures of flies were transferred to another room
with similar atmospheric conditions. I (Kramer, 1980) reported successful
transmission of E. muscae to house flies held in humid chambers at 14-
27°C and no transmission of this pathogen to flies held in a dry chamber at
21-27°C. I also noted that many conidia held on glass slides in a humid
chamber germinated whilst few conidia held under similar circumstances in
a dry chamber did so. The present study defines with greater precision the
effect of humidity in the ambient environment on the infectivity of E. mus-
cae conidia and on the germination of E. muscae conidia.

VOLUME LXXXVIII, NUMBER 4

237

Materials and Methods

I collected about 30 living cluster flies Pollenia rudis from the window
panes of a heated building where a natural epizootic of the mycosis was in
progress on December 8, 1979. Cluster flies from this collection dying of the
disease over the next six days served as a source of inoculum used to
establish in vivo cultures of E. muscae in house flies by a method described
previously (Kramer 1980). Conidia-bearing cadavers from these cultures
served as the source of inoculum for the studies described herein. Primary
conidia were uniformly bell-shaped with an apical point while the smaller
secondary conidia lacked the apical point; clearly the fungus was E. muscae
(see MacLeod et al. 1976).

Infectivity Tests. — Batches of 20 to 25 young healthy flies were placed in
cylindrical, clear plastic cages (height 35 mm, width 30 mm) having two
large mesh-covered windows and mesh-covered tops and bottoms. Three
fresh cadavers of M. domestica with E. muscae infections were affixed with
petrolatum to the inner surface of small plastic dishes. A dish bearing ca-
davers was placed over each cage to allow the conidia discharged to fall
through the mesh-covered top into the cage of flies. These cages with dishes
were placed in glass battery jars tightly covered with layers of polyethylene
wrap and waxed paper. The relative humidities maintained in these cham-
bers are given in Table 1. After 24 hours the test flies were fed honey and
the dishes with cadavers were removed. After an additional 24 hours the
flies were transferred to glass-covered carton cages, and each cage was
provisioned with a water bottle and a mixture of dried milk and sugar. These
cages were held at 21 ± 3°C and 45% R.H. with a 18:6 photoperiod. The
cages were checked for infected flies at daily intervals. The findings are
given in Table 1.

Germination of Conidia. — Three or four freshly dead or dying house flies
with E. muscae infections were affixed to the inner surface of small plastic
dishes as described above. These dishes were suspended over glass cover
slips in closed chambers maintained under the conditions given in Table 2.
Twenty-four hours later the slips, mounted in Colley’s solution (Colley,
1925), were examined microscopically. The patterns of germination ob-
served are given in Table 2.

Humidities. — The relative humidities within the tightly closed jars were
achieved with saturated solutions of the following compounds: Pyrocatechol
(94%), KC1 (85%), NaCl (75%), NaNO, (65%), glucose (55%), KN02 (48%),
Nal (38%), and NaOH (6%). Since a means to measure the actual humidities
was not available, the percentages were taken from a table given by Winston
and Bates (1960). Distilled H20 provided a relative humidity of 100%; an-
hydrous CaCl2 in a dry jar maintained a relative humidity at or near zero
percent.

238

NEW YORK ENTOMOLOGICAL SOCIETY

Table 1. Infectivity of Entomophthora muscae conidia falling upon healthy Musca do-
mestica held at various relative humidities at 21°C.

Relative

humidity

No. flies
at risk

Post-exposure days

with percentage dying of infection

Total

percentage

fatally

infected

1-4

5

6

7

8

100

20

0

5

50

45

0

100

94

20

0

5

70

25

0

100

85

21

0

0

86

14

0

100

75

23

0

0

61

30

9

100

65

24

0

0

38

29

12

79

55

22

0

10

36

36

0

82

48

20

0

0

60

0

0

60

38

20

0

10

50

20

0

80

6

21

0

14

33

10

14

71

0

15

0

0

7

13

0

20

Results and Discussion

Flies exposed to conidial showers falling through atmospheres with rel-
ative humidities ranging from zero to 100% acquired fatal E. muscae infec-
tions (Table 1). At relative humidities of 75% and above all flies at risk
became infected, while at 65% R.H. and below some flies apparently es-
caped infection. Why some flies did not acquire the infection is unclear.
Inherent resistance is probably not the explanation, since uninfected sur-
vivors from these tests later proved to be quite susceptible when exposed
to conidial showers at 75% R.H.

Primary conidia were produced and discharged from cadavers of flies held
at relative humidities ranging from zero to 100%. Conidial showers from
cadavers in atmospheres with low relative humidities (6 and 38%) were
markedly scanty compared to those from cadavers at 55% R.H. or higher.
That some conidia are produced at 0% R.H. suggests that the cadaver of a
well-nourished fly supplies moisture adequate for the development of a few
of these asexual spores. The data in Table 2 show that some primary conidia
produced secondary conidia at all humidities considered. Often these sec-
ondary conidia developed small lateral bulges. Other secondary conidia pro-
duced germ tubes about as long as a secondary conidium at humidities of
48% and above. Only at 100% R.H. were germ tubes more than twice as
long as a conidium formed from secondary conidia. These long tubes were
often branched and sometimes two or more tubes arose from one conidium.
The significance of these variations is unknown. Long tubes completely
separated from the secondary conidium of origin were observed only at
100% R.H. In no instances were forms clearly identifiable as tertiary conidia
observed in any of the slide preparations.

VOLUME LXXXVIII, NUMBER 4

239

Table 2. Germination patterns among conidia of Entomophthora muscae discharged onto
glass slips from house-fly cadavers during a 24-hour period at various relative humidities at
21°C.

Percentages of forms present*

Secondaries germinating

Relative

humidity

Primary

alone

Primary

with

secondary

Secondary

alone

Secondary
with bulge

Germ tube
short**

Germ tube
long***

Long germ
tubes alone

100

27

29

24

0

7

16

7

94

22

21

13

7

37

0

0

85

17

14

29

27

13

0

0

75

16

3

47

23

11

0

0

65

29

8

51

5

7

0

0

55

33

5

43

8

11

0

0

48

41

4

52

1

2

0

0

38

24

11

61

4

0

0

0

6

48

18

33

1

0

0

0

0

73

17

9

1

0

0

0

* Average of four replicates, 100 forms per replication.

** Germ Tube Short = Length no greater than length of conidium.
*** Germ Tube Long = Length greater than length of conidium.

The foregoing results indicate that: 1 , conidia falling through atmospheres
with relative humidities ranging from zero to 100% do give rise to infections
in flies (Table 1); and 2, long germ tubes, the invasive form of the fungus,
are produced only at 100% R.H. (Table 2). When viewed together these
findings suggest that the relative humidity within the boundary layer sur-
rounding the fly’s body is at or very near the saturation point. From an
epizootiological viewpoint these findings also suggest that E. muscae can
be transmitted within house-fly populations during periods of comparatively
dry and cool weather. Low humidities in themselves do not prevent the
spread of E. muscae infections in fly populations.

Acknowledgment

This study was supported in part by the UNDP/World Bank/WHO Special
Programme for Research and Training in Tropical Diseases.

Literature Cited

Baird, R. B. 1957. Notes on a laboratory infection of Diptera caused by the fungus Empusa
muscae Cohn. Can. Entomol. 89:432-435.

Berisford, Y. C. and C. H. Tsao. 1974. Field and laboratory observations of an entomogenous
infection of the adult seedcorn maggot, Hylemya platura (Diptera: Anthomyiidae). J.
Georgia Entomol. Soc. 9:104-110.

Colley, R. H. 1925. A biometric comparison of the uredinospores of Cronartium ribicola and
Cronartium occidentale. J. Agric. Res. 30:283-291.

240

NEW YORK ENTOMOLOGICAL SOCIETY

Kramer, J. P. 1980. Entomophthora muscae : Moisture as a factor affecting its transmission
and conidial germination. Acta Mycologica (in press).

MacLeod, D. M., E. Miiller-Kogler and N. Wilding. 1976. Entomophthora species with E.
muscae- like conidia. Mycologia 68:1-29.

Wilding, N. and F. B. Lauckner. 1974. Entomophthora infecting wheat bulb fly at Rothamsted,
Hertfordshire, 1967-71. Ann. Appl. Biol. 76:161-170.

Winston, P. W. and D. H. Bates. 1960. Saturated solutions for the control of humidity in
biological research. Ecology 41:232-237.

Department of Entomology Cornell University Ithaca, New York 14853.
Received for Publication July 2, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(4), 1980, pp. 241-251

STUDIES ON THE INCORPORATION OF [32P]-PHOSPHATE
INTO LOW MOLECULAR WEIGHT RNA OF DROSOPHILA
MELANOGASTER CELL LINES

M. Matouschek, A. Diibendorfer and E. Kubli

Abstract. — In higher organisms the specific activity of the RNA obtained
after feeding radioactive precursors to the animals is very low. Therefore,
we have systematically studied the incorporation of [32P]-phosphate into low
molecular weight RNA of the following four Drosophila melanogaster cell
lines: Kc, D1? D2, and line 3 of Schneider. After phenol-extraction the RNA
was loaded on a 10% polyacrylamide gel. The tRNA region and the region
between 4S and 5S ( = tRNA “precursor” region) were eluted and the ra-
dioactivity measured in a liquid scintillation counter.

Three labeling procedures were used: 1) Addition of [32P]-phosphate to
the cell cultures in normal medium containing [31P]-phosphate; 2) Culture
of cells for one hour in phosphate-free medium before addition of [32P]-
phosphate; 3) Three hours without phosphates prior to the addition of [32P]-
phosphate. The [32P]-phosphate pulse was two hours for all three types of
experiments. The optimum incorporation was found with the cells of line 3
incubated under conditions 2.

The specific activities obtained were 108'000 cpm/A260 for the RNA region
and 9'700 cpm/A26() for the “precursor” region. Modified nucleosides spe-
cific for tRNA were present in the region between 4S and 5S and also in the
RNAs moving slower than 5S RNA on a 10% polyacrylamide gel. This
suggests that tRNA precursors are localized in these regions. Schneider-3
cells were incubated for 2, 6, 12 and 24 hours with [32P]-phosphate in order
to determine the optimal incubation time for the labeling of stable tRNA.
Twelve hours was found to give the highest specific activity of 670'000 cpm/
^260*

Introduction

The discovery that the primary transcription products of tRNA genes are
precursor molecules larger than the tRNAs engaged in protein synthesis
was originally made in mammalian cells (Burdon et al. 1967). It was shown
that RNA of about 4.5S size could be converted into 4S molecules by crude
cell extracts. At the same time these molecules acquired the nucleoside
modifications characteristic for tRNAs. The detailed elucidation of the pro-
cessing and nucleoside modifications leading to a mature tRNA were worked
out in prokaryotes and most recently in lower eukaryotic cells (Altman 1978;
Hopper et al. 1978; Knapp 1978). The major reasons for this success were
the ease of selecting mutants, e.g. mutants blocking the processing steps

242

NEW YORK ENTOMOLOGICAL SOCIETY

(Schedl and Primakoff 1973), the use of transducing phages (Altman and
Smith 1971), and — for sequencing purposes — the high specific activities ob-
tained after [32P]-labeling.

Two major problems are encountered in studies of the primary transcrip-
tion products of tRNA genes from higher organisms. Firstly, only low spe-
cific activities of the RNA are obtained by feeding or injecting radioactive
precursors into animals. Secondly, the isolation and identification of specific
tRNAs out of a mixture of many transcription products of about equal size
is difficult. The first problem might be circumvented by the use of cell
cultures, where higher specific activities can be obtained, the second prob-
lem by using organs synthesizing specific tRNAs in great amount, e.g.,
Bombyx mori silk glands (Garber et al. 1978), or isolation techniques able
to pick up specific tRNA isoacceptors (Grosjean et al. 1973; Vogeli et al.
1975).

Drosophila melanogaster offers two advantages for studying the structure
and function of tRNA genes in a higher organism: the possibility of localizing
by “in situ’ ’-hybridization genes that code for tRNAs (for a review see
Kubli 1981) and sophisticated genetic techniques allowing the manipulation
of the genome. Recently plasmids containing Drosophila melanogaster
tRNA genes have been isolated and transcribed in Xenopus laevis oocyte
nuclei (Schmidt et al. 1978). Precursor tRNAs of about 5S RNA size could
be characterized. However, a careful comparison with products transcribed
in the homologous “in vivo” system is still needed. The full nucleotide
sequences of tRNA^, tRNALvs, and tRNA2phe from Drosophila melano-
gaster have been determined by post-labeling techniques (Silverman et al.
1979a; Silverman et al. 1979b; Altwegg and Kubli 1979). The high degree of
nucleoside modification found in tRNAs from eukaryotes sets some limits
to these procedures. Conventional sequencing techniques (Brownlee 1972)
using “in vivo” uniformly labeled [32P]-tRNA are therefore still necessary
to complement these recently developed elegant techniques. We have,
therefore, decided to conduct a preliminary study on the incorporation of
[32P] into the low molecular weight RNA of four Drosophila melanogaster
cell lines under various labeling conditions and to apply a specific affinity
chromatography (Grosjean et al. 1973) for the isolation of a tRNA precursor.

Materials and Methods

Cell lines. — The following Drosophila melanogaster cell lines were used:
Kc, D,, D2, and S (Schneider’s line 3) (Schneider and Blumenthal 1978).
The Kc cells were maintained in medium D22 (Echalier and Ohanessian
1970), the other cell lines in the medium of Shields and Sang (1977) at 25°C
in 30 cm3 Falcon flasks. The cells were subcultured after reaching a density
of 107 cells/ml. The cell lines D, , D2 and S were free of viruses whereas the
line Kc contained DXV-viruses (Plus 1978).

VOLUME LXXXVIII, NUMBER 4

243

Labeling procedures. — For the labeling experiments the cells were trans-
ferred into capped tubes on a roller drum or into spinner flasks, depending
on the number of cells needed. Under these conditions the cells stayed in
suspension and could be grown to a density of 15 x 106 cells/ml. Three
procedures were used for the pulse-label experiments. 1.) Addition of [32P]-
phosphate to the cell cultures in normal medium containing [31P]-phosphate.
2.) Culture of cells for 1 hour in phosphate-free medium before addition of
[32P]-phosphate. 3.) Three hours in phosphate-free medium prior to the ad-
dition of [32P]-phosphate. The [32P]-phosphate pulse was 2 hours for all three
experiments.

RNA extraction. — The cells were lysed in 50 mM Tris-HCl, pH 7.5, con-
taining 0.5% sodiumdodecylsulfate and 0.1% diethylpyrocarbonate at 0°C
for 5 min.. The DNA was digested by addition of DNAse (40 ptg/ml) at 0°C
for 5 min. and the RNA subsequently extracted according to Kirby (1956).
The high molecular weight RNA was removed by high-salt precipitation
(Kern 1975), and the low molecular weight RNA in the supernatant was
precipitated with ethanol. This RNA preparation was used for polyacryl-
amide gel electrophoresis and affinity chromatography. The aminoacyl-
tRNA synthetases were prepared according to Twardzik et al. (1971) and
the aminoacylation conditions were those of White and Tener (1973).

Polyacrylamide gel electrophoresis and affinity chromatography . — For
the polyacrylamide gel electrophoresis the protocol of Fradin et al. (1975)
was followed in detail. After autoradiography the RNA was eluted. The gel
pieces were forced through the orifice of a 2 ml syringe and eluted over
night at 37°C in 0.1 M Tris-HCl, pH 7.5, containing 0.5 M NaCl and 0.001
M EDTA. The gel particles were removed on a G-25 Sephadex column.
After addition of rRNA as a carrier (40 /^g/probe) the RNA was ethanol
precipitated. The procedure described by Grosjean et al. (1973) was used
for the affinity chromatography.

Results

Pulse-labeling experiments. — Precursor tRNAs (ptRNAs) in Bombyx
mori have been found on polyacrylamide gels to move between mature
tRNA and 5S RNA (Garber et al. 1978). The gel region containing the mature
tRNA was therefore studied separately from the region between mature
tRNA and 5S RNA. This region will be referred to as “ptRNA” since it
possibly contains tRNA precursors.

A comparison of the specific activities of the tRNA isolated from the four
cell lines by the three different labeling procedures is shown in Figure la.
The highest specific activity was obtained with S cells under labeling con-
ditions 2. The low value for the D2 cell line under these conditions may be
an artifact. The lowest incorporation was measured with the Kc cells
throughout all procedures.

244

NEW YORK ENTOMOLOGICAL SOCIETY

D

8

o

co

Q ‘

O 6

E

o 2

-A L

Kc Dt D2 S

3)

CELL LINES

12

10

o

CD

cT 8

o

co*

o 6

X

E

CL 4
o

2 L

1)

Kc D2 S

2)

12

12.

-

•

10

10.

•

•

8

8 .

6

6 .

•

4

4 -

.

•

2

2 .

•

.

•

. , ■

Kc D1 D2 S
CELL LINES

3)

Kc D, D2 S

Fig. 1 . a.) Specific activity of the RN A isolated from the tRNA region of a one-dimensional

polyacrylamide gel under 3 labeling conditions: 1.) 0.1 ml monosodium [32PJ-phosphate (2.5
mCi; pH 5-6; NEX 063, NEN, Dreieichenhein, BRD) and 0.1 ml double concentrated medium
was added to 4-9 ml cells in suspension (cell density 8-13 x 10'Vml). 2.) 3.5-7 ml cell suspen-
sion (15 x 10<!/ml) were centrifuged and washed with 3 ml medium without [31P]-phosphate.
The pellet was resuspended in the same medium to obtain a cell density of 15 x 10“ cells/ml.
The cells were kept in this medium for 1 hour on a roller drum (30 rev. /min.). Then 0.1 ml

Fig. 2. One-dimensional polyacrylamide gel of RNA extracted from different cell lines
labeled according to the protocol described in Figure la.

The specific activities of the “ptRNA” regions, though following a similar
trend (Fig. lb), were about an order of magnitude lower. Again the D2 cells
labeled under conditions 2.) showed reduced incorporation in comparison
with the D, and S cells. Under these conditions the specific activity was
slightly higher in D1 cells than in S cells.

Polyacrylamide gel electrophoresis. — The results of the one-dimensional
separation of the low molecular weight [32P]-RNAs from the different cell
lines are shown in Figure 2. No major differences in the pattern can be

monosodium [32P]-phosphate and 0.1 ml double concentrated medium was added. 3.) Same as
2.) but incubation of the cells in [31P]-phosphate-free medium for 3 hours. After 2 hours of
[32P]-phosphate labeling the RNA was extracted and fractionated on a one-dimensional poly-
acrylamide gel. b.) Specific activity of the RNA isolated from the “ptRNA” region. Labeling
conditions as described under a.).

246 NEW YORK ENTOMOLOGICAL SOCIETY

Hours

Fig. 3. Long term labeling of tRNA and “ptRNA.” S cells were labeled under the condi-
tions 2.) (see legend Fig. la) for 2, 6, 12 and 24 hours. The RNA was extracted and fractionated
on a one-dimensional polyacrylamide gel. • • tRNA region, O O “ptRNA” region.

detected in the “ptRNA” and tRNA regions. The RNAs moving slower
than 5S RNA differ markedly in the different cell lines. However, the la-
beling procedure seems not to influence the labeling pattern.

Long term labeling experiments. — Considering the high incorporation val-
ues into the tRNA region obtained with the S cells we chose these for the
long-term incorporation studies. Figure 3 shows that after 12 hours of la-
beling the specific activity levels off in the tRNA region. The maximal ac-
tivity in the “ptRNA” region is already reached after 6 hours, but remains
about an order of magnitude lower than the specific activity measured in the
tRNA region.

Affinity chromatography . — The principle of the anticodon-anticodon af-
finity chromatography is based on the observation that some tRNAs with
complementary anticodons can form a stable complex at low temperature
(Eisinger 1971). A pure tRNA (e.g., tRNAphe of yeast or tRNAGlu of E. coli )
is covalently bound to a supporting material and poured into a column. In
a mixture of tRNAs loaded on the column only those RNAs with the com-
plementary anticodon form a stable complex at 4°C, the rest flows through.
Increasing the temperature to 30°C and adding EDTA to the elution buffer
destabilizes the complex: the formerly bound tRNA elutes as a single peak
(Grosjean et al. 1973). Control experiments demonstrate that only the tRNA
with the complementary anticodon is retained on the column, e.g., tRNAphe
on a tRNAGlu column (Fig. 4a). An example of an affinity chromatography

VOLUME LXXXVIII, NUMBER 4

247

a

b

Fig. 4. a). [3H]-Phe-tRNA (A A) and [14C]-Glu-tRNA (▲ ▲) were cochromato-

graphed on a tRNAGlu affinity column. The [3H]-Phe-tRNA with the complementary anticodon
is retained on the column and eluted at 30°C, the non-complementary [14C]-Glu-tRNA is eluted
in the first peak. b). Affinity chromatography of [32P]-phosphate labeled tRNA on a tRNAGlu
column. The second peak contains [32P]-tRNAPhe. Labeling conditions: 95 x 106 cells were
labeled with 25 mCi [32P] according to conditions 2.) (see legend Fig. la) for 1 hour.

248

NEW YORK ENTOMOLOGICAL SOCIETY

t RNAPhe

5S RNA

t RNA

Fig. 5. One-dimensional polyacrylamide gel electrophoresis of [32P]-phosphate labeled
tRNAphe isolated by affinity chromatography (second peak Fig. 4b).

elution profile with pulse-labeled [32P]-tRNA isolated from the S cell line is
shown in Figure 4b. The second peak contains the tRNAphe with a comple-
mentary anticodon to the tRNAGlu fixed on the column (Altwegg and Kubli,
1979). After precipitation this peak was loaded on a 10% polyacrylamide
gel. The autoradiograph revealed about 8 bands in the tRNA region (Fig.
5). The bands were cut out and eluted. However, the radioactivity recovered
was too low to allow a characterization of the different bands by finger-
printing. The identification was therefore not possible.

VOLUME LXXXVIII, NUMBER 4

249

Discussion

The pulse labeling experiments show that the highest specific activity of
the tRNA region is obtained with S cells under labeling conditions 2 (Fig.
la). On the other hand, no similarly high specific activity is found in the
“ptRNA” region of S cells. These seemingly contradictory results are
understood in the light of the fact that some UV-absorbing material always
co-elutes when the RNAs are eluted from the gels for measuring the specific
activities. Considering the low concentration of RNAs in the ptRNA-region
this UV-absorbing material creates a relatively high background that inter-
feres with the calculation of the specific activity.

A comparison of the RNA pattern of the different cell lines on a one-
dimensional polyacrylamide gel reveals that the tRNA regions do not differ
considerably. However, major differences can be detected in the regions
above 5S RNA (Fig. 2). This might reflect differential RNA extraction, but
could also be due to the fact that some cell lines can be virus infected (Plus
1978) or aneuploid (Schneider and Blumenthal 1978). It has been reported
that viruslike particles (VLP) of cultured Drosophila melanogaster cell lines
contain a low molecular RNA which is believed to be the tRNA since this
RNA can be aminoacylated (Shiba et al. 1980). However, the differences in
the RNA patterns described here are not due to virus infections since the
observed extra fractions of RNA all show up in the virus-free cell lines.
On the other hand many cell lines do not contain a normal diploid set of
chromosomes (Schneider and Blumenthal 1978), and since the tRNA genes
are distributed over the entire Drosophila genome some tRNA genes might
be missing and others increased in number. These problems have to be
considered when tRNA preparations isolated from cell cultures are used for
biochemical studies, such as the comparison of isoacceptor patterns, or the
screening of plasmids containing tRNA genes. A further aspect deserving
attention is the degree of modification of the tRNA isolated from cell cul-
tures. For example, it has been shown that the tRNAHis and the tRNAAsN
isolated from Drosophila melanogaster cell cultures (S cells) do not contain
the hypermodified base Q in the first position of their anticodon (Schmidt
and Kubli 1980).

Affinity chromatography provides an elegant means of isolating tRNAs
and their precursors (Grosjean et al. 1973; Vogeli et al. 1975). This method
can only be used for the isolation of precursors if they do not contain in-
sertion sequences near their anticodon. We obtained several bands upon
separation of the second peak of the affinity column on a one-dimensional
polyacrylamide gel (Fig. 5). This result can be interpreted in different ways,
e.g., as unspecific binding to the tRNA hooked on the column, or as nu-
clease degradation of tRNA leaving the anticodon intact. Unfortunately the

250

NEW YORK ENTOMOLOGICAL SOCIETY

activities obtained do not allow us to decide whether tRNA precursors were
actually isolated. Pulse-label experiments with higher [32P]-concentrations
might give the necessary RNA activities to allow an adequate investigation
of these problems.

Acknowledgments

We wish to thank our colleague Martin Altwegg for providing technical
help and Johanna Nageli for assistance with the cell cultures. The work was
supported by the Swiss National Science Foundation, grants No. 3.024.76
and 3.741-0.76.

Literature Cited

Altman, S. and J. D. Smith. 1971. Tyrosine tRNA precursor molecule polynucleotide se-
quence. Nature New Biology 233:35-39.

. 1978. Biosynthesis of tRNA. In: S. Altman, ed., Transfer RNA. MIT Press, Cam-
bridge, pp. 48-77.

Altwegg, M. and E. Kubli. 1979. The nucleotide sequence of phenylalanine tRNA2 of Dro-
sophila melanogaster. four isoacceptors with one basic sequence. Nucl. Acids Res.
7:93-105.

Burdon, R. H., B. T. Martin and B. M. Lai. 1967. Synthesis of low molecular weight ribo-
nucleic acid in tumor cells. J. Mol. Biol. 28:357-371.

Brownlee, G. G. 1972. Determination of Sequences in RNA. North Holland/American Else-
vier, Amsterdam/New York.

Eisinger, J. 1971. Visible gel electrophoresis and the determination of association constants.
Biochem. Biophys. Res. Comm. 43:854-861.

Echalier, G. and A. Ohanessian. 1970. In vitro culture of Drosophila melanogaster embryonic
cells. In Vitro 6:162-172.

Fradin, A., H. Gruhl and H. Feldman. 1975. Mapping of yeast tRNAs by two dimensional
electrophoresis on polyacrylamide gels. FEBS Letters 50:185-189.

Garber, R. L., M. A. Q. Siddiqui and S. Altman. 1978. Identification of precursor molecules
to individual tRNA species from Bombyx mori. Proc. Natl. Acad. Sci. USA 75:635-639.

Grosjean, H., C. Takada, and J. Petre. 1973. Complex formation between tRNAs with com-
plementary anticodons: use of matrix bound tRNA. Bioch. Biophys. Res. Comm.
53:882-893.

Hopper, A. K., F. Banks and V. Evangelidis. 1978. A yeast mutant which accumulates
precursor tRNAs. Cell 14:211-220.

Kern, H. 1975. Fractionation of nucleic acids with special regard to rapidly labeled RNA.
Anal. Biochem. 67:147-156.

Kirby, J. D. 1956. A new method for the isolation of ribonucleic acids for mammalian tissues.
Biochem. J. 64:405-408.

Knapp, G., J. S. Beckman, P. F. Johnson, S. A. Fuhrman and J. Abelson. 1978. Transcription
and processing of intervening sequences in yeast tRNA genes. Cell 14:221-236.

Kubli, E. 1981. The genetics of transfer RNA in Drosophila . Adv. Genetics, in press.

Plus, N. 1978. Endogenous viruses of Drosophila melanogaster cell lines: their frequency,
identification and origin. In Vitro 14:1015-1021.

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Schedl, P. and P. Primakoff. 1973. Mutants of Escherichia coli thermosensitive for the syn-
thesis of transfer RNA. Proc. Natl. Acad. Sci. USA 70:2091-2095.

Schmidt, T., A. H. Egg and E. Kubli. 1978. The localization of tRNA2Asp genes from Dro-
sophila melanogaster by “in situ” hybridization. Molec. Gen. Genet. 164:249-254.

and E. Kubli. 1980. The localization of tRNA5AsN, tRNAHis, and tRNAAla genes from

Drosophila melanogaster by “in situ" hybridization to polytene salivary gland chro-
mosomes. Chromosoma 80:277-287.

Schneider, I. and A. B. Blumenthal. 1978. In: M. Ashburner and T. R. F. Wright, eds., The
Genetics and Biology of Drosophila Vol. Ila, Academic Press, London, New York, San
Francisco, pp. 266-315.

Shiba, T., K. Saigo, T. Marunouchi and T. Miyake. 1980. Isolation and characterization of
low molecular weight RNAs in Drosophila virus-like particles. In: E. Kurstak, K. Mar-
amorosch and A. Dubendorfer, eds., Invertebrate Systems in Vitro. Elsevier/North
Holland Biomedical Press, Amsterdam.

Shields, G. and J. H. Sang. 1977. Improved medium for Drosophila embryonic cells. Dros.
Info. Service 52: 161.

Silverman, S., J. Heckman, G. J. Cowling, A. D. Delaney, R. J. Dunn, I. C. Gillam, G. M.
Tener, D. Soil, and U. L. Raj Bhandary. 1979a. The nucleotide sequence of the initiator
tRNA from Drosophila. Nucl. Acids Res. 6:421-433.

, I. C. Gillam, G. M. Tener and D. Soil. 1979b. The nucleotide sequence of lysine

tRNA2 from Drosophila. Nucl. Acids Res. 6:435-442.

Twardzik, D. R., E. H. Grell, and K. B. Jacobson. 1971. Mechanisms of suppression in
Drosophila: a change in tyrosine transfer RNA. J. Mol. Biol. 57:231-245.

Vogeli, G., H. Grosjean and D. Soli. 1975. A method for the isolation of specific tRNA
precursors. Proc. Natl. Acad. Sci. USA 72:4790-4794.

White, B. N. and G. M. Tener. 1973. Chromatography of Drosophila tRNA on BD-cellulose.
Can. J. Biochem. 51:896-902.

Institute of Zoology, University of Zurich, Switzerland.
Received for Publication August 17, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(4), 1980, pp. 252-256

CONTINUOUS PRODUCTION OF PREDACIOUS
MITES IN THE GREENHOUSE

R. M. Hendrickson, Jr.

Abstract. — The predacious mite species Phytoseiulus persimilis Athias-
Henriot and Neoseiulus californicus (McGregor) (Acari: Phytoseiidae) were
reared on potted red clover, Trifolium pratense L., in the greenhouse at
ambient temperatures (min. 12°C). Twospotted spider mites, Tetranychus
urticae Koch (Acari: Tetranychidae), were placed on the plants as prey.
Red clover leaflets, on which were predators and prey, were removed and
placed on potted alfalfa, Medicago sativa L., in the same greenhouse. One
pot of red clover produced 5,000-8,000 predators per year, sufficient for
satisfactory control of T. urticae on 50 pots of alfalfa. The technique re-
quires minimal time, greenhouse space, and expense.

The purpose of this paper is to describe an unsophisticated technique for
rearing predacious mites in a greenhouse used for arthropod research, with
a minimum of time, space, and expense. The predators were used in routine
biological control of twospotted spider mite, Tetranychus urticae Koch,1’2
on potted alfalfa in the same greenhouse.

Many excellent methods for rearing predacious mites have been pub-
lished: for example, Ristich 1956; McMurtry and Scriven 1965; Kamburov
1966; and Gilstrap 1977. However, these techniques are relatively sophis-
ticated and are most suitable for biological control workers specializing in
the study of predacious mites. Burnett (1970a, b, 1971, 1977) has examined
in detail the population relationships between the predacious mite Ambly-
seius fallacis (Garman) and T. urticae on alfalfa in greenhouse and field.

The primary reason for rearing alfalfa in the greenhouse was as a host
plant for the introduced pest, the alfalfa blotch leafminer, Agromyza fron-
tella (Rondani).3 (We reared 4 European parasite species on this host (Hen-
drickson and Barth 1979)). It was necessary to attempt biological control of
T. urticae to avoid problems in insect rearing when alfalfa plants were
treated with acaricides. Some common acaricides, at recommended dos-
ages, were ineffective against T. urticae ; others interfered with the rearing

1 Acari: Tetranychidae.

2 Determined by R. L. Smiley, R. J. Gagne, or P. M. Marsh, Systematic Entomology
Laboratory, USDA-SEA-AR, % U.S. National Museum, Washington, DC 20560.

3 Diptera: Agromyzidae.

VOLUME LXXXVIII, NUMBER 4

253

of A. frontella. For example, the effective acaricide oxythioquinox,4’5 ap-
parently rendered alfalfa partially repellant to A. frontella for 3-4 weeks.
As a result, sprayed plants produced ca. 10% of the number of A. frontella
that untreated controls produced.

We reared the predacious mites Phytoseiulus persimilis Athias-Henriot6
and Neoseiulus californicus (McGregor)6 together on the same plants as an
experiment in competitive displacement. Although P. persimilis has a prov-
en reputation for effectiveness in greenhouses, it may not be the best pred-
ator under all greenhouse conditions. Since little has been published on the
biological characteristics of N. californicus, we had no way to estimate its
relative competitive ability vis-a-vis P. persimilis in the greenhouse. Oatman
et al. (1977) showed that P. persimilis was more effective than N. califor-
nicus ( =Amblyseius californicus ) in field releases against T. urticae on
strawberries. If the 2 species have the same ecological niche in the green-
house, competitive displacement should take place. On the other hand, both
species of predators may coexist based on exploitation of slightly differing
ecological niches, with a net effect in the control of T. urticae superior to
that of either predator species alone. Although preliminary results indicate
that P. persimilis has rapidly increased in numbers relative to N. califor-
nicus, the experiment is incomplete and is not discussed further in this
paper.

Prey culture. — The T. urticae culture was maintained in the laboratory
at 22-28°C. We could not rear prey mites in the same greenhouse with
predators because of the danger of contamination. Prey mites were placed
on 3-5 day old snap bean seedlings, of which we planted 200 per week. The
bean plants were grown to the 5-leaf stage and cut when mite populations
had become abundant. One-half of the mite-infested cut bean plants were
used to infest snap bean seedlings, and mites on the other half were used
as prey for predators.

Predator culture. — Predators were maintained in the greenhouse on pot-
ted red clover plants, Trifolium pratense L. Red clover is a vigorous grower
in the greenhouse even at low winter temperatures, withstands severe pop-
ulations of T. urticae and frequent removal of foliage, produces leaflets of
convenient size for production of predators, and is not a productive host
for greenhouse whitefly, Trialeurodes vaporariorum (Westwood)7.

4 Cyclic 5’A-(6-methyl-2,3-quinoxalinediyl)dithiocarbonate.

5 This paper reports the results of research only. Mention of a pesticide in this paper does
not constitute a recommendation for use by the U.S. Department of Agriculture nor does it
imply registration under FIFRA as amended.

Acari: Phytoseiidae. Determined by H. A. Denmark, Division of Plant Industry, Florida
Department of Agriculture and Consumer Services, Gainesville, FL 32602.

7 Homoptera: Aleyrodidae.

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Prey-infested snap bean plants were cut at the 5-leaf stage and placed on
predator-infested red clover plants. Foliage from 5-6 snap bean plants were
placed on a single red clover plant each week. Occasionally a red clover
plant was excessively infested with T. urticae, necessitating a recuperative
period of 1-2 weeks during which no mites were placed on it.

Prey- and predator-infested red clover leaflets were trimmed from plants
and placed on target alfalfa plants for T. urticae control. The older red
clover leaflets, usually the most severely damaged by T. urticae feeding,
sustained the highest populations of predators and the lowest populations
of prey. Therefore, these leaves were preferentially removed.

To determine the approximate rate of predator production, we selected
40 leaflets at random and counted mite populations on them under the micro-
scope. The numbers of predators ( P . persimilis and N. calif ornicus com-
bined) per leaflet were: adults, 2.6 x, 3.0 s; immatures, 4.0 x, 4.3 s; and
eggs, 3.1 x, 3.8 s. The number of T. urticae per leaflet were: adults, 7.3 x,
7.1 s; immatures, 44.4 x, 59.4 s; and eggs, 73.2 x, 99.7 s. Based on a count
of the number of leaflets regularly removed from each pot, we calculate
annual predator production (all stages) at 5,000-8,000 per pot of red clover.

How predators were used in the greenhouse. — We placed a predator and
prey-infested red clover leaf, comprised of 3 leaflets, on each of 80-160 pots
of alfalfa, which were used in insect rearing programs and which were cut
back weekly to ca. 5 cm height. Thus, the actual numbers of predators and
prey placed on each pot of alfalfa were 3 times the averages listed above.
Cut-back plants may have carried predators from earlier inoculations, which
would enhance the effect of newly added predators.

Predators were placed on potted alfalfa only once in each 5-10 week
usage cycle unless problems developed with pesticides applied for control
of greenhouse whitefly. (Attempts to use the parasite Encarsia formosa
Gahan8 9 for T. vaporariorum control were unsuccessful because even par-
asitized immature whiteflies produced honeydew, which apparently ren-
dered the plants unacceptable for oviposition by A. front ella.) We applied
resmethrin59 at 2-3 week intervals in the warm seasons of the year for
greenhouse whitefly control. Applications of resmethrin appeared to kill
nearly all adult and immature predacious mites, but left predator eggs alive.
Twospotted spider mites were unaffected by the pesticide, so that spotty
outbreaks resulted. To prevent pesticide-induced upsets of T. urticae , we
placed a single predator- and prey-infested red clover leaflet on each pot of

8 Hymenoptera: Aphelinidae. Kindly provided by M. J. Tauber, Dept. Entomology, Cornell
Univ., Ithaca, NY.

9 [5-pheny lmethyl)-3-furanyl]methyl cis-trans-( ±)-2,2-dimethyl-3-(2-methyl- 1-
propenyl)cyclopropanecarboxy!ate.

VOLUME LXXXVIII, NUMBER 4

255

alfalfa after the period of residual activity of resmethrin (3 days in warm
seasons).

One pot of red clover provided enough predators for 50 pots of alfalfa. In
general, the predacious mites gave us good control of T. urticae, better than
previously attained with acaricides. The bench area required for 16 red
clover plants in the greenhouse was 1.5 m2.

On about one pot of alfalfa in 100, we observed T. urticae feeding damage
on the first 15 cm of alfalfa regrowth, but at about this height, the predators
built up sufficiently high populations so that the remaining alfalfa growth
had no or very slight mite damage. Potted alfalfa used in rearing the alfalfa
blotch leafminer was 35-50 cm in height, and mite damage on the bottom
15 cm was unimportant.

We estimated that 1 Vi h per week were required in this technique for
successful biological control of T. urticae on 800 alfalfa plants. The time
estimate includes planting beans, watering, movement of prey on bean
leaves to red clover plants, and movement of prey and predators on red
clover leaflets to alfalfa plants. The time spent was ca. 3 times that formerly
required for the application of miticides, but was not excessive.

Discussion

The technique is useful for biological control of T. urticae in small green-
houses (ours is 100 m2). Its simplicity lends itself to routine application; it
requires relatively little time, and only enough greenhouse bench space for
one pot with predators per 50 pots with target prey; and it avoids compli-
cating factors in research induced by acaricide applications.

The technique may have applicability in commercial greenhouses, since
it is analogous to Gould et al. (1969) description of a ‘banker’ plant as a
source for predacious mites in greenhouses used for cucumber production.
In their concept, T. urticae on one cucumber plant in 100 was allowed to
reach heavy population levels, then P. persimilis was added. As many as
5,000 predators were produced on a single cucumber plant for the remainder
of the greenhouse. In the technique described in the present paper, prey
and predators were introduced simultaneously on red clover ‘banker’ plants,
with frequent addition of prey mites. The number of predacious mites pro-
duced (all stages), counted at the time of removal, was 5,000-8,000 per pot
per year.

Other predators of T. urticae encountered in the greenhouse were Anystis
agilis (Banks)2,10 and Feltiella Carolina (Felt),2,11 but both species were un-

10 Acari: Anystidae.

11 Diptera: Cecidomyiidae.

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NEW YORK ENTOMOLOGICAL SOCIETY

common and contributed little to control of T. urticae. Larvae of F. Carolina
fed on eggs and immotile forms of T. urticae. (A single individual of the
parasite Aphanogmus sp.2’12 emerged from a puparium of F. Carolina.)

Acknowledgment

This research was conducted with the able technical assistance of Susan
E. Barth.

Literature Cited

Burnett, T. 1970a. Effect of temperature on a greenhouse acarine predator-prey population.
Can. J. Zool. 48:555-562.

. 1970b. Effect of simulated natural temperatures on an acarine predator-prey popu-
lation. Physiol. Zool. 43:155-165.

. 1971. Prey consumption in acarine predator-prey populations reared in the green-
house. Can. J. Zool. 49:903-913.

. 1977. Dispersal of an acarine predator among greenhouse populations of the two-

spotted mite. Proc. Symp. Pest Management in Protected Culture Crops, XV Intern.
Congr. Entomol., Washington, DC (1976). USDA ARS-NE-85, pp. 49-64.

Gilstrap, F. E. 1977. Table-top production of tetranychid mites (Acarina) and their phytoseiid
natural enemies. J. Kans. Entomol. Soc. 50:229-233.

Gould, H. J. et al. 1969. Large scale commercial control of T. urticae on cucumbers by the
predator Phytoseiulus riegeli. Proc. 2nd Intern. Congr. Acarol. (1967), pp. 383-388.

Hendrickson, R. M. Jr. and S. E. Barth. 1979. Introduced parasites of Agromyza frontella
(Rondani) in the USA. J. N.Y. Entomol. Soc. 87:167-174.

Kamburov, S. S. 1966. Methods of rearing and transporting predacious mites. J. Econ. Ento-
mol. 59:875-877.

McMurtry, J. A. and G. T. Scriven. 1965. Insectary production of phytoseiid mites. J. Econ.
Entomol. 58:282-284.

Oatman, E. R., et al. 1977. Effect of releases of Amblyseius californicus, Phytoseiulus per-
similis, and Typhlodromus occidentalis on the twospotted spider mite on strawberry in
southern California. J. Econ. Entomol. 70:45-47.

Ristich, S. S. 1956. Mass rearing and testing techniques for Typhlodromus fallacis (Gar.). J.
Econ. Entomol. 49:476-479.

Beneficial Insects Research Laboratory, Agricultural Research, Science
and Education Administration, USDA, 501 S. Chapel St., Newark, Dela-
ware 19713.

Received for publication September 11, 1980.

12 Hymenoptera: Ceraphronidae.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXVIII(4), 1980, pp. 257-272

CONSPECTUS OF PENTATOMINI GENERA OF THE WESTERN
HEMISPHERE— PART 2 (HEMIPTERA: PENTATOMIDAE)

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

Abstract. — A key is provided to separate the genera of Pentatomini of the
Western Hemisphere that have a median tubercle or spine at the base of the
abdominal venter which is unapposed apically by the posterior margin of
the metasternum. New genera in this group are Aleixus McDonald, Grazia
Rolston, Kermana Rolston and Roferta Rolston. One new species is de-
scribed: Aleixus virgatus McDonald. New combinations are Grazia tincta
(Distant, 1890), Kermana bucera (Stal, 1860), K. imbuta (Walker, 1867), K.
fucosa (Berg, 1892) and Roferta marginalis (Herrich-Schaffer, 1836). A di-
agnosis is given for Zorcadium Bergroth, and Z. truncation (Fallou, 1888)
is redescribed.

In a preceding paper the tribe Pentatomini in the Western Hemisphere
was divided into 3 sections. A key was provided for the genera of section
3, viz. those genera with a median tubercle at the base of the abdominal
venter and with the metasternum produced ventrad in apposition to the apex
of that tubercle (Rolston et al. 1980).

In this paper those genera are keyed that have a median tubercle or spine
at the base of the abdominal venter but do not have the metasternum pro-
duced ventrad in apposition to the apex of that tubercle or spine. In fact,
the abdominal spine often projects under the metasternum and may reach
as far as the base of the head.

New genera added to this group are Aleixus McDonald, Grazia Rolston,
Kermana Rolston and Roferta Rolston. Aleixus in monotypic, based on A.
virgatus McDonald, n. sp. Grazia is also monotypic, based on Piezodorus
tinctus Distant, 1890. Kermana contains 3 species, all previously named
and in new combinations of K. bucera (Stal, 1860), K. fucosa (Berg, 1892)
and K. imbuta (Walker, 1867), of which the last is type species. Roferta is
monotypic, based on Pentatoma marginale Herrich-Schaffer, 1836.

The rare Zorcadium truncatum (Fallou, 1888) is redescribed and a diag-
nosis given for the monotypic genus. Since Fallou’s type is missing a vouch-
er specimen is designated.

Key to Genera of Pentatomini, Section 2

1. Stout pair of preapical spines present on inferior surface of poste-
rior femora Modicia Stal

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NEW YORK ENTOMOLOGICAL SOCIETY

Figs. 1-8. 1. Ramosiana insignis. Head, lateral view. 2. Brepholoxa heidmanni. Left buc-

cula, lateral view. 3. Nezara viridula. Left buccula, lateral view. 4. Brepholoxa heidmanni.
Left metathoracic stink gland orifice. 5. Odmalea basalis. Right fore femur, superior surface.
6. Nezara viridula. Left metathoracic stink gland orifice. 7. Acrosternum hilare. Left meta-
thoracic stink gland orifice. 8. Pellaea sticta. Pygophore, caudal view, legend: antennal seg-
ment 1 (a.x), buccula (b.), paramere (cl.), osteolar sulcus (o.), spine (s.).

- Femora not so armed 2

2. Median spine at base of abdominal venter projecting cephalad to

procoxae Disderia Bergroth

- Median spine at base of abdominal venter not projecting as far

cephalad as procoxae, sometimes reduced to tubercle 3

3. Distal end of first antennal segment clearly exceeding apex of head

(Fig. 1) 4

- Distal end of first antennal segment not surpassing apex of head 5

4. Distal end of first rostral segment clearly surpassing bucculae; scu-

tellar width at base about % length Ramosiana Kormilev

VOLUME LXXXVIII, NUMBER 4

259

- First rostral segment lying entirely between bucculae; scutellar

width and length subequal Vulsirea Spinola

5. Bucculae lobed posteriorly from lateral view (Fig. 2) 6

- Bucculae evanescent posteriorly (Fig. 3) 12

6. Ostiolar canal extending less than halfway from inner margin of

ostiole to lateral margin of metapleuron (Fig. 4); antennal segment
2 usually longer than or as long as each succeeding seg-
ment Brepholoxa Van Duzee

- Ostiolar canal extending more than halfway from inner margin of

ostiole to lateral margin of metapleuron; antennal segment 2 shorter
than each succeeding segment (except Aleixus ) 7

7. Superior surface of femora prolonged distally as small spine (Fig.

5) 8

- Femora not so armed 1 1

8. Juga contiguous before tylus 10

- Juga usually separated apically, if contiguous then coria decidedly

bicolored, stramineous and castaneous 9

9. Humeri bearing large dorsal tubercle (Fig. 10); second antennal
segment longer than each succeeding segment

Aleixus McDonald, n. gen.

- Humeri not tuberculate; second antennal segment shorter than each

succeeding segment Odmalea Bergroth

Humeri cornute (Fig. 32); costal angle of coria extending caudad
well beyond apex of scutellum Zorcadium Bergroth

Humeri angulate or spinose; costal angle of coria extending caudad
little if any farther than apex of scutellum Thoreyella Spinola

Median spine at base of abdominal venter projecting cephalad to

or beyond anterior limit of mesocoxae; scutellum at least 1 tenth
longer than basal width Rio Stal

- Abdominal spine shorter; scutellar width and length sub-
equal Dendrocoris Bergroth

12. Ostiolar sulcus short, length about twice diameter of orifice (Fig.

6) 18

- Ostiolar sulcus reaching about halfway or more from inner margin

of ostiole to lateral margin of metapleuron (Fig. 7) 13

13. Mesosternal carina compressed anteriorly, forming thin blade be-
tween procoxae Piezodoms Fieber

- Mesosternal carina nearly uniform in size and shape, sometimes

obsolete on xyphus between mesocoxae 14

14. Superior surface of femora prolonged distally into small angulate

tooth; parameres trilobed, posterior lobe curving mesoventrad and
exposed in mesial depression in posterior margin of pygophore
(Fig. 8). Pellaea Stal

10.

11

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NEW YORK ENTOMOLOGICAL SOCIETY

- Femora unarmed; parameres simple or bilobed (Figs. 16, 22, 29),

almost entirely concealed in genital cup 15

15. Lateral walls of genital cup bearing large tubercles near rim of cup

(Figs. 21, 25, 28); scutellum strongly convex at least basally 17

- Tubercles on lateral walls of genital cup small or absent; scutellum

weakly convex 16

16. Tibiae asculcate or weakly sulcate distally Grazia Rolston, new genus

- Tibiae clearly sulcate Acrosternum Fieber

17. Parameres with tubercle on cephalic margin between base and apex
(Fig. 29); spermathecal bulb with 2 unequal diverticula (Fig. 31);
median process at base of abdominal venter tuberculate Roferta
Rolston, new genus

- Parameres with ventrolateral process (Fig. 22); spermathecal bulb

bent into U (Fig. 24); abdominal spine projecting cephalad beyond
metacoxae Kermana Rolston, new genus

18. Abdominal spine surpassing anterior limit of metacoxae; each spi-
racle on yellowish callus Acrosternum abnorme (Berg)

- Abdominal tubercle reaching only to posterior limit of metacoxae;

spiracles unaccompanied by callus Nezara viridula (L.) )

Aleixus McDonald, new genus

Type species. — Aleixus virgatus McDonald, n. sp.

Diagnosis. — Abdominal sternite 3 (second visible) with small median i
spine reaching metacoxae or nearly so (Fig. 9). Dorsal surface of each hu-
merus bearing large tubercle (Fig. 10). Jugae narrowly rounded and sepa-
rated apically, surpassing tylus (Fig. 11). First antennal segment not reach-
ing apex of head; second segment longer than any other segment. First
rostral segment lying entirely between bucculae, these projecting as lobes
posteriorly onto prosternum (Fig. 12). Thoracic sterna nearly flat, not pro-
duced. Ostiolar sulcus extending about 6 tenths of distance from mesial
margin of ostiole to lateral margin of metapleuron (Fig. 13). Superior surface
of femora produced distally as small tooth, femora otherwise unarmed; tib-
iae sulcate; tarsi 3-segmented.

First gonocoxae lying mainly under sternite 7, visible only as small tri-
angular sclerites.

Aleixus virgatus McDonald, n. sp.

Dorsal surface fulvid overlaid with dark brown punctation; tubercles on
each humerus black. Dark brown stripe on each side of meson beginning at
apex of pronotum, skirting cicatrice mesially, continuing onto scutellum,
the 2 stripes converging about 3 quarters way down scutellum (Fig. 14);
narrower brown stripes on each hemelytron, one along frenum, another

VOLUME LXXXVIII, NUMBER 4

261

9

1-5mm

o.

Figs. 9-14. Aleixus virgatus. 9. Ventral view. 10. Left humeral angle, dorsal view. 11.
Head, dorsal view. 12. Buccula, lateral view. 13. Left metathoracic stink gland orifice. 14.
Dorsal view, legend: osteolar sulcus (o.).

along radial vein. Membrane smoky brown. Venter amber overlaid with
dark brown punctation. Length from apex of head to apex of abdomen 5.8
mm; width across humeri 4.5 mm.

Margins of jugae except mesial margin basally outlined in black (Fig. 11).
Eyes rusty brown. Antennifers prominent. Antennal segments 1-3 reddish
brown, slender; segment 4 reddish brown and slender basally, becoming
paler and swollen apically; segment 5 amber, swollen; length of segments
0.31, 0.62, 0.43, 0.56, 0.50 mm. Rostrum reaching metacoxae (Fig. 9); basal

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NEW YORK ENTOMOLOGICAL SOCIETY

segment shorter than bucculae; segments 2-3 pale brown, segment 4 dark
brown; length of segments 0.53, 0.77, 0.50, 0.43 mm. Width of head across
eyes 1.45 mm, between eyes 0.8 mm; length of head 1.4 mm.

Pronotum steeply declivous anteriorly. Humeral angles produced into
small knob; short cylindrical tubercle on dorsal surface of each humerus i
with distinct spine on lateral margin. Anterolateral margins of pronotum
rounded dorsoventrally, entire; anterolateral angles projecting as small
acute tooth. Pronotal length at meson 2.1 mm. Scutellum 2.9 mm wide at
base, equally long; apex broadly rounded. Basal half of coria with reddish
suffusion. Mesial impunctate line covering pronotum and scutellum.

Evaporative area on thorax granulose without fine rugae. Coxae, tro-
chanters and femora yellow brown; tibiae and tarsi darker brown. Abdom-
inal venter less heavily punctate than thorax.

Paratergites 8 medially fused. Paratergites 9 oblong, separated by second
gonocoxae. Male unknown.

Holotype. — Female labeled “Km 8 Est. do Aleixo Manaus AM. BV. 26-
VI-76. Nilu.” Deposited in U.S. National Museum. Type no. 72138. No
paratypes.

Grazia Rolston, new genus

Type species. — Piezodorus tinctus Distant, 1890.

Diagnosis. — Median spine on abdominal sternite 3 (second visible) pro-
jecting cephalad past mesocoxae, not attaining procoxae, largely obsuring
metasternum, compressed, acute apically. Mesial carinae of mesosternum
confined to anterior portion, weakly produced, widening near anterior me-
sosternal margin, not projecting onto prosternum. Ostiolar sulcus and ruga
on each side reaching about 7 tenths of distance from mesial margin of
ostiole to lateral margin of metapleuron. Femora unarmed; tibiae asulcate
or very weakly sulcate, superior surface rounded. Bucculae little produced
caudad of obtuse tooth, evanescent posteriorly; first rostral segment lying
entirely between bucculae. First segment of antennae not reaching apex of
head. Scutellum weakly convex basally.

Posterior surface of pygophore convex without median projection; genital
cup of normal size, lateral walls without tubercle. Thecal appendages ab-
sent.

Comments. — This genus superficially resembles Piezodorus but differs in
having a weak mesosternal carina, in lacking a median projection on the
posterior pygophoral surface and in having the opening of the genital cup
of normal size. The shape of the parameres and asulcate or very weakly
sulcate tibiae distinguish Grazia from Acrosternum.

We are pleased to dedicate this genus to Dr. Jocelia Grazia of the Uni-
versidade Estadual de Campinas in recognition of her contributions to the
taxonomy of neotropical pentatomids.

VOLUME LXXXVIII, NUMBER 4

263

Since the holotype of Piezodorus tinctus, type species, is of limited value
as a reference specimen by reason of being female and because some critical
characters were destroyed in pinning, this species is redescribed from a
series of 8 females and 4 males.

Grazia tine t a (Distant, 1890) New Combination

Tan to light castaneous with brown to black and reddish markings. Red-
dish submarginal line on lateral border of juga and line on anterolateral
margins of pronotum sometimes present. Fuscous to black markings often
present but varying in frequency are: suture between juga (mandibular
plates) and maxillary plates, thin line on lateral margin of juga, antennifers
and first antennal segment laterally, antennal segments 2-4, few to many
punctures plus surrounding areas on posterior disk of pronotum and base
of coria, 2 or 4 macules at base of scutellum, small spot on disk of each
corium and at distal end of each frenum. Black markings apparently present
consistently are: posterolateral angles of sternites 3-7 and corresponding
connexival angles, the latter mark sometimes extending partially or com-
pletely along posterior margin of each connexival segment.

Dorsum of head flat; punctation concolorous, rather fine and dense, ar-
ranged in subtle striae running anterolaterad from basal portion of tylus to
lateral margin of juga and apparently randomly elsewhere. Lateral jugal
margins sigmoid; apex of head moderately rounded; tylus a little longer than
juga. Width across eyes 2. 0-2. 3 mm, length 1. 6-2.0 mm; interocular dis-
tance 1.2-1. 4 mm. Antennal segments 0.4-0. 5, 0.45-0.5, 1.0-1. 3, 1.15-1.5,
1.15-1.5 mm long. Rostral segments 2-4 about 0.9-1. 1, 0.8-1. 1 , 0.7-0. 8 mm
long.

Pronotum more coarsely and sparsely punctate than head, punctation
usually a little stronger on posterior disk. Anterolateral margins straight.
Humeral angles slightly produced, rounded, anterolateral and posterolateral
margins forming right angle. Width at humeri 5. 1-6.2 mm, length at meson
2. 0-3.0 mm.

Scutellar apex narrowly rounded to subacute; punctation similar to that
on anterior pronotal disk; width at base 3. 2-3. 8 mm, length 3. 6-4. 5 mm;
frena extending along basal 5-6 tenths of lateral margins. Coria shallowly
and rather densely punctate, semitransparent, with posterolateral angle
reaching from about middle of fifth to anterior part of sixth abdominal seg-
ments; membranes vitreous, their boundary with coria sinuous. Connexiva
broadly exposed; posterolateral angle of each segment acutely produced.

Evaporative area matte, poorly defined. Abdominal venter tectiform, ob-
tuse median ridge forming continuous profile with basal spine.

Posterior margin of pygophore moderately concave from both dorsal and
caudoventral view. Head of parameres cupped, apical projection curving

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NEW YORK ENTOMOLOGICAL SOCIETY

V. p.

22

0- 3 mm

25 v.p.

27

0'5mm

Figs. 15-27. 15-20. Grazia tincta. 15. Pygophore, dorsal view. 16. Left paramere, outer

view. 17. Left paramere, inner view. 18. Aedeagus, lateral view. 19. Aedeagus. 20. Female
genitalia. 21-24. Kermana imbuta. 21. Pygophore, dorsal view. 22. Right paramere, outer
view. 23. Aedeagus, lateral view. 24. Spermatheca. 25. Kermana sp. Pygophore, dorsal view.
26-27. Kermana fucosa. 26. Right paramere, outer view. 27. Spermatheca, legend: paramere
(cl.), conjunctival lobe (co. 1.), dorsal conjunctival lobe (d.co.l.), process of genital cup (pr.),
ventral process of paramere (v.p.).

VOLUME LXXXVIII, NUMBER 4

265

dorsolaterad and cephalad, obscuring small dorsolateral projection on rim
of cup (Figs. 15, 16, 17). Conjunctiva with 3 pairs of lobes, each member
of dorsal pair long, digitiform (Figs. 18, 19). Endophalic duct sigmoid from
lateral view.

Genital plates as in Figure 20.

Distribution. — The 12 specimens seen came from the Dominican Repub-
lic, Panama, Venezuela, Ecuador, Brazil (Goias) and Paraguay.

Kermana Rolston, new genus

Type species. — Rhaphigaster imbutus Walker, 1867.

Diagnosis. — Median spine on abdominal sternite 3 projecting cephalad to
mesocoxae. Basal half of scutellum strongly convex. First antennal segment
not attaining apex of head. Bucculae evanescent at base of head; first rostral
segment lying entirely between bucculae. Ostiolar ruga on each side ex-
tending about 6-8 tenths distance from mesial margin of ostiole to lateral
margin of metapleuron. Mesosternum mildly tumescent on each side of me-
son; median carina weak, evanescent posteriorly. Metasternum not pro-
duced. Femora unarmed.

Genital cup bearing large process near rim on each lateral wall (Figs. 21,
25). Proctiger lacking tubercles. Ventrolateral process present on para-
meres, varying in size from knob to digit (Figs. 22, 26). Theca ovoid, lightly
sclerotized. Conjunctiva entirely membraneous, bilobed on each side, each
lobe broadly rounded at apex (Fig. 23).

Spermathecal bulb wide at base, distally bent through 180 degrees (Figs.
24, 27).

Comment. — The most recent generic (or subgeneric) placement of the 3
species assigned to Kermana has been Banasa or Acrosternum. Kermana
differs most notably from Banasa in having the abdominal spine projecting
beneath the metasternum. In Banasa the metasternum slopes ventrad in an
anterior to posterior direction and its posterior margin apposes the apex of
the abdominal tubercle. Kermana differs especially from Acrosternum in
possessing a large process on each lateral wall of the genital cup and in the
much greater convexity of the scutellum.

The synonymy of the 3 species and a key for their separation follow.

Kermana bucera (Stal, 1860). New Combination

1860 Rhaphigaster bucera Stal, Sv. Vet. Akad. Handl. 2(7):23.

1872 Banasa bucera : Stal, Sv. Vet. Akad. Handl. 10(4):43.

1909 Nezara (Banasa) bucera : Kirkaldy, Cat. Hem. 1:122.

Kermana fucosa (Berg, 1892). New Combination
1892 Nezara fucosa Berg, Ann. Soc. Cient. Arg. 33:9.

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NEW YORK ENTOMOLOGICAL SOCIETY

1909 Nezara (Acrosternum) fucosa: Kirkaldy, Cat. Hem. 1:118.

1948 Acrosternum fucosa : Piran, Acta Zool. Lill. 5:9.

Kermana imbuta (Walker, 1867). New Combination
1867 Rhaphigaster imbuta Walker, Cat. Het. 2:358.

1880 Banasa imbuta : Distant, Biol. Cent. Amer. Rhyn. 1:80, PI. 7, fig. 10.

1909 Nezara ( Atomosira ) imbuta : Kirkaldy, Cat. Hem. 1:122.

1910 Banasa imbuta : Banks, Cat. Nearctic Hem.-Het.:83.

1916 Banasa imbuta : Van Duzee, Check List Hem. Amer.:8.

1917 Banasa imbuta : Van Duzee, Cat. Hem. N. Amer.:62.

Key to Species of Kermana

1. Humeral angles strongly produced laterad, narrowly rounded api-

cally . . . (Brazil) bucera (Stal)

- Humeri scarcely produced 2

2. Punctation and color of scutellum almost uniform; excised para-
meres somewhat Y-shaped with long ventrolateral process (Fig. 26)

. . . (Argentina, Brazil, Uruguay) fucosa (Berg)

- Punctation of scutellum weakest and most sparse in light colored
areas at base and apex; excised parameres L-shaped with small ven-
trolateral process (Fig. 22) . . . (Texas to Costa Rica) imbuta (Walker)

Roferta Rolston, new genus

Type species. — Pentatoma marginale Herrich-Schaffer, 1836.

Diagnosis. — Median tubercle on abdominal sternite 3 short, obtuse,
scarcely reaching metacoxae. Scutellum strongly convex. First antennal
segment not attaining apex of head. Bucculae arcuate anteriorly, weakly
produced caudad of arch, evanescent at base of head; first rostral segment
lying entirely between bucculae. Ostiolar ruga on each side extending about
3 fourths of distance from mesial margin of ostiole to lateral margin of
metapleuron. Mesosternum mildly tumescent on each side of meson; median
carina moderately developed, extending full length of mesosternum. Meta-
sternum similarly but less strongly carinate posteriorly. Femora unarmed.

Genital cup bearing large process on each lateral wall near rim (Fig. 28).
Proctiger somewhat tuberculate subapically on each side. Parameres with
tubercle on cephalic margin. Median penial plates and penisfilum short (Fig.
30).

Spermathecal bulb with 2 arms (Fig. 31).

Comment. — This genus is similar to Kermana but differs especially in the
form of the parameres and spermathecal bulb. The type species, and only
known member of the genus, is readily recognized by the dorsal color:

VOLUME LXXXVIII, NUMBER 4

267

Figs. 28-31. Rofertia margincilis. 28. Pygophore, dorsal view. 29. Right paramere, mesial
face. 30. Aedeagus, lateral view. 31. Spermatheca, legend: process of genital cup (pr.).

fuscous with a broad orange-yellow border around the body (but not the
head) and median stripe of the same color on the pronotum and scutellum.
The synonymy of the species follow.

Roferta marginalis (Herrich-Schaffer, 1836) New Combination

1836 Pentatoma marginale Herrich-Schaffer, Wanz. Ins. 3:59, fig. 320.

1837 Pentatoma nitida Westwood, Cat. Hope 1:33-34 (synonymized by
Dallas, 1851, & Distant, 1900a).

1845 Rhaphigaster marginalis : Herrich-Schaffer, Wanz. Ins. 8:16.

1851 Rhaphigaster marginalis: Dallas, List Hem. 1:281.

1861 Ptilarmus marginalis: Stal, Stett, Ent. Zeit. 22:141.

1867 Strachia olivacea Walker, Cat. Het. 2:322 (synonymized by Distant,
1900a).

1867 Rhaphigaster marginalis: Walker, Cat. Het. 2:359.

1868 Strachia marginalis: Walker, Cat. Het. 3:561 (not 2:343-344).

1872 Nezara marginalis: Stal, Sv. Vet. Ak. Handl. 10(4): 40.

1892 Nezara marginalis: Berg, Anal. Soc. Cient. Arg. 33:6.

1893 Arocera olivacea: Lethierry & Severin, Cat. Hem. 1:159.

1893 Nezara marginalis: Lethierry & Severin, Cat. Hem. 1:166.

1900a Nezara marginalis: Distant, Ann. Mag. Nat. Hist. (7)5:392.

1900b Nezara marginalis: Distant, Proc. Zool. Soc. London: 823.

1909 Nezara (Nezara) nitida: Kirkaldy, Cat. Hem. 1:116.

1910 Nezara marginale: Valdes, Anal. Acad. Cien. Med., Fis. y Nat. Ha-
bana, Cuba. 46:429 (laps. cal.).

1932 Nezara nitida: Barber & Bruner, J. Dept. Agr. Puerto Rico 16(3):262-
263.

1935 Acrosternum nitida: Fennah, Trop. Agr. Trinidad 12:193.

1946 Acrosternum marginale: Barber & Bruner, Brooklyn Entomol. Soc.
40(2): 52-53.

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NEW YORK ENTOMOLOGICAL SOCIETY

35

33

39

34

d.c.

Figs. 32-39. Zorcadium truncatum. 32. Dorsal view. 33. Humeral angle of pronotum,
dorsal view. 34. Pygophore, dorsal view. 35. Pygophore, ventral border. 36. Right paramere,
lateral view; apices, dorsal view. 37. Aedeagus, lateral view. 38. Aedeagus, ventral view. 39.
Female genitalia, legend: dorsal conjunctival appendage (d.c.), ventral arm of paramere (v.a.).

1948 Acrosternum nitida : Callan, Proc. R. Entomol. Soc. London (B) 17
(9/10): 117.

1949 Acrosternum elegans Bruner & Barber, Mem. Soc. Cubana Hist. Nat.
19:160-161. NEW SYNONYMY.

1967 Acrosternum elegans : Alayo, Mus. Felipe Poey Trab. Divul. 43:27,
28, PI. 5, fig. 3.

1976 Acrosternum nitidum : Rolston, J. N.Y. Entomol. Soc. 84(1):3.

VOLUME LXXXVIII, NUMBER 4

269

The syntypes of Pentatoma nitida Westwood, one syntype of Strachia
olivacea Walker (the second syntype was not located) and the holotype of
Acrosternum elegans Bruner & Barber were examined. These specimens
appear to be conspecific.

Barber and Bruner (1946) noted that Kirkaldy (1909) apparently erred in
rejecting Pentatoma marginale Herrich- Schaffer, 1936, as preoccupied. I
have not found a prior usage of this name.

Zorcadium Bergroth, 1918

Zorcadium Bergroth, 1918, Ann. Mus. Natl. Hung. 16:307-308.

Type species. — Euschistus truncatus Fallou, 1888, by monotypy.

Diagnosis. — Median abdominal spine on sternite 3 stout, reaching me-
socoxae. Humeral angles greatly produced, cornute, much elevated (Fig.
33). Prosternum essentially flat; mesosternum slightly tumescent on each
side of meson, without carina; metasternum flat, not produced. Bucculae
prolonged as lobe at base of head, surpassing distal end of first rostral
segment; apex of rostrum reaching mesocoxae. Femora armed only by distal
extension of superior surface into small acute spine. Ostiolar canal long,
curved. Juga contiguous before tylus. First segment of antennae not reach-
ing apex of head, subequal in length to segment 2, half or less length of each
of last 3 segments. Costal angle of coria extending well past apex of scu-
tellum.

Zorcadium truncatum (Fallou, 1888)

Euschistus truncatus Fallou, 1888, Naturaliste p. 36.

Zorcadium truncatum : Bergroth, 1918, Ann. Mus. Natl. Hung. 16:308 (re-
description)

Yellowish brown base color; irregularly punctate with dark castaneous to
black, rather thickly so on dorsum (Fig. 32).

Head a little wider than long, 2.2 mm across eyes; 2.0 mm long. Lateral
margins of juga curving sinuously to narrowly rounded apex. Punctation
somewhat clustered on each side of tylus toward base, in part arranged in
irregular longitudinal lines on each side of vertex. Distance between eyes
1.4 mm, across ocelli 1.2 mm. Antenniferous tubercles largely exposed from
above; length of antennal segments 0.5, 0.5, 1.1, 1.1, 1.3 mm.

Pronotum 8.3 mm wide across humeri, 2.5 mm long at meson. Antero-
lateral angles produced into small tooth; anterolateral margin obtuse with
scattered black denticles. Humeral angles distally curving posteriorly with
posterolateral angle farther produced as subacute spine (Fig. 33). Anterior
pronotal disk strongly deflexed, calloused and impunctate mesially, behind
cicatrices subcalloused and somewhat sparsely castaneously punctate ex-
cepting 4 clusters of punctures arranged equidistantly in a nearly transverse

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NEW YORK ENTOMOLOGICAL SOCIETY

line; each cicatrice with large pale callus near middle; punctation in and
surrounding cicatrices mostly fuscous to black; punctation dense along front
portion of posterior disk.

Scutellum 3.8 mm wide at base, 3.4 mm long, with a pale subcalloused
basal spot near each angle; punctation sparse along midline; distal end of
each frenum marked by fuscous spot; lateral margins convex along frena, i
beyond frena parabolic with narrowly rounded apex. Punctation of coria
finer than on scutellum, lacking in lacuna near distal end of radial vein.
Membrane of hemelytra fuscous; veins few, darker, simple with occasional
spur. Connexivum narrowly exposed, finely punctate, a few punctures at
sutures black, others brown.

Evaporative areas sparsely and finely black punctate. Venter of head,
thorax and abdomen on sternites 2-4 between spiracular line and middle of
disk rather coarsely punctate; abdomen more finely punctate laterad of spi-
racular line, nearly impunctate and with broad dark streak down middle of
abdominal disk. Spiracles concolorous with surrounding part of sternites.

Ventral margin of pygophore with deep median U-shaped emargination
(Fig. 35), border developed into two distinct flattened lobes on each side of
emargination (Fig. 34); dorsal margin broadly arched. Proctiger box shaped,
flat dorsally, free margins narrow and vertical. Claspers C-shaped, com-
pressed; ventral arm longest, bifid, forming 2 short heavily pigmented blunt
fingers; dorsal arm short, concealed under dorsal margin of pygophore, api-
cally blunt and heavily pigmented (Fig. 36).

Theca oval, produced apically into narrow rim bearing pair of rounded
lobes dorsally. Dorsal conjunctival appendages forming large lightly scler-
otized shield surrounding ventral conjunctival appendages and endophalic
duct; shield produced laterally into flap on each side (Fig. 37); ventral con-
junctival appendages when not expanded enclosed by dorsal appendages,
bifid, forming 2 heavily pigmented horns, the ventral horn larger (Fig. 38).
Endophalic duct short, almost straight, projecting centrally at base of con-
junctiva. Median penial lobes absent.

Basal plates of female concealed; tenth sternite subquadrate (Fig. 39).

Voucher specimen. — Female labeled (a) “Forested eastern foothills of the
Andes, 2000 ft.” (b) “Peru: Tingo Maria 1 km E. of town. At edge of
woodland, 5. viii. 1971.” In the British Museum (Natural History).

Comment. — The holotype of Euschistus truncatus was in the Museum
National d’Histoire Naturelle, Paris, but has been misplaced or lost accord-
ing to Prof. J. Carayon (personal communication). In addition to the voucher
specimen, a male is in the private collection of Dr. H. Dodge Engleman.

Acknowledgments

We are grateful to Mssrs. W. R. Dolling of the British Museum (Natural
History), H. Dodge Engleman, R. C. Froeschner of the U.S. National Mu-

VOLUME LXXXVIII, NUMBER 4

271

seum, Luis de Santis of the Universidad Nacional de La Plata, Facultad de
Ciencias y Museo, Randall T. Schuh and P. Wygodzinsky, both of the
American Museum of Natural History for loans of specimens essential for
this work.

We wish to acknowledge especially the labor of Prof. J. Carayon in
searching fruitlessly for a Bergroth type and the critical reviews of parts of
the manuscript by Dr. Jocelia Grazia and Mssrs. H. Dodge Engleman and
Donald B. Thomas.

Dr. P. Alayo’s help was indispensible in solving the mystery of Valdes’
listing of Nezara marginale.

Literature Cited

Alayo D., P. 1967. Catalogo de la fauna de Cuba. XVIII. Los hemipteros de Cuba. II. Familia
Pentatomidae. Mus. “Felipe Poey” Acad. Cien. Cuba Trabajos Divulgation 43:1-47.
Banks, N. 1910. Catalogue of the nearctic Hemiptera-Heteroptera. Amer. Entomol. Soc.,
Philadelphia. 103 pp + viii.

Barber, H. G. and S. C. Bruner. 1932. The Cydnidae and Pentatomidae of Cuba. J. Dept.
Agr. Puerto Rico 16(3):23 1—280 + 3 pis.

and S. C. Bruner. 1946. Records and descriptions of miscellaneous Cuban Hemiptera.

Bull. Brooklyn Entomol. Soc. 4 1(2):52— 6 1 .

Berg, C. 1892. Nova Hemiptera faunarum Argentina et Uruguayenses. Ann. Soc. Cient. Arg.
33:7-11.

Bergroth, E. 1918. Hendecas generum Hemipterorum novorum vel subnovorum. Ann. Mus.
Natl. Hung. 16:298-314.

Bruner, S. C. and H. G. Barber. 1949. List of the Pentatomidae of Cuba, with the description
of a new species. Mem. Soc. Cubana Hist. Nat. 19(2): 155-165.

Callan, E. McC. 1948. The Pentatomidae, Cydnidae and Scutelleridae of Trinidad, B. W. I.

Proc. R. Entomol. Soc. London (B) 17:115-124.

Dallas, W. S. 1951. List of the specimens of hemipterous insects in the collection of the
British Museum. London. 592 pp.

Distant, W. L. 1880-1893. Insecta. Rhynchota, Hemiptera-Heteroptera. In Godman, F. D.
and O. Salvin. Biologia Centrali- Americana. Vol. 1. London.

. 1900. LII-Rhynchotal notes IV. Heteroptera: Pentatominae (part). Ann. Mag. Nat.

Hist. (7)5:386-397, 420-435.

. 1900. Revision of the Rhynchota belonging to the family Pentatomidae in the Hope

collection at Oxford. Proc. Zool. Soc. London (1900): 807-824 + 2 pis.

Fallou, J. 1888. Hemipteres nouveaux recueilles a Minas Geraes. Le Naturaliste (2)1:36. (not
seen)

Fennah, R. G. 1935. A preliminary list of the Pentatomidae of Trinidad, B. W. I. Trop. Agr.
(Trinidad) 12(7): 192-194.

Herrich-Schaffer, G. A. W. Wanz. Ins. 3:17-114 (1836) and 8:1-48 (1845).

Kirkaldy, G. 1909. Catalogue of the Hemiptera (Heteroptera). Vol. 1. Cimicidae. Berlin.
Lethierry, L. and G. Severin. 1893. Catalogue general des Hemipteres. Vol. 1. Brussels and
Berlin.

Piran, A. A. 1948. Contribution al conocimiento de la despersion geografica de los hemip-
teros neotropicales. Acta Zool. Lilloana 5:5-17.

Rolston, L. H. 1976. An evaluation of the generic assignment of some American Pentatomini
(Hemiptera: Pentatomidae). J. N.Y. Entomol. Soc. 84(l):2-8.

Rolston, L. H., F. J. D. McDonald and Donald B. Thomas, Jr. 1980. A conspectus of Penta-

272

NEW YORK ENTOMOLOGICAL SOCIETY

tomine genera of the Western Hemisphere-Part 1 (Hemiptera: Pentatomidae). J. N.Y.
Entomol. Soc. 88:120-132.

Stal, C. 1860. Bidrag till Rio Janeiro-traktens, Hemipter-fauna. K. Svenska Vet.-Akad. Handl.
2(7): 1-84.

. 1861. Miscellanea hemopterologica. Ent. Zeit. Stettin. 22:129-153.

. 1867. Bidrag till Hemiptererans systematik. Conspectus generum Pentatomidum

Americae. Ofv. K. Svenska Vet.-Akad. Forh. 24(7):522-534.

. 1872. Enumeratio Hemipterorum 2. Enumeratio Cimicinorum Americae. K. Svenska

Vet.-Akad. Handl. 10(4):3-65.

Valdes R., P. 1910. Clasificacion Gundlach de los hemipteros cubanos. Ann. Acad. Cien.
Med., Fis. y Nat. Habana, Cuba. 46:425-446.

Van Duzee, E. P. 1916. Check list of the Hemiptera (except Aphididae, Aleurodidae and
Coccidae) of America, north of Mexico. N.Y. Entomol. Soc. xi + 111 pp.

. 1917. Catalogue of the Hemiptera of America north of Mexico. Univ. Calif. Publ.,

Techn. Bull. 2. xiv + 902 pp.

Walker, F. 1867-1873. Catalogue of the specimens of Hemiptera Heteroptera in the collection
of the British Museum. 2:241-417 (1867) and 3:418-599 (1868).

Westwood, J. O. in F. W. Hope. 1837. A catalogue of Hemiptera in the collection of the Rev.
F. W. Hope, M. A. with short Latin diagnoses of the new species. Part 1. London 46

pp.

(LHR) Department of Entomology, Louisiana State University, Louisiana
Agricultural Experiment Station, Baton Rouge, Louisiana 70803 and
(FJDM) Department of Plant Pathology and Agricultural Entomology, Uni-
versity of Sydney, Sydney, N.S.W. Australia 2006.

Received for publication October 1, 1980.

VOLUME LXXXVIII, NUMBER 4

273

ACKNOWLEDGMENT

The Editors wish to express their appreciation to all those who have
helped in reviewing manuscripts submitted during 1980 for publication in
the Journal: D. F. Caccamise, W. H. Day, Robert Denno, Arnold Droz,
A. Forgash, H. Fowler, A. Gupta, F. O. Holmes, A. H. Hower, Jr., R.
Jeanne, A. Klots, R. K. Lindquist, G. Mailloux, R. W. Matthews, G. A.
Mount, H. D. Niemczyk, B. Puttier, L. H. Rolston, J. C. Schaffner, P. P.
Schubek, James Slater, Daniel E. Sonenshine, Daniel J. Sullivan, T. G.
Whitman, William Wolfe, T. Wood, and P. Wygodzinski.

HONORARY, LIFE, AND SUSTAINING MEMBERS

Charles P. Alexander, Joseph Bequaert, Raymond Brush, F. S. Chew,
Lucy Clausen, Howard E. Evans, Su Zan Swain Firmage, S. W. Frost,
Irving Granek, Mark Indenbaum, Alexander B. Klots, Kikumaro Okano,
Nellie Payne, L. L. Pechuman, Ruth R. Ralston, Hubert J. Thelan, George
Townes, Roman Vishniac, Kenneth D. Roeder (deceased), Edwin WayTeale
(deceased).

INDEX TO SCIENTIFIC NAMES OF ANIMALS AND PLANTS
VOLUME LXXXVIII

Generic names begin with capital letters. New genera, species, subspecies and varieties are
printed in italics. The following items and articles are not indexed:

1. Habitat association of adult cicadas in northeastern Costa Rica, below 250 m elevation,
p. 91.

2. Key to genera of Pentatomini, Section 3. pp. 121-123.

3. Insects employing plant tissue to transduce vibratory signals, p. 211.

4. Characteristics of the postcubital stridulitrum of some primitive Pentatomoidea. p. 232.

5. Key to genera of Pentatomini, Section 2. pp. 254-260.

6. Synonymy of three species of Kermana. pp. 265-266.

7. Synonymy of Roferta marginalis . pp. 267-268.

8. Authors and abstracts of papers presented at the 51st Annual Meeting. Eastern Branch,
Entomological Society of America, pp. 33-82.

Acacia karroo, 80
Acanthomyops plumipilosus, 207
Acer rubrum, 188
saccharum, 188
Acheta domesticus, 79
Acizzia russellae, 80
Acromyrmex versicolor, 102
Acrosternum, 262
elegans, 269
Actia, 51

autumnalis, 52
diffidens, 52
interrupta, 52
labellata, 52
pauciseta, 52
rufescens, 52
Acutalis anticonigra, 183
inornata, 174
semicrema, 174
tart area, 174
tartarea inornata, 182
tartarea semicrema, 182
tartarea tartarea, 182
Aedes aegypti, 49, 73
canadensis, 82
cantator, 82
excrucians, 82
sollicitans, 51
stimulans, 47
vexans, 82
Aenictus gracilis, 15
Agromyza frontella, 56, 61, 252
Aleixus, 257

virgatus, 257

Aleurocanthus woglumi, 52
Alsophila pometaria, 70

Amblyseius andersoni, 77
fallacis, 252

Ambrosia artemisiifolia, 174
Amnestus, 231
Anaphes flavipes, 39
Anaplasma marginale, 76
Anoetus gibsoni, 61
Anomma wilverthi, 24
Anystis agilis, 255
Apantales fumiferanae, 38
Aphanogmus, 256
Aphantorhapha, 51
Aphis neri, 1 15
Apis, 203

mellifera, 43, 73
Archips, 68

Argyrotaenia velutinana, 197
Aristolochia, 226
Ascosphaera apis, 74
Atta mexicana, 102
Autographa californica, 55
Azteca chartifex, 104

Babesia caballi, 76
Banasa, 265
Bathycoelia, 120
Bathyplectes anurus, 151
curculionis, 151
Beauveria bassiana, 40
Bethyloids, 204
Betula lenta, 188
Bidens bipinnata, 176
Blattella germanica, 33, 36, 40, 46, 52
Bombus, 205
Braula coeca, 74

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NEW YORK ENTOMOLOGICAL SOCIETY

276

Callidea duodecimpunctata, 233
Callistephus chinensis, 174
Calotrips procera, 115
Camponotus beebei, 104
femoratus, 104
Carduus acanthoides, 44
nutans, 44, 78
Carineta indecora, 94
Carya cordiformis, 188
tomentosa, 188
Catharanthus roseus, 174
Catocala, 72

Cecidomyia serotinae, 68
Ceranthia, 51
Cercaris watlingensis, 45
zonata, 170
Ceromya, 51

Ceroplastes ceriferus, 70
Chaetostigmoptera, 51
Cheliomyrmex morosus, 24
Chiastosternum, 234
Chilomenes sexmaculata, 115
Choristoneura fumiferana, 38
Chrysops, 58
atlanticus, 50
Cinara, 42

Cirsium arvense, 211

Coccinella transversoguttata richardsoni, 57

Colaspis brunnea, 45

Coleomegilla maculata, 57

Colletes thoracicus, 66

Coquillettidia perturbans, 82

Cordia alliodora, 94

Crematogaster limata parabiotica, 104

Cucumis melo, 189

Culex pipens morestus, 118

Dactylis glomerata, 210
Delia, 170

Delphacodes lutulenta, 36
Dendrocoris, 120
Dermacentor albipictus, 75
Diamma bicolor, 205
Diaparsis, 39
carinifer, 39

Diaphnocoris chlorionis, 81
Dirofilaria immitis, 51, 82
Dolichovespula arenaria, 146
Doratora stylata, 36
Dorylus labiatus, 16, 133
Dorylus (Anomma) nigricans, 133
wilverthi, 133

Dorymyrmex emmaericaellus, 16
Drosophila, 68
grimshawi, 68
melanogaster, 58, 241

Eciton, 24

burchelli, 24
hamatum, 24, 133
Elanela hevera, 120
Empoasca, 81
Encarsia formosa, 63, 254
Entomophthora muscae, 236
Ephemerella, 44
Epilachna varivestis, 76
Epistylis cambari, 113
Epitrix hirtipennis, 72
Euschistus truncatus, 269

Fagus grandifolia, 188
Fecilia biorbis, 29
minor, 29
nigridens, 29
Feltiella Carolina, 255
Fidicina amoena, 94
mannifera, 86
pronoe, 86
sericans, 86
spinocosta, 86
Fiorinia externa, 56
Formica rufa, 16
Fraxinus americana, 188
nigra, 188

Gasterophius intestinalis, 41
Glaucias, 127

Glaucioides englemani , 120
Glycobius speciosus, 64
Glypta fumiferanae, 38
Goethalsia meiantha, 87
Gomphus quadricolor, 113
Graminella nigrifrons, 36
Granadilla, 217
Grazzia, 257
tincta, 257

Hamamelis virginiana, 188
Heliconius cydno, 217
heurippa, 224
melpomene, 224
pachinus, 224
sapho, 217

Heliothis virescens, 53, 55, 72
zea, 55
Hevea, 218

Hippodamia convergens, 57
Hoplocampa testudinea, 69
Hybomitra, 58

lasiophthalma, 58
Hylaeus, 199
Hylotrupes bajulus, 39
Hypera postica, 151

VOLUME LXXXVIII, NUMBER 4

Inga, 94

Ilex cornuta var. bufordii, 70
Incisitermes minor, 36

Kalmia latifolia, 188
Kermana, 257
bucera, 257
fucosa, 257
imbuta, 257

Labidus, 26
Larrea tridentata, 102
Lemophagus curtus, 39
Leptinotarsa decemlineata, 40, 55
Leptothrips mali, 64
Linaria vulgaris, 211
Lindera benzion, 188
Liriodendron tulipifera, 188
Lotus corniculatus, 43
Lymantria dispar, 33, 60

Machaerium seemanii, 226
Macropsis fumipennis, 81
Majeorona bovilla, 94
Malacosoma americanum, 46
Mallota posticata, 59
Manduca sexta, 72
Medicago sativa, 252
Microtonus aethiopoides, 151
colesi, 151
Microstigmus, 206
comes, 206

Micrutalis malleifera, 174
semicrea, 183
tart area, 183
Monacis debilis, 104
Morpho peleides, 226
Musca domestica, 33, 236
Myristica fragans, 90
Myrmecocystus mexicanus, 7
mimicus, 9
Myrmica rubra, 16
ruginodis, 16

Myzus persicae, 35, 57, 65, 74

Neivamyrmex, 26
harrisi, 133

Neodiprion lecontei, 42
Neoseiulus californicus, 252
fallacis, 77
mckenzei, 77
Nocheta adda, 120

Oecanthus, 210
nigricornis, 210
Olneya tesota, 102

Ophullela pelidna pelidna, 170
Orientus ishidae, 81
Osmerus mordax, 189
Ostrinia nubilalis, 48
Oulema melanopus, 39

Pachydiplax longipennis, 113
Parides, 217

Passiflora guazumaefolia, 218
vitifolia, 218
Pelecinus, 201

Pemphigus populitransversus, 106
Pentaclethra macroloba, 87
Pentatoma marginale, 257
nitida, 269
Peribaea, 51

Periplaneta americana, 36, 66
australasiae, 66
brunnea, 66
fuliginosa, 66
japonica, 66
Philaethria dido, 223
Phineus, 120

Phyllonorycter blancardella, 81
crataegella, 81
Phytophthora infestans, 40
Phytoseiulus persimilis, 252
Piezodorus tinctus, 257
Pinus rigida, 42
Plagiodera versicolor, 67
Plantago insularis, 102
Platanthera cristata, 197
Plathypena scabra, 63
Plodia interpunctella, 78
Podisus, 46

maculiventris, 46
modestus, 46
placidus, 46

Pogonomyrmex occidentalis, 12
Polistes fuscatus, 143, 146
fuscatus aurifer, 146
Pollenia rudis, 237
Populus deltoides, 106
sargentii, 106
Porthetria dispar, 52, 69
Prionyx thomae, 170
Proarna sallei, 94
Prosopis velutina, 102
Prunus serotina, 67
Psammotettix lividellus, 36
Pseudosiphona, 51
Psylla pyricola, 48
Pyrus communis, 48
ussuriensis, 48

278

NEW YORK ENTOMOLOGICAL SOCIETY

Quercus prinus, 188
rubra, 188
velutina, 188
Quesada gigas, 86

Reticulitermes, 36
Rhagoletis pomonella, 34, 69
Rhaphigaster imbutus, 265
Rhinocyllus conicus, 44, 78
Rhododendron nudiflorum, 188
Rhyacionia, 42
Rhytidoponera metallica, 16
Roferta, 257

marginalis , 257
Rubus, 210

Salix babylonica, 67
Sarracenia purpurea, 60
Scleroderma, 204
Scutellera nobilis, 233
Sehirus, 231
Simulium venustum, 53
verecundum, 53
Siphona, 51
Sitophilus oryzae, 62
Solanum melongena, 174
tuberosum, 57
Solenopsis invicta, 16
Solidago, 211
Somula decora, 59
Spodoptera frugiperda, 55
Steganocerus, 234
Strachia olivacea, 269
Stragania apicalis, 81
Sus scrofa, 194

Tabanus, 58
atratus, 58
lineola, 58
nigrovittatus, 75
quiquevittatus, 58

Tachytes crassus, 171
distinctus, 170
tricinctus, 170
validus, 172
Tessaratoma, 231
papillosa, 231
Tetranychus urticae, 252
Tetraopes tetraophthalmus, 42
Tetrastichus incertus, 151
julis, 39

Thaumastella aradoides, 233
namaquensis, 233
Theobroma cacao, 218
Toumeyella, 42
Tragopa brunnea, 183
Trialeurodes vaporariorum, 63, 253
Trichoplusia ni, 55
Trifolium pratense, 252
Triglyphothrix striatidens, 206
Troglodytes aedon, 143
Tsugaspidiotus tsugae, 56
Typhus latifolia, 21 1

Vaccinium, 188
Verbascum thapsis, 65
Vernonia patens, 88
Vespula pennsylvanica, 203
Viburnum acerifolium, 188
Vidada dollingi, 120
Vismia guianensis, 88

Xenopus laevis, 242

Zammara smaragdina, 86
Zanthoxylum procerum, 93
Zootermopsis angusticollis, 50
Zorcadium, 257
truncatum, 257

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Vol. LXXXIX

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The New York Entomological Society

Incorporating The Brooklyn Entomological Society

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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

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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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Officers for the Year 1981

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President , Dr. Joseph M. Cerreta

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Columbia University, New York 10032

Vice-president , Dr. Gerard Iwantsch

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Fordham University, New York 10458

Secretary , Dr. Henry M. Knizeski, Jr.

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Mercy College, Dobbs Ferry, New York 10522

Assistant Secretary , Dr. Irene Matejko

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- Secaucus, New Jersey

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Treasurer , Mr. Louis Sorkin

American Museum of Natural History, New York 10024
Assistant Treasurer , Dr. Sally B. Padhi

Rutgers University, New Brunswick, New Jersey 08903

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' ' / x A' ,• / I \ . 'jT, \ ■ , . y v 'W-

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Journal of the

New York Entomological Society

VOLUME LXXXIX MARCH 1981 NO. 1

EDITORIAL BOARD

Editor Publication Committee

Dr. Karl Maramorosch Dr. Randall T. Schuh

Waksman Institute of Microbiology American Museum of Natural History

Rutgers University Dr Louis Trombetta

New Brunswick, New Jersey 08903 St Johns University

Associate Editors
Dr. Lois J. Keller, RSM
Dr. Herbert T. Streu

CONTENTS

The occurrence and relevance of arthropods of medical and veterinary importance
captured during a survey on Plum Island, New York

Dennis J. White and Cherylann P. White 2-15

Control of Dermacentor variabilis. 1 . Larval and adult susceptibility to selected
insecticides Dennis J. White and Jorge L. Benach 16-22

Control of Dermacentor variabilis. 3. An analytical study of the effect of low volume
spray frequency on insecticide-stressed and nonstressed populations

Dennis J. White, Jorge L. Benach, Laurel A. Smith and Soo P. Ouyang 23-33

Further evidence for chlorpyrifos tolerance and partial resistance by the Japanese
beetle (Coleoptera: Scarabaeidae) Sami Ahmad and Yuen-Shaung Ng 34-39

Ochlerini, a new tribe in Discocephalinae (Hemiptera: Pentatomidae) L. H. Rolston 40-42

Notes on the biology of a common sunflower bee, Melissodes ( Eumelissodes )
agilis Cresson F. D. Parker, V. J. Tepedino, and G. E. Bohart 43-52

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(l), 1981, pp. 2-15

THE OCCURRENCE AND RELEVANCE OF ARTHROPODS OF
MEDICAL AND VETERINARY IMPORTANCE CAPTURED
DURING A SURVEY ON PLUM ISLAND, NEW YORK

Dennis J. White and Cherylann P. White

Abstract. — Since 1954, the United States Department of Agriculture has
maintained a high containment biological research laboratory on Plum Is-
land, New York, involved with research and disgnoses of exotic commu-
nicable animal diseases. Isolation from the Long Island mainland provides
a physical barrier to foreign disease transmission to mainland livestock and
of domestic animal diseases to Plum Island livestock. At the request of the
Plum Island Animal Disease Center administration, this survey was con-
ducted to define the entomological fauna of medical and veterinary impor-
tance and to determine potential vectors of both native and migrating
species. In addition to the insect survey, mosquito pools were used for
native virus isolation attempts, sera from mammalian and avian specimens
were used for detection of antibodies to arboviruses and engorged mosqui-
toes were used for blood meal identification.

Twenty native culicid and fifteen tabanid species predominated in the
collections in addition to at least three species of migrants from the Orient
Point, New York salt marshes. All mosquito pools and avian sera were
negative when screened for CEV, EEE, SLE, WEE and POW viruses or
antibodies. Two mammalian sera reacted by HI to California and St. Louis
encephalitis viruses at low titers. Mosquito blood meal analyses showed an
avian host preference for the Culiseta species and a rodent host preference
for the Aedes species.

Introduction

The United States Department of Agriculture animal disease research
facility (Anonymous 1975) has been located on Plum Island since 1954. The
center is engaged in both basic and applied research on exotic communicable
diseases of animals and is the only facility in the United States so desig-
nated. In addition to educational and technical services provided to United
States and foreign governments, the Plum Island Animal Disease Center’s
(PIADC) responsibilities are threefold: develop diagnostic capability of ex-
otic animal diseases, conduct basic and applied research on these diseases
and their causative organisms, and develop procedures for safe importation
of foreign wild and domestic animals, semen, meat and other animal prod-
ucts. The central objective of the facility is to assist in the prevention of the

VOLUME LXXXIX, NUMBER 1

3

Y-8

Fig. 1. Habitat differentiation and quadrant delineation for Plum Island survey. R = old
estuarine and predominant lentic vegetation; B = dominant upland vegetation; Y = coastal
and upland vegetation in close proximity.

introduction of animal disease that could result in death or serious economic
losses in susceptible livestock populations in the United States.

Due to security measures necessary for P3 level biological containment,
rigid safety regulations have been established. All movement to, from and
on the island is strictly monitored to eliminate all chances of unintentional
transmission of microorganisms or contaminated material. In order to en-
hance biological containment, Congress stipulated that the Center be located
on an island entirely under Federal control and surrounded by deep navi-
gable water so as to provide a physical barrier for transmission of foreign
diseases from Plum Island to the mainland and domestic diseases from the
mainland to Plum Island. Plum Island is located 2.1 km from the eastern tip
of Long Island’s north fork and covers 320 ha. At its widest, the island is
1.6 km wide, which tapers to 0.1 km at the narrowest part and is 4.3 km
long (Fig. 1).

The announcement of a proposal to study Rift Valley Fever, a mosquito-
borne disease presently epidemic in Africa, prompted local public officials
to visit the facility. In December 1978, New York State Department of
Health officials toured the PIADC. During the course of this visit, State
Health Department officials were invited to conduct an entomological sur-
vey of the island. The purpose of this report is to define the entomological
fauna native to, or captured on, Plum Island that is of medical or veterinary
significance and present the results of a basic arbovirus survey.

4

NEW YORK ENTOMOLOGICAL SOCIETY

on/s « J

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PLUM I S L{A N D

RESTRICTED
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DANGEROUS ANIMAL DISEASES

Plum Gut
Har.bor

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Fig. 2. Topographic map of Plum Island.

Materials and Methods

Licenses were received from the United States Department of the Inte-
rior, Fish and Wildlife Service for avian collection and from the New York
State Department of Environmental Conservation for mammal collection.
The survey began on 21 May 1979 and terminated 25 September 1979. Prior
to specimen collection, maps were drawn on a grid system to readily provide
locality descriptions and habitat definitions.

The island was divided into 3 sections (Fig. 1) based on predominant
topographical features (Fig. 2). The R sector, ca. 150 ha, is distinguishable
by the predominant old estuarine and lentic vegetation associated with lower
elevations (Table 1). The B sector, ca. 130 ha, is characterized by deciduous
vegetation associated with higher elevations. The Y sector, ca. 40 ha, in-
cluded the eastern peninsula and is characterized by upland and coastal
vegetation in close proximity.

Diverse collection techniques were employed. Phototropic mosquitoes
were collected by 2 permanent New Jersey light traps and four portable
CDC light traps. Resting boxes were distributed throughout the island. A
malaise trap was placed in an area where tabanids were abundant. Ixodid
ticks were captured by dragging a 1 m2 piece of white flannel cloth along

VOLUME LXXXIX, NUMBER 1

5

Table 1.

Dominant vegetation found in each of the three major habitats on Plum Island.

R-Sector

Baccharis halimifolia L. — Groundsel bush

Iva oraria (Bartlett) — Marsh elder

Hibiscus palustris (L.) — Rose mallow

Phragmites communis Trin. — Ditch reed

Rhus radicans L. — Poison ivy

Myrica pensylvanica Loisel — Bayberry

Rhus copallinum L. — Shining sumac

Pinus resinosa Ait. — Red pine

Chamaecyparis thyoides (L.) — Atlantic white cedar

Ammophila breviligulata Fern. — Dune grass

Typha angustifolia L. — Cattail

B-Sector

Quercus velutinci Lam. — Black oak
Quercus coccinea Muenchh. — Scarlet oak
Quercus palustris Muenchh. — Pin oak
Acer platanoides L. — Norway maple
Salix babylonica L. — Weeping willow
Populus tremuloides Michx. — Quaking aspen
Populus alba L.— White poplar
Nyssa sylvatica Marsh — Black tupelo
Prunus serotina Ehrh. — Black cherry
Sassafras albidum (Nutt.) — Sassafras
Rhus copallinum L. — Shining sumac
Rhus typhina L. — Staghorn sumac
Rhus radicans L. — Poison ivy
Lonicera japonica Thunb. — Honeysuckle
Myrica pensylvanica Loisel — Bayberry
Rubus cuneifolius Pursh. — Blackberry

Y-Sector

Rosa virginiana Mill. — Wild rose

Phragmites communis Trin. — Ditch reed

Rhus radicans L. — Poison ivy

Myrica pensylvanica Loisel — Bayberry family

Rhus copallinum L. — Shining sumac

Rhus typhina L. — Staghorn sumac

Rubus cuneifolius Pursh. — Blackberry

Quercus velutina Lam. — Black oak

Quercus coccinea Muenchh. — Scarlet oak

Quercus palustris Muenchh. — Pin oak

Prunus serotina Ehrh. — Black cherry

Ammophila breviligulata Fern. — Dune grass

Chamaecyparis thyoides (L.) — Atlantic white cedar

6

NEW YORK ENTOMOLOGICAL SOCIETY

vegetation. Attempts to locate argasid ticks were done by sifting sand sur-
rounding seagull nests (H. Hoogstraal, NAMRU-3, Cairo, Egypt, personal
communication). Immature mosquitoes were collected by dipping. Sweep
nets were used to capture blood seeking specimens of mosquitoes, deer
flies, horse flies, stable flies and other filth-breeding muscids. Biting species
too small to be retained in a sweep net (e.g. Culicoides ) were collected only
as they were feeding. Active ectoparasites (ticks, fleas, lice, etc.) were re-
moved from hosts captured in Hav-a-hart mammal traps or avian mist nets.

In addition to the arthropod specimens, 0.1 to 0.9 cc of blood was drawn
from each of the captured mammals and birds to detect presence of antibody
to arboviruses. Giemsa-stained blood smears of 23 mammals were made for
detection of protozoal parasites (i.e. Babesia microti). The hemolymph test
(Burgdorfer 1970) was performed on 80 captured Dermacentor variabilis
(Say). Arbovirus isolation attempts using suckling mice were made from
frozen pools of mosquitoes collected in CDC traps. Engorged mosquitoes
captured by any method were used for blood meal identification using the
capillary-precipitin technique (Edman 1971). Thirty-five avian and 23 mam-
malian sera were tested by hemagglutination-inhibition tests (Clark and Cas-
als 1958).

Selected specimens of all invertebrates collected have been presented to
the PIADC for reference purposes.

Results

Island topography. — The island ranges in elevation from 0 to 30.5 m
above sea level (Fig. 2) and is surrounded by a rocky-sandy beach ca. 10-
20 m wide at high tide. A once existent salt marsh at the extreme southwest
part of the island had been diked to prevent tidal inundation. Routine water
analyses performed by the PIADC staff confirmed our observations that
no collections of salt or brackish water are present on the island. Three ma-
jor fresh water swamps cover ca. 50 ha in the western half of the island and
a few small fresh water ponds are scattered over the island. The sandy na-
ture of the island allows quick percolation or run off of rain water from the
surface. The upland areas are densely covered with scrub vegetation
(Table 1).

Arthropod specimens. — A summary of all captured specimens is included
in Table 2. Twenty-two species of mosquitoes were collected during the
study, 8 as larvae and 19 as adults. Ten species were collected from the
fresh water swamp group, only 4 of which were captured as immatures. The
relatively large proportion of specimens of this group (74.5% of adult mos-
quitoes) illustrates the extent of breeding that occurred in these swamps.
Most of these species were captured in CDC light traps.

Many discarded tires used around a sewage settling pond in the R-6 quad-

VOLUME LXXXIX, NUMBER 1

7

Table 2. Number and location of specimens captured on Plum Island.

Imma-

Collected species tures Location Adults Location

Diptera

Culicidae

— Woodland pool group

Aedes canadensis (Theobald)

12

R-17

Ae. stimulans (Walker)

9

R-17; B- 11,16

Ae. aurifer (Coquillett)

6

R-13,17

Ae. cinereus Meigen

58

R- 13,17

Ae. excrucians (Walk.)

29

R-17, 19; B-11,16

Ae.fitchii (Felt and Young)

40

R-6, 13, 16, 17; B-10,11

— Floodwater group

Ae. vexans (Meig.)

9

Y-6

410

R-6,7; B-3, 11, 15, 16

— Salt marsh group

Ae. cantator (Coq.)

5

R-17

Ae. sollicitans (Walk.)

14

R-6, 12, 16, 17;

B-3,4,15; Y-6

Culex salinarius Coq.

153

R-6,7; B-3

1

B-15

— Fresh water swamp group

Cx. pipiens L.

106

R-6; B-3

5

R-17; B-15

Cx. restuans Theo.

176

B-3

Cx. territans Walk.
Culiseta melanura (Coq.)

2

R-16

13

R-17; B-15

Cs. morsitans dyari (Coq.)

837

R-6, 10,17; B- 1,3,11,

Cs. silvestris minnesotae

15,16; Y-6

(Barr)

Coquillettidia perturbans

291

R-6,7, 17; B- 1,3,1 1,15

(Walk.)

130

R-6,7, 14, 17;

Uranotaenia sapphirina

B- 1,3,11,15

(Osten Sacken)

11

R-17

585

R-17; B-15

Anopheles crucians

1

B-l 1

Wiedemann
An. earlei Vargas

1

R-17

— Treehole group

Ae. triseriatus (Say)

48

R-6; B-3

52

R-6,7, 16, 17; B-3, 6, 11

— Rockhole group

Ae. atropalpus (Coq.)

253

B-3

Ceratopogonidae

Culicoides hollensis

2*

R-l

(Melander and Brues)
Cul. crepuscularis Malloch

1*

B-l 1

Tabanidae

Tabanus nigrovittatus

1

R-l 8

Macquart

T. lineola Fabricius

1

B-15

8

NEW YORK ENTOMOLOGICAL SOCIETY

Table 2. Continued.

Imma-

Collected species tures Location Adults Location

T. sackeni Fairchild

1

B-l 5

T. fulvulus Wied.

1

B-15

Hybomitra lurida (Fallen)

4

R- 17

Chrysops vittatus Wied.

2

R-17; B-10

C. montanus Osten Sacken

8

B-6; R-17

C. flavidus Wied.

3

R-17; B-6, 10, 13

C.frigidus O.S.

17

R-17; B-6, 15

C. pikei Whitney

1

B-6

C. callidus O.S.

1

R-17

C. univittatus Macq.

1

B-3

C. obsoletus Wied.

413

R-l ,6,7,12,17,18;
B-3, 6, 10, 1 1

C. sackeni Hine

2

B-10

Muscidae

Musca domestica L.

innumer-

able

throughout island

Stomoxys calcitrans (L.)

16

R-17; B-16; Y-1,3,5,6

Sarcophagidae

innumer-

able

throughout island

Calliphoridae

Phormia regina (Meig.)

2* R-4

innumer-

able

throughout island

Cuterebridae

1* Y-6

3

R-16; B-15

Mallophaga

Philopteridae

6*

B-6, 10, 15

Siphonaptera

Hystrichopsyllidae

Orchopeas howardii (Baker)

21

R-6,17; B-l 1

Ctenophthalmus pseudagyrates

6

R-6; B-l 1

Baker

Acarina

Ixodidae

Dermacentor variabilis (Say)
Ixodes dammini Spielman,

6* B-l 1

80

R-17, 18; B-l 1; Y-6

Clifford, Piesman and Corwin
Mammalia
Rodentia

Peromyscus leucopus

17

R-6, 10, 17; B-l 1; Y-6

Microtus pennsylvanicus

6

R-6; B-l 1; Y-6

VOLUME LXXXIX, NUMBER 1

9

Table 2. Continued.

Imma-

Collected species tures

Location

Adults

Location

Avian

Passeriformes

Dumetella carolinensis (L.)

14

B-6,16; Y-3,5

Empidonax minimus (Baird

1

Y-5

and Baird)

Dendroica striata (Forester)

2

Y-3,5

Dendroica pinus (Wilson)

1

B-16

Pipilo erythrophthalmus (L.)

2

Y-3,5

Melospiza georgiana (Latham)

1

Y-5

Turdus migratorius L.

3

B-6,15

Icterus galbula (L.)

2

Y-3

Sturnus vulgaris L.

2

B-15

Colaptes auratus (L.)

7

B-10,15

* Ectoparasites: removed from host.

rant, and discarded animal cages and tires in a dump within the B-3 quadrant
supported larval development of both Aedes triseriatus (Say), and Ae. atro-
palpus (Coquillett) (White and White 1979). This represented a new county
record for Ae. atropalpus.

With the exception of one Tabanus nigrovittcitus Macquart, all captured
tabanids were fresh water swamp breeders. The extensive swamp land pro-
vides an ideal larval breeding habitat, especially during the mid-summer dry
spell when vast areas of mud become exposed.

Eighty adult D. variabilis were collected during the survey, 42 males and
38 females, predominantly located at the narrowest part of the island (Y-6).
The adults persisted at this area until 20 August. All D. variabilis were
negative for rickettsiae. Six immature Ixodes dammini Spielman, Clifford,
Piesman and Corwin were removed from both Microtus pennsylvanicus and
Peromyscus lecopus hosts captured at B- 11, a dense woodland. Despite
extensive efforts to locate members of the argasid family, none were found.

The house fly, Musca domestica L. found throughout the island in the
adult stage. Since it was common, collection efforts were not extensive.
The stable fly, Stomoxys calcitrans (L.) was found in many areas of Plum
Island. Most specimens were collected near the animal holding facility in
the Y sector.

The flesh and blow flies were also extremely numerous. The common
larval breeding source proved to be carcasses of dead birds, especially gulls.

10

NEW YORK ENTOMOLOGICAL SOCIETY

The large gull population (2,500-3,000 birds) is reaching an “over-popula-
tion” point and many of the young birds do not survive the nesting period.
Various roadkills also provided excellent oviposition and breeding habitat
for members of both families. No attempt was made to enumerate the prev-
alence of these flies.

In addition to 3 adult cuterebrid specimens captured by sweeps, a Mi-
crotus pennsylvanicus vole was captured that was parasitized by a larva of
this family. Avian ectoparasites were not numerous. We collected 6 Phil-
opteridae soley for inclusion into the insect collection. Only 3 specimens of
Culicoides were captured as they attempted to feed.

All fleas were collected from their anesthetized hosts; 6 Ctenophthalmus
pseudagyrates Baker and 21 Orchopeas howardii (Baker) were removed
from meadow vole and white-footed mouse hosts.

All mosquito pools ( 10-100 mosq./pool) submitted for viral isolations were
negative when screened for presence of CE, EEE, SLE, WEE and POW
viruses. Blood meal analyses showed, as expected, that the avian popula-
tions were supporting the Culiseta species and that the rodent population
was supporting the Aedes species.

Mammal survey. — Despite 500 trap-nights throughout Plum Island, only
2 species of mammals were captured. In addition to these widespread and
fairly numerous Peromyscus leucopus and M. pennsylvanicus populations,
a few wild house cats, Felis domestica, and bats exist on the island. Re-
spectively, these offer a natural control against rodent and flying insect
populations. Other than these four mammals, no other wild animal was
found on Plum Island.

All blood smears taken from the captured mammals were negative for
intracellular protozoal parasites. The serum from one P. leucopus reacted
at 1:40 to California encephalitis virus; another P. leucopus serum reacted
at 1:20 to St. Louis encephalitis virus. All other serological tests were neg-
ative.

Cold-blooded animals seen on the island included a few garter snakes and
numerous box and spotted turtles.

Avian survey. — Birds provide the most common source of blood on Plum
Island. Since such large seagull ( Larus argentatus Pontoppidan and Larus
marinus L.) and goose ( Branta canadensis (L.)) populations coexist with
a small mammal host pool, the majority of mosquitoes on Plum Island are
preferentially bird feeders (Crans 1964; Downe 1962; Edman et al. 1972).

A total of 35 passerine birds was sampled for antibodies to domestic
arboviruses (EEE, WEE and SLE), and all were negative. Almost one-half
of those sampled were catbirds ( Dumetella carolinensis (L.)), an abundant
temporary Plum Island inhabitant. Other species sampled are listed in Table
2. Numerous sightings of a visiting Bald eagle ( Haliaeetus leucocephalus

VOLUME LXXXIX, NUMBER 1

11

(L.)) were made in mid-August. There were also 7 nesting pair of Osprey
( Pandion haliaetus (L.)) inhabiting the island. None of the shore birds,
geese or raptorials were sampled in this project.

Discussion

Of the 55 infectious animal diseases contained at the PIADC, only 13
(possibly 14) viral disease agents may be transmitted by arthropods. The
known natural and experimental vectors of these diseases include the fol-
lowing (Berge 1975; R. Shope, YARU, New Haven, Ct., personal com-
munication).

DISEASE

VECTOR

African Horse Sickness (AHS)

African Swine Fever (ASF)

Akabane (AKA)

Bluetongue (BLU)

Bovine Ephemeral Fever (BEF)
Eastern Equine Encephalitis (EEE)

Epizootic Hemorrhagic Disease
(EHD)

Ibaraki (IBA)

Louping 111 (LI)

Nairobi Sheep Disease (NSD)

Rift Valley Fever (RVF)

Venezuelan Equine Encephalitis

Vesicular Stomatitis Virus (VSV)

Western Equine Encephalitis (WEE)

Culicoides, Aedes, Anopheles, Cu-
lex spp.

Ornithidoros spp.

Aedes, Culex, Culicoides spp.

Culicoides spp.

Culicoides, Aedes, Culex spp.

Aedes, Culex, Culiseta Anopheles
spp.

Culicoides spp., Aedes aegypti

Not determined

Ixodes, Rhipicephalus spp.

Rhipicephalus spp.

Aedes, Culex, Eretmapodites spp.

Aedes, Culex, Anopheles, Mansonia,
Psorophora spp.

Aedes, Anopheles, Culex, Lutzo-
myia spp.

Aedes, Culex, Culiseta, Anopheles
spp.

The vectors for 2 of these diseases (ASF and NSD) have not been found on
Plum Island. Furthermore, 6 diseases (BLU, EEE, EHD, VEE, VSV and
WEE) can be considered domestic. Therefore, there are only 6 exotic dis-
ease agents of concern on Plum Island, i.e. AHS, AKA, BEF, IBA, LI, and
RVF. Only 6 arboviral disease agents are being used in current research,
i.e. AHS, ASF, BLU and RVF with AKA and IBA being studied minimally.
The remaining 8 arboviruses are presently only in storage. With the excep-

12

NEW YORK ENTOMOLOGICAL SOCIETY

tion of the vectors of ASF and NSD, all other disease vectors are repre-
sented on the island by one or more species of the above mentioned genera.
That is, of course, not to assume that species of mosquitoes, ticks or biting
midges from Plum Island are equally capable of transmitting the diseases as
the natural vectors or those used in experimental transmission studies.
There are species of Aedes, Culex, Culiseta and Anopheles present on Plum
Island which are considered natural vectors of 3 diseases (i.e. EEE, VEE
and WEE, all considered domestic diseases). Vectors of the remaining 9
diseases (AHS, AKA, BEF, BLU, EHD, IBA, LI, RVF, and VSV) have
only nonspecific representatives of the genera implicated as natural or ex-
perimental vectors.

A potential problem exists concerning the inter-island migration of vector
insect species. The absence of adequate breeding areas for Ae. sollicitans
(Walker), Ae. cantator (Coquillett), Culicoides hollensis (Melander and
Brues) and T. nigrovittatus Macquart on Plum Island indicates that these
species are crossing the 2 km of open sea from the closest land mass. Ex-
tensive salt marshes exist at Orient Point, the easternmost tip of Long Is-
land’s north fork and at Gardiner’s Island, ca. 8 km to the south of Plum
Island. Prevailing westerly winds may assist these insects in crossing Plum
Gut.

Despite the simultaneous presence of disease agents and potential vec-
tors, the prevalence of an active arbovirus in the invertebrate population
becomes medically important only when there exists a coincident suscep-
tible host population for virus amplification. With the exception of a few
animals destined for research that are quarantined as they arrive on Plum
Island, all research animals are housed within the high containment labo-
ratory buildings. Other than the two rodent species, the few cats and bats,
no other wild mammals exist on Plum Island. Therefore, the relative absence
of suitable large animal host populations would result in nullifying a potential
epizootic in a mammal population. For example, despite the abundant ta-
banid populations on Plum Island and the ability of certain tabanids to trans-
mit diseases studied at the PIADC (e.g. rinderpest), if only as a mechanical
vector (Bhatia 1935; Tidwell et al. 1972), the absence of a susceptible host
pool renders their presence insignificant. Indeed, as Krinsky (1976) states,
the role that vectors play in virus transmission maintenance is to “increase
the size of epizootics in localized situations in which large numbers of acute-
ly infected animals are in proximity to susceptible hosts.’’ The relative ab-
sence of mammalian hosts in this particular circumstance, coinciding with
an abundant avian population on Plum Island, creates a situation where the
majority of the native mosquito species are feeding on birds. Thus, it ap-
pears that the relative absence of susceptible mammalian hosts and the
abundance of avian hosts has resulted in influencing the abundance of po-
tential vectors within certain genera (cf. Aedes, Culiseta). Therefore, one
would expect that if Plum Island were susceptible to an arbovirus epizootic,

VOLUME LXXXIX, NUMBER 1

13

it would have to be through an avian route. This would be true for both
native arboviruses and arboviruses used in research.

Of the arboviral agents used for research at the PIADC that can be passed
through an avian rounte, EEE and WEE (eastern US isolates of WEE are
now referred to as Highlands J virus), are generally seen annually in arbo-
virus surveys conducted along the east coast (Altman et al. 1967; Bast et
al. 1973; Lord and Calisher 1970; Morris et ah 1973; Morris et al. 1975;
Srihongse et al. 1978). Therefore, should EEE or HJ be isolated from res-
ident or migrating birds of Plum Island, obviously it cannot be assumed that
there had been a break in containment. What could be of concern is the fact
that the extensive shore bird population on the island could potentially play
a role in amplifying native virus (along the flyway) and supporting an avian
arbovirus epizootic. In such a case, the proliferation of a native arbovirus
is much more likely than a breakdown in containment of exotic disease
agents.

None of the passerines sampled in this survey possessed detectable ar-
bovirus antibodies (EEE, HJ or SLE) at a time when adjacent states re-
ported numerous mosquito isolations (W. Crans, Rutgers University, New
Brunswick, NJ and H. Maxfield, Massachusetts Department of Public
Health, Middleboro, MA personal communications). Attempts to detect an-
tibodies in both the mammal and avian specimens to diseases studied on
Plum Island (except EEE and WEE) were not performed during this survey.
A comprehensive attempt should be made to routinely monitor sentinel
pheasants and rabbits for antibodies to arboviruses used in research at the
PIADC. The proliferation of EEE or HJ viruses through the mosquito or
avian cycles should not cause concern for the research facility since this
can occur anywhere along the east coast of the United States where vector
mosquito species and potential avian host populations overlap. What could
be of concern to the facility is the possible amplification of a disease organ-
ism used in research in the prolific native avian host pool with the bird-
feeding mosquitoes ( Culex , Culiseta spp., etc.) acting as vectors.

In order to circumvent such an occurrence, recommendations for chem-
ical, biological and mechanical control have been made to the PIADC staff.
Incorporation of certain of these suggestions may help to alleviate potential
problems. The unique proximity of foreign disease agents to native host and
vector species justifiably causes concern. However, should a concerted vec-
tor control program be established on Plum Island in conjunction with reg-
ular screening of sentinel species for antibodies to diseases used in research,
the chances for an epizootic become even more remote.

Acknowledgments

The authors would like to extend their sincere appreciation to Drs. Jerry
Callis, Jerry Walker and Jonathan Richmond and the staff of the PIADC for

14

NEW YORK ENTOMOLOGICAL SOCIETY

their gracious cooperation and permission for access to the entire island.
We would like to offer our deep gratitude to Dr. John Edman of the Uni-
versity of Massachusetts at Amherst for examining the engorged mosquitoes
for blood meal identification. We are also grateful to Dr. Margaret Grayson
and Ms. Judy Wethers of the New York State Department of Health, Di-
vision of Laboratories and Research for conducting the arbovirus isolation
and serological analyses of the Plum Island specimens.

Literature Cited

Altman, R., M. Goldfield and O. Sussman. 1967. The impact of vector-borne viral diseases
in the Middle Atlantic States. Med. Clin. North Am. 51:661-671.

Anonymous. Plum Island Animal Disease Center. Research on foreign diseases of animals.
1975. USDA Agricultural Research Service, U.S. Government Printing Office, Wash-
ington, D.C.

Bast, T. F., E. Whitney and J. L. Benach. 1973. Considerations on the ecology of several
arboviruses in eastern Long Island. Am. J. Trop. Med. Hyg. 22:109-115.

Berge, T. O., Ed. 1975. International Catalogue of arboviruses including certain other viruses
of vertebrates. DHEW Pub. No. (CDC) 75-8301.

Bhatia, H. L. 1935. The role of Tabanus orientis Wlk. and Stomoxys calcitrans Linn, in the
mechanical transmission of rinderpest. Indian J. Vet. Sci. Anim. Husb. 5:2-22.
Burgdorfer, W. 1970. Hemolymph test. A technique for detection of rickettsiae in ticks. Am.

J. Trop. Med. Hyg. 19:1010-1014.

Clark, D. H. and J. Casals. 1958. Techniques for hemagglutination and hemagglutination
inhibition with arthropod-borne viruses. Am. J. Trop. Med. Hyg. 7:561-573.

Crans, W. J. 1964. Continued host preference studies with New Jersey mosquitoes, 1963.

Proceedings of 51st Ann. Mtg. N.J. Mosq. Extermin. Assoc. Pp. 50-58.

Downe, A. E. R. 1962. Some aspects of host selection by Mansonia perturbans (Walk.)
(Diptera: Culicidae). Can. J. Zool. 40:725-732.

Edman, J. D. 1971. Host feeding patterns of Florida mosquitoes. I. Aedes, Anopheles, Co-
quillettidia, Mansonia and Psorophora. J. Med. Entomol. 8:687-695.

, L. A. Walker and H. W. Kale. II. 1972. Host-feeding patterns of Florida mosquitoes, i

II. Culiseta. J. Med. Entomol. 9:429-434.

Krinsky, W. L. 1976. Animal disease agents transmitted by horse flies and deer flies (Diptera:
Tabanidae). J. Med. Entomol. 13:225-275.

Lord, R. D. and C. L. Calisher. 1970. Further evidence of southward transport of arboviruses
by migratory birds. Am. J. Epidemiol. 92:73-78.

Morris, C. D., E. Whitney, T. F. Bast and R. Deibel. 1973. An outbreak of eastern equine
encephalomyelitis in upstate New York during 1971. Am. J. Trop. Med. Hyg. 22:561-
566.

, A. R. Caines, J. P. Woodall and T. F. Bast. 1975. Eastern equine encephalomyelitis

in upstate New York, 1972-1974. Am. J. Trop. Med. Hyg. 24:986-991.

Srihongse, S., M. A. Grayson, C. D. Morris, R. Deibel and C. S. Duncan. 1978. Eastern
equine encephalomyelitis in upstate New York: Studies of a 1976 epizootic by a modified
serologic technique, hemagglutination reduction, for rapid detection of virus infections.
Am. J. Trop. Med. Hyg. 27:1240-1245.

Tidwell, M. A., W. D. Dean, M. A. Tidwell, G. P. Coombs, D. W. Anderson, W. O. Cowart
and R. C. Axtell. 1972. Transmission of hog cholera virus by horseflies (Tabanidae:
Diptera). Amer. J. Vet. Res. 33:615-622.

VOLUME LXXXIX, NUMBER 1

15

White, D. J. and C. P. White. 1979. Aedes atropalpus (Coquillett) breeding in artificial con-
tainers in Suffolk County, New York. Mosq. News. 40:106-107.

New York State Department of Health, Saranac Lake District Office,
Saranac Lake, New York 12983.

Received for publication June 23, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(l), 1981, pp. 16-22

CONTROL OF DERMACENTOR VARIABILIS.1 1. LARVAL
AND ADULT SUSCEPTIBILITY TO
SELECTED INSECTICIDES

Dennis J. White and Jorge L. Benach

Abstract. — The relative susceptibility of the American dog tick, Derma-
centor variabilis, to selected insecticides was determined in the laboratory
by larval exposure to insecticide-treated surfaces and by adult exposure to
topically applied insecticides. Subsequent field experiments were conducted
with caged ticks to determine the potential of these insecticides for use in
area control of D. variabilis adults. Results from both field and laboratory
experiments indicate that emulsifiable formulations of naled and chlorpyri-
fos were highly effective acaricides, more toxic to larval and adult ticks
than acephate, propoxur, pyrethrins and ronnel, as demonstrated by LC50
and LC90 values.

Due to the prevalence of the American dog tick, Dermacentor variabilis
(Say), on Long Island, New York (Anastos 1947; Collins et al. 1949; Good
1973), and the recent increase of Rocky Mountain Spotted Fever (RMSF)
cases on Long Island (Benach et al. 1977), we wished to determine the
relative susceptibility of larval and adult D. variabilis to certain insecticides
under laboratory conditions and to determine the efficacy of field appli-
cations of these insecticides for the control of D. variabilis (White et al.
1980a, b).

Local environmental legislation and court actions precluded the use of
some insecticides and acaricides already labeled for area tick control. Thus,
our aim in studying the susceptibility of D. variabilis adults and larvae was
to attempt to increase the choice of insecticides available for area control.
Field applications of nonlabeled chemicals for D. variabilis control were
made possible by experimental use permits from the New York State De-
partment of Environmental Conservation, as per Part 172 of the Federal
Insecticide, Fungicide and Rodenticide Act of 1972.

This is the first of 3 reports aimed at the control of questing D. variabilis
in areas where attachment to humans and domestic animals is most likely
to occur. In this paper, we report the results of laboratory and field efficacy
trials of 6 insecticides against D. variabilis.

1 Acari: Ixodidae.

VOLUME LXXXIX, NUMBER 1

17

Materials and Methods

Larval bioassays. — Larvae utilized for bioassay experiments were reared
in the laboratory from eggs obtained from field-collected engorged D. var-
iabilis females. Larvae were kept in moist cotton-stoppered plastic tubes
within desiccator jars containing a saturated solution of NH4C1. Adult ticks
were collected for bioassay experiments by dragging a piece of white flannel
cloth (1 m2) along vegetation adjacent to roadsides and paths in various
areas of Long Island. Captured ticks were placed in moist cotton-stoppered
tubes, brought to the laboratory, and maintained in the same manner as
larvae.

Larval exposure to each chemical dilution was accomplished by slightly
modifying techniques reported by Hansens (1956). Filter paper (12.9 cm2)
was used to line the insides of 3.7 ml screw-cap vials. A uniform amount
(0.15 ml) of each insecticide dilution was added directly to the filter paper
in each vial. Control vials received 0.15 ml of distilled water. Ten 8- to 10-
week-old unfed larvae were placed with needle-point forceps directly on the
moist filter paper in each of the 12 vials. Two replicates of 5 chemical
concentrations and of controls represented a test series for the laboratory
bioassays.

The treated surface experiments involved emulsifiable concentrate for-
mulations of chlorphyrifos (0,0-diethyl 0-(3,5,6-trichloro-2-pyridl) phos-
phorothioate) from 36 mg to 1.5 mg a.i./vial and ronnel (0,0-dimethyl
0-2,4,5-trichlorophenyl phosphorothioate) from 108 mg to 10.8 /xg a. i. /vial,
courtesy of Dow Chemical Co.; propoxur (O-isopropoxyphenyl methylcar-
bamate) from 20 mg to 20 ng a.i./vial, courtesy Chemagro naled (1,2-dibromo-
2,2-dichloroethyl demethyl phosphate) from 840 mg to 840 ng a. i. /vial, cour-
tesy Chevron Chemical Co., and pyrethrins with 10% piperonyl butoxide
from 1.44 mg to 144 ng a. i. /vial, courtesy FMC. In addition to the above, a
water soluble powder formulation of acephate (0,5- dimethyl acetylphos-
phoramidothioate), courtesy Chevron Chemical Co., was used in the topical
application experiments. The same chemicals were used for field experi-
ments.

Adult bioassays. — Topical applications of various dilutions of the insec-
ticide emulsions proved to be the most efficient method of adult exposure
to the chemical in the laboratory. Filter paper (45.1 cm2) was used to line
the inside of 29.6 ml screw-cap vials. The paper in each vial was moistened
with 0.5 ml distilled water to provide a source of humidity. One microliter
of a chemical dilution was placed on the dorsum of each of 20 ticks (of both
sexes) to be treated with that dilution. Individual ticks were restrained by
forceps while the chemical was applied and held for 10 s before they were
placed into the vials. Ten treated adult ticks were contained in each vial. A
regimen of 2 replicates of each of the 5 chemical dilutions and controls

18

NEW YORK ENTOMOLOGICAL SOCIETY

represented a test series for the adults. Concentrations of the applied chem-
icals ranged from 40,000-3,000 ppm for acephate, 3,600-1.5 ppm for chlor-
pyrifos, 43,500-0.4 ppm for naled, 8,200-0.82 ppm for propoxur, 50,000-
500 ppm for pyrethrins with piperonyl butoxide and 7,500-75 ppm for ron-
nel.

Mortality, defined as the failure of a tick to respond to being breathed
upon (Mount et al. 1970), was determined in all vials 24 and 48 h after initial
exposure. Since ticks are negatively geotropic, all vials were maintained in
a horizontal position at 24°C and a 10: 14 h lightrdark photoperiod throughout
the 48 h test period.

Field tests. — During field experiments, postspray dragging proved to be
a variable and inefficient tool (White et al. 1980b) for monitoring insecticide
spray efficiency. Therefore, caged ticks were utilized to determine the ef-
ficacy of the insecticide applied under field conditions. The plots used for
the field trials were open, ca. rectangular fields, ranging in size from 0.4 to
1.6 ha and dominated by 10 to 25 cm high grass vegetation. Ten adult ticks
were placed inside each of 12 cylindrical aluminum wire screen cages (ca.
85 cm3, 4 cm diam x 9 cm long, mesh opening 2.25 mm2). Cages were
supported by stakes 15 cm above the ground (to approximate the position
of questing ticks) in rows parallel to the line of travel of the sprayer. A
Buffalo Turbine mist blower, model CS, was operated at 14.1 kg/cm2 (200
psi) with four 0.64-cm (lA") Tee Jet nozzles (i.e. 2, lA TT 6504; 1, lA TT
6503; and 1, 14 TT 6502) and calibrated to deliver 28.2 1/ha (3 gal/acre). The
vehicle upon which the blower was mounted had a forward speed of 8 km/
h (5 mph), during spray activities. Sprays were delivered downwind when
wind speed was no greater than 8 km/h. Immediately after treatment, ticks
were transferred to moist cotton-stoppered plastic tubes. Mortality was then
determined 24 h after chemical exposure.

Probit analyses (Finney 1964) were conducted on laboratory bioassay
mortality data taken at 24 and 48 h.

Results and Discussion

The mean LC50 and LC90 values, as well as the standard error of the
mean, determined at 24 and 48 h after exposure of the tick larvae to the
corresponding insecticide, are shown in Table 1. Results of the larval bioas-
says indicate that, based on LC50 and LC90 values, naled was the most toxic
of the 5 insecticides to D. variabilis larvae. According to the 48 h LC90
values, naled is from 32 x to 554 x more toxic to D. variabilis than the
remaining compounds. The 48 h LC90 values of ronnel and synergized py-
rethrins indicate that a substantially higher dose was necessary to result in
similar morality figures.

Data derived from the topical application experiments on adults are shown

VOLUME LXXXIX, NUMBER 1

19

Table 1. Mean LC50 and LC90 values (ppm) with standard error (SE) of the mean, deter-
mined at 24 and 48 h after exposure of D. variabilis larvae to surfaces treated with various
concentrations of 5 insecticides.

Chemical

lc50

lc90

Time

Mean

SE

Time

Mean

SE

Chlorpyrifos

24

2.39

1.15

24

1.99

1.32

48

0.21

—

48

1.29

—

Naled

24

0.02

0.01

24

0.09

0.05

48

0.01

0.006

48

0.04

0.01

Propoxur

24

0.64

—

24

3.48

—

48

0.58

48

2.87

—

Pyrethrins

24

2.64

0. If)

24

44.44

3.79

48

0.58

0.41

48

22.14

8.76

Ronnel

24

24.70

11.60

24

60.39

10.09

48

2.97

1.25

48

13.36

2.93

in Table 2. The mean LC50 and LC90 values for naled and chlorpyrifos are
very similar, indicating that relatively low concentrations of these materials
were necessary for effective control. Synergized pyrethrins was the third
most active material against topically treated adults followed by ronnel,
propoxur and acephate in order of decreasing toxicity. As with the larval
experiments, acephate required a very high concentration to produce mor-
tality in D. variabilis adults.

Table 2. Mean LC50 and LC90 values (ppm) with standard error (SE) of the mean, as
determined by probit analysis at 24 and 48 h after exposure of D. variabilis adults to various
concentrations of 6 topically applied insecticides.

Chemical

lc50

lc90

Time

Mean

SE

Time

Mean

SE

Acephate

24

4,941.83

356.77

24

28,248.99

3,775.48

48

2,185.00

1,441.89

48

9,745.27

448.71

Chlorpyrifos

24

7.75

0.50

24

29.84

3.65

48

6.32

1.16

48

19.08

5.92

Naled

24

8.76

0.64

24

57.50

20.04

48

7.26

1.04

48

27.30

6.88

Propoxur

24

182.39

15.57

24

673.72

42.52

48

166.18

14.29

48

574.78

61.47

Pyrethrins

24

26.60

6.28

24

106.24

21.94

48

25.32

7.56

48

97.13

12.83

Ronnel

24

185.05

35.16

24

519.87

212.73

48

158.41

26.34

48

418.09

30.38

20

NEW YORK ENTOMOLOGICAL SOCIETY

Table 3. Percentage mean mortality at 24 h of caged adult ticks placed at various distances
from the insecticide sprayer and treated with a specific concentration of 1 of 6 insecticides.

Distance
(m) from
sprayer

Chemical and concentration (actual insecticide)

Acephate

Chlorpyrifos

Naled

Propoxur

Pyrethrins

Ronnel

6%

3%

0.12%

0.06%

0.36%

0.18%

0.09%

0.22%

0.11%

0.05%

0.01%

0.25%

0.12%

6

100

100

100

100

100

100

100

100

100

100

100

100

100

12

100

75

100

100

100

100

100

100

100

100

80

100

90

18

80

70

100

100

100

100

100

100

90

95

50

100

90

24

75

70

90

100

100

100

100

100

70

100

20

100

65

30

45

55

100

85

100

40

95

100

60

80

10

100

40

Control

10

5

10

5

5

10

5

0

10

0

10

5

5

As illustrated in Table 3, these same emulsifiable chemicals (acephate was
formulated as WP) were used to determine relative efficacy for control of
caged D. variabilis under field conditions. The concentrations were selected
arbitrarily based on information already on the label for use under area wide
outdoor application and either diluted by Vi or doubled for additional data
against the American dog tick. The higher concentrations of each insecticide
produced 100% mortality in those caged ticks nearest the point of applica-
tion in all cases. All concentrations of naled and chlorphyrifos were shown
to be very active even when dispersed up to 24 m from the point of appli-
cation. High concentrations of ronnel and propoxur were effective up to 30
m from the application point, but these concentrations may not be accept-
able for area wide outdoor application. Similarly acephate required exceed-
ingly high concentrations to produce mortality. Synergized pyrethrins at
0.05% may be effective for the control of D. variabilis but potential appli-
cators should consider the cost/benefit ratios of all materials.

In addition to assessing the utility of the mist blower, the data in Table
3 indicate that both naled and chlorpyrifos are highly active for D. variabilis .
Choice of material to be used in a control program will depend on availability
of the individual materials, registration of the materials in the particular
geographic area and chemical cost. These parameters aside, three com-
pounds used in this study, chlorpyrifos, naled, and synergized pyre-
thrins, can be applied at acceptably low concentrations. The mortality fig-
ures produced by propoxur and ronnel show that they require slightly higher
concentrations, but if the materials are available and are registered for area
use, they may be effective agents for the control of D. variabilis. The con-
centrations required for acephate for effective applications were very high,
and as such, does not provide a justification for using acephate against the
American dog tick.

Our findings support those of Drummond et al. (1971) who also stated

VOLUME LXXXIX, NUMBER 1

21

that ronnel was less effective than chlorpyrifos for control of D. albopictus
Packard. Mount et al. (1970) showed that naled was ca. 10 times as toxic
as chlorpyrifos to 1- to 2-month-old Amblyomma americanum (L.) nymphs.
However, Drummond and Medley (1965) reported that a 0.25% naled for-
mulation was less effective than a 0.75% ronnel in controlling adult A.
americanum on cattle. Rawlins and Mansingh (1977) indicated that naled
exhibited acaricidal effects on 3 Jamaican species of livestock ticks.

The results of laboratory and field insecticide applications have shown
that naled and chlorpyrifos are toxic to D. variabilis and, when applied
under field conditions, will kill them even at concentrations of 0.09% and
0.06%, respectively.

The field caged tick study, as well as the laboratory toxicity study, strong-
ly suggest that naled and chlorpyrifos, when applied by high-pressure, low-
volume equipment, can provide control of American dog tick questing pop-
ulations. Since ticks congregate along paths or roadsides (Smith et al. 1946),
an effective and economical control program can be instituted with the ob-
jective of controlling populations of D. variabilis in these areas. If insecti-
cide applications are made to coincide with peak tick populations (Good
1973; Smith et al. 1946) in areas of endemic RMSF activity, the objective
of reducing the probability of human exposure to infected ticks can be re-
alized (White et al. 1980a). Control efforts directed against questing ticks in
infected foci could contribute to the decrease of human RMSF cases in
given areas.

Literature Cited

Anastos, G. 1947. Elosts of certain New York ticks. Psyche 54:178-180.

Benach, J. L., D. J. White, W. Burgdorfer, T. Keelan, S. Guirgis and R. Altieri. 1977. Chang-
ing patterns in the incidence of Rocky Mountain Spotted Fever on Long Island (1971-
1976). Am. J. Epidemiol. 106:380-387.

Collins, D. L., R. V. Nardy and R. D. Glasgow. 1949. Some host relationships of Long Island
ticks. J. Econ. Entomol. 42:110-112.

Drummond, R. O., W. J. Gladney, T. M. Whetstone and S. E. Ernst. 1971. Laboratory testing
of insecticides for the control of the winter tick. J. Econ. Entomol. 64:686-688.

and J. G. Medley. 1965. Field tests with insecticides for the control of ticks on live-
stock. J. Econ. Entomol. 58:1131-1136.

Finney, D. J. 1964. Probit analysis — a statistical treatment of the sigmoid response curve. 2nd
ed. Cambridge at the University Press. 318 pp.

Good, N. E. 1973. Ticks of eastern Long Island: notes on host relations and seasonal distri-
bution. Ann. Entomol. Soc. Am. 66:240-243.

Elansens, E. J. 1956. Chlordane resistant brown dog ticks and their control. J. Econ. Entomol.
49:281-283.

Mount, G. A., C. C. Lofgren and N. W. Pierce. 1970. Effect of age on the susceptibility of
nymphs of the lone star tick to insecticides. J. Econ. Entomol. 63:1681-1682.

Rawlins, S. C. and A. Mansingh. 1977. Relative susceptibility of Jamaican strains of Boophilus
microplus, Amblyomma cajennense and Anocentor nitens to various acaricides. J. Econ.
Entomol. 70:697-698.

22

NEW YORK ENTOMOLOGICAL SOCIETY

Smith, C. N., M. M. Cole and H. K. Gouck. 1946. Biology and control of the American dog
tick. U.S. Dep. Agric. Tech. Bull. 905. 74 pp.

White, D. J., J. L. Benach, L. A. Smith and S. P. Ouyang. 1980a. Control of Dermacentor
variabilis. 3. An analytical study of the effect of low volume spray frequency on insec-
ticide-stressed and nonstressed populations. J. N.Y. Entomol. Soc. 89:23-33.

, L. A. Smith and J. L. Benach. 1980b. Control of Dermacentor variabilis. 2. Analysis

of a mark-recapture study of a seasonal questing population. Submitted.

New York State Department of Health, Saranac Lake, New York 12983
and New York State Department of Health, State University of New York
at Stony Brook, Stony Brook, New York 11794.

Received for publication June 23, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(l), 1981, pp. 23-33

CONTROL OF DERMACENTOR VARIABILIS.1 3. AN
ANALYTICAL STUDY OF THE EFFECT OF LOW VOLUME
SPRAY FREQUENCY ON INSECTICIDE-STRESSED
AND NONSTRESSED POPULATIONS

Dennis J. White, Jorge L. Benach, Laurel A. Smith and Soo P. Ouyang

Abstract. — Nine field plots (each 0.4-0. 6 ha) were selected along the
south central part of Long Island, New York, to study the effect of insec-
ticide spray frequency on a questing Dermacentor variabilis population.
Assignment of treatment was made according to dominant vegetation, lo-
cation and tick preferential habitat. Three plots received no treatment, 3
plots were sprayed with water (3 once, 2 twice, and one 3 times) and 3 plots
were similarly sprayed with an emulsifiable formulation of naled. Plots were
randomly monitored for tick density at weekly intervals. A total of 2,872
ticks was collected from all plots by dragging. Multiple and simple regres-
sion and exploratory data analyses were employed to determine normal or
estimated tick populations at each plot throughout the adult tick season,
rate of seasonal weekly decline of tick population, and the effect of insec-
ticide spray frequency on the questing American dog tick population. Re-
sults indicated a 20% weekly exponential decay in numbers of questing adult
ticks over the 1978 season. No effect was seen due to the water treatment.
One spray of naled produced an 82% reduction of the questing tick popu-
lation; 2 sprays, a 95% reduction; and 3 sprays, a 96% population reduction.

Investigations on the effect of insecticides or acaricides on tick popula-
tions in the field have been reported (Hoch et al. 1971; Mount et al. 1968,
1971; Smith et al. 1946; Wharton et al. 1976). Yet, little work has been done
to delineate the effect produced on seasonal populations of questing Amer-
ican dog ticks by a series of well timed insecticide applications. The elab-
orate work reported by Smith et al. (1946) is the classic description of the
biology of Dermacentor variabilis and has provided a foundation for further
research into bionomics and improved control techniques.

Extensive tick control has been most commonly practiced in areas where
ticks affect livestock (Drummond 1977; Teel et al. 1977; Wharton 1976).
Area control of tick species affecting humans has become of recent interest
to public health agencies along the east coast of the United States due to

Acari: Ixodidae.

24

NEW YORK ENTOMOLOGICAL SOCIETY

the increase of reported Rocky Mountain spotted fever cases (Benach et al.
1977; D’Angelo et al. 1978; Hattwick 1971; Loving et al. 1978).

Preliminary laboratory and field experimentation (White and Benach 1980)
indicated the susceptibility of D. variabilis to naled (l,2-dibromo-2,2-di-
chloroethyl dimethyl phosphate), a nonpersistent insecticide. The devel-
opment of fast-acting non-residual insecticides has benefitted public health
agencies concerned with vector control. The advantages derived from the
use of nonpersistent chemicals can be enhanced by a schedule of chemical
applications during a particular tick season (Clymer et al. 1970).

Because control of the American dog tick is directed to the questing
population, a certain percentage of ticks may be protected from the spray
technique, e.g. in the soil, on hosts, or at the bases of vegetation. The
population monitoring technique most often used for D. variabilis (i.e.,
dragging) measures only the questing population. The effect of chemical
sprays may not be felt by the majority of ticks in a field or plot at any
particular time. Dragging has been shown to be an inefficient (White et al.
1980) tool for monitoring actual field populations, and actual estimates of
field populations have to be extracted from a combination of prior seasonal
population data, abiotic factors affecting a particular plot (weather, vehicle
traffic, etc.), potential favorable habitat, host availability and residents’
complaints. A series of chemical applications can be made to compensate
for the inherent weaknesses of field tick population estimates. Ticks that
happen to be at the base of some vegetation may successfully escape contact
with a chemical application. However, if these and other ticks are exposed
at the time of subsequent applications, chances are greater that most ticks
on the plot will, at some time, be subjected to contact with the acaricide.

A series of well timed chemical applications could lead to an effective D.
variabilis control program. The present paper discusses the effect of 3 mist
applications of naled at 4-week intervals on the questing population of D.
variabilis over a 3-month season.

Materials and Methods

During the early spring of 1978, nine field plots (0.4-0. 6 ha) were selected
along the south shore of central Long Island, New York. Plot selection was
based on dominant vegetation, location, and likelihood of supporting large
numbers of D. variabilis. Three plots were chosen for their grass-dominant
habitat, 3 for their grass-forb mixture habitat and 3 for their grass-forb-
deciduous combination habitat. All plots were grouped into triplets or blocks
that were similar in composition for the 3 parameters of vegetation, location
and tick preferential habitat (Smith et al. 1946; Sonenshine et al. 1972).
Treatment of each of the 9 plots was allocated a random, subject to the
following constraints: (1) Three plots received no treatment (Cl, C2, C3);

VOLUME LXXXIX, NUMBER 1

25

(2) Three plots received water sprays (as a control for the application pro-
cedure), 3 plots sprayed once (Wl), 2 plots sprayed twice (W2) and 1 plot
sprayed 3 times (W3) during the season; (3) Three plots received naled
sprays, 3 plots sprayed once (Dl), 2 plots sprayed twice (D2) and 1 plot
sprayed 3 times (D3) through the adult tick season. Assignment of spray
frequency among the 3 water-sprayed plots and 3 naled-sprayed plots was
made at random. (See Table 1.)

A 24 (c) registration was received by Chevron Chemical Co. for the use
of Dibrom 8 (naled) to control the American dog tick (EPA SLN No.
780008). Treatment for the Dl, D2 and D3 plots consisted of application of
168 g actual Dibrom per ha (0.15 lbs/A). The naled formulation (1 ,2-dibromo-
2,2-dichloroethyl dimethyl phosphate) was supplied courtesy of Chevron
Chemical Co. Both water and chemical applications were delivered as pre-
viously described (White and Benach 1980). Plots to be sprayed once were
sprayed during wk 1; plots sprayed twice were treated during wk 1 and 5;
the plot sprayed 3 times was treated during wk 1, 5 and 9.

Weekly tick collections were made at each plot. Ticks collected at plot
C3 were returned to the laboratory. On those plots that were treated during
a specific week, collections for that week were made 24 h after the spray.
White flannel cloth (1 m2) was used to collect questing ticks by dragging.
Ticks attached to the flannel drag were counted every 10 m along a prede-
termined line of travel at the roadside edge of each plot (60-100 m).

All 9 plots could not be monitored for tick population on 1 day. Therefore,
the group of plots was divided into 3 sets of 3 plots, each of which would
be randomly monitored on 3 successive days of the week. In order to control
for the effect of measuring the plots at different times of day and on different
days of the week, predetermined random schedules were made resulting in
72 combinations of tick collection sequences in control, water-sprayed and
naled-sprayed plots.

Statistical analyses were conducted using a randomized block analysis of
variance model where blocks were similar for plot location, dominant vege-
tation and tick density. For analysis purposes, actual tick counts in each
plot were normalized and transformed to base 10 logarithms. Analysis of
data was conducted by 3 separate techniques; exploratory data analysis
(Tukey 1977), multiple, and simple regression utilizing the analysis of vari-
ance model.

Actual field questing tick population data (Table 2) obtained by dragging
were subjected to the following transformation:

Yijk = (Xijk/Nij) x 100

Where i = indicator for treatments, i.e., i = 1 for control groups, i = 2 for
water-treatment groups, and i = 3 for naled-treatment groups; j = the indi-
cator for blocks, i.e., j = 1 for block 1, j — 2 for block 2, etc; and k is the

26

NEW YORK ENTOMOLOGICAL SOCIETY

Table 1. Vegetational composition and number of Dermacentor variabilis collected at each
of the 9 study plots over the entire season.

Plot

Vegetation*

No. ticks collected

Total no./treatment

C-l

g-f

261

C-2

g

316

C-3

g-f-d

526

1,103

W-l

g

1 1

W-2

g-f

1,034

W-3

g-f-d

80

1,125

D-l

g-f

65

D-2

g

185

D-3

g-f-d

394

644

total

2,872

* g = grass, f = forb, d = deciduous.

index for week sequence. Therefore, Xijk is the original total count of ticks
in treatment i, block j and week k; Ny is the number of intervals in block
j, treatment i from which ticks were collected and Yijk is the transformed
total count of ticks in treatment i, block j and week k.

Then, in order to stabilize the week-to-week variability within plots, the
transformed data were normalized using a log transformation:

ziik = log10(Yuk + 1).

In view of the fact that each plot had varied original (pre-experimental)
questing tick populations due to the dominant vegetation, and the plots were
minitored for tick counts on a random basis over a 3-day period, a median
polish process was utilized to produce a range of values about a 0.0 residual
value (expected questing tick counts at a given point in time, Tukey 1977).

Because both Cl and W1 plots supported very few ticks, the data gathered
during the same season from plot MR (White et al. 1980) replaced the data
from Cl. Data from plot W1 were dropped completely so that all analyses
of effect of water were conducted on data from 2 plots, W2 and W3.

Results and Discussion

Dominant vegetational characteristics of each of the 9 plots and numbers
of ticks collected in each plot throughout the 12- week experiment are listed
in Table 1. All captured ticks (other than C3) were released in the 10 m
section of each plot from which they were collected.

Exploratory data analysis. — By analyzing the transformed data for effect
of week on the questing tick population (i.e., numbers of ticks over time),

VOLUME LXXXIX, NUMBER 1

27

Table 2. Actual numbers of questing Dermacentor variabilis collected from each plot during
the 12- week study.

Week

Plot

MR*

Cl

C2

C3

W1

W2

W3

D1

D2

D3

1

49

4

102

56

2

69

15

3

12

35

2

33

4

46

60

6

107

21

7

38

128

3

28

0

45

145

1

98

14

7

27

120

4

20

0

1

50

1

101

6

6

41

62

5

23

4

47

114

1

109

4

6

1

2

6

25

2

23

36

0

141

4

10

3

12

7

10

0

13

21

0

145

3

2

13

11

8

16

0

17

12

0

76

3

7

21

18

9

25

0

7

20

0

84

3

5

15

1

10

15

0

4

5

0

53

4

2

5

1

11

4

0

4

4

0

33

3

5

3

1

12

13

0

7

3

0

18

0

5

6

3

* MR = mark-recapture (White et al. 1980).

the fitted equation produced by exploratory data analysis is Y = 0.5549 -
0.0913X which indicates an exponential decay rate of 19% (10 0 0913 - 1 =
0.8104. 1 - 0.8104 = 0.1896 = 19%) on the original scale (actual field col-
lection data) which explains 88% of the variability of week effects (F =
74.312, d.f. = 11, 0.01, R2 = 0.88). By holding week and plot effect

fixed, a noticeable drop of tick populations is apparent in plots D2 and D3
after treatment while control plots and water-treatment plots show no such
relationship. The residual value indicates the transformed tick count that,
based on data gathered from the control plots, would be expected to be
present during each week. Values in excess of this threshold value indicate
a population of ticks larger than that which would be expected; negative
values indicate populations less than that which would be expected. Also
evident is a resurgence of tick populations in the D block of plots shortly
after the spray until the next chemical treatment.

Multiple regression. — By employing a more definitive analysis in a mul-
tiple regression process (F = 3.369, 75 df, P = 0.00, R2 = 0.4733), further
significant relationships can be shown to exist. Although the model is ex-
tremely complex and encompasses effects of individual treatments in each
plot, the overall week effect and treatment effect are not easily distinguish-
able. The same transformed and normalized data were also used for this
approach. The effect of week on tick populations is similar to that which
resulted from exploratory data analysis with a 21% exponential decay rate
of actual tick numbers (F = 96.48, 11 df, a = 0.01, P < 0.001, R2 = 0.906).
However, a rebound (due to recruitment) of the tick population occurs dur-

NEW YORK ENTOMOLOGICAL SOCIETY

28

3.4 r
3.2 -

O

cr>

o

3.0 -
2.8 -
2.6 -
2.4 -
2.2 -

2.0 -
1 .8 -
16 -
1.4-
1 .2 -
1.0 -

• MR
x C2

o

X

o

J I I 1_

5 6 7 8

WEEK

_i i_

10 11

12

Fig. 1. Simple regression estimation of week effect on questing Dermacentor variabilis
population transformed data.

ing weeks 5 and 8. The effect seen on the field tick populations by the water
treatments was variable and probably due to the week effect included in
this analysis. However, an 82% decrease of the questing tick population
was seen after 1 application of naled on all 3 plots. On the two plots, D2
and D3, that were sprayed twice (during weeks 1 and 5), a reduction of 96%
occurred. Likewise, an 87% reduction occurred on the plot that was sprayed
for the 3rd time during week 9.

Simple regression. — By using a simple regression analysis the results be-
come more clarified because each effect (week or treatment) is removed
individually. For example, through use of the control groups to estimate the
week effect on tick populations, the week effect can be extracted from the
water-treatment group and the naled-treatment group to determine more
precisely the effect of such treatment. Therefore, because the week effect
has now been determined without the introduction of insecticide or me-
chanical stress on the population used for analysis, the actual results are
more practical. Now the equation fitted for week effect becomes:

Zijk = 2.9252 - 0.0976X (See Fig. 1)

F = 33.2983. 35 df, a = 0.05, P < 0.01

VOLUME LXXXIX, NUMBER 1

29

WEEK

Fig. 2. Effect of water treatment on questing D. variabilis populations.

or, that the week effect on the tick population is exponential decay with a
rate of 20% (lO"0 0976 - 1 = 0.7987. 1 - 0.7987 = 0.2013 = 20%).

By extracting the week effect from the data gathered from the water
treatment analysis, the effect of water on the tick population is nonexistent
(Fig. 2). By extracting the seasonal tick reduction figures (week effects)
from the data gathered from the naled treatment analysis, the effect on the
questing tick population is pronounced (Fig. 3). Obviously, the effect of the
1st naled spray cannot be determined by simple regression because exten-
sive tick sampling was not conducted prior to week 1 . The 2nd naled spray
reduced the population by 95% while the 3rd spray resulted in a 96% tick
population reduction.

The preliminary laboratory and field experiments performed by White and
Benach (1980) led to the conclusion that naled could prove effective in a
large scale control program directed against D. variabilis. However, be-
cause naled is nonpersistent, a control program would have to rely on a
series of well timed chemical applications over an entire season. For ex-
ample, although all ticks were removed from plot C3 each week (Table 2),
a large number of ticks were captured the following week, indicating 1 or
both of the following: a large immigration factor or a poor sampling tech-
nique (White et al. 1980). In order to reduce tick numbers over an entire
season, control should be carried out in a series of applications timed so

30

NEW YORK ENTOMOLOGICAL SOCIETY

0.8

A PLOT SPRAYED

< o

X

< -Q

i

o

o

x

D1

Fig. 3. Effect of naled treatment on questing D. variabilis populations.

VOLUME LXXXIX, NUMBER 1

31

that ticks do not reach a threshold value and so that chemicals are applied
cost-effectively.

As can be seen in Table 1, tick densities were greatest in either the grass-
forb or the grass-forb-deciduous mixed habitat than in grass alone. The
greatest proportion of ticks present in the plots was seen in the first 4-7
weeks of the study, again dependent on the habitat. By incorporating all
population data from the control plots (MR, C2 and C3, Table 2) a regres-
sional analysis resulted in the week effect, or effect of time within a season
on the natural or nonstressed tick population. Over the season of study, a
weekly exponential decay rate of 20% explained the general tick population
depletion through mid-August and therefore represented our weekly thresh-
old value.

By removing this week effect from the treatment effect (simple regres-
sion), a more precise description can be made. After each application of
naled, there was an immediate decrease in the tick population followed by
a gradual resurgence over the next 3 weeks. This would be expected because
naled is efficient but nonpersistent. As the tick population increases to the
expected value (0.0, Fig. 3), the effect of the previous naled spray has been
eliminated at that point. An increase beyond the expected value necessitates
a reapplication of naled to further reduce the questing population. The 2nd
naled application resulted in a 95% and 97% reduction on plots D2 and D3
while the tick population continued to increase in Dl.

After the 2nd spray, tick populations again started to increase to the
expected value at a more rapid recovery. Within 2 weeks, the tick popula-
tion recovered to the point where they exceeded the expected values in
both plots D2 and D3. During week 6 and week 9 plot Dl was mowed,
which brought the tick population below what was expected, an uninten-
tional control mechanism. Plot D3 received the 3rd application of naled
during week 9, which resulted in a 96% reduction of questing ticks. The
recovery was now much slower, from week 9 through 12. The effect of the
physical act of treating the ticks along roadside edges with water (W2, W3,
Fig. 2) is negligible.

Although each individual analysis provided both significant statements
and generalizations concerning the experiments, one analysis followed the
other in a logical sequence to determine the most significant statements.
Exploratory data analysis provided the justification for the transformation
(Tukey 1977) of the raw data to stabilize the variances (plot to plot, week
to week, etc.), and for the median polish to account for differernces of pre-
experimental tick populations. This analysis provided an insight into results
that were confirmed by classical regression techniques. The multiple regres-
sion and simple regression resulted in almost identical statements (e.g. 20%

32

NEW YORK ENTOMOLOGICAL SOCIETY

population decay rate). Simple regression assumed variable independence
while the multiple regression did not.

Conclusions derived from all analyses are in agreement. There is an over-
all decline of the tick population during a season of about 20% per week,
within the range reported by White et al. (1980). Spraying water on ticks
had no effect on the population so treated. Low- volume sprays of naled
resulted in dramatic drops of the population (82-97%) followed by gradual
recovery, consistent with the mark-recapture study (White et al. 1980) re-
cruitment phenomenon. Cutting of grass along tick habitats was statistically
shown to reduce the expected questing tick population for a short term.

Acknowledgments

We wish to acknowledge Hilary Summer, Robert Dickeson and A. Lee
Holcomb for their valuable technical assistance.

Literature Cited

Benach, J. L., D. J. White, W. Burgdorfer, T. Keelan, S. Guirgis and R. Altieri. 1977. Chang-
ing patterns in the incidence of Rocky Mountain Spotted Fever on Long Island (1970—
1976). Am. J. Epidemiol. 106:380-387.

Clymer, B. C., D. E. Elowell and J. A. Hair. 1970. Environmental alteration in recreational
areas by mechanical and chemical treatment as a means of Lone-star tick control. J.
Econ. Entomol. 63:504-509.

D’Angelo, L. J., W. G. Winkler and D. J. Bregman. 1978. Rocky Mountain Spotted Fever in
the United States 1975-1978. J. Infect. Dis. 138:273-276.

Hattwick, M. A. W. 1971. Rocky Mountain Spotted Fever in the United States 1920-1970.
J. Infect. Dis. 124:112-114.

Hoch, A. L., R. W. Barker and J. A. Hair. 1971. Further observations on the control of Lone
star ticks (Acarina: Ixodidae) through integrated control procedures. J. Med. Entomol.
8:731-734.

Loving, S. M., A. B. Smith, A. F. DiSalvo and W. Burgdorfer. 1978. Distribution and prev-
alence of spotted fever group rickettsiae in ticks from South Carolina, with an epide-
miological survey of persons bitten by infected ticks. Am. J. Trop. Med. Hyg. 27:1255-
1260.

Mount, G. A., J. G. Williams, C. S. Lofgren and S. A. White. 1968. Insecticides for control
of the Lone star tick tested in the laboratory and as high and ultra-low volume sprays
in wooded areas. J. Econ. Entomol. 61:1005-1007.

, C. S. Lofgren and N. W. Pierce. 1971. Effectiveness of 22 promising insecticides for

control of the lone star tick. J. Econ. Entomol. 64:262-263.

Smith, C. N., M. M. Cole and H. K. Gouck. 1946. Biology and control of the American dog
tick. U.S. Dep. Agric. Tech. Bull. 905. 74 pp.

Sonenshine, D. E., A. H. Peters and G. E. Levy. 1972. Rocky Mountain Spotted Fever in
relation to vegetation in the eastern United States. 1951-1971. Am. J. Epidemiol.
96:59-69.

Teel, D. D., J. A. Hair and T. C. Randolph Jr. 1977. Continuous administration of famphur
for control of ticks and bed bugs feeding on ruminants. J. Econ. Entomol. 70:664-666.

VOLUME LXXXIX, NUMBER 1

33

Tukey, J. W. 1977. Exploratory data analysis. Addison-Welsey Publishing Co., Reading,
Mass. 506 pp.

Wharton, R. H. 1976. Tick-borne livestock diseases and their vectors. Part 5. Acaricide
resistance and alternative methods of tick control. World Animal Rev. 20:8-15.

White, D. J. and J. L. Benach. 1980. Control of Dermacentor variabilis. 1. Larval and adult
susceptibility to selected insecticides. J. N.Y. Entomol. Soc. 89:16-22.

, L. A. Smith and J. L. Benach. 1980. Control of Dermacentor variabilis. 2. Analysis

of a mark-recapture study of a seasonal questing population. Submitted.

(DJW) New York State Department of Health, Saranac Lake, New York
12983; (JLB) New York State Department of Health, State University of
New York at Stony Brook, Stony Brook, New York 11794; and (LAS and
SPO) Department of Applied Mathematics, State University of New York
at Stony Brook, Stony Brook, New York 11794.

Received for publication June 23, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(l), 1981, pp. 34-39

FURTHER EVIDENCE FOR CHLORPYRIFOS TOLERANCE AND
PARTIAL RESISTANCE BY THE JAPANESE BEETLE
(COLEOPTERA: SCARABAEIDAE)

Sami Ahmad and Yuen-Shaung Ng

Abstract. — Dosage-mortality responses of third-instar grubs of the Jap-
anese beetle, Popillia japonica Newman, showed that a population from
Fairfield (Conn.), when compared at the LD95, is ca. 42x as tolerant to
chlorpyrifos as those from Adelphia (central N.J.; a standard susceptible
population). Moreover, the 95% conf. intervals of the LD50 and LD95 of the
Fairfield population did not overlap with those of the Adelphia population.
Another population from Rivervale (northern N.J.) also showed 6.8-fold
greater tolerance to chlorpyrifos, however, the conf. intervals of its LDs
did overlap with those of the susceptible strain. The Fairfield population
was characterized as “partially-resistant,” and that from Rivervale as “tol-
erant,” to chlorpyrifos.

The grubs of the Japanese beetle, Popillia japonica Newman, are highly
destructive to turfgrass in several states east of Mississippi River (Fleming
1972). Chlordane was first used for grub control in 1947 (Fleming 1950), and
since then for nearly twenty-five years chlordane and other cyclodiene in-
secticides, e.g., dieldrin and heptachlor, afforded good protection against
Japanese beetles. Beetle resistance to cyclodienes was first reported in 1973
from Ohio (Niemczyk and Lawrence 1973), New York (Tashiro and Neu-
hauser 1973), and Pennsylvania (Anon. 1973). Subsequently, cyclodiene re-
sistance among beetle populations of Connecticut (Dunbar and Beard 1975),
and New Jersey (Ahmad and Das 1978) also were documented. Further-
more, because of environmental concerns the use of highly persistent cy-
clodiene insecticides has been restricted by the Environmental Protection
Agency. Consquently, nonpersistent organophosphorus insecticides, chlor-
pyrifos, diazinon and trichlorfon have replaced cyclodienes for control of
the Japanese beetle.

In 1977, a population of the Japanese beetle from New Jersey (Rivervale)
was suspected of increased tolerance to chlorpyrifos (Ahmad and Das 1978).
The following year, the greater tolerance of chlorpyrifos by the Rivervale
beetle was confirmed by more extensive tests, using susceptible beetles for
comparison (Ng and Ahmad 1979). We now provide additional evidence for
a much greater chlorpyrifos tolerance in a Connecticut population of the
beetle, compared to the New Jersey insects.

VOLUME LXXXIX, NUMBER 1

35

Materials and Methods

Third-instar grubs of the Japanese beetle from Fairfield, Conn. (Brook-
lawn Country Club golf course) were supplied by A. M. Radko (U.S. Golf
Assocn., Far Hills, N.J.) on October 1, 1979. The grubs were held in the
laboratory for ca. 48 hr period and then tested for susceptibility to chlor-
pyrifos. The procedure and laboratory conditions for the maintenance of
the grubs were exactly the same as reported earlier (Ahmad and Das 1978).
For comparisons of susceptibility levels, grubs were collected from a chlor-
pyrifos-tolerant population in Rivervale (northern N.J.; Country Club golf
course), and a standard susceptible population from Adelphia (central N.J.;
Soils and Crops Res. Ctr., Rutgers Agric. Exp. Stn.) on October 7 and 13,
respectively. Grubs from these two populations also were held in the lab-
oratory for ca. 48 hr prior to screening for chlorpyrifos susceptibility.

The grubs were treated topically with one-microliter drops of chlorpyrifos
solutions, in 5 replicates, 10 grubs per replicate. Each test was repeated
twice and the data were pooled for analyses. Solvent controls were included.
The composition of the insecticide solvent, and the technique for the bioas-
say including that for grub rearing, were as reported earlier in detail (Ahmad
and Das 1978). Mortality was recorded on 8th post-treatment day since the
duration of the acute toxicity of chlorpyrifos in the Japanese beetle is 7 days
(loc. cit.). Mortality data were corrected for deaths in the control group by
Abbott’s formula, and subjected to probit analysis according to Finney
(1964). The regression coefficients (slopes) for the log dose-probit ( Idp ) lines
of the 3 populations also were analyzed for statistical difference.

Results and Discussion

The dosage-mortality responses of the Japanese beetle grubs from Adel-
phia (N.J.), Rivervale (N.J.) and Fairfield (Conn.) are shown in Fig. 1. The
LD values and their 95% confidence intervals are presented in Table 1. The
Rivervale population shows a 6.8-fold greater tolerance to chlorpyrifos at
the LD95 level to that of the susceptible population. Tests with adult beetles
also have shown the Rivervale population to be more tolerant to chlorpyrifos
than the Adelphia insects (Ng and Ahmad 1979). A 4.3 x greater tolerance
was discerned with a significant difference between the regression coeffi-
cients (b) of the Idp lines of the two populations (loc. cit.). In the present
investigation involving third-instar grubs, the regression coefficients of the
Idp lines of the Rivervale (b = 0.96) and Adelphia (b = 2.43) populations
also were found to be significantly different [t {df, 5) = 11.315, P < 0.05].
Thus our earlier conclusion that there were more chlorpyrifos-tolerant in-
dividuals in the Rivervale population than in the Adelphia population (Ng
and Ahmad 1979) was reaffirmed. The data on chlorpyrifos tolerance of the

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NEW YORK ENTOMOLOGICAL SOCIETY

pg CHLORPYRiFOSCLOG SCALE) PER LARVA

Fig. 1. Dosage-mortality responses of third-instar grubs of the Japanese beetles from Fair-
field (Conn.), Rivervale (northern N.J.), and Adelphia (central N.J.); 1979.

Rivervale beetles are typical of a “tolerant” insect population showing in-
cipient resistance to an insecticide (Busvine 1971).

Table 1 also indicates that by comparison to the Rivervale population,
grubs from Fairfield are exceedingly tolerant to chlorpyrifos. At the LD95
level the Fairfield insects are ca. 42 x as tolerant as the susceptible Adelphia
population, and ca. 6x more tolerant than the Rivervale population. As can
be expected the regression coefficient of the Idp line of the Fairfield (b =
0.96) grubs was significantly different to that of the Adelphia (b = 2.43)
population [t {df, 5) = 10.145; P < 0.05]. Moreover the 95% C.I.s of the
LD50 and LD95 of the Fairfield population showed no overlap to the corre-
sponding C.I.s of the Adelphia insects. Taken together, these data clearly
show that the selection towards more chlorpyrifos-tolerant individuals in
the Fairfield population has progressed further than in the chlorpyrifos-tol-
erant Rivervale population.

According to Dyte and Blackman (1967) if a population of insects is re-

VOLUME LXXXIX, NUMBER 1

37

Table 1. Comparison of chlorpyrifos susceptibility of third-instar grubs of the Japanese
beetle from Fairfield (Conn.), Rivervale (northern N.J.), and Adelphia (central N.J.); 1979.

Population3

LD50b

LD95b

LD95 ratio

Adelphia

1.90

9.17

1

(1.45-2.56)

(5.50-15.42)

Rivervale

1.20

62.45

6.8

(0.50-2.84)

(13.29-293.50)

Fairfield

7.50

388.06

42.2

(4.42-12.58)

(45.29-3,324.30)

a Mean weight of the grubs (N = 10) were 208 mg (±25 SD) for Adelphia, 196 mg (±39 SD)
for Rivervale, and 185 mg (±24 SD) for Fairfield populations.
b Confidence intervals ( P = 0.05) in parentheses.

sistant to an insecticide, the LD99 99 of the resistant strain must be signifi-
cantly greater than that of the susceptible strain. From the dosage-mortality
data in Table 1, we computed the lower C.I. for the LD99 99 of the Fairfield
grubs to be 579 /xg/grub; this did not overlap with the upper C.I. of the
LD99 99 of the Adelphia population, 221 /xg/grub. Thus the Fairfield popu-
lation appeared to meet the requirement for characterization as a resistant
population. On the other hand, as mentioned above the regression coeffi-
cient of the Idp line for the Fairfield population was considerably flatter
than that of the Adelphia population. A flat Idp line is often indicative of
the genetic variability in insecticide susceptibility of a population. A truly
resistant population typically depicts steep slope as is usually the case with
a homogeneous susceptible population (Busvine 1971). Therefore, it would
be premature to assume that the entire population from Fairfield is resistant
to chlorpyrifos, but rather it is an example of a partially-resistant field pop-
ulation, with more resistant insects in the population than susceptible ones.
The Fairfield population obviously has the potential for rapidly becoming
a highly resistant field population under further selection against chlorpy-
rifos.

Selection of a population against an organophosphorus insecticide often
confers resistance to other organophosphorus compounds (Perry and Agosin
1974). There is some indication that Japanese beetles may develop resis-
tance to organophosphorus compounds other than chlorpyrifos. For ex-
ample, trichlorfon and diazinon were as ineffective as chlorpyrifos during
1979 against the grubs of the Fairfield population (A. M. Radko, pers. com-
mun., 1979). On the other hand, trichlorfon afforded good protection against
the Rivervale grubs that are substantially more susceptible to chlorpyrifos
than the Fairfield population (W. Gaydosh, pers. commun., 1979).

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NEW YORK ENTOMOLOGICAL SOCIETY

In a previous report (Ng and Ahmad 1979), we had also shown the greater
inherent tolerance (2. 1 x) by the Rivervale beetles to bendiocarb, a potential
carbamate insecticide for grub control. Although we did not test bendiocarb
against the Fairfield strain, the possibility exists, that with time, the Jap-
anese beetle populations also may become resistant to carbamate insecti-
cides. This would not be surprising since the mode of action and detoxi-
cation mechanisms associated with tolerance and resistance are common to
both organophosphorus and carbamate insecticides (Perry and Agosin 1974;
Plapp 1976).

Acknowledgments

Paper of the Journal Series, New Jersey Agricultural Station, New Bruns-
wick, New Jersey. This work was performed under NJAES Project No.
08128 supported by Hatch Funds and grants from New Jersey Turfgrass
Association and Dow Chemical Company.

Literature Cited

Ahmad, S. and Y. T. Das. 1978. Japanese beetle grub: dosage-mortality response and symp-
toms of poisoning following treatments with chlorpyrifos and dieldrin. J. Econ. Entomol.
71:939-942.

Anonymous. 1973. Japanese beetle grubs developing resistance to chlordane. Turf-Grass
Times 9:35 pp.

Busvine, J. R. 1971. A Critical Review of the Techniques for Testing Insecticides. 2nd ed.

Commonwealth Agricultural Bureaux, Slough, England. 345 pp.

Dunbar, D. M. and R. L. Beard. 1975. Status of control of Japanese beetle and Oriental
beetles in Connecticut. Agric. Exp. Stn. Bull. 757:1-5.

Dyte, C. E. and D. G. Blackman. 1967. Selection of a DDT-resistant strain of Tribolium
castaneam (Herbst). J. Stored Prod. Res. 2:211-228.

Finney, D. J. 1964. Probit Analysis. 2nd ed. Cambridge University Press, London, England.
318 pp.

Fleming, W. E. 1950. Protection of turf from damage by Japanese beetle grubs. USDA Leafl.
290:8 pp.

Ng, Y.-S. and S. Ahmad. 1979. Resistance to dieldrin and tolerance to chlorpyrifos and
bendiocarb in a northern New Jersey population of Japanese beetle. J. Econ. Entomol.
72:698-700.

Niemczyk, H. D. and K. O. Lawrence. 1973. Japanese beetle: evidence of resistance to
cyclodiene insecticides in larvae and adults in Ohio. J. Econ. Entomol. 66:520-521.
Perry, A. S. and M. Agosin. 1974. The physiology of insecticide resistance by insects, p. 3-
121. In M. Rockstein (ed). The Physiology of Insects. 2nd ed., Vol. 6. Academic Press,
New York. 548 pp.

Plapp, F. W. 1976. Biochemical genetics of insecticide resistance. Annu. Rev. Entomol.
21:179-197.

Tashiro, H. and W. Neuhauser. 1973. Chlordane-resistant Japanese beetle in New York.
Search Agric. (Geneva, N.Y.) 3:6 pp.

VOLUME LXXXIX, NUMBER 1

39

Department of Entomology and Economic Zoology, New Jersey Agri-
cultural Experiment Station, Cook College, Rutgers — The State University
of New Jersey, New Brunswick, New Jersey 08903.

Received for publication July 8, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(l), 1981, pp. 40-42

OCHLERINI, A NEW TRIBE IN DISCOCEPHALINAE
(HEMIPTERA: PENTATOMIDAE)

L. H. Rolston

Abstract. — Ocherini is proposed as a new tribe of Discocephalinae. The
tribe is characterized and 23 genera assigned to it.

The discocephalines, which are restricted to the Western Hemisphere,
have been variously treated as a subfamily of Pentatomidae or a tribe of
Pentatominae. Rolston and McDonald (1979) regarded this group as a
subfamily and provided a diagnosis based primarily on the apparent origin
of the labium, usually caudad of the anterior limit of the eyes, and the
position of the trichobothria on sternite vii, the mesial member of each pair
being on or, most often, laterad of an imaginary line projected tangentially
along the lateral margins of the spiracular openings on sternites vi and vii.
These authors removed all Western Hemisphere genera except Brochymena
from Halyini in Pentatominae, placing Caracia Stal, Marghita Ruckes and
Janeirona Distant in Pentatomini and the remaining genera in Disco-
cephalinae. Those genera placed in Discocephalinae form a new tribe, for
which a name and diagnosis are here provided, that is characterized pri-
marily by the shallowly excavated or flattened superior surface of the third
tarsal segment of the hind legs in females and sometimes in males as well.
Members of this tribe are with few exceptions recognizable by their dull
black or fuscous coloration, while the remaining genera of discocephalines,
which constitute the nominate tribe, are brown, often mottled with black,
or shiny black.

Ochlerini Rolston, new tribe

Type genus. — Ochlerus Spinola, 1837.

Diagnosis. — Superior surface of third tarsal segment of hind legs shallow-
ly excavated in females and sometimes in males (only flattened in Adoxo-
platys ).

Trichobothria on at least last sternite laterad of adjacent spiracles, rarely
with mesial trichobothrium of each pair on last sternite on line tangential to
spiracle openings on last two sternites. Labium usually arising on or pos-
terior to plane transecting head at right angle to longitudinal body axis and
at anterior limit of eyes.

Basal segment of rostrum projecting caudad of bucculae, terminating on
prosternum. Mesosternum thinly carinate mesially, metasternum usually so

VOLUME LXXXIX, NUMBER 1

41

but sometimes weakly tectiform, flat or sulcate. Metapleural ostioles each
accompanied by auricle, this often somewhat elongated but not drawn out
into ruga. Scutellum longer than wide at base (brachypterous forms ex-
cepted). All tibiae broadly sulcate. Spiracles present on paratergite 8 of
females, on sternite 8 of males.

Comments: Among Western Hemisphere pentatomids the tarsal char-

acter state appears unique to this tribe. The location of the trichobothria
laterad of the adjacent spiracle on the last sternite is characteristic of Dis-
cocephalinae, but this character state is also found in a few members of
Pentatominae. The labial origin posterior to the anterior limit of the eyes is
apparently constant in Discocephalini but variable in Ochlerini. All of the
character states enumerated in the last paragraph of the diagnosis are con-
stant in Ochlerini, as far as known, but various combinations of them appear
elsewhere.

The following genera are placed in Ochlerini:

Adoxoplatys Breddin
Alathesus Dallas
Alitocoris Sailer
Audintella Spinola
Brachelytron Ruckes
Eritrachys Ruckes
Herrichella Distant
Lincus Stal
Macropygium Spinola
Melambyrsus Breddin
Melanodermus Stal
Minilincus Ruckes

Miopygium Breddin
Moncus Stal

Neoadoxoplatys Kormilev
Ochlerus Spinola
Orbatina Ruckes
Paralincus Distant
Parochlerus Breddin
Pherecles Stal
Schaefferella Spinola
Schaderia Ruckes
Tetrochlerus Breddin

I have examined species, usually the type species, of all of the above
genera except Audintella and Melambyrsus. These two monotypic genera
are included in Ochlerini on the basis of their descriptions.

The 23 genera of Ochlerini contain only 70 described species, but exam-
ination of a few collections reveals several undescribed genera and species.

Acknowledgments

I am indebted to Drs. H. Dodge Engleman, R. C. Froeschner (U.S. Na-
tional Museum), Jocelia Grazia (Universidade Estadual de Campinas), Ran-
dall T. Schuh (American Museum of Natural History) and P. van Doesburg
(Riksmuseum van Natuurlijke Historie) for the loan of specimens relevant
to this study, and especially to W. R. Dolling (British Museum [Natural
History]), A. Kaltenbach (Naturhistorisches Museum, Wien), G. Petersen

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NEW YORK ENTOMOLOGICAL SOCIETY

(Akademie der Landwirtshaftwissenshaften der D. D. R.) and Randall T.
Schuh for the loan of type material.

Dr. F. J. D. McDonald’s skill in dissection and knowledge of pentatomoid
genitalia contributed much to this work.

Literature Cited

Rolston, L. H. and F. J. D. McDonald. 1979. Keys and diagnosis for the families of Western
Hemisphere Pentatomoidea, subfamilies of Pentatomidae and tribes of Pentatominae
(Hemiptera). J. N.Y. Entomol. Soc. 87(3): 189-207.

Department of Entomology, Louisiana Agricultural Experiment Station,
Louisiana State University, Baton Rouge, LA 70803.

Received for publication August 4, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(l), 1981, pp. 43-52

NOTES ON THE BIOLOGY OF A COMMON SUNFLOWER BEE,
MELISSODES ( EUMELISSODES ) AGILIS CRESSON

F. D. Parker, V. J. Tepedino and G. E. Bohart

Abstract. — Melissodes agilis, an anthophorid bee that is oligolectic on
sunflowers, was found nesting in and adjacent to a planting of commercial
sunflowers in northern Utah. Observations and excavation of nests yielded
information on nest architecture, larval morphology, foraging activity, and
parasites. The species will probably be most valuable as a commercial
pollinator where minimum- or no-tillage agricultural practices are appropri-
ate.

Introduction

The commonest native bee species frequenting sunflower ( Helianthus
spp.) in the western United States is Melissodes agilis Cresson, a ground-
nesting member of the Anthophoridae. Although M. agilis is probably re-
sponsible for a significant proportion of sunflower pollination throughout
the west (Parker 1981a, b), biological information on this commerically im-
portant species is limited to a brief note on the nest tumulus and length of
the main burrow (Rau 1922) and a comment by Custer (1928) that a female
Melissodes (presumed to be M. agilis ) entered a nest of Svastra obliqua
(Say), another anthophorid bee. During the summer of 1979, large numbers
of M. agilis were observed visiting and nesting in our plantings of com-
mercial sunflowers in northern Utah; and we took the opportunity to study
the biology of this species. Specifically, we provide information on nest
architecture, larval morphology, nest associates, seasonal occurrence, for-
aging activities, and sleeping aggregations of males.

Nesting Site

Melissodes agilis nested in and between irrigation furrows in a 1-acre
(0. 4047-ha) plot planted to sunflowers ( Helianthus annuus variety macro-
carpus (D.C.)) and summer squash ( Cucurbita pepo L.) near Logan, Utah,
during July and August 1979. The site was nearly flat and the soil type was
Millville Silt loam. More than 20 nests of M. agilis were marked with stakes
and some were excavated in September. The nests appeared randomly dis-
tributed throughout the sunflower and zucchini plots; a few were as close
together as 12 cm. The following description of the nests was taken from 6
excavated nests with cells.

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NEW YORK ENTOMOLOGICAL SOCIETY

VOLUME LXXXIX, NUMBER 1

45

O Cells
® Old cell (14)

Fig. 3. Diagram of typical Melissodes nest. A. Arrangement of cells. B. Cross section of
nest with depth of cells indicated.

Nest Architecture

No bees were observed initiating nest excavation; nests were located by
observing bees entering or leaving their nests or by observing the newly
excavated crater-like tumuli. The mound of soil from a typical nest exca-
vation measured 5 cm across and 1.5 cm high. However, tumuli were not
always evident. The entrance hole was in the middle of the crater and av-
eraged 7 mm in diameter (Fig. 1). The entrance hole was not plugged during
nest provisioning.

When we excavated the nests, the main and lateral burrows were filled
with soil, but the main burrow could be traced by blowing out the soil and
refilling it with plaster of Paris powder. Typically, the main burrow was 7
mm in diameter and spiraled downward for 4 cm before descending nearly
vertically to a depth of about 12 cm (range 11-17 cm), where the first cell
was placed. The main burrow was unlined, but the walls were smooth. It

Figs. 1, 2, 4-6. 1. Nest entrance and tumulus of Melissodes. 2. Compacted soil in filled

Melissodes burrow. 4. Waxed lined cell of Melissodes. 5. Tops of 2 Melissodes cells exposed
during nest excavation. 6. Overwintering Melissodes larva in cell; cocoon cut away to expose
layers of the cocoon.

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NEW YORK ENTOMOLOGICAL SOCIETY

was not possible to trace the route of the lateral branches because they were
filled with tightly packed soil (Fig. 2). Lateral burrows were from 1 to 10
cm in length and radiated away from the main burrow (Fig. 3). In one nest,
the cells were distributed in a pattern indicating that at least 12 lateral bur-
rows diverged from the central main burrow. Since the cells were found at
8 levels (Fig. 3), it appeared that most of the laterals were begun at different
levels.

The number of cells/nest ranged from 1 to 27 (x = 11.5). Cells were found
at depths ranging from 11 to 19 cm below the soil surface. In multi-celled
nests, some cells were about 1 cm below an adjacent one, an indication that
some lateral burrows contained more than one cell.

Cell Structure and Morphology

The oval cells were vertical in orientation. Each cell was 7-8 mm wide
at its greatest diameter and 8-14 mm long (Fig. 4). Cell walls were made by
the bee compacting about 2 mm of soil against the “roughed-out” original
excavations. Following this, the inner walls (Except the cell cap) were coat-
ed with wax. The wax lining was evenly applied to all cell walls and was
thick enough to be readily scraped off with an insect pin. The upper V3 of
each cell was paler and slightly rougher than the basal portion. The area of
overlap between these color bands was darkened. The cell cap was com-
posed of rings of soil, but the precise pattern could not be determined since
the top of every cell was at least partially destroyed during excavation. In
all cells examined, the pollen provisions had been consumed.

Most fecal material was incorporated into the cocoon layers, but at the
top of the cell were some loose, concentric, tubular strands of fecal pellets
in a 3 mm thick layer (Fig. 5). The larva started the cocoon by first lining
the cell walls (except the top) with a thin varnish of silk. Over this layer it
smeared successive, longitudinally directed strands of yellowish fecal ma-
terial, separated but held together by layers of silk. Inside the fecal layer
the larva produced a shiny inner cocoon of thin, transparent, yellowish silk
(Fig. 6), in which it embedded a network of larger, darker strands. This
inner envelope was entire and of uniform thickness and its inner surface
was smooth and shiny. It was surmounted by a dome-shaped cap composed
of several sheets of silk loosely held together.

The overwintering prepupal larvae were creamy white and rather flaccid.

Figs. 7-14. 7-8. Overwintering Melissodes larvae. 7. Lateral view. 8. Larvae tilted to

illustrate lateral lobes. 9-10. Triepeolus prepupae. 9. Ventral view. Note head capsule of 1st
instar larva (arrow). 10. Lateral view. 1 1-12. Cells with Triepeolus. 11. Basal ring (arrow) that
holds prepupa. 12. Prepupa surrounded by fecal pellets. 13. Puncture in cell wall made by
Triepeolus. 14. Egg chamber of Triepeolus. Note shriveled egg chorion.

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NEW YORK ENTOMOLOGICAL SOCIETY

They were C-shaped and vertical with their weight resting on the postero-
dorsal area (Figs. 7, 8).

Nest Associates

Three nests contained a total of 7 cells parasitized by the cuckoo bee
Triepeolus helianthi (Robertson). The distinctive rigid prepupae (Figs. 9,
10) were held erect in the center of the cell by a basal ring of feces or pollen
(Figs. 11, 12). The tubular-shaped fecal pellets were yellowish and were
deposited in short strands, but in layers, on the upper % of the walls and
beneath the cell cap. The egg chamber (0.5 mm wide and 2 mm long) was
located 40% of the way up the side wall and perpendicular to the cell walls
(Figs. 13, 14). There was no evidence that the larvae deposited any silk.
Hurd and Linsley (1959) reported that adults of T. helianthi entered nests
of Melissodes composita Tucker, but they did not find any parasitized host
cells.

One cell contained the coarctate prepupal larva of the meloid beetle Nem-
ognatha. Another two Melissodes cells contained castings of earthworms,
which presumably were deposited in the cell before the bee larvae matured.

Observations of Adults

Seasonal occurrence. — Since sunflowers were planted at three different
times, bloom was available for the entire season. We counted all bees on
the flower heads at 0900, 1100, 1300 hr every Monday, Wednesday and
Friday from 25 July to 4 September. Both male and female M. agilis were
present on July 25 when bloom began. At this time their wing margins were
entire and their body hairs were not worn, indicating recent emergence.
Both sexes were abundant throughout the summer. The pattern of their
seasonal occurence suggests a single generation/year. Their abundance from
week to week closely followed the abundance of sunflower bloom (Fig. 15).

Daily activities. — Males were recorded more frequently from flower heads
than females (2.43/1.0$). Initially, males were more abundant at 0900 hr
than at any other time, but in late August and early September, when morn-
ing temperatures began to be quite cool, their abundance was similar in all
count periods.

Males gathered on sunflower heads in sleeping aggregations of from 2 to
20 bees/head in late afternoon and during inclement weather. No observa-
tions were made to determine if they chose the same sleeping cluster or the
same flower head each night. They clustered in partially opened heads under
the large ray petals. (The heads had to be tilted back before the bees could
be seen.)

Female Melissodes were most abundant on the flowers early in the day.
For example, at 0900 hr twice as many females were present as at 1100 hr,

VOLUME LXXXIX, NUMBER I

49

Fig. 15. Seasonal abundance of sunflower heads in bloom, number of Melissodes, and
percentage of female Melissodes.

and at 1300 hr, about lA as many females were present as at 1100 hr. Ap-
parently, most of the females remained in their nests in the afternoon; this
was corroborated by observations of the activity of individual females at
their nests (see below). Female M. agilis provisioned their nests exclusively
with sunflower pollen. The maximum pollen load carried by a female bee
was estimated at 330,000 grains (estimated by washing the pollen from the
body hairs and counting the pollen grains with a haemocytometer) (Parker
1981b).

Triepeolus helianthi was present throughout the observation period. The
activity pattern of this cuckoo bee varied with the time of day. In the morn-
ing, females were often observed flying slowly several inches above the soil
and investigating all types of holes. Several were seen entering Melissodes
nests where they remained only a few minutes (see below). In the after-
noon, Triepeolus spent more time on the flowers than in the morning. Their
abundance on flower heads at 1300 hr was nearly double that at 0900 hr.
Thus, this parasite appears to search for and invade host nests primarily in
the morning when most host adults are foraging and to forage in the after-
noon when Melissodes remain in their nests.

50

NEW YORK ENTOMOLOGICAL SOCIETY

Table 1. Mean time of foraging trips and within-nest periods of two Melissodes agilis
females on three observation dates. Initial foraging trips for each day and extended within-nest
periods (4, see text) excluded. Numbers in parentheses are standard deviation and sample size,
respectively.

Observation

date

(August)

Mean time/foraging trip (sec)

Mean time within nest (sec)

S

M

S

M

2

231.5 (±306.5;

10)

293.8 (±225.6; 9)

43.3 (±13.0; 9)

57.8 (±11.8; 8)

7

501.1 (±360.6;

11)

594.3 (±408.1; 9)

65.6 (±15.6; 10)

99.6 (±83.5; 9)

13

950.5 (±969.5;

11)

1,086.8 (±715.0; 9)

84.8 (±28.0; 11)

112.3 (±90.3; 8)

Foraging Activity

We monitored the activity of two female Melissodes bees at the nesting
site on three mornings, beginning at 0600 hr and using a stopwatch and
portable tape recorder. The nests were within 50 cm of each other near the
top of a furrow that separated a row of zucchini from the periphery of the
planting. We recorded times for commencement of activity and the number
and duration of trips from the nest as well as periods spent within the nest.

Both nests were unplugged and were without tumuli each morning when
observations began. Bee activity commenced between 0630 and 0745 hr.
Activity began earlier when temperatures were warm and cloud cover slight.
Bee S always became active first and made her initial trips from the nest 5-
15 minutes before bee M. The first trip was invariably an extended one for
both bees; it varied from 21 to 32 minutes in duration. Most other trips were
of shorter duration (Table 1). Bees returning from short trips invariably
carried pollen. Bee S also consistently completed her foraging trips more
rapidly than did bee M. Bees entered their respective nests from return
flights without hesitation and never investigated each others’ nests.

There was no discernible difference between the duration of early and
later foraging trips for either bee on any date, as might be expected if pollen
or nectar were becoming limiting later in the morning. However, foraging
trips became longer as the month progressed for both bees. The relatively
extended trips of 13 August were associated with sporadic rainfall and heavy
cloud cover throughout the morning. Both bees were trapped in the field by
rain on several occasions and did not return until the rain had ceased. Al-
though cloud cover remained heavy, both bees left for additional foraging
trips after brief periods in the nest.

Periods spent within the nest after the bees returned from foraging trips
were usually short (30-150 sec); presumably the pollen and nectar that the
bees had collected were deposited during these brief periods in the nest.
Occasionally, within-nest periods were much longer; four such periods rang-
ing from 21 to 41 minutes were recorded. We believe that pollen loaf prep-

VOLUME LXXXIX, NUMBER 1

51

aration, egg laying and cell closure occurred during these extended within-
nest periods. If this is so, then the number of foraging trips necessary to
provision a single cell can be estimated from the ratio of short/long within-
nest periods. For the two bees, we recorded 55 short and 4 long periods,
for an estimate of approximately 14 foraging trips/cell.

On the first two dates, we terminated observations at 1000 hr, when the
bees had been in their nests for at least 30 consecutive minutes. Foraging
appeared to cease on both days by approximately 0915 hr. By this time, bee
S had made 11 trips on 2 August and 12 on 7 August; bee M had made 10
trips on each day. Subsequent foraging activity was minimal, as previously
shown by comparisons of numbers of females recorded on flowers at the
different time periods. An exception was on 13 August, when the bees
foraged in sporadic rain until just after 1 100 hr. Although we continued ob-
servations until 1200 hr, neither bee emerged again. It is interesting to note
that at 1100 hr, bee S returned from her 12th trip and M returned from her
10th trip, the same number as had been made on previous days.

We observed a single intrusion by Triepeolus helianthi into the nest of
bee S on 2 August. The cuckoo bee was first seen flying around the nesting
site at 0755 hr, while both Melissodes were active. The Triepeolus remained
in the vicinity resting on the ground or on nearby vegetation. At 0803 hr,
she briefly investigated the entrance of S’s nest while the latter was out and
then moved to about 10 cm west of the entrance. S returned, remained in
the nest for a brief period, and left at 0807 hr. As S was leaving, Triepeolus
began to move towards the entrance and entered less than one second after
S had left. Triepeolus remained in the nest for 90 sec, then crawled out and
down the furrow. She then flew to a nearby grass blade, where she re-
mained, pulsated her abdomen for several seconds and then left. When the
nest was excavated in September, two larvae of Triepeolus were found.

Discussion

This study and those of Parker (1981a, b) have shown that Melissodes
agilis possesses numerous characteristics which make it a valuable polli-
nator in commercial sunflower plantings. The emergence of M. agilis adults
was closely synchronized with the initiation of sunflower bloom, and num-
bers of bees followed the number of flowers available (Fig. 15). Females
exclusively used sunflower pollen to provision their cells (Parker 1981b). In
contrast to some other species of anthophorid bees in which the nesting site
is patrolled by males in their search for females (e.g. Centris, Alcock et al.
1977), male M. agilis patrol the flowers for prospective mates. Thus, both
sexes are important pollinators. Finally, because females nested in or ad-
jacent to the planting, foraging trips were of short duration, and females
were probably able to complete approximately 1 cell/day. Elsewhere (Parker

52

NEW YORK ENTOMOLOGICAL SOCIETY

1981a, b) has shown that M. agilis is also a more efficient pollinator than
the commonly used honey bee. Indeed, although the honey bee is usually
credited for most sunflower pollination (McGregor 1976), natural field pop-
ulations of M. agilis, which are usually ignored, may be more important.

The tendency of M. agilis to nest in sunflower fields makes their popu-
lations vulnerable to certain tillage practices. Since cells were constructed
at soil depths between 11 and 19 cm, tillage practices that disrupt the soil
below 10 cm are likely to have a devastating effect on populations of M.
agilis. A zero or minimum tillage system (Robinson 1978), where appropri-
ate, would be least hazardous to Melissodes populations.

Acknowledgments

Appreciation is extended to our technichians, D. Veirs, C. Hatley, and
K. Ruggeri and part-time help G. Trostle and J. Higginson, for their assis-
tance in conducting field work and nest excavations. Thanks are also due to
E. Gorton Linsley, Univ. California-Berkeley and W. E. LaBerge, Illinois
Natural History Survey, for reviewing the manuscript.

Literature Cited

Alcock, J., C. E. Jones and S. L. Buchmann. 1977. Male mating strategies in the bee Centris
pallida Fox (Hymenoptera: Anthophoridae). Am. Nat. 111:145-155.

Custer, C. P. 1928. On the nesting habits of Melissodes Latr. Can. Entomol. 60:28-31.
Hurd, P. D., Jr. 1979. Apoidea, p. 1741-2209. In: Catalog of Hymenoptera in America North
of Mexico. Vol. 2. K. V. Krombein, P. D. Hurd, Jr., D. P. Smith and B. D. Burks
(eds.). Smithsonian Institution Press, Washington, D.C.

and E. G. Linsley. 1959. Observations on the nest-site behavior of Melissodes com-

posita Tucker and its parasites, with notes on the communal use of nest entrances.
Entomol. News 70:141-146.

McGregor, S. E. 1976. Insect pollination of cultivated crop plants. U.S. Dep. Agric., Agric.

Res. Serv. Agric. Hdbk. No. 496, Washington, D.C.

Parker, F. D. 1981a. Sunflower pollination: abundance, diversity, and seasonality of bees and
their effect on yields. J. Apic. Res. (In press).

. 1981b. How efficient are bees in pollinating sunflowers? J. Kans. Entomol. Soc. (In

press).

Rau, P. 1922. Ecological and behavioral notes on Missouri insects. Trans. Acad. Sci. St.
Louis 24: 1-71.

Robinson, R. G. 1978. Production and culture, p. 89-143. In: Sunflower Science and Tech-
nology, J. F. Carter (ed.). Amer. Soc. Agron., Madison, Wise.

Bee Biology and Systematics Laboratory, Agricultural Research Science
and Education Admin., USDA, Utah State University, UMC 53, Logan,
Utah 84322.

Received for publication October 30, 1980.

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Journal of the

New York Entomological Society

VOLUME LXXXIX JUNE 1981 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. Streu

Publication Committee
Dr. Randall T. Schuh
American Museum of Natural History
Dr. Louis Trombetta
St. Johns University

CONTENTS

Edwin Way Teale 1899-1980 Alexander B. Klots 54-55

Hydropsyche larvae (Trichoptera: Hydropsychidae) from a lake weedbed

Bernard A. Marcus 56-58

The larva of Agrotis volubilis Harvey (Lepidoptera: Noctuidae)

Timothy L. McCabe 59-64

Histological changes in the antenna of Tenebrio molitor L. after imaginal eclosion
(Coleoptera: Tenebrionidae) Louis D. Trombetta and James Forbes 65-79

Breeding experiments with Morpho peleides insularis (Lepidoptera: Nymphalidae)

in Trinidad, W.I. Malcolm Barcant 80-88

Parasitism of single and multiple egg clusters of Euphydryas phaeton (Nymphalidae)

Nancy E. Stamp 89-97

Some observations on soaring flight in the mourning cloak butterfly ( Nymphalis
antiopa L.) in southern Ontario David L. Gibo 98-101

Observations on the spring and summer behavior of the 12-spotted ladybird beetle,
Coleomegilla maculata (DeGeer) (Coleoptera: Coccinellidae)

Allen H. Benton and Andrew J. Crump 102-108

Principal components analysis of biogeographic patterns among Heliconia insect
communities Richard P. Seifert 109-122

Notes on seasonality and habitat associations of tropical cicadas ( Homoptera: Cicadi-
dae) in premontane and montane tropical moist forests in Costa Rica

Allen M. Young 123-142

A little-known, anonymous work on American and European butterflies and moths
(1906), which should be attributed to William Beutenmiiller (Lepidoptera: Nym-
phalidae) Cyril F. dos Passos 143-145

Book review
Announcement

146-147

148

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(2), 1981, pp. 54-55

EDWIN WAY TEALE 1899-1980
Alexander B. Klots

Edwin Way Teale, one of the oldest and most distinguished members of
the New York and Brooklyn Entomological Societies, died in Norwich,
Connecticut on 18 October, 1980. A member of each society since the
1930’s, he received the John Burroughs Medal and a Pulitzer Prize for his
natural history writings. He served as President of the New York Society
in 1944 and President of the Brooklyn Society in 1950-51 and 1953-54.

Born in Joliet, Illinois in 1899, Mr. Teale attended Earlham College in
Indiana (where he met his wife, then Nellie Donovan) and later Columbia
University. During his career Nellie, herself a keen and devoted naturalist,
was always with him. Perhaps she specialized on plants more than Ed did,
but together they made an ever-close team for the observation and under-
standing of nature.

The Teales soon came to New York, where Ed became a staff writer for
Popular Science magazine and '‘free-lanced” for some years. Eventually he
wrote his first book, Grassroot Jungles (1937) and from then on concen-
trated on photography and writing books, and never looked back. Eventu-
ally he published more than 20 books “of his own” in addition to scholarly
and sensitive studies of the lives and works of such great naturalists as
Thoreau, Hudson, Audubon, Fabre, Burroughs and Muir.

Many of the Teale books show a marked preoccupation with seasonal
changes. His most famous series: North With The Spring , Journey Into
Summer, Autumn Across America and Wandering Through Winter tells of
how the Teales followed the changing seasons throughout the United States
and Canada. It was for this series that he received the Pulitzer Prize in 1966.

Entomology played a very large part in the Teale writings, much more so
than in those of other authors. Even as a boy he was fascinated by the
complexity of the lives of insects; and this continued throughout his life. A
Book About Bees (1940), originally titled The Golden Throng, is a very
thorough description of the complexity of life in a honeybee hive. Like his
other books it was illustrated with his own photographs.

Photography indeed played a very large part in Teale’ s life. Starting with
what today seems a crude and cumbersome apparatus, a big bellows-exten-
sion camera and a loose flash-powder gun, Teale constantly photographed
his subjects. His photographs covered all phases of nature; and many of
them are superb by any standards.

Living in Baldwin, Long Island, Teale could be a faithful attendant at
meetings of the two entomological societies, and constantly showed up with
an interesting note on some observation, or a remarkable photograph. Many
of his observations were real contributions to knowledge. Then, in 1959,

VOLUME LXXXIX, NUMBER 2

55

encroaching urbanization dictated a move. Partly on this writer’s suggestion
a large area with a very old house was bought in largely rural Hampton,
Windham County, Connecticut. “Trail Wood,” as the place was christened,
is almost ideal for the naturalist, with many acres of unsullied old and young
woodlands, fields, pastures, hillsides, a beaver pond, a brook and, a recent
addition, a large, clear pond. Here for many years Ed and Nellie came to
know the most intimate details of the ever-abundant, ever-changing wildlife.

Despite his great familiarity with all phases of natural history Ed never
wavered in his primary interest in insects and their ways. After the move
to Connecticut he usually managed to have business affairs and confer-
ences with his publishers in New York scheduled for Tuesdays so that
he could attend the dinners and meetings of the New York Entomological
Society. He always arrived with some new and interesting observation
to tell about.

I particularly remember once when Ed and I were watching and photo-
graphing close-up a large bullfrog floating just below the surface at the
edge of his pond. Ed noticed and commented on the fact that the frog
kept only one eyeball at a time above the surface. The above-water eye
would be pulled down as the other came up inregular sequence. How
delighted he was when we noticed that tiny biting midges, or “no-see-ums”
(Ceratopogonidae) were feeding on the protruding eyeballs. When one was
pulled down they simply shifted to the other as it came up. We later
learned that there are species that specialize on amphibians! With the in-
sects, always something new!

“Trail Wood” will be taken over and maintained as the Edwin Way
Teale Memorial Sanctuary (and nature center) by the Connecticut Audubon
Society when Nellie no longer wishes to live there. Ed’s manuscripts and
books will go the Library of the University of Connecticut at Storrs
and be maintained there in a special memorial section.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(2), 1981, pp. 56-58

HYDROPSYCHE LARVAE (TRICHOPTERA: HYDROPS YCHIDAE)
FROM A LAKE WEEDBED

Bernard A. Marcus1

Abstract. — In February of 1979, two individual larvae of Hydropsyche
sp. (Trichoptera: Hydropsychidae) were collected from submerged aquatic
plants in Conesus Lake, N.Y. at 1.8 m in depth where they may have
survived for as long as ten months. No previous report on these organisms
suggests that they occur in such a situation. It is possible that their absence
from lake weed beds is due to factors other than simply unfavorable envi-
ronment.

Published information on the larvae of hydropsychid caddisflies (Trichop-
tera) generalizes that these organisms are commonly found in lotic waters
where they spin nets on the substrate and capture food materials that are
carried by the current (Denning 1968; Hynes 1970; MacKay 1978). In Great
Britain, hydropsychid larvae are found only in rivers and streams, not in
lakes or ponds (Eddington 1968).

In North America, hydropsychid larvae are found in the wave-washed
shoreline areas of large lakes. Barton and Hynes (1978) specified that they
can manage in standing water if there is sufficient movement to deliver food
materials to their nets. These investigators collected specimens from the
exposed Canadian shores of the Great Lakes. Much earlier, Ross (1944)
reported finding Hydropsyche recurvata Banks larvae along the Illinois
shore of Lake Michigan, at Evanston, living among the rocks in less than
one meter of water. Wiggins (1977) reported finding the same species, along
with larvae of the genus Cheumatopsyche, along the shore of Lake Win-
nipegosis in Mannitoba, Canada.

On February 19, 1979, I captured two larvae of Hydropsyche sp. from a
bed of submerged aquatic macrophytes at the Dacola Shores (inlet) area of
Conesus Lake, New York. The organisms were found among a sample
containing 75% Heteranthera dubia (Jacq.) MacM.; 20% Ceratophylum de-
mersum L.; and 5% Elodea canadensis Michx. The plants, growing in 1.8
m of water, were collected through the ice by means of a rake. Both larvae
appeared to be later instars. To my knowledge, this is the first time these
organisms have been reported from such conditions. Between January and
August, 1979, monthly collections were made at weedbeds at five separate

1 Present address: Division of Mathematics and Science, Genesee Community College, Ba-
tavia, New York 14020.

VOLUME LXXXIX, NUMBER 2

57

sites on Conesus Lake at depths from 1.5 to 4.0 m. Similar collections were
taken from six sites at nearby Honeoye Lake at depths from 2.0 to 3.0 m.
Hydropsyche larvae were found only in the February 19 collection at 1.8 m
at Dacola Shores in Conesus Lake. The general absense of the larvae from
the lake vegetation is consistent with the knowledge on the family; the single
find is an unusual, but still interesting, event.

Dacola Shores is in the southernmost part of Conesus Lake. Conesus
Inlet, the lake’s major tributary, empties into this area. Although the inlet
is the principal contributor of water to the lake (Forest, Wade and Maxwell
1978), it is sluggish near its mouth and does not appear to offer the type of
habitat preferred by Hydropsychidae. Additionally, any hydropsychid lar-
vae among the invertebrate drift (Waters 1965) from the inlet would have
ample opportunity to settle in the mouth of the stream well before even
entering the lake. Furthermore, the sampling site at Dacola Shores is beyond
the zone affected by water movement from the inlet.

The most plausible explanation, then, for the occurrence of typically run-
ning water invertebrates in a lake weedbed is that they were carried in from
the inlet stream during an episode of unusually high water. Organisms usu-
ally do not survive that experience for very long (Dendy 1944). If this is the
case, however, the two larvae collected would have survived for up to ten
months in the lake environment, to which they are supposedly poorly adapt-
ed. The previous high waters would have been in the Spring of 1978.

While one case does not refute the general rule on the ecology of immature
hydropsychids, it does offer an interesting note on their survivability, and
suggests that similar possibilities should not be overlooked in field studies.
If survival under these conditions is more common than previously known,
then the absence of hydropsychid larvae from the submerged weedbeds of
lakes may be attributed to reasons other than their simple inability to tol-
erate that kind of environment.

Acknowledgments

I am indebted to Dr. Herman S. Forest, State University College at Ge-
neseo, N.Y. for his criticisms of the manuscript and to an anonymous re-
viewer for pertinent references.

Literature Cited

Barton, D. R. and H. B. N. Hynes. 1978. Wave-zone macrobenthos of the exposed Canadian
shores of the St. Lawrence Great Lakes. Jour. Great Lakes Res. 4:27-45.

Denning, D. G. 1968. Trichoptera, pp. 237-270. In R. L. Usinger (Ed.), Aquatic insects of
California. U. California Press, Berkeley.

Dendy, J. S. 1944. The fate of animals in stream drift when carried into lakes. Ecol. Monogr.
14:333-357.

58

NEW YORK ENTOMOLOGICAL SOCIETY

Eddington, J. M. 1968. Habitat preferences in net-apinning caddis larvae with special reference
to the influence of water velocity. Jour. Anim. Ecol. 37:675-692.

Forest, H. S., J. Q. Wade and T. F. Maxwell. 1978. The limnology of Conesus Lake, pp.
122-225. In J. A. Bloomfield (Ed.), Lakes of New York State, Vol. I: Ecology of the
Finger Lakes. Academic Press, New York.

Hynes, H. B. N. 1970. The ecology of running waters. U. Toronto Press, Toronto. 555 pp.
Mac Kay, R. J. 1978. Larval identification and instar association in some species of Hydro-
psyche and Cheumatopsyche (Trichoptera: Hydropsychidae). Ann. Entomol. Soc.
Amer. 71:499-509.

Ross, H. R. 1944. The caddisflies, or Trichoptera, of Illinois. 111. Nat. Hist. Surv. Bull. 23:1—
326.

Waters, T. F. 1965. Interpretation of invertebrate drift in streams. Ecology 46:327-334.
Wiggins, G. B. 1977. Larvae of the North American caddisfly genera (Trichoptera). U. To-
ronto Press, Toronto. 401 pp.

Environmental Resource Center, State University of New York College
of Arts and Science, Geneseo, New York 14454.

Received for publication May 16, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(2), 1981, pp. 59-64

THE LARVA OF AGROTIS VOLUBILIS HARVEY
(LEPIDOPTERA: NOCTUIDAE)1

Timothy L. McCabe

Abstract. — The mature larva of Agrotis volubilis Harvey (Lepidoptera:
Noctuidae) is described and illustrated. Achillea millefolium L. was found
to be an acceptable host. Late instar larvae became more general, accepting
Vaccinium vacillans Torr. and Oenothera biennis L. Larvae were full-
grown by the 20th of July (52 days), but would not pupate. They remained
healthy through October, when they were placed in out-of-door cages; the
larvae failed to overwinter.

Despite the economic significance of the genus Agrotis, many species’
biologies are unknown and identification of the adults of several species is
hampered by the paucity of species-specific genitalic characters and the
variability shown in adult habitus. Agrotis volubilis Harvey (1874) can not
presently be satisfactorily separated from A. stigmosa Morrison (1874) or
even Feltia [Agrotis] musa Smith (1910) on the basis of adult morphology
(Ahmadi, 1977).

The genitalia of the A. volubilis parental female, whose progeny form the
basis of this paper, has posterior apophyses which extend to the base of the
anterior apophysis, as in Ahmadi’ s (1977) illustration of A. stigmosa ; his
specimen was from North Dakota. I have seen paratypes of A. stigmosa,
described from Massachusetts, and these may represent a very pale form
of A. volubilis, but it seems likely that it is a valid coastal species. The
parental female, from the Pine Bush, Albany County, New York, resembles
the inland and not the coastal species in habitus and I have called it A.
volubilis. It compares well with Holland’s (1903) figure of Feltia [ Agrotis ]
volubilis.

The only biological information on A. volubilis is a paper by Hawkins
(1930). This is a table of ratios of the tarsal claw of various noctuids. No
information is available on the larva of A. musa or A. stigmosa.

A female A. volubilis was collected on May 28th, 1978, and ova were
obtained by the 30th. The first instar larvae were confined with Taraxacum
officinale L., Prunus virginiana L., Achillea millefolium L., and Populus
tremuloides Michx., but fed only upon the Achillea. Later instar larvae
would feed on Vaccinium vacillans Torr., and Oenothera biennis L., but

1 Published by permission of the Director, New York State Museum, Journal Series No.
306.

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NEW YORK ENTOMOLOGICAL SOCIETY

Figs. 1-3. Agrotis volubilis, Pine Bush, Albany Co., New York: 1, frontal aspect of head;
2, hypopharyngeal complex; 3, oval aspect of left mandible. Scale lines equal 0.5 mm.

only sparingly on the latter. All larvae were ultimate instars by 20 July. At
this point there was a marked decrease in feeding and the larvae crawled
beneath the litter at the bottom of the rearing tray and fed only enough to
maintain fluids and weight. This torpid state lasted through the remainder
of the summer and fall and on 20 October 100 larvae that had not been

VOLUME LXXXIX, NUMBER 2

61

Fig. 4. Agrotis volubilis , Pine Bush, Albany Co., New York: photograph of living, sixth
instar larva.

preserved were left in out-of-door cages. The larvae did not survive the
winter.

Larvae were reared in continuous darkness, interrupted by addition of
fresh leaves every two days. This procedure usually causes larvae to bypass
a diapause, and, with night feeding larvae, accelerates development. Ap-
parently this species has a summer aestivation followed by an obligatory
larval diapause to overwinter. This is supported by the spring appearance
of adults and univoltinism in nature throughout its range. Crumb (1929)
reported similar adaptive strategies in species of Feltia and in another
Agrotis species.

Agrotis venerabilis Walker (1856) and A. volubilis larvae are extremely
similar and no consistent larval mouthparts or head capsule differences were
found. The size and relation of the Abl LI pinaculum to the Abl spiracle
seem to offer reliable differentiating characters: the pinaculum and the spi-
racle are the same size in A. volubilis , that of A. venerabilis have the
pinaculum 50% larger than the spiracle; the spiracle of A. volubilis is 0.52
mm high, that of A. venerabilis is 0.36 mm high on average (the full grown
length is comparable in the larvae). Considering the above similarity in two

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NEW YORK ENTOMOLOGICAL SOCIETY

Figs. 5-6. Agrotis volubilis. Pine Bush, Albany Co., New York: 5, prothorax; 6, first
abdominal segment. Scale line equals 1.0 mm.

“distant” Agrotis species, it may not be possible to distinguish A. musa,
A. stigmosa , and A. volubilis larvae.

Larval setal nomenclature follows Hinton (1946) with the exception of the
ultraposterior coronal punctures which are labelled with an X.

First Instar Larva. — First instar larva with primary body setae tipped
with a knob; proleg Ab3 rudimentary, Ab4 one-half the size of Ab5; proleg
Ab5, Ab6, and AblO all well developed.

Sixth Instar Larva. — General (Fig. 4): Average head width 3.20 mm; av-
erage total length (fully extended) 42.0 mm; abdominal prolegs present on

VOLUME LXXXIX, NUMBER 2

63

3rd through 6th and 10th segments; setae simple; crochets a uniordinal me-
soseries; spiracle A8 0.50 mm high on average (N = 18).

Coloration (living material): Head (Fig. 1) with a brown, oblique band
from vertex and then expanding ventrally to encompass entire adfrontal
line; reticulate pattern present dorsal to band and between band and stem-
mata. Body gray-brown with conspicuous lateral series of black spots, two
on each segment comprising the spiracle and the 1st lateral (LI) pinaculum.

Head (Fig. 1): Epicranial suture 0.50 mm long; height of frons 1.03 mm.
Second adfrontal (Af2) anterior and 1st adfrontal (Afl) posterior to apex of
frons. Coronal punctures Aa, pa, pb, and 3 ultraposterior (2 visible in frontal
aspect) present as illustrated.

Mouthparts: Hypopharyngeal complex (Fig. 2): spinneret shorter than
labial palpus, apex lacking setae; stipular seta (S) at anterodorsal region of
prementum; distal region of hypopharynx bears numerous fine spines, fol-
lowed by a single row of 25-30 spines which extends to the posterior apex
of the prementum. Mandible (Fig. 3) with inner ridges distinct, lacking basal
tooth. Sixth outer tooth low, divided into smaller subteeth.

Thoracic segments: Prothorax (Fig. 5): cervical shield weakly sclerotized.
Subdorsal setae (SD1 & SD2) lacking pinaculi; prespiracular setae (LI &
L2) also lacking pinaculi. Subventral setae share common pinaculum. Meso-
and metathorax with dorsal and subdorsal setae (Dl, D2, SD2, and SD1) all
in a vertical line through center of segment and each on its own pinaculum;
SD1 peculiarly modified, surrounded by an inverted horseshoe-shaped pin-
aculum and with a very large, black setal base which is twice the diameter
of other setal bases although the seta itself is less than half the size of other
primary setae. Angle of L3-L1-L2 120°.

Abdominal segments: Abl (Fig. 6): two subventral setae (SV1 & SV3);
LI posterior to spiracle; SD2 seta reduced or absent, but setal base always
present anterodorsal to spiracle, rarely contiguous with upper margin of
spiracle; spiracle 0.52 mm high. Ab2-6 with 3 subventral setae; SD2 setal
base anterior to spiracle. Ab7&8 with only 1 seta in subventral group.

Crochets a uniordinal mesoseries: 9-15 on Ab3 (mode = 12), 12-19 (15)
on Ab4, 13-20 (16) on Ab5, 14-18 (17) on Ab6, and 17-23 (20) on AblO.
Material examined. Eighteen specimens, Pine Bush, lat. 42°34'07", long.
73°52'53", Albany County, New York, elev. 91 meters, 30 July 1978, from
ova of female collected and determined by T. L. McCabe. PI 9 and all larvae
are coded tlm 78-65.

Acknowledgments

I thank Linnea Johnson for the illustrations. Voucher specimens of the
larvae will be deposited in the New York State Museum and the United
States National Museum.

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NEW YORK ENTOMOLOGICAL SOCIETY

Literature Cited

Ahmadi, A. A. 1977. A revision and review of the North American species of Agrotis and
Feltia known to occur north of the Mexican border (Lepidoptera: Noctuidae). Ph.D.
dissertation, Cornell University, Ithaca, N.Y.

Crumb, S. E. 1929. Tobacco Cutworms. U.S.D.A. Tech. Bull. 88:1-179.

Harvey, L. F. 1874. Observations on North American moths. Bull. Buffalo Soc. Nat. Sci.,
2:118-121.

Hawkins, J. H. 1930. Tarsal claws of noctuid larvae. Ann. Ent. Soc. Amer. 23:393-396.

Hinton, H. E. 1946. On the homology and nomenclature of the setae of lepidopterous larvae,
with some notes on the phylogeny of the Lepidoptera. Trans. Roy. Ent. Soc. London
97:1-37.

Holland, W. J. 1903. The moth book. New York: Doubleday, Page & Co., 479 pp.

Morrison, H. K. 1874. Descriptions of new Noctuidae. Proc. Boston Soc. Nat. Hist. 17:131-
166.

Smith, J. B. 1910. Feltia musa, new species, Porosagrotis catenuloides , new species. Journ.
N.Y. Ent. Soc. 16:87.

Walker, F. 1856. Catalogue of the Lepidoptera Heterocera in the British Museum. 10:328.

New York State Museum, The State Education Department, Cultural
Education Center, Albany, New York 12230.

Received for publication April 9, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(2), 1981, pp. 65-79

HISTOLOGICAL CHANGES IN THE ANTENNA OF TENEBRIO
MOLITOR L. AFTER IMAGINAL ECLOSION
(COLEOPTERA: TENEBRIONIDAE)

Louis D. Trombetta and James Forbes

Abstract. — The histology of the antenna of T. molitor and the changes
which take place in its maturation from imaginal eclosion to one week of
age are compared. The cuticle, epidermis, nerves and tracheae are de-
scribed. A mesocuticle is present in the cuticle, and a blood vessel is de-
scribed for the first time in the antenna of a beetle. In the one week old
insect the endocuticle becomes thicker, the underlying epithelial cells lower
in height, the layer of nerve fibers beneath the epithelium more compact,
and the walls of the tracheal trunks and blood vessel thinner.

Only two studies include observations on the anatomy of the coleopteran
antenna: Borden (1968) on the antennae of the scolytid Ips confusus and
Chang and Jensen (1973) on the curculionid Rhabdocelus obscurus. Most
investigations of this organ have concentrated on the morphology and phys-
iology of its sensilla (Roth and Willis 1951; Slifer and Sekhon 1964; Moeck
1968; Ernst 1969; and Borg and Morris 1971). The sensory receptors of T.
molitor have been studied by Valentine (1931), Pielou (1940), Doyen (1966),
Pierantoni (1974), and Harbach and Larsen (1977a, b).

This paper is the first description of the histology of the antenna of T.
molitor and describes the changes which take place during the first week
after imaginal eclosion.

Newly emerged and one week old adults were removed from individual
cultures and immediately placed in alcoholic Bourn’ s solution where the
antennae with their musculature were removed. After fixation for 12 hours,
they were washed and stored in 80% ethyl alcohol. The antennae were
processed and sectioned by a technique previously described (Trombetta
and Forbes 1977). The sections were stained with Harris’ haematoxylin
counterstained with eosin Y, Sharp’s modification of Mallory’s triple stain
(Kennedy 1932), safranin counterstained with fast green, and an argentaffin
staining technique (Lillie 1954).

Observations

One week old adult. The antenna of T. molitor is moniliform in shape and
consists of the scape, the pedicel, and a flagellum that has nine subsegments
or annuli joined to each other by membranes. These annuli give the flagellum

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NEW YORK ENTOMOLOGICAL SOCIETY

Abbreviations: BV — blood vessel, E — epidermis, En — endocuticle, Ex — exocuticle, I — in-
tersegmental membrane, J — sensillum of Johnston’s organ, M — mesocuticle, N — nerve trunk,
S — sense cell, Sn — sensory receptor, T — trachea.

VOLUME LXXXIX, NUMBER 2

67

a segmented appearance, but, because they do not contain muscles, are not
true segments (Schneider 1964). Longitudinal sections of the antennae show
a segmental arrangement of the annuli, and in recording the observations
the older description of the antenna, that it consists of 1 1 segments, is used.
The procuticle of the adult antenna consists of three layers: the exocuticle,
the mesocuticle and the endocuticle (Fig. 1). The exocuticle is heavily scler-
otized, and in sections it is amber in color and does not readily stain. The
thickness of this layer varies in each segment from 23 fx at the base to 17
f. i at its terminal end. The only exception to this is in the distal segments
where because of the numerous sensory receptors it can be as thin as 9 /x.
In the intersegmental regions the proximal segment forms a depressed ring
into which the base of the next distal segment fits. On the inner surface of
the ring, the exterior of the exocuticle is serrated (Fig. 2).

A mesocuticle is present beneath the exocuticle from the scape to segment
ten. This layer varies in thickness, stains with safranin and eosin, and except
for some red blotches does not stain with Mallory’s stain (Figs. 1 and 2).
No mesocuticle is seen in the eleventh segment.

The endocuticle lies beneath the mesocuticle, except in the eleventh seg-
ment where it lies directly beneath the exocuticle. It stains lightly with
haematoxylin and with fast green and red with Mallory’s stain. This layer
is laminated and is thickest at the bases of segments six to eleven (Fig. 1).
Here as many as ten lamellae are visible, and it is twice as thick as the
exocuticle. In the remaining segments the thickness of the endocuticle var-
ies, but generally it is thickest in the center of the segment and is thinnest
at the articulatory regions; slightly thicker at the proximal than at the distal.
Where numerous sensory receptors are located, the endocuticle appears as
a very thin layer, about 3 /x in thickness (Fig. 3). In other regions the
dendrites of nerves traverse the thick endocuticle to innervate receptors. In
the scape the endocuticle is thin and never exceeds five lamellae in thick-
ness.

Pore canals appear under the oil immersion lens as very fine striations
extending through the endo-, meso-, and exocuticle. When stained with the
argentaffin staining technique, they appear tubular in structure, and are
straight, not helical, and end in a small bulb beneath the epicuticle.

Fig. 1. Cross section of a one week old adult antenna showing the arrangement of the
cuticular layers and the epidermis. Haematoxylin and eosin. x400.

Fig. 2. Longitudinal section through the intersegmental membrane between two antennal
segments of a one week old adult antenna. The endocuticle appears dark because it is stained
red with Mallory’s. Note the serrated edge on the depressed ring of the proximal segment
(curved arrow). The blood vessel appears as a smooth-surfaced tube (straight arrow). Mallory’s
stain. x400.

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69

The inter segmental membrane between the antennal segments is a thin,
unsclerotized membrane, which appears white in color when unstained (Fig.
2). It stains with fast green and appears as a single layer continuous with
the endocuticle of the adjoining segments. Mallory’s stains two layers of the
intersegmental membrane, an exterior layer which stains red and the interior
layer which stains blue. In some of these intersegmental membranes random
areas of the interior layer have an amber color not unlike that of the exo-
cuticle.

The epidermis underlying the cuticle is composed of a simple epithelium
(Fig. 1) that varies from columnar to squamous depending on location and
density of the nervous tissue. In the proximal segments, where the sensory
receptors and the underlying nerve fibers are not numerous, the cells are
cuboidal or columnar. In the distal segments where these nervous elements
are numerous the cells are lower in height and are scattered among the
sensory cells. At the bases of the distal segments the cells are cuboidal and
appear small in comparison to the overlying cuticle. In the intersegmental
membranes the cells are also cuboidal. These epidermal cells have a large,
centrally located nucleus with peripheral chromatin, and their cytoplasm is
extremely basophilic in staining reaction. These cells stain strongly with
safranin and with Mallory’s stain they appear brown yellow in the proximal
segments but brown red in the distal.

In some regions of the antenna, between the epidermal cells and the
endocuticle a very thin, blue green layer can be seen in sections stained
with Mallory’s stain. Beneath the epidermis the basement membrane is seen
as a fine, blue green layer when stained with Mallory’s stain. However,
when nerve fibers and fine tracheae underlie the epidermal cells, the base-
ment membrane is not evident.

The sensory cells of the adult antenna are typical for insects. They have
short, distal dendrites and long axons that join to form large branches. The
axons and dendrites stain with fast green and stain light blue with Mallory’s
stain. The cytoplasm of the cell body of these cells stains deep red with
safranin and blue with Mallory’s stain. The nucleus stains darker than the
cytoplasm and contains peripheral chromatin. Sensory cells of the receptors
are found either singly or in clusters (Figs. 3, 4) throughout the antenna.

Fig. 3. Cross section through numerous sensory receptors in a distal segment of a one week
old adult antenna. Note the elongated and scattered epidermal cells (straight arrows) and a
nerve branching to the sense cells (curved arrow). The endocuticle appears as a thin layer.
Mallory’s stain, x 1,000.

Fig. 4. Longitudinal section through a distal segment of a one week old adult antenna
showing a branch of the nerve trunk (straight arrow) extending toward an aggregation of
sensory receptors. Mallory’s stain. x400.

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VOLUME LXXXIX, NUMBER 2

71

A large antennal nerve extends from the deutocerebrum of the brain to
the antenna. In the scape it innervates the intrinsic antennal muscles and
then divides to form two nerve trunks that extend up the antenna and end
in terminal arborizations in the eleventh segment. These nerve trunks give
off many fine branches which innervate the solitary sensory receptors, while
they extend up toward the distal segments. In these distal segments where
there are aggregations of sensory receptors, larger nerve branches split from
the antennal nerve trunks and innervate them (Figs. 4, 5). Each nerve trunk
is surrounded by a thin nerve sheath with flat nuclei (Fig. 6). Sections of
the nerve trunks show an intense safranin staining central area, that stains
darker than the peripheral nerve sheath nuclei.

Johnston’s organ can be seen in cross sections of the pedicel as clusters
of sensory cells encircling the nerve trunks, the blood vessel and the tra-
cheae (Fig. 7). These clusters of sensory cells are attached by their axons
to branches of the nerve trunks, and their dendrites extend into the inter-
segmental membrane between the pedicel and third segment (Fig. 8).

The nervous tissue in the antenna of the one week old adult is associated
with the epidermis in three ways. First, sensory cells may lie next to epi-
dermal cells with their axonal fibers extending to an antennal nerve branch
as in the case of a solitary receptor or a small aggregation of receptors (Fig.
3). Second, a nervous layer lies immediately beneath the epidermal cells.
This layer consists of nerve branches extending from the nerve trunks and
from which dendrites extend to the receptors (Fig. 4). Third, nerves have
almost completely displaced the epidermal cells as in the eleventh segment.

A large trachea enters the antenna, and in the third segment it bifurcates.
These two tracheae course through the lumen of the antenna and send tra-
cheoles to the epidermal cells and to the sensory cells (Figs. 6, 7). The tra-
cheoles are extremely numerous in regions where there are numerous re-
ceptors. They vary in size and some of the smallest lie against the basement
membrane of the epidermis. They give the appearance of being rigid and
circular in shape or collapsed. The tracheae are composed of extremely thin
cells with few nuclei and these protrude into the antennal lumen. Internal
to this epidermal lining a very thin cuticle with taenidia surrounds the tra-
cheal lumen (Figs. 2, 6).

The blood vessel is found among the nerve trunks and tracheae and ex-
tends straight through the antenna with no branches. It is always closely

Fig. 5. Oblique section through the last antennal segment of a one week old adult antenna
showing the arborization of an antennal nerve. Argentaffin stain. x80.

Fig. 6. High power view of a cross section of a one week old adult antenna showing the
arrangement of nerve trunks, tracheae and blood vessel. Arrows indicate the nuclei of the
nerve sheath and tracheae. Haematoxylin and eosin. xl,400.

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73

associated with one of the nerve trunks and usually lies directly against it.
The diameter of the blood vessel varies throughout the antenna, but it is
always largest in the third segment. No pores or openings in the vessel were
seen, not even at the tip. The wall of the blood vessel has many nuclei and
generally appears thicker than that of the tracheae.

Newly emerged adult. — The antenna of the newly emerged adult is not
heavily sclerotized. The exocuticle ranges in thickness from 23 /jl at the
bases of the antennal segments to 17 /x in the terminal regions of the prox-
imal segments and 14 /jl in the terminal regions of the distal segments. This
layer is similar to that of the one week old adult in that it is thinnest where
there are aggregations of sensory receptors. In unstained sections its color
is light yellow in the proximal segments and amber in the distal regions
(Figs. 9, 10). Only when the antenna is stained with Mallory’s stain can light
red blotches be seen in the exocuticle. The mesocuticle appears as a very
thin layer throughout the antenna when stained with safranin and eosin but
not with Mallory’s stain (Figs. 9, 10). The endocuticle ranges in thickness
from 2 /jl to 3 /a. It is thickest at the bases of the segments as well as in the
terminal regions of the proximal segments. It is present as a very thin layer
in areas of the antenna where there are aggregates of sensory receptors. It
stains with haematoxylin, fast green, and red with Mallory’s stain. Where
epidermal cells have pulled away from the endocuticle in sections of the
newly emerged adult, its inner edge is highly irregular but fits into the sur-
face contours of these cells. When this artifact is seen in sections of the one
week old adult beetle, the inner edge of the endocuticle is relatively smooth.
The pore canals are present and are similar in arrangement and position to
those in the cuticle of the one week old adult beetle.

The cuticle of the intersegmental membrane is thinner than that of the
segments and consists of two layers. The inner layer, 10 [± thick, stains blue
with Mallory’s stain while the outer, only 3 fx thick, ranges in color from
clear, to faint purple, to dark purple. The cuticle of this membrane, when
stained with fast green, appears to be a single layer.

The epidermis throughout the antenna is composed of tall columnar cells
that narrow slightly at their bases (Figs. 9, 10). In the proximal segments
the cells are taller than the thickness of the cuticle, and in the basal regions

Fig. 7. Cross section of the pedicel of a one week old adult showing the sensory cells of
Johnston’s organ (straight arrows). Curved arrows point to branches of the nerve trunk. Mal-
lory’s stain. x540.

Fig. 8. Longitudinal section through the pedicel of a one week old adult showing sensory
cells of Johnston’s organ. Arrow indicates attachment of the fibers of these cells to the inter-
segmental membrane between the pedicel and the third segment. Mallory’s stain, x 1,000.

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Fig. 11. High power view of a cross section of the antenna of a newly emerged adult
showing the arrangement of the nerve trunks, tracheae and blood vessel. Straight arrows
indicate nuclei of these various structures and the curved arrow the cuticle of the trachea.
Haematoxylin and eosin. x 1,000.

of the distal segments these cells are as tall as the thickness of the cuticle.
In the terminal region of the distal segments they are taller than the cuticle.
The epidermis is never displaced by nervous tissue even in areas where
there are numerous sensory cells. These cells have a large, centrally located
nucleus that has both peripheral chromatin and large, round, dense chro-
matin granules scattered throughout the karyoplasm. The cells stain intense-
ly with safranin and brown yellow to brown red with Mallory’s stain. The
cytoplasm close to the endocuticle is filled with basophilic granules which
also stain faintly blue with Mallory’s stain. A very thin basement membrane
is visible, which stains faintly blue green with Mallory’s stain. The epider-
mal cells of the intersegmental membrane are very small, cuboidal in shape,

Fig. 9. High power view of a cross section of the antenna of a newly emerged adult showing
the arrangement of the cuticular layers and the epidermis. Haematoxylin and eosin. x 1,000.

Fig. 10. Longitudinal section of the pedicel of a newly emerged adult showing the arrange-
ment of the antennal structures and sensory cells of Johnston’s organ. The arrow points to a
nerve innervating these sensory cells. Haematoxylin and eosin. x400.

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One Week Old

Newly Emergfd

Exocuticle

does not stain

generally does not stain
but shows some light
red blotches with
Mallory's stain

Mesocuticle

safranin ,

eosin, red blotches
with Mallory's stain

safranin ,
eosin

Endocuticle

haematoxy lin ,
fast green,
red with
Mallory's stain

haematoxy lin ,
fast green,
red with Mallory's
stain

Intersegmental

fast green,

fast green,

Membranes

red and
blue with
Mallory's stain

blue and purple
with Mallory's
stain

Epidermis

safranin ,

brown yellow

and brown red

with Mallory's stain

safranin ,
brown yellow and
brown red with
Mallory's stain

Nerves

soma: red with

safranin and blue
with Mallory's stain

axons and dendrites:
fast green, light
blue with Mallory's
stain

soma, axons and
dendrites: safranin,

brown yellow and
brown red with
Mallory ' s stain

Table 1. Summary of staining characteristics of antennal structures between one week old
and newly emerged adults.

and have a large, centrally located nucleus, similar to the other epidermal
nuclei. The basophilic area above the nucleus is also present but it is not as
clearly defined as in the cells beneath the endocuticle of a segment. The
basement membrane is not discernible beneath these cells.

The central body and axons of the sensory cells resemble the bipolar cells
of the one week old adult, except that the dendritic processes are thicker
and not as long. In the regions where there are aggregations of sensory
receptors the dendrites are very short, while the dendrites for the solitary
sense receptors are longer and more closely resemble the dendrites of the
older beetle. The cuticle at the apex of the receptor surface of the newly
emerged adult is not as sharply defined but appears more blunted than in
the one week old adult. The nerve trunk (Fig. 1 1), the fibers, and the sensory
cells all stain readily with safranin and with Mallory’s stain, similar to the
epidermal cells. At this age there is no way to distinguish nerve cells from
epidermal cells other than position and shape.

In the newly emerged adult the nerve trunks and branches are more loose
in their arrangement; they are not as compact as are those in the older
beetle. However, they are well developed, and many fine nerve branches
extend to the sensory cells. Johnston’s organ is fully developed in the newly
emerged adult (Fig. 10).

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77

The tracheae in this stage are arranged in the same manner as are those
in the one week adult. Their walls are thicker, the epithelial cells are taller
and the nuclei are larger and not as flat as are those of the older beetle. The
tracheal cuticle is clearly visible but its inner surface is not as smooth as
that of the older beetle (Fig. 1 1). The wall of the blood vessel is thicker than
that of the one week old beetle, and the nuclei do not bulge into the antennal
lumen (Fig. 11). Throughout the antennal lumen, but not within the blood
vessel, there is a granular substance which stains with fast green and blue
with Mallory’s stain.

Discussion

This study shows that the procuticle of the antenna of Tenebrio has three
layers, and that the mesocuticle is deposited before the endocuticle. The
variations of staining characteristics of the different cuticular layers (Table
1) indicates changes in chemical composition. Wigglesworth (1948), Dela-
chambre (1967), and Zlotkin and Levinson (1969) have described only two
layers, the exocuticle and endocuticle, in the abdomen of Tenebrio. The
cuticle of the pupal abdomen was described by Delachambre (1967), Zlotkin
and Levinson (1969), and Caveney (1970). Both Delachambre and Caveney
mention that the procuticle is composed of two layers, and that the endo-
cuticle is completely deposited within twenty-four hours of pupation.

The epidermis of the antenna undergoes extreme changes during its de-
velopment. After secretion of the adult cuticle, the epidermal cells are re-
duced in size and their reduction accompanies the increased development
of the nervous tissue. The large, dense, chromatin granules of the epidermal
cells of the developing antenna have been seen also in the cells of the
abdominal cuticle (Zlotkin and Levinson 1968). The basement membrane is
never clearly or completely seen in the antenna of Tenebrio because tra-
cheae and nerve fibers adhere closely to the epidermis. The basement mem-
brane is similar to that in the antenna of Bombyx mori (Schneider and
Kaissling 1959).

Johnston’s organ in Tenebrio is similar to the one described for the beetle,
Melolontha vulgaris (Child, 1894). Neither Borden (1968) nor Chang and
Jensen (1973) mention this organ in the antennae of the beetles they de-
scribe.

Except for the changes in cell thickness, the histology of the tracheae
found within the antennal lumen of Tenebrio is typical for insects. An an-
tennal blood vessel is described for a coleopteran for the first time and
appears similar to antennal blood vessels in other insects. The dilated region
of the blood vessel in the third segment may represent an accessory pul-
sating organ. Schneider and Kaissling (1959) describe a blood vessel in the
antenna of Bombyx mori that had an opening only at its distal end. The lack
of any openings in the blood vessel in Tenebrio might be explained on the

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NEW YORK ENTOMOLOGICAL SOCIETY

basis that small pores may be present in the wall which cannot be resolved
by the light microscope. The close association of the blood vessel and the
antennal nerve trunk suggests either mutual support or that the blood vessel
is innervated.

Literature Cited

Borden, J. H. 1968. Antennal morphology of Ips confusus (Coleoptera: Scolytidae). Ann.
Entomol. Soc. Amer. 61:10-13.

Borg, T. K. and D. M. Morris. 1971. Ultrastructure of sensory receptors on the antennae of
Scolytus multistriatus (Marsh.) Z. Zellforsch. 113:13-28.

Caveney, S. 1970. Juvenile hormone and wound modeling of Tenebrio cuticle architecture.
J. Insect Physiol. 16:1087-1107.

Chang, V. C. S. and L. Jensen. 1973. Antennal morphology of the New Guinea sugarcane
weevil, Rhabdocelus obscurus. Ann. Entomol. Soc. Amer. 66:117-121.

Child, C. M. 1894. Beitrage zur Kenntnis der Antennalen Sinnesorgane des Insekten. Zeitschr.
Wiss. Zool. 58:475-528.

Delachambre, J. 1967. Origin et nature de la membrane exuviale chez la nymphe de Tenebrio
molitor. Z. Zellforsch. Mik. Anat. 81:114-134.

Doyen, J. T. 1966. The skeletal anatomy of Tenebrio molitor (Coleoptera: Tenebrionidae).

Misc. Pub. Entomol. Soc. of Amer. 5:103-150.

Ernst, K. D. 1969. Die Feinstruktur von Riechsensillen auf der Antenne des Aaskafers, Nec-
rophorus. Z. Zellforsch. Mik. Anat. 94:72-102.

Harbach, R. E. and J. R. Larsen. 1977a. Humidity behavior and the identification of hygro-
receptors in the adult mealworm, Tenebrio molitor. J. Insect Physiol. 23:1121-1134.

and . 1977b. Fine structure of antennal sensilla of the adult mealworm beetle,

Tenebrio molitor L. (Coleoptera: Tenebrionidae). Int. J. Insect Morphol. Embryol.
6:41-60.

Kennedy, C. H. 1932. Methods for the Study of the Internal Anatomy of Insects. Mimeo-
graphed by H. L. Hendrick. Columbus, Ohio. 103 pp.

Lillie, R. D. 1954. Histopathologic Technic and Practical Histochemistry. The Blakiston Com-
pany, New York. 715 pp.

Moeck, H. A. 1968. Electron microscope studies of antennal sensilla in the ambrosia beetle,
Trypodendron lineatum (Oliver) (Scolytidae). Can. J. Zool. 46:521-575.

Pielou, D. P. 1940. The humidity behavior of the mealworm beetle, Tenebrio molitor L. II.

The humidity receptors. J. Exp. Bio. 17:295-306.

Pierantoni, R. 1974. Electron scanning microscopy of the antennal receptors of Tenebrio
molitor. A stereoscopic analysis. Cell Tissue Res. 148:127-142.

Roth, L. M. and E. E. Willis. 1951. Hygroreceptors in Coleoptera. J. Exp. Zool. 117:451-
487.

Schneider, D. and K. Kaissling. 1959. Der Bau der Antenne des Seidenspinners Bombyx mori
L. III. Das Bindegwebe und das Blutgefass. Zool. Jahrb. Abt. Anat. Ontog. Tierre.
77:111-132.

. 1964. Insect antennae. Ann. Rev. Entomol. 9:103-122.

Slifer, E. H. and S. S. Sekhon. 1964. Fine structure of the thin-walled sensory pegs on the
antenna of a beetle Popilius disjunct us (Coleoptera: Passalidae). Ann. Entomol. Soc.
Am. 57:541-548.

Trombetta, L. and J. Forbes. 1977. A double infiltration technic for serial sectioning of heavily
sclerotized insects. J. Histotech. 1:25-26.

Valentine, J. M. 1931. The olfactory sense of the adult mealworm beetle Tenebrio molitor L.
J. Exp. Zool. 58:165-228.

VOLUME LXXXIX, NUMBER 2

79

Wigglesworth, V. B. 1948. The structure and deposition of the cuticle in the adult mealworm,
Tenebrio molitor L. (Coleoptera). Quart. J. Micro. Sci. 89:197-217.

Zlotkin, E. and H. Z. Levinson. 1968. Influence of cecropia oil on the epidermis of Tenebrio
molitor. J. Insect Physiol. 14:1195-1204.

and . 1969. Influence of cecropia oil on the cuticle of Tenebrio molitor. J. Insect

Physiol. 15:105-110.

(LDT) College of Pharmacy, St. John’s University, Jamaica, New York
11439; and (JF) Department of Biological Sciences, Fordham University,
Bronx, New York 10458.

Received for publication June 6, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(2), 1981, pp. 80-88

BREEDING EXPERIMENTS WITH MORPHO PELEIDES INSULARIS
(LEPIDOPTERA: NYMPHALIDAE) IN TRINIDAD, W.I.

Malcolm Barcant

Abstract. — A breeding program of Morpho peleides insularis Fruhstorfer
in Trinidad led to successful rearing of large broods of this butterfly on
Paragonia pyramidata (Bignoniaceae). Crossing with M. achillaena Hiib-
ner from Rio de Janeiro, Brazil, gave good fertility to the F4, while crossing
with Morpho achdles Fruhst. from Belem, Para, Brazil did not lead to
generations beyond the FI. The appearance of melanic forms of peleides,
which occurred on two separate occasions in captivity, was followed by
means of selective breeding; this revealed that the melanism had a genetic
basis but was probably also influenced by the physical environment (espe-
cially excessive sun and heat exposure of juveniles in the breeding cage),
and that at least two melanic characters (and possibly three) were segre-
gating independently.

Morpho peleides insularis Fruhstorfer, the only member of its family in
Trinidad,1 is not rare on the island. It is well distributed in all forested areas,
particularly along the moist valleys and footpaths of the mountainous North-
ern Range (Barcant 1970). Very little study has been given to the life history
of this species in Trinidad; with the exception of a paper by Stollmeyer
(1932), giving details of the early stages obtained from two eggs extruded
from the abdomen of a captured female, no further work has been published.
Many biological data have recently been published, however, on central
American M. peleides (Young and Muyshondt 1973; Young and Thomason
1974; Young 1974).

In January 1964, I became interested in breeding Trinidad M. peleides in
captivity, under near-natural conditions. The following problems had to be
solved initially:

a. The butterfly had never been kept alive in captivity in Trinidad; its
adaptability to such a life was speculative.

b. The female was a known recluse; only a few existed in collections. It
was uncertain whether sufficient numbers in gravid condition could be lured
to bait, or whether they would oviposit in captivity.

1 I have followed Fruhstorfer (in Seitz, 1907) for the nomenclature of Morpho peleides
insularis. I consider LeMoult's application of the term tobagoensis [LeMoult & Real (1962)]
to be a misnomer; his recording of subspecies of M. trojana and M. corydon in Trinidad is
also not in accordance with fact.

VOLUME LXXXIX, NUMBER 2

81

c. The host plant, Paragonia pyramidata Rich. (Bignoniaceae) was
known to only two people in Trinidad (it was incorrectly quoted in Stoll-
meyer’s paper), and unknown to me except by vague description — “a forest
wild vine with purplish young leaves and not easy to detect.”

By mid- 1965 these obstacles had been overcome. A fine mesh cage 10' x
5' x 8' was built; the host plant, located and planted within it, grew rapidly
in leafy abundance; and ‘‘Emperor Valley” near Port-of- Spain had produced
numerous gravid females caught on rotting breadfruit, all willing to lay pro-
fusely in captivity. At the end of 1965 two good broods of over 80 adults
emerged and the way was open to further study. In February 1966 mating
of M. peleides in captivity was achieved for the first time in Trinidad; this
led to a successful program of breedings and crossings. Further breeding
space was also added with another cage of 20' x 10' x 8'.

In early 1966, a visit of Keith Brown, Jr. to Trinidad paved the way for
trials in cross-breeding M. peleides with M. achillaena Hiibner from Rio
de Janeiro. With his help in the supply of live males from that area, these
crossings were performed successfully during 1966. During the year 1967,
repeated broods of local peleides were raised in one cage and crossing
experiments with achillaena were continued in the other (until June). To-
wards the end of the year, the smaller cage was devoted to eventually
successful crossings between M. peleides and M. achilles amazonica Fruhs-
torfer, from the lower Amazon (Belem). Although the progeny of the latter
cross proved sterile, those from the achillaena crosses were taken through
four generations. In my opinion, this offers strong evidence that M. peleides
and M. achillaena are one species, with morphological differences merely
the result of geographical separation, without effect on potential for inter-
breeding. The relation of both to Amazonian M. achilles must still remain
an open question, in view of the negative results of the single experiment
performed.

Emergence of a melanic form of m. peleides. — In September 1966, dur-
ing the emergence of the third brood of inbred peleides, three males ap-
peared (on 20/IX, 27/IX and 29/IX) with the following characteristics:

a. Total absence of all white spots in the black forewing submarginal area
of the upperside (marginal white spots were still present round the edge of
both wings).

b. Total absence of red spottings and cream bands in the submarginal
areas of both wings, on the underside.

I expected to get females of this melanic form in the brood, but none
emerged, so I was unable to select a melanic pair for inbreeding.

An attempt was made to mate one melanic male with a normal female.
This proved unsuccessful, and the event passed by with a notation, the
other two melanic males being added to my collection.

No further melanic peleides emerged at any time during 1966-1967; it was

All specimens figured were reared in captivity at 19 San Diego Park, Diego Martin, Trinidad,
and were selected from Brood 3 described in the text, between November 22, 1968 and January
10, 1969; they are now in the author’s collection.

VOLUME LXXXIX, NUMBER 2

83

not until early 1968 that melanism again appeared in captive Trinidad Mor-
pho.

In March 1968, with the host-plants fully refoliated in the now empty
larger cage (after the crossing experiments), five gravid females were cap-
tured in the field and set for oviposition, with a full diet of overripe breadfruit
(this sweet smelling succulent is abundant in Trinidad, and forms an ideal
diet in captivity for all indigenous sugar suckers). Within a week these
females had produced 200 eggs, 180 of which hatched. From this successful
brood, there emerged 1 1 melanic males and 10 melanic females during June
2 to 25. The following points were noticeable in these specimens:

a. The males were constant among themselves and identical with the
original melanic males produced in September 1966 (Figs. 6, 10).

b. The females, also constant, retained an inner submarginal row of white
elliptical spots on the forewing, not as pronounced as in normal females.
The outer marginal row was absent. The underside submarginal region of
both wings showed the same absence of the three red and cream bands as
in the male (Figs. 7, 11).

The constancy of the 21 melanic forms suggested the presence of a gene
linkage of upperside submarginal white spots and underside submarginal
cream and red bands. Selective breeding was carried forward successfully
with the melanic individuals from this brood.

Further broods of melanic m. peleides. — On June 9, 1968, four melanic
males were put into the cage with four normal females. Matings took place
between June 10 and June 14. On June 15, two melanic females were put
in with the melanic males. Matings took place within the next two days.
Unfortunately, due to space and host-plant limitations, the two groups could
not be segregated, and the caterpillars were reared together, leaving a pure
black selection for the third brood.

In a total of 83 adults obtained from this “joint venture,” 16 (about 20%)
were melanic, again constant in phenotype. Due to the mixed nature of this
brood little importance can be given to this figure, but it might indicate that

Figs. 1-7. (uppersides) Fig. 1. n — Normal male as found in the field; arrows show marginal
intervenal white spots on both wings doubled on the hindwing (best noted against the light
background by the deeper serrations of the wing margins), and seven submarginal white spots
on the forewing. Fig. 2. n — Normal female as found in the field; arrows show marginal inter-
venal white spots as in the male but more pronounced, and eight submarginal spots on the FW,
larger than those of the male. Fig. 3. nrs — “Semi-normal’' male, with submarginal FW spots
reduced to four (two faint), and intervenal spots present on both wings but not pronounced;
the underside is normal. Figs. 4-5. b — Totally black female and male respectively, with both
marginal and submarginal spots absent; in the female the inner discal elliptical spots are present
but reduced in size. Figs. 6-7. bw — Black male and female respectively, similar to 4-5 except
that the intervenal marginal spots are present as in 1-3.

Figs. 8-13. (undersides) Figs. 8-9. n — Normal male and female respectively, with three
full submarginal cream and red bands. Figs. 10-11. bw — Black male and female respectively;
the three submarginal bands are entirely missing except for a trace at the female HW apex.
Figs. 12-13. bc — Totally black (upperside) male and female, respectively; on the underside
only the inner of the three submarginal bands remains. Form BCW is similar to 12-13, but
with intervenal marginal white spots present.

VOLUME LXXXIX, NUMBER 2

85

genetic factors were showing an influence in the appearance of the char-
acter. The 16 melanics broke evenly between the sexes and all were used
for a third selective brood.

Five matings were obtained between August 31 and September 15, 1968,
and egg-laying extended throughout September and the first week of Oc-
tober. No account can be given for the reluctance of three of the females
to mate. Surprising deviations appeared in the resulting progeny, which
emerged between November 22, 1968 and January 10, 1969. The six phe-
notypes which appeared in this brood are as follows:

a. Normal: N (Figures 1, 2, 8 and 9) — normal peleides.

NRS (Figure 3) — near-normal peleides with fore wing sub-
marginal white spots reduced in size and number; other-
wise normal with intervenal spots and all underside
submarginal bands and red spottings present; this
phenotype only in males.

b. Melanie: BW (Figures 6, 7, 10, and II) — black form with small

intervenal white spots (doubled in female) on the mar-
gins.

B (Figures 4 and 5) — totally blackened form, without the
intervenal white marginal spots.

BCW (not figured) — black form with white intervenal
spots, and one (the inner) instead of three cream bands
in the submarginal area of the underside; red spottings
absent.

BC (Figures 12 and 13) — same as BCW but intervenal
marginal white spots absent.

The following chart indicates the numbers obtained for each phenotype:

Phenotype

Male

Female

Total

N

30

44

74 ]

1

NRS

18

18 j

> Normal (69%)

BW

11

6

17

B

9

5

14

BCW

1

2

3

= Melanie (31%)

BC

3

5

8

Totals

72 (57%)

62 (43%)

134

The percentage of melanism increased to 31% in this second selective
breeding. While the submarginal spots of the forewing upperside and the
two outer submarginal bands of the underside were always absent in the

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NEW YORK ENTOMOLOGICAL SOCIETY

melanic types produced, two types of deviations appeared in the presence
or absence of intervenal white marginal spots (W) and the third (inner) cream
band on the underside (C). Total disappearance of red from the underside
marginal area was observed in all the melanic forms.

The normal cream and red marginal bands expressed themselves as 27%
with a single cream band (the inner of the three; BC) and 73% with all three
bands completely absent (B). The intervenal white spots divided nearly
equally between present (BW, 48%) and absent (B, 52%). The fact that the
two characters can be absent together (33%), individually present (W =
41%, C = 19%), or present together (CW = 7%), indicates independence of
segregation of the respective genes concerned with cream underside bands
and white intervenal spots.

At this stage, it would have been desirable to have unlimited space and
host plant available to carry on a growing number of possible combinations.
Unfortunately this could not be done, but a fourth brood was obtained on
very limited food supply from matings between BW pairs, with the following
results:

Phenotype

Male

Female

Total

B + BW

8

6

14 1

BC + BCW

4

1

Melanic (50%)

N

12

7

19

Normal (50%)

Totals

24 (63%)

14 (37%)

38

The brood was a small one, and an attempt to score the intervenal spots
versus bands was not made, but it may be noted that the single cream band
(C) in melanics was again present at 26%.

Due to a change in my residence and the consequent disruption of the
breeding experiments, the program had to be discontinued at this stage.

Conclusions and Discussion

I realize that these initial experiments have only scratched the surface of
the phenomenon of melanism in Trinidad M. peleides; in the absence of
more detailed selections and further broods, under controlled environmental
conditions, definite conclusions are difficult to make. The evidence from the
four broods does indicate, however, that:

a. Melanism does occur, potentially, in Trinidad peleides , although it has
never, to my knowledge, been recorded in the field; it was discovered
for the first time in breeding experiments in September 1966.

b. Genetic factors play a predominant role in the expression of melanism
in Trinidad peleides : it seems probable that a 100% pure melanic strain
could be developed by repeated selective breeding.

VOLUME LXXXIX, NUMBER 2

87

c. Selective breeding does bring out more than one expression of melanism,
with varying combinations of the presence or absence of the three normal
pattern features.

d. Morphologically, in habits and in lifespan, the early stages of the melanic
peleides are identical to those of the normal form.

In the absence of more detailed experiments, the initial cause for the
development of melanism in the captive populations can only be speculative.
In the face of a 10% development in the first brood, from normal parents
obtained directly in the field, where the phenomenon has not been recorded,
it is necessary to consider that the color modification may be due to envi-
ronmental stress under captive conditions.

In all the broods, it was observed that the host plant climbing against the
mesh roof of the cage was mostly sparse in growth, not only letting through
sunlight but in many areas having only a single thin layer of older leaves
exposed to the direct rays of the sun. Caterpillars which in the field would
invariably be well shaded by thick overhead forest growth throughout their
five instars, therefore found themselves exposed to a much higher degree
of direct sunlight than they would normally experience. In many cases, they
found large leaves at lower levels on the sides of the cage, where they could
be sheltered under more normal conditions of temperature and shade, but
some persisted at higher levels near the roof of the cage. In the heat of the
day, these could be seen hanging downwards by their hind pair of fleshy
prolegs, possibly in response to heat stress or to escape the high temperature
and sun’s rays which were penetrating and heating the leaves below which
they had been resting.

It is possible that the existence of such abnormal conditions of heat and
light during the first four instars, perhaps to a lesser degree in the fifth when
the caterpillars became gregarious and hid along the stems of the vine, but
also in the pupal stage when they were often on the cage roof again (though
in seclusion), could affect the expression of some genes controlling the
marginal spots and underside bands, and that this effect could be continued
increasingly by selective breeding in successive generations.

The proportional numerical increase in melanic emergence through selec-
tive breeding suggests that recessives, normally obscure in the field, were
concentrated in the later broods, though their expression may also have
been triggered by abnormal environmental conditions. It is difficult, how-
ever, to propose a mechanism whereby additional heat, sunlight and ex-
posure could promote melanism in the adults.

Acknowledgments

I would like to thank Clive Urich of Sangre Grande, Trinidad, for valuable
assistance in host plant identification and provision of live plant material;
Dr. Lee D. Miller of the Allyn Museum, Sarasota, Florida, for reading the

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NEW YORK ENTOMOLOGICAL SOCIETY

text of this paper and suggesting alterations; and Dr. Keith S. Brown, Jr.,
of Campinas, Sao Paulo, Brazil, for live specimens supplied for cross-breed-
ing, advice given through many years, and editing of this paper.

Literature Cited

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

Fruhstorfer, H. 1907. Morphidae. In Seitz, A. (ed.), Die Grossschmetterlinge der Erde, vol.
V. Kernen, Stuttgart, pp. 333-356.

LeMoult, E. and P. Real. 1962. Les Morpho d’Amerique du Sud et Centrale, Historique -
Morphologie - Systematique. Paris, Editions du Cab. Ent. E. LeMoult (Suppl. a Nov-
itates Entomologicae), xiv + 296 pp., 92 plates.

Stollmeyer, C. R. 1932. Notes on the life history of Morpho peleides insularis Fruhstorfer.
J. New York Entom. Soc ., 40:523-527.

Young, A. M. 1974. The rearing of the neotropical butterfly Morpho peleides (Nymphalidae)
on peanuts. J. Lepid. Soc., 28:90-99.

Young, A. M. & A. Muyshondt. 1973. Notes on the biology of Morpho peleides in Central
America. Carib. J. Sci., 13:1-49.

& J. H. Thomason. 1974. The demography of a confined population of the butterfly

Morpho peleides during a tropical dry season. Stud. Neotrop. Fauna, 9:1-34.

731 West Daughtery Road, Lakeland, Florida 33805.
Received for publication June 23, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(2), 1981, pp. 89-97

PARASITISM OF SINGLE AND MULTIPLE EGG CLUSTERS OF
EUPHYDRYAS PHAETON (NYMPHALIDAE)

Nancy E. Stamp1

Abstract. — The effect of egg numbers on parasitism of Euphydryas phae-
ton (Nymphalidae) was examined by comparing single and multiple egg
clusters on leaves. Some 65 to 80% of the clusters lost eggs to parasitism
and predation, with 5 to 9% of the eggs lost per cluster. For total clusters,
single clusters and multiple clusters, the numbers of parasitized clusters
were significantly different between years. It appeared that when the overall
level of parasitism was greater one year, multiple clusters were attacked
more frequently than single clusters. However, the number of egg clusters
per leaf had no effect on the number of parasitized eggs per cluster. The
sequence of egg clusters deposited on a leaf did not affect the level of
parasitism and number of parasitized eggs per cluster. Thus, these egg clus-
ters composed of several hundred eggs each lost only a small fraction of
eggs to parasitoids and predators and did not appear to benefit from aggre-
gation of egg clusters.

Introduction

The patterns of egg distribution by insects are varied; with some insects
depositing eggs singly, others depositing many small clusters of eggs, and
others laying a few large clusters. Eggs deposited in clusters may have
higher survivorship than those laid singly (Stamp 1980a). Furthermore, para-
sitoids may differ in their efficiency at exploiting various sizes of egg clusters
(Hokyo and Kiritani 1966). Egg parasitoids specializing on a host depositing
eggs singly, attacked fewer eggs when those eggs were clustered, as a con-
sequence of reduced encounters with total host eggs and limited mature
eggs for oviposition (Hirose et al. 1976). In contrast, for parasitoids adapted
to eggs occurring in clusters, large clusters were found more frequently and
had higher parasitism than small clusters (Lyons 1962). This may be a con-
sequence of the searching behavior of animals which concentrate their ef-
forts in areas where they recently found prey (Laing 1937, Tinbergen et al.
1967, Royama 1970). Also, as immobile host patches, large clusters may be
advantageous to parasitoids and predators as the overall distances between
clusters increases (e.g. Janzen 1975). Thus, egg parasitoids searching for
large egg clusters may deposit all or most of their eggs in one or a few places

Department of Zoology, University of Florida, Gainesville, Florida 32611.

90

NEW YORK ENTOMOLOGICAL SOCIETY

Fig. 1. Egg clusters of E. phaeton. A. Five clusters deposited on the same leaf by five
females. Scale is in cm. B. Eggs are laid in layers. C. The egg cluster on the right is 9 or more
days old and has some parasitized (black) eggs. The egg cluster on the left is 4 days old.

and, thus, benefit by spending less time searching and exposing themselves
to mortality factors.

Whether the hosts or parasitoids benefit more from clustered eggs may
be examined in differing but naturally-occurring group sizes of eggs. The
Baltimore checkerspot ( Euphydryas phaeton Drury: Nymphalidae) is suit-
able for this study because it deposits large egg clusters (274 eggs ±23 SD

VOLUME LXXXIX, NUMBER 2

91

per cluster, Stamp 1980b). Frequently females lay eggs with those of other
females (Fig. 1A). Of 483 leaves with egg clusters, 18% had multiple (two
or more) egg clusters (Stamp 1980b). The eggs are parasitized by an unde-
scribed trichogrammatid wasp (David Vincent and Carll Goodpasture, un-
published). Thus, the advantages of clustering eggs can be evaluated by
examining parasitism of single and multiple egg clusters on leaves. For com-
parison, numbers of missing eggs as an indicator of egg predation were
determined also.

Materials and Methods

A population of E. phaeton was observed at the Conservation and Re-
search Center of the National Zoological Park at Front Royal, Virginia in
1978 and 1979. Turtlehead ( Chelone glabra L.: Scrophulariaceae) the larval
host plant grew in wet meadows along one drainage. In each year about 500
egg clusters were deposited in this study area. Egg clusters consisted of
eggs deposited usually in two layers, with a typical nymphalid egg shell and
moderate adhesion to the leaf (Fig. IB).

New egg clusters on tagged plants were photographed with Kodachrome
film. The bright yellow coloration of new clusters differentiated them from
older (5 to 24 days), tan to red egg clusters. Females sometimes deposited
eggs adjacent to those of another female. The newer egg clusters were easily
distinguished from the older clusters by coloration. The egg clusters were
rephotographed 15 to 20 days later. At that time the black, parasitized eggs
were readily distinguishable from the red, unparasitized eggs (Fig. 1C).
From the photographs I counted the eggs to determine the original numbers
deposited. Due to layering of eggs in the clusters, the margin of error for
egg counts was 5.5% of the mean eggs per cluster. This was determined by
photographing 32 clusters, estimating the number of eggs per cluster from
the photograph, and then counting the number of eggs per cluster using a
dissecting microscope.

The number of missing eggs was calculated by counting eggs in the re-
photographed clusters and comparing that to the original numbers deposit-
ed. Empty egg shells and partial egg shells were classified as missing eggs.
Although some of the missing eggs may have been lost due to abiotic factors
(e.g. Dempster 1971, Root and Chaplin 1976), most were probably lost to
predators (e.g. indirect evidence from Ives 1967; Owen 1971; Ehrlich and
Gilbert 1973; Gilbert 1975; Rausher 1979a, 1979b; Stamp, pers. observ.).
Thus, I considered missing eggs as representative of those eaten by pred-
ators.

To determine egg parasitism, the black eggs in the rephotographed clus-
ters were counted. These numbers were not affected by layering of eggs
because the parasitized eggs were only found in the exposed layers of the

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NEW YORK ENTOMOLOGICAL SOCIETY

Fig. 2. Total egg loss between years. Numbers above bars are total egg clusters, and ±
one standard error is indicated for mean egg loss. The proportion of clusters with egg losses
(missing and parasitized) differed significantly (x2-test, P < .05). Mean egg loss for all clusters
and mean egg loss for only those clusters with losses were both significantly different between
years (all clusters, two-sample t test, P < .001; clusters with loss, two-sample t test, P < .05).

clusters. In addition, egg clusters rephotographed in late June 1979 (n =
102) and clusters collected two weeks later in July (n = 120) were examined
for parasitism. For 29 leaves the sequence of clusters deposited was deter-
mined based on egg coloration. Fifteen of these leaves had one or more
parasitized cluster, for a total of 19 parasitized clusters. Parasitism rates for
the first cluster and clusters deposited later were compared.

VOLUME LXXXIX, NUMBER 2

93

1978

CLUSTERS

1979

Fig. 3. Percentages of egg clusters with missing eggs and parasitized eggs. Percentages are
given for total clusters, single clusters, and multiple clusters. Numbers of clusters are indicated
above bars. Of the 150 clusters in 1978, 73 were classified as single or multiple clusters. A.
Missing eggs: for single clusters between years, x2-test, P < .025; for single and multiple
clusters in 1979, x2-test, P < .001. B. Parasitized eggs: for single clusters between years and
for multiple clusters between years, x2-tests, P < .001 and P < .005, respectively. No signif-
icant differences occurred between single and multiple clusters within years (x2-tests, P > .25).

Results

Total clusters. — Some 65 to 80% of the clusters lost eggs (parasitism and
predation), and from 5 to 9% of the eggs per cluster were lost (n = 73 and
102 clusters for 1978 and 1979, respectively; Fig. 2). Most clusters with egg

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NEW YORK ENTOMOLOGICAL SOCIETY

A. MISSING EGGS

Fig. 4. Mean numbers of missing and parasitized eggs per cluster with egg loss. Means are
given for total clusters, single clusters, and multiple clusters, with ± one standard error.
Numbers of clusters with egg loss are indicated above bars. Of 67 parasitized clusters in 1978,
54 were classified as single or multiple clusters. High and low refer to a relatively high or low
overall rate of parasitism between years as indicated in Fig. 2B. A. Missing eggs: for single
and multiple clusters in 1979, and for multiple clusters between years, Mann- Whitney U test,
P < .05. B. Parasitized eggs: for single and multiple clusters in 1978, Mann-Whitney U test,
P < .001 : but for single and multiple clusters in 1979, Mann-Whitney U test was not significant
( P > .05).

loss lost a few eggs (less than 20), but a few lost as many as 40 or more.
The proportion of total clusters with missing eggs were similar between
years, as were the mean losses per cluster (Figs. 3 A and 4A). Some 4% of
the eggs per cluster were missing. Chrysopid larvae, coccinellid larvae and
pentatomid nymphs were observed at egg clusters. In contrast to missing
eggs, the numbers of total clusters with parasitized eggs and the mean losses
per cluster were significantly different between years (Figs. 3B and 4B).
Generally, less than 5% of the total eggs were parasitized. During the egg

VOLUME LXXXIX, NUMBER 2

95

deposition period (June), clusters laid later in the month had a level of
parasitism similar to those laid earlier in June (x2-test, P > .10), but mean
loss per cluster was higher in July than in June (normal approximation to
Mann- Whitney U test, nx = 28 and n2 = 102, P < .001). Thus, egg clusters
attacked by predators and parasitoids and which were composed of several
hundred eggs lost only about a tenth of the eggs (e.g. 27 eggs lost ±26 SD
in 1978).

Single and multiple clusters. — The proportion of egg clusters attacked by
predators varied considerably between years for single clusters and between
single and multiple clusters in 1979 (Fig. 3A). Egg losses due to predation
also varied from year to year, with less on multiple clusters than on single
clusters in 1979 (Fig. 4A).

The numbers of parasitized clusters were significantly different between
years for single clusters and for multiple clusters (Fig. 3B). However, the
number of egg clusters per leaf (that is, single compared to multiple clusters)
had no disproportionate effect on the number of parasitized eggs per cluster
(Mann- Whitney U test for 1978 and 1979 combined; nj = 20 and n2 = 22 for
single and multiple clusters, respectively; P > .05). The sequence of egg
clusters deposited did not affect the level of parasitism and number of eggs
parasitized (for clusters attacked, x2_test, P > .10; for parasitized eggs,
Mann- Whitney U test, P > .10).

Discussion

The number of parasitized eggs among E. phaeton clusters was probably
a function mainly of three factors. 1) The distribution of E. phaeton clusters
in these wet meadows was clumped (Stamp 1980b) and most likely was
clumped relative to the parasitoids’ habitat (e.g. Flanders 1937). Conse-
quently, some egg clusters escaped parasitism and predation. More para-
sitized eggs in multiple clusters than in single clusters when overall egg
parasitism was high suggests that these egg parasitoids may cue on the large,
clumped resources of clusters. Cheke (1974) found that the area of discovery
was higher for randomly dispersed egg clusters when parasitoid ( Alaptus
fusculus Haliday; Mymaridae) densities were low whereas when parasitoid
densities were high, multiple clusters were attacked more efficiently. 2) The
variation in numbers of eggs attacked (range 1 to 93 parasitized eggs per
cluster) indicates that some clusters were used by two or more parasitoid
females. 3) The age of the eggs when located by parasitoids may determine
host acceptance (Lewis and Redlinger 1969, Vinson 1976). Furthermore, the
change in egg coloration (yellow to red) followed by the pale green color-
ation of the newly-hatched larvae suggests that these eggs may contain
toxins (e.g. Stamp 1980a) and, thus, such toxins may restrict the period
when eggs are acceptable to parasitoids and predators.

The variation in clusters attacked by predators and loss of eggs was prob-

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NEW YORK ENTOMOLOGICAL SOCIETY

ably a result of the clumped distribution of egg clusters in the habitat, mo-
bility of the egg predators (mainly nymphs and larvae rather than winged
insects), and wide variety of generalist predators using these eggs. For ex-
ample, the greater loss of eggs from single clusters compared to multiple
clusters in 1979 but similar losses between single and multiple clusters in
1978 appeared to be a consequence of the random distribution of the set of
egg predators in this habitat. Ives (1967) suggested that overall egg mortality
of the larch sawfly Pristiphora erichsonii (Hartig) was unrelated to predator
density due to the random distribution of the predators in contrast to the
clumped distribution of eggs, but that once egg predators located clusters
they remained near the eggs and fed periodically. The clumped distribution
of missing eggs within clusters of E. phaeton supports this.

Finally, for E. phaeton females, depositing eggs with those of other fe-
males appeared to have no effect on levels of egg parasitism and predation.
Perhaps later-deposited clusters benefited from nearby predators satiating
themselves on the first cluster, but in one case the second cluster deposited
of three on a leaf was almost completely destroyed by a predator whereas
the other two clusters lost only a few eggs each. Similarly, later-deposited
clusters had the same level of parasitism as the first clusters deposited per
leaf. Consequently, the advantage of depositing clusters with those of other
females was unrelated to the sequence of cluster deposition on a leaf. Thus,
in this host-parasitoid system, egg clusters composed of several hundred
eggs lost a small fraction of eggs to parasitoids and did not benefit from
aggregation of egg clusters.

Acknowledgments

I thank an anonymous reviewer and Bob Fritz (University of Maryland)
for criticizing the manuscript and Doug Morse (Brown University) for com-
menting on an early draft. I am grateful to John Kemper (University of
Maryland) for advice and comments on the research and help with the field
work. The computer time was supported by the Computer Science Center
of the University of Maryland. The research was funded by National
Science Foundation Doctoral Dissertation Improvement Grant No.
DEB-7907618, Xerces Society and Sigma Xi. I am grateful to the Conser-
vation and Research Center of the National Zoological Park at Front Royal,
Virginia for use of the study area and living accommodations. I thank Dave
Vincent and Carll Goodpasture of the Systematic Entomology Laboratory,
U.S.D.A. for identification of the parasitoid.

Literature Cited

Cheke, R. A. 1974. Experiments on the effect of host spatial distribution on the numerical
response of parasitoids. J. Anim. Ecol. 43:107-1 14.

VOLUME LXXXIX, NUMBER 2

97

Dempster, J. P. 197 L The population ecology of the cinnabar moth, Tyria jacobaeae L.
(Lepidoptera, Arctiidae). Oecologia 7:26-67.

Ehrlich, P. R. and L. E. Gilbert. 1973. Population structure and dynamics of the tropical
butterfly Heliconius ethilla. Biotropica 5:69-82.

Flanders, S. E. 1937. Habitat selection by Trichogramma. Ann. Entomol. Soc. Amer. 30:208-
210.

Gilbert, L. E. 1975. Ecological consequences of a coevolved mutualism between butterflies
and plants, p. 210-240. In L. E. Gilbert and P. H. Raven, eds. Coevolution of animals
and plants. Univ. of Texas, Austin.

Hirose, Y., H. Kimoto and K. Hiehata. 1976. The effect of host aggregation on parasitism by
Trichogramma papilionis Nagarkatti (Hymenoptera: Trichogrammatidae), an egg para-
sitoid of Papilio xuthus Linne (Lepidoptera: Papilionidae). Appl. Entomol. Zool.
11:116-125.

Hokyo, N. and K. Kiritani. 1966. Oviposition behavior of two egg parasites, Asolcus mit-
sukurii Ashmead and Telenomus nakagaxvai Watanabe (Hym., Proctotrupoidea, Sce-
lionidae). Entomophaga 11:191-201.

Ives, W. G. H. 1967. Relations between invertebrate predators and prey associated with larch
sawfly eggs and larvae on tamarack. Can. Entomol. 99:607-622.

Janzen, D. H. 1975. Interactions of seeds and their insect predators/parasitoids in a tropical
deciduous forest, p. 154-186. In P. W. Price, ed. Evolutionary strategies of parasitic
insects and mites. Plenum, New York.

Laing, J. 1937. Host finding by insect parasites. I. Observations on the finding of hosts by
Alysia manducator, Mormoniella vitripennis and Trichogramma evanescens. J. Anim.
Ecol. 6:298-317.

Lewis, W. J. and L. M. Redlinger. 1969. Stability of eggs of the almond moth, Cadra cautella,
of various ages for parasitism by Trichogramma evanescens. Ann. Entomol. Soc. Am.
62:1482-1485.

Lyons, L. A. 1962. The effect of aggregation on egg and larval survival in Neodiprion swainei
Midd. (Hymenoptera: Diprionidae). Can. Entomol. 94:49-58.

Owen, D. F. 1971. Tropical butterflies. Clarendon, Oxford. 214 pp.

Rausher, M. D. 1979a. Egg recognition: its advantage to a butterfly. Anim. Behav. 27:1034-
1040.

. 1979b. Larval habitat suitability and oviposition preference in three related butterflies.

Ecology 60:503-511.

Root, R. B. and S. J. Chaplin. 1976. The life styles of tropical milkweed bugs, Oncopeltus
(Hemiptera: Lygaeidae) utilizing the same hosts. Ecology 57:132-140.

Royama, T. 1970. Factors governing the hunting behaviour and food selection of the great tit
( Parus major L.). J. Anim. Ecol. 39:619-688.

Stamp, N. E. 1980a. Egg deposition patterns in butterflies: why do some species cluster their
eggs rather than deposit them singly? Amer. Natur. 115:367-380.

. 1980b. Effect of group size on an egg-clustering butterfly. Ph.D. Dissertation. Univ.

of Maryland. 94 pp.

Tinbergen, N., M. Impekoven and D. Franck. 1967. An experiment on spacing-out as a
defence against predation. Behaviour 28:307-321.

Vinson, S. B. 1976. Host selection by insect parasitoids. Ann. Rev. Entomol. 21:109-133.

Department of Zoology, University of Maryland, College Park, Maryland
20742.

Received for publication August 12, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(2), 1981, pp. 98-101

SOME OBSERVATIONS ON SOARING FLIGHT IN THE
MOURNING CLOAK BUTTERFLY ( NYMPHALIS
ANTIOPA L.) IN SOUTHERN ONTARIO

David L. Gibo

Abstract. — The Mourning Cloak butterfly, Nymphalis antiopa L., is ca-
pable of soaring flight. The adults appear to recognise rising air currents and
exhibit complex behaviour patterns to stay inside these systems. The soar-
ing behaviour of N. antiopa is similar to that of monarch butterflies, Danaus
plexippus L.

Soaring flight consists of maintaining or gaining altitude by gliding in rising
air currents (Conway 1969; Gibo and Pallett 1979). Consequently, a soaring
animal is actually remaining aloft by extracting energy from the atmosphere.
Although soaring flight is the most efficient form of flight possible in terms
of energy expenditure per unit of distance travelled (Pennycuick 1969), de-
liberate use of this flight technique has only been reported for two insects,
the monarch butterfly, Danaus plexippus L. (Gibo 1981; Gibo and Pallett
1979; Schmidt-Koenig 1979), and the dragonfly Pantala flavescens Fabricius
(Hankin 1921). The apparent rarity of this flight technique among insects
may be due to the need for complex behaviour patterns (Gibo and Pallett
1979; Hankin 1921; Pennycuick 1969, 1972, 1975), special sensory systems
(Johnson 1969), and specific meteorological conditions (OSTIV 1978). Dur-
ing a 9 day period in the late summer of 1979, while observing the flight of
migrating monarch butterflies, Danaus plexippus L., in southern Ontario,
I was also able to observe the flight techniques of mourning cloak butteries,
Nymphalis antiopa L. Three of the 11 N. antiopa observed were soaring.
This note is a preliminary report on the soaring behaviour of N. antiopa.

One observation was made in an open area on the Erindale College Cam-
pus of the University of Toronto and two were made in an open area 3 km
to the west of the campus. The times ranged from noon to midafternoon.
Wind velocities at 1 m above the ground were measured with a wind meter.
The courses of the butterflies were determined with a compass. The air
temperature at 1 m was also measured. The estimated altitude of the but-
terfly above the ground was recorded and any significant manoeuvers per-
formed by the soaring butterflies, such as circling, were noted.

Two of the soaring N. antiopa were observed to enter thermals (rising
bubbles or columns of relatively warm air produced by convection). The
first individual, observed on 4 September at 12:03 P.M. (E.S.T.), flew over-
head at an altitude of 20 m. The wind direction was 165° and the velocity

VOLUME LXXXIX, NUMBER 2

99

was 5 km/hr. Initially the butterfly employed powered flight and maintained
a 225° (SW) course. Suddenly, presumably upon encountering a thermal,
the butterfly stopped beating its wings and began soaring in circles 1 m in
diameter. After gaining approximately 10 m in altitude and simultaneously
drifting approximately 50 m downwind on a 325° (NW by W) track, the
butterfly abruptly resumed powered flight and reestablished its former SW
course.

The second individual, observed on 5 September, gave an impressive
demonstration of its ability to locate and exploit a weak thermal. When the
butterfly was first observed (11:45 P.M., E.S.T.), it was flying under power
at a height of 1 m and was maintaining a SW course. There was no apparent
wind within 2 m of the ground. As the N. antiopa passed over a paved
parking area 5 m from the observation point it suddenly began to climb, still
beating its wings, until it reached a height of 3 m. It then stopped beating
its wings and began soaring in circles. The butterfly did not beat its wings
again for the rest of the observation: a period of 2 minutes. Initially the
butterfly flew in circles approximately 3 m in diameter remaining directly
over the parking area. It gained altitude gradually and required about 1.5 m
to reach a height of 15 m. Apparently there was a light northeast breeze at
this height because the butterfly then began to drift on a 215° (SW by S)
course, reducing the size of its circles to approximately 1 m in diameter.
For the remainder of the observation the butterfly continued to soar in 1 m
circles and drift on a 215° (SW by S) track. When the N. antiopa reached
an approximate altitude of 25 m and a distance of 80 m, I could no longer
determine whether it was still gliding (i.e. not beating its wings) and ter-
minated the observation.

The third individual was observed on 8 September. The N. antiopa was
already soaring overhead in a thermal when first noticed. The time was 2:38
P.M. (E.S.T.). The wind direction was 105°, and the velocity was 5 km/hr.
This butterfly was at an altitude of 20 m and soared in circles of approxi-
mately 1 m in diameter while drifting overhead on a 285° (W by N) course.
This last specimen neither gained nor lost altitude, but simply drifted down-
wind until it was lost from sight.

The mourning cloak has long been known to migrate in Europe (Williams
1942, 1958) and there is evidence that at least part of the population in
eastern North America migrates (Shannon 1917). Frequent episodes of soar-
ing would significantly reduce the energy expenditure of this activity. It is
interesting to note that the mean course and angular deviation (see Bat-
schelet 1965) of the 11 N. antiopa in this study was 246° ± 30.3 (WSW),
which is similar to the SW to SSW courses frequently reported for migrating
D. plexippus (Beall 1941; Schmidt-Koenig 1979; Urquhart 1960; Urquhart
and Urquhart 1978).

The mourning cloak butterfly is the second butterfly found to exhibit

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NEW YORK ENTOMOLOGICAL SOCIETY

specialized soaring behaviour. The soaring behaviour observed for N. an-
tiopa in thermals was similar to the behaviour reported by Gibo and Pallet
(1979) for D. plexippus. Individuals of both species are apparently able to
detect and enter thermals both near the ground and well above the ground.
In addition, adults of both species are able to remain within thermals by
soaring in circles. The presence of this highly specialized behaviour in mem-
bers of two distinct families of butterflies (Danaidae and Nymphalidae) sug-
gests that the ability to soar may be widespread among butterflies.

Acknowledgments

I would like to thank Thomas Alio way for helpful assistance, and Cath-
erine A. Neal and Bonnie M. Soutar for critically reading the manuscript.

Literature Cited

Batschelet, E. 1965. Statistical methods for the analysis of problems in animal orientation and
certain biological rhythms. America Institute of Biological Sciences, Washington D.C.
57 p.

Conway, C. 1969. The Joy of Soaring. The Soaring Society of America, Inc., Los Angeles,
Calif. 134 p.

Gibo, D. L. 1981. Altitudes attained by migrating monarch butterflies, Danaus p. plexippus
(Lepidotera: Danaidae), as reported by glider pilots. Can. J. Zool. 80:In Press.

Gibo, D. L. and M. J. Pallett. 1979. Soaring flight of monarch butterflies, Danaus plexippus
(Lepidoptera: Danaidae), during late summer migration in southern Ontario. Can. J.
Zool. 57(7): 1393-1401.

Hankin, E. H. 1921. The soaring flight of dragonflies. Cambrid. Phil. Soc. Proc. 20(4):460-
465.

Johnson, C. G. 1969. The Migration and Dispersal of Insects by Flight. Methuen and Co.
Ltd., London. 763 p.

The Organisation Scientifique et Technique Internationale du Vol a Voile — OSTIV — in collab-
oration with WMO (World Meteorological Organization). 1978. Handbook of Meteo-
rological Forecasting for Soaring Flight. Secretariat of the World Meteorological Or-
ganization, Geneva, Switzerland. 101 p.

Pennycuick, D. J. 1969. The mechanics of bird migration. Ibis 11:525-526.

. 1972. Soaring behaviour and performance of some east African birds observed from

a motorglider. Ibis 114:178-218.

. 1975. The mechanics of flight. In Avian biology, D. S. Farmer, J. R. King, and K.

C. Parkes, eds.. Academic Press, New York 5:1-75.

Shannon, H. J. 1917. Autumn migration of butterflies. Amer. Mus. J. 17( 1):32— 40.
Schmidt-Koenig, K. 1979. Directions of migrating monarch butterflies ( Danaus plexippus ;
Danaidae; Lepidoptera) in some parts of the Eastern United States. Behav. Proc.
4:73-78.

Urquhart, F. A. 1960. The Monarch Butterfly. University of Toronto Press. Toronto, Ont.
361 p.

and N. R. Urquhart. 1979. Breeding Areas and Overnight Roosting Locations in the

Northern Range of the Monarch Butterfly ( Danaus plexippus plexippus) with a Summary
of Associated Migratory Routes. Can. Field Nat. 93:41-47.

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101

Williams, C. B. 1958. Insect Migration. Wilmer Brothers and Haram Ltd., London and Glas-
gow. 235 p.

, G. F. Cockbill, M. E. Gibbs and J. A. Downes. 1942. Studies in the migration of

Lepidoptera. Tran. R. Ent. Soc. Lond. 92:101-283.

Wolters, R. A. 1971. The Art and Technique of Soaring. McGraw-Hill, New York. 197 p.

Department of Zoology, Erindale College, University of Toronto, Missis-
sauga, Ontario L5L 1C6, Canada.

Received for publication November 10, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(2), 1981, pp. 102-108

OBSERVATIONS ON THE SPRING AND SUMMER BEHAVIOR OF
THE 12-SPOTTED LADYBIRD BEETLE, COLEOMEGILLA
MACULATA (DEGEER) (COLEOPTERA: COCCINELLIDAE)

Allen H. Benton and Andrew J. Crump

Abstract. — The twelve-spotted ladybird beetle, Coleomegilla maculata
(DeGeer) was studied at sites in western and central New York from 1973
to 1980. Aggregations of overwintering beetles begin to be active in April,
and dispersal occurs gradually from then until late May. During this time,
beetles feed on pollen of a variety of flowers in and near the aggregation
site. After dispersal and a period of feeding, mating occurred near the end
of May. Summer generations moved into cornfields when the corn was large
enough to provide pollen and/or aphids. Sweet corn in this area may reach
this size by early July, field corn by early August. The daily activity cycle
of the beetles appeared to be based on geotaxis, with upward movement
occurring in the early daylight hours and downward movement occurring in
late morning. A positive hygrotaxis may also be involved in the downward
movement, as the higher locations begin to become drier as the day ad-
vances.

During dispersal, beetles showed a strong preference for yellow in color
preference tests in the laboratory, and were commonly found on yellow
flowers in the field. Once dispersed, this preference disappears and no clear
pattern of color preference is shown. This phenomenon may have some
bearing on the fact that dispersing beetles tend to fly in the direction of the
sun.

Introduction

The twelve-spotted ladybird beetle, Coleomegilla maculata (DeGeer), is
a common Coccinellid throughout temperate North America and northern
Europe. The species has been well studied, beginning with the pioneering
study by Riley (1893). More recent studies include those of Hodek (1967)
in Europe, Balduf (1935), Hagen (1962), Ewert and Chiang (1966) and Smith
(1971). In a previous paper from this laboratory (Benton and Crump, 1979)
we have discussed overwintering aggregations in this species. The present
paper discusses the behavior of this beetle from the time of spring dispersal
until the formation of the overwintering aggregation.

Methods and Materials

Winter aggregations of C. maculata were observed in the town of Pom-
fret, Chautauqua County, New York, and in the town of Victory, Cayuga

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103

County, New York. Most of the observations on dispersal were made at the
Cayuga County site. Post-dispersal observations were made at both loca-
tions, while summer generations were studied near the Chautauqua county
site. Several hundred beetles were used in a series of laboratory experiments
related to sensory perception, including reaction to light, temperature, rel-
ative humidity, color and odors. Dispersal patterns were observed in the
laboratory by releasing beetles in a greenhouse at the appropriate time.

Spring Dispersal

While the process of aggregation in the autumn occurs within a week or
two (Benton and Crump, 1979), dispersal is a much more prolonged process.
The beetles become active after a few days of warm weather, even if they
occur during the winter months. Usually the first few warm days in early -
to mid-April cause the beetles to move up grass stems or other plants in
preparation for dispersal from the overwintering site. If pollen is available
at or near the aggregation area, they will begin to feed. In early May, many
beetles were found on male catkins which had fallen from the cottonwood
(Populus deltoides) at the base of which an aggregation had formed and
shortly thereafter on dandelions up to ten meters from the aggregation site.
At other locations, we have found large numbers (up to 10 per flower) on
the cowslip (C alt ha palustris ), and smaller numbers on a variety of other
spring flowers. No preference for specific flowers was noted, although the
greatest numbers have been found on bright yellow flowers such as cowslips
and dandelions. However, these are often the most abundant large flowers
in the vicinity of aggregations, so they are obvious choices for efficient
pollen feeding. The preference of dispersing beetles for yellow, as noted
below, may or may not be related to the presence of yellow flowers during
the gradual dissolution of the aggregation.

On May 4, 1980, beetles were counted on dandelion flowers within 10
meters of the Cayuga County aggregation. Feeding began shortly after sun-
rise, with the first beetle observed on a flower at 5:30 a.m./EST. From then
until 9:30, the numbers increased steadily, reaching a peak of 32 individuals
at 9:30/EST. By 10:00 a.m., the number had dropped to 19 and counting
was discontinued. This agrees well with summer observations noted below
(see Fig. 1).

Fanwise dispersal from the aggregation has been reported for this species
(Allee, 1949). A possible mechanism for attainment of this sort of dispersal
might be flight in the direction of the sun. As the day advanced, beetles
would fly successively in a more westerly direction, and would thus be dis-
persed over a wide area. To investigate this, we placed a group of 50 beetles
in a darkened box at the time when the aggregation was dispersing. The box
was opened in a greenhouse, and the beetles crawled to the top of the box
and took off in the direction of the sun. For several hours, the beetles re-

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EASTERN STANDARD TIME

Fig. 1. Vertical distribution of adult Coleomegilla maculata during daylight hours under
laboratory conditions. See text for details.

mained clustered at the top of the greenhouse in a direct line from the box
toward the sun.

As noted above, beetles emerging from the aggregation occurred most
commonly on yellow flowers. Coupled with the direct sunward flight on
dispersal, this suggested an attraction for bright yellow at this time. We
tested this in the laboratory by placing colored cards of equal size in an area
30 cm square and 10 cm high. The reflectivity of the cards was not mea-
sured. Groups of beetles were placed in the arena, which was lighted from
above by fluorescent lights, and their locations checked at intervals. At the
time of dispersal, 90% of the beetles remained on yellow and white cards
after one hour, indicating initial preference for light colors. When similar
tests were conducted with free-ranging beetles collected in June and August,
no clear color preference was evident.

Mating may occur before the beetles leave the aggregation, though long
term observations have not revealed it. It is more probable that a period of
breeding is necessary before mating. Adult beetles collected from flowers in
Chautauqua County on May 15, 1979, were placed in a large jar as they were
collected. By the time they were transferred to the laboratory, most of them
were mating. We maintained these beetles in the laboratory under natural

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105

1H Before addition of water
□ After addition of water

LIGHT

DARK

j

LIGHT DARK

12 HOURS 36 HOURS

Length of water deprivation

Fig. 2. Vertical distribution of adult Coleomegilla maculata after water deprivation, before
and after addition of water to the substrate. See text for details.

conditions of light and temperature, but no oviposition was observed and
all beetles died within 30 days.

Although C. maculata may be found in small numbers in a variety of
habitats in summer, most individuals tend to congregate in corn fields, mak-
ing them readily available for study. By early July, local fields of corn ( Zea
mays)' have reached a size adequate to supply aphids, while sweet corn
fields may supply abundant pollen by mid-July. Both adult and larval beetles
may be found on the corn in great numbers, feeding on both aphids and
pollen of the corn, for the rest of the summer, leaving the corn only to move
to the aggregation site.

Adult beetles were found on exposed leaves and stems only during the
morning hours (7-11 EST). At other times, they could be found in rolled-up

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NEW YORK ENTOMOLOGICAL SOCIETY

leaves, leaf axils or on the ground around the bases of the corn plants. Field
observations showed that beetles climbed or flew to the upper levels of the
corn in the early morning hours, and descended to lower levels or to leaf
axils before noon. Several possible reasons could explain this vertical move-
ment: 1) Positive phototaxis might cause upward movement as the sun rose.
This appeared unlikely, since downward movement was completed while
the sun was still moving toward its zenith. 2) Rising temperature might
stimulate feeding activity. This, too, seemed unlikely, since the downward
movement occurred while the temperature was still rising. 3) Negative geo-
taxis might initiate upward movement at sunrise. Observations in both field
and laboratory showed that beetles tended to move upward on any vertical
object available to them, suggesting a geotactic response. The most obvious
explanation for the downward movement was a positive hygrotaxis of suf-
ficient strength to overcome the climbing tendency. Obviously, a reversal
of the geotaxis or a negative phototaxis would have the same result.

This phenomenon was investigated in the laboratory. Ten groups of ten
beetles each were placed in 10 cm x 24 cm glass bell jars with netting over
them. The jars were about lA filled with moist soil and leaf litter, with small
sticks extending to the top of the jar. The jars were placed in natural light
and temperature conditions. At intervals throughout the daylight hours, the
numbers of beetles in the top five cm of the jar (i.e., near the top of the
vertical stick) was counted. This experiment was repeated monthly for three
months, in July, August and September. The mean results (Fig. 1) show
that under conditions of natural light and temperature, and constant high
humidity, there was a general early morning increase in locomotor activity
or a tendency to climb, leading to a peak shortly before 10 a.m. EST. This
agreed precisely with the field observations in corn fields. Furthermore, the
time pattern agrees with that noted above when beetles move out from the
aggregation to feed on nearby flower pollen.

The relationship between the effects of light, gravity and humidity was
further investigated by measuring the vertical distribution of beetles in bell
jars in constant light (3 K lux) and constant darkness after 12 or 36 hours
of water deprivation, and again after the addition of water to the soil at the
bottom of the containers. The results (Fig. 2) show that a) the beetles were
distributed at higher levels in darkness than in light irrespective of their
state of desiccation; and b) that, when desiccated, beetles kept in darkness
were distributed at markedly lower levels than when they were supplied
with water. However, this difference did not appear when water was added
to desiccated beetles kept in constant light.

Discussion

It is known that C. maculata has a low tolerance for desiccation, and
Ewert and Chiang (1965) have shown that the beetles move up a moisture

VOLUME LXXXIX, NUMBER 2

107

gradient and exhibit negative phototaxis. During extensive testing in an
insect olfactometer, with many stimuli, we could find no evidence of chem-
otaxis, but confirmed that C. maculata could detect and move toward areas
of high moisture level. This positive hygrotaxis appears to be the dominant
factor in the behavior of the beetle. It plays an important role in the for-
mation of aggregations (Benton and Crump 1979) and is a major influence
in the distribution of beetles in their summer habitats. The importance of
phototactic and geotactic elements in the distribution of the beetles are
difficult to elucidate due to their changing nature and interaction. We have
observed that C. maculata shows apparent positive phototaxis, probably
complemented by a negative geotaxis, in order to effect dispersal from the
overwintering site, the same individuals having shown the opposite response
during the formation of the aggregation some months earlier. The same sort
of shift occurs in summer between early morning and late morning. Avoid-
ance of desiccation is of primary importance to this species, however, and
it may well be that the aforementioned downward movement of late morning
is the result of a positive hygrotaxis as the air around the upper strata of
corn becomes drier. This would act to guide the beetles down toward the
basal regions of the corn. Since a good deal of pollen accumulates in leaf
axils and on the ground, this would permit pollen feeding to continue even
though the beetles were relatively inactive. Considerable refinements of
experiments are essential to eliminate all responses except the one being
studied before we can understand the movements of this species, since
responses appear to alter substantially with subtle changes in environmental
conditions.

Acknowledgments

For assistance in the field work connected with this study, we are indebted
to James Hays, Joseph Moore, Richard Northrup and Marlon Zielinski. This
research was sponsored, in part, by National Science Foundation Grant
B036435.

Literature Cited

Allee, W. C., A. E. Emerson, O. Park, T. Park and K. P. Schmidt. 1949. Principles of Animal
Ecology. W. B. Saunders & Co., Philadelphia.

Balduf, W. V. 1926. The bionomics of Entomophagous Coleoptera. J. S. Swift Co., Inc., St.
Louis.

Benton, Allen H. and Andrew J. Crump. 1979. Observations on aggregations and overwin-
tering in the Coccinellid beetle Coleomegilla maculata (DeGeer). J. N.Y. Entomol.
Soc., 87(2): 154-159.

Ewert, M. A. and H. C. Chiang. 1966. Dispersal of three Coccinellids in a corn field. Canad.
Entomol. 98:999-1003.

Hagen, K. S. 1962. Biology and ecology of predaceous Coccinellids. Ann. Rev. Entomol.
7:289-326.

108

NEW YORK ENTOMOLOGICAL SOCIETY

Hodek, I. 1967. Bionomics and ecology of predaceous Coccinellids. Ann. Rev. Entomol.
12:79-104.

Riley, C. V. 1893. Note on the life habits of Megilla maculata. Proc. Entomol. Soc. Wash-
ington 2: 168-170.

Smith, R. C. 1971. Effects of various factors on the local distribution and density of Coccinellid
adults. Canad. Entomol. 103:1115-1124.

State University College, Fredonia, New York 14063.

Received for publication November 18, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(2), 1981, pp. 109-122

PRINCIPAL COMPONENTS ANALYSIS OF BIOGEOGRAPHIC
PATTERNS AMONG HELICONIA INSECT COMMUNITIES

Richard P. Seifert

Abstract. — This study explores biogeographic patterns among insect com-
munities which live in the water-filled floral bracts of Heliconia plants.
Twenty-five collections from 13 Neotropical locations are studied using a
principal components analysis where the collections are ordinated on the
basis of the frequency of occurrence of the 23 most common insect morpho
species. Heliconia species from the French Antilles show a depauperate
insect community, apparently due to island isolation. Floral structure is
important in determining the insect community: Heliconia species with
small floral bracts have low species richness and only one Heliconia species
with a pendant inflorescence has an aquatic or semi-aquatic insect com-
munity. Different Heliconia species collected from the same location do not
necessarily have comparable insect species communities.

Introduction

Biogeographic studies have held an important position in the growth and
understanding of evolutionary ecology since the work of Darwin (1859) and
Wallace (1876). These studies have attempted to discern patterns of distri-
butions and to identify the factors which determine the distributions of either
taxonomically or ecologically related organisms (Price 1975; Pianka 1978).
This paper is a biogeographic study based on a principal components anal-
ysis (Pielou 1977) of collections of 25 different insect communities living in
the water-filled floral bracts of Heliconia (Zingiberales: Musaceae) plants
collected from 13 different locations in Costa Rica, northern South America
and the Antilles made over a period of 8 years. Through this analysis I tried
to discern basic biogeographic patterns of Heliconia insect communities, to
determine if Heliconia species in the same location had similar insect com-
munities even if they bloomed during different seasons and to determine if
similar insect communities lived in the water-filled bracts of Heliconia
species with similar bract morphologies.

Plants in the genus Heliconia are common members of low- and mid-
elevation Neotropical wet forests. These plants are most frequent in sunlit
areas. Detailed studies on the natural history of these plants have been
published by Stiles (1975, 1979). Many Heliconia species have inflores-
cences consisting of a series of large erect cup like floral bracts, each bract
containing several flowers (Fig. 1). The erect bracts collect and hold both
rainwater and water actively pumped into the bracts by the plant (Seifert

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NEW YORK ENTOMOLOGICAL SOCIETY

Fig. 1 . An inflorescence of Heliconia cf. caribaea sp. 2 from Limoncocha, Ecuador.

and Seifert 1976). A diverse group of insects live all or part of their lives in
the water-filled Heliconia bracts (Seifert 1975; Seifert and Seifert 1976). The
insect species composition varies with Heliconia species and depends in
part on the flowering season of the Heliconia species as well as the pro-
portion of the bract which has standing water (Seifert 1980). Some Heliconia

VOLUME LXXXIX, NUMBER 2

111

species have inflorescences which are pendant rather than erect. In most
cases, the bracts of pendant species do not hold water or harbor an aquatic
insect community. However, one pendant species ( H . ro strata Ruiz and
Pavon) has tightly compressed bracts which retain water and maintain a
community of insects similar to that found in some erect bract Heliconia
species.

Materials and Methods

Twenty-five Heliconia insect communities were collected from different
Heliconia species from Costa Rica (2 locations), Venezuela (5 locations),
Ecuador (3 locations), Trinidad (1 location), Martinique (1 location) and
Guadeloupe (1 location) (Table 1). In 3 cases I collected insect communities
from the same Heliconia species at the same location in both the early and
late portions of the blooming season. At some sites I was able to make
collections of more than 1 Heliconia species. In each collection I sampled
at least 20 inflorescences, carefully dissected each bract and recorded the
number of individuals of each insect species in each inflorescence. After all
collections were made, I identified (to genus or family) the 23 insect ”mor-
pho species” found most commonly in the collections. For this study, insect
morpho species were recorded as ecological equivalents. Thus, for example,
there was a category for small hispine beetle (genus Cephaloleia) and I
recorded this as present from several collections even though different his-
pine species existed at different locations. I assumed that the small hispines
represented organisms foraging in the same manner even though their Latin
binomens differed. Further, in the case of hispine beetles, I recorded adults
and larvae separately since in some hispine species only the larvae feed on
the inflorescences (Strong 1977a, b; Seifert and Seifert 1979). Next I com-
puted the frequency of occurrence of each insect morpho species in each
collection. This figure was simply the number of inflorescences which had
at least one specimen of a given insect morpho species divided by the total
number of inflorescences in the collection. Following this, a matrix of insect
morpho species frequencies (columns) for each Heliconia collection (rows)
was constructed (Table 2). These data were analyzed using a principal com-
ponents analysis which examined the similarity of collections based on their
insect morpho species frequencies (R technique of Pielou 1977).

Principal components analysis is an ordination method which is used to
examine the pattern of variation among variates where no a priori patterns
of causality are suggested. This technique shows relationships among sam-
ples which are not apparent from a simple inspection of the data. The orig-
inal data are projected onto a plot (or series of plots) of few dimensions in
such a way that the arrangement of points suffers the least possible distor-
tion. Each principal component is a linear combination of the original vari-

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NEW YORK ENTOMOLOGICAL SOCIETY

Table 1. Collection label, Heliconia species resemblance, collection location and season
of collection for 25 Heliconia collections used in the principle components analysis.

Collection

label

Heliconia
species
resemblance
and describer

Collection

location

Approximate

co-ordinants

Season

collected

1

aurea

Rodriguez

Rancho Grande
Venezuela

10°21'N

65°4LW

wet season

2

cf. humilis
sp. 1
Jacq.

La Escalera
Venezuela

05°45'N

61°25'W

wet season

3

cf. humilis
sp. 2
Jacq.

El Dorado
Venezuela

06°4LN

61°35'W

wet season

4

cf. caribaea
sp. 2
Lamarck

Limoncocha

Ecuador

0°34'S

76°38'W

dry season

5

cf. humilis
sp. 3
Jacq.

Primavera

Ecuador

0°32'S

76°45'W

dry season

6

episcopalis

Velloso

Primavera

Ecuador

0°32'S

76°45'W

dry season

7

bihai

Linn.

Les Nuages
Martinique

14°42'N

61°06'W

wet season

8

cf. caribaea
sp. 1
Lamarck

Panaquire

Venezuela

10°21'N

66°15'W

wet season

9

bihai

Linn.

Bains Juanes
Guadeloupe

16°03'N

61°40'W

wet season

10

caribaea

Lamarck

Les Nuages
Martinique

14°42'N

61°06'W

wet season

11

caribaea

Lamarck

Bains Juanes
Guadeloupe

16°03'N

61°40'W

wet season

12

cf. caribaea
sp. 1
Lamarck

La Trilla
Venezuela

10°22'N

65°42'W

wet season

13

rostrata
Ruiz and
Pavon

Limoncocha

Ecuador

0°34'S

76°38'W

dry season

14

latispatha

Bentham

Tinalandia

Ecuador

0°14'S

79°07'W

dry season

15

cf. caribaea

sp. 1

Lamarck

Guatopo

Venezuela

09°45'N

66°28'W

wet season

16

rodriguensis

Aristeguieta

Guatopo

Venezuela

09°45'N

66°28'W

wet season

VOLUME LXXXIX, NUMBER 2

113

Table 1.

Continued.

Collection label

Heliconia
species
resemblance
and describer

Collection

location

Approximate

co-ordinants

Season

collected

17

bihai

Linn.

Rancho Grande
Venezuela

10°2LN

65°41'W

dry season
(early)

18

bihai

Linn.

Rancho Grande
Venezuela

10°21'N

65°41'W

wet season
(late)

19

imbricata

(Kuntze)

Baker

Rincon de Osa
Costa Rica

08°42'N

83°30'W

wet season
(early)

20

imbricata

(Kuntze)

Baker

Rincon de Osa
Costa Rica

08°42'N

83°30'W

wet season
(late)

21

wagneriana

Petersen

Rincon de Osa
Costa Rica

08°42'N

83°30'W

dry season
(early)

22

wagneriana

Petersen

Rincon de Osa
Costa Rica

08°42'N

83°30'W

dry season
(late)

23

latispatha

Bentham

Sirena
Costa Rica

08°40'N

83°40'W

dry season

24

wagneriana

Petersen

Sirena
Costa Rica

08°40'N

83°40'W

dry season

25

cf. caribaea
sp. 1
Lamarck

Simla

Trinidad

10°45'N

61°22'W

wet season

NOTE: The taxonomic status and nomenclature of many members of the genus Heliconia
are uncertain. Identifications follow, when possible, Aristeguieta (1961) and Daniels and Stiles
(1979). Heliconia species designated as H. cf. caribaea and H. cf. humilis represent morpho-
logical forms, some of which may be undescribed, which appear similar to members of the H.
caribaea and H. humilis species complexes. However, only the forms from Martinique and
Guadeloupe represent true populations of the species originally described as H. caribaea
Lamarck.

ates where the first principal component accounts for the maximum possible
variance and each subsequent component accounts for a decreasing amount
of the residual variance. In this study each of the 25 Heliconia collections
is characterized by the frequency of occurrence of the 23 morpho species.
The Heliconia collections represent the variates to be ordinated; the insect
morpho species frequencies are the attributes on which the ordination is
based. Principal components analysis has been used frequently in taxonomic
studies (Sneath and Sokal 1973) as well as in ecological studies (Pielou 1977).
Often the characters used are standardized when the variates are measured
in different units. However, in ecological work when the variates are mea-
sured in the same units (here, frequencies of insect morpho species) stan-
dardization is not appropriate (Pielou 1977). Further, some workers, partic-

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NEW YORK ENTOMOLOGICAL SOCIETY

Table 2. Frequency of occurrence of the 23 most common insect “morpho species” (listed
by letters) found in 25 Heliconia inflorescences collections.

Heliconia

collection

label

Insect morpho species frequencies

a

b

c

d

e

f

g

h

i

j

k

1

1

0.91

0.14

0.84

0.02

0.28

0.97

0.14

0.00

0.22

0.09

0.03

0.02

2

0.85

0.40

0.35

0.00

0.20

0.00

0.00

0.00

0.00

0.00

0.05

0.00

3

1.00

0.55

0.10

0.65

0.15

0.00

0.00

0.00

0.00

0.05

0.00

0.05

4

0.79

0.88

0.09

0.02

0.21

0.40

0.26

0.54

0.37

0.26

0.09

0.00

5

0.77

0.60

0.00

0.13

0.57

0.10

0.43

0.27

0.00

0.13

0.00

0.00

6

0.00

0.61

0.00

0.00

0.79

0.03

0.00

0.00

0.00

0.00

0.00

0.00

7

0.00

0.00

0.00

0.18

0.00

0.00

0.00

0.00

0.00

0.00

0.02

0.00

8

1.00

0.17

0.27

0.67

0.57

0.73

0.30

0.00

0.20

0.03

0.00

0.00

9

0.00

0.00

0.48

0.31

0.00

0.00

0.00

0.00

0.00

0.10

0.10

0.00

10

0.52

0.00

0.00

0.77

0.00

0.00

0.00

0.00

0.00

0.00

0.00

0.00

11

0.00

0.00

0.58

0.93

0.00

0.00

0.00

0.00

0.00

0.04

0.00

0.00

12

1.00

0.83

0.64

0.93

0.80

0.23

0.00

0.03

0.00

0.17

0.00

0.00

13

0.00

0.79

0.00

0.10

0.69

0.00

0.07

0.21

0.07

0.45

0.00

0.03

14

0.68

0.77

0.10

0.00

0.39

0.00

0.32

0.00

0.32

0.06

0.16

0.00

15

1.00

0.85

0.50

0.80

0.25

0.30

0.05

0.00

0.00

0.10

0.00

0.00

16

0.82

0.71

0.52

0.94

0.09

0.15

0.03

0.12

0.35

0.12

0.03

0.03

17

0.59

0.66

0.94

0.28

0.03

0.88

0.41

0.12

0.19

0.59

0.28

0.00

18

0.12

0.70

0.79

0.18

0.00

1.00

0.24

0.09

0.42

0.39

0.09

0.03

19

0.83

0.80

0.30

0.93

0.47

0.70

0.50

0.03

0.03

0.23

0.20

0.73

20

0.40

0.80

0.37

0.87

0.33

0.83

0.17

0.43

0.00

0.27

0.13

0.53

21

0.10

0.97

0.43

0.03

0.00

0.07

0.37

0.00

0.00

0.13

0.03

0.03

22

0.83

0.67

0.77

0.37

0.97

0.00

0.07

0.67

0.00

0.60

0.07

0.27

23

0.60

0.07

0.13

0.00

0.27

0.00

0.73

0.00

0.00

0.13

0.00

0.00

24

0.95

0.90

0.90

0.05

0.90

0.10

0.16

0.32

0.00

0.26

0.00

0.00

25

0.85

0.20

0.10

0.05

0.20

0.10

0.00

0.00

0.35

0.10

0.05

0.00

NOTE: The Heliconia collection labels are the same as those listed in Table 1. Insects
represented by letters are: a = Quichuana (Syrphidae), b = Gillisius (Hydrophilidae), c =
copestylum (Syrphidae), d = Merosargus (Stratiomyidae), e = Beebeomyia (Richardiidae),
f = Cephaloleia larva (Chrysomelidae), g = Cephaloleia adult (Chrysomelidae), h = small
Gillisius (Hydrophilidae), i = large hispine (Chrysomelidae), j = Odontolinus (Staphylinidae),

ularly in taxonomic studies, remove highly correlated characters. Once
again this procedure is less commonly used in ecological work where high
correlations of insect frequencies may indicate a particular and regular pat-
tern of community structure. I have constructed a correlation matrix based
on insect morpho species frequencies (Table 3). However, insect morpho
species whose frequencies are highly correlated were not removed from the
data set used to produce this principal components analysis. My procedure
(no standardization, no removal of correlated characters) follows Pielou
(1977) who can be referred to for a more detailed treatment of principal
components analysis in ecological studies. Thus, the data set which I use
in this principal components analysis is the Heliconia collection by insect
morpho species frequencies listed in Table 2.

VOLUME LXXXIX, NUMBER 2

115

Table 2. Continued.

Heliconia

collection

label

Insect morpho species frequencies

m

n

0

p

q

r

s

t

u

V

w

1

0.00

0.00

0.02

0.00

0.00

0.00

0.00

1.00

1.00

0.57

0.00

2

0.05

0.00

0.00

0.00

0.15

0.00

0.00

0.55

0.00

0.00

0.00

3

0.05

0.00

0.00

0.00

0.00

0.00

0.00

0.05

0.20

0.00

0.40

4

0.02

0.00

0.16

0.00

0.19

0.02

0.09

0.59

0.00

0.00

0.00

5

0.03

0.00

0.00

0.00

0.07

0.00

0.03

0.90

0.00

0.00

0.00

6

0.03

0.06

0.00

0.00

0.00

0.00

0.15

0.00

0.00

0.00

0.00

7

0.00

0.00

0.00

0.00

0.00

0.00

0.00

0.00

0.68

0.00

0.00

8

0.00

0.00

0.00

0.00

0.00

0.00

0.07

0.97

0.20

0.00

0.17

9

0.00

0.00

0.00

0.00

0.00

0.00

0.00

0.00

0.19

0.00

0.00

10

0.00

0.03

0.03

0.00

0.00

0.03

0.00

0.03

0.87

0.00

0.00

11

0.00

0.00

0.00

0.00

0.00

0.02

0.00

0.00

1.00

0.00

0.00

12

0.00

0.30

0.33

0.03

0.00

0.00

0.00

0.97

0.00

0.00

0.00

13

0.00

0.03

0.21

0.00

0.00

0.07

0.00

0.00

0.00

0.00

0.00

14

0.03

0.00

0.39

0.00

0.00

0.00

0.00

0.00

0.00

0.00

0.00

15

0.00

0.10

0.00

0.00

0.00

0.00

0.00

0.60

0.00

0.00

0.00

16

0.00

0.09

0.00

0.00

0.00

0.00

0.00

0.91

0.00

0.00

0.00

17

0.00

0.00

0.09

0.00

0.00

0.03

0.00

0.00

0.00

0.00

0.00

18

0.00

0.00

0.03

0.00

0.00

0.00

0.00

0.00

0.00

0.00

0.00

19

0.00

0.00

0.17

0.07

0.00

0.00

0.17

0.77

0.00

0.00

0.00

20

0.00

0.00

0.17

0.03

0.00

0.03

0.10

0.73

0.00

0.00

0.00

21

0.00

0.17

0.00

0.00

0.00

0.00

0.07

0.00

0.00

0.00

0.00

22

0.20

0.07

0.43

0.00

0.00

0.03

0.03

0.83

0.00

0.00

0.00

23

0.07

0.20

0.47

0.00

0.00

0.00

0.00

0.00

0.00

0.00

0.00

24

0.00

0.21

0.05

0.00

0.05

0.00

0.21

0.73

0.79

0.00

0.00

25

0.15

0.00

0.00

0.00

0.00

0.00

0.00

0.75

0.00

0.00

0.15

k = Carcinophora (Carcinophoridae), 1 = cockroach (Blattidae), m = Noctuidae, n = Pyr-
alidae, o = small Staphylinidae, p = tiny Staphylinidae, q = Scarabidae, r = Curculionidae,
s = Carabidae, t = Wyeomyia (Culicidae), u = Culex (Culicidae), v = Trichoprosopon (Culici-
dae), w = Toxorhynchites (Culicidae).

Results

The first 3 components accounted for a total of 68.13% of the variation
of Heliconia insect communities. The component weights for each of the
first 3 components are listed in Table 4. Component 1 weights most heavily,
in decreasing order of weight, on Quichuana (Diptera: Syrphidae), Gillisius
(Coleoptera: Hydrophilidae) and Wyeomyia (Diptera: Culicidae). These are
the most frequently encountered and abundant insect species in many Hel-
iconia inflorescences. The frequencies of Quichuana and Wyeomyia are
positively correlated in Heliconia collections (Table 3). Component 2
weighs most heavily on Culex (Diptera: Culicidae), Merosargus (Diptera:
Stratiomyidae) (a negative weight) and Copestylum (Diptera: Syrphidae) (a
negative weight). This component weighs most heavily on a second group

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NEW YORK ENTOMOLOGICAL SOCIETY

Table 3. Matrix of pairwise correlations among the frequencies of occurrence of 23 insect
morpho species from 25 different Heliconia collections.

Insect

morpho

species

Insect morpho species

a

b

c

d

e

f

g

h

i

j

k

1

a

1.000

b

0.306

1.000

c

0.176

0.262

1.000

d

0.197

0.077

0.139

1.000

e

0.313

0.287

0.043

-0.071

1.000

f

0.170

0.201

0.467

0.159

-0.104

1.000

g

0.115

0.214

-0.023

-0.228

0.013

0.309

1.000

h

0.142

0.466

0.172

-0.066

0.438

0.085

0.052

1.000

i

0.154

0.153

0.103

-0.202

-0.247

0.420

0.098

0.046

1.000

j

-0.015

0.518

0.485

-0.076

0.281

0.347

0.239

0.652

0.155

1.000

k

-0.014

0.286

0.276

-0.010

-0.207

0.496

0.360

0.157

0.313

0.493

1.000

1

0.080

0.301

0.042

0.418

0.189

0.361

0.238

0.308

-0.186

0.284

0.460

1.000

m

0.264

-0.084

-0.077

-0.261

0.302

-0.327

-0.062

0.368

0.039

0.236

-0.041

0.055

n

0.188

0.292

0.257

0.064

0.388

-0.258

0.105

-0.010

-0.313

0.036

-0.376

-0.166

o

0.160

0.259

0.004

-0.064

0.441

-0.108

0.397

0.368

-0.035

0.418

0.204

0.243

P

0.167

0.309

0.036

0.516

0.217

0.350

0.216

0.038

-0.199

0.106

0.378

0.843

q

0.246

0.205

-0.139

-0.351

0.016

-0.083

0.033

0.377

0.132

-0.042

0.002

-0.150

r

-0.294

0.148

-0.059

0.081

0.129

-0.001

-0.120

0.491

-0.086

0.578

0.139

0.157

s

0.070

0.289

0.036

-0.039

0.496

0.147

0.210

0.331

-0.180

0.082

0.090

0.462

t

0.788

0.278

0.221

0.305

0.440

0.282

-0.069

0.325

0.099

0.007

-0.136

0.281

u

-0.115

-0.497

0.180

0.092

-0.182

-0.035

-0.303

-0.193

-0.185

-0.325

-0.314

-0.205

V

0.179

-0.215

0.315

-0.196

-0.033

0.411

-0.033

0.125

0.170

-0.104

-0.055

-0.060

w

0.336

-0.108

-0.244

0.143

-0.073

-0.068

-0.168

-0.201

0.041

-0.225

-0.198

-0.077

of commonly encountered species all of which are mosquito and fly species.
Component 3 weighs most heavily on insects associated with the first two
components: Copestylum (a negative weight), Gillisius (a negative weight)
and Wyeomyia.

The results of the principal components analysis are presented graphically
in Figures 2, 3 and 4. Figure 2 shows a plot of component 1 versus com-
ponent 2. The collections from the French Antilles (10, 7, 9, 11) are grouped
in the lower left region of the graph. These collections have low insect
species richness. In particular, hydrophilid, hispine and staphylinid beetles
which are common members of many Heliconia insect communities, are
not found in the island collections. Further, in 3 of the collections (7, 9, 1 1)
only 1 genus of mosquito ( Culex ) was recorded, while up to 4 genera are
sometimes found in mainland collections. On the right edge of the graph are
collections (24, 12, 15, 8, 3) from various locations which, with the exception
of H. wagneriana Petersen (24), have morphologies similar to H. cf. cari-
baea Lamarck (Fig. 1). This graph also shows close association of insect
community structure among collections of the same Heliconia species from
the same locations. Thus, both the early (17) and late (18) collections of H.
bihai L. from Rancho Grande (Venezuela), the early (19) and late (20) col-
lections of H. imbricata (Kuntze) Baker and Rincon de Osa (Costa Rica)

VOLUME LXXXIX, NUMBER 2

117

Table 3. Continued.

Insect Insect morpho species

morpho —

species mnopqrstuvw

a

b

c

d

e

f

g

h

i

j

k

1

m

1.000

n

-0.035

1.000

o

0.385

0.375

1.000

P

-0.180

0.104

0.249

q

0.043

-0.123

-0.077

r

0.046

-0.187

0.264

s

-0.121

0.166

-0.040

t

0.205

0.118

-0.013

u

-0.274

-0.077

-0.326

V

-0.107

-0.126

-0.117

w

0.229

-0.203

-0.227

1.000

-0.133

1.000

-0.029

-0.006

1.000

0.410

0.194

-0.092

1.000

0.324

0.197

-0.224

0.222

-0.197

-0.118

0.014

-0.005

-0.071

-0.079

0.114

-0.125

-0.114

-0.127

-0.184

-0.117

1.000

-0.068

1.000

-0.289

0.474

1.000

-0.187

-0.037

0.067 1.000

and the early (21) and late (22) collections of H. wagnerianci from Rincon
de Osa are located close to one another.

Considering Figure 3 next, which plots component 1 versus component
3, much of the same clustering as in Figure 2 is shown. Heliconia species
which were collected twice from the same location cluster together, Heli-
conia species with morphologies similar to H. cf. caribaea cluster together
and the Antillean collections border the left side of the graph. Here, how-
ever, the Antillean collections (10, 7, 11,9) enclosed the collection of H.
episcopalis Velloso from Primavera, Ecuador (6). H. episcopalis is a species
with small floral bracts and contains few insects. Thus, the generation of
low insect diversity Heliconia plants occurs both because of the isolation
of some Heliconia species on islands and because of the small inflorescence
size of H. episcopalis.

The final graph, Figure 4, which compares component 2 versus compo-
nent 3, again shows groupings of the species from the Antilles, clustering
of collections of the same Heliconia species collected at different times and
clustering of Heliconia species with similar floral morphologies. Figure 4,
however, also shows that H. rostrata from Limoncocha, Ecuador (13) lies
on the right side of the graph rather distant from most other collections. H.
rostrata represents the only Heliconia species studies which had pendant

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Table 4. Component weights on 23 insect morpho species from 25 Heliconia collections.

Component number

species

1

2

3

4

a

0.667

-0.034

0.125

-0.049

b

0.546

0.097

-0.188

0.074

c

0.360

-0.128

-0.191

0.006

d

0.374

-0.245

0.031

0.011

e

0.340

0.083

0.056

0.204

f

0.270

-0.055

-0.167

-0.150

g

0.162

0.053

-0.065

0.002

h

0.126

0.042

-0.029

0.035

i

0.103

0.014

-0.034

-0.050

j

0.169

0.028

-0.114

0.037

k

0.052

-0.000

-0.044

-0.013

1

0.080

-0.001

-0.030

-0.013

m

0.027

0.011

0.016

0.006

n

0.052

0.005

-0.006

0.035

o

0.100

0.047

-0.017

0.056

P

0.007

-0.001

-0.002

-0.001

q

0.023

0.010

0.007

-0.003

r

0.008

-0.000

-0.006

0.007

s

0.037

0.007

-0.003

0.020

t

0.527

-0.039

0.142

-0.079

u

0.117

-0.290

0.039

0.113

V

0.020

-0.024

0.007

-0.018

w

0.035

-0.010

0.029

-0.015

NOTE: Insects listed by letters follow the same sequence as that listed in Table 2.

inflorescences but still contained an aquatic insect community. While this
community includes many of the common insects, it is unusual in that nei-
ther syrphid fly genera nor hispine beetle larvae were represented in the
collection.

Discussion

The principal components weigh heavily on some of the most common
fly, mosquito and beetle species. This implies that much of the variation in
these insect communities from the 25 Heliconia collections is dependent on
differences in Quichuana , Gillisius, Wyeomyia, Culex, Merosargus and
Copestylum frequencies. The resulting placement of the 25 Heliconia col-
lections on the principal components analysis graphs (Figs. 2, 3, 4) can be
interpreted in relationship to knowledge about Heliconia collection locali-
ties, bract morphologies and flowering phenologies. The principal compo-
nents analysis shows most clearly the similarity of the depauperate fauna
of Heliconia inflorescences from the Antilles. Low species richness among

VOLUME LXXXIX, NUMBER 2

119

FIRST COMPONENT

Fig. 2. A plot of component 1 versus component 2 for 25 collections of Heliconia insect
communities. The numbers are the same as those listed in Table 1.

island biota is a commonly observed phenomenon (MacArthur and Wilson
1967; Carlquist 1974). In the case of Heliconia insect communities, this
reduction of species richness is largely the result of a lack of beetle species
and the presence of only 1, instead of several, mosquito species. I suspect
that the lack of these insects is simply the result of an inability of some
insects to locate, invade and colonize the islands. In Guadeloupe I was able
to maintain Ecuadorian hispine species in petri dishes by feeding them
bracts of both H. bihai Linn, and H. caribaea from local Heliconia popu-
lations. Thus, it seems clear that these beetles could form a breeding pop-
ulation on Guadeloupe. That they have not done so is probably attributable
to their low rates of movement. Other research (Beaman 1980) has shown
that Heliconia feeding hispines move only short distances among Heliconia
clumps and do not exhibit long range dispersal.

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Fig. 3. A plot of component 1 versus component 3 for 25 collections of Heliconia insect
communities. The numbers are the same as those listed in Table 1.

Further, from this biogeographic work we can interpret that floral mor-
phologies are important in generating particular Heliconia insect community
structure. Small Heliconia species, such as H. episcopalis, not only have
low species richness but also are devoid of some of the most frequent Hel-
iconia inquilines, syrphid fly larvae. Clearly, the kind of floral morphology
of pendant Heliconia species is also important in determining the Heliconia
insect community. While most pendant species do not contain an aquatic
insect community (Seifert and Seifert 1979; personal observation), H. ros-
trata, because of its compressed bracts, does contain water and harbors
some of the insect species associated with other Heliconia species. Finally,
Heliconia with similar morphologies, such as H. cf. caribaea species, may
contain largely similar Heliconia insect communities.

The results of the principal components analysis also point out that Hel-
iconia inflorescences of the same species at the same location have similar
Heliconia insect communities even if they are collected during different
portions of the blooming season. The insect community structure, based on

VOLUME LXXXIX, NUMBER 2

121

Fig. 4. A plot of component 2 versus component 3 for 25 collections of Heliconia insect
communities. The numbers are the same as those listed in Table 1.

the presence of at least one individual of each insect species, remains ap-
proximately the same throughout much of the blooming season of a Heli-
conia species.

This principal components analysis did not show clustering of different
Heliconia species from the same location. Thus, H. aurea Rodriguez from
Rancho Grande (1) did not cluster close to H. bihai from Rancho Grande
(17, 18), and H. imbricata from Rincon de Osa (19, 20) did not cluster with
H. wagneriana from the same location (21, 22). Only in the cases of the
collections from the Antilles, where isolation is important in determining
insect species richness, did different Heliconia species from the same lo-
cation show affinities.

In conclusion, this study has shown how principal components analysis
can be used to examine the similarity of insect communities living in asso-
ciation with different species of closely related plants from different loca-
tions. The results of the analysis indicate the importance of plant isolation
and floral morphology in determining insect community structure.

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NEW YORK ENTOMOLOGICAL SOCIETY

Acknowledgments

I thank the many people who helped me with the field work: Florence H.
Seifert (Costa Rica, Venezuela, Ecuador), Guido Pereira S. (Venezuela),
Nancy J. Beaman (Costa Rica, Venezuela) and Roberto Barrera R. (Vene-
zuela). I also thank Nancy J. Beaman for tabulating data, Joseph L. Russo
for computer assistance and Florence H. Seifert for comments on the manu-
script.

This research was supported financially by National Science Foundation
grants GB-30542 (Costa Rica) and DEB-79-06593 (Venezuela), The Tropical
Science Center (Costa Rica) and The George Washington University Com-
mittee on Research (Venezuela, Ecuador, Martinique, Guadeloupe).

Literature Cited

Aristeguieta, L. 1961. El genero Heliconia en Venezuela. Instituto Botanico. Direccion de
Recursos Naturales Renovables. Ministerio de Agricultura y Cria. Caracas.

Beaman, N. J. 1980. Dispersion, dispersal and phenotypic variability of Xenarescus mono-
cerus, a Neotropical rolled-leaf hispine beetle. M.S. thesis. The George Washington
University.

Carlquist, S. 1974. Island biology. Columbia University Press, New York.

Daniels, G. S. and F. G. Stiles. 1979. The Heliconia taxa of Costa Rica. Keys and descrip-
tions. Brenesia 15:1-150.

Darwin, C. 1959. The origin of species by means of natural selection. Murray, London.
MacArthur, R. H. and E. O. Wilson. 1967. The theory of island biogeography. Princeton
University Press, Princeton, New Jersey.

Pianka, E. R. 1978. Evolutionary ecology. Harper and Row, New York.

Pielou, E. C. 1977. Mathematical ecology. John Wiley and Sons, New York.

Price, P. 1975. Insect ecology. John Wiley and Sons, New York.

Seifert, R. P. 1975. Clumps of Heliconia inflorescences as ecological islands. Ecology
56:1416-1422.

. 1980. Mosquito fauna of Heliconia aurea. Journal of Animal Ecology 49:687-697.

and F. H. Seifert. 1976. Natural history of insects living in inflorescences of two

species of Heliconia. Journal of the New York Entomological Society. 84:233-242.

and . 1979. Utilization of Heliconia (Musaceae) by the beetle Xenarescus mon-

ocerus (Oliver) (Chrysomelidae: Hispinae) in a Venezuelan forest. Biotropica 11:51-59.
Sneath, P. H. A. and R. R. Sokal. 1973. Numerical taxonomy. W. H. Freeman and Co., San
Francisco.

Stiles, F. G. 1975. Ecology, flowering phenology, and hummingbird pollination of some Costa
Rican Heliconia species. Ecology 56:285-301.

. 1979. Notes on the natural history of Heliconia (Musaceae) in Costa Rica. Brenesia

15: 151-180.

Strong, D. R., Jr. 1977a. Insect species richness: hispine beetles of Heliconia latispatha.
Ecology 58:573-582.

. 1977b. Rolled leaf hispine beetles (Chrysomelidae) and their Zingiberales host plants

in Middle America. Biotropica 9:156-169.

Wallace, A. R. 1876. The geographical distribution of animals. 2 vols. Harper, New York.

Department of Biological Sciences, The George Washington University,
Washington, D.C. 20052.

Received for publication November 26, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(2), 1981, pp. 123-142

NOTES ON SEASONALITY AND HABITAT ASSOCIATIONS OF
TROPICAL CICADAS (HOMOPTERA: CICADIDAE) IN
PREMONTANE AND MONTANE TROPICAL MOIST
FORESTS IN COSTA RICA

Allen M. Young

Abstract — Annual patterns of adult emergences and habitat associations
for several species of tropical cicadas (Homoptera: Cicadidae) were studied
within the premontane-to-montane tropical moist forest zone northwesterly
of the Meseta Centrale region of Costa Rica. The region largely consists of
expansive coffee plantations, secondary vegetation, and strips of forest
along streams, and the censusing of cicadas at four widely separated local-
ities (Grecia, Naranjo, San Ramon, and Esparta) covered an elevational
range of about 200-1,100 meters. Coffee plantations in this region have an
overstory (shade cover) of Inga spp. trees (Leguminosae) and sometimes
(Grecia) a mixture of these trees with another legume tree, Zygia longifolia.
Z. longifolia is also the dominant tree in forest remnants along streams in
this region. Given the pronounced patterns of annual seasonal precipitation
at all four localities and the mosaic of secondary coffee plantations and
forest remnants at three of them, and forest remnants and secondary vege-
tation at the fourth (Esparta), it was expected that the census of nymphal
skins and adult cicadas would indicate distinctive seasonal adult emergence
cycles and habitat associations. The data confirmed this. Fidicina amoena
emerges during the dry season near stream-edge Zygia trees in coffee plan-
tations (Grecia) and near other legume trees in forest remnants (Esparta).
Fidicina semilata and F. “ coffea ” emerge near Inga trees in coffee plan-
tations (Naranjo and San Ramon) during the dry season as does F. pronoe
and F. spinocosta in secondary vegetation (Esparta). Fidicina mannifera
emerges during the wet season in primary forest remnants at about 200
meters elevation (Esparta). Zammara smaragdula emerges in the wet sea-
son near Zygia trees in coffee plantations (Grecia) and forest remnants along
streams (Naranjo and San Ramon) and Conibosa sp. emerges during the
wet season in coffee plantations (Grecia, Naranjo, and San Ramon). Que-
sada gigas emerges during the dry season near Zygia trees along the borders
of coffee plantations and in stream-edge forest remnants. The data are dis-
cussed from the standpoint of adaptations in tropical cicadas to seasonality
and habitats, the latter particularly in terms of the exploitation of legume
trees as feeding sites (root crowns) of nymphs.

Introduction

A field survey of the cicadas across northern Costa Rica, extending from
lowland tropical wet forest into montane wet forest, montane moist forest

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and into lowland tropical dry forest, revealed distinctive distributional pat-
terns for many genera and species (Young 1976). Such a transect slices
through a series of distinctive climatic and vegetation zones (Holdridge et
al. 1971) and varying degrees of seasonality in terms of the length and se-
vereness of the dry season (Janzen 1967). Several previous studies have
revealed that tropical cicadas may exhibit very pronounced adult seasonal
emergences annually, and that genera and species sometimes also partition
the local environment by habitat (Young 1972, 1974, 1975, 1980a, b, 1981a).
These studies were conducted for the common genera and species found in
these regions of Costa Rica: lowland tropical wet forest, premontane trop-
ical wet forest, highland moist forest, and lowland tropical dry forest. The
purpose of the present paper is to summarize the annual peak adult emer-
gence periods and habitat associations for several genera and species of
tropical cicadas in yet another major climatic belt of northern Costa Rica,
the premontane-to-montane tropical moist forest zone. The data, although
discontinuous, indicate that some cicadas of this zone emerge during the
lengthy dry season, others during the wet season, and that most species are
differentially associated with (a) forest remnants forming the borders of
coffee plantations and grazing pastures, (b) shade trees in coffee plantations,
and (c) secondary vegetation. The data are consistent with observations on
cicada emergences at other sites in Costa Rica in that emergences are strik-
ingly seasonal, and that a variety of habitats, including agricultural ones,
are exploited by cicadas in the tropics.

Materials and Methods

I examined the temporal and spatial properties of adult emergences of
cicadas by selecting four widely separated localities along the northwest
axis through the highlands and adjacent mountains northwest of the Meseta
Centrale region of Costa Rica: habitats located at the cities of Rosario de
Grecia (Grecia), Naranjo, San Ramon, and Esparta, sites within Alajuela
and Puntarenas Provinces (Fig. 1). This transect covers an elevational gra-
dient from 825 meters (Grecia), 1042 meters (Naranjo), 1116 meters (San
Ramon) down to 208 meters (Esparta). The first three localities are very
similar topographically and floristically, consisting of rolling hills covered
with extensive coffee plantations at least 100 years old in most parts, and
with strips or remnants of forest vegetation along streams and rivers (Figs.
2, 3, 4). The coffee plantations have an overstory of primarily Inga spp.
trees (Leguminosae) (Figs. 2, 3) and the forest remnants consist chiefly of
adult-size Zygia longifolia trees (Leguminosae) (Fig. 4). The coffee plan-
tation at Rosario de Grecia consisted of adult-size Zygia trees along streams
and drainage ditches running through the plantings, and Inga trees as shade
on slopes. Patches of grasslands are interspersed with the coffee agricultural

VOLUME LXXXIX, NUMBER 2

125

Fig. 1 . The distribution of localities within the premontane-to-montane tropical moist forest
zone in Costa Rica, northwest of the Meseta Centrale and San Jose, used to study cicada
emergences. These localities are: (1) Grecia, (2) Naranjo, (3) San Ramon, and (4) Esparta.

system of the region. The fourth site, Esparta, is a very steep (incline about
60° in most places) forested ravine surrounded by patches of secondary
growth and open pastures lightly to heavily grazed by Brahma cattle (Fig.
5). In this forest bordering a stream system, several genera of legume trees
predominate, of which Pithecollobium species predominates. A marked dry
season, of similar duration and severeness, is a major component of the
climatic regime of all four sites (Fig. 6). Areas within each habitat studied
at each locality are representative of the habitats in general.

A cicada census program was deployed at these four localities to test the
hypothesis that representative cicada species at each one exhibit distinct
annual emergence patterns somehow related to the annual seasonal cycle
of precipitation (Fig. 6). A second hypothesis examined, also in a prelimi-
nary fashion, was that cicada emergences occur in different kinds of habitats
at each locality, habitats that are representative of each locality overall. As
with previous similar studies (Young 1972, 1974, 1975, 1980a, b, 1981a)
cicada emergence patterns were studied by making collections of freshly

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NEW YORK ENTOMOLOGICAL SOCIETY

Fig. 2. The overstory of Inga spp. trees in the coffee plantation at Naranjo where emer-
gences of Fidicina semilata and Conibosa sp. were studied. These shade trees, easily recog-
nizable by the whitish bark, are rather uniformly distributed over large areas of the plantations
in this locality.

discarded nymphal skins from marked plots in different habitats at each
locality. The skins were collected, identified in the field to species, sexed,
and counted every time a census was made. Whenever possible, adult spec-
imens were also collected for voucher material to confirm species deter-
minations later. Sample plots for collecting nymphal skins were usually 5 x
6 meters and in coffee plantations, about half were enclosing one Inga shade
tree and the others only coffee bushes. Plots with shade trees also have
coffee bushes, and the DBH for the shade trees is 20-35 cm. Nymphal skins
were collected exhaustively from tree trunks, bushes, leaf litter, and ground
surface. For most species, skins are readily identified to species based on
gross morphological features (size, form, coloration). The reliability of
species determinations of skins in this manner has been discussed elsewhere
(Young 1980a). Because skins remaining from a previous year’s emergence
period are very discolored, broken, and crumbly, they are readily distin-
guished from fresh skins of the current emergence thereby eliminating a bias

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127

Fig. 3. Inga spp. shade trees interspersed with coffee bushes in a plantation at San Ramon.
Cicada nymphal skins were collected from the ground litter, coffee bushes, and trunks of shade
trees, and special care was taken to examine branches cut away as part of pruning of coffee
bushes as shown here.

of confusing two or more annual emergence periods. Records were also
kept for adult cicadas heard or seen at times when nymphal skins were
censused.

For the Rosario de Grecia locality the study plots were situated within
two areas of a large coffee plantation, the selection of the plots being based
on previous observations of nymphal skins in them. Prior visits to all four
localities indicated that at least two large-bodied species of cicadas were
active at each one. The Grecia coffee plantation has a mixed shade cover
of Z. longifolia and Inga spp. trees, while the Naranjo and San Ramon
plantations have several species of Inga trees as ahde. In addition to setting
up sample plots in coffee in Naranjo and San Ramon, I also established a
large study plot along a forest remnant in San Ramon, adjacent to a coffee
plantation. Z. longifolia is the only tree species in this plot. At Esparta,
cicada nymphal skin distributions were examined by selecting plots around
some adult-size legume trees where skins had been seen. Details of the sizes

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Fig. 4. Stream-edge habitat near San Ramon showing a large Zygia longifolia tree (on right
bank of stream), a typical emergence spot for cicadas such as Zammara smaragdula.

of plots, total areas represented by sample sites, and census dates are sum-
marized along with the data. The census program spanned a period of about
two years and included samples taken in both wet and dry seasons.

Because my samples of nymphal skins are very discontinuous over each
year of the study, one source of error in estimating density of emerging
cicadas from my census program is the possibility that some nymphal skins
were blown into the plots from other areas of the habitats studied, although
more than 80% of the skins collected from all plots throughout the study
were found attached to vegetation. Unlike some temperate zone cicadas,
clean emergence holes are not commonly found in tropical species.

Results

The cicadas associated with each locality are shown in Figures 7 and 8
and can be described further as follows:

(A) GRECIA: Fidicina amoena • medium to large-bodied cicada (body

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129

Fig. 5. An overview of the canopy of the primary forest remnant in the steep ravines at
Esparta during the dry season. There are also surrounding patches of secondary vegetation in
addition to pastures. The sides of the ravines with forest are steep (45-60°).

length 30 mm), greenish body all over, clear wings, bell-like chirp for court-
ship song. Also found at Naranjo, San Ramon, and Esparta, although not
abundant. Common in some parts of the Meseta Centrale (Young 1980a),
but not common in lowland and premontane tropical rain forest (Young
1972, 1980b). Probably scarce in lowland tropical dry forest, although a
population occurs at Santa Rosa National Park (Young 1981a).

Zammara smaragdula • medium-sized cicada (body length 25 mm), dis-
tinctly mottled green and black body all over, bright green in male, more
olive green in female; wings clear but with large black spots; “hoarse buzz”
for courtship song. Also found at other localities mentioned above for F.
amoena but excluding tropical rain forest sites in the northeast where it is
replaced by Z. smaragdina ; also found at Finca La Taboga, near Canas in
lowland Guanacaste Province as well as at the Barranca Forest site (Young
1981a).

Quesada gigas • large-bodied cicada (44 mm), body all over streaked in
shades of brown, olive green, and black; clear wings; very distinctive “lo-

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Aug. Sep. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May Jun. Jul.

SUCCESSIVE MONTHS

Fig. 6. The distribution of precipitation at the four localities within the premontane-to-
montane tropical moist forest zone, showing a distinct and similar dry season between Decem-
ber and April for all localities. Data are means for the period 1943-1973. The standard devia-
tions and coefficients of variation for monthly mean values are strikingly similar among all four
localities, as expected for the very similar mean values for each month among all localities.
Data is courtesy of the Meteorology Service of Costa Rica.

comotive whistle ’ -like courtship song, sometimes sings at dusk as do the
others mentioned here but to lesser degree; widely-distributed throughout
Costa Rica (Young 1976, 1980a, b).

Conibosa sp. • small-bodied cicada (10 mm) not figured in this paper, but
with greenish head and thorax and maroon-colored abdomen and clear
wings; courtship song resembles “zzzst” repeated over and over; widely
distributed in the Meseta Centrale north of San Jose, and through the three
coffee-growing localities discussed in this paper (see also Young 1980a), but
not found to my knowledge in lowland to premontane tropical wet and dry

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131

Fig. 7. Cicadas from the montane tropical moist forest zone in Costa Rica, northwest of
the Meseta Centrale. Top row, left to right: Quesada gigas , Zammara smaragdula, and Fi-
dicina amoena, three of the four common cicadas associated with predominantly coffee x
Zygia tree plantations in the Meseta Centrale, and lower montane moist forest region adjacent
to it; Bottom row, left to right; Fidicina semilata and Fidicina “ coffea two of three common
cicadas associated primarily with coffee x Inga tree plantations in the upper montane moist
forest region. It both series, the cicada not shown is th small Conibosa sp., but it is figured
in Young (1980a).

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Fig. 8. Cicadas from the premontane tropical moist forest zone in Costa Rica, northwest
of the Meseta Centrale. Top row, left to right: Fidicina mannifera and Fidicina sericans, two
of four common primary forest species. Bottom, left to right: Fidicina pronoe and Fidicina
spinocosta, the two common species associated with certain types of secondary vegetation in
this region. The third common primary forest species of this region is Zammara smaragdula,
figured in Fig. 7. For both Fig. 7 and 8, the vertical black line to the right of each cicada shown
gives the actual body length relative to the magnified figure. See text for actual body length

measures.

VOLUME LXXXIX, NUMBER 2

133

forest sites in Costa Rica. Appears to be distinctly associated with coffee
plantations, perhaps more so than other cicadas.

(B) NARANJO: Fidicina semilata • species determination tentative as it
may be a new species also tentatively called F. “ guayabana ” by A. M.
Young and T. E. Moore; medium-sized cicada (body length 22 mm) with
body generally greenish all over and wings crystal-clear; monotonous buzz-
like courtship song and sings in coffee bushes and shade trees in coffee
plantations; as to my knowledge found only at San Ramon in Costa Rica in
addition to Naranjo.

(C) SAN RAMON: Fidicina “ coffea ” • probably a new species, tenta-
tively determined by A. M. Young and T. E. Moore; medium-sized cicada
(body length 23 mm), greenish but with some mottling (faint and thin) in
brown, similar to F. semilata but with very distinctive courtship song con-
sisting of pulsating buzz from coffee bushes and shade trees in plantations;
not collected from other localities in Costa Rica; along with F. semilata
appears somewhat restricted to the coffee habitat between Naranjo and San
Ramon. May occur at Naranjo but not found.

(D) ESPARTA: Fidicina mannifera • large-bodied cicada (42 mm body
length), dark brown with some mottling in green, particularly on prothorax
dorsum; wings clear but with somewhat diffuse brown tinges along veins;
very distinctive strong pulsating buzz, particularly at dusk; sings from
trunks of large forest trees; widely distributed in northern Costa Rica out-
side of the Meseta Centrale and adjacent foothills (see Young 1972, 1976,
1980a, b).

Fidicina pronoe • medium-sized cicada (body length 25 mm) with co-
coa-brown meso- and meta-thoracic dorsum and greenish head and pro-
thoracic dorsum and lateral borders of meso- and meta-thoracic dorsum
lined with white; wings clear and abdomen thickly banded laterally with
black and brown; courtship song a high pulsating whistle heard mostly in
open secondary vegetation where the canopy height is less than 7 meters;
widely-distributed across northern Costa Rica (Young 1976, 1980b) and
probably occurring at at least Naranjo and Grecia localities in the present
study although most likely patchy and scarce as it is in the Meseta Centrale
(Young 1980a). Probably not found in most localities in lowland tropical dry
forest in Costa Rica but perhaps in Santa Rosa National Park (Young 1981a).

Fidicina spinocostam at lower end of medium-sized range (body length 20
mm), body with head and thoracic areas bright green and abdomen maroon
with silvery pubescence; body short and thick relative to other Fidicinas
and wing length; wings crystal clear; courtship song is strongly pulsating
short whistles mostly at dusk and from thick secondary vegetation; to my
knowledge only occurs at this locality and also in the premontane tropical
wet forest zone in the northeast (Young 1980a).

Fidicina sericansm large-bodied cicada (body length 30 mm) with black

Table 1 . Notes on the seasonal distribution of adult cicadas along a transect through the Cordillera Central northwest of San Jose, San Jose
Province, Costa Rica.

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NEW YORK ENTOMOLOGICAL SOCIETY

* The localities examined are: Rosaria de Grecia (RG), Naranjo (N), Esparta (E), San Ramon (SR).

** Species designations unconfirmed due to lack of information on genus Fidicina in general. Fidicina semilata is also coded as F. “ guaya-
bana ” in MPM and UWMZ Collections, and F. “ coffea ” is a temporary designation.

VOLUME LXXXIX, NUMBER 2

135

body mottled with dark green; wings slightly smokey with tinges of brown;
courtship song a steady buzz; associated with primary and advanced sec-
ondary wet to moist forest habitats in northeast premontane, lower mon-
tane, and lowlands with nymphal skins invariably found near adult-size can-
opy legume tree species (Young 1972, 1980b, unpublished).

Another small-bodied cicada, Pacarina spp., also occurs at Esparta and
perhaps at San Ramon, where they are associated with grassland habitats
(Young 1974) but these species were not studied here. A summary of the
distributions, localities, habitats or adult cicadas and nymphal skins, are
given for these species in Table 1.

As shown in Table 1, forms such as F. amoena, F. “ coffea , ” F. semilata
F. sericans, and F. pronoe are active in the dry season at their respective
localities and many of these species occur in coffee plantations and other
disturbed habitats. Z. smaragdula is the common wet season cicada at
several localities and it is associated with forest remnants, places where its
nymphal skins are found beneath Zygia trees. There is considerable overlap
for some species of Fidicina (those associated with coffee plantations)
among several coffee-growing localities and Z. smaragdula is found at all
localities studied (Table 2). Forest-associated Fidicinas are restricted to
Esparta (Table 2). Although the data on emergences are discontinuous, peak
numbers of nymphal skins for several species censused agree with seasonal
activity patterns as seen from adults (Fig. 9). Densities of nymphal skins
generally fall well below one nymphal skin per square meter, although cen-
suses of the same species at different sites within a locality generate great
differences in estimates of density (Table 3). In coffee plantations at Naranjo
and San Ramon, the greatest abundances of nymphal skins of F. semilata
and F. “ coffea ” occur in plots with Inga spp. shade trees (one tree per
plot) and plots with only coffee bushes beyond the immediate vicinity of
shade trees disclose few skins (Table 4).

Density estimates for cicada nymphal skins will be very sensitive to areas
involved, particularly when data from two or more sites are polled when
individual sample sizes are low. Yet for a species such as F. semilata, the
low estimate of about 0.12 nymphal skins per square meter may be biolog-
ically significant as suggested by the rather continous data on nymphal skins
of this cicada obtained from a small patch of coffee (840 m2) within the city
of Naranjo for the period 30 December 1972 through 28 April 1973, a time
span covering the major portion of the emergence period of this species at
this locality. A total of 97 fresh skins were collected from this coffee patch,
which contained a living (freshly-cut) stump of an Inga tree (DBH-25 cm)
and about 50 coffee bushes giving a density estimate of about 0. 12 skins per
square meter. More than 70% of the skins collected were located within a
3-meter radius of the tree stump. The wood of the stump was still fresh and
green as the tree had been cut down about a month before my census was
commenced.

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Table 2. The distribution of cicada species in the Central Cordillera northwest of San Jose,
in Costa Rica, based on records of adults and nymphal skins.

Cicada

Localities

Rosaria
de Grecia
(coffee farms)

Naranjo
(coffee farms)

San Ramon
(coffee farms)

E sparta
(forested
ravines;
second
growth)

Zammara smaragdula

+

+

+

+

Quesada gigas

+

?

?

+

Conibosa sp.

+

+

+

Fidicina amoena

+

+

+

F. semilata

+

+

+

F. “ coffea ”

+

+

F. pronoe

+

F. spinocosta

+

F. mannifera

+

F. sericans

+

Pacarina sp.

+

+

There is some evidence of habitat partitioning at Rosaria de Grecia in
terms of cicadas associated with Zygia trees: F. amoena, Z. smaragdula,
and Q. gigas all emerge near these trees but not at other trees used as shade
here. Furthermore, Z. smaragdula exhibits a tendency to emerge close to
streams: of 174 nymphal skins of this cicada collected on 23 July 1973 along
five parallel rows of coffee bushes running along a stream bordered with

Table 3. Abundance and densities of nymphal skins of cicadas from study plots at several
localities in the Central Cordillera of Costa Rica (1973).

Cicada species

Localities studied

Habitat studied

Total area
examined

Num-

ber

of

sites

Densities of
nymphal skins

Fidicina amoena

Rosaria de Grecia

coffee plantation

2,030 m2

2

0.17, 0.27/m2

Fidicina semilata

Naranjo

coffee plantation

1,256 m2

2

0.12, 0.40/m2

San Ramon

coffee plantation;
riveredge forest
remnant

1,128 m2

2

0.12, 0.28/m2

Fidicina “coffea”

San Ramon

coffee plantation;
riveredge forest
remnant

1,128 m2

2

0.08, 0.28/m2

Zammara smaragdula

Rosario de Grecia

coffee plantation

2,030 m2

2

0.30, 0.42/m2

Conibosa sp.

Rosario de Grecia

coffee plantation

2,030 m2

2

0.06, 1.61/m2

San Ramon

coffee plantation;
riveredge forest
remnant

688 m2

2

0.41, 0.78/m2

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137

Table 4. Collections* of nymphal skins of Fidicina spp. cicadas proximal and distal to Inga
spp. shade trees in coffee plantations at Naranjo and San Ramon in the central highlands of
Costa Rica northwest of the Meseta Centrale.

No. Inga
trees**
censused

DBH range

Tree

trunks

Coffee

bushes

within

Range 16 m2 area

trunk (proximal)

Coffee
bushes
beyond
16 m2 area
(distal)

Ground

Total nymphal skins

trees
+ coffee)

oo

oo

T

Naranjo: Fidicina semilata

26

25-35 cm

35

0-12 108

5

12

93

67

160

San Ramon: Fidicina semilata

33

25-35 cm

23

0-2 127

8

11

87

59

146

Fidicina “ coffea ”

33

25-35 cm

16

0-2 23

2

10

22

19

41

* Over 60% of the shade trees sampled were I. aff. fissicalyx and the remaining ones were
I. goldmannii and I. leptoloba.

** Census dates for Naranjo: 29 December 1972, 11 February 1973, 17 & 28 April 1973;
census dates for San Ramon: 6-9 January 1973, 7 February 1973.

Zygia, 100 skins were taken from the row of bushes nearest to the stream,
and with a marked decline in other rows, with none being found in the fourth
and fifth rows away from the stream. In areas of coffee at Grecia where
Inga trees are mixed with Zygia for shade, nymphal skins of Q. gigas are
found beneath both kinds of trees.

In one stream-edge plot (600 m2) containing six large Zygia trees and 25
coffee bushes on a steep incline of about 50°, two census dates (6 January
and 7 February 1973) disclosed 69 fresh skins of F. semilata and 100 fresh
skins of F. “ coffea ” with the distributions of the skins of the two species
mixed within the plot but not measured. These data and the accompanying
estimates of density (0.12 for semilata and 0.28 for “coffea") suggest co-
occurrence of both species over a relatively small portion of the coffee
plantation-border transitional habitat. Nymphal skins of these species are
also mixed under Inga spp. trees in upland coffee, usually with 1-2 skins
of each species per tree. Although my samples are small, the data suggest
that Z. smaragdula is associated with Zygia trees only in the context that
at both the Rosario de Grecia coffee habitat and San Ramon stream-edge
habitat, nymphal skins of this species are found beneath Zygia trees. When
Inga is used as a shade of upland coffee, only nymphal skins of F. semilata
and F. “ coffea ” and Conibosa sp. are found near these trees, but not those
of Z. smaragdula. Likewise, although Fidicinas are commonly found be-
neath Inga shade trees and even near the borders of Zygia with coffee,
most Zygia habitats lack Fidicina nymphal skins.

The habitat complex of narrow valley strips of primary forest and adjacent
patches of hillside secondary vegetation interspersed with grasslands at Es-

NUMBERS OF NYMPHAL SKINS

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200 -

A

/■»

115

100

85

60

45

30

15

0

200

100

85

60

45

30

15

0

500

100

85

60

45

30

15

SAN RAMON COFFEE PLANTATION
AND RIVER-EDGE FOREST

Fidicina
semilata T-

Fidicina

coffea

NO DATA

l l >

Conibosa sp.

NO DATA AVAILABLE
For Fidicina spp.

* I i

NARANJO COFFEE PLANTATION

Fidicina semilata

NO DATA

GRECIA COFFEE PLANTATION

Fidicina

amoena

313 SKINS

4

464 SKINS

4

Zammara

smaragdula

Quesada
gigas

JSSO Li

1 229 SKINS

JL|4

Conibosa sp.

Conibosa sp.

Dec Jan Feb Mar Apr May Jun Jul Aug

1972 1973-

Jan Feb Mar Apr May Jun Jul Aug

•1974

MONTHS

VOLUME LXXXIX, NUMBER 2

139

parta (Fig. 5) is characterized by rather low density emergences of cicadas,
some, such as F. pronoe and F. spinocosta associated with secondary vege-
tation and others such as F. mannifera, F. sericans and F. amoena asso-
ciated with primary forest. In one approximately 210 m2 primary forest plot
containing one canopy-like Pithecollobium tree censused once in the dry
season (10 January 1973), fresh nymphal skin abundances were: amoena :
20 and sericans : 12. A second smaller plot in the same ravine produced six
nymphal skins of sericans and one amoena. The larger plot during the wet
season (13 July 1974 census) produced two fresh skins of F. mannifera and
one fresh skin of Z. smaragdula. Nymphal skins of F. pronoe and F. spi-
nocosta were not searched for.

Discussion

Based on the limited data from this study, it is concluded that cicadas
within the premontane-to-montane tropical moist forest zone in Costa Rica
exhibit distinctive seasonal emergence patterns. Two seasonal adult emer-
gence patterns are recognized based on my study: "dry season or tropical
summer” cicadas, which include F. semilata, F. " coffea , ” F. amoena, F.
pronoe, F. spinocosta, F. sericans, and Quesada gigas, and "wet season
or tropical winter” cicadas such as F. mannifera, Z. smaragdula and Con-
ibosa sp. Therefore there is some evidence for allochronic annual emergence
strategies in tropical cicadas within this major climatic and vegetation zone
of Costa Rica, and presumably, to varying degrees, elsewhere in Central
America. Extensive data on the geographical distribution of cicadas else-
where in Central America are lacking. With the exceptions of F. semilata
and F. " coffea , ” all of the other cicadas studied have been observed to
have the same seasonal adult emergence patterns in other major climatic
and vegetation zones of Costa Rica (Young 1972, 1976, 1980a, b, 1981a; A.
M. Young, unpublished data).

With the exceptions of but a few species, the genus Fidicina is predom-
inantly associated with tropical dry seasons while Zammara with the wet
seasons. Fidicina also represents the largest genus in Central America in
terms of numbers of determined and partially-determined species, while
Zammara is represented by two, possibly three, species based on current
information. The monotypic genus Quesada is widely distributed in Central
and South America, and also filters into the subtropical zone of North
America, and it is a dry season form. Whereas the two species of Zammara

Fig. 9. Collections of cicada nymphal skins over a two-year period from the three localities
(lower and upper montane tropical moist forest) within the extensive coffee district northwest
of the Meseta Centrale in Costa Rica. Data is for fresh skins only.

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generally replace one another geographically in Costa Rica (A. M. Young,
unpublished), there is considerable sympatry for the species of Fidicina.
Because our knowledge of cicada natural history in the tropics is very mea-
ger, I hesitate to suggest that selection pressures such as competition have
resulted in the evolutionary diversification and ecological differentiation of
the genus Fidicina. Partial, tentative support of this view, however, is seen
in the observed patterns of habitat partitioning in areas where several
species occur together. An illustrative example would be the association of
F. pronoe and F. spinocosta with secondary vegetation and the association
of sympatric species such as F. mannifera, F. amoena, and F. sericans
with primary forests. Such a pattern of ecological separation also occurs for
these cicadas within the premontane tropical wet forest zone of Costa Rica
(Young 1980b).

The proximal cues regulating the timing of annual emergences in tropical
cicadas have not been determined and little is known about the adaptive
significance of allochronic annual adult emergence patterns in these insects.
Because many tropical habitats have assemblages of several species, al-
lochrony may reduce competition for acoustical signal space, chorusing
sites, and other space requirements related to reproduction. Patterns of
habitat separation may be more related to resource requirements of nymphs.

Viewed within the context of ecological separation, examples of appar-
ently microsympatric species of Fidicina such as F. semilata and F. “cof-
fea beneath Inga trees in coffee plantations may have resulted from recent
colonizations of these agricultural habitats when the original forest cover
was cleared. Elsewhere (Young, unpublished manuscript) I have suggested
that there is strong selection pressure favoring the exploitation of canopy-
size Leguminosae trees in tropical forests because (a) various genera and
species of Leguminosae are locally abundant in some forest habitats in dif-
ferent climatic and vegetational zones in Costa Rica (Holdridge et al. 1971)
and elsewhere in the tropics, (b) the nymphal skins of many species of
cicadas in tropical forests are very abundant beneath these trees (e.g.,
Young 1972, 1980a, b, 1981a; A. M. Young, unpublished) and abundances
decline rapidly with increasing distance from these trees, and (c) the xylem
fluids of root crowns of Leguminosae trees in tropical forests may have
higher concentrations of nitrogen-containing nutrients such as amino acids
than roots of other trees in the same habitats as a result of the nitrogen-
fixing bacteria association with many legumes, thereby making these trees
optimal feeding patches for developing cicadas. Cicada nymphs feed on
xylem fluids (Cheung and Marshall 1973; White and Strehl 1978).

In a tropical region such as premontane-to-montane moist forest zone in
Costa Rica northwest of the Meseta Centrale, one of the major coffee-grow-
ing districts of that country, intense removal of the forest cover and replac-
ing it with a mosaic of coffee bushes and legume shade trees may have

VOLUME LXXXIX, NUMBER 2

141

allowed for the colonization of these areas by some species of cicadas al-
ready adapted for legume-feeding or to generalized legume associations in
the original forest. From my observations of Z. smaragdula in this region
and elsewhere in Costa Rica (Young 1980a; A. M. Young, unpublished), I
predict that the apparent restriction of this cicada to Zygia trees along
streams is due to sensitivity to low moisture availability in the soil rather
than due to the kind of tree. Such an effect is particularly noticeable in my
study from the observed distribution of Z. smaragdula nymphal skins near
Zygia trees near water or low topographic points, while those of other
legume-associated cicadas such as F. semilata and F. “ coffea ” are found
with Inga trees on the upper slopes of coffee plantations. I doubt very
much, if the legume-feeding hypothesis is correct, that Zygia root crowns
are terribly different in terms of physical and nutritive properties from those
of Inga, save for the difference in size (age) of the trees.

The observed densities of nymphal skins in coffee plantations for both
Fidicina and Zammara falls well below densities for closely related species
in relatively undisturbed forest habitats within the premontane tropical wet
forest zone in Costa Rica (Young 1980b). The low densities are possibly due
to the combined effects of (a) Inga trees, used as shade, being young or
small relative to legume trees in natural forests, thereby providing less nu-
trients and other resources, (b) lower survival of cicada nymphs in the soil
of coffee plantations and their very disturbed borders, and (c) low density
courtship and oviposition by adult cicadas in the coffee plantation environ-
ment resulting from the even distribution of Inga trees over very large areas
of the hills. Chorusing strategies in tropical cicadas are greatly influenced
by the structure of the vegetation (Young 1980c, 1981b) which in turn may
influence the oviposition behavior. If resources are abundant but rather
uniformly distributed over large areas of habitat, the oviposition effort per
resource patch may be low thereby generating low density distributions of
developing cicadas.

Acknowledgments

This research was funded by National Science Foundation Grant
GB-33060 from the government of the United States of America. I am grate-
ful for this support and also for the assistance with determinations of cicadas
given to me by Dr. Thomas E. Moore (University of Michigan). Dr. Moore
recorded courtship songs and disturbance squawks of several of the cicadas
studied here while accompanying me in the field work during January 1973.
I thank Dr. Dieter C. Wasshausen (National Museum of Natural History,
U.S.A.), and Messrs. Luis Diego Gomez P. and Luis Poveda of the National
Museum of Costa Rica for their assistance with identifications of tree
species. Several undergraduate students from Lawrence University in Ap-

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pleton, Wisconsin assisted with all aspects of the field work. Dr. Ridgway
Satterthwaite, then Director of The Costa Rican Field Studies Program of
The Associated Colleges of the Midwest (U.S.A.), assisted with field logis-
tics. Voucher specimens of the cicadas studied are deposited in the per-
manent collections in the Section of Invertebrate Zoology at The Milwaukee
Public Museum, and in The Museum of Zoology at The University of Mich-
igan. I thank Karen Heerhold for typing the manuscript. This paper is ded-
icated to the kind folks of Grecia, Naranjo, and San Ramon who always
remember fondly that big cicada of “Semana Santa,” Q. gigas.

Literature Cited

Cheung, W. W. K. and A. T. Marshall. 1973. Water and ion regulation in cicadas in relation
to xylem feeding. Journal of Insect Physiology 19:1801-1816.

Holdridge, L. R., W. C. Grenke, W. H. Hatheway, T. Liang and J. A. Tosi, Jr. 1971. Forest
environments in tropical life zones. A pilot study. Oxford and New York: Pergamon,
747 pp.

Janzen, D. H. 1967. Why mountain passes are higher in the tropics. American Naturalist
101:233-249.

White, J. and C. E. Strehl. 1978. Xylem feeding by periodical cicada nymphs on tree roots.
Ecological Entomology 3:323-327.

Young, A. M. 1972. Cicada ecology in a Costa Rican tropical rain forest. Biotropica 4:152-
159.

. 1974. The population biology of Neotropical cicadas. III. Notes on the behavioral

natural history of Pacarina cicadas in some Costa Rican grasslands. Entomological
News 85:239-256.

. 1975. The population biology of Neotropical cicadas. I. Emergences of Procollina

and Carineta in a mountain forest. Biotropica 7:248-258.

. 1976. Notes on the faunistic complexity of cicadas (Homoptera: Cicadidae) in northern

Costa Rica. Revista de Biologia Tropical (Costa Rica) 24:267-279.

. 1980a. Habitat and seasonal relationships of some cicadas (Homoptera: Cicadidae) in

central Costa Rica. American Midland Naturalist 103:155-166.

. 1980b. Environmental partitioning in lowland tropical rain forest cicadas. Journal of

the New York Entomological Society 88:86-101.

. 1980c. Observations on the aggregation of adult cicadas (Homoptera: Cicadidae) in

tropical forests. Canadian Journal of Zoology 58:711-722.

. 1981a. Seasonal emergences of cicadas (Homoptera: Cicadidae) in northwestern Costa

Rica. Milwaukee Public Museum Contributions in Biology and Geology, No. 40, 29 p.

. 1981b. Temporal selection for communicatory optimization: the dawn-dusk chorus as

an adaptation in tropical cicadas. American Naturalist 117:826-829.

Section of Invertebrate Zoology, Milwaukee Public Museum, Milwaukee,
Wisconsin 53233.

Received for publication December 18, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(2), 1981, pp. 143-145

A LITTLE-KNOWN, ANONYMOUS WORK ON AMERICAN AND
EUROPEAN BUTTERFLIES AND MOTHS (1906),

WHICH SHOULD BE ATTRIBUTED TO WILLIAM
BEUTENMULLER (LEPIDOPTERA: NYMPHALIDAE)

Cyril F. dos Passos

Abstract . — This paper calls attention to an anonymous publication on
butterflies and satisfactorily establishes the authors thereof.

It all began several years ago when my two friends, Mr. and Mrs. Allen
L. Freas of Westfield, New Jersey, came to visit us and brought with them
as a present to the author a small book on butterflies that they had found
in their library and thought would be of interest. When it was noticed that
this small book had been published under the auspices of the late William
Beutenmuller, it occurred to the author to write to Miss Nina J. Root,
Librarian of The American Museum of Natural History. She responded that,
while the book was carded, it had disappeared from the shelves. Then com-
menced a search for another copy to replace it. One of the first responses
was from a friend, Mr. William D. Field of the U.S. National Museum, who
kindly loaned to the author his copy. But the greatest piece of luck was
when in a conversation with Mr. Fred Lemmer, then president of the New-
ark Entomological Society, he disclosed that he himself had a copy actually
given to him by Beutenmuller many years ago when Lemmer was a boy.
But let Mr. Lemmer tell his own story:

“Enclosed find the Beutenmuller Booklet.”

“About 47 years ago I used to accompany my Uncle Frederick Lemmer
to the meetings of the Entomological Societies at the New York and
Brooklin [sic] Museums and it was on these occasions that I met Mr.
Beutenmuller and many of the other old time collectors.

“The little Booklet is a souvenir of one of those meetings.

“There is a possibility that Mrs. Beutenmuller made the pictures, as
she was an artist and supposedly made all the pictures for the book of
American Catocalas [s/c].”

This work, entitled A Manual of American and European Butterflies and
Moths Reproduced in Natural Colors with Their Common and Scientific
Names to which is sometimes added, “prepared under the supervision of
William Beutenmuller, Curator of the Department of Entomology, American
Museum of Natural History, New York” was published about 1906 in at
least two countries, the New York edition being by Funk and Wagnall’s

144

NEW YORK ENTOMOLOGICAL SOCIETY

Company of New York and London. It was designed as a volume of the
Standard Nature Series, and copies doubtless exist in many old-time private
libraries.

Mr. Lemmer kindly gave his beautiful copy in original dust cover to the
author, who in turn presented it to the Library of The American Museum
of Natural History to replace the lost copy, where it is now kept in the Rare
Book Room.

Mrs. Edna Hyatt Beutenmiiller was an excellent artist, who assisted her
husband in much of his work, and it is probable that she prepared the
illustrations for the twenty-four plates of this manual, while her husband
prepared the text explaining the figures, and one or both probably prepared
the index. Mrs. Beutenmiiller is described by Weiss (1943, p. 286) as an
“entomological artist,” and Barnes and McDunnough (1918) remarked:

“Ten plates of excellent water colors — drawings of the various species

and two plates containing colored figures of the larvae, all drawn by the

accomplished hand of Mrs. Beutenmiiller.”

This case is similar to that of the Genera of Diurnal Lepidoptera ( 1 846—
52), the title page of which lists the authors as Edward Doubleday and John
O. Westwood followed by “Illustrated with eighty-six plates by William C.
Hewitson. ” The latter phrase did not make Hewitson an author of the work
or scientific names proposed in the work (see Hemming, 1941).

The work under consideration may be collated as follows: 1) dust cover
with title, 2) title pages with different texts, only one seen with a year date
(1860), 3) plates 1-24 with colored figures, 47 being butterflies and 67 moths,
and 4) index, 3 pages with common names and references to plates and
figures. The pages are about \Al/i by 8V2 mm.

No new names are introduced. Plate 13 figures the ova and larvae of the
Chinese silkworm, Bombyx mori, natural size.

Acknowledgments

I am indebted to Mr. and Mrs. Allen L. Freas, Mr. Fred Lemmer, Miss
Nina J. Root, Dr. Frederick H. Rindge, and Mr. John C. Pallister, all three
of The American Museum of Natural History, Mr. William D. Field of the
U.S. National Museum, and to Mr. Joseph Muller for assistance in the
preparation of this paper for which my appreciation and thanks are hereby
tendered.

Literature Cited

Barnes, William and James Halliday McDunmough. 1918. Illustrations of the North American
species of the genus Catocala by Wm. Beutenmuller, with additional plates and text.
Mem. Amer. Mus. Nat. Hist., new ser., 3(1): 1-47, pis. 1-22.

VOLUME LXXXIX, NUMBER 2

145

Hemming, Arthur Francis. 1941 . The dates of publication of the several portions of Doubleday
(E.) Genera of Diurnal Lepidoptera and of the continuation thereof Westwood (J. O.).
Jour. Soc. Bibl. Nat. Hist. 1(1 1):335-41 1.

Weiss, Harry B. 1943. The Journal of the New York Entomological Society 1893-1942. Jour.
N.Y. Ent. Soc. 5 1(4): 285-294.

Washington Corners, Mendham, NJ 07945.
Received for publication September 29, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(2), 1981, pp. 146-147

BOOK REVIEW

Butterflies and Moths of Newfoundland and Labrador: The Macrolepidop-
tera. Ray F. Morris. 1980. Publication 1691, Agriculture Canada. 407 pp.,
34 color plates. Available for $15 (Canada) or $18 (outside Canada) from
Canadian Government Publishing Centre, Supply and Services, Hull,
P.Q. K1A 0S9, Canada.

This book represents an opulently produced documentation of a restrict-
ed, insular, boreal or near-boreal fauna, based on the published records of
over a century and the author’s personal collecting experience of some 25
years plus a large number of private and institutional collections. After a
cursory survey of Lepidopterology in the region, its physical geography,
and the seemingly obligatory material on basic biology, classification, and
collection methodology, the book settles down to individual species ac-
counts of all the butterflies and macro-moths. This is where the principal
interest of the work lies, and this section deserves close scrutiny.

The book will be much sought-after for the color plates of moths, many
of which have never been illustrated before — page after page of obscurely
marked (and often obscure!) Noctuids and Geometrids in faithful color; this
alone is worth the price of the book, and more. Unfortunately the butterfly
plates are as consistently unpleasing to the eye as the moths are pleasing;
a great many of the specimens are of inferior quality, and every inperfection
is emphasized by the photographic technique — the pictures are back-lit.
Several specimens are rubbed, a few are greasy, a number are poorly set,
and the Pierids on plate 2 combine these flaws with poor color reproduction,
the only such plate in the book. These problems will not bother the moth
specialist. But both butterfly and moth collectors may be annoyed that no
data are provided for the individual specimens illustrated, although there
was plenty of room to do so; and the plate legends do not give page refer-
ences to the text, forcing the reader constantly back to the index.

The species descriptions contain considerable new biological information,
particularly on early stages, host plants, and distribution. Again, however,
Morris’ book resembles W. T. M. Forbes’ Macrolepidoptera of New York
in being fine for the moths but generally unsatisfactory for butterflies. For
example, one of the strangest things in the Newfoundland literature is the
assertion by Austin and Leila Clark (inexplicably, and repeatedly, spelled
“Clarke” by Morris) in Butterflies of Virginia (1951) that the black form of
the Tiger Swallowtail occurs there. One would think this book would either
support or refute this claim; instead it merely quotes it (p. 36) — in the pro-
cess listing both P. glaucus glaucus and P. glaucus canadensis as occurring
on the island. This treatment of subspecies, like Austin Clark’s own, betrays
a fundamental misunderstanding of the subspecies concept, and it reappears

VOLUME LXXXIX, NUMBER 2

147

under Nymphalis milberti, where we learn that subspecies milberti and
viola both occur on Newfoundland: “ milberti is more prevalent in the north,
whereas viola predominates in the south.” And further, “the life-history of
the two subspecies is probably very similar” (pp. 54-55).

Those whose interests are limited to diumals will probably not want to
buy this book, while moth collectors, especially in the Northeast and in
eastern Canada, will find much of great value in it. Those who have pub-
lished or are working up data for regional faunas may rightly turn green
with envy over the plates.

Arthur M. Shapiro, Department of Zoology, University of California,
Davis.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(2), 1981, p. 148

ANNOUNCEMENT

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The Smithsonian Foreign Currency Program, a national research grants
program, offers opportunities for support of research in Burma, Guinea,
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Grants in the local currencies of the above listed countries are awarded to
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programs involving host country institutions are welcome. Awards are de-
termined on the basis of competitive scholarly review. The deadline for
submission is November 1 annually. For further information write the For-
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stitution, Washington, D.C. 20560, or call (202) 287-3321.

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The New York Entomological Society

Organized June 29, 1892 — Incorporated February 25, 1893

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Mercy College, Dobbs Ferry, New York 10522
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Journal of the

New York Entomological Society

VOLUME LXXXIX SEPTEMBER 1981

NO. 3

CONTENTS

Dr. Roman Vishniac Karl Maramorosch 150-151

Prepupation behavior and pupation of the predaceous diving beetle Dytiscus verti-
calis Say (Coleoptera: Dytiscidae)

Daniel R. Formanowicz Jr. and Edmund D. Brodie Jr. 152-157

Insect succession in the decomposition of a mammal in Costa Rica

Luis Fernando Jiron and Victor M. Carlin 158-165

Comparison of two sampling methods for Leptothrips mali in Virginia apple orchards

M. P. Parrella, J. P. McCaffrey and R. L. Horsburgh 166-169

The European katydid Meconema thalassinum (DeGeer): new locality records for
North America (Orthoptera: Tettigoniidae) E. Richard Hoebeke 170-171

Synonymy and distribution records in the genus Eucerceris (Hymenoptera: Phil-
anthidae) George R. Ferguson 172-183

Some observations on slope soaring in Pantala flavescens (Odonata: Libellulidae)

David L. Gibo 184-187

A revision of the genus Pephysena with descriptions of two new species (Hemiptera:
Lygaeidae) B. J. Harrington 188-196

Aggregation behavior of Aplomyiopsis xylota (Diptera: Tachinidae)

Frank J. Messina 197-201

Hesperiidae as prey of Stictia Carolina A. Hook 202-203

Notes on the moth Pericopus leucophaea Walker (Lepidoptera: Pericopidae) as a
defoliator of the tree Vernonia patens H. B. K. (Compositae) in northeastern
Costa Rica Allen M. Young 204-213

214-217

Book Reviews

150

NEW YORK ENTOMOLOGICAL SOCIETY

DR. ROMAN VISHNIAC

Honorary Life Member, New York Entomological Society

Roman Vishniac was born in Pavlosk near St. Petersburg, Russia, on
August 19, 1897. He received his M.D. and Ph.D. (biology) degrees in Mos-
cow in 1920. Shortly afterwards, he fled to Berlin, where he conducted
research in endocrinology, working at the same time as a photojournalist.
From 1933 to 1939 he produced a photographic record of Jewish commu-
nities in Central and Western Europe and part of this unique work was
published in 1947 under the title “Polish Jews.” His activities attracted the
attention of the Gestapo and during the Nazi regime he was imprisoned 1 1
times, kept twice in concentration camps, and forced to do hard labor. He
was sentenced to death twice but fortunately managed to escape and emi-
grated to the United States in 1940. In New York he became a research
associate in anatomy at Albert Einstein College of Medicine and, in 1961,
professor in the Department of Biology Education at Yeshiva University.
At the same time he also served as project director and film maker for the
Living Biology film series of the National Science Foundation, applying his
pioneering time-lapse cinematography and light-interruption techniques as

VOLUME LXXXIX, NUMBER 3

151

well as color photomicroscopy of living organisms to produce a classic and
fascinating film series. His chief biological researches, in addition to ento-
mology, have been in protozoology, marine microbiology, physiology of
ciliates and circulation systems in unicellular plants. He originated the hy-
pothesis of the polyphyletic origin of life, according to which the first living
organisms were multicellular structures that emerged many times in many
places by different biochemical pathways. In 1971 a volume of his splendid
color micrographs of proteins, vitamins, and hormones, entitled “Building
Blocks of Life” was published.

Dr. Vishniac is fluent in most European and several Asian languages. He
is an expert in Far Eastern art and philosophy, and a recognized art historian
and science historian as well as a bibliographer. He has one of the largest
collections of science books, including many old and very rare volumes. He
has taught, among others, at City University of New York and Case Western
Reserve University and he is Chevron Professor of Creativity, teaching at
present at the Pratt Institute in Brooklyn and in Manhattan.

Dr. Vishniac served for two years as President of the New York Ento-
mological Society. Among his many honors and awards was the Eastman
Kodak Gold Medal, which he received “in recognition of outstanding con-
tributions which lead to new and unique educational programs, utilizing
motion pictures and instrumentation photography.” At present, Dr. Vish-
niac is working on a new book, entitled “The World that Disappeared.”

Karl Maramorosch

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(3), 1981, pp. 152-157

PREPUPATION BEHAVIOR AND PUPATION OF THE
PREDACEOUS DIVING BEETLE DYTISCUS VERTICALIS
SAY (COLEOPTERA: DYTISCIDAE)

Daniel R. Formanowicz Jr. and Edmund D. Brodie Jr.

Abstract. — Preceding pupation, Dytiscus verticalis larvae burrow into
soil above the waterline. They construct pupal cells using mandibles, legs,
and body wriggling. Pupal cells are nearly spherical and the openings are
sealed from the inside with soil placed there by mandibles. Beetles spent 20
to 34 days in pupal cells. Newly eclosed adults are white but darken in 8 to
14 h. Pupae are described.

Introduction

Information concerning prepupation behavior and pupal stages of North
American Dytiscidae is either lacking or fragmentary. In his review of dy-
tiscid biology, Bertrand (1972) presented descriptions of larvae and pupae
of some species. Matheson (1914), Wilson (1923), and Leech and Chandler
(1956) have presented accounts of dytiscids which form pupal cells by bur-
rowing into mud. James (1969) described eggs, larvae, and pupae of five
species of dytiscids, all of which apparently construct pupal cells in mud
along pond margins. However, actual construction of a pupal cell was only
observed for one species, Acilius semisulcatus Aube (James 1969).

Dytiscus verticalis occur in shallow, weedy, temporary and permanent
ponds in woodlands and open fields. The larva was described by Wilson
(1923) but the pupa is undescribed.

This paper describes pupal cell construction behavior, pupal cells, pupal
stages, length of pupation, and length of time to full coloration for Dytiscus
verticalis reared under laboratory conditions.

Methods

Two hundred and fifty-six D. verticalis larvae were collected from four
ponds on or near the E. N. Huyck Preserve, Rensselaerville, N.Y.; 50 of
these were used in the lab rearing study. Observations from two pupae
found along margins of two of these ponds are also included. Larvae were
held in circular plastic chambers 15 cm in diameter, 6.5 cm in depth, and
contained 4 to 5 cm of water. Each larva was fed several frog tadpoles
( Rana , Hyla ) and/or salamander larvae (Amby stoma) daily.

Beetle larvae which appeared ready to pupate (ceased feeding, became
lethargic, and did not strike at prey) were measured by displacement volume

VOLUME LXXXIX, NUMBER 3

153

(this method was used to minimize the risk of injury during handling, Brodie
et al. 1978) and transferred into water of one of the three types of chambers
described below.

(a) 10 gal. glass aquaria (26 x 51 cm). — These chambers had a maximum
soil depth of 15-20 cm, a soil slope 30 cm long (approx. 30°), and water
added at one end to a depth of 3 to 4 cm.

(b) 20 gal. glass aquaria (30 x 76 cm). — The slope was 60 cm long (ap-
prox. 20°), maximum soil depth was 11 cm, and water depth was 5 cm.

(c) Circular plastic chambers (15 cm diameter). — These chambers con-
tained no standing water but had a level layer of moist soil 4.5 cm deep.

Behavior of each larva was observed continuously for two hours after
introduction into one of the chambers. For descriptive purposes represen-
tative pupae were uncovered and preserved at 7 and 14 days (4 and 1 re-
spectively) after the larval skin was shed. Length of pupation and time from
eclosion to attainment of adult coloration were recorded. Dimensions of
some pupal cells were measured after the adult had left or was removed.
Pupal cells of two field collected pupae were also measured and the pupae
immediately preserved. Newly eclosed and fully colored beetles were sexed
and preserved.

Results

A) Pupal cell construction. — Five categories of behavior were sequen-
tially involved in the construction of a pupal cell after the beetle crawled
onto the soil. 1) The initial hole was dug by moving mud pellets with scoop-
ing motions of the legs and pushing motions of the head (observed in 21
larvae). This hole was usually as wide as the larva and at least one-half its
length deep. 2) Eleven larvae were observed to push their bodies head first
into this initial hole which they widened and deepened by body wriggling.
3) During the widening process described above, 12 beetle larvae were ob-
served to move pellets of mud out of the hole by grasping them with their
mandibles. 4) Mud pellet grasping was used by 12 larvae to form the pupal
cell into a roughly spherical shape. 5) Finally, 12 larvae closed the cell from
the inside by placing pieces of mud in the hole with the mandibles.

Pupal cell construction was completed in 20 min to 12 h. Variability in
amount of time needed to complete a pupal cell may have been the result
of differences in soil texture or moistness within the chambers. The size of
the Dytiscus larvae placed into chambers to pupate varied from 1.4 to 2.0
cc ( x = 1.79 cc).

Mean vertical diameter of pupal cells in the lab was 35 mm and mean
horizontal diameter 36 mm. Pupal cells found in the field were 26 x 35 mm
and 28 x 32 mm. Cells in chambers a and b were located from 220 to 590
mm (x = 338 mm) upslope from the water at soil depths of 64 to 110 mm

154

NEW YORK ENTOMOLOGICAL SOCIETY

Table 1. Description of 5 pupae, 7 (a, b, c, d) and 14 (e) days after shed of the larval skin.

Pupa

a

b

c

d

e

Total length (mm)

31

35

36

37

36

Max. width (mm)

12

15

9

13

16

Setae number

Head

60

47

53

43

62

Pronotum

148

134

144

131

141

Mesonotum

9-M-9

9-M-7

7-M-6

6-M-6

8-M-7

Metanotum

6-M-6

5-M-6

6-M-5

7-M-6

6-M-7

Abdominal segments

1

7-M-7

6-M-8

7-M-7

5-M-6

6-M-6

2

9-M-10

10-M-9

9-M-10

8-M-7

7-M-6

3

10-M-8

10-M-6

9-M-8

6-M-8

8-M-7

4

8-M-9

9-M-6

9-M-9

7-M-8

10-M-9

5

8-M-7

9-M-9

8-M-9

6-M-6

8-M-9

6

8-M-8

9-M-7

8-M-8

6-M-5

9-M-8

7

17-M-18

20-M-14

16-M-13

17-M-14

19-M-12

8

26-M-26

29-M-25

27-M-33

27-M-24

25-M-30

Sex

3

9

9

6

9

( x = 89 mm). The floor of three pupal cells was below the waterline and one
of these three pupae successfully eclosed. There were 44% males and 56%
females among the successfully reared beetles. There were no consistent
differences between the three types of chambers in the parameters that were
measured.

B) Description of pupa. — The following description is based on the com-
bined observations on two male and two female 7-day pupae and one female
14-day pupa (Table 1).

Measurements of lab reared pupae: length — 31 to 37 mm (jc = 34.8 mm);
maximum width — 9 to 16 mm (jc = 12.3 mm). Measurements of field col-
lected pupae: length — 32 to 36 mm; maximum widths — 11 to 14 mm. Caudal
cerci 2 mm long in all pupae.

Head setae arranged as follows (Jc = 50.8): 14 to 18 along continuous
anterior curve between eyes; 1 to 3 at inner posterior corner of each eye;
2 to 3 in each of two groups ventromedial to each eye; and 21 to 32 across
posterior margin of head.

Pronotum with 131 to 148 setae along the lateral, posterolateral, and an-
terior margins and along disc. Setae on mesonotum, metanotum, and eight
abdominal segments variable and unequal on opposite halves of pupae (see
Table 1) (formula used to indicate number of setae on left and right halves
of each pupa is from Spangler (1973), midline is indicated by — M — between

VOLUME LXXXIX, NUMBER 3

155

Fig. 1. Newly eclosed adult Dytiscus verticalis in pupal cell.

the numbers). Ninth abdominal segment terminates in two cylindrical cerci,
each bearing 40 to 50 setae. One pair of spiracles on first six abdominal
segments, located on each anterolateral corner.

Antennae directed ventrally underneath head, between the wing pads and
the femora. Tibiae of first two pair of legs folded against femora with tarsi

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NEW YORK ENTOMOLOGICAL SOCIETY

parallel to body axis. Metafemur and metatibia under hind wing pads and
not folded; metafemora are directed away from the tibia toward the midline.

Dorsal setae associated with thick spine-like projections which are largest
along midline of fifth abdominal segment.

C) Coloration. — Immediately after shedding larval skin (7 days after cell
construction), pupa was white but changed to yellowish- white within a few
hours. Legs, antennae, and mouth parts darken 7 days after eclosion. Pupae
light brown with dark brown appendages just prior to eclosion. Length of
time in pupal cell varied from 20 to 34 days (x = 24.9 days).

Newly emerged adults were white (Fig. 1). Wing covers expanded to
cover the abdomen within five minutes of eclosion and were white with
longitudinal brown streaks. Eyes and tip of abdomen were dark brown.
Adult pigmentation developed dorsally and ventrally from posterior to an-
terior in a V-shaped pattern and took from 8 to 14 h (x = 10.5 h). Beetles
remained in pupal cells throughout this process. Subsequent period of time
spent in pupal cells was not determined. Fully colored adults were trans-
ferred to circular plastic chambers (half filled with water and half with soil).
All newly eclosed adults fed within 24 h of eclosion (0.2 cc Hyla tadpoles).

Discussion

Dytiscus verticalis has a larger pupal cell and a longer pupation period
than either D. fasciventris Say or Acilius semisulcatus which James (1969)
described. This is probably because D. verticalis is larger in all life stages
than either A. semisulcatus or D. fasciventris. Acilius semisulcatus also
exhibits at least one of the categories of pupal cell construction behavior
that we observed in D. verticalis. James (1969) described soil grasping using
the mandibles by A. semisulcatus during pupal cell construction.

There was no apparent difference in pupal cells between lab reared and
field collected D. verticalis pupae. However, eclosion success of beetle
larvae which had constructed cells in the lab (50%) may have been different
than under field conditions. Other factors such as drying and predation (one
field collected pupae found by us had been killed by ants) may lower eclo-
sion success in natural situations.

Acknowledgments

This study was conducted while the authors were supported by NSF-
DEB78-11196 and the E. N. Huyck Preserve; we thank R. C. Dagleish,
Executive Director, for courtesies extended and help in the research. Stu-
dent members of Earth Watch helped collect and observe beetles. We thank
P. J. Spangler of the Smithsonian Institution for his identification of the
beetles. We also thank M. S. Bobka and J. C. Bodenwiser for their com-
ments on the manuscript.

VOLUME LXXXIX, NUMBER 3

157

Literature Cited

Bertrand, H. 1972. Larves et Nymphes des Coleopteres Aquatiques du Globe. Imprimerie F.
Paillart, Paris. 799 pp.

Brodie, E. D., Jr., D. R. Formanowicz Jr. and E. D. Brodie III. 1978. The development of
noxiousness of Bufo americanus tadpoles to aquatic insect predators. Herpetologica
34:302-304.

James, H. G. 1969. Immature stages of five diving beetles (Coleoptera: Dytiscidae), notes on
their habits and life history, and a key to aquatic beetles of vernal woodland ponds in
southern Ontario. Proc. Entomol. Soc. Ont. 100:52-97.

Leech, H. B. and H. A. Chandler. 1956. Aquatic Coleoptera. In: Aquatic insects of California,
R. L. Usinger (ed.). University of California Press, Berkeley, Calif.

Mathieson, R. 1914. Life history of a dytiscid beetle ( Hydroporus septentrionalis ). Canad.
Entomol. 46:37-40.

Spangler, P. J. 1973. The bionomics, immature stages, and distribution of the rare predaceous
water beetle Hoperius planatus (Coleoptera: Dytiscidae). Proc. Biol. Soc. Wash.
86:423-434.

Wilson, C. B. 1923. Water beetles in relation to pond fish culture with life histories of those
found in Fairport, Iowa. Bull. U.S. Bureau of Fisheries 39:232-245.

(DRF) Biology Department, State University of New York at Albany,
Albany, New York 12222 and (EDB) Biology Department, Adelphi Univer-
sity, Garden City, New York 11530.

Received for publication September 4, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(3), 1981, pp. 158-165

INSECT SUCCESSION IN THE DECOMPOSITION OF
A MAMMAL IN COSTA RICA1

Luis Fernando Jiron2 and Victor M. Cartin2

Abstract. — Observations were made on the decomposition of a dead dog
during the dry season of 1977 in the Central Valley of Costa Rica, Central
America. The terrain is classified as premontane humid forest and the ob-
servations were made in a secondary forest. The general pattern of decom-
position was basically the same as has been described by authors in other
latitudes, but different in the ecological complexity and the insect fauna
involved. The classification used for human cadavers by forensic patholo-
gists in Costa Rica and other countries in the American tropics was utilized
in this study. It includes the following stages: discoloration, emphysematic
(bloated), liquefaction and skeletal remains. The succession of different
species appeared to depend on their specific feeding preferences, interspe-
cific competition, and the microclimate provided by the substratum. Marked
changes in the activity of populations during crepuscular periods coincided
with an increase in relative humidity and a decline in temperature in the ma-
croenvironment of the surrounding forest. Included among the principal
insect consumers of the remains were the calliphorid dipterans Phaenicia
eximia Wiedemann and Hemilucilla segment aria Fabricius, the piophilid
dipteran Prochyliza azteca McAlpine and the coleopteran Dermestes car-
nivorus Fabricius. The most important predators were the histerids Eu-
spilotus aenicollis Marshall, Hister punctiger Paykal and Geomysaprinus
(Priscosaprinus) belioculus Marshall. Some of these species have also been
associated with a similar type of substratum in the tropical rain forest and
tropical dry forest in Costa Rica.

Introduction

Much is known about the behavior and taxonomy of the principal groups
of insects that participate in the successional stages of decomposition in
temperate zones, but relatively few studies have been made in tropical zones
(Bohart and Gressit 1951; Cornaby 1974; Payne 1965). Previous studies have
shown that certain general patterns exist during the decomposition process
that accounts for the natural or slightly altered ecological conditions. The
decomposition phenomenon, although it presents a continuum of changes,
has been divided into various phases or stages in order to facilitate its study

1 This publication was supported by funds provided by Consejo Nacional de Investigaciones
Cientificas y Tecnologicas (CONICIT) de Costa Rica.

2 Temporary address: Department of Entomology, University of Wisconsin, Madison, Wis-
consin 53706, U.S.A.

VOLUME LXXXIX, NUMBER 3

159

(Johnson 1975; Payne 1965; Reed 1958). Given the limited knowledge of this
process in the neotropical region, the objective of our study was to interpret
and characterize the decomposition process in a dead dog, and to determine
the principal insect species that were present.

Materials and Methods

Field observations were made during the dry season of 1977 in a second-
ary forest on the campus of the University of Costa Rica, in San Jose, Costa
Rica, Central America. The altitude of the study area is 1,200 m. The sub-
strate utilized was a dead female dog that had been euthanized with nem-
butal (Pentobarbital). The carcass was placed on the floor of the forest and
observations were made each day for 28 days, every four hours from 0600
hours to 1800 hours with an additional observation at 2000 hours. From the
28th through 48th day, observations were made every 4th day, and subse-
quent observations were made on days 58, 70, 80 and 90. At each observa-
tion period, recordings on temperature, relative humidity, state of decom-
position of the dog, dominant insect population and predation were made.
Samples of adult and larval insects were collected for later identification.
Larvae were reared to the adult stage for identification. Particular attention
was given to the collection of the most abundant species. Rarer species
were grouped according to the similarity of their ecological function and
identified only to family level.

Results and Discussion

Our observations indicated that decomposition in the tropics has the same
general pattern as described for temperate zones. The differences are prin-
cipally in the number and duration of the phases, the insect faunae involved
and the degree of ecological complexity (Johnson 1975; Payne 1965; Reed
1958). In our study we utilized the classification used by forensic patholo-
gists in Costa Rica and other Latin American countries for human cadavers
(Lopez and Gisbert 1962; Vargas 1977). This classification divides the de-
composition process into four phases as follows:

Discoloration. — Color changes begin immediately following death and in-
clude pallid color of the lips, nose, feet and abdominal region and green
color of superficial veins. Hobson (1932) found that during this initial period
the tissues are acid and not suitable for feeding by fly larvae, which feed on
the liquid between muscle fibers. Later, the tissues become alkaline, the
sarcolemma dissolves and the intermuscular tissue is attacked by the larvae.
In our study this phase lasted approximately three days (Fig. 1). During this
period we observed the presence of ants ( Camponotus sp.), muscoid, sar-
cophagid and drosophilid flies and a large number of the calliphorid, Phoen-
icia eximia Wiedemann, which oviposited in the throat region near the nasal
fosae, and in the eyes, mouth, vagina and anus.

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NEW YORK ENTOMOLOGICAL SOCIETY

CAMPONOTUS SPP.

PHAENICIA EXIMIA

HEMILUCILIA
SEGMENT ARIA

COCHLIOMYIA
M ACELL ARIA

PROCH YLIZA
AZTEC A?

OTHER MUSCOIDS

GEOTRUPINES

EUSPILOTUS

AENICOLLIS

HISTER PUNCTIGER

GEOMYSAPRINUS

BELIOCULUS

TRIGONA SPP.

DERMESTES

CARNIVORUS

DAYS

Fig. 1. Appearance of insect populations thru the necrologic stages of decomposition of a
dead dog in the Central Valley of Costa Rica, during dry season of 1977. ■ High population
density, | | | | low population density.

Emphysematic state. — This state of degradation is characterized by
marked tumefaction of the abdomen and other body regions and protrusion
of the eyes and tongue. This condition is caused by anaerobic decomposition
of the proteins and is subject to individual variation (Johnson 1975; Payne
1965; Reed 1958). Duration of this period is variable and probably depends
on the temperature and other environmental conditions.

In this study, this phase lasted approximately four days. During this time
the main insects observed were large foci of active larvae in the anal and
mandibular regions. The majority of the specimens collected from these two
regions were P. eximia. At night, adults of the histerid beetle Euspilotus
aenicollis Marshall, and certain geotrupine scarabs comprised the majority
of the active faunae. This group of Coleoptera hid during daylight hours in
holes dug into the soil near or beneath the body of the animal. Similar
behavior was also observed by Payne (1965) in the temperate zone. During
the final part of this phase, bare areas developed near the soil interphase
due to hair slippage. Several muscoid flies were observed ovipositing on
these fleshy exposed regions.

Liquefactive stage. — This stage is characterized by the liquefaction of the

VOLUME LXXXIX, NUMBER 3

161

viscera and soft tissues. It begins with breakage of the skin in one or more
places, permitting entry of air and leading to aerobic decomposition (Reed
1958). Johnson (1975) interpreted this phase as a general deterioration of the
tissues until only dried tissues and bones remained, with a loss of 90 percent
of the original weight of the carcass. This degenerative phase corresponded to
what Payne (1965) divided into two steps: active decay stage and advanced
decay stage.

In the present study, this phase began on the 8th day and lasted approx-
imately 19 days. During this phase we observed the greatest activity by
necrophilic insects, both in the size of population present and the variety
of species.

In the initial part of this phase we observed that the remains of the animal
underwent rapid deterioration. Muscoid fly larvae of various species, in
various stages of development, constituted the most conspicuous group of
insects, but the majority of the pupae collected were of P. eximia, the
pioneer species. It is important to note that this calliphorid completed two
or perhaps three generations during degradation of the carcass, and,
together with Hemilucilia segmentaria Fabricius and the piophilid Prochy-
liza azteca McAlpine were the most abundant species during this phase of
decomposition. Adult insects, including silphids and other Coleoptera were
observed but not identified to species level.

The abundance of fly eggs and larvae attracted numerous predators. These
were most abundant at night and included staphylinids, and especially his-
terids ( E . aenicollis, Hister punctiger Paykal, and Geomysaprinus ( Pris -
cosaprinus) belioculus Marshall). Other species occasionally observed in-
cluded dermapterans, an ichneumonid, some forest roaches, wasps
(probably of the genus Stelopolybia), bees of the genus Trigona and lepi-
dopterans (Pyralidae and Noctuidae), none of which were identified to
species.

After twelve days there was a marked decline in the population of adult
muscoids visiting the carcass, although larval activity was still very intense.
We observed an increase in the quantity of pupae in the soil around the
remains of the animal. A new focus of larval activity occurred in the thoracic
region, which included larvae of P . eximia, H. segmentaria and Cochlio-
myia macellaria Fabricius. At night, histerids and certain staphylinids were
the most abundant predators.

After twenty days, the remains of the dog were dehydrated and the dy-
namics of the decomposition process began to slow. Activity of the larvae
of the necrophilic species was reduced and they began to migrate to the

3 On the 18th day we observed a recent human fecal deposit 15 meters from our study
material. Insects attracted to the feces included adult sarcophagid, muscid and calliphorid flies
with C. macellaria and P. eximia the most abundant species, although as noted above, they
were absent as adults on the remains of the dog at this time.

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lower and internal areas where moisture content was higher. Emergence of
adult forms, which began on the 18th day became more marked. Histerids,
staphylinids and some dermestids (this latter group had recently appeared)
were active in the same areas in which fly larvae were found.

During the following days the process of dessication became more accen-
tuated and larvae of calliphorids and other groups were restricted to small
more liquid zones. Recently emerged adults left the area. Numbers of P.
azteca and other small dipterans that had been constantly present during
most of the degradation process also diminished considerably. Populations
of predators during the final part of the liquefactive phase were eventually
limited to the histerids, E. aenicollis, H. punctiger and G. belioculus. Visits
by bees of the genus Trigona became most frequent as they collected ma-
terial for construction of their nests (Wille 1965).

Mummification. — The process of decomposition was virtually inter-
rupted in the liquefactive phase. Instead of continuing to a state termed
“skeletal remains’ ’ by forensic pathologists, however, a process of mum-
mification began. This phenomenon occurs frequently in tropical zones un-
der conditions of low humidity, high temperature and appropriate air cir-
culation (Vargas 1977).

During our study period (1-28 February 1977) there was no rain, and
warm air circulated over vegetational cover of the forest, contributing to a
high degree of mummification of the remains. A large proportion of the
muscle tissue dried and became hard, principally those areas in direct con-
tact with the sun’s rays. However, it was possible to observe a few larvae
and pupae of muscoid flies in these lower zones that were in contact with
the soil.

On the thirty-second day we began to observe the presence of adult der-
mestids (which had disappeared many days before) and on the fortieth day
there appeared the first larvae of Dermestes carnivorus Fabricius, which
later increased to high population levels. These larvae constituted the dom-
inant population at least until approximately day ninety. The remains were
consumed by the larvae of this coleopteran through the skeletal phase at the
end of the dry season, when our observations ceased.

Temperature and humidity. — The effects of temperature and humidity on
the behavior of populations of necrophilic insects have been observed by
various authors. Reed (1958) did not note appreciable changes in the activity
of necrophilic insects during periods of rainfall, while Payne (1965) stated
that the activities of insects involved in the putrefactive process were influ-
enced more by temperature than by other environmental factors.

We observed a marked change in the activity of larvae and adults at the
beginning and end of daylight period. During our initial 28 day observation
period, diurnal temperature was relatively constant, ranging from 18.5°C to
20°C between 0600 hours and 2000 hours. During the night, relative humidity
was usually 100% and temperature about 15°C.

VOLUME LXXXIX, NUMBER 3

163

phalanglds

ataphlllnlda

Fig. 2. Trophic relationships observed among arthropods associated with the carcass of a
mammal in a premontane humid forest. Trophic levels: A. animal under process of degradation,
B. Primary consumers, and C. Predatory insects.

During the dry season temperature and humidity are relatively stable dur-
ing daylight hours. However, atmospheric conditions are less stable during
the rainy season in the Central Valley of Costa Rica. During the months of
October and November of 1974 (rainy season) one of us (L. F. J.), working
in a preliminary study in the same forest, observed a clear reduction in the
activity of adult and larval muscoid flies associated with a dead armadillo
( Dasypus novemcinctus) on cold cloudy days.

Feeding relationships. — Trophic levels associated with the decomposition
of the vertebrate comprised three categories (Fig. 2). The first level was
comprised of the material undergoing degradation by bacteria and fungi.
The second level was comprised of primary consumers including larval and
adult coleopterans, principally of the families Silphidae, Scarabaeidae and
Dermestidae and dipterans of the families Calliphoridae, Drosophilidae,
Chloropidae, Otitidae, Piophilidae, Phoridae and Sarcophagidae. Other pri-
mary consumers included certain Hymenoptera, Lepidoptera and roaches
that occasionally fed or extracted material from the substrate. The third
level was comprised of secondary consumers that were predators of the
primary consumers. These predators were principally histerids, staphylinids,
vespids, ants, phalangids, chilopods and reduviids. In the primary study
mentioned above, during the rainy season we observed a greater variety of

164

NEW YORK ENTOMOLOGICAL SOCIETY

secondary consumers which included practically the same groups as seen
during the dry season plus other arthropods: carabids, spiders, abundant
mites Macrocheles muscadomesticae (Scopoli) and a braconid mycrohy-
menopteran which was a parasite of fly pupae.

Appreciable differences in the time of appearance of the different necro-
philic species were observed during the process of decomposition (Fig. 1).
The presence and abundance of a given species appeared to depend largely
on its microclimatic and feeding tolerances and preferences. There was not a
strict relationship between a given decompositional stage and the presence
of any particular species. For example, the piophilid P. azteca was present
for many days, independent of the stage of decomposition, probably because
this insect feeds on secretions and exudations of the carcass, and its envi-
ronmental and feeding preferences are apparently wide. The presence and
abundance of larval and adult stages of a given insect varied. For example,
the calliphorid P. eximia was present for a long period of time, although the
adults were not attracted to the carcass after the second half of the lique-
factive phase. However, they and other muscoid flies were present in the
area as was evidenced by attraction to fresh human feces in the vicinity.

Temperate zone studies have shown seasonal variation in the occurrence
of muscoid species in carcass substrates (Johnson 1975; Reed 1958). Denno
and Cothran (1975) indicated that in addition to seasonal variations, the size
of the carcass and intraspecific competition among the necrophilic insects
influence the presence of fly species. We found that some calliphorid species
occur throughout the year in the tropics, although there may be variations
in population densities related to season of the year and the types of sub-
strates available (Cornaby 1974; Jiron 1979).

The presence of Dermestes carnivorus also appears to depend on the
biochemical condition of the carcass and on the microclimate. Favorable
conditions for development of large dermestid populations were present
after the cessation of the liquefactive phase, when large portions of the
carcass began to dry. The absence of rain also appears to favor the pres-
ence of this coleopteran, which was present for several weeks in the dried
carcass. This requirement for dry conditions may explain the fact that der-
mestids appear sporadically during the liquefactive phase and later disap-
pear, before reappearing in the mummification stage.

The presence of the predators H. punctiger, E. aenicollis, G. belioculus
and others depends mainly upon the abundance of fly larvae, microclimatic
conditions, and probably interspecific competition. State of decomposition
in the carcass did not affect these predatory insects directly. It is important
to note that several of the principal species observed in our study in the
Central Valley of Costa Rica (premontane humid forest) were observed also
by Cornaby (1974) in tropical humid and tropical dry forest of the same
country. Also the calliphorids P. eximia, H. segmentaria and C. macellaria

VOLUME LXXXIX, NUMBER 3

165

have been found associated with human cadavers in the liquefactive stage
in several regions of Costa Rica (Jiron 1979).

Acknowledgments

The collaboration of several individuals is gratefully acknowledged. Drs.
John M. Kingsolver and Raymond J. Gagne (Systematic Entomology Lab-
oratory, USDA, Washington, D.C.); Dr. Rupert L. Wenzel (Field Museum
Natural History, Chicago, 111.); Drs. K. G. V. Smith and B. G. Cogan (Brit-
ish Museum Natural History) and Ing. Luis A. Salas (University of Costa
Rica) helped with identification of the material. Dr. Luis G. Vargas pre-
pared some of the specimens in the laboratory. Prof. Luis D. Gomez (Na-
tional Museum of Costa Rica); Drs. Rodrigo Zeledon and Eduardo Vargas
(University of Costa Rica) and Drs. Gene R. DeFoliart, Thomas M. Yuill
and Robert L. Jeanne (University of Wisconsin, Madison) critically re-
viewed the manuscript.

Literature Cited

Bohart, G. E. and J. L. Gressitt. 1951. Filth-inhabiting flies of Guam. Bernice P. Bishop Mus.,
Honolulu Bull. 104:1-152.

Cornaby, B. W. 1974. Carrion reduction by animals in contrasting tropical habitats. Biotropica
6:51-63.

Denno, R. F. and W. R. Cothran. 1975. Niche relationships of a guild of necrophagous flies.
Ann. Entom. Soc. Amer. 68:741-754.

Hobson, R. P. 1932. Studies on the nutrition of blow-fly larvae. III. The liquefaction of muscle.
J. Exp. Biol. 9:359-365.

Jiron, L. F. 1979. Sobre moscas califoridas de Costa Rica. Brenesia 16:221-222.

Johnson, M. D. 1975. Seasonal and microserai variations in the insect populations on carrion.
Amer. Midi. Natur. 93:79-90.

Lopez, L. and J. A. Gisbert. 1962. Tratado de medicina legal. Edit. Saber, Valencia. 827 pp.
Payne, J. A. 1965. A summer carrion study of a baby pig Sus scrofa Linnaeus. Ecology
46:592-602.

Reed, H. B. A. 1958. Study of dog carcass communities in Tennessee with special reference
to the insects. Amer. Biol. Natur. 59:213-295.

Wille, A. 1965. Las abejas atarra de la region mesoamericana del genero y subgenero Trigona.
Rev. Biol. Trop. 13:271-291.

Vargas, E. 1977. Medicina legal. Edit. Univer. Costa Rica, San Jose, Costa Rica. 386 pp.

(LFJ) Departamento de Paras itologia, Universidad de Costa Rica, San
Jose, Costa Rica, C.A. and (VMC) Escuela de Ciencias Agrarias, Univer-
sidad Nacional, Heredia, Costa Rica, C.A.

Received for publication September 2, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(3), 1981, pp. 166-169

COMPARISON OF TWO SAMPLING METHODS FOR
LEPTOTHRIPS MALI 1 IN VIRGINIA
APPLE ORCHARDS

M. P. Parrella,2 J. P. McCaffrey and R. L. Horsburgh

Abstract. — A destructive and time-consuming limb-tap sampling method
was demonstrated to be 88% efficient for capturing adult and 2nd larval
stage Leptothrips mali. This sampling method was compared to a simple,
non-destructive, but inefficient, visual sampling technique through regres-
sion analysis. A significant relationship was found which indicated that the
visual technique may be a practical sampling method.

Predaceous thrips have been sampled from deciduous fruit trees by tap-
ping limbs over a cloth covered tray (Putnam 1965; Parent 1967; Horsburgh
and Asquith 1968; Holds worth 1968) and by visual examination of the foliage
for timed periods (Clancy and Pollard 1952). Problems associated with the
limb-tap method are the amount of time required, damaged limbs and dis-
lodged fruit. Visual searching of the foliage for Leptothrips mali (Fitch) is
practical because the adults are black and 2nd larval stages are reddish-
brown which allows these stages to be observed against the green foliage.
Unfortunately, this technique greatly underestimates the thrips’ population
on the tree because these predators are not always on the leaves where they
can be observed.

To circumvent these problems, the efficiency of the limb-tap technique
for capturing thrips was evaluated and regression analysis was employed to
determine the relationship between thrips’ numbers obtained utilizing a vi-
sual search and the limb-tap technique.

Materials and Methods

The efficiency of the limb-tap technique for sampling adult and 2nd larval
stage L. mali was evaluated in 1977. An abandoned orchard which had not
been sprayed for ca. 10 years was used as the study area. Several cultivars
were present in the 3.2 ha orchard but Golden Delicious was the only cul-
tivar sampled; trees were 15 years old and ca. 4 m in height. Eight trees
were arbitrarily chosen within the orchard and three limbs (ca. 3.2 m high)
were selected from the northern, western and southeastern sectors of each

1 Thysanoptera: Phlaeothripidae.

2 Present address: Dept, of Entomology, University of California, Riverside, CA 92521.

VOLUME LXXXIX, NUMBER 3

167

Table 1. Efficiency of the limb-tap method for dislodging Leptothrips mali.

Mean numbers of L. mali

**

Tree

no.*

Dislodged

Remaining

Percent capture***

19-V

2- VI

21-VI

19-V

2- VI

21-V

19-V

2- VI

21-V

1

22

32

28

2

7

3

91.6

82.0

90.3

2

25

36

37

2

6

1

92.6

85.7

97.4

3

9

57

71

1

10

2

90.0

85.0

97.3

4

14

31

49

0

7

2

100.0

81.6

96.1

5

3

42

27

0

8

4

100.0

84.0

87.1

6

2

48

15

0

6

3

100.0

88.9

83.3

7

14

32

5

2

3

0

87.5

91.4

100.0

8

8

21

24

0

1

4

100.0

95.5

85.7

* 3 limbs per tree were tapped for thrips and subsequently examined in the lab for remaining
thrips.

** Means from 3 limbs per tree.

*** No. dislodged 4- no. dislodged + no. remaining.

tree. The outer meter of each limb was tapped (five taps per limb) with a
rubber-coated stick over a 1 m2 muslin cloth-covered tray. The tray was
searched for about 15 min. and the number of thrips dislodged per limb was
counted. Immediately after tapping, each limb was covered with a polyeth-
ylene bag, clipped, and taken to the laboratory. The limb and bag were
carefully examined for remaining thrips within 48 h. Sampling was on three
dates; separate trees were sampled only once.

In 1978 this study area was used to determine the relationship between
the numbers of thrips counted while visually searching the foliage and the
numbers obtained when the same foliage was tapped. Thrips were counted
visually on Golden Delicious trees while walking around one-half of a tree
for 3 min. The natural position of any part of the tree was not disturbed
during the inspection. The visually searched area was then tapped with a
rubber-coated stick over a 1 m2 muslin cloth-covered tray. L. mali falling
on the tray were collected and counted. Sampling was on five dates at 2-
week intervals beginning in mid- June. On each sampling date six trees were
arbitrarily chosen in the orchard; separate trees were sampled only once.

Regression analysis was calculated using the General Linear Model pro-
cedure (Barr et al. 1976) with the limb-tap and visual data as independent
and dependent variables, respectively.

Results

The efficiency of the limb-tap technique for capturing adult and 2nd larval
stage L. mali ranged from 82 to 100% (Jc ± S.D. = 88.7 ± 14.6) (Table 1).
The relationship between the mean numbers of L. mali obtained in a visual

168

NEW YORK ENTOMOLOGICAL SOCIETY

Fig. 1. Relationship between the number of L. mali observed and limb-tapped from 6
Golden Delicious trees on 5 sampling dates, 1978 (R2 = 0.76). 95% confidence bands are in-
cluded.

search and limb-tap is presented in Fig. 1. A significant ( P < 0.05) R2 value
of 0.76 (based on 5 sampling dates) was obtained indicating that 76% of the
variation in the numbers of L. mali obtained in the visual search is explained
by their linear relationship with the limb-tapped numbers of thrips.

Many factors can affect this relationship including observer efficiency,
tree height, cultivar, density of L. mali and its prey (mites), environmental
conditions, etc. These would have to be taken into consideration and eval-
uated before the relationship between the limb-tap and visual search can be
employed in a sampling program. The visual search is 6 times faster than
the limb-tap (3 min. vs 15-20 min.) and may be of use in a study of popu-
lation dynamics where an arbitrary rather than an optimum sampling plan
is acceptable (Morris 1960).

Literature Cited

Barr, A. J., J. H. Goodnight, J. P. Sail and J. T. Helwig. 1976. A user’s guide to SAS 76.
SAS Institutes Inc., Raleigh, N.C. 329 pp.

Clancy, D. W. and H. N. Pollard. 1952. The effect of DDT on mite and orchard predator
populations in apple orchards. J. Econ. Entomol. 45:108-114.

Holdsworth, R. P., Jr. 1968. Integrated control: effect on European red mite and its more
important predators. J. Econ. Entomol. 61:1602-1607.

Horsburgh, R. L. and D. Asquith. 1968. Initial survey of arthropod predators of the European
red mite in south-central Pennsylvania. J. Econ. Entomol. 61:1752-1754.

VOLUME LXXXIX, NUMBER 3

169

Morris, R. F. 1960. Sampling insect populations. Ann. Rev. Entomol. 5:243-264.

Parent, B. 1967. Population studies of phytophagous mites and predators on apple in south-
western Quebec. Can. Entomol. 99:771-778.

Putnam, W. L. 1965. The predacious thrips, Haplothrips faurei Hood (Thysanoptera:
Phloeothripidae) in Ontario peach orchards. Can. Entomol. 97:1208-1221.

Department of Entomology, Shenandoah Valley Research Station, Vir-
ginia Polytechnic Institute and State University, Steeles Tavern, Virginia
24476.

Received for publication October 8, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(3), 1981, pp. 170-171

THE EUROPEAN KATYDID MECONEMA THALASSINUM
(DEGEER): NEW LOCALITY RECORDS FOR
NORTH AMERICA (ORTHOPTERA: TETTIGONIIDAE)

E. Richard Hoebeke

Abstract. — Meconema thalassinum (DeGeer) is recorded from Rhode Is-
land and New York. These records represent new United States localities
for this European katydid which was first detected in North America in 1957
on Long Island, New York.

The presence of the European katydid, Meconema thalassinum (De-
Geer), in eastern North America was first reported by Gurney (1960, 1960a).
The first specimens (two males and two females) were collected in July and
August 1957, and again in July 1959 (one male and one female) at Little
Neck, Long Island (Queens County), New York. Johnstone (1970) reported
the second collection of this species with the capture of a single male spec-
imen in August 1968 at King’s Park, Long Island (Suffolk County). A third
collection of five specimens (three males and two females) at Scarsdale and
Larchmont (Westchester County), New York during July and August from
1974-77 was made by Sismondo (1978); he also observed that the nest pro-
visions of Sphex ichneumoneus (Linnaeus) (Hymenoptera: Sphecidae) were
comprised exclusively of Meconema thalassinum. A fourth collection of
thirteen specimens (ten males and three females) at Garden City, Long
Island (Nassau County) in July 1977 was documented by Smith (1979). It is
my purpose in this note to provide two new additional records for M. tha-
lassinum which extend the known distribution of this introduced species in
North America (Fig. 1).

As a result of the USDA- APHIS “High Hazard Pest Survey” program,
a single female specimen of M. thalassinum was recently submitted to me
for identification; this specimen was collected in a nursery at Middletown,
Rhode Island (Newport County) on 14 August 1980 by R. B. LaFrance.

In addition, a thorough search of the undetermined Tettigoniidae in the
Cornell University Insect Collection (Ithaca, New York) uncovered another
new record; one male specimen was collected on the campus of Cornell
University (Comstock Hall) in July 1974 by J. A. Schafrik.

Both specimens are deposited in the Cornell University collection.

VOLUME LXXXIX, NUMBER 3

171

Fig. 1. Known distribution of Meconema thalassinum (DeGeer) in eastern North America
(male habitus drawing adapted from Smith, 1979).

Literature Cited

Gurney, A. B. 1960. Meconema thalassinum, a European katydid new to the United States.
Proc. Ent. Soc. Wash. 62:95-96.

. 1960a. Meconema taken in the United States in 1957. (Orthoptera: Tettigoniidae).

Proc. Ent. Soc. Wash. 62:279.

Johnstone, D. E. 1970. Notes on the Palearctic grasshopper, Meconema thalassinum (De-
Geer), (Orthoptera: Tettigoniidae: Meconematinae) established in Long Island, New
York. Ent. News 81:62-65.

Sismondo, E. 1978. Meconema thalassinum (Orthoptera: Tettigoniidae), prey of Sphex ich-
neumoneus (Hymenoptera: Sphecidae) in Westchester County, New York. Ent. News
89:244.

Smith, B., Jr. 1979. European katydid Meconema thalassinum (DeGeer) recorded from new
location on Long Island, New York (Orthoptera: Tettigoniidae). Jour. N.Y. Ent. Soc.
87(1): 38-41.

Department of Entomology, Cornell University, Ithaca, New York 14853.
Received for publication November 14, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(3), 1981, pp. 172-183

SYNONYMY AND DISTRIBUTION RECORDS IN THE GENUS
EUCERCERIS (HYMENOPTERA: PHILANTHIDAE)1

George R. Ferguson

Abstract. — Eucerceris ferruginosa Scullen is removed from synonymy
with E. nevadensis (Dalla Torre). E. baja Scullen is the male and a junior
synonym of E. ferruginosa. E. nevadensis mojavensis Scullen (New Status)
is the correct name for the taxon north of Mexico previously referred by
authors to E. nevadensis ferruginosa nec Scullen. The following synonymy
is presented: E. lacunosa Scullen ( =E . lacunosa sabinasae Scullen, New
synonym); E. morula Scullen ( =E . morula albarenae Scullen, New syn-
onym); E. superba Cresson (=E. bicolor Cresson, =Cerceris dichroa Dalla
Torre, New synonyms); E. canaliculata (Say) ( =E . zimapanensis Scullen,
New synonym). The male of E. lapazensis Scullen is characterized, and the
distribution of several species is discussed.

Eucerceris is a North American genus distributed as far south as Panama.
A transverse depression on each of the middle gastral terga separates it from
the closely related and worldwide genus Cerceris. Scullen (1968) revised
the genus Eucerceris, but his key to species and subspecies relied heavily
on variable color characters which makes identification of many specimens
difficult. Bohart and Grissell (1975) revised the California species, but this
helpful work covered only ten of the more than thirty species known. I have
undertaken a review of the genus Eucerceris with the objective of clarifying
synonymy and developing a key to the species based on morphological
characters. This paper is the first in a planned series and presents some of
the results of these studies.

Eucerceris ferruginosa Scullen (New Status)

Eucerceris ferruginosa Scullen, 1939:45. [Female holotype, Angeles Bay,
Baja California Norte, Mexico; Calif. Acad. Sci., San Francisco]; Scullen
1968:30, in part.

Eucerceris baja Scullen, 1948:170. [Male holotype, 20 mi. n. Mesquital,
Baja California Norte, Mexico; Calif. Acad. Sci., San Francisco]; Scullen
1968:19; Bohart and Menke 1976:591. New Synonymy.

I have studied the three specimens comprising the type series of Eucer-
ceris ferruginosa Scullen. The holotype [CAS] and paratype [OSU] from

Oregon Agricultural Experiment Station Technical Paper No. 5717.

VOLUME LXXXIX, NUMBER 3

173

Baja California are not the same species as the paratype labelled Mojave
Desert n. Palmdale, California, June 22, 1931 (F. E. Lutz) [AMNH]. The
latter is the female of Eucerceris mojavensis Scullen which is recognized
below as a subspecies of Eucerceris nevadensis (Dalla Torre). Bohart and
Grissell (1975) synonymized both E. ferruginosa Scullen and E. mojavensis
Scullen with Eucerceris nevadensis (Dalla Torre) ( =Eucerceris elegans
Cresson)2 based on California and Nevada specimens, but the name ferru-
ginosa must be applied to the Baja California species as represented by the
holotype.

Females of E. ferruginosa are essentially all red in body color, but they
are morphologically distinct from E. nevadensis which also has all red fe-
males in a large part of its range. In E. ferruginosa the midsection of the
clypeus is slightly convex, on about the same plane as the lateral clypeal
lobes when viewed in profile, and its apical margin is shallowly biemarginate
between the lateral teeth, forming a short, broad, triangular tooth overlying
the bases of the pre-apical setae. In E. nevadensis the midsection of the
clypeus is flat and depressed below the plane connecting the outer margins
of the lateral clypeal lobes, and the bases of the pre-apical setae are visible,
not being covered by a median tooth.

The holotype and thirteen paratypes of Eucerceris baja Scullen have the
same collection labels as six females of E. ferruginosa Scullen, and five
paratypes of E. baja have the same collection data as the holotype of E.
ferruginosa. In addition Scullen (1968) recorded a female of E. ferruginosa
collected with 12 males of E. baja at 28 mi. s.e. El Arco, Baja California
[UCB], and I have seen a male and female collected at 33 mi. s. La Laguna,
June 30, 1968 [AZS]. These collection records leave no doubt that E. baja
is the male of E. ferruginosa.

This species has been collected at several localities from the vicinity of
San Felipe in Baja California Norte to San Carlos in Baja California Sur.
One female, possibly adventive, was collected 39 mi. n. Puerto Penasco,
Sonora, Mexico [AZS]. Collection dates are from April 1 to September 27
with all intervening months represented except August. This suggests two
generations per year. I have seen 24 males, including the holotype of E.
baja, and 14 females, including the holotype of E. ferruginosa [AZS, CAS,
CSU, OSU, UCB, UID].

2 When Dalla Torre (1890) transferred the species of Eucerceris to Cerceris, Eucerceris
elegans Cresson became a homonym of the earlier Cerceris elegans Eversmann. He renamed
Cresson’s species Cerceris nevadensis Dalla Torre. Many North American hymenopterists
have not recognized the secondary homonymy created by Dalla Torre, and Krombein (1979,
1980) continued to prefer the use of Eucerceris elegans Cresson. Bohart and Grissell (1975, p.
27) state that “Although the international rules of nomenclature on this point were changed in
1961, they were not retroactive and the names of . . . Dalla Torre must be used,” and I prefer
to follow this strict interpretation of the rules as the best long-range policy.

174

NEW YORK ENTOMOLOGICAL SOCIETY

Eucerceris nevadensis nevadensis (Dalla Torre)

Eucerceris elegans Cresson, 1879:xxiii. [Male holotype, Nevada; Acad.
Nat. Sci., Philadelphia]; Mickel 1916:413, males only; Scullen 1939:32,
males only; Scullen 1965:132.

Cerceris nevadensis Dalla Torre, 1890:200, new name for Eucerceris ele-
gans Cresson, nec Cerceris elegans Eversmann.

Eucerceris elegans elegans, Scullen 1968:27; Krombein 1979:1739.
Eucerceris elegans monoensis Scullen, 1968:28. [Female holotype, Grant
Lake, Mono County, California; Calif. Acad. Sci., San Francisco]; Krom-
bein 1979:1739.

Eucerceris nevadensis, Bohart and Grissell 1975:31, in part.

Eucerceris nevadensis nevadensis, Bohart and Menke 1976:591.

Eucerceris nevadensis monoensis, Bohart and Menke 1976:591.

Bohart and Grissell (1975) synonymized E. elegans monoensis with the
nominate form and its continued recognition by Bohart and Menke (1976)
and Krombein (1979) is apparently due to time lag between copy preparation
and final printing of the latter two publications.

E. nevadensis nevadensis occurs in Mono and Inyo Counties, California,
and Churchill, Esmeralda, Lyon, Mineral and Washoe Counties, Nevada.
It is apparently a Transition Zone form of the eastern slopes of the Sierra
Nevada Mountains. I have seen 59 males and 14 females. Collection dates
are June 20 to September 27 [AMNH, AZS, CAS, MCZ, OSU, UCB].

Eucerceris nevadensis mojavensis Scullen (New Status)

Eucerceris ferruginosa, Scullen 1939:45, in part, nec holotype; Scullen
1968:30, in part, nec Scullen 1939; Bohart and Grissell 1975:31, nec Scul-
len 1939.

Eucerceris mojavensis Scullen, 1968:44. [Male Holotype, 7 mi. e. Mojave,
California; Univ. of California, Davis]; Bohart and Grissell 1975:31.
Eucerceris nevadensis, Bohart and Grissell 1975:31, in part.

Eucerceris nevadensis ferruginosa, Bohart and Menke 1976:592, nec Scullen
1939.

Eucerceris nevadensis mohavensis (sic), Bohart and Menke 1976:592.
Eucerceris elegans ferruginosa, Krombein 1979:1739, nec Scullen 1939.

The confusion of the females of E. nevadensis mojavensis with the fe-
males of Eucerceris ferruginosa Scullen has been discussed under the latter
species. Bohart and Grissell (1975) did not formally recognize subspecies,
but they did concede that “a reasonable case could be made for E. neva-
densis ferruginosa with mojavensis as a synonym.” Since the name ferru-
ginosa Scullen must be used for quite a different species, the name moja-
vensis Scullen is available for this subspecies.

VOLUME LXXXIX, NUMBER 3

175

E. nevadensis mojavensis occurs in the Upper and Lower Sonoran Zones
of the southwestern states including Inyo, Kern, Los Angeles, San Bernar-
dino and San Diego Counties, California; Mohave and Yuma Counties, Ar-
izona; and Clark, Humboldt, Lincoln and Lyon Counties, Nevada. I have
also seen a specimen labelled Cholla Bay, Sonora, Mexico [WSU]. Dates
of capture are April 26 to June 28, and August 13 to October 20, indicating
the probability of two generations per year. Intergrades occur in Inyo Coun-
ty, California, and Lyon County, Nevada. I have seen 42 males, including
the holotype, and 17 females [AZS, AMNH, CAS, OSU, UCB, UCR, USU,
WSU].

The following key will separate the two subspecies of Eucerceris neva-
densis (Dalla Torre):

1. Males 2

- Females 3

2. Mesosternum largely or entirely black; anteromedial scutal marks

usually small and not fused with the anterolateral marks, sometimes
absent nevadensis nevadensis (Dalla Torre)

- Mesosternum largely or entirely white or light yellow; scutal marks
always present, usually large and fused with anterolateral marks

nevadensis mojavensis Scullen

3. Ground color of thorax and gaster black, or, at least with large black
areas; light colored tergal bands always present

nevadensis nevadensis (Dalla Torre)

- Ground color of thorax and gaster red; tergal bands usually absent,
but if present, background color red nevadensis mojavensis Scullen

Eucerceris lacunosa Scullen

Eucerceris lacunosa Scullen, 1939:19. [Male holotype, Bill Williams Fork,
Arizona; Univ. of Kansas, Lawrence].

Eucerceris arizonensis Scullen, 1939:20. [Female holotype, Oslar, Huachu-
ca Mtns., Arizona; Univ. of Kansas, Lawrence].

Eucerceris lacunosa lacunosa, Scullen 1968:34; Bohart and Menke
1976:591; Krombein 1979:1739.

Eucerceris lacunosa sabinasae Scullen, 1968:36. [Male holotype, 23 mi. n.
Sabinas, Coahuila, Mexico; Univ. of California, Davis]; Bohart and
Menke 1976:591; Krombein 1979:1739. New Synonymy .

The slightly darker specimens on which Scullen (1968) based the subspe-
cies sabinasae occur sporadically over most of the range of the species.
The two color forms are identical morphologically, and therefore no basis
exists for maintaining the name sabinasae.

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NEW YORK ENTOMOLOGICAL SOCIETY

I have seen 41 males, including 2 paratypes of E. sabinasae, and 12
females. All specimens were within the distribution range given by Scullen
(1968). Collection dates ranged from April 23 to September 27 with about
80% of captures in July and August [AMNH, COR, OSU, UAZ, USU].

Eucerceris morula Scullen

Eucerceris morula morula Scullen, 1968:49. [Female holotype, 17 mi. n.e.
San Luis Potosi, San Luis Potosi, Mexico; U.S. Natl. Mus., Washington,
D.C.]; Bohart and Menke 1976:591; Krombein 1979:1739.

Eucerceris morula albarenae Scullen, 1968:46. [Female holotype, 31 mi.
n.e. Las Cruces, Otero County, New Mexico; U.S. Natl. Mus., Wash-
ington, D.C.]; Bohart and Menke 1976:591; Krombein 1979:1739. New
Synonymy .

The two subspecies of E. morula were separated by Scullen (1968) on
rather tenuous and minor color differences. The tendency toward devel-
opment of a red or ferruginous color as partial replacement for a black
background is commonly encountered in the genus Eucerceris. In E. morula
the tendency is much less pronounced than in such well known species as
E. rubripes Cresson. Of the large number of specimens studied, no satis-
factory basis was found for recognizing subspecies.

I have seen 421 male paratypes and 117 female paratypes of E. morula
morula as well as 101 male paratypes and 14 female paratypes of E. morula
albarenae [OSU]. In addition to the distribution given by Scullen (1968), I
have seen a specimen from Lincoln County, New Mexico [UID]. Collection
dates are from June 15 to October 3 with a strong peak in September.

Eucerceris pimarum Rohwer

Eucerceris pimarum Rohwer, 1908:326. [Female holotype, Phoenix, Ari-
zona; U.S. Natl. Mus., Washington, D.C.]; Bohart and Grissell 1975:32;
Bohart and Menke 1976:592; Krombein 1979:1740.

Eucerceris apicata Banks, 1915:404. [Male holotype, Yuma, Arizona; Mus.
Comp. Zool., Cambridge, Massachusetts]; tentative synonymy by Bohart
and Grissell 1975; synonymy by Bohart in Bohart and Menke 1976.
Eucerceris elegans, Scullen 1939:32, nec Cresson 1879, females only.
Eucerceris pimarium (sic), Scullen 1939:32, as synonym of Eucerceris ele-
gans nec Cresson 1879, females only.

Eucerceris pimarum , Scullen 1965:135, females in part; Scullen 1968:52,
females in part.

Eucerceris apicata , Scullen 1965:132, males in part; Scullen 1968:11, males
in part.

Bohart and Grissell (1975) discussed the confusion surrounding this
species and pointed out that the E. apicata and E. pimarum of Scullen

VOLUME LXXXIX, NUMBER 3

177

(1965, 1968) were in fact composed of three species, i.e. Eucerceris bitrun-
cata Scullen ( =Eucerceris triciliata Scullen), Eucerceris conata Scullen
( ^Eucerceris hespera Scullen), and Eucerceris pimarum Rohwer ( =Eucer -
ceris apicata Banks).

Distribution of E. pimarum is limited to the Lower Sonoran deserts of
the southwest, and it is allopatric with both E. bitruncata and E. conata.
Bohart and Grissell (1975) reported E. pimarum from Imperial and Riverside
Counties, California, and Maricopa, Mohave, Pima and Yuma Counties,
Arizona. In addition I have seen specimens from Clark County, Nevada,
[USU], and 15 mi. s. Guaymas, Sonora, Mexico [OSU]. The latter is a male
without red markings as are some males from the Tucson area in Arizona.

The females of E. pimarum were at one time thought to be females of E.
elegans Cresson, and I have seen 15 males and 11 females standing under
the latter name [UAZ]. I have seen a total of 50 males, including the holo-
type of E. apicata Banks, and 40 females of this species. Dates of collection
are from May 29 (Clark County, Nevada) to January 10 (Sonora, Mexico)
with the bulk of collections in September-October [AZS, CAS, COR, MCZ,
OSU, UAZ, UCR, UID, USU].

Eucerceris bitruncata Scullen

Eucerceris bitruncata Scullen, 1939:35. [Female holotype, San Antonio,

Texas; U.S. Natl. Mus., Washington, D.C.]; Bohart and Grissell 1975:32;

Bohart and Menke 1976:591; Krombein 1979:1738.

Eucerceris triciliata Scullen, 1948:172. [Male holotype, 20 mi. n. El Paso,

Texas; Calif. Acad. Sci., San Francisco].

Eucerceris pimarum , Scullen 1965:135, nec Rohwer, females in part, males;

Scullen 1968:52, nec Rohwer, females in part, males.

Scullen (1965, 1968) interpreted his E. pimarum to include both sexes of
E. bitruncata as well as females of E. pimarum Rohwer, and his (1968)
distribution map must be interpreted accordingly. With the exception of the
Utah records, which represent an undescribed species to be described sep-
arately, all males determined by him as E. pimarum belong to E. bitruncata.

Peripheral distribution records from the United States are from Cochise
and Santa Cruz Counties, Arizona; north through Bernalillo and Guadelupe
Counties, New Mexico; Lubbock County, Texas; Cimarron County, Okla-
homa; to Barber County, Kansas. It is known from most of the border
counties from Hidalgo County, New Mexico, to Cameron County, Texas,
and north to Anderson County, Texas. In Mexico it has been collected in
the states of Chihuahua, Coahuila, Durango and Zacatecas.

Dates of capture are from May 30 to September 24 in Arizona and New
Mexico with a peak in August; from April 14 to October 2 in Texas, Okla-
homa and Kansas with a peak in June-July; and from June 24 to October
25 in Mexico with a peak in August. One specimen, a male, from Coahuila,

178

NEW YORK ENTOMOLOGICAL SOCIETY

Mexico, and the single male from Zacatecas, Mexico, are black and white
rather than black, yellow and red. I have seen a total of 548 males and 119
females from all parts of the range of the species [AMNH, AZS, CAS, COR,
CSU, MCZ, OSU, UAZ, UCB, UCR, UID, USU].

Eucerceris conata Scullen

Eucerceris elegans, Mickel 1916:413, nec Cresson 1879, females only.
Eucerceris conata Scullen, 1939:34. [Female holotype, Halsey, Nebraska;
Univ. of Nebraska, Lincoln]; Bohart and Grissell 1975:32; Bohart and
Menke 1976:591; Krombein 1979:1738.

Eucerceris hespera Scullen, 1948:171. [Male holotype, 25 mi. e. El Paso,
Texas; Calif. Acad. Sci., San Francisco].

Eucerceris apicata, Scullen 1965:132, nec Banks 1915, males in part, fe-
males; Scullen 1968:11, nec Banks 1915, males in part, females.

This species has had a tangled history. Mickel (1916) regarded it to be the
female of Eucerceris elegans Cresson known at that time only from males.
Scullen (1939) described E. conata for those females referred to elegans by
Mickel and assigned females to elegans that we now know to be E. pimarum
Rohwer. Scullen (1948) described E. hespera suggesting that it might be the
male of either E. bitruncata or E. conata. Scullen (1965) described the true
female of E. elegans Cresson but incorrectly synonymized E. conata and
E. hespera with E. apicata Banks. This interpretation, followed in Scullen
(1968), brought together both sexes of E. conata and the males of E. pi-
marum under the name E. apicata until clarified by Bohart and Grissell
(1975).

The distribution data for E. apicata given by Scullen (1968) must be
revised since E. pimarum males were included. The specimens recorded
from Pima County, Arizona, are all referable to E. pimarum Rohwer, and
the specimens recorded from Mono County, California, are over-cyanided
males of E. nevadensis nevadensis (Dalla Torre) [CAS]. I have not seen the
specimen from Coconino County, Arizona.

E. conata occurs in the Wilcox area of northern Cochise County, Arizona,
but I have not found it in large quantities of material collected in the Portal
area. It has also been collected in Navajo County, Arizona; Grand and San
Juan Counties, Utah; Pueblo County, Colorado; Goshen County, Wyoming;
Morrill and Thomas Counties, Nebraska; Fall River County, South Dakota;
Bernalillo, Chaves, Dona Ana, Lincoln and Socorro Counties, New Mexico;
and Culberson and El Paso Counties, Texas. Dates of collection are May
25 to October 7. I have seen two specimens from Samalayuca, in northern
Chihuahua, Mexico, collected on June 24 and October 6. I have seen 75
males and 107 females [AMNH, AZS, CAS, CSU, MCZ, OSU, UAZ, UID,
USU].

VOLUME LXXXIX, NUMBER 3

179

Eucerceris superba Cresson

Eucerceris superba Cresson, 1865:108. [Male lectotype, Rocky Mtns., Col-
orado Territory; Acad. Nat. Sci., Philadelphia].

Eucerceris fulviceps Cresson, 1879:xxiii. [Female holotype, New Mexico;
Acad. Nat. Sci., Philadelphia].

Eucerceris bicolor Cresson, 1881:xxxviii. [Female holotype, Montana;

Acad. Nat. Sci., Philadelphia]. New Synonymy.

Cerceris dichroa Dalla Torre, 1890:199. New name for Eucerceris bicolor
Cresson, nec Cerceris bicolor F. Smith. New Synonymy .

Eucerceris fulviceps var. rhodops Viereck and Cockerell, 1904:84. [Female
holotype, Pecos, New Mexico; Acad. Nat. Sci., Philadelphia].
Eucerceris superba var. bicolor, Scullen 1939:37.

Eucerceris superba bicolor, Scullen 1968:65; Krombein 1979:1740.
Eucerceris superba superba, Scullen 1968:66; Bohart and Menke 1976:592;
Krombein 1979:1740.

Eucerceris superba dichroa, Bohart and Menke 1976:592.

Eucerceris bicolor Cresson is a color variant of Eucerceris superba Cres-
son that is known only in females. E. bicolor has the posterior terga black,
whereas in E. superba all terga have broad yellow bands. Scullen (1939)
quoted at length from personal correspondence from Prof. O. A. Stevens
who had collected the two forms together in North Dakota, and he reduced
bicolor to the status of a variety. Later (1968) he treated it as a subspecies.

The frequency of the dark form decreases from north to south and from
east to west as follows (total females with number of dark form females in
parentheses): Alberta 48 (46); Montana 9 (9); North Dakota 17 (16); South
Dakota 7 (7); Wyoming 13 (9); Colorado 4 (2, 1 intermediate); Nebraska 4
(4); Kansas 1 (1); Idaho 10 (0); Utah 28 (0); New Mexico 5 (1, 1 interme-
diate); Arizona 2 (0); Washington 2 (0).

Of interest is the occurrence of this species on Orcas Island (Doebay),
San Juan County, Washington. Two males and two females were collected
by A. R. Gittins, VIII- 14- 1964 [UID, OSU]. The females are of the light
form. The University of Washington collection is now a part of the collec-
tions of the Systematic Entomology Laboratory, Oregon State University,
and it is rich in specimens collected on the San Juan Islands and in the Puget
Sound area during the early decades of this century. E. superba is not rep-
resented in that material. The nearest recorded capture is Lewiston, Idaho,
(Scullen 1968) near the southeastern border of Washington.

I have seen a total of 127 males and 68 females. Dates of collection are
July 2 to September 7 with a pronounced peak in August [COR, OSU, UCB,
UCR, UID, USU, WSU].

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NEW YORK ENTOMOLOGICAL SOCIETY

Eucerceris canaliculata (Say)

Philanthus canaliculatus Say, 1823:80. [Male holotype, Arkansas, de-
stroyed; male neotype, Kansas; Acad. Nat. Sci., Philadelphia].

Cerceris bident ata Say, 1823:80. [Female holotype, Arkansas, destroyed].
Eucerceris canaliculata, Cresson 1865:112; Scullen 1939:47; Bohart and
Menke 1976:591; Krombein 1979:1738.

Eucerceris canaliculata var. atronitida Scullen, 1939:50. [Male holotype,
Beaver Canyon, Utah; U.S. Natl. Mus., Washington, D.C.].

Eucerceris biconica Scullen, 1948:178. [Female holotype, 15 mi. n. El Paso,
Texas; Calif. Acad. Sci., San Francisco].

Eucerceris canaliculata canaliculata, Scullen 1968:23.

Eucerceris canaliculata atronitida, Scullen 1968:25.

Eucerceris zimapanensis Scullen, 1968:72. [Male holotype, 9 mi. n. Ojo
Caliente, Zacatecas, Mexico; Calif. Acad. Sci., San Francisco]; Bohart
and Menke 1976:592. New Synonymy .

Most specimens of Eucerceris canaliculata (Say) have yellow markings
on a red background, but specimens with a black scutum and black markings
on other parts of the body have been collected over much of its range. The
dark form from Utah was described by Scullen (1939) under the name atro-
nitida and was synonymized by Bohart and Grissell (1975). Scullen (1968)
described a similar form as Eucerceris zimapanensis from the states of
Zacatecas and Hidalgo, Mexico, that had slightly more extensive black
markings. The distinction was arbitrary since some specimens from Zima-
pan, Hidalgo, were determined as atronitida. Similar black-marked speci-
mens are common in the states of Durango and San Luis Potosi, Mexico.

With the exception of the Utah population, the frequency of the darker
forms increases from north to south. From Dr. Scullen’ s records I have
tabulated some 2,295 specimens determined by him over the years as follows
(total specimens shown with number of dark specimens in parentheses):
Montana, Wyoming, South Dakota, Nebraska 14 (0); Kansas, Oklahoma,
Arkansas 77 (0); Colorado 51 (2); Utah 29 (12); Texas 786 (11); New Mexico
429 (2); Arizona 532 (7); California, Nevada 53 (0); Chihuahua, Coahuila,
Nueva Leon 156 (0); Durango 22 (16); Zacatecas 5 (2); San Luis Potosi 125
(75); Queretaro 2 (2); Hidalgo 12 (10); Guatemala 1(1).

A specimen labelled Puerto Barrios, Guatemala, VIII- 16- 1965 (Alberto
Ortiz) [OSU] extends the known range southward.

A color form occurs in the southwestern deserts in which the terga are
entirely yellow, the usual transverse dark stripe of the tergal depressions
being absent. I have seen 9 males and 3 females of this form from Imperial,
Riverside and San Bernardino Counties, California; Lincoln County, Ne-
vada; and Yuma County, Arizona.

VOLUME LXXXIX, NUMBER 3

181

I have seen a total of 561 males, including a paratype of E. zimapanensis,
and 184 females. Collection dates are from April to October with a broad
peak in June, July and August.

Eucerceris lapazensis Scullen

Eucerceris lapazensis Scullen, 1968:37. [Female holotype, La Paz, Baja

California Sur, Mexico; Calif. Acad. Sci., San Francisco]; Bohart and

Menke 1976:591.

This species is closely related to Eucerceris canaliculata (Say) and Eu-
cerceris sonorae Scullen, but the females of E. lapazensis are easily sepa-
rated by the absence of the prominent projections on the lateral lobes of the
clypeus characteristic of the other two species. Males of E. lapazensis have
not heretofore been recognized.

I have examined a series of 8 males and 2 females collected 2 mi. s. La
Paz, Baja California Sur, VIII-5/7-1966 that had been previously identified
as E. canaliculata and E. lapazensis respectively [UCB], and a previously
unidentified series of 40 males and 6 females collected at La Paz, Baja
California Sur, IX-5/7-1967 and IX- 17/22- 1967 [UCB]. The females are E.
lapazensis and the males are undoubtedly the same species.

The males of E. lapazensis are very close to E. sonorae and E. canali-
culata in structure and coloration. Most males of E. canaliculata can be
separated by a rounded, somewhat ridgelike, swelling near the dorsolateral
margin of the lateral clypeal lobe that is absent in the other two species.
The subantennal sclerite and immediately adjacent areas of the clypeus and
lower face are rather sparsely punctate in most specimens of E. canaliculata
but more densely punctate in the other two species. In E. lapazensis the
vertex averages somewhat wider with respect to the ocellar triangle than in
the other two species. The least vertex width averaged 2.9, 2.8 and 2.7 times
the ocellar triangle width in E. lapazensis, E. canaliculata , and E. sonorae
respectively with considerable overlap. I have found no single morpholog-
ical character which is reliable in all cases for separating the males of these
three species. As presently known, the three species are allopatric, and
geographic distribution will separate them.

I have seen most of the specimens from Baja California that were deter-
mined as E. canaliculata by Scullen according to his records. These include
the two localities shown in his (1968) distribution map for E. canaliculata,
and I believe all of his determinations from Baja California are properly
referable to E. lapazensis.

In addition to the La Paz area, E. lapazensis has been collected at Todos
Santos, 20 mi. n. Comondu, 20 mi. s. El Arco, and El Centenario in Baja
California Sur; and 14 km s. Rosarito and 20 mi. n. Mezquital in Baja

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NEW YORK ENTOMOLOGICAL SOCIETY

California Norte. I have seen 57 males and 9 females with collection dates
from August 2 to October 3. The bulk of the collection dates are in Septem-
ber [CAS, OSU, UCB, UID].

Acknowledgments

The following individuals and institutions have generously loaned speci-
mens which have been studied in the course of this work [abbreviations in
brackets as used in the text]: M. S. Favreau, American Museum of Natural
History, New York [AMNH]; F. F. Hasbrouck, Arizona State University,
Tempe [AZS]; P. H. Arnaud, California Academy of Sciences, San Fran-
cisco [CAS]; L. L. Pechuman, Cornell University, Ithaca, New York
[COR]; H. E. Evans, Colorado State University, Fort Collins [CSU]; M. K.
Thayer, N. D. Stone, Museum of Comparative Zoology, Cambridge, Mas-
sachusetts [MCZ]; J. D. Lattin, Oregon State University, Corvallis [OSU];
F. G. Werner, University of Arizona, Tucson [UAZ]; J. A. Powell, Uni-
versity of California, Berkeley [UCB]; S. I. Frommer, University of Cali-
fornia, Riverside [UCR]; W. F. Barr, University of Idaho, Moscow [UID];
F. D. Parker, Utah State University, Logan [USU]; R. S. Zack, Washington
State University, Pullman [WSU].

The Systematic Entomology Laboratory, Oregon State University, con-
tains the extensive collection of cercerine wasps built up by Dr. H. A.
Scullen over a period in excess of forty years together with his unpublished
notes and records. I have relied heavily on these resources.

Literature Cited

Banks, N. 1915. New fossorial Hymenoptera. Can. Entomol. 47:400-406.

Bohart, R. M. and E. E. Grissell. 1975. California wasps of the subfamily Philanthinae. Bull.
Calif. Insect. Surv. 19:1-92.

and A. S. Menke. 1976. Sphecid wasps of the world. Univ. of California Press, Berke-
ley, ix + 695 pp.

Cresson, E. T. 1865. Monograph of the Philanthidae of North America. Proc. Entomol. Soc.
Philadelphia 5:85-132.

. 1879. “. . . descriptions of new Hymenoptera.” Trans. Am. Entomol. Soc. (Proc.)

7:xxii-xxiii.

. 1881. “. . . descriptions of new Hymenoptera.” Trans. Am. Entomol. Soc. (Proc.)

9:xxxviii-xxxix.

Dalla Torre, C. G. de. 1890. Hymenoptera Notizen. Wien Entomol. Z. 9:199-204.

Krombein, K. V. 1979. Sphecoidea. Pp. 1573-1740. in Krombein, K. V., Paul D. Hurd, Jr.,
David R. Smith and B. D. Burks, Catalog of Hymenoptera in America north of Mexico.
v.2: 1199-2209. Smithsonian Inst. Press, Washington, D.C.

. 1980. Personal communication.

Mickel, C. E. 1916. New species of Hymenoptera of the superfamily Sphecoidea. Trans. Am.
Entomol. Soc. 42:399-434.

Rohwer, S. A. 1908. New philanthid wasps. Can. Entomol. 40:322-327.

VOLUME LXXXIX, NUMBER 3

183

Say, T. 1823. A description of some new species of hymenopterous insects. Western Quarterly
Reporter 2:71-82.

Scullen, H. A. 1939. A review of the genus Eucerceris. Oregon State Monogr., Stud. Entomol.
1:1-80; Oregon State College, Corvallis.

. 1948. New species in the genus Eucerceris with notes on recorded species and a

revised key to the genus. Pan-Pac. Entomol. 24:155-180.

. 1965. Revised synonymy in the genus Eucerceris with a description of the true female

of E. elegans Cresson. Entomol. News 76:131-136.

. 1968. A revision of the genus Eucerceris Cresson. U.S. Natl. Mus. Bull. 268:1-97.

Viereck, H. L. and T. D. A. Cockerell. 1904. The Philanthidae of New Mexico — I. J. N.Y.
Entomol. Soc. 12:84-88.

Professor of Entomology, Systematic Entomology Laboratory, Depart-
ment of Entomology, Oregon State University, Corvallis, Oregon 97331.

Received for publication December 23, 1980.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(3), 1981, pp. 184-187

SOME OBSERVATIONS ON SLOPE SOARING IN
PANT ALA FLAVESCENS (ODONATA: LIBELLULIDAE)

David L. Gibo

Abstract. — Slope soaring behaviour is described for the dragonfly Pantala
flavescens (Fabricius). Some of the physical and airflow properties of a
soaring site are also described. The ability to slope soar may have a signif-
icant impact on the energy budget of foraging P. flavescens.

Powered flight is a highly energetic activity for insects. Because dragon-
flies perform most of their adult activities in the air (Corbet 1962; Needham
and Westfall 1955), any behaviour that reduces the energy expenditure of
flight would be advantageous. One pattern of behaviour that can result in
a large reduction in energy expenditure is soaring. Soaring occurs when
animals extract energy from the atmosphere by gliding in ascending air
currents or updraughts. Consequently, soaring animals can maintain or gain
altitude without beating their wings.

Rising air currents can be produced by several mechanisms. Updraughts
that are produced by convection are termed thermals (Pennycuick 1975;
Scorer 1978; Welch et al. 1977), while those produced by wind interacting
with an obstacle, such as a hill, are termed slope lift (Pennycuick 1975;
Welch et al. 1977; OSTIV 1978).

Although the dispersal of many insects is known to be enhanced by ther-
mals (Johnson 1969; Rainey 1976; Scorer 1978), the specialized behaviour
patterns necessary for sustained soaring flight have only been reported for
a few species. These species include the monarch butterfly, Danaus plex-
ippus L. (Gibo and Pallett 1979; Schmidt-Koenig 1979), the mourning cloak
butterfly, Nymphalis antiopa L. (Gibo 1981), and the dragonfly Pantala
flavescens (Fabricius) (Hankin 1921). Nymphalis antiopa and P. flavescens
have only been observed to soar in thermals, while migrating D. plexippus
exhibit behaviour patterns that allow them to soar in slope lift as well as
thermals (Gibo and Pallett 1979). This note is a report of slope soaring
behaviour in foraging P. flavescens and also describes the physical features
of a soaring site.

On August 11, 1980 a P. flavescens was observed slope soaring in an area
of lift in the lee of a beach house in Kill Devil Hills, North Carolina. The house
was on pilings and raised approximately 2 m above the ground. Although
air flow under the house was possible, it was impeded both by the pilings
and by a lattice work of slats that ran around the perimeter of the pilings.
As a result, air flow over and around the house would normally have a

VOLUME LXXXIX, NUMBER 3

185

greater velocity than air flow under the house. This flow pattern was ap-
parently producing a form of slope lift in the lee of the house. The exact
mechanism may have been due to turbulence (see Munn 1966), a poorly
understood three dimensional effect on airflow in the lee of an object
(see Scorer 1978), or the Bernoulli principle (see Sears and Zemansky 1970).

The location, dimensions and shape of the area of lift were determined
by releasing small bits of paper at appropriate locations and observing
whether they were carried upward. The area of lift was located directly in
the lee of the house, and was approximately 12 m long, 3 m wide and 2.5
m deep. It was first detectable at approximately 1.5 m above the ground.
Wind velocities within the band of lift, and at appropriate locations near the
house were measured with a wind meter. The wind direction was from the
north. The wind velocity at 1 m was 15 km/hr on either side of the house
(approximately 2 m outside the area of lift) and 15 km/hr 1 m above the roof.
Directly below the area of lift, at 1 m, the wind velocity was approximately
10 km/hr. The wind velocity within the area of lift, 2 m from the house and
2 m above the ground, was approximately 5 km/hr. The horizontal com-
ponent of the wind velocity, in this area, was still north (i.e. there was no
evidence of a counter current). During the observation period the tempera-
ture was 24°C and the sky was overcast with broken altocumulus.

The P. flavescens was observed from 10:25 to 10:55 A.M. (E.S.T.), while
it soared back and forth in the band of lift, generally paralleling the house.
The dragonfly tended to fly at a height of approximately 2 m and remain 2
to 3 m from the house. The P. flavescens usually soared in a straight line
for a distance of approximately 6 m before beating its wings a few times or
turning. The duration of the soaring part of the flight was usually 3 s, but
ranged between 2-5 s (based on 10 observations). The longer glides extend-
ed the entire length of the band of lift. Whenever the P. flavescens arrived
at the boundaries of the area of lift it made an abrupt 180° turn and usually
beat its wings a few times before resuming soaring flight. When the soaring
dragonfly was viewed from the front, it could be seen to rock from side to
side, indicating that the lift was turbulent.

Occasionally, the P. flavescens would suddenly accelerate by beating its
wings, and fly 5-10 m in pursuit of small insects. As these flights were
usually upward, it was twice possible to observe prey capture against the
sky. The dragonfly also employed powered flight to approach other dragon-
flies that flew near the area, although no overt aggressive encounters were
noted. After 30 minutes of observations the dragonfly was collected. Within
10 minutes a second P. flavescens flew into the band of lift and began
soaring back and forth in the same manner as the first specimen. However,
it only remained in the area of lift for approximately 2 minutes.

These observations show that foraging P. flavescens are able to locate

186

NEW YORK ENTOMOLOGICAL SOCIETY

and soar in restricted areas of lift. It is important to note that the potential
foraging area of a slope soaring dragonfly is larger than the narrow area of
lift actually being patrolled. Because air continuously flows through the site,
dispersing insects that are upwind of the site may drift towards the drag-
onfly, extending the effective foraging area.

If the energy metabolism for soaring is similar to the resting metabolism
(soaring dragonflies are essentially holding their wings immobile), then slope
soaring would substantially reduce the cost of patrolling an area. The rela-
tive advantage of slope soaring is indicated by studies of endothermic warm-
up in dragonflies. May (1979) found that during warm-up active dragonfly
species have maximum metabolic rates, that are approximately 19.5 to 29.5
times higher than their resting metabolism. The expected difference in the
two metabolic rates for P. flavescens can be calculated from the data pre-
sented in May (1979). At 30°C the mean resting metabolic rate, from Table
I in May (1979) is 0.0024 W. The mean warm-up metabolic rate, extrapolated
from Figure 3 in May (1979) and corrected by the recommended 15%, is
approximately 0.0680 W. Consequently, for P. flavescens, the metabolic
rate during endothermic warm-up is approximately 28 times the resting met-
abolic rate. If warm-up metabolism approximates flight metabolism, then it
is apparent that episodes of slope soaring could be very important in the
energy budget and ecology of P. flavescens.

Acknowledgments

I thank Miss C. A. Neal for reading the manuscript and for many helpful
suggestions. This research was supported in part by the National Research
Council of Canada Grant A5991.

Literature Cited

Corbet, P. S. 1962. A Biology of Dragonflies. Witherby, London. 247 pp.

Gibo, D. L. 1981. Some observations on soaring flight in the Mourning Cloak butterfly ( Nym -
phalis antiopa L.) in southern Ontario. J. N.Y. Ent. Soc. In press.

and M. J. Pallett. 1979. Soaring flight of monarch butterflies, Danaus plexippus (Lep-

idoptera: Danaidae) during the late summer migration in southern Ontario. Can. J. Zool.
57(7): 1393-1401.

Hankin, E. H. 1921. The soaring flight of dragon-flies. Proc. Camb. Phil. Soc. Bio. Sci. 20:460-
465.

Johnson, C. G. 1969. Migration and Dispersal of Insects by Flight. Methuen and Co. Ltd.,
London. 763 pp.

May, M. L. 1979. Energy metabolism of dragonflies (Odonata: Anisoptera) at rest and during
endothermic warm-up. J. Exp. Biol. 83:79-94.

Munn, R. E. 1966. Descriptive Micrometeorology. Academic Press, New York. 245 pp.
Needham, J. G. and M. J. Westfall, Jr. 1955. A Manual of the Dragonflies of North America
( Anisoptera ). Univ. of Calif. Press, Berkeley and Los Angeles. 615 pp.

VOLUME LXXXIX, NUMBER 3

187

The Organisation Scientifique et Technique International du Vol a Voile — OSTIV — in collabo-
ration with WMO (World Meteorological Organization). 1978. Handbook of Meteoro-
logical Forecasting for Soaring Flight. Secretariat of the World Meteorological Organi-
zation, Geneva, Switzerland. 101 pp.

Pennycuick, C. J. 1975. The mechanics of flight. In: Avian biology. Vol V. Edited by D. S.

Farner, J. R. King and K. C. Parkes, pp. 1-75. Academic Press, New York.

Rainey, R. C. 1976. Flight behaviour and features of the atmospheric environment. In: Insect
Flight. Edited by R. C. Rainey, pp. 75-112. Symp. R. Ent. Soc. Lond. 7. Blackwell
Scientific, London.

Schmidt- Koenig, K. 1979. Directions of migrating monarch butterflies ( Danaus plexippus\
Danaidae; Lepidoptera) in some parts of the eastern United States. Behavioural Pro-
cesses 4:73-78.

Scorer, R. S. 1978. Environmental Aerodynamics. Ellis Horwood Limited, Toronto. 488 pp.
Sears, F. W. and M. W. Zemansky. 1970. University Physics. Addison-Wesley Publishing
Company Inc. 702 pp.

Welch, A., L. Welch and F. Irving. 1977. New Soaring Pilot. John Murray Ltd, London. 412

pp.

Department of Zoology, Erindale College, University of Toronto, Missis-
sauga, Ontario, Canada L5L 1C6.

Received for publication January 5, 1981.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(3), 1981, pp. 188-196

A REVISION OF THE GENUS PEPHYSENA WITH
DESCRIPTIONS OF TWO NEW SPECIES
(HEMIPTERA: LYGAEIDAE)

B. J. Harrington

Abstract. — A diagnosis for the Neotropical genus Pephysena and a key
for identification of five species are presented. Two new species, P. longir-
hynchus and P. microlevis, both from Brazil and Surinam are described.
Dorsal views of both holotypes are provided.

The Neotropical lygaeid genus Pephysena Distant belongs to the tribe
Myodochini in the subfamily Rhyparochrominae. Distant’s (1882) descrip-
tion of the genus included the description of two species, P. levis and P.
insignis. There was no further taxonomic activity concerning this genus
until 1954 when H. G. Barber briefly redescribed Pephysena ; designated P.
levis as the type species; noted that P. insignis did not belong to the genus
Pephysena ; and described two new species, P. picta from Ecuador and P.
fuscosa from Brazil, Trinidad and Venezuela. After examining the type
specimen of P. insignis, Scudder (1962) removed P. insignis to the new
monotypic genus Distingphyses, which Harrington’s (1980) generic level
analysis of the tribe Myodochini recognizes as the sister group of Pephy-
sena.

Pephysena and Distingphyses both have elongate oval eyes, a generally
ant mimetic habitus and a unique characteristic texture of nearly contiguous
pits or punctures on the head. Yet these two genera are readily distinguish-
able. Pephysena can be recognized by the possession of a long, cylindrical,
stalk-like neck; the dome-like convex nature of the head between and im-
mediately behind the eyes; the very long slender antennae (but with the first
segment short and not attaining the apex of the tylus); and a markedly
impunctate band on the anterior one-third to one-half of the posterior pro-
notal lobe. Distingphyses has the head abruptly constricted behind the eyes
but lacks a stalk-like neck; the first antennal segment attains the apex of the
tylus; the head is markedly flattened and depressed between the eyes; and
the entire posterior pronotal lobe is uniformly punctate. In addition, the
lateral corial margin is beaded or crenate for most of its length in Disting-
physes but smooth in Pephysena.

The present paper provides the first key to the species of Pephysena
including two new species described here from Brazil and Surinam. In the
following key and descriptions all measurements are in mm and the Villa-
lobos color chart (Palmer, 1962) has been used as a standard.

Key to Species of Pephysena

1. Wing membrane entirely and uniformly dark (blackish brown);
femora of all legs uniformly dark; ground color of clavus and
corium dark, strongly contrastingly marked with white maculae
as in Figure 1 fuscosa

- Wing membrane, marked mesally or apically with a pale area, or
with one or more pale finger-like markings along veins (Figs. 2,

3 and 4); one or more pair of legs with femora pale on proximal
one-third to one-half, or all femora entirely light orangish buffy
brown or tawny; clavus and corium in part buffy yellow, not
marked exclusively in a contrasting black and white pattern 2

2(1). Meso- and metafemora uniformly light orangish buffy brown or
tawny; wing membrane dark save for a central oval pale spot
which is “enclosed” and does not approach posterior wing margin
(Fig. 3); head not declivent anteriorly, preocular portion porrect,

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NEW YORK ENTOMOLOGICAL SOCIETY

prolonged and snout-like with apex of tylus readily visible from
above (Fig. 3) longirhynchus

- Meso- and metafemora dark distally and contrastingly pale proxi-

mally; wing membrane with pale finger-like markings along veins
(Fig. 4) or with a pale median vitta extending forward from pos-
terior margin (Fig. 2); head declivent anteriorly, apex of tylus
generally not readily apparent in dorsal view (Fig. 4) 3

3(2). Wing membrane dark on basal (anterior) one-half, lighter on apical
(posterior) one-half, with several pale finger-like markings present
along veins (Fig. 4) 4

- Wing membrane without several pale finger-like markings along

veins, largely dark, patterned as in Figure 2 with a single pale
median vitta extending forward from posterior margin picta

4(3). Proximal ends of fore femora dark, concolorous with trochanters;
clavus with a slender elongate pale macula adjacent to anterior
one-half of claval suture and another elongate pale spot paralleling
claval commissure; total length in dorsal view 6. 5-8.0 mm levis

- Proximal ends of fore femora pale, contrasting with darker distal

one-half to two-thirds of femora and with trochanters; clavus uni-
formly dark, dusky brown or chestnut, lacking pale slender mac-
ulae described above (but with broader heavy gray pruinose light-
reflecting patches in the same positions giving a deceptively sim-
ilar light-marked appearance); total length in dorsal view 4. 5-6.0
mm microlevis

Pephysena longirhynchus Harrington, new species
Figure 3

Head, anterior pronotal lobe, scutellum basally and laterally and mem-
brane of hemelytra blackish brown; a medial oval spot in membrane and
ground color of clavus and corium very pale grayish cream; anterior pro-
notal collar, posterior pronotal lobe, tylus, juga, femora, tibiae, epimera,
episterna, medioapical portion of scutellum and transverse bands on corium
at apex and just posterior to claval commissure tawny; tarsi, labium, anten-
nal segment IV, distal portion of segment III and extreme distal end of
segment II fuscous; antennal segment I and major portions of segments II
and III as well as indistinctly marked areas on clavus and corium between
buffy yellow and pale tawny; abdomen light chestnut.

Antennae, legs, labium and abdomen smooth, impunctate and moderately
shining; head shining with numerous nearly contiguous micropunctures, this
surface reflecting the light in a characteristic broken pattern; pronotum,
scutellum and hemelytra including membrane dull, clothed with a pale gray
or whitish pruinosity which is densest or most apparent in a broad band
across anterior one-third of posterior pronotal lobe; anterior one-third of

VOLUME LXXXIX, NUMBER 3

Fig. 3. Pephysena longirhynchus Harrington. Holotype, dorsal view.

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NEW YORK ENTOMOLOGICAL SOCIETY

Fig. 4. Pephysena microlevis Harrington. Holotype, dorsal view.

VOLUME LXXXIX, NUMBER 3

193

posterior pronotal lobe impuctate; anterior pronotal lobe only sparsely punc-
tate with small shallow indistinct punctures; lines or grooves demarking
anterior pronotal collar and transverse impression with larger deeper more
closely spaced and distinct punctures; punctures equally large or larger on
scutellum, clavus, corium and a band across posterior two-thirds of poste-
rior pronotal lobe, excluding wide impunctate lateral areas at the humeral
angles (punctures large dark and extremely apparent against light back-
grounds of clavus, corium and posterior pronotal lobe); much of body sur-
face, particularly legs and antennae, clothed uniformly with fine short re-
cumbent hairs; fewer long distinctive erect hairs present over entire surface
of head, fore femora and scutellum and on dorsal surfaces of both pronotal
lobes.

Head little declivent anteriorly with pre-ocular portion prolonged and
snout-like; tylus readily apparent in dorsal view; post-ocular region of head
elongate, constricted posteriorly to form a stalk-like neck, post-ocular por-
tion of head immediately behind eyes convex and dome-like; eyes sessile,
longitudinally oval; length head 1.34; width 0.80; interocular distance 0.46.
Anterior pronotal lobe globose, with a distinct anterior collar demarked by
a line-like groove; posterior margin of pronotum straight across base of
scutellum, humeral angles squarely truncate; length anterior pronotal lobe
0.68; width 0.74; length posterior pronotal lobe 0.40; width across humeral
angles 1.02; width transverse impression 0.64. Length scutellum 0.54; width
0.48. Length corium 1.76; distance apex corium to apex membrane 0.72;
length claval commissure 0.32; distance apex clavus to apex corium 0.78.
Labium extending onto abdominal sternum II (first visible); length labial
segments I 0.72, II 0.92, III 0.94, IV 0.38; bucculae quite short but expanded
and leaf-like, projecting forward on either side of labial base. Antennae
slender; length antennal segments I 0.26, II 0.78, III 0.68, IV 0.88. Total
length 4.94.

Holotype: Surinam: S P. H. v. Doesburg Jr. In Leiden Museum No.
1054.

Paratypes: Surinam: IS Same data as holotype. Brazil: 1 (abdomen miss-
ing) Mato Grosso 12°15'S, 51°47'W. Campo 16-X-1968, O. W. Richards. In
Leiden Museum No. 1056 and J. A. Slater collection.

In the paratype from Surinam the anterior pronotal lobe is lighter than
that of the holotype and closer to chestnut in color, hence, contrasting less
markedly with the posterior pronotal lobe. In the Brazilian paratype the
head is slightly less elongate and a little broader.

P. longirhynchus is quite different from other described species of Pe-
physena. Specimens can be readily identified by the distinctive, pale, me-
dial, oval spot in an otherwise dark wing membrane and the characteristic
porrect, prolonged, snout-like preocular portion of the head. In addition,
the punctures on the posterior pronotal lobe, scutellum and clavus of P.

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NEW YORK ENTOMOLOGICAL SOCIETY

longirhynchus are larger and deeper than those of other species and are very
apparent against a light-colored background.

Pephysena microlevis Harrington, new species
Figure 4

Head blackish brown; anterior pronotal lobe and scutellum dusky brown;
anterior one-half of hemelytral membrane fuscous; posterior one-half of
membrane smoke gray with distinctive pale gray finger-like markings along
veins; posterior pronotal lobe, clavus, broad transverse bands on corium at
apex and at level of posterior end of claval commissure and a fine line along
entire membranal margin of corium chestnut; coxae, distal one-half to two-
thirds of all femora, proximal and distal portions of all tibiae, antennal seg-
ment IV, distal one-fourth to one-third of segments II and III, proximal one-
half of segment I and abdomen between light chestnut and tawny; other
portions of antennal segments and a sinuate vitta along medial fracture in
corium buffy yellow; corium narrowly marked with sordid white along lat-
eral margin to level of posterior end of claval commissure, a broad subapical
corial macula and three small spots at level of claval commissure (two along
claval suture and a third laterad of median fracture) also sordid white; labial
segments I, III and IV sepia; segment II sordid buffy yellow; distal end of
tarsal segment I and segment II light fuscous; remainder of tarsal segment
I, segment II, most of tibial length and proximal one-third to one-half of
femur on all legs pale buffy yellow.

Antennae, legs, labium and abdomen smooth, impunctate, moderately
shining and evenly clothed with fine recumbent hairs; longer erect hairs
present on forelegs, head, scutellum and dorsal surface of pronotum; head
shining with numerous nearly contiguous micropunctures, this surface re-
flecting the light in a characteristic broken pattern; pronotum, scutellum and
hemelytra including membrane dull, clothed with a pale gray pruinosity
which is densest and most apparent on anterior portion of posterior pronotal
lobe, in a pair of patches on base of scutellum and in elongate patches on
clavus, one along claval suture at level of scutellum and a smaller one
parallel to claval commissure (these areas of heavy pruinosity give a light
appearance to and partially mask the actual dark color of the integument
beneath); pruinosity of anterior pronotal lobe broken by irregular anasto-
mosing shining spots scattered over calli and by a shining lateral line on
propleuron; anterior pronotal lobe very faintly and sparsely punctate lat-
erad; a close row of distinct punctures marking transverse impression; pos-
terior pronotal lobe impunctate on anterior one-third; posterior two-thirds
of posterior pronotal lobe (save humeral angles), scutellum and hemelytra
distinctly punctate.

Head sharply decurved anteriorly from a dome-like vertex between ocelli;
juga and tylus not visible in dorsal view, even antenniferous tubercles barely

VOLUME LXXXIX, NUMBER 3

195

visible from above; post-ocular portion of head broad immediately behind
eyes, constricted more posteriorly to form a cylindrical stalk-like neck; eyes
rounded and protruding; length head 1.00; width 0.94; interocular distance
0.54. Anterior pronotal lobe globose; with a distinct anterior collar de-
marked by a line-like groove; transverse impression deeply line-like; pos-
terior pronotal margin straight across base of scutellum, humeral angles
rounded; length anterior pronotal lobe 0.46; width 0.70; length posterior
lobe 0.40; width across humeral angles 1.12; width transverse impression
0.64. Length scutellum 0.42; width 0.52. Length corium 1.96; distance apex
corium to apex membrane 0.74; length claval commissure 0.42; distance
apex clavus to apex corium 0.90. Labium extending onto anterior portion
of mesosternum, not attaining mesocoxae; length labial segments I 0.50, II

O. 50, III 0.32, IV 0.30; bucculae short and rounded, cupped around base of
labium. Antennae slender; length antennal segments I 0.36, II 0.78, III 0.72,
IV 0.88. Total length 4.70.

Holotype: Brazil: S Natal, Mann. coll. In American Museum of Natural
History.

Paratypes : Surinam: \S Paramaribo 26-11-1962 P. H. v. Doesburg Jr.
Brazil: 1$ Taperina; 29$, 3c 3d Santarem. In Leiden Museum, Carnegie
Museum (Acc. No. 2966), J. A. Slater and B. J. Harrington collections.

Color variation in the type series is slight and only one of degree. In
several of the paratypes the head is less abruptly decumbent anteriorly than
is that of the holotype so that the tylus is apparent in dorsal view.

P. microlevis is closest in appearance to, and might be mistaken for, P.
levis. However, P. microlevis is a smaller species, typically about 5 mm
long, with the basal one-third of the fore femora pale and contrasting with
the darker distal portion and the clavus uniformly chestnut although marked
by two elongate patches of pale gray pruinosity that occur in the same
general areas as the actual pale vittae on the clavus of P. levis. These
features will serve to distinguish specimens of P. microlevis from those of

P. levis which are generally about 7 mm in total length; show distinct pale
claval vittae paralleling the claval suture and claval commissure; and have
the fore femora not pale at the base but uniformly tawny. In both of these
species the proximal one-third to one-half of the meso- and metafemora are
pale in contrast to the distal portions.

Acknowledgments

I wish to express appreciation for the loan of materials for study by the
following individuals and institutions: Drs. R. T. Schuh and P. Wygodzinsky
(American Museum Natural History); Dr. R. C. Froeschner (National Mu-
seum of Natural History, Smithsonian Institution); Dr. G. E. Wallace (Car-
negie Museum); Dr. J. D. Lattin (Oregon State University); Mr. S. L. Szer-
lip (University of California, Berkeley); Drs. J. Becker and J. C. M.

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NEW YORK ENTOMOLOGICAL SOCIETY

Carvalho (Museau Nacional, Rio de Janeiro); Dr. P. H. van Doesburg (Lei-
den Museum); and J. A. Slater (personal collection). I thank Barbara M.
Kratochvil for her preparation of the dorsal view illustration of P. micro-
levis. This research was supported by the College of Agricultural and Life
Sciences, University of Wisconsin, Madison.

Literature Cited

Barber, H. G. 1954. Some neotropical “Lygaeidae." Rev. Brasil. Biol. 14(2):2 15— 224.
Distant, W. L. 1880-1893. Insecta. Rhynchota. Hemiptera-Heteroptera. Vol. I. Biol. Centr.
Amer. London (Lygaeidae 1882:173-220).

Harrington, B. J. 1980. A generic level revision and cladistic analysis of the Myodochini of
the world (Hemiptera, Lygaeidae, Rhyparochrominae). Bull. Amer. Mus. Nat. Hist.
167(2):45— 1 16.

Palmer, R. S. 1962. Handbook of North American birds. Vol. I. Loons through Flamingos.
New Haven, Yale Univ. Press, col. pi.

Scudder, G. G. E. 1962. The world Rhyparochrominae (Hemiptera: Lygaeidae) II. New genera
for previously described species. Canadian Ent. 94:981-989.

Department of Entomology, University of Wisconsin, Madison, Wiscon-
sin 53706.

Received for publication February 12, 1981.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(3), 1981, pp. 197-201

AGGREGATION BEHAVIOR OF APLOMYIOPSIS XYLOTA
(DIPTERA: TACHINIDAE)

Frank J. Messina

Abstract. — The tachinid Aplomyiopsis xylota is a common parasitoid of
larvae of Trirhabda virgata and T. borealis (Coleoptera: Chrysomelidae) in
central New York. Flies were observed to aggregate on herbaceous vege-
tation along the borders between old fields and more shaded habitats (shrub-
by fields or woodlots). Aggregations comprised from less than one hundred
to a few thousand individuals, almost all of which were male. Males con-
tinuously perched and moved about in sun flecks on the vegetation, and
frequently grappled with each other. In the old field adjacent to the primary
aggregation site, the tachinid sex ratio was strongly skewed toward females.
Aggregations of A. xylota are probably involved in mating.

Tachinid flies in the genus Aplomyiopsis Villenueve parasitize chryso-
melid larvae, particularly of the subfamily Galerucinae (Arnaud 1978; D. M.
Wood, pers. comm.).1 The life histories and hosts of most species of Aplo-
myiopsis are unknown. Sholes (1977) obtained an unidentified species of
Aplomyiopsis from the galerucine Trirhabda virgata LeConte. Hogue (1970)
reported that several western species of Trirhabda were parasitized by A.
xylota (Curran). During a study of Trirhabda virgata and T. borealis Blake
in central New York, I also found high rates of parasitization of these beetles
by A. xylota. This paper presents one aspect of the biology of this fly, its
aggregation behavior.

Several families of cyclorrhaphous Diptera are known to form adult ag-
gregations (Hunter and Webster 1973; Tychsen 1977). Aggregations typi-
cally consist almost entirely of males, in a kind of ‘‘waiting station” assem-
bly (Downes 1969). Females are presumed to visit aggregation sites to mate.
The site of assembly may be highly specific; certain tachinid species aggre-
gate on mountain summits, hilltops, man-made towers, and other vertical
prominences (Chapman 1954; Shields 1967; Lederhouse et al. 1976). I de-
scribe here the characteristics of aggregation sites of A. xylota and the
behavior of individuals in the aggregation.

Methods

Observations were made during 1979-1980 in old fields at the Whipple
Farm, 8 km NE of Ithaca, N.Y. (Fig. 1). The vegetation was dominated by

1 The species epilachnae (Aldrich) parasitizes coccinellid larvae, but is no longer placed in
Aplomyiopsis (D. M. Wood, pers. comm.).

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NEW YORK ENTOMOLOGICAL SOCIETY

Fig. 1. Map of the Whipple Farm. Arrows indicate aggregation sites. See text for expla-
nation of letter symbols.

co

Fig. 2. Seasonal densities (number/ 100 sweeps) of Trirhabda larvae and tachinid adults in
1979. Dotted lines indicate periods when aggregations were observed.

VOLUME LXXXIX, NUMBER 3

199

goldenrod ( Solidago spp.). In 1979, the seasonal abundance of Aplomyiop-
sis adults and Trirhabda larvae was estimated by taking weekly sweep sam-
ples (100 sweeps/date) along 4 permanent transects (site C in Fig. 1, further
described in Messina and Root, 1980). Voucher specimens of the 2 Tri-
rhabda spp. and A. xylota were placed in the Cornell University Collection
in Lot No. 1068.

Results

In 1979, adult flies were first collected in sweep samples in mid-May,
shortly after the appearance of their larval hosts (Fig. 2). The tachinid pop-
ulation remained at relatively low density until late June, when flies emerged
from Trirhabda larvae in peak numbers. Fly density in the old field then
declined rapidly, and flies were not found after July.

During 2 periods in 1979 and 1980 (Fig. 2), I observed flies aggregating at
several locations along the border between the old field vegetation and the
shrubby fields and woodlots (Fig. 1). Aggregations in May were small, con-
sisting of fewer than 50 individuals. Each aggregation occupied an area less
than 0.5 m2. In late June and July, aggregations were much larger, with
several thousand individuals clustered in 2 to 3 square meters at the primary
aggregation site (site A in Fig. 1). Flies were clustered on the lower vege-
tation, which included goldenrod ( Solidago spp.), aster ( Aster spp.), Vir-
ginia creeper ( Parthenocissus quinque folia), poison ivy ( Rhus toxicoden-
dron), and common barberry ( Berberis vulgaris). The taller vegetation at
the aggregation sites consisted of dogwood ( Cornus spp.), arrow- wood
( Viburnum spp.), shadbush ( Amelanchier sp.), white ash ( Fraxinus amer-
icana), shagbark hickory ( Carya ovata), black cherry ( Prunus serotina),
and apple ( Pyrus malus).

Flies were found particularly in the sun flecks on the leaf surfaces at these
sites. Individuals alternately perched in the sun flecks, made short, darting
flights, and returned to the leaves. Aggregations were maintained throughout
the day, and the precise location of the flies changed according to the shifting
positions of the sun flecks. In the densest aggregations, flies were tightly
clustered, with up to 15 individuals perched on a single goldenrod leaf. A
fly in such a cluster frequently oriented toward and lunged onto adjacent
individuals. These individuals invariably grappled for an instant and sepa-
rated. Individuals did not maintain a consistent position in the aggregation.
Vespula wasps occasionally hovered about and preyed on flies in the ag-
gregation.

On 9 July 1980, 99% of the individuals collected from the primary aggre-
gation site with a sweep net were male (Table 1). A simultaneous sweep
sample from several meters away in the open field (site B in Fig. 1) yielded

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NEW YORK ENTOMOLOGICAL SOCIETY

Table 1. Sex ratio of A. xylota populations in the field and laboratory.

Male

Percent

Female

N

p*

Reared from Trirhabda in laboratory

45

55

42

ns

At aggregation site (9 July 1980)

99

1

85

<0.001

At field site (9 July 1980)

15

85

59

<0.001

* Chi-square test.

mostly females. Females in the open field were observed ovipositing on the
few remaining Trirhabda larvae at this time. Sex determinations of flies
emerging from larval hosts in the laboratory indicated a primary sex ratio
of 1:1 (Table 1).

Discussion

Male aggregations of A. xylota differed from other reported muscoid ag-
gregations (e.g., Lederhouse et al. 1976) in that flies were not clustered at
or near the top of any vertical peak. Flies were located on the lower, her-
baceous vegetation, and the sites themselves occurred in an area of rela-
tively flat topography. While hilltops usually contain multi-species assem-
blages (Chapman 1954), only A. xylota was observed aggregating at the
Whipple Farm sites. It is not known how individuals choose where to ag-
gregate; they may orient toward the relatively abrupt changes in either light
intensity or vegetation structure. Individuals clearly respond on a smaller
scale to changes in light intensity or temperature by constantly positioning
themselves in sun flecks.

Aggregations of A. xylota are probably involved in mating; if so, this
activity would be spatially separated from host-finding or oviposition. Po-
tential food sources of larval hosts were not available at the primary aggre-
gation site, which produced the most abundant aggregation in each year.
The collection of a single female at this site suggests that females do visit
it occasionally. The observed lunging and grappling behavior resembled the
"trial and error” or "assault” mating behavior of the Cyclorrhapha (Thom-
as 1950; Spieth 1974).

Acknowledgments

I thank G. C. Eickwort and D. Bickel for suggestions, Dr. C. W. Sabrosky
(USNM) for identification of A. xylota, and P. L. Marks for plant identifi-
cations. This research was supported by NSF Grant No. DEB 77-25120 to
R. B. Root.

VOLUME LXXXIX, NUMBER 3

201

Literature Cited

Arnaud, P. H., Jr. 1978. A Host-Parasite Catalog of North American Tachinidae (Diptera).
SEA, USDA, Misc. Pub. No. 1319.

Chapman, J. A. 1954. Studies on summit-frequenting insects in western Montana. Ecology
35:41-49.

Downes, J. A. 1969. The swarming and mating flight of Diptera. Ann. Rev. Entomol. 14:27 1—
298.

Hogue, S. M. 1970. Biosystematics of the genus Trirhabda LeConte of America north of
Mexico (Chrysomelidae: Coleoptera). Ph.D. Diss., Univ. of Idaho. 212 pp.

Hunter, D. M. and J. M. Webster. 1973. Aggregation behavior of adult Cuterebra grisea and
C. tenebrosa (Diptera: Cuterebridae). Can. Entomol. 105:1301-1307.

Lederhouse, R. C., R. A. Morse, J. T. Ambrose, D. M. Burgett, W. E. Conner, L. Edwards,
R. D. Fell, R. Rutowski and M. Turell. 1976. Crepuscular mating aggregations in certain
Ormia and Sitophaga. Ann. Entomol. Soc. Amer. 69:656-658.

Messina, F. J. and R. B. Root. 1980. Association between leaf beetles and meadow goldenrods
(Solidago spp.) in central New York. Ann. Entomol. Soc. Amer. 73:641-646.

Shields, O. 1967. Hilltopping. J. Res. Lepid. 6:69-178.

Sholes, O. D. 1977. Parasitism of Trirhabda virgata LeConte (Coleoptera: Chrysomelidae)
by a species of Aplomyiopsis Villenueve (Diptera: Tachinidae). J. Wash. Acad. Sci.
67:157-158.

Spieth, H. T. 1974. Mating behavior and evolution of the Hawaiian Drosophila. In: Genetic
Mechanisms of Speciation in Insects. M. J. D. White, ed., pp. 94-101. Australia and
New Zealand Book Co., Sydney.

Thomas, H. T. 1950. Field notes on the mating habits of Sarcophaga Meigen (Diptera). Proc.
Roy. Ent. Soc. Lond. (A) 25:93-98.

Tychsen, P. H. 1977. Mating behaviour of the Queensland fruit fly, Dacus tryoni (Diptera:
Tephritidae), in field cages. J. Aust. Entomol. Soc. 16:459-465.

Section of Ecology and Systematics, Cornell University, Ithaca, New
York 14853.

Received for publication February 10, 1981.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(3), 1981, pp. 202-203

HESPERIIDAE AS PREY OF STICTIA CAROLINA 1
A. Hook

Abstract. — Many dipterous prey records exist for Stictia Carolina (F.).
Only one study reports the use of hesperiids and cicadas in addition to flies
as prey items. The present study confirms the use of hesperiids as prey of
S. Carolina ; of 9 unicellular nests examined, 4 contained hesperiids in ad-
dition to dipterous prey.

Abundant prey records exist for Stictia Carolina (F.), all but one reporting
Diptera, especially Tabanidae. Only Lin (1971) has noted the use of hes-
periids ( Atalopedes campestris Boisduval) and small cicadas, in addition to
Diptera, as prey of S. Carolina (studied in Oklahoma).

This paper confirms the use of Hesperiidae as prey of S. Carolina. Field
research was conducted during Aug 1979, on state forest land, situated 300
m west of Turkey Swamp State Park, Freehold, NJ. Two nesting aggrega-
tions, each with 20 to 40 females, were located in small, sandy clearings.
Surrounding habitat consisted of a mixed deciduous and evergreen forest
(pine barrens type flora).

Females had begun to nest and males were still active when these sites
were 1st located on 9 August. Five unicellular nests were excavated on 16
Aug and were found to contain dipterous prey, but 2 cells also had hesperiid
wings. Four additional nests excavated on 24 Aug revealed 2 cells bearing
skippers and flies. One skipper was determined as Epargyreus clarus (Cra-
mer). In 9 nests, 100% contained Diptera while 44% also had hesperiids (1
or 2 skippers per cell).

In the Bembicini, 6 genera are known to prey on Hesperiidae: Rubrica,
Stictia, Editha, Bembix, Zyzzyx and Stictiella (Evans 1966; Evans et al.
1974; Bohart and Menke 1976). Only in the case of some Stictiella do Hes-
periidae provide a major source of prey. Lin (1971) suggested that shortage
of dipterous prey had led S. Carolina to select skippers and cicadas in
addition to Diptera. Further comparative as well as experimental studies
are needed to clarify prey selection in this group.

Literature Cited

Bohart, R. M. and A. S. Menke. 1976. Sphecid Wasps of the World: a Generic Revision.
Univ. Calif. Press, Berkeley. 695 pp.

Evans, H. E. 1966. The Comparative Ethology and Evolution of the Sand Wasps. Harvard
Univ. Press, Cambridge, Mass. 526 pp.

VOLUME LXXXIX, NUMBER 3

203

, R. W. Matthews and E. McC. Callan. 1974. Observations on the nesting behavior of

Rubrica surinamensis (DeGeer) (Hymenoptera: Sphecidae). Psyche 81:334-352.

Lin, C. S. 1971. Bionomics of Stictia Carolina at Lake Texoma, with notes on some Neo-
tropical species (Hymenoptera: Sphecidae). Tex. J. Sci. 23:275-286.

Department of Zoology and Entomology, Colorado State University, Ft.
Collins, Colorado 80523.

Received for publication February 20, 1981.

1 This research was supported by a National Science Foundation Grant (BNS 79-26655)
awarded to H. E. Evans, and by a Sigma Xi R.E.S.A. Grant-in-Aid of Research.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(3), 1981, pp. 204-213

NOTES ON THE MOTH PERICOPUS LEUCOPHAEA WALKER
(LEPIDOPTERA: PERICOPIDAE) AS A DEFOLIATOR OF
THE TREE VERNON I A PATENS H.B.K. (COMPOSITAE)

IN NORTHEASTERN COSTA RICA

Allen M. Young

Abstract. — Two closely-spaced individuals of the woody composite Ver-
nonia patens H.B.K. were severely defoliated by the brightly-colored, gre-
garious larvae of the moth Pericopus leucophaea Walker (Lepidoptera: Pe-
ricopidae) in northeastern Costa Rica. Although many other individuals
were examined, no other V. patens were found to be defoliated. A group
of 66 larvae was found on the larger tree and 20 on the smaller one. At the
beginning of a three-day period of observation, about 70% of the larger tree
and 40% of the smaller tree was defoliated. Defoliation apparently begins
on the lower leaves and spreads towards the top. No larvae disappeared
during the study period, suggesting high survivorship. Although individuals
of the social paper wasp Polybia simillima Smith (Hymenoptera: Vespidae:
Polistinae: Polybiini) made repeated attempts to capture larvae, all were
unsuccessful. When disturbed by wasps, larvae drop from the plant, either
suspending themselves on silken threads or falling into the dense undercov-
er. Larvae thus dislodged successfully relocate the plant as they are capable
of very rapid locomotion. When prodded with forceps, larvae exhibit strong
thrashing movements and exude droplets of a clear fluid which may have
a defensive function. Both the aposematic appearance and behavior of the
larvae (and adults) suggest unpalatability. Unusually attractive conditions
of individual plants, and high survival of egg masses and larvae, probably
promote defoliation. Various factors are predicted to determine intensity of
defoliation and comprise a basis for further study of this interesting system.

Vernonia patens H.B.K. is a locally abundant woody composite of sec-
ondary-growth habitats in the premontane and lowland tropical wet forest
regions of northeastern Costa Rica. This small tree (canopy about 4-5
meters tall) of woody shrub often grows as small clumps or single individuals
in regenerating fields and along the edges of forest, habitats where it is used
as a feeding and chorusing site for various species of cicadas (Homoptera:
Cicadidae) (Young 1980a, b). The purpose of this brief communique is to
report the severe defoliation of two individuals of this tree species by the
gregarious larvae of the medium-sized (i.e. wingspan of 40 mm) moth Peric-
opus leucophaea Walker (Lepidoptera: Pericopidae) in this region of Costa

VOLUME LXXXIX, NUMBER 3

205

Fig. 1. The larger of the two Vernonia patens (Compositae) defoliated by the gregarious
larvae of the moth Pericopus leucophaea (Lepidoptera: Pericopidae) at one site in northeastern
Costa Rica, February 1981. Note the severe defoliation of the lower branches of this tree and
the relatively intact crown of young leaves. When disturbed by Polybia wasps the aposemat-
ically-colored larvae drop off the tree into the dense ground cover below, and then crawl back
to the same plant.

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NEW YORK ENTOMOLOGICAL SOCIETY

Rica during the dry season. The general pattern of defoliation of the twcn
trees, and the ineffectiveness of a social paper wasp, Polybia simillimai
Smith (Hymenoptera: Vespidae: Polistinae: Polybiini) as a predator on the
larvae of this species, are described for the first time.

Methods

On 3 February 1981, I discovered two closely-spaced Vernonia patens
trees, one about 1.5 meters tall and the other about 2.5 meters, in different!
stages of noticeable defoliation by the gregarious larvae of an unidentified
moth species. The location of these trees, about 3 meters apart, is along a
gravel road at “Finca La Tigra,” near La Virgen (10°23'N, 84°07'W; 220
m. elev.), Heredia Province, Costa Rica. The habitat is a road-side patch
of regenerating secondary vegetation, and no other individuals of V. patens
were found within 20 meters in any direction from this “island.” Between
3 and 5 February, I made daily observations on the number of larvae pres-
ent, their behavior, the amount of defoliation of each tree, and the inter-
action of the larvae with other arthropods. I checked the two trees at various
times each day, to determine larval feeding patterns, if any, and changes in
the location of larvae on the trees. A sample of 20 larvae was collected the
evening of 5 February and confined, along with fresh cuttings of the food
plant, to a clean, clear-plastic bag, for rearing to the adult stage since I had
to leave the site at this time. The captured larvae were taken to other sites
in Costa Rica, until 25 February, at which time they were taken to Milwau-
kee, Wisconsin (as pupae) to complete the rearing.

At several times during the field study, the larvae were prodded with
forceps to elicit defensive behavior.

Results i

At the time of discovery, approximately 70% of the leaves from the larger
of the two trees had been stripped from the trees (Fig. 1), and about 40%
of the leaves of the smaller tree were gone. That the leaves had been eaten
by the larvae was verified by examining the number of chewed petioles, and
distinguishing these from regular leaf-drop.

Based on voucher specimens of larvae, pupae, and adults, the moth was
identified later as P. leucophaea. No other lepidopterous larvae were found
on the trees, and with the exception of an occasional orthopteran, no other
herbivorous insects were found during daytime hours. A brief inspection of
an additional 20 individuals of V. patens of similar size, scattered over the
site, failed to turn up additional larvae of P. leucophaea. At the time of
discovery the larvae were easy to spot from a distance of about 5 meters
with unobstructed view from the road, owing to their bright color patchwork

VOLUME LXXXIX, NUMBER 3

207

Fig. 2. Gregarious behavior and defoliation by larval Pericopus leucophaea. Left: cluster
of larvae, forming a conspicuous mass (to the human observer) amidst the defoliated branches
of the food plant and just below the crown. Right: the larvae form tightly-packed aggregations
on branches, petioles, and leaf midribs, bared by defoliation.

of red, white and black, and their gregarious habit (Fig. 2). At this time the
larvae were about 15 mm long. During the three days of observation, there
were 66 larvae on the larger tree, and 20 on the smaller tree.

Larvae feed intermittently throughout the day, but become very quiescent
by 5:30 p.m. As many as ten Ectatomma ants were seen crawling over the
leaves of each tree during this study, sometimes on the same leaves where
larvae were resting or feeding. The ants did not attack the larvae. Defoliation
apparently started near the bottom of the tree since the crown of each tree
was largely intact (Fig. 1), and most of the Ectatomma were found on these
young leaves.

Tight clusters of larvae were seen on defoliated petioles and leaf midribs
(Fig. 2), rendering them very conspicuous to the human eye. When resting
in clusters (Fig. 2), individual larvae, when gently poked with a forceps,
exhibit rapid thrashing movements of the anterior half of the body, and
produce small clear droplets, which were first noticeable on some of the

208

NEW YORK ENTOMOLOGICAL SOCIETY

Fig. 3. Gregarious feeding and defensive behavior by the larvae of Pericopus leucophaea.
Top: a typical larval aggregation on the underside of a mature leaf of Vernonia patens. The
light areas on the bodies of the larvae are bands of white and orange interspersed with bands
of black. Below, left: gregarious feeding. Below, right: the larva in full view at the top has just
exuded a large droplet of clear fluid on the dorsal bristles just before the posteriorly-terminal
segments. This fluid may have a defensive function.

VOLUME LXXXIX, NUMBER 3

209

long hairs of body segments (Fig. 2). Sometimes clusters of many larvae
were seen on the undersides of leaves, and such leaves were usually quickly
defoliated (Fig. 3). The positions of groups varied greatly each day, presum-
ably as a result of feeding activity. Sometimes there were both solitary
larvae and several small clusters (Fig. 3) on the larger tree.

On 4 February, I observed from 1 to 3 individuals of the polybiine wasp
Polybia simillima make repeated attempts to capture larvae, either ones in
clusters or single individuals. From a total of 45 predatory-attack attempts
observed, none was successful. On a typical attack, a wasp would weave
among the leaves and upon discovering one with larvae on it, it would then
land on the dorsal side of the leaf, crawl a bit, and then quickly flip over to
the ventral side to capture a larva. But at the moment the wasp landed,
most of the larvae dropped off the leaf, some suspended on long silken
threads, while others (most) fell into the dense herbaceous layer beneath
the tree. For a total of twelve larvae followed after falling into the dense
cover on different occasions, each one returned to the food plant. The
larvae crawl very rapidly when disturbed, and one dislodged from the food
plant is usually back on it within five minutes after a fall. Larvae sus-
pended on threads usually dropped to the ground and then crawled to
the food plant.

Deliberate mechanical disturbance of larvae on leaves with forceps does
not elicit the “dropping off’ behavior. In the larger group of larvae, there
were two sized classes apparent, the larger sub-group of 45 individuals being
about 15 mm long, and the smaller ones, about 10 mm long. All individuals
in the group of twenty on the smaller tree were the same size (20 mm long).

In captivity, larvae pupated about 18 days after discovery, each one form-
ing a loose cocoon wrapped in a leaf. The pupa stage lasted about 18 days
in this study.

By the end of the field study, both trees were almost entirely defoliated,
save for small crown areas of young leaves on each one. Since it required
more than two additional weeks to bring larvae to pupation, both trees
would have been insufficient total food supply for the larvae under natural
conditions.

Discussion

The defoliation of individual food plants by a herbivorous insect species
in the tropics may be considered an “outbreak” if a large portion of the
local food plant population is defoliated. The defoliation of a few individuals
of such a population, however, may be a result of isolated patches where
survival of the herbivore is unusually high, and may not necessarily be an
(“outbreak.” A variety of ecological factors, including the degree to which

210

NEW YORK ENTOMOLOGICAL SOCIETY

patches of varying size of a plant species experience severe defoliation,
determine pest outbreak conditions in the tropics (e.g., Rey et al. 1981).

While the present study on the interaction of P. leucophaea with V:
patens is very preliminary, the data suggest that defoliation of a few closely
spaced individuals of the food plant resulted from an “escape” of a few egg
clusters of the moth from regulatory agents such as predators and parasit-
oids. There is, however, a growing body of evidence showing that the early
stages of Lepidoptera are sources of prey for a variety of arthropods (e.g. j
Gilbert and Singer 1975).

There have been very few studies of pericopid moths anywhere, save for
some systematic and survey studies (e.g., Kendall 1970, 1974; Beutelspach-
er 1976) and very little is known about the food plant associations of indi-
vidual species within and outside of the tropics. Presumably the moth places
large egg clusters on the food plant, and upon hatching, the larvae exhibit
pronounced gregariousness as shown in the present study. Studies of noc-
tuid and arctiid moths feeding on Vernonia species indicate considerable
food plant specificity in terms of species exploited (e.g., Tietz 1951; Burnett
et. al. 1974). To the human observer, the brightly-colored larvae of P. leu-
cophaea and their gregarious behavior on the food plant suggest aposematic
display and unpalatability. Some Vernonia contain high concentrations oi
bitter sesquiterpene lactones in the leaves (Abdel-Baset et al. 1971; Mabry
et al. 1975), which could be the basis for suspected unpalatability of some
herbivores such as P. leucophaea if these compounds are incorporated intc
the insects. These compounds have also been demonstrated to be a basis
for oviposition preferences in various Lepidoptera exploiting some species
of Vernonia (Burnett and Jones 1978).

Because sesquiterpene lactones in some Vernonia have been shown tc
deter feeding by some Lepidoptera (Burnett et al. 1974), the aposematic
coloration of both the larvae and adult stages of P. leucophaea may be
indirect evidence that this species is able to sequester and incorporate these
compounds. Relative to what is known about the feeding-deterring prop-
erties of these compounds for other Lepidoptera (Burnett et al. 1974), and
assuming these compounds to be present in the age-classes of leaves of V.
patens exploited by larval P. leucophaea, this moth may have entered into
a new adaptive zone (in the context of Ehrlich and Raven 1965). The vivid
color pattern of the adult moths, with wings banded in black, white, and
red, not only suggests unpalatability, but also mimetic resemblance to fe-|
male Parides (Lepidoptera: Papilionidae: Papilioninae: Troidiini) found in
the same habitats (e.g., P. iphidamas — see Young 1977b). Adult P. leuco -j
phaea fly during the daytime and could possibly participate in a Mullerian
mimicry association with these Aristolochia-feeding butterflies, even though
the moths are somewhat smaller than typical female Parides.

VOLUME LXXXIX, NUMBER 3

211

The association of P. leucophaea with V. patens in Costa Rica may be
a very specific interaction. It would be expected that a group-feeding bright-
ly-colored herbivore such as P. leucophaea would have evolved effective
defense mechanisms against predators and parasitoids. My preliminary ob-
servations on the ineffectiveness of the wasp P. simillima to successfully
capture the larvae supports the contention of effective defenses being op-
erative. Thus, in addition to the suspected systemic unpalatability, the com-
position and functional role of the observed droplets of fluid exuded by the
larvae when prodded warrants further study in this context.

A plausible working hypothesis for detailed study of the P. leucophaea x
V. patens interaction in Costa Rica may be that individual egg masses, if
escaping from predation or parasitism, result in large groups of larvae which
generally have high survival on the food plant. Although Polybia wasps at
this locality are successful at capturing the gregarious larvae of the butterfly
Mechanitis isthmia Bates (Lepidoptera: Ithomiidae) on Solanum spp. (So-
lanaceae) (Young 1978), presumably the larvae of P. leucophaea, although
also gregarious, have an effective means of making Polybia attacks very
unsuccessful, if my observations are representative of such interactions. In
the Mechanitis study, Ectatomma was also present in close proximity to
the larvae, and neither attacked the larvae nor provided an effective defense
against the Polybia attacks. In the present study, the effectiveness of P.
leucophaea larvae in escaping from Polybia attacks appears to be intrinsic
to the larvae, with Ectatomma having no apparent effects on the interaction.

When egg and larval survival is high, defoliation of individual food plants
may occur. The severity of such defoliation will be a function of the effective
size of the egg masses on each individual plant (i.e., the number of eggs
hatching) and the physiological receptivity of the individual plant to the
larvae. As shown by my preliminary data, smaller numbers of larvae on a
plant result in less damage than larger numbers on a larger plant. For P.
leucophaea, egg masses are presumably placed on lower leaves since the
defoliation begins at the bottom of the plant and spreads upwards. As my
I observations were made in the short and erratic dry season of this region,
the possibility exists that conditions resulting in localized defoliation of V.
\vatens may be seasonal.

, The ability of the larvae of P. leucophaea to crawl very rapidly may allow
them to encounter other individuals of the food plant when defoliation is
severe.

Under conditions of relaxed predation or parasitism on eggs and larvae,
one consequence of massive group-feeding is the rapid defoliation of the
! food plant. Enforced emigration to fresh plants, made possible by high lo-
comotory ability, allows the larval population to re-distribute on the food
plant population.

212

NEW YORK ENTOMOLOGICAL SOCIETY

Since V. patens often occurs in clumps of varying sizes abundant over
relatively small areas of habitat, when conditions are favorable for the sur-
vival of large numbers of eggs and larvae, larval dispersal behavior and
successful encounter of fresh food plants may prevent localized extinctions.

In the tropics in particular, defoliation by herbivorous insects should be
considered as highly transient and ephemeral conditions associated with
changes in food plant distribution, community structure, and kinds of con-
trol mechanisms operative on populations (e.g., Rey et al. 1981). The degree
to which an individual of V. patens will be defoliated by P. leucophaea is
therefore directly related to a wide variety of factors, including (a) attrac- ;
tiveness of the individual for oviposition by the moth and numbers of egg)
masses deposited, (b) survival patterns of eggs and larvae, (c) effects of
other herbivores on this individual plant, (d) the dispersal strategy of mated
female moths, (e) the degree of oviposition preference for V. patens , and
(f) the effects of seasonality on (a-e), if any. A complete understanding of
severe defoliation of plants other than crops in the tropics requires a con-
sideration of these factors. Such factors determine the carrying capacity of
the environment at any one point in time for herbivores such as P. leuco-
phaea on V. patens.

Acknowledgments

This research was funded by a grant from The American Cocoa Research
Institute. I thank Dr. J. Robert Hunter for allowing me to conduct these
studies at “Finca La Tigra.” Dr. R. W. Poole of the United States Depart-
ment of Agriculture identified the life stages of the moth, and specimens are
deposited in the USDA collections.

Literature Cited

Abdel-Baset, Z. H., L. Southwick, W. G. Padolina, H. Yoshioka, T. J. Mabry and S. B. Jones.
1971. Sesquiterpene lactones: a survey of 21 United States taxa from the genus Ver-
nonia (Compositae). Phytochemistry 10:2201-2204.

Beutelspacher, C. R. 1976. The family Pericopidae in Mexico (Lepidoptera: Heterocera). Soc.
Fol. Ent. Mex. 36:97-130.

Burnett, W. C. and S. B. Jones. 1978. Influence of sesquiterpene lactones of Vernonia (Com-
positae) on oviposition preferences of Lepidoptera. American Midland Naturalist
100:242-245.

, , T. J. Mabry and W. G. Padolina. 1974. Sesquiterpene lactones — insect feeding

deterrents in Vernonia. Biochemical Systematics and Ecology 2:25-29.

Ehrlich, P. R. and P. H. Raven. 1965. Butterflies and plants: A study in coevolution. Evolution
18:586-608.

Gilbert, L. E. and M. C. Singer. 1975. Butterfly ecology. Ann. Rev. Ecol. and Syst. 6:365-
398.

Kendall, R. O. 1970. A day-flying moth (Pericopidae) new to Texas and the United States. J.
Lepid. Soc. 24:301-303.

VOLUME LXXXIX, NUMBER 3

213

. 1974. Two moth species (Pericopidae and Notodontidae) new to Texas and the United

States. J. Lepid. Soc. 28:243-245.

Mabry, T. J., Z. Abdel-Baset, W. G. Padolina and S. B. Jones. 1975. Systematic implications
of flavonoids and sesquiterpene lactones in species of Vernonia. Biochem. Systematics
and Ecology 2:185-192.

Rey, J. R., E. D. McCoy and D. R. Strong, Jr. 1981. Herbivore pests, habitat islands, and
the species-area relation. Amer. Naturalist 117:611-622.

Tietz, H. M. 1951. The Lepidoptera of Pennsylvania. Amanual. Pa. Agric. Exp. Sta., 194 pp.

Young, A. M. 1977a. Possible coevolution of mutualism between Mechanitis caterpillars and
an ant in northeastern Costa Rica. Biotropica 10:77-78.

. 1977b. Studies on the biology of Parides iphidamas (Papilionidae: Troidini) in Costa

Rica. J. Lepid. Soc. 31:100-108.

. 1980a. Environmental partitioning in lowland tropical rain forest cicadas. J. N.Y. Ent.

Soc. 88:86-101.

. 1980b. Observations on the aggregation of adult cicadas (Homoptera: Cicadidae) in

tropical forests. Canad. J. Zool. 58:711-722.

Department of Invertebrate Zoology, Milwaukee Public Museum, Mil-
waukee, Wisconsin 53233.

Received for publication April 30, 1981.

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(3), 1981, pp. 214-217

BOOK REVIEWS

Fleas. Proceedings of the International Conference on Fleas, Ashton Wold,

Peterborough, U.K., 21-25 June 1977. R. Traub and H. Starcke, eds. A.

A. Balkema, Rotterdam. 1980. 420 pp. $49.50.

This book is unique in several aspects. Beautifully published, with 150
black and white illustrations, it represents the first multiauthor treatise on
fleas (Siphanoptera), one of the most important groups of vectors of human
and animal disease agents. The list of contributors reads like who is who in
flea research, and even like who was who because it starts, properly, with
a very interesting and personal chapter devoted to Nathaniel Charles Roths-
child (1877-1923), the foremost conservationist, collector of fleas and author
of 150 scientific publications (more than 100 dealing with fleas), who did his
scientific work “in his spare time” while working for the Rothschild banking
house. His famous daughter, the world’s renowned authority on fleas, Dr.
Miriam Louisa Rothschild, herself the author of more than 250 scientific
papers, together with her son, Dr. Charles Lane, hosted the 80 participants
of the international conference at Ashton Wold.

The first two chapters (pp. 13-172), by Prof. Traub, deal with new genera
of Siphonaptera and with adaptive modifications in fleas. The excellent il-
lustrations, prepared by the author, are very well reproduced in these chap-
ters, documenting the convergent evolution and flea taxonomy. Following
chapters, of various lengths, reflect the extensive experience of the authors !
and the high level of excellence in the area of flea research. R. L. C. Pilgrim
describes fleas of New Zealand. The New World host associations of Pulex
are dealt with by Cluff E. Hoppla. James R. Busvine illustrates his chapter
on the four Fs (fleas, fables, folklore and fantasies) with drawings of diverse
flea traps that I found most interesting and amusing. The control of flea
vectors of plague is described by Norman G. Gratz. Ecological interrela- .
tionships among fleas are covered by D. C. Cavanaugh and J. E. Williams, j
Harry Hoogstral describes flea vectors of rickettsiae and trypanosomes.
Flea studies in the U.S.S.R. are reviewed by V. A. Bibikova and I. F.
Zhovtyi, and a link between epizootics of sylvatic plague in California and
caves in the Lava Beds National Monument are outlined by B. C. Nelson
and C. R. Smith. Excellent electron micrographs of rickettsiae in the flea
mitdgut are provided by S. Ito and J. W. Vinson. The role of flea vectors
in murine typhus, tularemia, and myxomatosis are the subjects of shorter
reports. J. Goose describes chemical control of fleas and Rachel Galun the
specificity and biological significance of the feeding response. The last 16
contributions, two in French, deal with flea physiology, morphology, be-
havior and taxonomy. The presentation of each facet of a chapter is clear,

VOLUME LXXXIX, NUMBER 3

215

lucid, and to the point. Those desiring detailed information on fleas will find
for the first time in a single volume a complete up-to-date survey of the
subject.

This book will appeal to a very wide audience. I recommend it highly for
use in entomology departments and in medical schools.

Karl Maramorosch, Waksman Institute of Microbiology , Rutgers — The
State University , New Brunswick, New Jersey.

An Atlas of the Fleas of the Eastern United States. Allen H. Benton. Mar-
ginal Media, Fredonia, N.Y. 1980. 177 pp. 63 maps. Looseleaf, Soft Cov-
er. $6.50.

The book was designed to facilitate the study of the geographic distri-
bution of the fleas of the eastern United States by permitting the qualified
reader to spot the known records and to readily add and map new data as
they appear.

There are a few introductory pages on the ‘natural history and ecology’
of fleas and an outline of the history of Siphonapteran studies for the area.
A checklist of the fleas of the eastern U.S. is presented ahead of the atlas
of 63 pages of maps, on which dots are used to indicate the known records,
arranged by county. A set of larger maps near the front of the book shows
the counties for the states encompassed. A glance at the maps suffices to
grasp the known information on any of the species regarding its distribution
within the region in question. For each species listed, the author also pro-
vides a summary of available information on hosts, seasonal abundance,
and medical importance. One or two references are cited in each case as
sources for additional data. Pertinent state and regional references and some
general ones, are cited at the end of the book, preceding an appendix on
the distribution of the fleas of Minnesota. This book does not purport to be
a guide for the identification of fleas, nor is it a taxonomic work in any sense
of the term. Not even the original descriptions are cited.

The author worked diligently and accurately in checking references and
in tracking down and verifying certain critical specimens. None of the
species known to occur in the territory seem to have been omitted, and the
scientific names used agree with the standard literature. The comments on
bionomics and seasonal variations for the various species will be appreciated
by all who have had to comb scattered references seeking such meaningful
information. The appendix has valuable ecological notes on Minnesota.

The manner in which the book is organized presents problems for the
reader. The list of species included in the atlas (pp. 24-26) is arranged by
family and subfamily, but there is no indication of the pages where the

216

NEW YORK ENTOMOLOGICAL SOCIETY

particular maps and discussion are to be found. The species are not arranged
alphabetically (even by family) in the list or in the sequence of maps, and
it is difficult to find the appropriate page one seeks, even if he is familiar,
with the scheme of classification and the system used. The sequence em-
ployed is based upon that employed by Hopkins and Rothschild in the Cat-
alogues or by R. E. Lewis in his host-lists, but the Catalogue series had
indices and Lewis’ papers were arranged alphabetically and hence are easy
to use. Benton apparently relies upon the maps to indicate what he means
by “eastern United States,” but the territory included does not have eco-
logical or natural boundaries. Minnesota is treated, but not Iowa, Missis-
sippi, Arkansas and Louisiana, all of which have a western boundary that
is east of that of Minnesota. If Minnesota were not discussed, it could be
stated that the book was restricted to the states east of the Mississippi River.
The appendix on Minnesota is somewhat confusing regarding its biblio-
graphic citation, since the authors are not designated. By checking the bib-
liography it can be deduced that the ‘senior author’ mentioned here and
there in the text is Benton, and the ‘junior author,’ R. Timm.

There are inherent disadvantages in any atlas on fleas because of the
inadequacy of available data. Even in the U.S. there are vast areas where
little or no collecting has been done. In consequence, many species-maps
may unavoidably mislead the non-specialist into believing that the blanks
represent the absence of species instead of the absence of data. Many of
the fleas listed seem to be found only near certain university-towns harbor-
ing entomologists or mammalogists. The following example illustrates the
problem. Bird-fleas would be expected to have a broad range, considering
the extent of the territory covered by their hosts (both as species and as 1
individuals) but Ceratophyllus celsus is listed only from four states and in
three widely-separated areas.

The author must have been thinking of the U.S. and Canada when he
wrote “Flea students are extremely fortunate in having the entire literature
in their field quite well presented in a few excellent sources.” The statement
does not apply to Asia, South America, etc. The notes on medical signifi-
cance refer to the species in general and not merely to the eastern part of
this country. For example, unless the reader realizes that plague does not
occur in the east, he can be misled. It is regretted that no conclusions,
generalizations or interpretations of the data are presented, and that there
is no discussion of zoogeography, for example. It also would have been
edifying to compare the range of the host with that of its fleas. Thus, pocket
gophers occur in Georgia and Florida, but their characteristic fleas are un-
reported south or east of Illinois and western Indiana. Unfortunately, the
format employed precluded the presentation of data to substantiate the re-
marks. Nosopsyllus fasciatus is stated to be “most abundant in the sum-
mer.” but is this true throughout the eastern U.S. or only in the north? The
species has been considered a flea of temperate zones and the impression

VOLUME LXXXIX, NUMBER 3

217

exists that in many areas N. fasciatus is most common in cool weather.

This book will be extremely useful to those who are interested in review-
ing, compiling, or adding to data on the Siphonapteran fauna of the eastern
United States, or who want access to more information on those species.
It should also serve to encourage research along those lines, which would
be a boon, for a real need exists for such endeavors.

Robert Traub, Department of Microbiology, University of Maryland
School of Medicine, Baltimore, Maryland.

Leafhopper Vectors and Plant Disease Agents. K. Maramorosch and K. F.

Harris, eds. Academic Press, 111 Fifth Avenue, New York, NY 10003.

1979. 654 pp. Price $42.00.

This text, a companion volume to the excellent treatise on Aphids as
Virus Vectors, is part of a planned series on vectors and vector-borne dis-
eases. It is organized into the following five parts: Part I includes two chap-
ters on the taxonomy and bionomics of leafhoppers; Part II, three chapters
on the worldwide importance of leafhoppers and planthoppers as vectors;
Part III, four chapters on vector-disease agent-plant interactions; Part IV,
five chapters on experimental approaches to virus-vector and MLO-vector
research; and Part V concludes the volume with five chapters on the leaf-
hopper transmission of specific viruses and prokaryotes. These last chapters
give detailed information on rice viruses and MLOs and control of their
vectors, corn stunt, western X-disease, and xylem-borne plant pathogens.

Excellent reviews of leafhopper vector research have been contributed
by 25 authorities from eight countries. Many of the chapters are much more
than organized reviews or summaries of current research. The authors have
made a special effort to draw conclusions based on their own experiences
and have raised questions that hopefully will stimulate others to make fur-
ther contributions to leafhopper vector research. The text includes a number
of useful illustrations of good quality and many valuable summary tables.

Although information in portions of this text have appeared in past re-
view-type texts and series, this particular collection of papers will be an
excellent addition to the library of anyone concerned with vectors and vec-
tor-transmitted plant diseases. The editors are to be congratulated on putting
together a coherent volume with minimum duplication in spite of the nu-
merous contributors. The extensive lists of key references make this text
of great value to teachers of advanced entomology and plant pathology
courses.

E. H. Varney, Department of Plant Pathology, N.J. Agricultural Exper-
iment Station and Rutgers University.

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Ent.

DECEMBER 1981

No. 4

Vol. LXXXIX

of the

Devoted to Entomology in General

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The New York Entomological Society

Incorporating The Brooklyn Entomological Society

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Incorporated May 21, 1968

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Organized June 29, 1892 — Incorporated February 25, 1893
Reincorporated February 17, 1943

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The Journal of the New York Entomological Society (ISSN 0028-7199) is published quarterly for tl^e Society by Allen Press
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Journal

of the

New York

ENTOMOLOGICAL SOCIETY

Devoted to Entomology in General

VOLUME LXXXIX

published by the Society
New York, N.Y.

INDEX OF AUTHORS

AHMAD, S. and YUEN-SHAUNG NG. Further evidence for chlorpyrifos tolerance

and partial resistance by the Japanese beetle (Coleoptera: Scarabaeidae) 34

B ARC ANT, M. Breeding experiments with Morpho peleides insularis (Lepidoptera:

Nymphalidae) in Trinidad, West Indies 80

BENTON, A. H. and A. J. CRUMP. Observations on the spring and summer behavior
of the 12-spotted ladybird beetle, Coleomegilla maculata (DeGeer) (Coleoptera:

Coccinellidae) 102

dos PASSOS, C. F. A little-known, anonymous work on American and European but-
terflies and moths (1906), which should be attributed to William Beutenmuller (Lepi-
doptera: Nymphalidae) 143

FERGUSON, G. R. Synonymy and distribution records in the genus Eucerceris (Hy-

menoptera: Philanthidae) 172

FORMANOWICZ, D. R., JR. and E. D. BRODIE, JR. Prepupation behavior and pu-
pation of the predaceous diving beetle Dytiscus verticalis Say (Coleoptera: Dytiscidae)

152

GIBO, D. L. Some observations on soaring flight in the mourning cloak butterfly ( Nym -

phalis antiopa L.) in southern Ontario 98

GIBO, D. L. Some observations on slope soaring in Pantala flavescens (Odonata:

Libellulidae) 184

HARRINGTON, B. J. A revision of the genus Pephysena with descriptions of two new

species (Hemiptera: Lygaeidae) 188

HEPPNER, J. B. A World catalog associated with the Glyphipterigidae auctorum (Lep-
idoptera: Copromorphoidea) 220

HOEBEKE, E. R. The European katydid Meconema thalassinum (DeGeer): New lo-
cality records for North America (Orthoptera: Tettigoniidae) 170

HOOK, A. Hesperiidae as prey of Stictia Carolina 202

JIRON, L. F. and V. M. CARTIN. Insect succession in the decomposition of a mammal

in Costa Rica 159

KLOTS, A. B. and C. F. dos PASSOS. Studies on North American Erora (Scudder)

(Lepidoptera, Lycaenidae) 295

MARCUS, B. A. Hydropsyche larvae (Trichoptera: Hydropsychidae) from a lake
weedbed 56

i

McCABE, T. L. The larva of Agrotis volubilis Harvey (Lepidoptera: Noctuidae) 59

MESSINA, F. J. Aggregation behavior of Aplomyiopsis xylota (Diptera: Tachinidae) 197
PARKER, F. D., V. J. TEPEDINA, and G. E. BOHART. Notes on the biology of a

common sunflower bee, Melissodes ( Eumelissodes ) agilis Cresson 43

PARRELLA, M. P., J. P. McCAFFREY, and R. L. HORSBURGH. Comparison of

two sampling methods for Leptothrips mali in Virginia apple orchards 166

ROLSTON, L. H. Ochlerini, a new tribe in Discocephalinae (Hemiptera:

Pentatomidae) 40

SEIFERT, R. Principal components analysis of biogeographic patterns among Heli-

conia insect communities 109

STAMP, N. E. Parasitism of single and multiple egg clusters of Euphydryas phaeton

(Nymphalidae) 89

TROMBETTA, L. D. and J. FORBES. Histological changes in the antenna of Tenebrio

molitor after imaginal eclosion (Coleoptera: Tenebrionidae) 65

WHITE, D. J. and J. L. BENACH. Control of Dermacentor variabilis. 1. Larval and

adult susceptibility to selected insecticides 16

WHITE, D. J., J. L. BENACH, L. A. SMITH, and S. P. OUYANG. Control of Der-
macentor variabilis. 3. An analytical study of the effect of low volume spray frequency

on insecticide-stressed and nonstressed populations 23

WHITE, D. J. and C. P. WHITE. The occurrence and relevance of arthropods of
medical and veterinary importance captured during a survey on Plum Island, New

York 2

YOUNG, A. M. Notes on seasonality and habitat associations of tropical cicadas (Ho-
moptera: Cicadidae) in premontane and montane tropical moist forests in Costa Rica 123
YOUNG, A. M. Notes on the moth Pericopus leucophaea Walker (Lepidoptera: Per-
icopidae) as a defoliator of the tree Vernonia patens H.B.K. (Compositae) in north-
eastern Costa Rica 204

Book Reviews

MARAMOROSCH, K. Fleas. Proceedings of the International Conference on Fleas,
Ashton Wold, Peterborough, United Kingdom, 21-25 June 1977. R. Traub and H.

Starcke, eds — 214

SHAPIRO, A. M. Butterflies and moths of Newfoundland and Labrador: The Macro-

lepidoptera. R. F. Morris 146

TRAUB, R. An Atlas of the fleas of the eastern United States. A. H. Benton 215

VARNEY, E. H. Leafhopper vectors and plant disease agents. K. Maramorosch and
K. F. Harris, eds. 217

ii

Journal of the

New York Entomological Society

VOLUME LXXXIX DECEMBER 1981 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. Streu

Publication Committee
Dr. Randall T. Schuh
American Museum of Natural History
Dr. Louis Trombetta
St. Johns University

CONTENTS

A world catalog of genera associated with the Glyphipterigidae auctorum (Lepi-
doptera) John B. Heppner 220-294

Studies of North American Erora (Scudder) (Lepidoptera, Lycaenidae)

Alexander B. Klots and Cyril F. dos Passos 295-331

Acknowledgment 332

Honorary, Life, and Sustaining Members 332

Index to Scientific Names of Animals and Plants for Volume LXXXIX 333-335

Index of Authors for Volume LXXXIX i-ii

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(4), 1981, pp. 220-294

A WORLD CATALOG OF GENERA ASSOCIATED
WITH THE GLYPHIPTERIGIDAE AUCTORUM
(LEPIDOPTERA)

John B. Heppner

Abstract. — The catalog lists 272 generic-group names previously associ-
ated with the family Glyphipterigidae auctorum of the world and segregates
these generic names to Glyphipterigidae sensu stricto (Copromorphoidea),
Immidae (Immoidea), and Brachodidae and Choreutidae (Sesioidea), as well
as 21 other families. Transfers to other families are based on examination
of the type-species of each genus, in some cases only on literature references
and notes from colleagues. Generic synonyms, homonyms, and type-
species, with their designations, type localities and depositories, are given
when known. Lectotypes are designated for 19 of the generic type-species
listed. A complete bibliography is given for all included taxa, as well as
an index to cited species. V. O. Becker contributes the new name,
Rectiostoma Becker, replacing Setiostoma Zeller, 1875 [not Setiostoma
Felder and Rogenhofer, 1875] in Oecophoridae (Stenomatinae).

The following catalog lists generic names previously associated with the
family Glyphipterigidae auctorum and reassigns them to appropriate fami-
lies. Included are generic names originally described in Glyphipterigidae,
names subsequently transferred to the family, and generic names at one
time or another associated with the family. The Glyphipterigidae auctorum
was a family concept largely the result of generic and family groupings by
Edward Meyrick, a British lepidopterist active from 1875 until 1938. Mey-
rick’s (1914d) concept of the family encompassed what had often been con-
sidered two families prior to this century, Glyphipterigidae and Choreuti-
dae. These two families, as well as numerous other groups, Meyrick
combined into one chaotic family based upon morphological similarities and
a general superficial similarity in wing maculation, as far as various metallic
iridescent markings are concerned. These two families have more recently
again become recognized as distinct, in fact belonging to different superfam-
ilies, Copromorphoidea for Glyphipterigidae and Sesioidea for Choreutidae
(Heppner 1977a). In addition, further segregation of some genera has been
completed to form the family Immidae (Heppner 1977a) and a revised Brach-
odidae, formerly Atychiidae (Heppner 1979a; Heppner and Duckworth
1981).

This catalog notes the current family position of 272 generic names (in-
cluding synonyms and misspellings) previously associated with the Gly-

VOLUME LXXXIX, NUMBER 4

221

phipterigidae auctorum. Many genera have been found to belong to a wide
range of families, in fact a total of 24 families other than Gly phipterigidae,
including the following:

Acrolepiidae

Immidae

Agonoxenidae

Incurvariidae

Brachodidae

Limacodidae

Choreutidae

Noctuidae

Copromorphidae

Oecophoridae

Douglasiidae

Plutellidae

Elachistidae

Psychidae

Epermeniidae

Pyralidae

Gelechiidae

Tineidae

Geometridae

Tortricidae

Gracillariidae

Y ponomeutidae

Heliodinidae

Zygaenidae

The catalog is largely the result of investigations of generic type-species
completed in conjunction with a revision of the North American Glyphip-
terigidae and Choreutidae. In beginning the generic studies it quickly be-
came apparent that a review of the world genera associated with the Gly-
phipterigidae auctorum would be needed, both to determine the actual
parameters of Glyphipterigidae sensu stricto and Choreutidae, and also to
properly define each family in relation to other families, especially the re-
cently defined families of Brachodidae and Immidae.

Although all the type-species of genera herein restricted to Glyphipterig-
idae, Brachodidae, Choreutidae, Immidae, and the tribe Hilarographini of
Tortricidae (a group especially well associated with choreutids for a long
time) were studied, not all the type-species of other genera listed have like-
wise been studied. Some type-species were not examined because in many
cases well known genera were involved and there was no further need to
verify their family placement, although at some previous time they were
associated with the Glyphipterigidae auctorum. In only a few cases were
literature references or notes of colleagues relied upon, in some cases due
to inability to locate the holotypes for genera based upon uniques.

As presently restricted, out of a total of 272 generic names listed, 42
names remain in Glyphipterigidae sensu stricto, 24 names remain in Immi-
dae, 22 names remain in Brachodidae, and 62 names remain in Choreutidae.
These names are summarized and listed briefly for each family below. The
families as now perceived contain 1,016 described species: 326 in Glyphip-
terigidae (Heppner 1982c), 238 in Immidae (Heppner 1982b), and 96 in
Brachodidae and 356 in Choreutidae (Heppner 1981a).

222

NEW YORK ENTOMOLOGICAL SOCIETY

Glyphipterigidae (Copromorphoidea)

Chrysocentris Meyrick, 1914
Irinympha Meyrick, 1932
Ernolytis Meyrick, 1922
Carmentina Meyrick, 1930
Metapodistis Meyrick, 1933
Cotaena Walker, [1865]

Myrsila Boisduval, [1875]

Lepidotarphius Pryer, 1877
Desmidoloma Erschoff, 1892
Tetracmanthes Meyrick, 1925
Phalerarcha Meyrick, 1913
Cronicombra Meyrick, 1920
Taeniostolella Fletcher, 1940
Taeniostola Meyrick, 1920, preocc.

Machlotica Meyrick, 1909
Maclotica [sic] Busck, 1915, missp.

Abrenthia Busck, 1915
Neomachlotica Heppner, 1981
Trapeziophora Walsingham, 1892
Ussara Walker, 1864
Setiostoma Felder and Rogenhofer, 1875
Usara [sic] Busck, 1933, missp.

Electrographa Meyrick, 1912
Rhabdocrates Meyrick, 1931
Apistomorpha Meyrick, 1881
Phryganostola Meyrick, 1881
Pantosperma Meyrick, 1888
Circica Meyrick, 1888
Glyphipterix Hiibner, [1825]

Heribeia Stephens, 1829
Aechmia Treitschke, 1833
Aecimia [sic] Boisduval, 1836, missp.

Glyphipteryx Zeller, 1839, emend.

Glyphiteryx [sic] Fischer von Roslerstamm, 1841, missp.
Anacampsoides Bruand, 1850, nom. oblit.

Glyphopteryx Herrich-Schaffer, 1854, emend.
Glyphiptoryx [sic] Mann and Rogenhofer, 1878
Glyphptieryx [sic] Turati, 1879, missp.

Glyphipterys [sic] Christoph, 1882, missp.

Glyphyteryx [sic] Hampson, 1918, missp.

Glyphteryx [sic] Watt, 1920, missp.

Diploschizia Heppner, 1981

VOLUME LXXXIX, NUMBER 4

223

Immidae (Immoidea)

Ptochaula Meyrick, 1920
Bursadella Snellen, 1880
Scaptesylix Hampson, 1895, n. syn.
Cercosimma Diakonoff, 1948
Alampla Diakonoff, 1978
Moca Walker, 1863
Adricara Walker, 1863, n. syn.

Alicadra Walker, [1866], n. syn.

Jobula Walker, 1866, n. syn.

Callartona Hampson, [1893], n. syn.
Bryonympha Meyrick, 1930
Sthenistis Hampson, 1896
Loxotrochis Meyrick, 1906
Birthana Walker, [1865]

Me thy ps a Butler, 1875, n. syn.

Hyperperissa Walsingham, 1900, n. syn.

Imma Walker, [1859]

Pingrassa Walker, [1859]

Tortricomorpha C. Felder, 1861
Topaza Walker, 1864
Vinzela Walker, [1866]

Thylacopleura Meyrick, 1886
Davendra Moore, 1887
Pseudotortrix Turner, 1900

Brachodidae (Sesioidea)
Brachodinae

Brachodes Guenee, 1845
Chimaera Ochsenheimer, 1808, preocc.
Atychia Latreille, 1809, preocc.
Procerata Berthold, 1827, nom. oblit.
Chimera [sic] Feisthamel, 1833, missp.
Palamernis Meyrick, 1906
Bradyptesis sensu Kasy, 1979
Miscera Walker, 1863
Euthorybeta Turner, 1913
Synechodes Turner, 1913
Atractoceros Meyrick, 1936
Callatolmis Butler, 1877
Sisyroctenis Meyrick, 1936

224

NEW YORK ENTOMOLOGICAL SOCIETY

Sagalassa Walker, 1856
Gora Walker, 1862
Jonaca Walker, 1863
Polyphlebia Felder, 1874

Phycodinae

Phycodes Guenee, 1852
Tegna Walker, 1866
Nigilgia Walker, 1863
Nigilica [sic] Turner, 1929, missp.
Hoplophractis Meyrick, 1920

Choreutidae (Sesioidea)
Brenthiinae

Litobrenthia Diakonoff, 1978
Brenthia Clemens, 1860
Microaethia Chambers, 1878

Choreutinae

Anthophila Haworth, [1811]

Simaethis Leach, 1815
Xylopoda Berthold, 1827
Xylopoda Latreille, 1829, redesc.

Simoethis [sic] Desmarest, 1848, missp.

Symaethis [sic] Bruand, 1850, missp.

Xylopodo [sic] Morris, 1872, missp.

Simaetis [sic] Kautz, 1930, missp.

Simethis [sic] Bleszynski, Razowski, and Zukowski, 1965, missp.
Antophila [sic] Bleszynski, Razowski, and Zukowski, 1965, missp.
Siamethis [sic] Klimesch, 1968, missp.

Millieria Ragonot, 1874
Ripismia Wocke, [1876]

Rhipismia Reutti, 1898, emend.

Milliereia Spuler, 1910, emend.

Milliera [sic] Le Marchand, 1937, missp.

Millieroa [sic] Le Marchand, 1937, missp.

Peotyle Diakonoff, 1978
Prochoreutis Diakonoff and Heppner, 1980
Choreutis of authors [not Hiibner, 1825]

Prochoreutis Heppner, 1981, redesc.

Caloreas Heppner, 1977
Tebenna Billberg, 1820

VOLUME LXXXIX, NUMBER 4

225

Porpe Hiibner, [1825]

Tebeuna [sic] Danilevsky, 1969, missp.

Asterivora Dugdale, 1979

Aster ophaga Horning and Greenwood, 1977, nom. nud.
Choreutis Hiibner, [1825]

Hemerophila Hiibner, [1806], rejected

Eutromula Frolich, 1828

Chore utes Treitschke, 1835, emend.

Macropia O. Costa, [1836]

Chorentes [sic] Morris, 1871, missp.

Entomoloma Ragonot, 1875
Chorentis [sic] Turner, 1898, missp.

Orchemia sensu Fernald, 1900
Hemerophila sensu Fernald, 1900
Choreutidia Sauber, 1902
Allononyma Busck, 1904
Allonyma [sic] Fracker, 1915, missp.

Cloreutis [sic] Kautz, 1930, missp.

Choreuthis [sic] Hackman, 1947, missp.

Chloreutis [sic] Viette, 1948, missp.

Allonomyia [sic] Ferguson, 1975, missp.

Saptha Walker, 1864
Badera Walker, 1866
Chordates Snellen, 1877
Choredates [sic] Pagenstecher, 1884, missp.

Saphtha [sic] Walsingham, 1900, missp.

Tortyra Walker, 1863
Choregia Felder and Rogenhofer, 1875
Choregia Zeller, 1877, redesc.

Hemerophila Hiibner, [1817]

Gauris Hiibner, 1821
Walsinghamia Riley, 1889
Guaris [sic] Fernald, 1900, missp.

Zodia Heppner, 1979
Melanoxena Dognin, 1910
Rhobonda Walker, 1863

Family Distribution of Other Genera

Incurvariidae

Tegeticula Zeller, 1873
Thia H. Edwards, 1888, preocc.

Thelethia Dyar, 1893, repl. name

226

NEW YORK ENTOMOLOGICAL SOCIETY

Tineidae

Ereunetis Meyrick, 1881

Psychidae

Aprata Moore, 1883
Cebysa Walker, 1854
Sezeris Walker, 1863
Colpotorna Meyrick, 1920
Taleporia Hiibner, [1825]

Gracillariidae

Callisto Stephens, 1834
Euprophantis Meyrick, 1921
Philodoria Walsingham, 1907

Oecophoridae

Depressariinae

Callizyga Turner, 1894

Autostichinae

Lasiodictis Meyrick, 1912

Xyloryctinae

Amphimelas Turner, 1929
Mylocera Turner, 1897
Symphorostola Meyrick, 1927

Stenomatinae

Aproopta Turner, 1919
Rectiostoma Becker, 1982, repl. name
Setio stoma Zeller, 1875, preocc.

Oecophorinae

Aeolocosma Meyrick, 1881
Coridomorpha Meyrick, 1914
Eretmocera Zeller, 1852
Staintonia Staudinger, 1859
Heliostobes Zeller, 1874
Hierodoris Meyrick, 1912
Homoplastis Meyrick, 1926
Lamprystica Meyrick, 1914
Lygronoma Meyrick, 1913

VOLUME LXXXIX, NUMBER 4

227

Pachyphoenix Butler, 1883
Tyriomorpha Meyrick, 1918
Matte a Duckworth, 1966

Hypertrophinae

Allotropha Diakonoff, 1954
Epithetica Turner, 1923
Eupselia Meyrick, 1881
Hypertropha Meyrick, 1881
Oxytropha Diakonoff, 1954
Peritropha Diakonoff, 1954
Polygiton Diakonoff, 1955

Elachistidae

Perittia Stainton, 1854

Agonoxenidae

Chrysoclista Stainton, 1854
Glyphipteryx Curtis, 1827, nom. oblit.
Heinemannia Wocke, 1853
Tebenna Hiibner, [1825], preocc.

Gelechiidae

Dichomeris Hiibner, [1825]

Rhobonda Walker, 1864, preocc.

Carna Walker, 1864, repl. name; preocc.

Copromorphidae

Lotisma Busck, 1909
Ordrupia Busck, 1911
Ardrupia [sic] Busck, 1911, missp.

Ordupia [sic] Busck, 1911, missp.

Epermeniidae

Agiton Turner, 1926
Picrodoxa Meyrick, 1923

Plutellidae

Araeolepia Walsingham, 1881
Ellabella Busck, 1925
Probolacma Meyrick, 1927
Spilogenes Meyrick, 1938

228

NEW YORK ENTOMOLOGICAL SOCIETY

Homadaula Lower, 1899
Homadaula Meyrick, 1907, redesc.
Paraprays Rebel, 1910
Stichotactis Meyrick, 1930
Napecoetes Turner, 1913
Protosynaema Meyrick, 1886

Yponomeutidae

Charixena Meyrick, 1920, repl. name
Philpottia Meyrick, 1916, preocc.
Charizena [sic] Neave, 1939, missp.
Ditrigonophora Walsingham, 1897
Embryonopsis Eaton, 1876
Iridostoma Meyrick, 1909
Niphonympha Meyrick, 1914
Calantica Zeller, 1847, preocc.

Piestoceros Meyrick, 1907
Roeslerstammia Zeller, 1839
Tanaoctena Turner, 1913
Nesotropha Turner, 1926
Cylicophora Turner, 1927

Douglasiidae

Klimeschia Amsel, 1938
Tinagma Zeller, 1839
Douglasia Stainton, 1854

Acrolepiidae

Acrolepia Curtis, 1838
Antispastis Meyrick, 1926

Heliodinidae

Actinoscelis Meyrick, 1912
Amphiclada Meyrick, 1912
Corsocasis Meyrick, 1912
Encratora Meyrick, 1923
Epicroesa Meyrick, 1907
Lithariapteryx Chambers, 1876
Philocoristis Meyrick, 1927
Sobareutis Meyrick, 1910

VOLUME LXXXIX, NUMBER 4

229

Thrasydoxa Meyrick, 1912
Thriambeutis Meyrick, 1910
Trichothyrsa Meyrick, 1912

Tortricidae

Olethreutinae

Cryptophlebia Walsingham, 1899
Thaumatotibia Zacher, 1915
Cydia Hiibner, [1825]

Ore hernia Guenee, 1845
Dudua Walker, 1864
Ganabalia Diakonoff, 1975

Chlidanotinae

Archimaga Meyrick, 1905
Charitographa Diakonoff, 1979
Embolostoma Diakonoff, 1977
Hilarographa Zeller, 1877
Idiothauma Walsingham, 1897
Irianassa Meyrick, 1905
Mictocommosis Diakonoff, 1977
Mictopsichia Hiibner, [1825]

Mictopsychia [sic] Riley, 1889, missp.
Mictropsichia [sic] Heppner, 1978, missp.
Nexosa Diakonoff, 1977
Thaumatographa Walsingham, 1897
Hilarographa sensu Meyrick, 1886
Tharmatographa [sic] Diakonoff, 1977, missp.

Zygaenidae

Adscita Retzius, 1783
Atychia Ochsenheimer, 1808
Atichia [sic] Ochsenheimer, 1808, missp.
Brady pte sis Sodoffsky, 1837
Burlacena Walker, [1865]

Sesiomorpha Snellen, 1885
Cibdeloses Durrant, 1919

Limacodidae

Penthocrates Meyrick, 1934

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Pyralidae

Heliothela Guenee, 1854
Orosana Walker, 1863
Orosona [sic] Cotes, 1889, missp.

Geometridae

Menophra Moore, 1887
Hemerophila Stephens, 1829, preocc.

Noctuidae

Chalenata Walker, 1864
Porphyrinia Hiibner, [1821]

Anthophilae Hiibner, [1806], rejected
Antophila Hiibner, [1806], rejected
Anthophila Ochsenheimer, 1816, preocc.
Antophyla [sic] Duponchel, 1829, missp.
Anthrophila [sic] Treitschke, 1832, missp.
Heliomanes Sodoffsky, 1837
Thopelia Nye, 1975, repl. name
Plotheia Walker, 1863, preocc.

Discussion of Selected Genera

1. Agiton Turner, 1926: newly transferred to Epermeniidae. The mor-
phological characters of this genus are amenable to the family Eper-
meniidae, namely a naked haustellum, absence of ocelli and chaetose-
mata, small maxillary palpi, smooth head vestiture, upcurved labial
palpi with an elongated middle segment (typical for the family), wing
venational characters, and bristles on the hind tibiae (typical for the
family). The abdominal articulation is not typical for Copromorphoidea,
as it appears rather of the tortricoid type, however, there are elongated
ventral sclerotized rods on the anterior sternite but these do not actually
reach the apodemes as is normal in the tineoid form in Copromorphoi-
dea.

2. AUotropha Diakonoff, 1954: newly transferred to Oecophoridae by re-
lational characters conforming to other genera of the Hypertrophinae.

3. Antispastis Meyrick, 1926: newly transferred to Acrolepiidae due to
genital and wing venational characters typical for this family, e.g., the
long male genital saccus and the enlarged aedeagus, both unique to
Acrolepiidae among the Yponomeutoidea.

4. Aprata Moore, 1883: this genus conforms to Psychidae, as Moore orig-
inally noted.

5. Araeolepia Walsingham, 1881: transferred to Plutellidae by Heppner

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231

(1978a) by virtue of head morphology, among other characters, con-
forming to this family.

6. Burlacena Walker, [1865]: transferred to Zygaenidae. This genus is a
typical tropical zygaenid and why it was placed in Glyphipterigidae is
difficult to comprehend. There is no relationship to Yponomeutidae as
alluded to by Common (1970b).

7. Cebysa Walker, 1854: transferred to Psychidae by Common (1970b).
This genus conforms to other tropical psychids, although it is unusual.

8. Cercosimma Diakonoff, 1948: tentatively transferred to Immidae. The
described characters appear to conform to the family definition of Im-
midae (Heppner, 1977a, 1982b) but the unique holotype (supposedly in
the RMNL collection) has not been located.

9. Charixena Meyrick, 1920: newly transferred to Plutellidae. Meyrick’s
original description of Philpottia Meyrick (1916), a homonym replaced
by Charixena , is quite detailed and notes the reduced labial palpi and
minute haustellum, among other characters. There may be some rela-
tionship between this genus and such genera as the Japanese Rhabdo-
cosma and Bhadorcosma but Charixena is otherwise quite isolated in
the family.

10. Cibdeloses Durrant, 1919: transferred to Zygaenidae. Like Burlacena,
this is a typical tropical zygaenid.

11. Colpotorna Meyrick, 1920: newly transferred to Psychidae. This genus
was originally allied to Cebysa Walker, now in Psychidae, and its char-
acters appear to conform to this group of unusual Australian psychids.

12. Coridomorpha Meyrick, 1914: newly transferred to Oecophoridae by
virtue of a scaled haustellum and long upcurved labial palpi, among
other characters.

13. Cotaena Walker, [1865]: transferred to Glyphipterigidae by Heppner
(1981a). This genus includes species subequal to the largest known gly-
phipterigids and otherwise conforms to certain tropical members of the
family. It has no relation to Heliodinidae, as thought by Naumann
(1971).

14. Ditrigonophora Walsingham, 1897: tentatively transferred to Ypono-
meutidae. The two known specimens of the type-species both lack
heads and abdomens. However, these parts were apparently intact
when Walsingham described the species and from his description it
appears that the moths may have some affinity to yponomeutids like
Xyrosaris, which also have the unusual distally enlarged labial palpi
noted by Walsingham.

15. Ellabella Busck, 1925: transferred to Plutellidae by Heppner (1978a).
This genus has no relation to Glyphipterigidae and is in fact a senior
synonym of Spilogenes Meyrick, 1938, both conforming to Plutellidae.

16. Encratora Meyrick, 1923: newly transferred to Heliodinidae. The short
porrect labial palpi and naked haustellum are typical of Heliodinidae.

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17. Epicroesa Meyrick, 1907: newly transferred to Heliodinidae. The genus
has labial palpi and other head morphology typical of tropical Heliodi-
nidae.

18. Epithetica Turner, 1923: newly transferred to Oecophoridae. This genus
is a member of the Hypertrophinae (Common, pers. comm.).

19. Heliostibes Zeller, 1874: newly transferred to Oecophoridae. This genus
is typical for Oecophoridae and why it was associated with Glyphipte-
rigidae by Meyrick (1914d) is difficult to understand.

20. Hierodoris Meyrick, 1912: newly transferred to Oecophoridae. This ge-
nus also is a typical oecophorid among a group of New Zealand genera
related to Heliostibes.

21. Hilarographa Zeller, 1877: transferred to Tortricidae by Diakonoff
(1977a) and Heppner (1977a). As noted by Heppner (1978a), the naked
haustellum and unusual genitalia relate this genus and its relatives to
other Chlidanotinae.

22. Homadaula Lower, 1899: transferred to Plutellidae by Friese (1962).
There is no question that this genus is not a choreutid by virtue of a
naked haustellum and tineoid abdominal articulation, the latter placing
the genus outside of Sesioidea. It also is not related to Brachodidae by
virtue of this same articulation. There also are no characters to support
a placement in Copromorphoidea, or Glyphipterigidae in particular, but
rather a placement among some other tropical plutellids is amenable to
the characters of the genus. Homadaula is an unusual tropical plutellid,
although now introduced into some temperate regions, and appears re-
lated only to Prays to any degree. It may require a separate subfamily
but since the tropical Yponomeutoidea as a whole are so poorly known,
considerable research is yet required to determine its true nearest rel-
atives.

23. Homoplastis Meyrick, 1926: newly transferred to Oecophoridae. The
unique holotype has not been located. Meyrick’ s notation that the genus
appears related to Eupselia prompts a transfer to Oecophoridae pending
further study.

24. Irianassa Meyrick, 1905: newly transferred to Tortricidae. This genus
conforms to characters of other Hilarographini (Chlidanotinae).

25. Iridostoma Meyrick, 1909: transferred to Yponomeutidae by Heppner
(1981a). This genus has a naked haustellum and other characters placing
it in Yponomeutidae.

26. Lamprystica Meyrick, 1914: newly transferred to Oecophoridae. The
scaled haustellum and long labial palpi are typical of Oecophoridae and
conform, together with other characters, to the Stathmopodini.

27. Lotisma Busck, 1909: transferred to Copromorphidae by Heppner
(1978a). This genus has an array of characters, notably genital features
and wing venation, that align it with other copromorphids.

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233

28. Loxotrochis Meyrick, 1906: newly transferred to Immidae. This genus
appears to conform to characters of Immidae but should be studied
further whenever it is rediscovered (indeed the type locality should be
reconfirmed as well).

29. Lygronoma Meyrick, 1913: transferred to Oecophoridae by Heppner
(1981a). This genus has a scaled haustellum and shows other charac-
ters placing it in Oecophorinae (Becker, pers. comm.).

30. Melanoxena Dognin, 1910: transferred to Choreutidae by Heppner
(1981a). This genus has a scaled haustellum and wing venation similar
to other choreutids and appears very near to Rhobonda.

31. Mictopsichia Hiibner, [1825]: transferred to Tortricidae by Diakonoff
(1977a) and Heppner (1977a). This genus appears to be related to other
Hilarographini but needs further study to verify a possible relationship
to Archipini (Tortricinae).

32. Myrsila Boisduval, [1875]: transferred to Glyphipterigidae by Heppner
(1981a). This genus has no relationship to Sesiidae, (Eichlin, pers.
comm.) as originally thought by Boisduval ([1875]), but conforms to
certain tropical glyphipterigids, notably Cotaena.

33. Napecoetes Turner, 1913: newly transferred to Plutellidae. This genus
appears to conform to other plutellids, perhaps nearest Plutella (Com-
mon, pers. comm.), but is still known only from the unique holotype.

34. Nesotropha Turner, 1926: newly transferred to Yponomeutidae as a
junior synonym of Tanaoctena Turner, 1913, as confirmed by Common
(pers. comm.).

35. Pachyphoenix Butler, 1883: newly transferred to Oecophoridae. This
genus is the senior synonym of Tyriomorpha Meyrick, 1918, and Mat-
tea Duckworth, 1966, and conforms to characters for Oecophorinae
(Becker, pers. comm.).

36. Penthocrates Meyrick, 1934: newly transferred to Limacodidae. Al-
though transferred to the Zygaenoidea by Meyrick (1934) as a member
of the Heterogynidae, the genus appears more typical of the Limacod-
idae.

37. Philocoristis Meyrick, 1927: newly transferred to Heliodinidae. The ge-
nus exhibits the short porrect labial palpi and other characters typical
of tropical Heliodinidae. It may be related to Epicroesa.

38. Philpottia Meyrick, 1916: newly transferred to Plutellidae by synonymy
with Charixena Meyrick, 1920.

39. Picrodoxa Meyrick, 1923: newly transferred to Epermeniidae. This ge-
nus has characters similar to those noted for Agiton and may in fact be
relatively closely related.

40. Piestoceros Meyrick, 1907: newly transferred to Yponomeutidae. The
genus conforms to characters of Yponomeutidae, notably such head
characters as the labial palpi. The only known species, Piestoceros

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conjunctella (Walker), also makes a rolled-leaf pupal case extended on
a filament similar to that of Urodus species of the western hemisphere,
although in Urodus the case is filigreed.

41. Rhobonda Walker, 1863: transferred to Choreutidae. This genus con-
forms to characters of Choreutidae and is related to Melanoxena Dog-
nin.

42. Sesiomorpha Snellen, 1885: transferred to Zygaenidae by Heppner
(1981a). This genus is a junior synonym of Burlacena Walker, [1865],
and is typical of tropical zygaenids.

43. Setiostoma Felder and Rogenhofer, 1875: newly transferred to Gly-
phipterigidae. The only included species in the original description, Se-
tiostoma flaviceps Felder and Rogenhofer, is clearly a member of the
glyphipterigid genus Ussara Walker, 1864, precluding the use of Setio-
stoma Zeller, 1875 (described a few months after Felder and Ro-
genhofer) in the Stenomatinae (Oecophoridae).

44. Sezeris Walker, 1863: transferred to Psychidae by virtue of synonymy
with Cebysa Walker, 1854.

45. Spilogenes Meyrick, 1938: transferred to Plutellidae by virtue of syn-
onymy with Ellabella Busck, 1925.

46. Stichotactis Meyrick, 1930: transferred to Plutellidae by virtue of syn-
onymy with Homadaula Lower, 1899.

47. Symphorostola Meyrick, 1927: transferred to Oecophoridae (Heppner,
1981a). Morphological characters, including the unusual genitalia and
a scaled haustellum, conform to Xyloryctinae of Oecophoridae.

48. Thaumatotibia Zacher, 1915: transferred to Tortricidae by Heppner,
(1980). This long forgotten genus is a synonym of Cryptophlebia Wal-
singham, 1899, as far as can be determined from the original description
of Thaumatotibia. The unique holotype has not been located.

Acknowledgments

This catalog of generic names is an outcome of research originally begun
for a revision of North American Glyphipterigidae and, as such, was in part
supported by a Smithsonian Fellowship (1976-77), by a National Science
Foundation grant (76-12550 DEB) and by the University of Florida, De-
partment of Entomology and Nematology (IFAS), Gainesville, Florida, for
research at the British Museum (Natural History), London, during 1976-77
to study holotypes. A number of persons helped with specific information
on some of the genera or gave various advice, including V. O. Becker,
Brasilia, Brazil (currently at the British Museum (Natural History), London,
England); I. F. B. Common, Division of Entomology (CSIRO), Canberra,
Australia; J. S. Dugdale, Division of Entomology (DSIR), Auckland,
New Zealand; T. D. Eichlin, California Dept, of Food and Agriculture,
Sacramento, California; J. A. Powell, University of California, Berkeley,
California; D. S. Fletcher and K. Sattler, British Museum (Natural His-

VOLUME LXXXIX, NUMBER 4

235

tory), London; and J. F. G. Clarke, R. W. Hodges (USD A), and D. R.
Davis, of the Smithsonian Institution, Washington, D.C. I also wish to
especially acknowledge the numerous discussions and correspondence
provided by A. Diakonoff, Rijksmuseum van Natuurlijke Historie, Leiden,
Netherlands, on various aspects of this project over the last 6 years.

Catalog of Glyphipterigidae Auctorum Genera

The listing of each genus follows the sequence of genus name, author,
date and page citation, and present family position, followed by the type-
species and citation, mode of selection (subsequent designations show au-
thor, date, and citation), type locality, form of type (lectotypes have des-
ignator and year in parentheses, with citation), and present location of the
type specimen. Below this data it is noted in which family the genus was
originally described, when it was associated with the Glyphipterigidae auc-
torum (families are spelled as each author used them), and when and by
whom it was first transferred to its present family position if in a family
other than Glyphipterigidae. The original spellings are used: Meyrick and
predecessors spelled Glyphipterigidae as Glyphipterygidae, since they used
the emended spelling of Glyphipteryx Zeller, rather than the original Gly-
phipterix Hiibner. Lectotypes indicated as by present designation have the
actual designation data in the section on new lectotypes following the cat-
alog.

Transfers of genera already published have no symbol by the family name
at the right margin. New generic transfers are indicated by an asterisk (*)
before the family name. Current generic names are in bold letters; synonyms
or unavailable names are in italics. The currently accepted dates of various
older works are used (see Nye 1975; Heppner 1982a).

Collections containing types are abbreviated as follows:

AMS Australian Museum, Sydney, Australia
ANIC Australian National Insect Collection, Canberra, Australia
ANSP Academy of Natural Sciences, Philadelphia, Pennsylvania, USA
BMNH British Museum (Natural History), London, England
CAS California Academy of Sciences, San Francisco, California, USA
DEI Institut fur Pflanzenschutzforschung Kleinmachnow, Eberswalde,
DDR [formerly Deutsches Entomologisches Institut]

INHS Illinois Natural History Survey, Urbana, Illinois, USA
MCZ Museum of Comparative Zoology, Cambridge, Massachusetts,
USA

MNHP Museum National d’Histoire Naturelle, Paris, France
NHMV Naturhistorisches Museum, Vienna, Austria
RMNL Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands
SAM A South Australian Museum, Adelaide, Australia
SMF Senckenburg Museum, Frankfurt, West Germany

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SMS Sarawak Museum, Kuching Sarawak, Malaysia
SMW Staatliches Museum, Wiesbaden, West Germany
UMB Ubersee Museum, Bremen, West Germany
UMO University Museum, Oxford, England

USNM National Museum of Natural History, Smithsonian Institution,
Washington, D.C., USA

ZMC Zoological Museum, Copenhagen, Denmark

ZMHB Zoological Museum, Humboldt University, Berlin, DDR

Catalog of Genera Associated with Glyphipterigidae

Abrenthia Busck, 1915:87 Glyphipterigidae

Type-species: Abrenthia cuprea Busck, 1915:87, by monotypy. Type lo-
cality: USA: Roxboro, Pennsylvania [lectotype 6 (present designation),
USNM].

Originally described in Glyphipterygidae [sic].

Acrolepia Curtis, 1838:679 Acrolepiidae

Type-species: Acrolepia autumnitella Curtis, 1838:679 [=Tortrix pyg-
meana Haworth, [181 1] :439] , by original designation. Type locality ( autum-
nitella): [England] [lectotype, BMNH?].

Originally described without family reference; subsequently transferred
to Glyphipterygidae [sic] by Stainton (1854:169); transferred to Tineidae by
Meyrick (1895:771); transferred to Acrolepiidae by Spuler (1910:453).

Actinoscelis Meyrick, 1912a:59 Heliodinidae

Type-species: Actinoscelis irina Meyrick, 1912a:59, by monotypy. Type
locality: India: Bombay, Kanara [holotype 6 , BMNH].

Originally described in Glyphipterygidae [sic]; subsequently transferred
to Heliodinidae by Meyrick (1913b: 19).

Adricara Walker, 1863:114 *Immidae

Type-species: Adricara albodiscata Walker, 1863:115, by monotypy.
Type locality: Brazil, Ega, [=Tefe], Amazonas [holotype 9 , BMNH].

Originally described in Galleridae [=Pyralidae]; subsequently transferred
to Glyphipterygidae [sic] by Meyrick (1913b: 25), as a synonym of Imma
Walker, [1859]; hereby transferred to Immidae as a junior subjective syn-
onym of Moca Walker, 1863, new synonymy.

Aechmia Treitschke, 1833:69 Glyphipterigidae

Type-species: Phalaena ( Tortrix ) fyeslella [sic] Fabricius, 1798:493
[ =Phalaena {Tortrix) fueslella Fabricius, 1781:301; = Phalaena thrasonella
Scopoli, 1763:253], by subsequent designation of Westwood (1840:112).
Type locality {fueslella ): Germany: Kiel [lectotype 6 (Diakonoff,
1977c: 176), ZMC].

Originally described among the Tineina; subsequently transferred to Gly-

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237

phipterygidae [sic] by Stainton (1854:173), as a synonym of Glyphipteryx
[sic] auctorum [ =Glyphipterix Hiibner, 1825].

Aechmia sensu Zeller, 1847:881, included only Tinagma metallicella Du-
ponchel, 1840, now in Heliozela Herrich-Schaffer, 1853 (Heliozelidae).

Aechmia sensu Stainton, 1854:176, included only Aechmia dentella Zell-
er, 1839, now in Phaulernis Meyrick, 1895 (Epermeniidae).

Aecimia [sic] Boisduval, 1836:138, misspelling Glyphipterigidae

A misspelling of Aechmia Treitschke, 1833, now a synonym of Glyphip-
terix Hiibner, [1825].

Aeolocosma Meyrick, 1881:224 Oecophoridae

Type-species: Aeolocosma iridozona Meyrick, 1881:225, by subsequent
designation of Meyrick (1922b: 101). Type locality: Australia: Sydney, New
South Wales [lectotype 8 (Diakonoff, 1954:694), BMNH].

Originally described in Glyphipterygidae [sic]; subsequently transferred
to Oecophoridae (Oecophorinae) by Meyrick (1922b: 101).

Agiton Turner, 1926a: 145 * Epermeniidae

Type-species: Agiton idioptila Turner, 1926a: 145, by monotypy. Type
locality: Australia: [Lamington] National Park, Queensland [lectotype
8 (present designation), ANIC].

Originally described in Glyphipterygidae [sic]; subsequently transferred
to Heliodinidae sensu Turner [=Yponomeutoidea] by Turner (1941:24);
hereby transferred to Epermeniidae.

Alampla Diakonoff, 1978:36 Immidae

Type-species: Imma palaeodes Meyrick, 1914a:57, by original designa-
tion. Type locality: Taiwan: Shuisharyo [lectotype unselected, BMNH].
Originally described in Immidae.

Alicadra Walker, [1866]: 1192 *Immidae

Type-species: Alicadra vexatalis Walker, [1866]: 1192, by monotypy.
Type locality: Brazil [holotype 9 , UMO].

Originally described in Herminidae [=Noctuidae]; subsequently trans-
ferred to Glyphipterygidae [sic] as a synonym of Imma Walker, [1859], by
Meyrick, (1913b:25); hereby transferred to Immidae as a junior subjective
synonym of Moca Walker, 1863, new synonymy.

Allonomyia [sic] Ferguson, 1975:41, misspelling Choreutidae

A misspelling of Allononyma Busck, 1904, now a synonym of Choreutis
Hiibner, [1825].

Allononyma Busck, 1904:745 Choreutidae

Type-species: Tortrix diana Hiibner, [1819-22]: pi. 44, fig. 274, by mono-
typy. Type locality: Europe [type lost?].

Originally described in Yponomeutidae; subsequently transferred to Gly-

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phipterygidae [sic] by Meyrick (1913b:34); transferred to Choreutidae as a
junior subjective synonym of Eutromula Frolich, 1828, by Bradley
(1972:12); now a junior subjective synonym of Choreutis Hiibner, [1825]
(Heppner, 1981a:53).

Allonyma [sic] Fracker, 1915:77, misspelling Choreutidae

A misspelling of Allononyma Busck, 1904, now a synonym of Choreutis
Hiibner, [1825].

Allotropha Diakonoff, 1954:688 * Oecophoridae

Type-species: Orosana percussana Walker, 1864:998, by original desig-
nation. Type locality: Australia: Tasmania [holotype 6 , BMNH].

Originally described in Hypertrophinae of Glyphipterygidae [sic]; hereby
transferred to Oecophoridae: Hypertrophinae.

Amphiclada Meyrick, 1912a:60 Heliodinidae

Type-species: Amphiclada fervescens Meyrick, 1912a:60, by monotypy.
Type locality: Grenada: St. George’s [holotype 9, BMNH].

Originally described in Glyphipterygidae [sic]; subsequently transferred
to Heliodinidae by Meyrick (1913b: 18).

Amphimelas Turner, 1929:306 Oecophoridae

Type-species: Amphimelas argopasta Turner, 1929:307 [=Chereuta tin-
thalea Meyrick, 1906a:33], by monotypy. Type locality ( argopasta ): Aus-
tralia: New South Wales [lectotype unselected, ANIC?].

Originally described in Glyphipterygidae [sic]; Amphimelas Turner, 1929,
is now a junior subjective synonym of Chereuta Meyrick, 1906, in Oeco-
phoridae: Xyloryctinae.

Anacampsoides Bruand, 1850b:32 Glyphipterigidae

Type-species: Heribeia simpliciella Stephens, 1834:263, by monotypy.
Type locality: England [lectotype 6 (Diakonoff, 1977c: 174), BMNH].

Originally listed in a catalog under Tineidae; subsequently a nomen ob-
litum and a junior subjective synonym of Glyphipterix Hiibner, [1825], in
Glyphipterigidae .

Anthophila Haworth , [ 1 8 1 1 ] : 47 1 Choreutidae

Type-species: Anthophila fabricii Haworth, [1811]: 47 1 , emendation [ =
Phalaena ( Tortrix ) fabriciana Linnaeus, 1767:880], by subsequent desig-
nation of Fletcher (1929:16). Type locality {fabriciana ): Europe [type, Lin-
nean Society?, London].

Originally described without family placement; subsequently transferred
to Choreutidae by Stainton (1859:158), as a synonym of Simaethis Leach,
1815; now considered a valid genus in Choreutidae.

There has been some reference in the literature to a homonym of Antho-
phila Haworth, other than Anthophila Ochsenheimer, but this refers to the
genus Anthophilus in Hymenoptera which is spelled differently.

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239

Anthophila Ochsenheimer, 1816:93 Noctuidae

Type-species: Noctua purpurina [Denis and Schiffermiiller], 1775:88, by
subsequent designation of Duponchel (1829:72). Type locality: Austria: Vi-
enna [type lost?].

Preoccuppied by Anthophila Haworth, [1811] (Choreutidae); currently
considered to be a junior objective synonym of Porphyrinia Hiibner, [1821],
in Noctuidae.

Anthophilae Hiibner, [1806]: [2] Noctuidae

Rejected for nomenclatural purposes by Opinion 97 (ICZN, 1926). Noctua
purpurina [Denis and Schiffermiiller], 1775:88, was the only included
species, now in Porphyrinia Hiibner, [1821] (Noctuidae).

Anthrophila [sic] Treitschke, 1832:134 Noctuidae

A misspelling of Anthophila Ochsenheimer, 1816 [not Anthophila Ha-
worth, 1811], now a synonym of Porphyrinia Hiibner, [1821], in Noctuidae.

Antispastis Meyrick, 1926b: 307 *Acrolepiidae

Type-species: Antispastis xylophragma Meyrick, 1926b:307, by mono-
typy. Type locality: Peru: Cocapata [holotype 8 , BMNH].

Originally described in Glyphipterygidae [sic]; hereby transferred to Ac-
rolepiidae.

Antophila Hiibner, [1806]: [2] Noctuidae

An original multiple spelling of Anthophilae Hiibner, [1806]: [2], published
in a work rejected for nomenclatural purposes by Opinion 97 (ICZN, 1926);
now a synonym (although unavailable) of Porphyrinia Hiibner, [1821], in
Noctuidae.

Antophila [sic] Bleszynski, Razowski, Choreutidae

and Zukowski, 1965:413

A misspelling of Anthophila Haworth, [1811] [not Anthophila Ochsen-
heimer, 1816], now in Choreutidae.

Antophyla [sic] Duponchel, 1829:72 Noctuidae

A misspelling of Anthophila Ochsenheimer, 1816 [not Anthophila Ha-
worth, 1811], now synonym of Porphyrinia Hiibner, [1821], in Noctuidae.

Apistomorpha Meyrick, 1881:247 Glyphipterigidae

Type-species: Apistomorpha argyrosema Meyrick, 1881:247 by mono-
typy. Type locality: Australia: New South Wales [lectotype unselected,
BMNH].

Originally described in Glyphipterygidae [sic] ; considered a junior sub-
jective synonym of Glyphipterix Hiibner, [1825], by Meyrick (1913b:41);
here considered a distinct genus.

Aprata Moore, 1883:106 Psychidae

Type-species: Aprata mackwoodii Moore, 1883:106, by subsequent des-

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NEW YORK ENTOMOLOGICAL SOCIETY

ignation of Walsingham and Durrant, 1900b:582. Type locality: Ceylon
[= Sri Lanka] [holotype 6 , BMNH].

Originally described in Psychidae; subsequently transferred to Glyphip-
terygidae [sic] by Meyrick (19 13b: 32); hereby returned to Psychidae.

Aproopta Turner, 1919:171 Oecophoridae

Type-species: Aproopta melanchlaena Turner, 1919:172, by monotypy.
Type locality: Australia: Katoomba, New South Wales [holotype <$ , AMS].

Originally described in Gelechianae [=Gelechiidae]; subsequently includ-
ed in Glyphipterygidae [sic] by Fletcher (1929:20); transferred to Stenomi-
dae by Sattler (1973:171); now in Oecophoridae: Stenomatinae.

Araeolepia Walsingham, 1881:303 Plutellidae

Type-species: Araeolepia subfasciella Walsingham, 1881:303, by mono-
typy. Type locality: USA: Currant Creek, Grant Co., Oregon [lectotype
6 (present desgination), BMNH].

Originally described in Tineidae; subsequently transferred to Glyphipte-
rygidae [sic] by Busck (1925:46); transferred to Plutellidae by Heppner
(1978a:51).

Archimaga Meyrick, 1905:609 Tortricidae

Type-species: Archimaga pyractis Meyrick, 1905:609, by monotypy.
Type locality: Ceylon [=Sri Lanka]: Maskeliya [lectotype 6 (Clarke,
1963:71), BMNH].

Originally described in Plutellidae and noted to be “allied to Hilarogra-
pha”\ subsequently transferred to Chlidanotidae by Meyrick (1906b:411),
now a subfamily of Tortricidae.

Ardrupia [sic] Busck, 1911:228 Copromorphidae

One of two original spellings of Ordrupia Busck, 1911, corrected by
Busck (1912:8); now in Copromorphidae.

Asterivora Dugdale, 1979:461 Choreutidae

Type-species: Simaethis comhinatana Walker, 1863:456, by original des-
ignation. Type locality: New Zealand [lectotype unselected, BMNH].
Originally described in Choreutidae.

Asterophaga Horning and Greenwood, 1977:295 Choreutidae

A nomen nudum without family reference; now a synonym of Asterivora
Dugdale, 1979, in Choreutidae.

Atichia [sic] Ochsenheimer, 1808:11 Zygaenidae

A misspelling of Atychia Ochsenheimer, 1808:10, now a synonym of Ad-
scita Retzius, 1783, in Zygaenidae.

Atractoceros Meyrick, 1936a:40 Brachodidae

Type-species: Phy codes xanthoprocta Meyrick, 1914c:283, by original

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241

designation. Type locality: Nyasaland [= Malawi]: Mt. Mlanje [lectotype
unselected, BMNH].

Originally described in Glyphipterygidae [sic]; subsequently transferred
to Brachodidae by Heppner (1979a: 127).

Atychia Ochsenheimer, 1808:10 Zygaenidae

Type-species: Sphinx statices Linnaeus, 1758:495, by subsequent desig-
nation of Tremewan (1973:119). Type locality: Europe [type, Linnean So-
ciety, London].

Originally described in Sphinges; currently a junior synonym of Adscita
Retzius, 1783, in Zygaenidae.

Atychia Latreille, 1809:214 Brachodidae

Type-species: Sphinx chimaera Hiibner, 1796:pl. 1, fig. 1 [=Sphinx ap-
pendiculata Esper, 1783:227], by subsequent designation of Latreille
(1810:441). Type locality {chimaera): Europe [type lost?]; {appendicula-
ta ): Europe [lectotype 9 (present designation), SMW].

Originally described without family placement; subsequently transferred
to Atychides by Duponchel (1835:169); transferred to Tineidae by Staudin-
ger (1870:229); transferred to Plutellidae by Meyrick (1906c: 169); transferred
to Glyphipterygidae [sic] by Meyrick (19 13b: 29); transferred to Brachodidae
(formerly Atychiidae) by Heppner (1979a: 127), as a junior subjective syn-
onym of Brachodes Guenee, 1845.

Atychia Latreille, 1809, is a junior homonym of Atychia Ochsenheimer,
1808 (Zygaenidae).

Badera Walker, 1866:1819 Choreutidae

Type-species: Badera pretiosa Walker, 1866:1819, by subsequent desig-
nation of Meyrick (1914d:18). Type locality: [Indonesia]: Java [lectotype
8 (present designation), BMNH].

Originally described in Tineidae; subsequently transferred to Plutellidae
by Meyrick (1907:97); transferred to Glyphipterygidae [sic] Meyrick
(1913b:33) as a synonym of Tortyra Walker, 1863; transferred to Choreu-
tidae as a junior subjective synonym of Saptha Walker, 1864, by Heppner
(1981a:55).

Birthana Walker, [1865]: 145 *Immidae

Type-species: Birthana consocia Walker, [1865]: 145, by monotypy. Type
locality: India: Mysol [holotype 8 , UMO].

Originally described in Melameridae [=Noctuidae]; subsequently trans-
ferred to Glyphipterygidae [sic] as a synonym of Imma Walker, [1859], by
Meyrick (19 13b: 25); hereby transferred to Immidae as a distinct genus.

Brachodes Guenee, 1845:311 Brachodidae

Type-species: Brachodes vernetella Guenee, 1845:311 [=Chimera [sic]

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NEW YORK ENTOMOLOGICAL SOCIETY

funebris Feisthamel, 1833:259], by monotypy. Type locality ( vernetella ):
Persia [type, MNHP?].

Originally described in a catalog under Tineae; subsequently transferred
to Plutellidae by Mevrick (1906c: 169); transferred to Glyphipterygidae [sic]
as a synonym of Atychia Latreille, 1809, by Meyrick (1913b:29), and as
a distinct genus in Brachodinae by Agenjo (1966: [148]); transferred to
Brachodidae by Heppner (1979a: 127).

Bradyptesis Sodoffsky, 1837:83 Zygaenidae

An unjustified replacement name for Atychia Ochsenheimer, 1808 [not
Atychia Latreille, 1809], now a junior objective synonym of Adscita Ret-
zius, 1783, in Zygaenidae.

Bradyptesis sensu Kasy, 1979:5 Brachodidae

An incorrect use of Bradyptesis as a replacement name for Atychia La-
treille, 1809, now in Brachodidae as ajunior synonym of Brachodes Guenee.

Brenthia Clemens, 1860:172 Choreutidae

Type-species: Brenthia pavonacella Clemens, 1860:172, by monotypy.
Type locality: USA: [Pennsylvania] [holotype 9, ANSP].

Originally described in Tineina; subsequently transferred to Choreutina
by Zeller (1875:323); transferred to Plutellidae by Meyrick (1907:108); trans-
ferred to Choreutidae by Heppner (1977b: 633).

Bryonympha Meyrick, 1930a:560 *Immidae

Type-species: Bryonympha silvana Meyrick, 1930a:560, by monotypy.
Type locality: Comoro Is.: Grand Comoro Id. [holotype 9 , BMNH].

Originally described in Xyloryctidae; subsequently transferred to Gly-
phipterygidae [sic] by Viette (1954:13); hereby transferred to Immidae.

Burlacena Walker, [1865]: 80 Zygaenidae

Type-species: Burlacena aegerioides Walker, [1865]:80, by subsequent
designation of Meyrick (1914d:7). Type locality: New Guinea [holotype
6 , BMNH].

Originally described in Zygenidae [sic]; subsequently transferred to Gly-
phipterygidae [sic] by Meyrick (1913b:25); transferred to Yponomeutidae
by Common (1970b:813); hereby returned to Zygaenidae.

Bursadella Snellen, 1880:83 *Immidae

Type-species: Bursadella dichroalis Snellen, 1880:83, by subsequent des-
ignation of Meyrick (1906c: 170). Type locality: [Indonesia]: Silago, Sumatra
[holotype 6 , RMNL].

Originally described in Tineina; subsequently transferred as a synonym
of Imma Walker, [1859], in Plutellidae by Meyrick (1906c: 170), and in Gly-
phipterygidae [sic] by Meyrick (1913b:25); hereby transferred to Immidae
as a distinct genus.

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243

Calantica Zeller, 1847:811 Yponomeutidae

Type-species: Calantica albella Zeller, 1847:811 [ =Calantica dealbatella
Zeller, 1847:811], by subsequent designation of Friese (1960:38). Type lo-
cality {albella): Germany: Taunus [type?].

Originally described in Tineides; subsequently transferred to Hypono-
meutidae [sic] by Heinemann (1870:101); transferred to Hemerophilidae by
Walsingham (1914:319); transferred to Yponomeutidae as a junior synonym
of Niphonympha Meyrick, 1914, by Friese (1960:38).

Calantica Zeller, 1847, is a junior homonym of Calantica Gray, 1825
(Crustacea).

Callartona Hampson, [1893]:233 *Immidae

Type-species: Callartona purpurascens Hampson, [1893] :233, by original
designation. Type locality: India: Nilgiris [holotype 9, BMNH].

Originally described in Zygaenidae; subsequently transferred as a syn-
onym of Imma Walker, [1859], to Plutellidae by Meyrick (1906c: 170), and
to Glyphipterygidae [sic] by Meyrick (1913b:25); hereby transferred to Im-
midae as a junior subjective synonym of Moca Walker, 1863, new synonymy.

Callisto Stephens, 1834:276 Gracillariidae

Type-species: Gracillaria guttea Haworth, 1828:531 [ =Tinea denticulella
Thunberg, 1794:97], by subsequent designation of Bradley (1966a: 131). Type
locality {guttea): England [lectotype 8 (Bradley, 1966a: 131), UMO].

Originally described in Yponomeutidae; subsequently transferred to Gly-
phipterygidae [sic] as a synonym of Glyphipteryx [sic] Hiibner, [1825], by
Stainton (1854:173); now a distinct genus in Gracillariidae.

Callatolmis Butler, 1877:348 Brachodidae

Type-species: Lycomorpha coleoptrata Walker, 1854:288, by original
designation. Type locality: Brazil: Tapayos [holotype 8 , BMNH].

Originally described in Lithosiidae [=Arctiidae]; subsequently transferred
to Glyphipterygidae [sic] as a junior synonym of Sagalassa Walker, 1856,
by Meyrick (1913b:31); transferred to Brachodidae as a distinct genus by
Heppner (1979a: 127).

Callizyga Turner, 1894:132 Oecophoridae

Type-species: Callizyga dispar Turner, 1894:132, by monotypy. Type lo-
cality: Australia: Brisbane, Queensland [lectotype 8 (present designation),
ANIC].

Originally described in Oecophoridae; subsequently transferred to Hy-
ponomeutinae [sic] in Plutellidae together with Imma Walker, [1859], by
Turner (1913:208); transferred to Oecophoridae by Meyrick (1922b: 137);
now in Oecophoridae: Depressariinae (Amphisbatini).

Caloreas Heppner, 1977b:631 Choreutidae

Type-species: Choreutis apocynoglossa Heppner, 1976:256, by original

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NEW YORK ENTOMOLOGICAL SOCIETY

designation. Type locality: USA: Del Valle Lake, Alameda Co., California
[holotype 8 , CAS].

Originally described in Choreutidae.

Carmentina Meyrick, 1930b: 597 Glyphipterigidae

Type-species: Carmentina iridesma Meyrick, 1930b:598, by monotypy.
Type locality: Solomon Is.: Bougainville [holotype 9, BMNH].

Originally described in Yponomeutidae; subsequently transferred to
Glyphipterigidae as the senior subjective synonym of Metapodistis Meyrick,
1933, by Heppner (1982c).

Carna Walker, 1864:1038 Gelechiidae

Type-species: Rhobonda punctatella Walker, 1864:802, by monotypy of
Rhobonda. Type locality: Brazil: Ega [=Tefe], Amazonas [holotype 9,
BMNH].

Originally described in Gelechidae [sic] as a replacement name for Rho-
bonda Walker, 1864, a junior homonym of Rhobonda Walker, 1863, in Cho-
reutidae. Carna Walker, 1864, is a junior homonym of Carna Gistel, 1848

(Echinodermata), and is currently considered to be a junior synonym of

Dichomeris Hiibner, [1825], in Gelechiidae: Dichomerinae.

Cebysa Walker, 1854:486 Psychidae

Type-species: Cebysa leucotelus Walker, 1854:486, by monotypy. Type
locality: Australia: Sydney, New South Wales [holotype 9, BMNH].

Originally described in Lithosiidae [=Arctiidae]; subsequently transferred
to Plutellidae by Meyrick (1907:92); transferred to Glyphipterygidae [sic] by
Meyrick (1913b:32); transferred to Psychidae by Common (1970b:804).

Cercosimma Diakonoff, 1948:197 *Immidae?

Type-species: Cercosimma electrodes Diakonoff, 1948:199, by original
designation. Type locality: Indonesia: Burn Id. [holotype 8 , RMNL?].

Originally described in Glyphipterygidae [sic]; hereby tentatively trans-
ferred to Immidae.

Chalenata Walker, 1864:1001 Noctuidae

Type-species: Chalenata micaceella Walker, 1864:1001, by monotypy.
Type locality: Brazil, Ega [=Tefe], Amazonas [holotype 9, BMNH].
Originally described in Choreutidae; now in Noctuidae: Acontiinae.

Charitographa Diakonoff, 1979:291 Tortricidae

Type-species: Hilarographa mikadonis Stringer, 1930:418, by original
designation. Type locality: Japan: Sapporo [holotype 8 , BMNH].

Originally described in the tribe Hilarographini of Tortricidae: Chlidano-
tinae.

Charixena Meyrick, 1920a:279 *Plutellidae

Type-species: Philpottia iridoxa Meyrick, 1916:417, by original designa-

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245

tion. Type locality: New Zealand: Mt. Burns, Hunter Mts., South Island
[lectotype unselected, BMNH].

Originally described as a replacement name for Philpottia Meyrick, 1916,
in Glyphipterygidae [sic]; hereby transferred to Plutellidae.

Charixena Meyrick, 192 lb:335, is a redescription of Charixena Meyrick,
1920.

Charizena [sic] Neave, 1939:673 *Plutellidae

A misspelling of Charixena Meyrick, 1920, hereby in Plutellidae.

Chimaera Ochsenheimer, 1808:2 Brachodidae

Type-species: Sphinx chimaera Hiibner, 1796:pl. 1, fig. 1 [= Sphinx ap-
pendiculata Esper, 1783:227], by subsequent designation of Blanchard
(1840:474). Type locality (chimaera): Europe [type lost?]. Preoccupied by
Chimaera Linnaeus, 1758 (Pisces).

Originally described in Sphinges; subsequently transferred to Chimerites
by Blanchard (1840:474); transferred to Glyphipterygidae [sic] as a junior
synonym of Phy codes Guenee, 1852, by Meyrick (1913b:32); transferred to
Brachodidae as a junior synonym of Brachodes Guenee, 1845, by Heppner
(1979a: 127).

Walsingham (1900b: 568) designated Chimaera radiata Ochsenheimer,
1808:5, as the type-species of Chimaera Ochsenheimer, thus, making the
genus a synonym of Phy codes Guenee, 1852, due to homonymy with Chi-
maera Linnaeus, 1758, and equal type-species. However, the type-species
of Chimaera Ochsenheimer, 1808, is Sphinx chimaera Hiibner, 1796, by an
earlier designation (Blanchard, 1840), thus making the genus a junior syn-
onym of Brachodes Guenee, 1845.

Chimera [sic] Feisthamel, 1833:259 Brachodidae

A misspelling of Chimaera Ochsenheimer, 1808, now a junior synonym
of Brachodes Guenee, 1845, in Brachodidae.

Chloreutis [sic] Viette, 1948a: 40 Choreutidae

A misspelling of Choreutis Hiibner, [1825], in Choreutidae.

Chordates Snellen, 1877:49 Choreutidae

Type-species: Simaethis pronubana Snellen, 1877:48, by subsequent des-
ignation of Fletcher (1929:47). Type locality: [Indonesia]: Java [lectotype
9 (present designation), RMNL].

Originally described in Tineina; subsequently transferred to Hemerophil-
idae as a junior synonym of Tortyra Walker, 1863, by Walsingham
(1914:312); transferred to Choreutidae as a junior subjective synonym of
Saptha Walker, 1864, by Heppner (1981a:55).

Choredates [sic] Pagenstecher, 1884:289 Choreutidae

A misspelling of Chordates Snellen, 1877, now a synonym of Saptha
Walker, 1864, in Choreutidae.

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NEW YORK ENTOMOLOGICAL SOCIETY

Choregia Felder and Rogenhofer, 1875:6 Choreutidae

Type-species: Choregia fulgens Felder and Rogenhofer, 1875:6, by sub-
sequent designation of Heppner (1981a:56). Type locality: Colombia: Bogota
[lectotype 8 (present designation), BMNH].

Originally described in Tineidae; subsequently transferred to Choreutiden
by Zeller (1877:191) as Choregia Zeller; transferred to Hemerophilidae
as a junior synonym of Tortyra Walker, 1863, by Walsingham (1914:312);
transferred to Choreutidae as a junior subjective synonym of Tortyra
Walker, 1863, by Heppner (198 la: 56).

Felder and Rogenhofer (1875) attributed Choregia to Zeller but the Zeller
description did not appear until 1877.

Choregia Zeller, 1877:191 Choreutidae

Type-species: Choregia fulgens Felder and Rogenhofer, 1875:6, by sub-
sequent designation of Meyrick (1914d: 18). Type locality: Colombia: Bogota
[lectotype 8 (present designation), BMNH]. Preoccuppied by Choregia
Felder and Rogenhofer, 1875 (Choreutidae).

Originally described in Choreutiden; subsequently transferred to Hemero-
philidae as a junior synonym of Tortyra Walker, 1863, by Walsingham
(1914:312); transferred to Choreutidae as a junior subjective synonym
of Tortyra Walker, 1863, by Heppner (198 la: 56).

Chorentes [sic] Morris, 1871 :iv Choreutidae

A misspelling of Choreutis Hiibner, [1825], now in Choreutidae.

Chorentis [sic] Turner, 1898:203 Choreutidae

A misspelling of Choreutis Hiibner, [1825], now in Choreutidae.

Choreutes Treitschke, 1835:31 Choreutidae

An unjustified emendation of Choreutis Hiibner, [1825], now in Choreu-
tidae.

Choreutes Burmeister, 1838 (Collembola), is a junior homonym of Cho-
reutes Treitschke, 1835.

Choreuthis [sic] Hackman, 1947:71 Choreutidae

A misspelling of Choreutis Hiibner, [1825], now in Choreutidae.

Choreutidia Sauber, 1902:702 Choreutidae

Type-species: Choreutidia sexfasciella Sauber, 1902:702, by monotypy.
Type locality: Philippines: Luzon [holotype 8 , SMF].

Originally described in Hyponomeutidae [sic]; subsequently transferred
to Glyphipterygidae [sic] as a junior synonym of Choreutis Hiibner, [1825],
by Meyrick (1913b:38); transferred to Choreutidae as a junior subjective
synonym of Choreutis Hiibner, [1825], by Heppner (1981a:53).

Choreutis Hiibner, [1825]: 373 Choreutidae

Type-species: [ Phalaena ] pariana Clerck, 1759:pl. 10, fig, 9, by subse-

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247

quent designation of Walsingham (1908:987). Type locality: Europe [type
lost?].

Originally described in Tortrices; subsequently transferred (as Choreutes
[sic]) to Choreutidae by Stainton (1859:158); transferred to Glyphipterygidae
[sic] by Wocke (1871:265); now in Choreutidae.

Choreutis of authors [not Hiibner, 1825] Choreutidae

This name was used by most authors in reference to Pyralis myllerana
Fabricius, 1794, and congeners, especially since Meyrick (1914d), but the
first valid type-species designation for Choreutis Hiibner, [1825], was only
recently confirmed to be that of Walsingham (1908:987), being Phalaena
pariana Clerck, 1759, and not Pyralis myllerana Fabricius, 1794. Conse-
quently, the name Choreutis Hiibner, [1825], has been transferred to species
until recently referred to Eutromula Frolich, 1828, while Choreutis of au-
thors has been given the new name Prochoreutis Diakonoff and Heppner,
1980 (Diakonoff and Heppner, 1980:196).

Chrysocentris Meyrick, 19 14c: 284 Glyphipterigidae

Type-species: Chrysocentris clavaria Meyrick, 1914c:284, by monotypy.
Type locality: Nyasaland [=Malawi]: Mt. Mlanje [holotype 9, BMNH].
Originally described in Glyphipterygidae [sic].

Cibdeloses Durrant, 1919:121 Zygaenidae

Type-species: Cibdeloses dolopis Durrant, 1919:121, by original desig-
nation. Type locality: India: Khasis, Assam [holotype 9, BMNH].

Originally described in Hemerophilidae [= Choreutidae]; subsequently
transferred to Zygaenidae by Heppner (1981a: 15).

Circica Meyrick, 1888:88 Glyphipterigidae

Type-species: Circica cionophora Meyrick, 1888:88, by monotypy. Type
locality: New Zealand: Christchurch, South Island [lectotype S (present
designation), BMNH].

Originally described in Tineina; subsequently transferred to Glyphipte-
rygidae [sic] as a junior synonym of Glyphipteryx [sic] Hiibner, [1825], by
Meyrick (1913b:41); now considered as a distinct genus in Glyphipterigidae.

Cloreutis [sic] Kautz, 1930:28 Choreutidae

A misspelling of Choreutis Hiibner, [1825], in Choreutidae.

Colpotorna Meyrick, 1920b:325 *Psychidae

Type-species: Colpotorna lasiopa Meyrick, 1920b:326, by monotypy.
Type locality: Australia: Brisbane, Queensland [holotype 6 , BMNH].

Originally described in Glyphipterygidae [sic]; hereby transferred to Psy-
chidae.

Coridomorpha Meyrick, 1914b: 111 *Oecophoridae

Type-species: Coridomorpha Stella Meyrick, 1914b: 111, by monotypy.

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NEW YORK ENTOMOLOGICAL SOCIETY

Type locality: New Zealand [lectotype unselected, BMNH].

Originally described in Glyphipterygidae [sic]; hereby trasnferred to Oe-
cophoridae: Oecophorinae.

Corsocasis Meyrick, 1912a:59 Heliodinidae

Type-species: Corsocasis coronias Meyrick, 1912a:59, by monotypy.
Type locality: India and Sri Lanka [lectotype unselected, BMNH?].

Originally described in Glyphipterygidae [sic]; subsequently transferred
to Heliodinidae by Meyrick (1913b:20).

Cotaena Walker, [1865] :2 1 Glyphipterigidae

Type-species: Cotaena mediana Walker, [1865]:21, by monotypy. Type
locality: Brazil: Para [holotype 9, UMO].

Originally described in Aegeriidae [=Sesiidae]; subsequently transferred
to Heliodinidae by Naumann (1971:15); transferred to Glyphipterigidae by
Heppner (1981a:44).

Cronicombra Meyrick, 1920b: 3 27 Glyphipterigidae

Type-species: Cronicombra granulata Meyrick, 1920b: 3 27, by monotypy.
Type locality: Brazil: Para [holotype S , BMNH].

Originally described in Glyphipterygidae [sic].

Cylicophora Turner, 1927:156 Yponomeutidae

Type-species: Cylicophora collina Turner, 1927:156, by monotypy. Type
locality: Australia: Cradle Mt., Tasmania [holotype <3, ANIC].

Originally described in Glyphipterygidae [sic]; subsequently transferred
to Yponomeutidae as a junior subjective synonym of Tanaoctena Turner,
1913, by Clarke (1971:167).

Davendra Moore, 1887:520 *Immidae

Type-species: Davendra mackwoodii Moore, 1887:520, by original des-
ignation. Type locality: Ceylon [=Sri Lanka] [lectotype unselected,
BMNH].

Originally described in Gelechiidae; subsequently transferred to Plutelli-
dae as a junior synonym of Imma Walker, [1859], by Meyrick (1906c: 170);
hereby transferred to Immidae as a junior subjective synonym of Imma
Walker, [1859].

Desmidoloma Erschoff, 1892:671 Glyphipterigidae

Type-species: S taint onia fulgens Erschoff, 1877:347 [=Argyresthia per-
ornatella Walker, 1864:840], by monotypy. Type locality {fulgens ): China:
Amur region, [Manchuria] [holotype 9 , lost?].

Originally described without family reference; subsequently transferred
to Glyphipterygidae [sic] as a junior subjective synonym of Glyphipteryx
[sic] Hiibner, [1825], by Meyrick (1913b:41); here considered a junior sub-
jective synonym of Lepidotarphius Pryer, 1877, in Glyphipterigidae.

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249

Diploschizia Heppner, 1981b:311 Glyphipterigidae

Type-species: Glyphipteryx [sic] impigritella Clemens, 1863:9, by original
designation. Type locality: USA: [Easton, Northampton County,
Pennsylvania] [holotype 8 , ANSP].

Originally described in Glyphipterigidae.

Ditrigonophora Walsingham, 1897b: 117 *Yponomeutidae?

Type-species: Ditrigonophora marmoreipennis Walsingham, 1897b: 118,
by original designation. Type locality: Grenada: Balthazar [lectotype (pres-
ent designation), BMNH].

Originally described in Glyphipteryginae [sic] of Hyponomeutidae [sic];
subsequently transferred to Glyphipterygidae [sic] by Meyrick (1913b:40);
hereby tentatively transferred to Yponomeutidae, [syntypes lack abdomens
and heads].

Douglasia Stainton, 1854:179 Douglasiidae

Type-species: Gracilaria [sic] ocnerostomellum Stainton, 1850:6, by orig-
inal designation and monotypy. Type locality: England [lectotype unselect-
ed, BMNH].

Originally described in Glyphipterygidae [sic]; subsequently transferred
to Elachistidae by Meyrick (1895:684); transferred to Douglasiadae [sic] by
Meyrick (1928:721); now considered a junior subjective synonym of Tin-
agma Zeller, 1839, in Douglasiidae.

Dudua Walker, 1864:1000 Tortricidae

Type-species: Dudua hesperialis Walker, 1864:1000, by monotypy. Type
locality: Sarawak [holotype 8, BMNH].

Originally described in Choreutidae; subsequently transferred to Tortric-
idae by Walsingham (1900a: 135).

Electrographa Meyrick, 1912a:63 Glyphipterigidae

Type-species: Electrographa thiolychna Meyrick, 1912a:63, by mono-
typy. Type locality: Burma: Momeit [holotype 8 , BMNH].

Originally described in Glyphipterygidae [sic].

Ellabella Busck, 1925:46 Plutellidae

Type-species: Ellabella editha Busck, 1925:46, by original designation.
Type locality: Canada: Saanichton, British Columbia [holotype 9 , BMNH].

Originally described in Glyphipterygidae [sic]; subsequently transferred
to Plutellidae by Heppner (1978a: 50).

Embolostoma Diakonoff, 1977b:51 Tortricidae

Type-species: Embolostoma plutostola Diakonoff, 1977b:52, by original
designation. Type locality: Indonesia: West Java [holotype 9, RMNL].

Originally described in the tribe Hilarographini of Tortricidae: Chlidano-
tinae.

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Embryonopsis Eaton, 1876:61 Yponomeutidae

Type-species: Embryonopsis halticella Eaton, 1876:61, by monotypy.
Type locality: New Zealand: Heard Id. [lectotype 8 (Common, 1970a:231),
BMNH].

Originally described in Gelechiidae; subsequently transferred to Ypono-
meutidae by Viette (1948b: 16).

Encratora Meyrick, 1923:618 *Heliodinidae

Type-species: Encratora plumbigera Meyrick, 1923:618, by monotypy.
Type locality: India: Shillong, Assam [holotype 9, BMNH].

Originally described in Glyphipterygidae [sic]; hereby transferred to He-
liodinidae.

Entomoloma Ragonot, 1875:45 Choreutidae

Type-species: Tortrix nemorana Hiibner, [17 96—99] : pi . 1, fig. 3, by sub-
sequent designation of Walsingham (1908:988). Type locality: Europe [type
lost?].

Originally described in Choreutina [= Choreutidae]; subsequently trans-
ferred to Glyphipterygidae [sic] as a junior synonym of Simaethis Leach,
1815, by Meyrick (1913b:34); transferred to Choreutidae as a junior subjec-
tive synonym of Eutromula Leach, 1815, by Arita and Diakonoff (1979:8);
now a junior subjective synonym of Choreutis Hiibner, [1825] (Heppner,
1981a:53).

Epicroesa Meyrick, 1907:94 *Heliodinidae

Type-species: Epicroesa ambrosia Meyrick, 1907:96, by subsequent des-
ignation of Meyrick (1914d: 17). Type locality: Australia: Queensland [lec-
totype unselected, BMNH].

Originally described in Plutellidae; subsequently transferred to Glyphip-
terygidae [sic] by Meyrick (1912a:58); hereby transferred to Heliodinidae.

Epithetica Turner, 1923:165 *Oecophoridae

Type-species: Epithetica typhoscia Turner, 1923:165, by monotypy. Type
locality: Australia: Lismore, New South Wales [lectotype 8 (present des-
ignation), ANIC].

Originally described in Glyphipterygidae [sic]; hereby transferred to Oe-
cophoridae: Hypertrophinae.

Ereunetis Meyrick, 1881:258 Tineidae

Type-species: Ereunetis iuloptera Meyrick, 1881:260, by subsequent des-
ignation of Walsingham (1907:714). Type locality: Australia: Sydney, New
South Wales [lectotype unselected, BMNH].

Originally described in Erechthiadae [sic] [= Tineidae]; subsequently
transferred to Lyonetiadae [sic]; now considered in Tineidae.

Meyrick (1912b: 122) listed a new species of Ereunetis under Glyphipte-
rygidae [sic] but the genus has always been associated with the Tineoidea
and is now in Tineidae.

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251

Ernolytis Meyrick, 1922a:488 Glyphipterigidae

Type-species: Ernolytis chlorospora Meyrick, 1922a:488, by monotypy.
Type locality: Fiji: Lolotu [lectotype unselected, BMNH].

Originally described in Glyphipterygidae [sic].

Euprophantis Meyrick, 1921a: 191 Gracillariidae

Type-species: Euprophantis autoglypta Meyrick, 1921a: 191, by mono-

typy. Type locality: [Indonesia]: Pekalongan, Java [lectotype 6 (present
designation), RMNL].

Originally described in Gracilariadae [sic]; subsequently partially associ-
ated with Glyphipterix Hiibner, [1825]; now in Gracillariidae.

Eupselia Meyrick, 1881:216 Oecophoridae

Type-species: Eupselia satrapella Meyrick, 1881:220, by subsequent des-
ignation of Meyrick (1922b: 148). Type locality: Australia: Sydney, New
South Wales [lectotype unselected, BMNH].

Originally described in Glyphipterygidae [sic]; subsequently transferred
to Oecophoridae (Oecophorinae) by Meyrick (1922b: 148).

Euthorybeta Turner, 1913:200 Brachodidae

Type-species: Euthorybeta xanthoplaca Turner, 1913:201, by original
designation. Type locality: Australia: Stadbroke Id., Queensland [holotype
9 , ANIC].

Originally described in Plutellidae (Yponomeutinae); subsequently trans-
ferred to Brachodidae by Heppner (1979a: 127).

Eutromula Frolich, 1828:11 Choreutidae

Type-species: [Phalaena] pariana Clerck, 1759:pl. 10, fig. 9, by subse-
quent designation of Fletcher (1929:93). Type locality: Europe [type lost?].

Originally described without family placement; subsequently transferred
to Glyphipterygidae [sic] by Fletcher (1929:93) as a junior synonym of An-
thophila Haworth, [1811]; transferred to Choreutidae by Heppner
(1977b:633); now a junior obiective synonym of Choreutis Hiibner, [1825]
(Heppner, 1981a:53).

Ganabalia Diakonoff, 1975:316 Tortricidae

Type-species: Ganabalia planipes Diakonoff, 1975:317, by original des-
ignation. Type locality: Indonesia: Celebes [holotype S , BMNH].

Originally described in “Choreutinae” of Tortricidae; transferred to Tor-
tricidae: Olethreutinae by Diakonoff (1981:71).

Gauris Hiibner, 1821 : [1] Choreutidae

Type-species: Phalaena ( Tortrix ) albertiana Cramer, 1781:163, by mono-
typy. Type locality: Surinam [type lost?].

Originally listed in an index and described in Tineina by Hiibner
([1825]:374); subsequently transferred to Atychianae [sic] of Tineidae by
Walsingham (1892:529); transferred to Glyphipteryginae [sic] of Hypono-

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NEW YORK ENTOMOLOGICAL SOCIETY

meutidae [sic] by Walsingham (1897b: 120); transferred to Choreutidae as a
junior objective synonym of Hemerophila Hiibner, [1817], by Heppner
(1977b: 634).

Glyphipterix Hiibner, [1825]:421 Glyphipterigidae

Type-species: Glyphipterix linneella Hiibner, [1825] :42 1 [ =Tortrix li-
neana Hiibner, [1796-99]:pl. 14, fig. 8; =Tinea bergstraesserella Fabricius,
1781:302] [not Phalaena linneella Clerck, 1759:pl. 12, fig. 8 (Agonoxen-
idae)], by subsequent designation of Duponchel (1845:243). Type locality
(, linneella Hiibner): Europe [type lost?].

Originally described in Tineae; subsequently transferred to Glyphipte-
rygidae [sic] by Stainton (1854:173).

Glyphipterys [sic] Christoph, 1882:38 Glyphipterigidae

A misspelling of Glyphipterix Hiibner, [1825], now in Glyphipterigidae.

Glyphipteryx Curtis, 1827:152 Agonoxenidae

Type-species: [Phalaena] linneella Clerck, 1759:pl. 12, fig. 8, by original
designation. Type locality: Europe [type lost?].

Originally described without family placement; subsequently a nomen
oblitum until resurrected and transferred to Momphidae (Cosmopteriginae)
by Bradley (1972:25); here considered a junior objective synonym of Chry-
soclista Stainton, 1854, by suppression under plenary powers (case ICZN
2115) (Diakonoff and Heppner, 1977), in Agonoxenidae.

Glyphipteryx Zeller, 1839:203 Glyphipterigidae

An unjustified emendation of Glyphipterix Hiibner, [1825] [ not Glyphip-
teryx Curtis, 1827 (Agonoxenidae)], now in Glyphipterigidae.

Glyphiptoryx [sic] Mann and Rogenhofer, 1878:500 Glyphipterigidae

A misspelling of Glyphipterix Hiibner, [1825], now in Glyphipterigidae.

Glyphiteryx [sic] Fischer von Roslerstamm, 1841:233 Glyphipterigidae

A misspelling of Glyphipterix Hiibner, [1825], now in Glyphipterigidae.

Glyphopteryx Herrich-Schaffer, 1854:92 Glyphipterigidae

An unjustified emendation of Glyphipterix Hiibner, [1825], [ not Glyphip-
teryx Curtis, 1827 (Agonoxenidae)], now in Glyphipterigidae.

Glyphptieryx [sic] Turati, 1879:203 Glyphipterigidae

A misspelling of Glyphipterix Hiibner, [1825], now in Glyphipterigidae.

Glyphteryx [sic] Watt, 1920:439 Glyphipterigidae

A misspelling of Glyphipterix Hiibner, [1825], now in Glyphipterigidae.

Glyphyteryx [sic] Hampson, 1918:387 Glyphipterigidae

A misspelling of Glyphipterix Hiibner, [1825], now in Glyphipterigidae.

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253

Gora Walker, 1862:89 Brachodidae

Type-species: Gora aequalis Walker, 1862:90, by monotypy. Type local-
ity: Brazil [holotype 9, UMO].

Originally described in Heliothidae ONoctuidae]; subsequently trans-
ferred to Glyphipterygidae [sic] as a junior synonym of Sagalassa Walker,
1856, by Meyrick (1913b:31); transferred to Brachodidae as a junior subjec-
tive synonym of Sagalassa Walker, 1856, by Heppner (1981a: 14).

Guaris [sic] Fernald, 1900:236 Choreutidae

A misspelling of Gauris Hiibner, 1821, now a junior objective synonym
of Hemerophila Hiibner, [1817], in Choreutidae.

Heliomanes Sodoffsky, 1837:89 Noctuidae

An unjustified replacement name for Anthophila Ochsenheimer, 1816 [not
Anthophila Haworth, 1811], now a junior synonym of Porphyrinia Hiibner,
[1821], in Noctuidae.

Heliostibes Zeller, 1874:434 *Oecophoridae

Type-species: Heliostibes mathewi Zeller, 1874:435, by monotypy. Type
locality: Chile: Valparaiso [holotype 6, BMNH].

Originally described in Tineacea; subsequently transferred to Gelechiidae
by Butler (1883:76); transferred to Glyphipterygidae [sic] by Meyrick
(1913b :29); hereby transferred to Oecophoridae: Oecophorinae.

Hemerophila Hiibner, [1806]: [2] Choreutidae

Type-species: [ Phalaena ] pariana Clerck, 1759:pl. 10, fig. 9, by monotypy.
Type locality: Europe [type lost?].

Originally described in Tortrices; now an unavailable synonym of Cho-
reutis Hiibner, [1825], in Choreutidae.

Hemerophila Hiibner, [1806], is an unavailable name due to publication
in a work rejected for nomenclatural purposes by Opinion 97 (ICZN, 1926).

Hemerophila Hiibner, [18 17]: pi. 213, fig. 1 Choreutidae

Type-species: Phalaena ( Tortrix ) albertiana Cramer, 1781:163, by mono-
typy. Type locality: Surinam [type lost?].

Originally described in Tortrices; subsequently transferred to Hemero-
philidae by Walsingham (1910:257); transferred to Glyphipterygidae [sic] (as
Hemerophila sensu Fernald, 1900) as a junior synonym of Simaethis Leach,
1815, by Meyrick ( 1913b :34); transferred to Choreutidae by Heppner
(1977b:634) as a distinct genus.

Hemerophila sensu Fernald, 1900:239 Choreutidae

A redescription of Hemerophila Hiibner, [1806], including Simaethis
vicarialis Zeller, 1875:322 [ =Tortrix diana Hiibner, 1819-22]; now a junior
subjective synonym of Choreutis Hiibner, [1825], in Choreutidae.

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NEW YORK ENTOMOLOGICAL SOCIETY

Hemerophila Stephens, 1829a:43 Geometridae

Type-species: Phalaena abruptaria Thunberg, 1792:59, by monotypy.
Type locality: Sweden [type lost?]. Preoccuppied by Hemerophila Hiibner,
[1817] (Choreutidae).

Originally described in Geometrae [= Geometridae]; now a junior objec-
tive synonym of Menophra Moore, 1887, in Geometridae.

Hemerophila Stephens was redescribed twice by Stephens (Stephens,
1829b: 125; 1831:189).

Heribeia Stephens, 1829a:49 Glyphipterigidae

Type-species: Tinea forsterella Fabricius, 1787:252, by subsequent des-
ignation of Westwood (1840:112). Type locality: Germany: Hamburg [type
lost?].

Originally listed in 1829 and described in Yponomeutidae by Stephens
(1834:261); subsequently transferred to Glyphipterygidae [sic] by Stainton
(1854:163) as a junior subjective synonym of Glyphipterix Hiibner, [1825].

Heribeia Stephens, 1829b:207, a redescription, was published in July,
1829, while the 1829a date is June, 1829.

Hierodoris Meyrick, 1912a:41 *Oecophoridae

Type-species: Hierodoris iophanes Meyrick, 1912a:42, by monotypy.
Type locality: New Zealand: Wellington, North Island [holotype 6 ,
BMNH].

Originally described in Glyphipterygidae [sic]; hereby transferred to Oe-
cophoridae: Oecophorinae.

Hilarographa Zeller, 1877:187 Tortricidae

Type-species: Phalaena ( Tortrix ) swederiana Stoll, 1790:75, by subse-
quent designation of Walsingham (1897a:51). Type locality: Surinam [type
lost?].

Originally described as a subgenus of Setiostoma Zeller, 1875, in Tortri-
cinen [= Tortricidae]; subsequently transferred to Glyphipterygidae [sic] as
a distinct genus by Meyrick (1886: 286); transferred to the tribe Hilarogra-
phini of Tortricidae: Chlidanotinae by Diakonoff (1977a:76).

Hilarographa sensu Meyrick (1886:286) Tortricidae

A redescription of Hilarographa Zeller, 1877, based on Hilarographa
zapyra Meyrick (1886:286); now an objective synonym of Thaumatographa
Walsingham, 1897, in the tribe Hilarographini of Tortricidae: Chlidanotinae.

Homadaula Lower, 1899:115 Plutellidae

Type-species: Homadaula lasiochroa Lower, 1899:115, by monotypy.
Type locality: Australia: Broken Hill, New South Wales [lectotype unse-
lected, SAMA?].

Originally described in Plutellidae; subsequently transferred to Glyphip-
terygidae [sic] by Clarke (1943:206); transferred to Plutellidae by Friese
(1962:302).

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255

Homadaula Meyrick, 1907:73 Plutellidae

Type-species: Homadaula myriospila Meyrick, 1907:73, by original des-
ignation. Type locality: Australia: West Australia [lectotype unselected,
BMNH?].

A redescription of Homadaula Lower, 1899, in Plutellidae.

Homoplastis Meyrick, 1926a: 162 *Oecophoridae?

Type-species: Homoplastis agathoclea Meyrick, 1926a: 162, by mono-
typy. Type locality: [Malaysia]: Sarawak: Mt. Murud [holotype 9, SMS?].

Originally described in Glyphipterygidae [sic]; hereby tentatively trans-
ferred to Oecophoridae: Oecophorinae.

Hoplophractis Meyrick, 1920b: 3 26 Brachodidae

Type-species: Hoplophractis heptachalca Meyrick, 1920b:326, by mono-
typy. Type locality: Brazil: Obidos, Para [lectotype 6 (Clarke, 1969:99),
BMNH].

Originally described in Glyphipterygidae [sic]; subsequently transferred
to Brachodidae by Heppner (1979a: 127).

Hyperperissa Walsingham, 1900b: 546 *Immidae

Type-species: Sidyma aurantiaca Semper, 1899:501, by original desig-
nation. Type locality: Philippines: Mindanao [lectotype undesignated,
BMNH].

Originally described in Gelechiadae [sic]; subsequently transferred as a
junior synonym of Imma Walker, [1859], to Plutellidae by Meyrick
(1906c: 170) and to Glyphipterygidae [sic] by Meyrick (1913b:25); hereby
transferred to Immidae as a junior subjective synonym of Birthana Walker,
[1865], new synonymy.

Hypertropha Meyrick, 1881:208 Oecophoridae

Type-species: Hypertropha thesaurella Meyrick, 1881:209, by monotypy.
Type locality: Australia: Parramatta, New South Wales [lectotype 9 (Diako-
noff, 1954:474), BMNH].

Originally described in Glyphipterygidae [sic]; subsequently transferred
to Copromorphidae by Meyrick (1906a:51); transferred to Glyphipterygidae
[sic]: Hypertrophinae by Diakonoff (1954:274); transferred to Oecophoridae:
Hypertrophinae by Common (1970b:819).

Idiothauma Walsingham, 1897a:49 Tortricidae

Type-species: Idiothauma africanum Walsingham, 1897a:50, by original
designation. Type locality: French Congo [=Republic of the Congo]:
Kangwe, Ogowe River [lectotype 8 (present designation), BMNH].

Originally described in Glyphipteryginae of Hyponomeutidae [sic]; sub-
sequently transferred to Glyphipterygidae [sic] as a synonym of Hilarogra-
pha Zeller, 1877, by Meyrick (1913b:24); transferred to the tribe Hilarogra-
phini of Tortricidae: Chlidanotinae as a distinct genus by Heppner
(1978a:53).

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NEW YORK ENTOMOLOGICAL SOCIETY

Imma Walker, [1859]: 159 Immidae

Type-species: Imma rugosalis Walker, [1859]: 195, by monotypy. Type
locality: Ceylon [=Sri Lanka] [holotype 8 , BMNH].

Originally described in Herminidae [=Noctuidae]; subsequently trans-
ferred to Plutellidae sensu Meyrick (1906c: 169); transferred to Glyphip-
terygidae [sic] by Meyrick (1910:464); transferred to Immidae by Heppner
(1977a: 129).

Inapha Walker, 1864:999 Gelechiidae

Type-species: Inapha lampronialis Walker, 1864:1000 [ =Thubana bisig-
natella Walker, 1864:814], by monotypy. Type locality ( lampronialis ):
[Malaysia]: Sarawak [holotype 8, BMNH].

Originally described in Choreutidae; subsequently transferred to Gele-
chiidae by Meyrick (1925:234), as a junior subjective synonym of Thubana
Walker, 1864:814.

Irianassa Meyrick, 1905:609 *Tortricidae

Type-species: Irianassa sapphiropa Meyrick, 1905:609, by monotypy.
Type locality: Ceylon [=Sri Lanka]: Kandy [holotype 8 , BMNH].

Originally described in Plutellidae sensu Meyrick; subsequently trans-
ferred to Glyphipterygidae [sic] by Meyrick (1913b:23); hereby transferred
to the tribe Hilarographini of Tortricidae: Chlidanotinae,

Iridostoma Meyrick, 1909b:425 *Yponomeutidae

Type-species: Iridostoma ichthyopa Meyrick, 1909b:425, by monotypy.
Type locality: Ceylon [=Sri Lanka]: Peradeniya [holotype 9, BMNH].

Originally described in Plutellidae sensu Meyrick; subsequently trans-
ferred to Glyphipterygidae [sic] by Meyrick (1913b:23); transferred to Ypon-
omeutidae by Heppner (198 la:57).

Irinympha Meyrick, 1932:274 Glyphipterigidae

Type-species: Irinympha aglaograpta Meyrick, 1932:275, by monotypy.

Type locality: Uganda: Entebbe [holotype 8 , BMNH].

Originally described in Glyphipterygidae [sic].

Jobula Walker, 1866:1888 *Immidae

Type-species: Jobula semilinea Walker, 1866:1889, by monotypy. Type

locality: [Indonesia]: Sula [holotype 8, UMO].

Originally described in Lithosiidae [=Arctiidae]; subsequently transferred
to Hyponomeutidae [sic] by Walsingham (1897a:46); transferred to Plutel-
lidae sensu Meyrick as a synonym of Imma Walker, [1859], by Meyrick,
(1906c: 170); transferred to Glyphipterygidae [sic] as a synonym of Imma
Walker, [1859], by Meyrick (191 3b: 25); hereby transferred to Immidae
as a junior subjective synonym of Moca Walker, 1863, new synonymy.

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257

Jonaca Walker, 1863:457 Brachodidae

Type-species: Jonaca compulsana Walker, 1863:457 [ =SagaIassa valida
Walker, 1856:6], by monotypy. Type locality ( compulsana ): Brazil: Ega
[=Tefe] [holotype 8, BMNH].

Originally described in Choreutidae; subsequently transferred to Glyphip-
terygidae [sic] by Meyrick (1913b:31), as a synonym of Sagalassa Walker,
1856; transferred to Brachodidae as a junior subjective synonym of Saga-
lassa Walker, 1856, by Heppner (1981a: 14).

Klimeschia Amsel, 1938:89 Douglasiidae

Type-species: Klimeschia lutumella Amsel, 1938:89, by original desig-

nation. Type locality: [Israel]: Jerusalem [holotype 8 , UMB].

Originally described in Douglasiidae.

Lamprystica Meyrick, 1914a:58 *Oecophoridae

Type-species: Lamprystica purpurata Meyrick, 1914a:58, by monotypy.
Type locality: Formosa [^Taiwan]: Kosempo [lectotype unselected,
HZMB].

Originally described in Glyphipterygidae [sic]; hereby transferred to
Stathmopodini of Oecophoridae: Oecophorinae.

Lasiodictis Meyrick, 1912a:41 Oecophoridae

Type-species: Lasiodictis melistoma Meyrick, 1912a:41, by monotypy.

Type locality: [India]: Khasi Hills, Assam [lectotype 8 (Clarke, 1955b:438),
BMNH].

Originally described in Glyphipterygidae [sic]; subsequently transferred
to Xyloryctidae [sic] by Clarke (1955a: 19); transferred by Hodges ([1979]:8),
to Oecophoridae: Autostichinae.

Lepidotarphius Pryer, 1877:235 Glyphipterigidae

Type-species: Lepidotarphius splendens Pryer, 1877:235 [=Argyresthia
perornatella Walker, 1864:1040], by monotypy. Type locality ( splendens ):
China: Shanghai [lectotype unselected, BMNH].

Originally described without family reference but noted to be related to
Butalis (Gelechiidae); subsequently transferred to Glyphipterygidae [sic] as
a synonym of Glyphipteryx [sic] Hiibner, [1825], by Meyrick (1913b:41);
here considered a distinct genus in Glyphipterigidae.

Lithariapteryx Chambers, 1876:217 Heliodinidae

Type-species: Lithariapteryx abroniaeella Chambers, 1876:217, by mono-
typy. Type locality: USA: Edgerton, Colorado [holotype, MCZ?].

Originally described in Tineina; subsequently transferred to Glyphipte-
rygidae [sic] by Riley (1891:104); transferred to Heliodinidae as a synonym
of Heliodines Stainton, 1854, by Meyrick (1913b: 17); now considered a
distinct genus in Heliodinidae.

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NEW YORK ENTOMOLOGICAL SOCIETY

Litobrenthia Diakonoff, 1978:28 Choreutidae

Type-species: Brenthia japonica Issiki, 1930:424, by original designation.
Type locality: Japan: Hasimoto [lectotype (Diakonoff, in press), USNM].
Originally described in Choreutidae.

Lotisma Busck, 1909:98 Copromorphidae

Type-species: Sciaphila trigonana Walsingham, 1879:22, by original des-
ignation. Type locality: USA: Mendocino, California [lectotype 9 (present
designation), BMNH].

Originally described as being related to Hemerophila Hiibner; subse-
quently transferred to Glyphipterygidae [sic] by Busck (1925:46); transferred
to Copromorphidae by Heppner (1978a:49).

Loxotrochis Meyrick, 1906c:205 *Immidae

Type-species: Loxotrochis sepias Meyrick, 1906c: 205, by monotypy.
Type locality: Brazil: Espirito Santo [holotype 6 , BMNH].

Originally described in Plutellidae sensu Meyrick; subsequently trans-
ferred to Glyphipterygidae [sic] by Meyrick (1913b:25); hereby transferred
to Immidae.

Meyrick (1906c) originally indicated the type locality to be “Espiritu Santo,
New Hebrides” but he later corrected this (Meyrick, 1907:108) to read Es-
pirito Santo, Brazil.

Lygronoma Meyrick, 1913a: 100 Oecophoridae

Type-species: Lygronoma sporimaea Meyrick, 1913a: 100, by monotypy.
Type locality: British Guiana [= Guyana]: Bartica [lectotype S (Clarke,
1969:175), BMNH].

Originally described in Glyphipterygidae [sic]; subsequently transferred
by Heppner (1981a:57), to Oecophoridae: Stenomatinae; hereby transferred
to Oecophoridae: Oecophorinae (Becker, pers. comm.).

Machlotica Meyrick, 1909a: 36 Glyphipterigidae

Type-species: Machlotica chrysodeta Meyrick, 1909a:37, by original des-
ignation. Type locality: Bolivia: Songo [=Zongo] [holotype 9, BMNH].

Originally described in Plutellidae; subsequently transferred to Glyphip-
terygidae [sic] by Meyrick (1913b:23).

Maclotica [sic] Busck, 1915:87 Glyphipterigidae

A misspelling of Machlotica Meyrick, 1909.

Macropia Costa, [1836]: [196] Choreutidae

Type-species: Asopia incisalis Treitschke, 1829:157 [ =Tortrix nemorana
Hiibner, [ 1796-99] :pl. 1, fig. 3], by subsequent designation of Heppner
(1978b: 159). Type locality ( incisalis ): [Europe] [type lost?].

Originally described in Pyralidae; subsequently transferred to Choreuti-

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259

dae as a junior subjective synonym of Eutromula Frolich, 1828, by Heppner
(1978b: 159); now a junior subjective synonym of Choreutis Hiibner, [1825]
(Heppner, 198 la:53).

Macropia Costa, [1836], is a senior homonym of Macropia Malloch, 1930
(Diptera) (Heppner, 1978b).

Mattea Duckworth, 1966:2 Oecophoridae

Type-species: Cryptolechia phoenissa Butler, 1883:81, by original des-
ignation. Type locality: Chile: Corral [holotype 8 , BMNH].

Originally described in Oecophoridae. Mattea Duckworth, 1966, is a ju-
nior objective synonym of Tyriomorpha Meyrick, 1918, as indicated by
Clarke (1979:142), in Oecophoridae: Oecophorinae; now also a junior sub-
jective synonym of Pachyphoenix Butler, 1883, new synonymy (Becker,
pers. comm).

Melanoxena Dognin, 1910:121 Choreutidae

Type-species: Melanoxena falsissima Dognin, 1910:122, by original des-
ignation. Type locality: Colombia: San Antonio, near Cali [lectotype 8
(present designation), USNM].

Originally described in Tineidae; subsequently transferred to Glyphipte-
rygidae [sic] as a synonym of Rhobonda Walker, 1863, by Meyrick
(1913b:31); transferred to Choreutidae by Heppner (1981a:56), as a distinct
genus in Choreutidae.

Metapodistis Meyrick, 1933:372 Glyphipterigidae

Type-species: Metapodistis chrysosema Meyrick, 1933:372, by mono-
typy. Type locality: Solomon Is.: Tulagi [holotype 9, BMNH].

Originally described in Glyphipterygidae [sic]; now a junior subjective
synonym of Carmentina Meyrick, 1930 (Heppner, 1982c).

Methypsa Butler, 1875:324 *Immidae

Type-species: Hypsa saturata Walker, [ 1865] :2 17, by original designa-
tion. Type locality: [Malaysia?] [holotype 8 , BMNH].

Originally described in Lithosiidae [=Arctiidae]; subsequently transferred
to Plutellidae sensu Meyrick as a synonym of Imma Walker, [1859],
by Meyrick (1906c: 170); transferred to Glyphipterygidae [sic] as a synonym
of Imma Walker, [1859], by Meyrick (1913b:25); hereby transferred to Immi-
dae as junior subjective synonym of Birthana Walker, [1865], new synonymy.

Microaethia Chambers, 1878:76 Choreutidae

Type-species: Microaethia amphicarpeoeana Chambers, 1878:76, by
monotypy [nomen nudum, published in synonymy].

Microaethia Chambers, 1878, is not an available name as it was published
in synonymy under Brenthia Clemens, 1860, now in Choreutidae.

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Mictocommosis Diakonoff, 1977b: 8 Tortricidae

Type-species: Simaethis nigromaculata Issiki, 1930:423, by original des-
ignation. Type locality: Japan: Mt. Iwawakisan, Honshu [holotype 9 ,
USNM].

Originally described in the tribe Hilar ographini of Tortricidae: Chlidano-
tinae.

Mictopsichia Hiibner, [1825]: 3 74 Tortricidae

Type-species: Phalaena ( Tortrix ) hubneriana Stoll, 1787:41, by subse-
quent designation of Walsingham (1914:303). Type locality: [Surinam] [type
lost?].

Originally described in Tortrices Lascivae; subsequently transferred to
Glyphipteryginae [sic] of Hyponomeutidae [sic] by Walsingham (1897a:54);
transferred to Glyphipterygidae [sic] by Meyrick (19 13b: 24); transferred to
the tribe Hilarographini of Tortricidae: Chlidanotinae by Diakonoff
(1977a:76).

Mictopsychia [sic] Riley, 1889:158

A misspelling of Mictopsichia Hiibner, [1825].

Tortricidae

Mictropsichia [sic] Heppner, 1978a:53 Tortricidae

A misspelling of Mictopsichia Hiibner, [1825].

Milliera [sic] Le Marchand, 1937:192 Choreutidae

A misspelling of Millieria Ragonot, 1874.

Milliereia Spuler, 1910:298 Choreutidae

An unjustified emendation of Millieria Ragonot, 1874.

Millieria Ragonot, 1874:173 Choreutidae

Type-species: Choreutis dolosana Herrich-Schaffer, 1854:95, by original
designation. Type locality: Hungary [type lost?].

Originally described in Choreutidae.

Millieroa [sic] Le Marchand, 1937:192 Choreutidae

A misspelling of Millieria Ragonot, 1874.

Miscera Walker, 1863:457 Brachodidae

Type-species: Miscera resumptana Walker, 1863:458, by monotypy.
Type locality: Australia: Moreton Bay, Queensland [holotype 6 , BMNH].

Originally described in Choreutidae; subsequently transferred to Hypon-
omeutinae [sic] of Plutellidae by Meyrick (1907:100); transferred to Glyphip-
terygidae [sic] as a synonym of Sagalassa Walker, 1856, by Meyrick
(1913b:31); transferred to Brachodidae as a distinct genus by Heppner
(1979a: 127).

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261

Moca Walker, 1863:102 *Immidae

Type-species: Moca velutina Walker, 1863:102, by subsequent designa-
tion of Meyrick (1906c: 170). Type locality: Ceylon [=Sri Lanka] [holotype
6 , BMNH].

Originally described in Galleridae OPyralidae]; subsequently transferred
to Plutellidae sensu Meyrick by Meyrick (1906c: 170), as a synonym of
Imma Walker, [1859]; transferred to Glyphipterygidae [sic] as a synonym
of Imma Walker, [1859], by Meyrick ( 191 3b:25) ; hereby transferred to
Immidae as a distinct genus.

Mylocera Turner, 1897:27 Oecophoridae

Type-species: Mylocera tenebrifera Turner, 1897:27, by monotypy. Type
locality: Australia: Brisbane, Queensland [holotype <3, SAMA].

Originally described in Xyloryctidae; later associated with Glyphipterig-
idae; now in Oecophoridae: Xyloryctinae.

Myrsila Boisduval, [1875]:433 Glyphipterigidae

Type-species: Myrsila auripennis Boisduval, [1875]:433, by monotypy.
Type locality: Brazil: Para [holotype, UMO].

Originally described in Sesiidae; transferred to Glyphipterigidae by
Heppner (1981a:44).

Napecoetes Turner, 1913:218 * Plutellidae

Type-species: Napecoetes crossospila Turner, 1913:218, by monotypy.
Type locality: Australia: Montville, Queensland [holotype d, ANIC].

Originally described in Glyphipteryginae [sic] of Plutellidae; hereby trans-
ferred to Plutellidae.

Neomachlotica Heppner, 1981c:479 Glyphipterigidae

Type-species: Neomachlotica spiraea Heppner, 1981c:481, by original des-
ignation. Type locality: Florida: Fisheating Creek, Glades Co. [holotype
3 , USNM].

Originally described in Glyphipterigidae.

Nesotropha Turner, 1926b: 110 *Yponomeutidae

Type-species: Nesotropha pygmaeodes Turner, 1926b: 110, by monotypy.
Type locality: Australia: Cradle Mtn., Tasmania [lectotype unselected,
ANIC].

Originally described in Arctiadae [sic]; later associated with a Glyphip-
terigidae genus; hereby transferred to Yponomeutidae as a junior subjective
synonym of Tanaoctena Turner, 1913, new synonymy (Common, pers.
comm.).

Nexosa Diakonoff, 1977b: 12 Tortricidae

Type-species: Mictopsichia marmarastra Meyrick, 1932:273, by original
designation. Type locality: [Indonesia]: Java [holotype d, BMNH].

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NEW YORK ENTOMOLOGICAL SOCIETY

Originally described in the tribe Hilarographini of Tortricidae: Chlidano-
tinae.

Nigilgia Walker, 1863:511 Brachodidae

Type-species: Nigilgia adjectella Walker, 1863:512, by monotypy. Type
locality: Sierra Leone [holotype 9, BMNH].

Originally described in Tineidae; subsequently transferred to Atychiadae
[= Brachodidae] by Walsingham (1891:80), as a synonym of Phy codes Gue-
nee, 1852; transferred to Glyphipterygidae [sic] as a synonym of Phy codes
Guenee, 1852, by Meyrick (1913b:32); transferred to Brachodidae as a dis-
tinct genus by Heppner (1979a: 127).

Nigilica [sic] Turner, 1929:306 Brachodidae

A misspelling of Nigilgia Walker, 1863.

Orchemia Guenee, 1845:192 Tortricidae

Type-species: Orchemia gallicana Guenee, 1845:192, by subsequent des-
ignation of Bruand (1850a:96). Type locality: France [type, MNHP?].

Originally described in Tortrices; subsequently transferred to Glyphipter-
ygidae [sic] as a synonym of Simaethis Leach, 1815, by Meyrick (1913b:34);
currently considered to be a junior subjective synonym of Cydia Hiibner,
[1825], in Tortricidae.

Orchemia sensu Fernald, 1900:238 Choreutidae

Originally described in Choreutidae as Orchemia Guenee but with only
one included species ( Tortrix diana Hiibner, [1819-22]); now a junior sub-
jective synonym of Choreutis Hiibner, [1825], in Choreutidae.

Ordrupia Busck, 1911:228 Copromorphidae

Type-species: Ordrupia friserella Busck, 1911:228, by original designa-
tion. Type locality: French Guiana: St. Jean, Maroni River [holotype 9 ,
USNM].

Originally described in Hemerophilidae [= Choreutidae]; subsequently
transferred to Copromorphidae by Meyrick (1926:242).

Ordupia [sic] Busck, 1911:228 Copromorphidae

An original multiple spelling of Ordrupia Busck, 1911, corrected by Busck
(1912:8) and now in Copromorphidae.

Orosana Walker, 1863:458 Pyralidae

Type-species: Orosana ophideresana Walker, 1863:459, by subsequent
designation of Diakonoff (1954:285). Type locality: Australia, Hindustan,
and Ceylon [lectotype unselected, BMNH].

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263

Originally described in Choreutidae; subsequently transferred to Pyrali-
dae by Moore (1885:268) as a junior subjective synonym of Heliothela Gue-
nee, 1854, now in Pyralidae: Scopariinae.

Orosona [sic] Cotes, 1889:666 Pyralidae

A misspelling of Orosana Walker, 1863 [=Heliothela Guenee, 1854], now
in Pyralidae.

Oxytropha Diakonoff, 1954:488 Oecophoridae

Type-species: Hypertropha ametalla Turner, 1898:202, by original des-
ignation. Type locality: Australia: Armidale, New South Wales [holotype
8 , ANIC].

Originally described in Hypertrophinae of Glyphipterygidae [sic]; now in
Oecophoridae: Hypertrophinae.

Pachyphoenix Butler, 1883:81 *Oecophoridae

Type-species: Pachyphoenix sanguinea Butler, 1883:81, by monotypy.
Type locality: Chile: Corral [holotype 8, BMNH].

Originally described in Gelechiidae; subsequently associated with Gly-
phipterigidae; hereby transferred to Oecophoridae: Oecophorinae (Becker,
pers. comm.).

Palamernis Meyrick, 1906c: 205 Brachodidae

Type-species: Palamernis canonitis Meyrick, 1906c:206, by monotypy.
Type locality: India: Simla [lectotype 8 (Clarke, 1969:187), BMNH].

Originally described in Plutellidae sensu Meyrick subsequently transferred
to Glyphipterygidae [sic] by Meyrick (1913b:29); transferred to Brachodidae
by Heppner (1979a: 128), as a junior subjective synonym of Brachodes
Guenee, 1845.

Pantosperma Meyrick, 1888:89 Glyphipterigidae

Type-species: Pantosperma holochalca Meyrick, 1888:89, by monotypy.
Type locality: New Zealand: Makatoku [lectotype 8 (present designation),
BMNH].

Originally described in Glyphipterygidae [sic]; subsequently considered
a synonym of Glyphipterix Hiibner, [1825]; here considered as a distinct
genus.

Paraprays Rebel, 1910:13 Plutellidae

Type-species: Paraprays punctigera Rebel, 1910:13, by monotypy. Type
locality: [USSR]: Altai Mts. [Kirghiz SSR] [lectotype, NHMV?].

Originally described in Hyponomeutidae [sic]; subsequently transferred
to Plutellidae by Friese (1960:23); transferred to Glyphipterygidae [sic] as
a synonym of Homadaula Lower, 1899, by Clarke (1968:228); transferred
to Plutellidae as a junior subjective synonym of Homadaula Lower, 1899,
by Heppner and Dekle (1975:1).

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Penthocrates Meyrick, 1934:523 *Limacodidae

Type-species: Penthocrates bigenita Meyrick, 1934:523, by monotypy.
Type locality: [Indonesia]: Koetoardjo, Rio Awibowo, Java [holotype 9,
BMNH].

Originally described in Glyphipterygidae [sic]; transferred to Heteroge-
neidae [sic] by Meyrick (1934:523) following the original description; hereby
transferred to Limacodidae.

Peotyle Diakonoff, 1978:23 Choreutidae

Type-species: Choreutis atmodesma Meyrick, 1933:371, by original des-
ignation. Type locality: [India]: Killanmarg, Kashmir [lectotype 8 (Clarke,
1969:43), BMNH].

Originally described in Choreutidae.

Peritropha Diakonoff, 1954:278 Oecophoridae

Type-species: Peritropha oligodrachma Diakonoff, 1954:280, by original
designation. Type locality: Australia: Billopp [holotype 8 , BMNH].

Originally described in Hypertrophinae of Glyphipterygidae [sic]; now in
Oecophoridae: Hypertrophinae.

Perittia Stainton, 1854:177 Elachistidae

Type-species: Aphelosetia obscurepunctella Stainton, 1848:2164, by
monotypy. Type locality: England [type, BMNH?].

Originally described in Glyphipterygidae [sic]; subsequently transferred
to Elachistidae by Meyrick (1895:685).

Phalerarcha Meyrick, 1913a: 100 Glyphipterigidae

Type-species: Phalerarcha chrysorma Meyrick, 1913a: 101, by monotypy.
Type locality: British Guiana [= Guyana]: Bartica [holotype 9 , BMNH].
Originally described in Glyphipterygidae [sic].

Philocoristis Meyrick, 1927c: 102 *Heliodinidae

Type-species: Philocoristis catachalca Meyrick, 1927c: 102, by mono-
typy. Type locality: Samoa: Upolu, Malololelei [holotype 9, BMNH].

Originally described in Glyphipterygidae [sic], hereby transferred to He-
liodinidae.

Philodoria Walsingham, 1907:717 Gracillariidae

Type-species: Philodoria succedanea Walsingham, 1907:717, by original
designation. Type locality: Hawaii: Heleakala, Maui [lectotype unselected,
BMNH].

Originally described in Tineidae; subsequently transferred to Glyphipter-
ygidae [sic] by Meyrick (1913b:40); transferred to Gracilariadae [sic] by
Swezey (1934:524).

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265

Philpottia Meyrick, 1916:416 *Plutellidae

Type-species: Philpottia iridoxa Meyrick, 1916:417, by monotypy. Type
locality: New Zealand: Mt. Burns, Hunter Mts. [lectotype 9 , BMNH].

Originally described in Glyphipterygidae [sic]; hereby transferred to Plu-
tellidae.

Philpottia Meyrick, 1916, is a junior homonym of Philpottia Brown, 1915,
(Coleoptera) and is now replaced by Charixena Meyrick, 1920.

Phryganostola Meyrick, 1881:248 Glyphipterigidae

Type-species: Phryganostola drosophaes Meyrick, 1881:249, by subse-
quent designation of Meyrick (1914d:28). Type locality: Australia: Parra-
matta, [New South Wales] [lectotype unselected, BMNH].

Originally described in Glyphipterygidae [sic].

Phycodes Guenee, 1852:389 Brachodidae

Type-species: Phycodes hirudinicornis Guenee, 1852:389 [ =Chimaera
radiata Ochsenheimer, 1808:5], by monotypy. Type locality ( hirudinicornis ):
“Indies Orientals’ ’ [holotype 8, MNHP?].

Originally described in Hyblaeidae; subsequently transferred to Atychia-
dae [sic] [= Brachodidae] by Walsingham (1891:78); transferred to Glyphip-
terygidae [sic] by Meyrick (1913b:32); transferred to Brachodidae by
Heppner (1979a: 127).

Picrodoxa Meyrick, 1923:617 *Epermeniidae

Type-species: Picrodoxa harpodes Meyrick, 1923:617, by monotypy.

Type locality: India: Palnis [lectotype $ (Clarke, 1969:192), BMNH].

Originally described in Glyphipterygidae [sic]; hereby transferred to Eper-
meniidae.

Piestoceros Meyrick, 1907:94 *Yponomeutidae

Type-species: Incurvaria conjunctella Walker, 1863:491, by monotypy.
Type locality: Australia [holotype 8 , BMNH].

Originally described in Plutellidae; subsequently transferred to Glyphip-
terygidae [sic] by Meyrick (1913b:32); hereby transferred to Yponomeuti-
dae.

Pingrassa Walker, [1859]: 266 *Immidae

Type-species: Pingrassa accuralis Walker, [1859] :227, by monotypy.

Type locality: Ceylon [=Sri Lanka] [holotype 9, BMNH].

Originally described in Herminidae [=Noctuidae]; subsequently trans-
ferred to Plutellidae as a synonym of Imma Walker, [1859], by Meyrick
(1906c: 170); hereby transferred to Immidae as a junior subjective synonym
of Imma Walker, [1859].

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Plotheia Walker, 1863:460 Noctuidae

Type-species: Plotheia innotabilis Walker, 1863:461, by monotypy. Type
locality: Sarawak [holotype 9, BMNH].

Originally described in Choreutidae; now in Noctuidae as a junior objec-
tive synonym of Thopelia Nye, 1975:480, a replacement name.

Plotheia Walker, 1863, is a junior homonym of Plotheia Walker, [1858]
(Noctuidae).

Polygiton Diakonoff, 1955:26 Oecophoridae

Type-species: Polygiton pachypus Diakonoff, 1955:28, by original des-
ignation. Type locality: New Guinea: Araucaria Camp [holotype 6,
RMNL].

Originally described in Glyphipterygidae [sic], subfamily Hypertrophinae;
now a subfamily of Oecophoridae.

Polyphlebia Felder, 1874:8 (pi. 102, f. 38) Brachodidae

Type-species: Polyphlebia atychioides Felder, 1874:8 (plate 102, figure
38) [=Aclytia buprestoides Walker, [1865]: 101], by monotypy. Type locality
(atychioides): South America [type, NHMV?].

Originally described in Tineidae; subsequently transferred to Glyphipte-
rygidae [sic] as a synonym of Sagalassa Walker, 1856, by Meyrick
(1913b:31); transferred to Brachodidae as a distinct genus by Heppner
(1979a: 127).

Porpe Hiibner, [1825]:373 Choreutidae

Type-species: Porpe fibrana Hubner, [ 1825] :373 [= Tinea vibrana Hub-
ner, [1811-13]: plate 32, figure 202; = Tinea bjerkandrella Thunberg,
1784:24], by monotypy. Type locality (fibrana ): [Europe] [type lost?].

Originally described in Tortrices Lascivae; subsequently transferred to
Glyphipterygidae [sic] as a synonym of Choreutis Hubner, [1825], by Mey-
rick (19 13b: 38); transferred to Hemerophilidae [= Choreutidae] by Walsing-
ham (1914:309), as a distinct genus; transferred to Glyphipterigidae by Brad-
ley (1972:12) as a synonym of Tebenna Billberg, 1820; transferred to
Choreutidae as a junior subjective synonym of Tebenna Billberg, 1820, by
Heppner (1977b:635).

Probolacma Meyrick, 1927a:362 Plutellidae

Type-species: Probolacma melanoclista Meyrick, 1927a:362, by mono-
typy. Type locality: USA: Alpine, [Brewster Co.], Texas [holotype 9,
BMNH].

Originally described in Hyponomeutidae [sic]; subsequently transferred
to Glyphipterygidae [sic] by McDunnough (1939:84); transferred to Plutel-
lidae as a junior subjective synonym of Ellabella Busck, 1925, by Heppner
(1978a:50).

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267

Procerata Berthold, 1827:484 Brachodidae

Type-species: Pyralis saldonana Fabricius, 1787:232 [as “ soldana ” [sic]
Berthold] [= Sphinx appendiculata Esper, 1783:227], by monotypy. Type
locality ( saldonana ): Europe [type lost?].

Originally described in Pyrales; subsequently a nomen oblitum; treated
as a nomen oblitum in synonymy of Brachodes Guenee, 1845, in Brach-
odidae by Heppner (1981a: 13) [unused since Wocke (1871:266)].

Prochoreutis Diakonoff and Heppner, 1980:196 Choreutidae

Type-species: Pyralis myllerana Fabricius, 1794:277, by original desig-
nation. Type locality: Sweden [type lost?].

Originally described in Choreutidae as a new name for Choreutis of au-
thors [not Choreutis Hiibner, 1825].

Prochoreutis Heppner, 1981a:58 Choreutidae

Type-species: Pyralis myllerana Fabricius, 1794:277, by original desig-
nation. Type locality: Sweden [type lost?].

Redescription of Prochoreutis Diakonoff and Heppner, 1980, as a new
name for Choreutis of authors [not Choreutis Hiibner 1825].

Protosynaema Meyrick, 1886:173 Plutellidae

Type-species: Protosynaema eratopis Meyrick, 1886:174, by subsequent
designation of Fletcher (1929: 187). Type locality: New Zealand: Otira Gorge
[lectotype unselected, BMNH].

Originally described in Plutellidae and noted to be related to Glyphipteryx
[sic] Hiibner (Glyphipterigidae); now returned to Plutellidae.

Pseudotortrix Turner, 1900:15 *Immidae

Type-species: Pseudotortrix acosma Turner, 1900:16, by monotypy.
Type locality: Australia: Brisbane, Queensland [lectotype unselected,
ANIC].

Originally described in Plutellidae; subsequently transferred to Glyphip-
terygidae [sic] as a synonym of Imma Walker, [1859], by Meyrick
(1913b:25); hereby transferred to Immidae as a junior subjective synonym
of Imma Walker, [1859].

Ptochaula Meyrick, 1920b: 325 *Immidae

Type-species: Ptochaula niphadopa Meyrick, 1920b:325, by monotypy.
Type locality: [India]: Khasis, Assam [lectotype 6 (Clarke, 1969:195),
BMNH].

Originally described in Glyphipterygidae [sic]; hereby transferred to Im-
midae.

Rhabdocrates Meyrick, 1931:183 Glyphipterigidae

Type-species: Rhabdocrates sporomantis Meyrick, 1931:184, by mono-
typy. Type locality: Peru: Jurimaguas [holotype 6, BMNH].

Originally described in Glyphipterygidae [sic].

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Rhipismia Reutii, 1898:180 Choreutidae

An emendation of Ripismia Wocke, [1876] [= Millieria Ragonot, 1874].

Rhobonda Walker, 1863:424 Choreutidae

Type-species: Rhobonda gaurisana Walker, 1863:425, by monotypy.
Type locality: Brazil: Rio de Janeiro [holotype 9 , BMNH].

Originally described in Tortricidae; subsequently transferred to Glyphip-
terygidae [sic] by Meyrick (1913b:34); transferred to Hemerophilidae [=
Choreutidae] by Walsingham (1914:314); now in Choreutidae.

Rhobonda Walker, 1864:802 Gelechiidae

Type-species: Rhobonda punctatella Walker, 1864:802, by monotypy.
Type locality: Brazil: Ega [=Tefe], [Amazonas] [holotype 9, BMNH?].
Originally described in Gelechidae [sic].

Rhobonda Walker, 1864, is a junior homonym of Rhobonda Walker, 1863
(Choreutidae), and is now considered a junior subjective synonym of Di-
chomeris Hiibner, [1825], in Gelechiidae. The name Carna Walker,

1864:1038, proposed as a replacement name for Rhobonda Walker, 1864,

is a junior homonym of Carna Gistel, 1848 (Echinodermata).

Ripismia Wocke, [1876]: 3 99 * Choreutidae

Type-species: Chore utis dolosana Herrich-Schaffer, 1854:95, by mono-
typy. Type locality: Hungary [type lost?].

Originally described in Glyphipterygidae [sic]; subsequently considered
in Glyphipterygidae [sic] as a junior synonym of Millieria Ragonot, 1874,
by Meyrick (1913b:38); hereby transferred to Choreutidae as a junior sub-
jective synonym of Millieria Ragonot, 1874.

Roeslerstammia Zeller, 1839:202 Yponomeutidae

Type-species: Alucita erxlebella Fabricius, 1787:256, by monotypy. Type
locality: Germany: Gottingen [type?].

Originally described in Glossata (Alucita); subsequently transferred to
Glyphipterygidae [sic] by Stainton (1854:172); transferred to Tineidae by
Meyrick (1895:770); transferred to Acrolepiidae by Spuler (1910:454); trans-
ferred to Hyponomeutidae [sic] by Meyrick (1928:743).

Sagalassa Walker, 1856:5 Brachodidae

Type-species: Sagalassa robusta Walker, 1856:6, by subsequent desig-
nation of Meyrick ( 1914d: 15). Type locality: Brazil: Para [lectotype unse-
lected, BMNH].

Originally described in Stygiidae [=Cossidae]; subsequently transferred
to Glyphipterygidae [sic] by Meyrick (1913b:31); transferred to Brachodidae
by Heppner (1979a: 127).

Saphtha [sic] Walsingham, 1900b:567 Choreutidae

A misspelling of S apt ha Walker, 1864.

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269

Saptha Walker, 1864:1015 Choreutidae

Type-species: Saptha divitiosa Walker, 1864:1015, by monotypy. Type
locality: [Indonesia]: Ceram [holotype 9, BMNH].

Originally described in Tineidae; subsequently transferred to Plutellidae
by Meyrick (1905:610); transferred to Glyphipterygidae [sic] as a synonym
of Tortyra Walker, 1863, by Meyrick (1913b:33); transferred to Hemero-
philidae [= Choreutidae] as a synonym of Tortyra Walker, 1863, by Wal-
singham (1914:312); now a distinct genus in Choreutidae (Heppner, 1981a:55).

Scaptesylix Hampson, 1895:283 *Immidae

Type-species: Scaptesylix hemichryseis Hampson, 1895:283, by original
designation. Type locality: Burma: Donaut Range [holotype 9, BMNH].

Originally described in Zygaenidae; subsequently transferred as a syn-
onym of Imma Walker, [1859], to Plutellidae by Meyrick (1906c: 170) and
to Glyphipterygidae [sic] by Meyrick (19 13b: 25); hereby transferred to Im-
midae as a junior subjective synonym of Bursadella Snellen, 1880, new
synonymy.

Sesiomorpha Snellen, 1885:111 Zygaenidae

Type-species: Sesiomorpha abnormis Snellen, 1885:112 [= Syntomis vac-
ua Walker, [1865]:75], by monotypy. Type locality {abnormis): [Indonesia]:
Bantimoreng, Celebes [lectotype unselected, RMNL?].

Originally described in Tineidae; subsequently transferred to Glyphipte-
rygidae [sic] as a synonym of Burlacena Walker, [1865], by Meyrick
(1913b: 25); transferred to Zygaenidae as a junior subjective synonym of
Burlacena Walker, [1865], by Heppner (1981a: 15).

Setiostoma Felder and Rogenhofer, 1875:18 *Glyphipterigidae

Type-species: Setiostoma flaviceps Felder and Rogenhofer, 1875:18 (pi.
138, fig. 1), by monotypy. Type locality: Amazonas [Brazil] [lectotype un-
selected, BMNH].

Originally described in Tineae; subsequently transferred to Glyphipte-
rygidae [sic] (as Setiostoma Zeller, 1875) by Meyrick (1913b: 13); thereafter
forgotten in lieu of Setiostoma Zeller, 1875 (subsequently placed in Oeco-
phoridae: Stenomatinae); hereby placed in Glyphipterigidae as a junior sub-
jective synonym of Ussara Walker, 1864 (Becker, pers. comm.).

Setiostoma Zeller, 1875:324 Oecophoridae

Type-species: Setiostoma xanthobasis Zeller, 1875:325, by subsequent
designation of Meyrick (1914d:4). Type locality: USA: [Dallas], Texas [ho-
lotype 6 , MCZ].

Originally described in Choreutina [= Choreutidae]; subsequently trans-
ferred to Glyphipterygidae [sic] by Meyrick (1913b: 13); transferred to Ste-
nomidae by Busck (1922:279); now in the subfamily Stenomatinae of Oeco-

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phoridae as a junior objective synonym of Rectiostoma Becker, (Type-
species: Setiostoma xanthobasis Zeller, 1875 [by present designation]).

Setiostoma Zeller, 1875 (September 1875) is a junior homonym of Se-
tiostoma Felder and Rogenhofer, 1875 (June 1875) [ = Ussara Walker, 1864
(Glyphipterigidae)] .

Sezeris Walker, 1863:509 Psychidae

Type-species: Sezeris conflictella Walker, 1863:509 [=Cebysa leucotelus
Walker, 1854:486], by monotypy. Type locality ( conflictella ): Australia:
Tasmania [holotype 8, BMNH].

Originally described in Tineidae; subsequently transferred to Glyphipte-
rygidae [sic] as a synonym of Cebysa Walker, 1854, by Meyrick (1913b:32);
now considered to be a junior subjective synonym of Cebysa Walker, 1854,
in Psychidae.

Siamethis [sic] Klimesch, 1968:150 Choreutidae

A misspelling of Simaethis Leach, 1815 [—Anthophila Haworth, 1811).

Simaethis Leach, 1815:135 Choreutidae

Type-species: Tortrix dentana Hiibner, [1796-99]: plate 1, figure 4
[=Phalaena ( Tortrix ) fabriciana Linnaeus, 1767:880], by monotypy. Type
locality (dentana): [Europe] [type lost?].

Originally described in Tortricina [sic]; subsequently transferred to Py-
ralidae by Stephens (1829b: 161); transferred to Choreutidae by Walker
(1863:450); transferred to Glyphipterygidae [sic] by Meyrick (1913b:34);
transferred to Choreutidae as a junior subjective synonym of Anthophila
Haworth, [1811], by Heppner (1977b:633).

Simaetis [sic] Kautz, 1930:28 Choreutidae

A misspelling of Simaethis Leach, 1815 [= Anthophila Haworth, 1811].

Simethis [sic] Bleszynski, Razowski, Choreutidae

and Zukowski, 1965:413

A misspelling of Simaethis Leach, 1815 [= Anthophila Haworth, 1811].

Simoethis [sic] Desmarest, 1848:617 Choreutidae

A misspelling of Simaethis Leach, 1815 [= Anthophila Haworth, 1811].

Sisyroctenis Meyrick, 1936b: 106 Brachodidae

Type-species: Sisyroctenis hemicamina Meyrick, 1936b: 106, by mono-
typy. Type locality: Peru: Marcapata [holotype 9, DEI].

Originally described in Glyphipterygidae [sic]; subsequently transferred
to Brachodidae as a junior subjective synonym of Callatolmis Butler, 1877,
by Heppner (1981a: 14).

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271

Sobareutis Meyrick, 1910:469 Heliodinidae

Type-species: Sobareutis conchophanes Meyrick, 1910:470, by mono-
typy. Type locality: [Malaysia] Kuching, [Sarawak] [holotype 6, lost?].

Originally described in Glyphipterygidae [sic]; subsequently transferred
to Heliodinidae by Meyrick (1913b: 18).

(The unique holotype has not been located for study to determine if the
genus correctly belongs in Heliodinidae [it may belong to Oecophoridae:
Stathmopodini]).

Spilogenes Meyrick, 1938:19 Plutellidae

Type-species: Spilogenes chalazombra Meyrick, 1938:19, by monotypy.
Type locality: China: Likiang [lectotype 8 (Clarke, [1965]: 3 84), BMNH].

Originally described in Hyponomeutidae [sic]; subsequently transferred
to Xyloryctidae by Clarke (1955:24); transferred to Plutellidae as a junior
subjective synonym of Ellabella Busck, 1925, by Heppner (1978a:50).

Staintonia Staudinger, 1859:250 Oecophoridae

Type-species: Staintonia medinella Staudinger, 1859:250, by monotypy.
Type locality: Spain: Chiclana [lectotype unselected, ZMHB].

Originally described without family reference except as being related to
Butalis (Gelechiidae); subsequently associated with Glyphipteryx [sic] Hiib-
ner, [1825], by Erschoff (1877:347); transferred to Heliodinidae by Meyrick
(1913b: 15) as a junior subjective synonym of Eretmocera Zeller, 1852, now
a genus in Stathmopodini of Oecophoridae: Oecophorinae.

Sthenistis Hampson, 1896:541 *Immidae

Type-species: Sthenistis gyrtoniformis Hampson, 1896:541, by original
designation. Type locality: Ceylon [=Sri Lanka] [lectotype unselected,
BMNH].

Originally described in Noctuidae; subsequently transferred to Glyphip-
terygidae [sic] as a synonym of Imma Walker, [1859], by Meyrick
(1913b:25); hereby transferred to Immidae as a distinct genus.

Stichotactis Meyrick, 1930b:562 Plutellidae

Type-species: Stichotactis calamitosa Meyrick, 1930b:563, by monotypy.
Type locality: Sudan: Gendettu [lectotype 9 (Clarke, 1969:200), BMNH].

Originally described in Hyponomeutidae [sic]; subsequently transferred
to Glyphipterygidae [sic] by Clarke (1955:24); transferred to Plutellidae by
Heppner and Dekle (1975:1) as a junior subjective synonym of Homadaula
Lower, 1899.

Symaethis [sic] Bruand, 1850a: 101 Choreutidae

A misspelling of Simaethis Leach, 1815, now a synonym of Anthophila
Haworth, [1811], in Choreutidae.

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NEW YORK ENTOMOLOGICAL SOCIETY

Symphorostola Meyrick, 1927c: 3 76 Oecophoridae

Type-species: Symphorostola encomias Meyrick, 1927c:376, by mono-
typy. Type locality: [Indonesia]: Soekaranda Urwald, Sumatra [holotype
6 , BMNH].

Originally described in Glyphipterygidae [sic]; transferred to Oecophori-
dae: Xyloryctinae by Heppner (198 la: 57).

Synechodes Turner, 1913:200 Brachodidae

Type-species: Synechodes coniophora Turner, 1913:200, by monotypy.
Type locality: Australia: Kuranda, Queensland [holotype S , ANIC].

Originally described in Hyponomeutinae [sic] of Plutellidae as being re-
lated to Miscera and Tortyra; transferred to Brachodidae by Heppner
(1979a: 127).

Taeniostola Meyrick, 1920b:326 Glyphipterigidae

Type-species: Taeniostola celophora Meyrick, 1920b:327, by monotypy.
Type locality: Brazil: Rio Trombetas [holotype 9 , BMNH].

Originally described in Glyphipterygidae [sic].

Taeniostola Meyrick, 1920, is a junior homonym of Taeniostola Bezzi,
1913 (Diptera), and is now a synonym of Taeniostolella Fletcher, 1940, in
Glyphipterigidae .

Taeniostolella Fletcher, 1940:109 Glyphipterigidae

Type-species: Taeniostola celophora Meyrick, 1920b:327, by original des-
ignation. Type locality: Brazil: Rio Trombetas [holotype 9, BMNH].

Originally described in Glyphipterygidae [sic] as a replacement name for
Taeniostola Meyrick, 1920.

Taleporia Hiibner, [1825]: 400 Psychidae

Type-species: Tinea pseudobombycella Hiibner, 1796:pl. 31, fig. 212, by
subsequent designation of Fletcher (1929:215). Type locality: [Europe] [type
lost?].

Originally described in Tineae; subsequently considered in Glyphipteryg-
idae [sic] by Meyrick (1912b: 123); currently in Psychidae.

Tebenna Billberg, 1820:90 Choreutidae

Type-species: Tinea bjerkandrella Thunberg, 1784:24, by subsequent des-
ignation of Bradley (1966b:220). Type locality: Sweden [type, Univ.
Uppsala].

Originally described in Tortrices; subsequently transferred to Glyphipte-
rigidae by Bradley (1966b: 220); transferred to Choreutidae by Heppner
(1977b:632).

Tebenna Hiibner, [1825]:414 Agonoxenidae

Type-species: Tinea festivella [Denis and Schiffermiiller] , 1775:319, by
subsequent designation of Fletcher (1929:216). Type locality: [Austria]: Vi-
enna [type lost?].

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273

Originally described in Tineae Incertae; now a junior subjective synonym
of Heinemannia Wocke, 1853, in Agonoxenidae: Blastodacninae.

Tebenna Hiibner, [1825], is a junior homonym of Tebenna Billberg, 1820
(Choreutidae).

Tebeuna [sic] Danilevsky, 1969:922 Choreutidae

A misspelling of Tebenna Billberg, 1820, in Choreutidae.

Tegna Walker, 1866:1809 Brachodidae

Type-species: Tegna hyblaeella Walker, 1866:1810 [ =Chimaera radiata
Ochsenheimer, 1808:5], by monotypy. Type locality ( hyblaeella ): Nepal
[lectotype unselected, BMNH].

Originally described in Choreutidae; subsequently transferred to Glyphip-
terygidae [sic] as a synonym of Phy codes Guenee, 1852, by Meyrick
(1913b:32); transferred to Brachodidae by Heppner (1981a: 14) as a junior
subjective synonym of Phy codes Guenee, 1852.

Tetracmanthes Meyrick, 1925a: 136 Glyphipterigidae

Type-species: Tetracmanthes astrocosma Meyrick, 1925a: 136, by mono-
typy. Type locality: [South Africa]: Weenan, Natal [holotype 8 , BMNH].
Originally described in Glyphipterygidae [sic].

Tharmatographa [sic] Diakonoff, 1977b:51 Tortricidae

A misspelling of Thaumatographa Walsingham, 1897, now in the tribe
Hilarographini of Tortricidae: Chlidanotinae.

Thaumatographa Walsingham, 1897:52 Tortricidae

Type-species: Hilarographa zapyra Meyrick, 1886:286, by original des-

ignation. Type locality: [Papua]: Port Moresby, New Guinea [holotype 8 ,
BMNH].

Originally described in Glyphipteryginae [sic] of Hyponomeutidae [sic];
subsequently transferred to Glyphipterygidae [sic] as a synonym of Hilar-
ographa Zeller, 1877, by Meyrick (1913b:24) transferred to the tribe Hilar-
ographini of Tortricidae: Chlidanotinae by Diakonoff (1977b:22).

Thaumatotibia Zacher, 1915:529 Tortricidae

Type-species: Thaumatotibia roerigii Zacher, 1915:529 [=Argyroploce
leucotreta Meyrick, 1913], by monotypy. Type locality (roerigii): Togo:
Sokode, Bassari [type lost?].

Originally described without family placement; subsequently transferred
to Tortricidae: Olethreutinae by Heppner (1980:334), as a junior subjective
synonym of Cryptophlebia Walsingham, 1899.

Thelethia Dyar, 1893:301 Incurvariidae

Type-species: Thia extranea H. Edwards, 1888:181, by original designa-
tion. Type locality: USA: Los Angeles, California [lectotype 8 (Davis,
1967:44), INHS].

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NEW YORK ENTOMOLOGICAL SOCIETY

Originally described in Tineidae as a replacement name for Thia H. Ed-
wards, 1888; subsequently transferred to Glyphipterygidae [sic] by Meyrick
(1913b:29); transferred to Incurvariidae as a junior subjective synonym of
Tegeticula Zeller, 1873, by Davis (1967:29).

Thia H. Edwards, 1888:181 Incurvariidae

Type-species: Thia extranea H. Edwards, 1888:181, by monotypy. Type
locality: USA: Los Angeles, California [lectotype 6 (Davis, 1967:44),
INHS].

Originally described in Heterogynidae; subsequently transferred to Gly-
phipterygidae [sic] in synonymy with Thelethia Dyar, 1893, by Meyrick
(1913b:29); transferred to Incurvariidae as a junior subjective synonym of
Tegeticula Zeller, 1873, by Davis (1967:29).

Thia H. Edwards, 1888, is a junior homonym of Thia Leach, 1815 (Crus-
tacea), Thia Oken, 1815 (Polychaeta), and Thia Newman, 1840 (Coleop-
tera).

Thrasydoxa Meyrick, 1912a:60 Heliodinidae

Type-species: Thrasydoxa tyrocopa Meyrick, 1912a:60, by monotypy.
Type locality: Colombia: San Antonio [holotype 6 , BMNH].

Originally described in Glyphipterygidae [sic]; subsequently transferred
to Heliodinidae by Meyrick (1913b: 18).

Thriambeutis Meyrick, 1910:470 Heliodinidae

Type-species: Thriambeutis hemicausta Meyrick, 1910:470, by mono-
typy. Type locality: Solomon Is.: Isabel Id. [holotype 9, BMNH].

Originally described in Glyphipterygidae [sic]; subsequently transferred
to Heliodinidae by Meyrick (1913b: 19).

Thylacopleura Meyrick, 1886:284 *Immidae

Type-species: Thylacopleura autodoxa Meyrick, 1886:285, by monotypy.
Type locality: Fiji [lectotype unselected, BMNH].

Originally described in Glyphipterygidae [sic]; subsequently transferred
as a synonym of Imma Walker, [1859], to Plutellidae by Meyrick
(1906c: 170); hereby transferred to Immidae as a junior subjective synonym
of Imma Walker, [1859].

Tinagma Zeller, 1839:203 Douglasiidae

Type-species: Tinagma perdicellum Zeller, 1839:204, by subsequent des-
ignation of Fletcher (1929:223). Type locality: [Germany]: Spitzberg [lec-
totype unselected, BMNH].

Originally described in Tineana; subsequently transferred to Glyphipte-
rygidae [sic] by Stainton (1854:178); transferred to Douglasiidae by Meyrick
(1928:721).

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275

Topaza Walker, 1864:808 *Immidae

Type-species: Topaza alienella Walker, 1864:808, by monotypy. Type
locality: [Malaysia]: Sarawak [holotype 8, BMNH].

Originally described in Gelechidae [sic]; subsequently transferred as a
synonym of Imma Walker, [1859], to Plutellidae by Meyrick (1906c: 170)
and to Glyphipterygidae [sic] by Meyrick (1913b:25); hereby transferred to
Immidae as a junior subjective synonym of Imma Walker, [1859].

Tortricomorpha C. Felder, 1861:34 *Immidae

Type-species: Tortricomorpha atrosignata C. Felder, 1861:35, by subse-
quent designation of Meyrick (1906c: 170). Type locality: [Indonesia]: Am-
boina, [Moluccas Is.] [holotype $, BMNH].

Originally described in Arctiidae; subsequently transferred to Atychiadae
[sic] [=Brachodidae] by Lower (1903:69); transferred to Plutellidae by
Meyrick (1905:611); transferred to Glyphipterygidae [sic] as a synonym of
Imma Walker, [1859], by Meyrick (1913b:25); hereby transferred to Immi-
dae as a junior subjective synonym of Imma Walker, [1859].

Tortyra Walker, 1863:510 Choreutidae

Type-species: Tortyra spectabilis Walker, 1863:510, by subsequent des-
ignation of Busck (1914:57). Type locality: Brazil: Ega [=Tefe] [lectotype
8 (present designation), BMNH].

Originally described in Tineidae; subsequently transferred to Glyphipte-
rygidae [sic] by Meyrick (1910:462); transferred to Choreutidae by Heppner
(1977b:633).

Trapeziophora Walsingham, 1892:529 Glyphipterigidae

Type-species: Trapeziophora gemmula Walsingham, 1892:530, by origi-
nal designation. Type locality: St. Vincent [lectotype 8 (present designa-
tion), BMNH].

Originally described in Glyphipteryginae [sic] of Tineidae; subsequently
transferred to Glyphipterygidae [sic] by Meyrick (1913b:40) as a synonym
of Ussara Walker, 1864; here considered a distinct genus.

Trichothyrsa Meyrick, 1912a:61 Heliodinidae

Type-species: Trichothyrsa flammivola Meyrick, 1912a:61, by original
designation. Type locality: India: Coorg [holotype 8 , BMNH].

Originally described in Glyphipterygidae [sic]; subsequently transferred
to Heliodinidae by Meyrick (1913b: 19).

Tyriomorpha Meyrick, 1918:191 Oecophoridae

Type-species: Cryptolechia phoenissa Butler, 1883:81, by original des-
ignation. Type locality: Chile: Corral [holotype 8 , BMNH].

Originally described in Glyphipterygidae [sic]; subsequently transferred

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NEW YORK ENTOMOLOGICAL SOCIETY

to Oecophoridae as Matte a Duckworth, 1966, a junior objective synonym
of Tyriomorpha Meyrick, 1918; Clarke (1979:142) transferred Tyriomorpha
to Oecophoridae: Oecophorinae, now a junior subjective synonym of Pa-
chyphoenix Butler, 1883, new synonymy (Becker, pers. comm.).

Usara [sic] Busck, [1934a]: 182 Glyphipterigidae

A misspelling of Ussara Walker, 1864.

Ussara Walker, 1864:800 Glyphipterigidae

Type-species: Ussara decoratella Walker, 1864:801, by monotypy. Type
locality: Brazil: Ega [=Tefe] [holotype 9, BMNH].

Originally described in Gelechidae [sic]; subsequently transferred to Gly-
phipterygidae [sic] by Meyrick (1913b:40).

Vinzela Walker, [1866]: 1260 *Immidae

Type-species: Vinzela inaptalis Walker, [1866]: 1261, by monotypy. Type
locality: [Malaysia]: Sarawak [holotype 6, UMO].

Originally described in Pyralidae; subsequently transferred to Gelechia-
dae [sic] as a synonym of Tortricomorpha C. Felder, 1861, by Walsingham
(1900a:78); transferred as a synonym of Imma Walker, [1859], to Plutellidae
by Meyrick (1906c: 170) and to Glyphipterygidae [sic] by Meyrick
(1913b:25); hereby transferred to Immidae as a junior subjective synonym
of Imma Walker, [1859].

Walsinghamia Riley, 1889:157 Choreutidae

Type-species: Walsinghamia diva Riley, 1889:158, by monotypy. Type
locality: USA: Cocoanut Grove, [Miami], Florida [lectotype $ (present des-
ignation), USNM].

Originally described in Noctuidae; subsequently transferred to Glyphip-
terygidae [sic] as a synonym of Tortyra Walker, 1863, by Meyrick
(1913b:33); transferred to Choreutidae as a junior subjective synonym of
Hemerophila Hiibner, [1817], by Heppner (1977b:634).

Xylopoda Latreille, 1829:412 Choreutidae

Type-species: Tortrix dentana Hiibner, [1796-99]:pl. 1, fig. 4 [ =Phalaena
( Tortrix ) fabriciana Linnaeus, 1767:880], by monotypy. Type locality ( den-
tana): [Europe] [type lost?].

Originally described in Tortrices; subsequently transferred to Glyphipte-
rygidae [sic] as a synonym of Simaethis Leach, 1815, by Meyrick
(19 13b: 34); transferred to Choreutidae as a junior subjective synonym of
Anthophila Haworth, [1811], by Heppner (1981a:50).

Xylopoda Latreille, 1829, is a junior homonym of Xylopoda Berthold,
1827.

Xylopoda Berthold, 1827:484 Choreutidae

Type-species: Tortrix dentana Hiibner, [ 1 796—99] :pl . 1, fig. 4 [=Pha-

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277

laena (Tortrix) fabriciana Linnaeus, 1767:880], by monotypy. Type locality
( dentana ): [Europe] [type lost?].

Originally described in Tortrices; subsequently forgotten in lieu of Xy-
lopoda Latreille, 1829, in Glyphipterygidae [sic] and as a synonym of Si-
maethis Leach, 1815, by Meyrick (1913b:34); now a junior subjective syn-
onym of Anthophila Haworth, [1811].

Xylopoda Berthold, 1827, was published ahead of Latreille (1829) in a
German edition of Latreille’ s work, with additions by Berthold.

Xylopode Latreille, 1825:476 Choreutidae

A vernacular spelling, later corrected to Xylopoda Latreille, 1829; now
an invalid name under Anthophila Haworth, [1811].

Xylopodo [sic] Morris, 1872:iv Choreutidae

A misspelling of Xylopoda Latreille, 1829 [=Xylopoda Berthold, 1827],
now a junior subjective synonym of Anthophila Haworth, [1811], in Choreu-
tidae.

Zodia Heppner, 1979b:685 Choreutidae

Type-species: Simaethis plutusana Walker, 1863:453, by original desig-
nation. Type locality: Brazil: Ega [=Tefe] [lectotype 6 (Heppner,
1979b: 689), BMNH].

Originally described in Choreutidae.

Lectotype Designations

Specimens of species listed below have been examined and are selected
as lectotypes by present designation. The slash mark (/) indicates separate
lines on labels; each separate label is distinguished by a semicolon. The
notations “Durrant label” below indicate large black-bordered labels at-
tached to specimens by Durrant at the British Museum (Natural History)
on specimens considered the type or on syntypes.

Abrenthia cuprea Busck, 1915:87 Glyphipterigidae

Lectotype 8 [USNM] with the following labels: Roxboro/VI. 27 Pa.; Coll.
by/F. Haimbach; Type/No. 19239/U.S.N.M. [red label]; Abrenthia/cuprea/
Type Busck; Genitalia Slide/By Heppner ’76/ USNM 77247 8 [green label].

Paralectotypes: 2 — Roxboro, Pennsylvania (1 c5); Falls Church, Virginia
(1 8) [USNM].

Agiton idioptila Turner, 1926a: 145 Epermeniidae

Lectotype 8 [ANIC] with the following labels (teste Common): National
Pk., Q., 47, 1-1-21; Agiton idioptila Turn./TYPE.

Paralectotypes: 3 — [Lamington] National Pk., Queensland], Australia (2
d,19) [ANIC].

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NEW YORK ENTOMOLOGICAL SOCIETY

Araeolepia subfasciella Walsingham, 1881:303 Plutellidae

Lectotype 6 [BMNH] with the following labels: Currant Creek/Grant Co./
OREGON/16. IV. 1872 Wlsm.; Walsingham/Collection/19 10-427; B.M. 61
Genitalia Slide/No. 20209.

Paralectotypes: 16 — (same locality) (4 6,2 9) [BMNH]; 2 $ (ex Fernald
Coll.) [USNM]; 1 S (ex Chambers Coll.) [MCZ]; 7 not located.

Badera pretiosa Walker, 1866:1819 Choreutidae

Lectotype $ [BMNH] with the following labels (teste Sattler): Type
[round, red edge]; 60- 1 5/E. I. C.; 771. [small square]; Type S [Durrant label],
[abdomen missing].

Paralectotypes: 1 S — Java [BMNH].

Callizyga dispar Turner, 1894:132 Oecophoridae

Lectotype 6 [ANIC] with the following labels (teste Common): Brisbane;
Callizyga dispar/ Turn./ TYPE; Genitalia Slide; G463.

Paralectotypes: 1 9 — Brisbane [Queensland, Australia] [ANIC].

Choregia fulgens Felder and Rogenhofer, 1875:6 Choreutidae

Lectotype 6 [BMNH] with the following labels: Type [round, red edge];
312 [small square]; Felder Coll. /Rothschild/1913-86; Novara CXL/f. 17.
Choregia/fulgens F. /Bogota S [Felder label]; [blank Durrant label] TYPE;
FELDER’S TYPE [reversed]. [S abdomen glued on is incorrect one].

Paralectotypes: none [2 syntypes are from Brazil (Cuyabe [NHMV]) but
belong to a different species of Tortyra ].

Circica cionophora Meyrick, 1888:88 Glyphipterigidae

Lectotype 6 [BMNH] with the following labels: Christchurch/New Zea-
land/24-11-1882/Meyrick 1888/1740; Walsingham/Collection/19 10-427; Circi-
ca/cionophora Meyr. /Named by Meyrick; B.M. 6 /Genitalia Slide/No.
20236.

Paralectotypes: 12 — (same locality, 24.2.82) (11 6, 1 9) [BMNH].

Ditrigonophora marmoriepennis Walsingham, 1897:118 Yponomeutidae
Lectotype [sex unknown] [BMNH] with the following labels: Balthazar/
Grenada.

Paralectotypes: 1 [BMNH]. [Both syntypes are without heads or abdo-
mens.]

Epithetica typhoscia Turner, 1923:165 Oecophoridae

Lectotype 6 [ANIC] with the following labels (teste Common): Lismore,
N.S.W./22-10-22; Epithetica/typhoscia Turn. /TYPE.

Paralectotypes: 1 6 — (same locality and date) [ANIC].

Euprophantis autoglypta Meyrick, 1921a: 191 Gracillariidae

Lectotype 6 [RMNL] with the following labels (teste Diakonoff): Java/

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279

Pekalongan/v. Deventer; Coll. Piepers/Snellen, Java; [genitalia mini-slides]
2122 & 2122a; Type [red label].

Paralectotypes: 2 (same locality) [specimens not located].

Idiothauma africanum Walsingham, 1897:49 Tortricidae

Lectotype 8 [BMNH] with the following labels: Type [round, red’ edge];
Kangwe/Ogowe River/FRENCH CONGO/Good-Holland, 6967; Walsing-
ham/Collection/ 19 10-427; [Durrant label] TYPE 8; B.M. 8 /Genitalia Slide/
No. 20233.

Paralectotypes: 2 — (same locality) [specimens not located; not at
BMNH].

Melanoxena falsissima Dognin, 1910:121 Choreutidae

Lectotype 8 [USNM] with the following labels: Colombie/Fassel; Melan-
oxena/falsissima/Dgn./type 8 ; Durrant/12.6.12; 6554/Wlsm. 1911; “8/4.08 S.
Antonio” [folded]; (Melanoxena)/gen. nov./(falsissima)/sp. nov. /Warren
1.7.09; Dognin/Collection ; Type No. /32355/U.S.N.M. [red label].
Paralectotypes: 28 — (same locality) [USNM].

Pantosperma holochalca Meyrick, 1888:89 Glyphipterigidae

Lectotype 8 [BMNH] with the following labels: Makotaku/ N. Zealand /
8-III-1883/Meyrick 1888, 1691; Walsingham/Collection/ 19 10-427; Pantosper-
ma/holochalca Meyr. /Named by Meyrick; B.M. 8/ Genitalia Slide/No.
20238.

Paralectotypes: 11 — (same locality) 10 8, 1 9 [BMNH].

Sciaphila trigonana Walsingham, 1879:22 Copromorphidae

Lectotype 9 [BMNH] with the following labels: N. nr. Mendocino City /
CALIFORNIA/3-5 VI. 1871/Wlsm. 91861; Brit. Mus./1880-75; [Durrant
label] TYPE 8 [sic]; B.M. 9/Genitalia Slide/No. 20212.

Paralectotypes: 5 — (same locality) 1 8 , 4 9 [BMNH].

Simaethis pronubana Snellen, 1877:48 Choreutidae

Lectotype 9 [RMNL] with the following labels (teste Diakonoff): Java [in
Pieper’s handwriting] [abdomen missing].

Paralectotypes: 4 — (same locality) [specimens not located; not at RMNL].

Sphinx appendiculata Esper, 1783:227 Brachodidae

Lectotype 9 [SMW] with the following labels: [no data; Coll. Gerning].
Paralectotypes: 3 9 — [no data; Coll. Gerning]. (The Gerning Collection
dates from the late 1700’s to early 1800’s and contains the remaining Esper
types.)

Tortyra spectabilis Walker, 1863:510 Choreutidae

Lectotype 8 [BMNH] with the following labels: Ega/57, 125 [reverse]
[blue, round label]; [Durrant label] Tortyra spectabilis Wkr./“28/510 e”/
PARATYPE “5/6”; B.M. /Genitalia Slide/No. 20223.

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NEW YORK ENTOMOLOGICAL SOCIETY

Paralectotypes: 3 $ — (same locality [Tefe, Amazonas, Brazil]) [BMNH].

[Two other syntypes belong to another species of Tortyra .]

Trapeziophora gemmula Walsingham, 1892:530 Glyphipterigidae

Lectotype 6 [BMNH] with the following labels: Type H.T. [round, red

edge]; St. Vincent, W.I./H.H. Smith; Walsingham/Collection/19 10-427/

65356; [Durrant label] TYPE 9 [sic]; B.M. 6 /Genitalia Slide/No. 20235.
Paralectotypes: 1 6 — “Windward side/St. Vincent, W.I./H.H. Smith”;

(Walsingham Coll. 65355) [BMNH].

Walsinghamia diva Riley, 1889:158 Choreutidae

Lectotype 9 [USNM] with the following labels: Type/No. 400/U.S.N.M.

[red]; Walsinghamia/diva/Type Riley, [abdomen missing].

Paralectotypes: 2 — (same locality) [Cocoanut [sic] Grove, Florida], (1

cJ, 1 9) [USNM].

Literature Cited

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Lederer, J. 1853. Lepidopterologisches aus Sibirien. Verh. Zool.-Bot. Ver. Wien 3:351-386.
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Linnaeus, C. 1758. Systema naturae per regna tria naturae, secundum classes, ordines,
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. 1767. Ibid. (12th ed.). Vol. 1, Pt. 2. Laurentii Salvii, Holmiae [Stockholm]. Pp. 533—

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Lower, O. B. 1899. Descriptions of new Australian Lepidoptera. Proc. Linn. Soc. N. S. W.
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. 1903. Descriptions of new Australian Noctuina, &c. Trans. Roy. Soc. So. Austr.

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Mann, J. and A. F. Rogenhofer. 1878. Zur Lepidopteren-Fauna des Dolomiten-Gebietes.
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. 1886. Descriptions of Lepidoptera from the South Pacific. Trans. Ent. Soc. Lond.

1886:189-296.

. 1888. Descriptions of New Zealand Tineina. Trans. New Zealand Inst. 20:77-106.

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Department of Entomology, Smithsonian Institution, Washington, D.C.
20560.

Received for publication March 10, 1981.

292

NEW YORK ENTOMOLOGICAL SOCIETY

Index to Species

abnormis Snellen, Sesiomorpha
abroniaeella Chambers, Lithariarpteryx
abruptaria (Thunberg), Hemerophila
accuralis Walker, Pingrassa
acosma Turner, Pseudotortrix
adjectella Walker, Nigilgia
aegerioides Walker, Burlacena
aequalis Walker, Gora
africanum Walsingham, Idiothauma
agathoclea Meyrick, Homoplastis
aglaograpta Meyrick, Irinympha
albella Zeller, Calantica
albertiana (Cramer), Gauris
albertiana (Cramer), Hemerophila
albodiscata Walker, Adricara
alienella Walker, Topaza
ambrosia Meyrick, Epicroesa
ametalla (Turner), Oxytropha
amphicarpeoeana Chambers, Microaethia
apocynoglossa (Heppner), Caloreas
appendiculata (Esper), Atychia
appendiculata (Esper), Chimaera
appendiculata (Esper), Procerata
argopasta Turner, Amphimelas
argyrosema Meyrick, Apistomorpha
astrocosma Meyrick, Tetracmanthes
atmodesma (Meyrick), Peotyle
atrosignata Felder, Tortricomorpha
atychioides Felder, Polyphlebia
aurantiaca (Semper), Hyperperissa
auripennis Boisduval, Myrsila
autodoxa Meyrick, Thylacopleura
autoglypta Meyrick, Euprophantis
autumnitella Curtis, Acrolepia
bergstraesserella (Fabricius), Glyphipterix
bigenita Meyrick, Penthocrates
bisignatella (Walker), Inapha
bjerkandrella (Thunberg), Porpe
bjerkandrella (Thunberg), Tebenna
buprestoides (Walker), Polyphlebia
calamitosa Meyrick, Stichotactis
canonitis Meyrick, Palamernis
catachalca Meyrick, Philocoristis
celophora Meyrick, Taeniostola
celophora Meyrick, Taeniostolella
chalazombra Meyrick, Spilogenes
chimaera (Hiibner), Atychia
chimaera (Hiibner), Chimaera

chlorospora Meyrick, Ernolytis
chrysodeta Meyrick, Machlotica
chrysorma Meyrick, Phalerarcha
chrysosema Meyrick, Metapodistis
cionophora Meyrick, Circica
clavaria Meyrick, Chrysocentris
coleoptrata (Walker), Callatolmis
collina Turner, Cylicophora
combinatana (Walker), Asterivora
compulsana Walker, Jonaca
conchophanes Meyrick, Sobareutis
conflict ell a Walker, Sezeris
coniophora Turner, Synechodes
conjunctella (Walker), Piestoceros
consocia Walker, Birthana
coronias Meyrick, Corsocasis
crossospila Turner, Napecoetes
cuprea Busck, Abrenthia
dealbatella Zeller, Calantica
decoratella Walker, Ussara
dentana (Hiibner), Simaethis
dentana (Hiibner), Xylopoda
dentella Zeller, Aechmia
denticulella (Thunberg), Callisto
diana (Hiibner), Allononyma
diana (Hiibner), Hemerophila
diana (Hiibner), Orchemia
dichroalis Snellen, Bursadella
dispar Turner, Callizyga
diva Riley, Walsinghamia
divitiosa Walker, Saptha
dolopis Durrant, Cibdeloses
dolosana (Herrich-Schaffer), Millieria
dolosana (Herrich-Schaffer) Ripismia
drosophaes Meyrick, Phyrganostola
editha Busck, Ellabella
electrodes Diakonoff, Cercosimma
encomias Meyrick, Symphorostola
eratopis Meyrick, Protosynaema
erxlebella (Fabricius), Roeslerstammia
extrahea (H. Edwards), Thelethia
extranea H. Edwards, Thia
fabriciana (Linnaeus), Anthophila
fabriciana (Linnaeus), Simaethis
fabriciana (Linnaeus), Xylopoda
fabricii Haworth, Anthophila
falsissima Dognin, Melanoxena
fervescens Meyrick, Amphiclada

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293

festivella (D. & S.), Tebenna
fibrana Hiibner, Porpe
flammivola Meyrick, Trichothyrsa
flaviceps (F. & R.), Setiostoma
forsterella (Fabricius), Heribeia
friserella Busck, Ordrupia
fueslella (Fabricius), Aechmia
fulgens (Erschoff), Desmidoloma
fulgens F. & R., Choregia
funebris (Feisthamel), Brachodes
fyeslella [sic] (Fabricius), Aechmia
gallicana Guenee, Orchemia
gaurisana Walker, Rhobonda
gemmula Walsingham, Trapeziophora
granulata Meyrick, Cronicombra
gut tea (Haworth), Callisto
gyrtoniformis Hampson, Sthenistis
halticella Eaton, Embryonopsis
harp odes Meyrick, Picrodoxa
hemicamina Meyrick, Sisyroctenis
hemicausta Meyrick, Thriambeutis
hemichryseis Hampson, Scaptesylix
heptachalca Meyrick, Hoplophractis
hesperialis Walker, Dudua
hirudinicornis Guenee, Phy codes
holochalca Meyrick, Pantosperma
hubneriana (Stoll), Mictopsichia
hyblaeella Walker, Tegna
ichthyopa Meyrick, Iridostoma
idioptila Turner, Agiton
impigritella (Clemens), Diploschizia
inaptalis Walker, Vinzela
incisalis (Treitschke), Macropia
innotabilis Walker, Plotheia
iophanes Meyrick, Hierodoris
iridesma Meyrick, Carmentina
iridoxa (Meyrick), Charixena
iridoxa Meyrick, Philpottia
iridozona Meyrick, Aeolocosma
irina Meyrick, Actinoscelis
iuloptera Meyrick, Ereunetis
japonica (Issiki), Litobrenthia
lampronialis Walker, Inapha
lasiochroa Lower, Homadaula
lasiopa Meyrick, Colpotorna
leucotelus Walker, Cebysa
leucotelus Walker, Sezeris
lineana (Hiibner), Glyphipterix
linneella (Clerck), Glyphipteryx
linneella Hiibner, Glyphipterix
lutumella Amsel, Klimeschia

mackwoodii Moore, Aprata
mackwoodii Moore, Davendra
marmarastra (Meyrick), Nexosa
marmoreipennis Walsingham, Ditrigonophora
mathewi Zeller, Heliostibes
mediana Walker, Cotaena
medinella Staudinger, Staintonia
melanochlaena Turner, Aproopta
melanoclista Meyrick, Probolacma
melistoma Meyrick, Lasiodictis
metallicella (Duponchel), Aechmia
micaceella Walker, Chalenata
mikadonis (Stringer), Charitographa
myllerana (Fabricius), Choreutis
myllerana (Fabricius), Prochoreutis
myriospila Meyrick, Homadaula
nemorana (Hiibner), Entomoloma
nemorana (Hiibner), Macropia
nigromaculata (Issiki), Mictocommosis
niphadopa Meyrick, Ptochaula
obscurepunctella (Stainton), Perittia
ocnerostomellum (Stainton), Douglasia
oligodrachma Diakonoff, Peritropha
ophideresana Walker, Orosana
pachypus Diakonoff, Polygiton
palaeodes (Meyrick), Alampla
pariana (Clerck), Choreutis
pariana (Clerck), Eutromula
pariana (Clerck), Hemerophila
pavonacella Clemens, Brenthia
percussana (Walker), Allotropha
perdicellum Zeller, Tinagma
perornatella (Walker), Desmidoloma
perornatella (Walker), Lepidotarphius
phoenissa (Butler), Mattea
phoenissa (Butler), Tyriomorpha
planipes Diakonoff, Ganabalia
plumbigera Meyrick, Encratora
plutostola Diakonoff, Embolostoma
plutusana (Walker), Zodia
pretiosa Walker, Badera
pronubana (Snellen), Chordates
pseudobombycella (Hiibner), Taleporia
punctatella (Walker), Carna
punctatella Walker, Rhobonda
punctigera Rebel, Paraprays
purpurascens Hampson, Callartona
purpurata Meyrick, Lamprystica
purpurina (D. & S.), Anthophila
purpurina (D. & S.), Anthophilae
pygmaeodes Turner, Nesotropha

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pygmeana (Haworth), Acrolepia
pyractis Meyrick, Archimaga
radiata Ochsenheimer, Chimaera
radiata (Ochsenheimer), Phy codes
radiata (Ochsenheimer), Tegna
resumptana Walker, Miscera
robusta Walker, Sagalassa
roerigii Zacher, Thaumatotibia
rugosalis Walker, Imma
saldonana (Fabricius), Procerata
sanguinea Butler, Pachyphoenix
sapphiropa Meyrick, Irianassa
satrapella Meyrick, Eupselia
saturata (Walker), Methypsa
semilinea Walker, Jobula
sepias Meyrick, Loxotrochis
sexfasciella Sauber, Choreutidia
silvana Meyrick, Bryonympha
simpliciella (Stephens), Anacampsoides
soldana [sic] Berthold, Procerata
spectabilis Walker, Tortyra
spiraea Heppner, Neomachlotica
splendens Pryer, Lepidotarphius
sporimaea Meyrick, Lygronoma
sporomantis Meyrick, Rhabdocrates
statices (Linnaeus), Atychia

Stella Meyrick, Coridomorpha
subfasciella Walsingham, Araeolepia
succedanea Walsingham, Philodoria
swederiana (Stoll), Hilarographa
tenebrifera Turner, Mylocera
thesaurella Meyrick, Hypertropha
thiolychna Meyrick, Electrographa
thrasonella (Scopoli), Aechmia
tinthalea (Meyrick), Amphimelas
trigonana (Walsingham), Lotisma
typhoscia Turner, Epithetica
tyrocopa Meyrick, Thrasydoxa
vacua (Walker), Sesiomorpha
valida (Walker), Jonaca
velutina Walker, Moca
vernetella Guenee, Brae bodes
vexatalis Walker, Alicadra
vibrana (Hiibner), Porpe
vicarialis Zeller, Hemerophila
xanthobasis Zeller, Setiostoma
xanthoplaca Turner, Euthorybeta
xanthoprocta (Meyrick), Atractoceros
xylophragma Meyrick, Antispastis
zapyra Meyrick, Hilarographa
zapyra (Meyrick), Thaumatographa

NEW YORK ENTOMOLOGICAL SOCIETY
LXXXIX(4), 1981, pp. 295-331

STUDIES OF NORTH AMERICAN ERORA (SCUDDER)
(LEPIDOPTERA, LYCAENIDAE)

Alexander B. Klots1 and Cyril F. dos Passos2

Abstract. — The genus Erora Scudder is characterized and discussed.
Characteristics of E. laeta (Edwards), E. quaderna quaderna (Hewitson)
and E. quaderna sanfordi dos Passos are given and evaluated. The early
stages, ethology, ecology and geographic distributions of E. laeta and E.
quaderna sanfordi are described. The peculiar dense vesture of the larvae,
and a “bald” area (the calvarium) on the prothorax, are potentially signif-
icant in generic taxonomy. Full, annotated synonymies and lists of distri-
butional records are given. Possible phytogenies are discussed.

Introduction

For many years the authors pursued studies of the North American Erora,
first of E. laeta and more recently of E. quaderna sanfordi. The work has
consisted of three main parts: field studies of the life histories, ecology and
ethology; compilations of records and data from museum specimens, the
published literature and personal communications; and comparative taxo-
nomic studies. This article deals with these taxa in North America north of
Mexico; but it also contains some notes and comments on the still little-
known E. q. quaderna and E. caudata Miller in Mexico.

Acknowledgments

We are greatly indebted to many persons for information and assistance.
From John H. Cook, A. C. Frederick and James Sanford we learned about
the two localities in Vermont where most of our eastern field studies were
made. The late Sidney Hessel furnished notes and records, and generously
refrained from collecting in the areas where we were doing life-history work.
(What greater sacrifice can a devoted collector make?) Joseph J. Copeland
checked many plant identifications. T. G. Howarth of the British Museum
(Natural History) picked out and dissected for our study the types of E.
attalion (Godman and Salvin), E. quaderna (Hewitson) and possibly related
species. W. D. Field of the U.S. National Museum gave us many data about
genitalic structures and neotropical species. Harry Clench of the Carnegie

1 Research Associate, American Museum of Natural History.

2 Research Associate, American Museum of Natural History and Carnegie Museum.

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Museum very kindly gave us transcripts of his and Lee D. Miller’s field data
on E. quaderna in Mexico, and data on the differentiation of E. q. quaderna
and E. q. sanfordi. A. C. Allyn, Director and Lee D. Miller, Curator, of
the Allyn Museum of Entomology gave us very valuable photographic aid
and sent us some needed data. Vincent Roth and Kilian Roever gave much
helpful advice at the Southwestern Research Station of the American Mu-
seum of Natural History near Portal, Arizona, where E. q. sanfordi was
studied. Numerous other individuals who have collected E. laeta very kind-
ly answered our queries about the circumstances, or looked up specimens
and records in collections.

Erora Scudder, 1872

4th Ann. Rept. Peabody Acad. Sci., 1871:53

Type species by original designation: Thecla laeta Edwards; includes also
T. clothilde Edwards, Proc. Acad. Nat. Sci. Philadelphia, 1862 (14):55.
Erora Scudder, 1872, p. 169.

Erora Scudder, 1875, 10:166.

Erora Scudder, 1876, p. 106.

Erora Scudder, 1889, 2:815.

Erora Scudder, Dyar, “1902” [1903], p. 40.

Erora Scudder, Barnes and Benjamin, 1926, p. 18.

Erora Scudder, McDunnough, 1938, p. 25.

Erora Scudder, Procter, 1938, p. 186.

Erora Scudder, Field, 1941, p. 303.

Erora Scudder, dos Passos, 1964, p. 55.

Erora Scudder, Hemming, 1967, p. 169.

Erora Scudder, dos Passos, 1970, p. 35.

Erora Scudder, Miller, 1980, p. 209.

Field (1941) added Thecla quaderna Hewitson, 1868 to Erora so that the
genus consisted of two species, laeta and quaderna ; in 1980 Miller added
E. caudata.

Scudder’s original description of Erora read as follows: “Lower Canada
to Virginia. Head moderately large; front as broad as the eyes on a front
view and scarcely half as high again as broad; eyes thinly pilose; palpi
slender, but little longer than the eye. Fore tibiae about four-fifths the length
of the hind tibiae; middle and hind tibiae of about equal length; first superior
branch of the subcostal nervure [RJ arising, at least in the female, a very
little beyond the middle of the upper border of the cell; the second [R2]
about midway between this and apex, cell slightly more than half as long as
the wing.”

To Scudder’s original characterization we add the following, based on

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297

Figs. 1-8. Erora laeta. 1. 1st instar larva, left lateral view. 2. 2nd instar larva, cephalic
view, head. 3-8. 5th instar larvae, coronate chalazae.

both laeta and quaderna, and on the characters given by Miller for caudata.
Very likely some of the characters cited will prove of secondary importance
when more is known about the neotropical Theclini.

Male wings lacking stigma or scent pad: Both sexes of laeta and quaderna
with outer margin of hindwing even, with no tails, anal projections or lobes;

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Figs. 9-14. Erora laeta. 9. Egg, lateral view. 10. Same egg, terminal view. 11. A different
egg, terminal view. 12. 2nd instar larva, lateral view. 13. The same larva, dorsal view, showing
characteristic skeletonizing of leaf. 14. 3rd instar larva, dorsal view, disproportionately greatly
enlarged.

caudata with a short tail at CU2. Wings deep fuscous above with some
iridescent blue in male, much more in female. Hindwings and at least apical
area of fore wings beneath pigmented green with transverse rows of spots;
these, and to some degree fringes, orange to orange-red.

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299

Male genitalia with labides broad and rounded: A longitudinal sclerotized
strip in fultura superior. Falces strongly curved and pointed, wholly well
separated from each other. Tegumen with no processes from ventro-caudal
angle. Valvae well separated from each other, broad basally, terminally bent
strongly dorsad and pointed, terminal margins not toothed. Aedeagus long
and thick, its basal opening nearly half the length of the penis; penis with
a short terminal spine. Cornutus normally well basad in penis, thick, basally
blunt, tapering to a point, about Vs the length of penis.

Female genitalia with papillae anales long, slender, setose: A single pair
of apophyses (apophyses posteriores) bluntly angled or curved ventrad, cau-
dad of middle. Ostium bursae well sclerotized, flat, its dorsal wall (lamella
postvaginalis) split and extending farther caudad than its ventral wall (la-
mella antevaginalis), subterminally widened and then tapering caudad to a
blunt, rounded end. Ductus bursae short, wide, containing a well sclero-
tized, doubled structure with a median series of short, transverse grooves;
ductus seminalis coming off dorso-caudally from this, bending laterad. Bur-
sa copulatrix elongate, with a pair of sclerotized signa, each an oval plate
with a longitudinal, projecting keel tapering gradually cephalad to a fine,
single or double point. Field (1951) gives excellent figures of the genitalia
of laeta and quadema, and Miller (1980) of caudata, which we do not think
it necessary to duplicate.

Egg ( laeta only, Figs. 9-11): Not strongly flattened, laterally rounded,
with a great many short, very fine, conical projections arising from the
junctions of very fine, raised ridges.

Larva, mature ( laeta and sanfordi ): Light greenish, patterned with dark
reddish spots {laeta) or unmarked {sanfordi). Very thickly covered with
various sizes of coronate chalazae, each bearing a single terminal, very
finely spiculate seta (Figs. 3-8). Prothorax with a median dorsal, diamond-
shaped, or scutelliform, area of very minute vesture which we term a “cal-
varium” (Latin — calvus, bald). Prominent dorsolateral and ventrolateral
ridges, the areas between these slightly concave. Swellings in each segment
along dorsolateral ridges give the larva a somewhat serrate aspect in lateral
profile. A pair of more prominent swellings on mesothorax. No honey gland
or associated tubercles. No silk girdle seen (three larvae pupated in captiv-
ity): cremasteral pads and bristles normal.

Pupa (two of laeta, one of sanfordi ): Short, rounded, with darker and
lighter areas caused by the relative amount of sclerotization in the irregular
mesh work of ridges. Vesture sparse, of single, minutely spiculate setae; no
complex projecting chalazae. Stridulatory organ with the grating surface
heavily sclerotized, of many short, fine, parallel, almost uniform ridges.

Foodplants: Fagaceae {Fagus and Quercus) as far as known; perhaps
also Corylaceae {Corylus).

Phylogeny: For phylogenetic speculations we are concerned here with

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only laeta and sanfordi, omitting caudata. Obviously laeta and sanfordi
are very closely related to each other, yet the wide separation of their
geographic ranges presents something of a problem. E. laeta occurs from
southern Canada southward to Virginia, Tennessee, Kentucky and Georgia
along the Appalachian highlands. If considered alone it might well be judged
a species of Nearctic origin with southward extensions along the mountains,
as is the case with many northern species. E. quaderna quaderna, on the
other hand, occurs from south-central Mexico southward to Guatemala and
Costa Rica, but has an isolated subspecies ( sanfordi ) in the southern Rocky
Mountains and northern Mexico. This distribution is characteristic of many
neotropical species. Attempting to reconcile the close relationship of the
two species with their very different distributions, we see two possibilities:

(1) . Erora may have evolved as an oak-feeding quaderna- like stock in the
central or southern Mexican highlands. Then, as a Pleistocene glaciation
(probably the Wisconsin) pushed the biota southward, the northern beech-
birch-maple forest reached this region. Most of the quaderna- like popula-
tion would be forced farther southward along the Central American Cor-
dillera; but some of it, lagging behind, became adapted to a beech-feeding
life in the deciduous (and less xeric) forest, and evolved changes that led to
laeta. Then as the glaciation subsided this differentiating laeta accompanied
the boreal forest northward to its present northern range. Meanwhile, the
southern quaderna -like stock evolved into the present quaderna (and per-
haps other species) and moved northward in Mexico again, and along the
Sierra Madre Occidental (and perhaps the Sierra Madre Oriental) into the
southern Rocky Mountains of New Mexico and Arizona along with the
“live-oaks.” After this, we know, followed a period of climatic desiccation
(which still continues) that formed the Sonoran and Chihuahuan deserts.
This pinched off the northern population, which evolved into E. quaderna
sanfordi , separated by desert from E. q. quaderna and by the arid Great
Plains from E. laeta.

(2) . A second possibility is that Erora may have originally evolved as a
laeta- like stock in the northern deciduous forest. During the Wisconsin
glaciation this would have been pushed far southward into Mexico. On the
recession of the glacier this population would have returned northward as
laeta. Left behind in Mexico were populations that evolved there into the
live-oak feeding quaderna ; and from these, as postulated above, evolved
the northern isolate, E. quaderna sanfordi.

We consider the first of these alternatives to be the more likely, chiefly
because of the existence in Mexico and southward of other species (e.g. E.
caudata Miller) very similar to, and possibly congeneric with, quaderna ;
and also because of the differentiation of quaderna into two subspecies,
while the more recently evolved laeta has remained homogeneous. Cer-

VOLUME LXXXIX, NUMBER 4

301

tainly a great deal more fieldwork and collecting is needed along the Sierra
Madre Occidental, Sierra Madre Oriental and Sierra Madre Transversal,
and also southward along the highlands of Central America, to learn both
more about quaderna itself, and also more about the possible other Erora
and Erora- like species in this region. E. caudata was named “from cloud
forest,” and there may be yet-undescribed species in this environment;
while undescribed species of a quaderna stock may exist in the more xeric
highlands of southern Mexico and southward.

Erora laeta (Edwards) 8, 1862
Erora clothilde (Edwards) 9, 1863

Thecla laeta Edwards, 1863, [14] (4):55 (London, [Ontario], leg. W. Saun-
ders) (our figures 21-22).

Thecla clothilde Edwards, 1863, 2(1): 15 (Quebec, C. E., leg. Rev. M. Pro-
vancher).

Thecla laeta Edwards, Weidemeyer, “1863” [1864], 2(2):534 (British
N[orth] A[merica]).

Thecla clothilde Edwards, Scudder, 1868, 11(24):377 (Streaked Mountain,
near Paris, Maine, July 22, leg. Smith).

Thecla laeta Edwards, Scudder, 1868, 11(24):377 (Streaked Mountain, near
Paris, Maine, July 22, leg. Smith).

Thecla laeta Edwards, “1868-72” [1869], 1(3):[139], pi. Thecla 1, figs. 1,
2 3, 4 $ (London, [Ontario] Canada, leg. Saunders, 1861; Coalburgh,

Kanawha Co., West Virginia, April 1868; Paris, Maine, July 22, leg. W.
Saunders).

T[hecla ] laeta Edwards, Kirby, 1871, p. 401, no. 371 (Amer. Sept.).
T[hecla ] clothilde Edwards, Kirby, 1871, p. 401, no. 371 (Amer. Sept.).
Erora laeta (Edwards) ( =Thecla clothilde Edwards), Scudder, 1872, p. 32
(lower Canada to Virginia).

Erora Scudder (1872) laeta (Edwards) ( =clothilde ) (Edwards), Scudder,
1876, 3(13): 106, no. 203 (Ontario, Quebec and Maine [along the Appala-
chians?) to West Virginia).

Thecla laeta Edwards ( =clothilde Edwards), 1877, p. 42, no. 327 (Maine to
West Virginia, Ontario, Quebec).

Thecla laeta Edwards (= 9 Thecla clothilde Edwards), Strecker 1878, p.

90, no. 108 (Canada, Atlantic states, Maine to Virginia).

Thecla laeta Edwards, Anon., 1881, p. 3, no. 358.

Thecla laeta Mead, 1882, 2(1): 18 (Coalburgh, West Virginia, July 1881, leg.
Theodore L. Mead).

Thecla laeta Edwards, Edwards (partim ), 1883, 3(1):8 (Pr. Quebec, Ontario,
Maine, Catskills, New York; White Sulphur, Coalburgh, West Virginia).

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NEW YORK ENTOMOLOGICAL SOCIETY

Thecla laeta, 1883, 3(9): 123 (Coalburgh, West Virginia, April 17, 1883).
T[hecla] laeta Aaron, 1884, 4(1): 22 (Sand Hills below Atlantic City, New
Jersey, 1 July [1883]).

Thecla laeta Edwards, Fernald, 1884, p. 83, no. 45 (Orono, 18 May).
Thecla laeta Edwards (=9 clothilde Edwards), Edwards {partim ), 1884,
p. 299, no. 374 (Maine to West Virginia; Atlantic City, New Jersey; On-
tario, Quebec).

Thecla laeta Edwards {partim ), 1884, p. [336] (Quebec, Ontario, Maine,
New York, West Virginia).

Thecla laeta Maynard, 1886, p. 37, no. 47 (Canada, Maine, West Virginia).
Thecla laeta Edwards, French {partim ), 1886, p. 277, no. 114 (Atlantic City,
New Jersey; Maine to West Virginia).

Erora laeta Scudder {partim), 1889, 2:819, 3: PI. 14, figs. 6, 9; PI. 23, fig.
2; PI. 39, fig. 17; PI. 55, fig. 2; PI. 65, fig. 8 (St. Joachim 25 mi from
Quebec, leg. Bowles; London, Ontario, leg. Saunders; Catskills, New
York, leg. Edwards; Atlantic City, New Jersey, leg. Aaron; Coalburgh,
Kanawha Co., West Virginia, leg. Edwards; Streaked Mt. near Paris, leg.
Smith; Orono, Maine, leg. Fernald; Graylock Hopper, Williamstown,
Massachusetts, leg. Scudder).

Thecla laeta Edwards, Winn, 1891, 23(5):96 (Beloeil Mt. 22 mi east Mon-
treal, May 24, 1888, leg. Albert F. Winn).

Thecla laeta Edwards, Maynard {partim), 1891, p. 37, no. 47 (Atlantic City,
New Jersey).

Thecla laeta Edwards, Smith et al., 1891, p. 12, no. 296.

Thecla laeta, Maynard, 1891, p. 37, no. 47 (Canada, Maine, West Virginia).
Thecla laeta Edwards, Fyles, 1893, p. 31 (Beloeil Mt., [Quebec] May, leg.
Albert F. Winn).

Thecla laeta, Scudder, 1893, p. 123 (“widely distant places”).

Thecla laeta Edwards {Erora laeta), Bethune, 1894, p. 36, no. 56 (London
and York Mills, Ontario, Beloeil Mt., St. Joachim, St. Hilaire and Que-
bec, May).

Thecla laeta Edwards, Fyles, 1897, p. 12 (Sherbrook, May 25, 1895, leg.
Rev. Abbe Begin).

Thecla laeta Edwards (= 9 clothilde Edwards), Skinner {partim) 1898, p.

50, no. 306 (Maine to West Virginia; Ontario, Quebec).

Thecla laeta Edwards, Holland {partim ), 1898, p. 249, (36) pi. 29, figs. 23
8 , 24 8 (Quebec to southern New Jersey westward to West Virginia).
Thecla laeta, Grant, 1898, p. 76 (cedar swamp on cedar bush near Orillia
[Ontario], May 12, leg. James Walker).

Thecla laeta Edwards, Skinner, {partim ), 1898, Synonymic Cat. N.A.

Rhop., p. 50, no. 306 (Atlantic City, New Jersey).

Thecla laeta Edwards, Weed and Fiske, 1901, p. 47 (New Hampshire).

VOLUME LXXXIX, NUMBER 4

303

Thecla laeta Edwards, Smith et al., 1903, p. 7, no. 325.

Erora laeta (Edwards) ( =clothilde Edwards), Dyar, “1902” [1903], p. 40,
no. 383 (Montana, Colorado).

Thecla laeta, Young, 1904, p. 19 (Ottawa District, leg. C. H. Young).

Erora laeta (Edwards), Fletcher et al., 1904, p. 91, no. 383 (Meach Lake,
Quebec, May 18, leg. C. H. Young).

Thecla laeta Edwards, Wright {partim ), 1905, p. 62 (E[astern] states N[ew]
J[ersey], Canada to Ariz[ona]).

Thecla laeta Edwards, Skinner, [1905], p. 18 (Maine to West Virginia; On-
tario, Quebec).

Erora laeta (Edwards), Stevenson, 1905, p. 54 (St. Hilaire, Quebec, May
24, leg. E. C. Barwick).

Erora laeta (Edwards), Elrod {partim ), 1906, p. 131 (?Montana and Colo-
rado fide Dyar).

T[hecla ] laeta, Fletcher, 1906, pp. 27, 92, no. 383 (Digby, [Nova Scotia]
leg. John Russell).

Erora laeta (Edwards), Fletcher and Gibson, 1907, p. 119, no. 383 (Meach
Lake, Quebec, June 14, 15, leg. C. H. Young).

T[hecla] laeta Edwards, Coolidge {partim ), 1910, p. 374 (Quebec, S to West
Virginia and W to Montana, then S to Sonora).

Erora laeta (Edwards), Perrin and Russell, 1912, 12:262 (Mt. Beaman, Dig-
by, June 19, 1905, June 7, 1906, leg. Ben Lemond).

Thecla laeta Edwards, Winn, 1912, p. 15 (Quebec, leg. G. J. Bowles; St.
Joachim, May, leg. T. W. Fyles; St. Hilaire, May, leg. Albert F. Winn;
Lost River, May, leg. Lachlan Gibb; Meach Lake, leg. Charles H.
Young).

Erora Scudder laeta (Edwards) (= 9 clothilde Edwards), Barnes and
McDunnough, 1917, p. 15, no. 393.

T[hecla ] laeta Edwards {partim ) (= $ clothilde Edwards), Draudt in Seitz,
“1924” 1907-1924 (1920), 5:783, pi. 1044, ibid. 1924, pi. 1044 (eastern
United States).

Erora laeta (Edwards), Criddle, 1922, p. 58, no. 393 (16 Island Lake, Que-
bec, May 18, leg. Miss Ina B. Muir) [Argenteuil Co., May 19 or 20].

Erora laeta (Edwards), Mousley, 1923, 55(2):26 (Hatley, Quebec, Mt. Or-
ford, 2860, April 30, May 21, 1922 [beech]).

Erora laeta, Mousley, 1926, 58(12):293 (Quebec, April 1921 [beech] [ovum
on Fagus]).

Erora Scudder laeta (Edwards) (= 9 clothilde [Edwards]), Barnes and
Benjamin, 1926, 25(1): 18, no. 398.

E[rora] laeta (Edwards), Forbes in Leonard, 1928, p. 680, no. 393 (Keene
Valley, Not [man]; Cortland, Wickwire, leg. Angle; Ithaca, leg. Eyer —
CU; Catskill, New York, May, June, leg. Edwards).

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NEW YORK ENTOMOLOGICAL SOCIETY

Thecla (Erora Scudder) laeta (Edwards), Holland {partim ), 1931, p. 239,
no. (55), pi. 29, figs. 23 6, 24 $ underside (Quebec to southern New
Jersey westward to West Virginia).

Erora laeta (Edwards), dos Passos and Grey, 1934, 66(8): 191 (Paris, leg.
Scudder; Lincoln, leg. Clayton; Mt. Desert, leg. Mrs. A. E. Brower;
Maine [beech]).

E[rora ] laeta (Edwards), Procter, “1937” [1938], p. 186 (418) (Bar Harbor,
May 19, leg. Br[ower]).

T[hecla ] laeta Edwards, Davenport and Dethier, “1937” [1938], 17(4): 173
(ova, larva, food plant Fagus).

Erora Scudder laeta (Edwards (= 9 clothilde Edwards), McDunnough,
1938, p. 25, no. 418.

Erora laeta (Edwards), Comstock {partim ); 1940, 48(1):83, (Atlantic City,
New Jersey, July 1, leg. Aa[ron], 2 broods, Pennsylvania, New York,
New England states, eastern Canada).

Erora laeta (Edwards), Field, 1941, 34(2) : 309—3 11, pi. 1, fig. 1; pi. 2, figs.
1-4; pi. 3, figs. 9 and 10 (Atlantic City and Cape May, New Jersey fide
Murray-Aaron; Huntington, West Virginia; Mt. Equinox, Manchester,
Vermont, 1935; Mt. Killington, Vermont, 1937; Little River, Great Smoky
Mts., Nat. Park, Tennessee, April 15, 1938, leg. Arthur Stupka; Mt.
Washington, New Hampshire, leg. L. W. Sweat; London, Ontario; City
of Quebec, St. Joachim, St. Hilaire, Sherbrooke, Meach Lake, 16 Island
Lake, Hatley, Mt. Orford, Quebec; Digby, Nova Scotia; Paris, Bar Har-
bor, Lincoln, Orono, Norway, Maine; Catskill Mts., Keene Valley, Cort-
land, Ithaca, New York; Atlantic City, Cape May, New Jersey; Williams-
town, Massachusetts; Coalburgh, West Virginia; White Sulphur,
Huntington, Virginia).

Erora laeta (Edwards), Clench, 1943, 51(3):223 (Lake St. Joseph, Portneuf
Co., Quebec, June 1932, leg. G. B. Fairchild).

Erora laeta Remington and Clench, 1947, 1(4):42 (Great Smoky Mts., North
Carolina and Tennessee).

Erora laeta, Carl Cook, 1948, 2(2):22 (Crailhope, Kentucky, April 1941,
May 6, 1947).

Erora laeta (Edwards), Klots, 1951, pp. 16, 24, pi. 16, fig. 14, pp. 129, 142
(eastern Canada, New England south to Kentucky, chiefly Canad. Zone
Forest).

Erora laeta (Edwards), Clark and Clark, 1951, 15:85 frontispiece, fig. 8; pi.
12, a. b. (Mt. Lakes Biol. Stat., Giles Co., Virginia, June 23, 1938, leg.
Lorus J. Milne).

Erora laeta (Edwards), Hessel, 1952, 6(l-3):34 (Cragway Spring, Mt. Wash-
ington Toll Road, New Hampshire, 4,660').

E[rora ] laeta (Edwards), Ferguson, 1954, 23(3): 197 (Mt. Beaman, leg. John

VOLUME LXXXIX, NUMBER 4

305

Russell; June 19, 1905; June 2, 1908; June 7, 1906; near Digby, June 16,
1931; Armdale, May 14, 1944, leg. D. C. Ferguson).

Erora laeta (Edwards), Voss and Wagner, 1956, 10(1 -2): 20, fig. 1, 5th row;
21 (Bliss Township, Emmet Co., Michigan, May 14, 1955).

Erora laeta (Edwards), Lycaenidae, Clench, “1956” [1957], 10(5): 161,
(Powdermill Nature Reserve, 4 mi south Rector, eastern Westmoreland
Co., Pennsylvania, May 3, 1956).

Erora laeta, Clench, 1958, p. 7 (Calverley Lodge, Powdermill Nature Re-
serve, Pennsylvania, May 3, 1956, leg. H. K. Clench).

T[hecla] laeta Edwards, Forbes, 1960, p. 131, pi. 1, fig. c (northeastern
states west to Michigan, mts. of Tennessee).

Erora laeta Edwards, Moore, 1960, p. 17 (Bliss Township, Emmet Co.,
Michigan, May 14, 1955).

Erora laeta Edwards, Dunlop, 1960 (Algonquin Provincial Park, May 20,
1960; June 19, 1960; Fagus present in both places); (also records specimen
in Royal Ontario Museum, Orillia, Ontario, leg. J. Walker).

Erora laeta (Edwards), Clench in Ehrlich and Ehrlich, [1961], p. 218, fig.
419 (Bear Mt., Vermont; south Nova Scotia, south Quebec to Appala-
chians to Tennessee, west to north Michigan, south-central Kentucky).

Erora laeta (Edwards), Smith, “1960” [1961], 14(4):239 (Benton Twp., Graf-
ton Co., New Hampshire 1,800', June 7, 1960).

Erora laeta (Edwards), Riotte, 1961, 15(2): 92 (Algonquin Park, Ontario,
May 20, 1960).

Erora laeta (Edwards), Roever ( partim ), 1962, 16(1): 1, 4 (Ontario and Nova
Scotia south to Virginia, 9 Little River, Great Smoky Mts. Nat. Park,
Sevier Co., Tennessee, 3,000', April 15, 1938, leg. Arthur Stupka; An-
drew’s Bald, 5,860', July 17, 1936, Great Smoky Mts. Nat. Park, Swain
Co., North Carolina, July 17, 1936, leg. Siebert and Evans).

Erora laeta (Edwards), Small “1962” [1963], 16(3): 195 (New Hampshire).

Erora laeta, Hessel, 1963, 17(1):43.

Erora laeta (Edwards) (=9 clothilde [Edwards]), dos Passos, 1964, p. 55,
no. 375.

Erora laeta, Clench, 1966, p. 9 (Powdermill Nat. Res., Pennsylvania).

Erora laeta (Edwards), Clench, 1968, p. 4 (Powdermill Nat. Res., Pennsyl-
vania).

Erora laeta (Edwards), dos Passos, 1970, p. 35.

Erora laeta (Edwards), F. M. Brown and P. A. Opler, 1970, p. 19-77, (Type
in Acad. Nat. Sci. Philadelphia).

Erora Scudder laeta (Edwards), Riotte, 1970, p. 3.

Erora laeta in A. B. Klots, 1970, the rarest of Northern butterflies, news-
letter Mich. Ent. Soc. 18(2): 1-3.

Erora laeta, Patterson, 1971, 25(3):222; “on May 19, 1968 while I was walk-

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NEW YORK ENTOMOLOGICAL SOCIETY

ing along a grassy wood road about 8 mi SW of Wellsboro [Northern
Pennsylvania], in an area much grown up to mixed hardwood brush, a
fresh female Erora laeta literally dropped into the road in front of me.
Later that same year on July 29, a worn male was collected and another
sighted on blossoming hardhack [Spiraea tomentosa ] in a nearby wet
field.”

Erora laeta (Edwards), Sullivan, 1971, Journ. Lep. Soc. 25(4): 295 (Alleghe-
ny Co., NW corner of state while catching Speyeria idalia).

Erora laeta , Sullivan, 1971 ibid. 25(4):296 (Allegheny Co., North Carolina,
July 1, 2,700' $ , Ash Co., 2,700' 8).

Erora laeta (Edwards), Lewis, 1973 ( partim ), p. 218, Lycaenidae pi. 18,
figs. 14, 18 (Quebec to Arizona, Mexico and Central America).

Erora laeta Edwards, Brower, 1974, p. 20 (Streaked Mt., near Paris, 22 July
(Smith); Dallas Pit. (Rangeley) 16 June, 9; T2R12 (Greenville), 4 July,
(3; Orono, 18 May; Lincoln (Clayton); Bar Harbor, 19 May, 8 , 25 May,
9 all Maine.

Erora laeta (Edwards), T. C. Emmel in Howe, 1975, p. 306, pi. 50, fig. 21
8 (range does not approach that of quaderna).

Erora laeta, Bowers, 1978 (abundance, Carroll County, nr. Bartlett, New
Hampshire; habits).

Erora laeta (Edwards), Drees and Butler, 1978, p. 202 (Cabell, Greenbriar,
Kanawha, Pendleton and Randolph Counties, West Virginia, V-5 to VII-
26).

Erora laeta, Oosting, 1979, p. 160 (Ontonagon Co., Michigan, 27 May 1975
(one female; no Fagus present; Corylus present).

Erora laeta Miller, 1980, pp. 209-216 (general discussion and differentia-
tion).

Taxonomic notes: The type of Thecla laeta Edwards (Figs. 21-22) is in
the Carnegie Museum. It was discussed by Brown and Opler (1970). The
type locality is London [Ontario]; the label on the type reads only “Cana-
da.” (At that time what is now called Ontario was known as “Upper Can-
ada.”)

The type of Thecla clothilde Edwards cannot be found in the Carnegie
Museum. It was taken near Quebec by Rev. M. Provancher presumably in
1862 and probably at Cap Rouge where he lived. Provancher’ s collections
of insects are in the College of Levis and the Quebec Public Museum, but
Rev. J. C. E. Riotte who kindly examined these collections advises us (in
litt.) that the type of clothilde is not in either of these collections. Conse-
quently it must be deemed to have been lost or destroyed. We see no ne-
cessity to erect a neotype.

For many years the butterfly that we now call E. quaderna sanfordi dos

VOLUME LXXXIX, NUMBER 4

307

Passos was known, but classified as E. laeta. This has caused considerable
confusion, and specimens of both were mixed together. We note that a
record of “laeta” by Aaron (1884) from Atlantic City, New Jersey was the
basis for various authors (Edwards 1884; French 1886; Scudder 1889; May-
nard 1891; Holland 1898; Skinner 1898; Holland 1931; Comstock 1940; Field
1941) attributing laeta to New Jersey. We have always mistrusted this rec-
ord, which is from an entirely wrong environment. A note from Harry
Clench at the Carnegie Museum (in litt. 24 July 1978) reads: “We have a
female Erora that is labelled Atlantic City, N.J., ex coll. Skinner, ex coll.
A.N.S.P. The A.N.S.P. collection had much (all?) of E. Murray Aaron’s
collection, and I believe he was the source of “Atlantic City” as a locality
for laeta. This specimen, however, is a quaderna sanfordi\” We believe
that this Atlantic City record and any New Jersey records based on it (e.g.
“Cape May”) are erroneous.

Some additional records follow, based on specimens in various museums
and private collections. We are indebted to J. Donald Lafontaine and A. C.
Sheppard for most of these. BM = British Museum (Natural History).
CNC = Canadian National Collection. LEM = Lyman Entomological Mu-
seum, McGill. UM = University of Montreal. ACS = Collection, A. C.
Sheppard. PH = Collection, Peter Hall. RL = Collection, Ross Layberry.

New York. 1 8, Long Lake, Adirondack Forest Preserve, Upper New
York State, May 22, 1977, leg. P. Hall (CNC). New Brunswick. Edmund-
ston, leg. Henry Hensel, 1 8 June 12, 4 $ $ June 13-16 (BM). Quebec. St.
Hilaire (1,500 ft) 24.V.88, 24.V.02, 24.V.10, 24.V.11, ex A. F. Winn Coll.,
(LEM). 1 9 Meach Lake, 12 mi NW Ottawa, Ont., 25. V. 1968, leg. A. Hanes
(CNC). 1 9 Duncan Lake, 7 mi WNW Wakefield, Masham Twp. 1.V.1977,
leg. D. M. Wood (CNC). 1 8 Gatineau Park, 6 mi N. of Aylmer, 27. V. 1978,
leg. P. Hall (PH). 19, Gatineau Park, 29. V. 1978, leg. R. Layberry (RL).
Parc du Mont Tremblant, 27. V. 1957, 1 9 and 2. VI. 1958, 1 9 , leg. A. Robert
(UM). Vernet (Papineau Co.) 24.VII.1941, 1 9 , on Spiraea blossom, leg.
A. C. Sheppard (ACS). 1 8, 4 mi S. of Lost River (Argenteuil Co.)
12. V. 1957, on Dandelion blossom, leg. A. C. Sheppard (ACS). Maryland.
Dargan, Washington County, 14 April, 1977, Robert S. Simmons.

Field studies; localities: Field studies were made by the authors separately
and together, and accompanied at times by others, beginning in 1934 and
ending in 1968. These were at two localities; others were visited but with
no results for laeta.

1. Near West Bridgewater, Windsor County, Vermont, up a woodroad
toward Killington Peak (Rutland and Woodstock topographic quadrangles).
The woodroad ascends generally westward and eventually runs into trails
up Killington Peak. E. laeta was found along the woodroad between 1,600
and 1,900 ft altitude. This locality was first found by A. C. Frederick and

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NEW YORK ENTOMOLOGICAL SOCIETY

L. J. Sanford. It has been referred to by them and others, and specimens
from there labelled, as “Bear Mt. near Bridgewater Corners” and “Mt.
Killington” (Field, 1941, pp. 305, 315).

2. Near Sandgate, Bennington County, Vermont (Equinox topographic
quadrangle). We refer to this as “Mt. Equinox,” as have other collectors.
From Sandgate a country road runs northeastward and eastward through a
community called Beartown; and from this a woodroad ascends to the sad-
dle (2,300 ft altitude) between Mt. Equinox and Mother Meyrick Mountain.
E. laeta was found most commonly at this saddle, but also below it along
woodroads on both sides down to about 1,900 ft altitude. This locality was
first found by the veteran student of lycaenids, J. C. Cook; and A. C.
Frederick also collected there. Their material was mostly labelled “Mt.
Equinox, near Manchester, Vermont, 2000 ft.” and was so referred to by
Field, loc. cit.

We learned about these localities through the kindness of Messrs. Cook,
Frederick and Sanford. During our fieldwork considerable plant-succession-
al changes took place at both localities as old meadows and clearings were
invaded by trees and shrubs and the woodroads became more shaded. In
addition the woodroads at both localities were heavily bulldozed because
of logging operations higher up. However, these changes did not appear to
affect the laeta except, perhaps, to concentrate them in open spaces.

In all, the authors put in, separately or together, a total of 52 days at
these localities. E. laeta was taken or merely seen on 16 occasions, as noted
below.

1934, 22-23 May, Mt. Equinox, none
14 June, Mt. Equinox, none
1936, 29 May-1 June, Mt. Equinox, none
1938, 4 June, Mt. Equinox, none
5 June, W. Bridgewater, none
1947, 29 May, Mt. Equinox, none

I June, W. Bridgewater, none

1949, 4 June, Mt. Equinox, none

5 June, W. Bridgewater, 1 3 , 1 9 , both worn

10 June, Mt. Equinox, 1 $ , 2 9 9 (1 very fresh), 2 seen

1950, 9 June, Mt. Equinox, none
10 June, Mt. Equinox, none

II June, W. Bridgewater, none
12 June, Mt. Equinox, none

1951, 19 May, Mt. Equinox, 1 9 (worn), 1 9 (very fresh), 1 6 seen
26 May, Mt. Equinox, 2 $ $ (very fresh), 1 seen

28 May, W. Bridgewater, none

VOLUME LXXXIX, NUMBER 4

309

5 June, Mt. Equinox, 5 S S , 9 9 9, mostly very fresh
20-21 June, Mt. Equinox, 1 9 (worn), 1 9 (very fresh)

1952, 23-24 May, Mt. Equinox, none

6 June, Mt. Equinox, none

1953, 29 May, Mt. Equinox, none

14-15 June, W. Bridgewater, 1 $ seen

1955, 7-9 June, Mt. Equinox, none

1956, 16 June, Mt. Equinox, none

17-18 June, W. Bridgewater, 1 8 , 1 S seen

1957, 9 June, W. Bridgewater, none

10-11 June, Mt. Equinox, 20 S 6 and 9 9 , mostly fresh to very fresh,
several more seen

1960, 17-19 June, Mt. Equinox, none

1961, 17-18 June, Mt. Equinox, 14, mostly 9 9 and very fresh, 2 seen
19 June, W. Bridgewater, 2 seen

1963, 11 June, Mt. Equinox, 1 seen

1966, 12 June, Mt. Equinox, 16 6 6 and 9 9 , some very fresh, others seen

1967, 12-15 June, Mt. Equinox, none

1968, 31 May, Mt. Equinox, none
3 June, Mt. Equinox, none

Only days of favorable weather when laeta might have been flying are
listed. All records are, of course, of the first generation. No consistent
efforts could be made to find the second generation; on two visits to Mt.
Equinox in mid- July none were seen. We found laeta in only 8/18 years,
but suppose that it was present in all years. Only four years: 1951, 1957,
1961 and 1966 were “good.” The record for 1951 at Equinox is notable, for
the first specimen, a somewhat worn 9 , was taken on 19 May and a very
fresh one on 21 June, a flight period of more than a month. During the first
part of the flight period Shadbush Amelanchier canadensis (L.) comes into
bloom, followed by the Wild Cherries Prunus serotina Ehrh. and pensyl-
vanica L.

Associated butterflies: E. laeta appeared to be preceded in flight, at the
same altitude levels, by Celastrina ladon (Cramer) and Erynnis brizo (Bois-
duval and LeConte) and Juvenalis (Fabricius). A few individuals of Pieris
napi oleracea Harris precede laeta, but occasional very heavy flights of this
species may coincide with flights of laeta. Pieris virginiensis Edwards was
taken only once (1951). Fresh Poanes hobomok (Harris) and Amblyscirtes
samoset (Scudder) and vialis (Edwards) flew consistently with fresh laeta.
Lower down on the woodroads fresh Papilio glaucus L. of the small north-
ern form, Boloria selene (Denis and Schiffermueller) and bellona (Fabri-
cius), Melitaea harrisii (Scudder) and Phyciodes tharos (Drury) were be-

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ginning to fly and occasionally strayed up into laeta territory. Toward the
latter part of the laeta flight season a phenotypically very mixed population
of Limenitis arthemis (Drury) and astyanax (Fabricius) became common.
We conclude that the flight period of laeta lasts about a month to five weeks
beginning a week to ten days after the first flight of C. ladon and E. brizo
and Juvenalis.

Our studies and the observations of other collectors indicate that laeta
populations show considerable fluctuation from year to year, with occa-
sional years when there is almost an outburst with large numbers occurring.
Opposed to this is the possibility that, since laeta is essentially a tree-living
species, a year of apparent sudden abundance is merely one when a chance
combination of circumstances brings (or keeps) a large number of individ-
uals down at ground level for hardening the wings, drinking and basking.
Very likely much of the seemingly abnormal abundance is due to the for-
tuitous presence of the collector at just the right time.

The two laeta localities are in what is essentially the Beech-Birch-Maple
hardwood forest that covers large areas in the Canadian Life Zone (Mer-
riam), Coniferous Forest Biome (Shelford) and Canadian Biotic Province
(Dice). The localities contain, however, large admixtures of the Transition
Life Zone, particularly in the warmer localities at lower elevations. This is
in accordance with the known geographic distribution of laeta, since such
an environment occurs far southward along the Appalachian and Allegheny
mountains and highlands.

Associated plants: The chief trees and shrubs serve as an index to the
environment. Some of these, such as Red Spruce {Pice a rube ns Sargent),
Sugar Maple {Acer saccharum L.), Mountain Maple {Acer spicatum La-
marck), Striped Maple {Acer pensylvanicum L.) and White Birch {Be tula
papyrifera Marshall) are characteristic dominants of this lower Canadian
Zone formation. Other important woody plants were willow ( Salix ) two
species, Quaking Aspen {Populus tremuloides Michaux), Beaked Hazelnut
{Corylus cornuta Marshall), Gray Birch {Be tula populifolia Marshall), Cher-
ry Birch {Be tula lenta L.), Yellow Birch {Betula lutea Michaux), Beech
{Fagus grandifolia Ehrhart), Gooseberry {Ribes sp.), Shadbush {Amelan-
chier canadensis L.), Raspberry {Rubus idaeus L.), Flowering Raspberry
{Rubus odoratus L.), Blackberry {Rubus allegheniensis Porter), Wild Black
Cherry {Prunus serotina Ehrhart), Wild Red Cherry {Prunus pensylvanica
L.), Red Maple {Acer rubrum L.), Maple-leaved Viburnum {Viburnum ac-
erifolium L.), Hobblebush {Viburnum alnifolium Marshall) and Common
Elder {Sambucus canadensis L.).

The mixed Canadian-Transition zonal status of the region studied at Mt.
Equinox is also indicated by other butterflies found there. Cercyonis pegala
(Fabricius) occurred in grassy meadows about a half mile below the laeta
area as a very mixed population. In a considerable series (taken later in the

VOLUME LXXXIX, NUMBER 4

311

season) specimens range all the way from the northern “ nephele ” type
lacking an orange patch on the forewing to the more southern “ alope ” type
with a large orange patch. We personally consider these two types, alope
(Fabricius) and nephele (Kirby) to rank as subspecies. In any event, they
here intergraded in a narrow tension zone. The same was true of the pop-
ulation of Limenitis arthemis (Drury) which occurred very commonly, chief-
ly below the restricted laeta area, as a very mixed population of L. arthemis
arthemis, the northern, white-banded subspecies, and L. arthemis astyanax
(Fabricius), the more southern, non-banded subspecies. In a large series
collected here occurred almost every possible intergradation between these
two phenotypes. In the West Bridgewater area a large series of Melitaea
harrisii Scudder was taken over several years in a meadow below the laeta
area. They show a great variation between lightly marked individuals like
the southern population liggetii Avinoff and dark individuals like the north-
ern albomontana Avinoff. We have some doubt about recognizing liggetii
and albomontana as “good” geographic subspecies; there is too much in-
tergradation all along the line between them.

Foodplant: For many years only the record of Mousely (1923) pointed to
beech as the laeta foodplant. On Mt. Orford, Quebec, Mousley watched a
laeta female deposit an egg on the underside of a beech leaf near the base
and midrib. The egg was taken; the larva began feeding on beech, but soon
died. We therefore paid special attention to beech, but never saw females
on it. A number of captive females were confined on it both in sleeves in
the field and in cages at home. Other potential woody foodplants were also
tried, but with no luck. Most of the females were very fresh, and probably
had never mated; one worn one had apparently already deposited her eggs.
Then in 1951 a female laid (8-10 June) about 20 eggs on both beech and
beaked hazel leaves and also the screen of the cage. The eggs hatched 14-
19 June. A number of the larvae were lost, but several ate either beech or
beaked hazel leaves through the first instar. After that those on beech ceased
to feed. One of these, transferred to beaked hazel, ate that and survived;
the other beech-eaters died. One larva on beaked hazel died in its second
instar, three survived into the third instar, and two others survived to pu-
pate. One of the pupae died, but the other survived to eclose as a rather
small female on 4 August, under the care of Dr. Frederick Rindge. We
afterward guessed that some of the beech leaves, which came from Pelham,
New York, were polluted. (This we know to have been the case with some
pine foliage from there used for other larvae.) The beaked hazel, brought
from Vermont and Connecticut, was clean enough. It was noted that three
of the larvae, including the one that developed into an adult, fed partly on
developing hazel fruits, which is consistent with Whittaker’s record of find-
ing a laeta larva on a beech fruit (see below). During later fieldwork in 1951
and subsequent years a great deal of time was spent in examining both beech

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and hazel minutely, but no eggs or larvae were found on either. There was
abundant beaked hazel, as well as beech, in the laeta areas.

In 1972 or 1973 Dr. R. H. Whittaker found a larva on beech in the Hubbard
Brook Experimental Forest in the vicinity of West Thornton, 22 km north
of Plymouth, Grafton County, New Hampshire. This was in the course of
a Hubbard Brook ecosystem study, forest part (Whittaker, R. H. et al. 1974,
Ecological Monographs, 44:233-254). Dr. Whittaker kindly sent us the fol-
lowing information: “We were felling trees, including beech, for detailed
dimensional analysis; a part of this was the separation of sample branches
into wood, twigs with leaves, and fruit. I was plucking and bagging fruits,
and had tossed a fruit with larva into the bag when my mind connected two
points; it was a lycaenid larva, and the larva was associated with beech and
Erora. The larva, retrieved, was found to have been feeding on the soft
papillae of the beech fruit. It continued to feed on these for a few days.”
The larva then died and was preserved and deposited in the Peabody Mu-
seum of Yale University. Sent to us through the kindness of Sidney Hessel,
it was compared with our preserved laeta larvae and photographs and pos-
itively identified as laeta.

We sent a questionnaire to a number of collectors who had taken laeta.
Many of these answered that there had been beech in the environment; none
stated that there was no beech; fewer had noticed beaked hazel. Edward
Voss kindly showed us the locality in Bliss Township, Emmet Co., Michigan
where he and Wagner had collected laeta (Voss and Wagner 1956). Beech
was present, but beaked hazel was not. The fact that our larvae were reared
through on beaked hazel does not indicate that this is a normal foodplant —
only that it is an acceptable substitute in vitro. Unfortunately our mention
of laeta having been reared on beaked hazel has crept into print (via the
grapevine) in Forbes, 1960, pp. 131-132. However, Oosting (1979) has re-
cently recorded the presence of laeta at a locality in Ontonagon Co., Mich-
igan where there is abundant beaked hazel but apparently no beech. So, the
matter stands open; we consider beech to be the normal foodplant over at
least most of the range, with beaked hazel perhaps a natural foodplant in
some localities and circumstances.

Newly hatched larvae probably do a certain amount of leaf skeletonizing
at first but are likely to switch to flowers and developing fruits. If they do
not find any of these they may continue on leaves. As the rearings took
place the following durations were noted: egg, 6-6.5 days; 1st instar, 4-5
days; 2nd instar, 4-5 days; 3rd instar, 4^1.5 days; 4th instar, 5.5 days, 5th
instar, 11.5 days; pupa, 13 days.

Egg (Figs. 9-11): Pale, slightly greenish yellow, considerably flattened;
height 0.30-0.34 mm, diameter 0.69-0.75 mm, with a somewhat variable,
shallow depression surrounding micropyle. Micropyle of 5-6 unequal po-
lygonal cells surrounded by many smaller ones. Surface of egg thickly stud-

VOLUME LXXXIX, NUMBER 4

313

ded with small, conical projections irregularly placed, basally connected by
very fine, raised ridges. There are about 25 of these projections along an
irregular line from base to termen; those around the base are slightly longer.
Duration of egg stage (5 eggs) averaging slightly more than 6 days. Cf.
Scudder 1889, Vol. Ill, pi. 65, fig. 8.

Larva, first instar (Fig. 1): Length 1.9-2. 2 mm (5 larvae). Pale, greenish
yellow, elongate, subcylindrical, not flattened. Head small, its connection
with prothorax very lightly sclerotized. A finely reticulate area on dorsum
of prothorax marking the site of the calvarium (see below) in later instars.
All body segments with very long, finely spiculate setae as shown. Heavily
sclerotized black lenticles as shown on prothorax, mesothorax and abdom-
inal segments 1-7 (two each) and 8 (one). Prolegs with 4 crotchets on each
side. From a preserved specimen. Average duration of instar (4 larvae) 4.5
days.

Larva, second instar (Figs. 2, 12-13): Length 2.8-4. 1 mm. Pale greenish
yellow with an indistinct, darker, middorsal line. Head small, almost col-
orless; a round dark spot at each cluster of stemmata; mandibles and bases
of antennae brown. Head greatly retracted beneath prothorax. Body cov-
ered with very small, coronate chalazae, each consisting of a small, basal
swelling bearing a corona of 5-9 radiating spines or teeth and a central,
longer spine usually minutely spiculate and more or less curved. These
chalazae are essentially colorless and almost hide the underlying skin. The
slightly darker appearance of the dorsal and lateral lines is caused by the
darkness of the large chalazal spines. A series of paired, slightly protuberant
dorsolateral swellings on mesothorax, metathorax and abdominal segments
1-7, which bear noticeably longer central spines. Dorsal surface between
these swellings slightly concave, and lateral surfaces between dorsolateral
swellings and ventrolateral ridges almost flat or slightly concave. Average
duration of instar (5 larvae) 4.5 days.

Larva, third instar (Fig. 14): Length 4. 8-5. 9 mm. Very much as in second
instar, essentially colorless, the very dense coronate chalazae masking the
skin color. Central spines of chalazae along dorsolateral swellings and ven-
trolateral ridges especially long. Average duration of instar (5 larvae) 4.2
days.

Larva, fourth instar: (Pressure of other work prevented the recording of
a detailed description.) In this instar the larvae had developed a definite
pattern of dark spots and except in size resembled the fifth instar larva
described below.

Larva, fifth instar (Figs. 3-8, 15-19): Length when mature 11.5-12.3 mm.
Vesture of prothorax very short and sparse in central calvarium. This is
diamond-shaped, its cephalic angle very attenuate and sharp, with concave
sides, its caudal angle shorter and truncate. Skin yellowish green showing
more or less through vesture depending on degree of distention of larva.

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NEW YORK ENTOMOLOGICAL SOCIETY

Figs. 15-20. Errora laeta. 15-18. Mature larvae, lateral and laterodorsal views, showing
normal variation. 19. Mature larva, lateral view, a lightly marked specimen. 20. Prepupal larva,
dorsal view; the prothoracic calvarium shows plainly.

VOLUME LXXXIX, NUMBER 4

315

Collection

ofix. cf{L Cj^

j Collection |

W. H. Edwards

W. H. Edwards ;

fit* •CunJa.

23

—

Collection of
C.F. dos Passos

«L\. \* 193 b

Arizona

V-C\Oftt\

L.SK*>?0«.V\

193

Cyril F. dosPasaos

White Mts. ,
$OOftft. Alt. \

d-

Collection of
C.F. dos Passos

n.\. \J\ 193 b

Arizona

White Mts..
$OO0Tt Alt. •

L.S**>?o*V\

**8^* 193
Cyril F. dosPassos

25

<1

Collection of j

C.F. doa f ’i

VN-'A mb

Arizona J
White .Mts., I
<XOCOft. Alt. |

Det. 193

Cyril F. dos Pass'

26

' rs.
r<>

*>

J

ection of

wise

Arizona

lite^Mts. ,

“7

*A.v.oTN*fc

Det. 193

Cyril F. dosPassos

<*

Figs. 21-26. 21-22. Erora laeta (Edwards), type 8 , upper and under sides. 23-24. Erora

laeta sanfordi dos Passos, holotype 8 , upper and under sides. 25-26. Erora laeta sanfordi dos
Passos, allotype $ , upper and under sides.

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NEW YORK ENTOMOLOGICAL SOCIETY

Figs. 27-34. 27-28. Erora laeta, pupa, dorsal and lateral views. 29-30. Erora quaderna

sanfordi, mature larva, lateral views; Chiracahua Mts., Arizona. 31-32. E. q. sanfordi, pupa,
dorsal and lateral views. 33-34. E. laeta 9 9 displaying, from life, Sandgate, Vermont.

VOLUME LXXXIX, NUMBER 4

317

Vesture of coronate chalazae with their central spines particularly long along
ventrolateral ridges, especially on thoracic and last abdominal segments.
Dorsolateral ridges each consisting of a swelling of each segment from me-
sothorax posterad; these give the larva a subserrate outline. The most an-
terior of these swellings, on the mesothorax, is especially prominent. Similar
lateroventral ridges, weak on mesothorax, extending caudad to around end
of abdomen. Legs and prolegs largely hidden below these. Honey gland not
present.

Dark markings quite consistent in pattern but varying in color and extent.
In one larva they were bright reddish; in another darker brownish red. On
or about each mesothoracic swelling a small dark area. On first abdominal
segment a pair of dorsolateral patches more or less running down on each
side. On each side of fourth and fifth abdominal segments a patch slightly
above ventrolateral ridge with traces of dark running cephalad on third, and
caudad on sixth abdominal segments. On or slightly below each ventrolat-
eral ridge a narrow dark shade on abdominal segments 1-6, diffusing out on
7. On sixth abdominal segment a pair of wide dark patches almost confluent
together dorsally, extending down on each side to about the level of the
spiracles, and slightly confluent with the ventrolateral patch on sixth seg-
ment, tapering caudad on seventh and eighth abdominal segments. There
was some variation in the size and extent of these patches, but they were
present in all of the five mature larvae studied.

Prepupal larva (Fig. 20): The general color became darker and dingier,
with the dark markings less contrasty. The larva became shorter and as-
sumed a more broad-oval shape, more rounded laterally and dorsally. The
calvarium was very noticeable. No silk girdle was spun. Duration (2 larvae)
2 days.

Pupa (Figs. 27-28): Short, blunt and rounded, characteristically lycaenid.
Integument more or less covered with fine, raised reticulations which, being
more heavily sclerotized than the rest of the cuticula, account for the darker
appearing areas. Setae arising almost entirely from chalazae at the intersec-
tions of reticulations. Setae rather sparse, there being, for example, about
nine along the middorsum of the third abdominal segment, which measures
0.91 mm; setal lengths 0.10-0.21 mm. All setae simple, very finely spiculate,
gradually tapering to fine tips. None of the specialized and complex sensillae
present in some lycaenoid pupae (cf. Downey and Allyn 1973, many figs.)
were noted. (The pupa of E. quaderna sanfordi had a number of “sensillae
companiformia” (ibid., Fig. 62) chiefly in small clusters about the abdominal
spiracles). We were also unable to find any of the small, paired pores noted
by Downey and Allyn dorso-mesad of the abdominal spiracles. The strid-
ulating structures were not studied in detail because of our reluctance to
dissect the specimens. They have the caudal margin of the 5th abdominal

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NEW YORK ENTOMOLOGICAL SOCIETY

segment raised and more strongly sclerotized than the remainder of the
segment. Entad from this is the grating surface, which is heavily sclerotized
and bears many fine, uniform, parallel ridges running entad (cf. Downey
and Allyn, op. cit., Fig. 24). This extends down on each side to slightly
below the level of the spiracle. The file opposed to these ridges was not
studied. No trace of a honey gland was found on the dorsum of segment
seven. Cremasteral hooks are of the conventional form, each with a flat-
tened, rounded tip bent strongly basad, on a low, rounded protuberance on
each side of the anal slit. Each of the two pupae was formed on a leaf, but
no silk girdle was seen.

Adult behavior (Figs. 33-34): A majority of other collectors’ records, as
well as our own, mention specimens of laeta as having alighted on bare
ground and being seen only when they flew up, often to the collectors’
surprise because their cryptic coloration made them inconspicuous. In 18/
19 of our records of this the specimen was on dry earth or gravel, only one
being on damp earth. Quite a number of other collectors, however, mention
damp earth or mud. The late J. H. Cook described to us (in litt.) a number
of laeta in the Mt. Equinox area flying up from a damp place where he
believed there was horse urine. We tried several times to attract laeta in
areas where they were flying by wetting the ground with water or urine, but
with no success, although we did attract numbers of Celastrina ladon. Per-
haps we should have had a horse. The majority of the specimens seen on
the ground were females, and the majority of these were teneral, i.e. very
recently emerged from the pupa, and emitting meconium when kept alive.
They were also very tame, crawling about slowly, so that we were able to
crawl up, watch them very closely and photograph them from only a few
inches away. Some were making probing motions with the proboscis (per-
haps only adjusting it) but others were not. We have no doubt that individ-
uals on wet earth or mud may drink, but we did not see this. Those on dry
ground seemed to be basking, sometimes orienting themselves across the
line of the sun, sometimes opening the wings so as to receive a maximum
of sun on the upper surface. Such an action by a female may also serve for
visual attraction of a male (Figs. 33-34). The large proportion of teneral
females to males probably resulted from the males having emerged earlier
from the pupae and having flown to rest on foliage higher up where they
were not seen.

An extraordinary experience of Bowers (1978) is highly pertinent. Mr.
Bowers and Reginald Webster, collecting near Bartlett, Carroll County,
New Hampshire on 21 May 1977, along abandoned dirt roads through beech
woods, saw over 80 laeta of which all but two were females, some quite
worn, while others looked freshly emerged. The females were resting on the
ground, chiefly on mud, where they were presumably drinking, and are
described as very easy to catch, some not even flying up when the net was

VOLUME LXXXIX, NUMBER 4

319

clapped over them. The day was sunny, hot (33°C) and humid, so that the
laeta were presumably thirsty. The only two males seen flew much more
actively than the females, only occasionally alighting on vegetation.

Of 27 of our specimens only 8 were caught as they rested upon or flew
from the leaves of various plants. In 4/8 of these cases the leaves were of
such low species as Wood Nettle ( Laportea canadensis L.), Canada violet
( Viola canadensis L.) and Raspberry ( Rubus idaeus L.). In the other four
cases the individuals were resting on the leaves of Mountain and Sugar
Maples ( Acer spicatum and saccharum ) five to eight feet above the ground.
One of these specimens, on a maple leaf, was caught entirely unexpectedly
(in the true laeta tradition) when a collector made a sweep at a high-flying
geometrid and found not only the moth but a scale-perfect laeta in his net —
his first! Perhaps the individuals resting on high leaves were basking where
there was more direct sunlight — a phenomenon we have noticed in some
Callophrys ( Incisalia ) henrici (Grote and Robinson), which are much given
to basking on the shiny leaves of Holly and Laurel.

This persistent habit of laeta, especially when teneral, of resting on the
ground may largely explain the irregular and unexpected records of large
numbers. Perhaps the adults are on the ground for only a short time when
the right combination of heat, sun and thirst keeps them there. Most of the
time large numbers may be present, but unseen, high in the trees. We think
that the species is more a dweller in the forest canopy than has been real-
ized.

Only once was anything like a courtship observed; a male and a female
were fluttering about each other in a small, sunny spot along a shaded
woodroad. After about two minutes they flew up high together and disap-
peared among beech foliage overhead. Perhaps most courtships and matings
take place high up.

We were long puzzled by an apparent absence of flower-visiting. Captive
individuals fed well on honey and sugar water. Numerous flowers such as
those of Shadbush, Wild Cherries, Violet, Blackberry, Raspberry and Win-
tercress ( Barbarea vulgaris Brown) were available in abundance but were
not visited. Finally, on 12 June, 1966 a large number of laeta were seen
visiting the flowers of a Wild Cherry several feet above the ground; 16
specimens were taken by W. D. Field and the junior author. Ferguson (1954,
p. 197) records laeta in Nova Scotia visiting Blackberry flowers; Sullivan
(1971) saw it on Hardhack ( Spiraea ) flowers in Pennsylvania; Coveil (in
litt.) records it visiting Oxeye Daisy and Wild Hydrangea in Kentucky; and
Sheppard on Spiraea and Dandelion in Quebec. For comparison be it noted
that E. q. sanfordi is an eager flower visitor, especially on Amelanchier and
Ceanothus.

Fluctuation in numbers: Our own records, as well as those of other col-
lectors, seem to indicate considerable variation in numbers from year to

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NEW YORK ENTOMOLOGICAL SOCIETY

year. Most records are of only one or two individuals; but occasionally and
unexpectedly the species will be found very common or abundant. It is
always possible, as noted above, that individuals are present, but unob-
served, being high in the beech foliage, courting or ovipositing. We suspect
that when collectors watch among high foliage they will see many more
laeta than otherwise.

Voltinism: Our own data on voltinism are meagre. Our one reared laeta
emerged in August. Two visits to the Mt. Equinox locality in mid-July re-
vealed no laeta. Enough records are available, however, to indicate that
laeta is bivoltine, with a second generation in July-August. Dr. Charles
Coveil tells us (in litt.) of over a hundred laeta taken on and near Big Black
Mountain, alt. 4,145 ft, Harlan County, Kentucky in early to middle July,
obviously second generation; only two Spring specimens have been taken,
24 April, 1976.

Distribution, summary: As discussed above, laeta is essentially a species
of the beech-birch-maple forest that characterizes lower Canadian Zone,
and of the very extensive ecotonal regions where this mixes with upper
Transition Zone. In Canada it is found in Nova Scotia, New Brunswick,
Quebec and Ontario, not extending northward into the Canadian Zone re-
gions of dominant coniferous forest. In northern Michigan it flies where
there are strong Canadian Zone elements, and “Great Lakes Forest.” In
New England it occurs in Maine, New Hampshire, Vermont and western
Massachusetts, following Canadian Zone through the Green Mts. and Berk-
shires. It is not recorded from Connecticut or Rhode Island. We suspect,
however, that it may occur in extreme western Connecticut and perhaps
eastern New York where the high ridge of the Taconics runs southward
from the Berkshires, carrying considerable Canadian Zone and beech, at
altitudes up to 2,316 ft. Its occurrence in the Catskill Mts. is quite as ex-
pected; it must occur widely in the Adirondacks. The outlying records from
the Finger Lakes region (Ithaca, Cortland, Wickwire) are in a region where
some Canadian Zone elements occur on high hills, but do show that laeta
has been able to maintain colonies that are offshoots of the main lower
Canadian Zone pattern. It has not been reliably recorded from New Jersey
(we regard the Aaron record from Atlantic City and Cape May as false) but
it may well occur along the Kittatinny Ridge in the north. It occurs in the
highlands of Pennsylvania. The single record from Maryland (Dargan,
Washington County, 14 April 1977, leg. Robert S. Simmons, in litt.) is in
semi-highland environment with beech present. Thence laeta expectedly
follows the Appalachian and Allegheny highlands southward through West
Virginia, Kentucky, Tennessee and North Carolina to Georgia. It may occur
in the Cumberland Plateau in Tennessee and the Ozark Plateau in Missouri
and Arkansas.

VOLUME LXXXIX, NUMBER 4

321

Erora quaderna quaderna (Hewitson)

Thecla quaderna Hewitson, 1868, p. 35.

Thecla attalion Godman and Salvin, 1887, 2:60; 3:pl.35, Figs. 19-20.

The present paper deals with E. quaderna only because of its relationship
with E. sanfordi dos Passos, which we place as a subspecies of quaderna.
The nominate subspecies occurs in central Mexico and southward. Specu-
lations as to its phylogeny have been covered above under the genus Erora.
Some data about E. q. quaderna have turned up, however, in the course of
our studies and these are presented below.

Type material: The types (both unique) of Theda quaderna and T. at-
talion were studied at the British Museum (Natural History) where T. G.
Howarth very kindly picked them out and dissected them for us. That of
quaderna, a female, is dissection T.G.H. 656 (1964); that of attalion, a
male, is dissection T.G.H. 657 (1964). We compared these with specimens
of quaderna from the United States National Museum very kindly dissected
and sent on loan by W. D. Field. These were: a male, dissection W.D.T.
475 from Guatemala; and three females, dissections W.D.F. 476, 477 and
478 from Mexico and Guatemala. In coloration and pattern the specimens
agreed well with the types of the corresponding sexes (making some allow-
ance for discoloration and wear) and similarly with each other. In the gen-
italia we found only minor differences which seem well within the limits of
individual variation. The male differed from the attalion type only slightly
in the amount of angulation of the ventrocaudal angle of the valva. The
females differed from the quaderna type only in minor details of the fine
toothing and terminal spines of the signa. We consider these specimens and
the types of quaderna and attalion to be conspecific, and attalion to be a
junior synonym of quaderna.

Clench (1943, p. 223) “selected” the type locality of quaderna as Tan-
citaro, Michoacan, Mexico. We accept that designation because that locality
is within the known range of the species and is one from which specimens
identified with the species have been taken (Code, 72E). In addition that
locality is about 365 miles from Orizaba, the type locality of attalion which
we consider the female of quaderna.

We note, incidentally, that the types of Theda aura Godman and Salvin
(1887) a female from Irazu, Costa Rica and of Erora gillottae Riley (1924)
a male, from Mt. Irazu, Costa Rica, were also compared with our material
and with the types of quaderna and attalion, since it was believed that they
might be related. They are certainly not conspecific with quaderna or cau-
data, and show such genitalic differences that we believe that they do not
belong in Erora.

Environmental data: Mr. Harry K. Clench very kindly made available to

322

NEW YORK ENTOMOLOGICAL SOCIETY

us some of his unpublished notes and observations of E. quaderna quad-
erna. These were made 5 air miles north of Zimapan, Hidalgo, Mexico,
about 2,140 m altitude, 12 and 21 January 1966 by him and Lee D. Miller.
In all, 16 quaderna were taken. The area is in a small, northward-facing
valley clothed with oak-pine-juniper “chaparral” (low, open, scrubby for-
est). The quaderna were almost exclusively above 2,140 m alt. They were
noted as being associated with one or more species of the small-leaved
Quercus; and Clench (correctly, it now appears) believed that this was the
most likely foodplant. These data agree well with our own from Arizona
and with remarks by others. It seems reasonable to assume that this very
widespread live oak-pine-juniper habitat is the home of quaderna wherever
it occurs. This contrasts with the “cloud forest” habitat cited by Miller for
caudata.

Erora quaderna sanfordi dos Passos, 1940
Figures 23-26, 29-32

Thecla laeta Edwards (partim ), 1883, 3(1):8 (Ft. Grant, Cochise Co., Gra-
ham Mt., Arizona, leg. H. K. Morrison).

Thecla laeta Edwards {partim ), 1884, p. [336] (Mt. Graham, Arizona, leg.
Morrison).

Thecla laeta Edwards (= 9 clothilde Edwards), Edwards (partim ), 1884,
p. 299, no. 374 (Arizona).

Thecla laeta Edwards, French (partim ), 1886, p. 277, no. 114 (Arizona).
Erora laeta, Scudder (partim ), 1889, 2:821, pi. 23, fig. 2 (Mt. Graham, Ar-
izona).

Thecla laeta Edwards, Skinner (=9 clothilde Edwards) (partim) 1898, p.
50, no. 306 (Arizona).

Thecla laeta Edwards, Holland (partim ), 1898, p. 249, no. (36), pi. 29, figs.
23 6 , 24 S (Mt. Graham, Arizona).

Erora laeta (Edwards), Elrod (partim), 1906, p. 131 (Montana and Colorado,
fide Dyar; Arizona).

T[hecla\ laeta Edwards, Coolidge (partim ), 1910, 42(11):374 (Huachuca
Mts., Cochise Co., July; Montezuma Canyon, Mt. Graham, Graham Co.;
Chiricahua Mts., Arizona.

T[hecla ] (Erora Scudder) laeta Edwards (partim) (=9 clothilde Edwards),
Draudt in Seitz, “1924” [1907-1924] (1920), 5:783, pi. 155i, 1044 (1924)
(Arizona).

Thecla (Erora Scudder) laeta Edwards, Holland (partim ), 1931 p. 239, no.

(55), pi. 29, figs. 23 6 , 24 6 (Mt. Graham, Arizona).

Erora laeta (Edwards), Comstock (partim ), 1940, 48(1 ):83 (Arizona).

Erora laeta sanfordi dos Passos, 1940, p. I (White Mts. 8,000', June 21,
1936; Santa Catalina Mts. 8,500'; Chiricahua Mts., April 7; Chiricahua,

VOLUME LXXXIX, NUMBER 4

323

Arizona, July 7, 1933, leg. ex Sternitzky; southern Arizona, leg. Poling,
ex J. Dahl; Mud Springs, Santa Catalina Mts., Arizona, 6,500', July 17-
20, 1916; Silver City, New Mexico, leg. R. T. Kellogg).

Erora quaderna, Field ( partim ), 1941, 34(2):303, pi. 1, figs. 4, 5, 6, 8; pi. 2,
figs. 5-8; pi. 3, figs. 11-13 (Mt. Graham, Chiricahua Mts., Prescott, Oak
Creek Canyon, Santa Rita Mts., Paradise, Redington, Palmerlee, Senator,
Santa Catalina Mts., Huachuca Mts., Arizona; New Mexico; Utah).
Erora quaderna sanfordi dos Passos, Clench, 1943, p. 223.

Erora quaderna, Bauer, 1954, 26(3):95— 102, pi. 101 (Mingus Mt., Arizona).
Erora Scudder laeta Edwards quaderna Hewitson, Martin and Truxal,
1955, p. 23 (Arizona, March-July; California, March).

T[hecla ] laeta Edwards, Forbes {partim ), 1960, p. 131 (Arizona).

Erora quaderna Hewitson, Clench, 1961, p. 218, fig. 420 {partim).

Erora laeta sanfordi dos Passos, Roever {partim ), 1962, 16(1):4 (Apache
Co., Trout Creek Road, 7,500', July 4, 1958, 3 <3c3, July 4, 1959 ASS,
June 25 and 26, 1960 4 S S , 1 $ , July 22, 1961, 1 ^,3 9 9; Cochise Co.,
E. Turkey Creek Canyon, 6,400', April 10, 1959 2 S S , 10 9 9, April 15,
1960 12 9 9 ; Pinery Canyon, Chiricahua Mts., April 15, 1960, 2 S S , 4
9 9 , Chiricahua Mts., April 15, 1960 4 9 9, June 19, 1960, 2 S S; Cocon-
ino Co., Oak Creek Canyon, March 28, 1959 1 S , July 16, 1961 1 9 ; Gila
Co., Peterson Ranch, 7,000', July 2, 1960 2 9 9; Graham Co., Wet Can-
yon, 6,000', April 23, 1961 2 9 9; Greenlee Co., Rte. 666, July 5, 1959 6
S S , Gray’s Peak Road Camp, July 5, 1959 5 S S ; 1 9 Pima Co., Madera
Canyon, Santa Rita Mts. 4,400^1,600', March 8, 1959 1 S , March 19,

1960 3 SS , Summerhaven, 7,800', Santa Catalina Mts., May 24, 1959 1

5 , July 9, 1961 1 9 ; Pinal Co., Peppersauce Wash, 5,000', Santa Catalina
Mts., April 18, 1961 1 9; Santa Cruz Co., Madera Canyon, Santa Rita
Mts., 5,600-6,400', March 29, 1959 3 SS, April 9, 1959 4 (3d, 28 9 9,
April 10, 1959 1 S , 32 9 9 , April 3, 1960 8 9 9, April 9, 1960 3 9 9, July

6, 1960 2 33,7 9 9, July 9, 1960 4 9 9, July 31, 1960 2 9 9, April 6,

1961 12 9 9 , Arizona).

Erora quaderna, Field, 1962 (Arizona).

Erora quaderna, (Hewitson), Hubbard 1965, Journ. Lep. Soc. 19(4), 232
(Pintos Altos Mts., New Mexico).

Erora quaderna sanfordi dos Passos, dos Passos, 1970, p .35.

Erora quaderna sanfordi dos Passos, Emmel in Howe, 1975, p. 307, pi. 50,
fig. 22 9 (Madera Canyon, Santa Cruz Co., Arizona A.M.N.H.).

Erora quaderna (Hewitson), Emmel and Emmel, 1973, p. 94 (doubtfully in
California).

Erora quaderna sanfordi dos Passos, Ferris, 1976, Journ. Lep. Soc.
30( 1):38 — 49 (Grand Co., N. Mexico, February-May). Sierra Co., July
1976. (The specimens were absolutely fresh so that it appears this species
is bi-voltine. Normally collected from February to May.)

324

NEW YORK ENTOMOLOGICAL SOCIETY

Erora quaderna sanfordi dos Passos, Miller, 1980, p. 214 (characteristics;

New Mexico, Arizona, southern Utah, Mexico: Madera, Chihuahua; Lo-

beras Summit, Sinaloa).

The citations before 1940 given above consist of references to what we
now classify as Erora quaderna sanfordi dos Passos, known only from Utah
(?), New Mexico, Arizona, northern Mexico and California (?). We regard
the nominate subspecies, E. q. quaderna, as occurring from central Mexico
southward, as noted above (under that name) and in the discussion of Erora
phylogeny. It was known to Edwards and others of the older authors, but
regarded as being merely laeta, so that some of their references to “ laeta ”
are partly attributable to sanfordi when its western localities are given. It
was named by dos Passos in 1940 in the combination Erora laeta sanfordi
but since then has been variously placed as laeta, quaderna and quaderna
sanfordi, the last combination by Clench (1943, p. 223) being in our judge-
ment correct.

Taxonomic notes: Although Field (1941) placed sanfordi as a junior syn-
onym of quaderna we believe it sufficiently distinct to deserve subspecific
rank. For much of the following we are indebted to Harry Clench, since our
own material of q. quaderna is scanty. On the forewings of sanfordi females
the blue is more restricted, with only a sprinkling in the discal cell and only
traces in the base of cell Cuj ; and in cell Cu2 the blue extends distad to only
a bit beyond the middle of the inner margin. On the fore wings of nominate
quaderna females the blue largely fills the discal cell; almost entirely fills
cell Cux; and extends distad in cell Cu2 to above about % out on the inner
margin. On the hindwings of sanfordi females, however, the blue is more
extensive than in nominate quaderna females, especially in cell M3 where
it nearly reaches the outer margin; and the veins through this blue are more
extensively lined with black. The blue of sanfordi is more purplish. Males
of sanfordi have the red spots of the hindwings beneath a little smaller and
duller than those of nominate quaderna. We have been unable to note any
consistent difference in the hue of the wings beneath — not that there may
not be a difference, but this green, a pigment color, seems to be rather
unstable in Erora, fading during only a few days in flight. The same seems
to be true of the color of the fringes; apart from wear these seem to fade
easily from a definite red to a pale orange-red.

Field studies: Field studies of E. quaderna sanfordi were made 17 April-
10 May, 1969, in the Chiricahua Mts. near Portal, Cochise County, Arizona,
while working at the Southwestern Research Station of the American Mu-
seum of Natural History. Adults were found to be common at many places
from 5,400 to 7,000 ft altitude, always in the open evergreen oak-juniper-
pine scrub that covers very large areas in the lower parts of this and other
Arizona mountain ranges. This represents Upper Sonoran Life Zone. The

VOLUME LXXXIX, NUMBER 4

325

adults were always associated with Quercus arizonica Sargent and Q.
Emoryi Torrey, the dominant oaks of this habitat.

The adults were eager flower visitors, especially at Berberis wilcoxi Kear-
ny, Ceanothus integerrimus Hook, and Arn. and Amelanchier in company
with Celastrina ladon cinerea (Edwards), Callophrys ( Incisalia ) augustinus
annetteae (dos Passos), Callophrys (Mitoura) spine torum (Hewitson) and
siva (Edwards) and Erynnis Juvenalis clitus (Edwards). Zestusa dorus (Ed-
wards) was also a common associate, but did not visit flowers. Its larvae
were found on the same tree of Q. Emoryi on which a Larva of sanfordi was
found, and were reared through (Klots 1971).

When not visiting flowers the sanfordi adults spent much time flying about
and alighting on the leaves of both the species of Quercus ; only one indi-
vidual was seen to alight on the ground, and that for only a few seconds.
Even when wet soil was available the adults did not alight on it, as adults
of C. ladon and Z. dorus often did. There may have been some basking on
Quercus leaves. After a great deal of work examining Quercus, with the
kind help of Kilian Roever (during which several Z. dorus larvae were
found) a presumably sanfordi larva was found, 29 April, on Q. Emoryi .
This was successfully reared (partly while driving to New York), pupated
on 1 June and emerged as an adult sanfordi on 11 June. The specimen with
its larval and pupal exuvia is in the American Museum of Natural History.

Mature larva (Figs. 29-30): Length with head extended forward 15 mm.
Skin pale translucent green, more dilute green dorsally, showing where not
covered by vesture. Head yellowish brown, darker anteriorly, paler poste-
riorly, darker toward mouthparts, usually almost entirely retracted beneath
prothorax. Prothorax rounded, with the middorsal calvarium more or less
scutelliform and somewhat depressed. Legs and prolegs pale green, yellow-
ish toward tips. General shape onisciform, rather flattened by the lateral
production of the ventrolateral ridges. Sides above these ridges slightly con-
cave up to a pair of dorsolateral ridges along mesothorax, metathorax and
abdominal segments inclusive. On mesothorax a pair of rounded protuber-
ances at the cephalic edge, i.e. at the cephalic ends of the dorsolateral
ridges; these look yellowish brown because of the concentration on them
of the vesture units. Middorsal surface slightly concave between the dor-
solateral ridges. Behind abdominal segment 6 the dorsolateral ridges diverge
strongly to the lateral margins of the 8th abdominal segment, followed by
the quite flat last segment. The ridges are most prominent at the caudal edge
of each abdominal segment, giving a subserrate lateral profile. On metatho-
rax also a pair of slightly protruding humps between the dorsolateral pro-
trusions and the lateral margins. No dark markings; color light brown be-
cause of thick covering of brown vesture.

Most units of the vesture are, as in E. laeta, short coronate chalazae, i.e.
each is a short chalaza with a subterminal ring of short, radiating, triangular

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NEW YORK ENTOMOLOGICAL SOCIETY

teeth and usually a slightly curved, stout, minutely spiculate spine termi-
nally. Many of the pale brown vesture units have the terminal spine very
short or absent. The longest terminal spines are along the ventrolateral and
dorsolateral ridges. Scattered among the pale brown vesture units are a
smaller number of smaller, darker, red-brown ones, not so protruding, look-
ing like tiny red dots. No honey gland or associated structures seen. A slight
silk pad was spun on a leaf, but no silk girdle was seen.

Pupa (Figs. 31-32): Exceedingly like laeta pupa, differing only in minor
respects. Slightly more setose. Pattern of darker and lighter areas similar
to, but in general slightly darker than those in laeta due to greater sclero-
tization of the raised areas and integument. A number of sensilla compan-
iformia (Downey and Alleyn 1973, fig. 62) were noted in small clusters about
the abdominal spiracles; these were not seen in laeta. Stridulatory apparatus
like that of laeta.

Voltinism: E. q. sanfordi is clearly bivoltine, with an early Spring gen-
eration (March, April, May) followed by a Summer one (June, July) as is
shown by the records cited by dos Passos (1940), Roever (1962) and Ferris
(1976).

Distribution: E. q. sanfordi occurs in many localities in Arizona and New
Mexico. Dyar (1903, p. 40) listed it from Montana and Colorado (as E. laeta)
and gave no other localities, East or West! Elrod (1906) merely repeated
Dyar. The species cannot be credited to Montana and Colorado on this
basis. Field (1941) cites a Barnes Collection specimen from Utah, leg.
Bruce. This is not necessarily authentic. Similarly, a specimen in the Los
Angeles County Museum from Providence Mts., San Bernardino County,
California, III-22-40, leg. T. B. Blevins, is regarded by Emmel and Emmel
(1973, p. 94) as mislabelled. Possibly this is the same specimen cited by
Martin and Truxal (1955, p. 23) as “California, March.” In any event the
presence of the species in California needs verification. It is highly desirable
that its extent in northern Mexico, and its southern limits there, should be
investigated. Likewise, the northern extent of E. q. quaderna, and any
possible contact between it and E. q. sanfordi should be investigated. Miller
(1980, p. 214) gives the two localities in northern Mexico.

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VOLUME LXXXIX, NUMBER 4

327

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VOLUME LXXXIX, NUMBER 4

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330

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14(4):239-240.

VOLUME LXXXIX, NUMBER 4

331

Stevenson, C. 1906. Notes on the season 1905 (Western Quebec), 36th Ann. Rep. Ent. Soc.
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American Museum of Natural History, 79th Street & Central Park West,
New York, New York 10024.

Received for publication March 6, 1981.

332

NEW YORK ENTOMOLOGICAL SOCIETY

ACKNOWLEDGMENT

The Editors wish to express their appreciation to all those who have
helped in reviewing the manuscripts submitted during 1981 for publication
in the Journal: E. W. Baker, J. W. Brewer, W. L. Brown, Jr., R. L. Detra,

E. E. Grissell, A. P. Gupta, R. Hoy, S. P. Hubbell, A. B. Klots, F. E.
Kurczewski, M. L. May, H. D. Niemczyk, R. B. Roberts, R. K. Sofield,

F. C. Swift.

HONORARY, LIFE, AND SUSTAINING MEMBERS

Charles P. Alexander, Joseph Bequaert, Raymond Brush, F. S. Chew,
Lucy Clausen, Howard E. Evans, Su Zan Swain Firmage, S. W. Frost,
Irving Granek, Mark Indenbaum, Alexander B. Klots, Kimumaro Okano,
Nellie Payne, L. L. Pechuman, Ruth R. Ralston, Hubert J. Thelen, George
Townes, Roman Vishniac.

VOLUME LXXXIX, NUMBER 4

333

INDEX TO SCIENTIFIC NAMES OF ANIMALS AND PLANTS
VOLUME LXXXIX

Generic names begin with capital letters. New genera, species, subspecies and varieties are
printed in italics. The following items and articles are not indexed:

1. Dominant vegetation found in each of the three major habitats on Plum Island, p. 5.

2. Number and location of specimens captured on Plum Island, pp. 7-9.

3. Arthropod-borne infectious animal diseases contained at the Plum Island Animal Disease
Center, p. 11.

4. Collection label, Heliconia species resemblance, collection location and season of collec-
tion for 25 Heliconia collections used in the principal components analysis, p. 112.

5. Frequency of occurrence of the 23 most common insect “morpho species” (listed by
letters) found in 25 Heliconia inflorescences collections, p. 114.

6. Genera associated with Glyphipterigidae auctorum. pp. 220-290.

7. North American Erora. pp. 291-327.

Achillea millefolium, 59
Acilius semisulcatus, 152
Adoxoplatys, 41
Aedes, 2

atropalpus, 9
cantator, 12
sollicitans, 12
triseriatus, 9
Agrotis musa, 59
stigmosa, 59
venerabilis, 61
volubilis, 59
Alathesus, 41
Alitocoris, 41

Amblyomma americanum, 21
Ambystoma, 152
Amelanchier, 199
Anopheles, 12
Aplomyiopsis, 197
xylota, 196
Aster, 199

Atalopedes campestris, 202
Audintella, 41

Babesia microti, 6
Bembix, 202
Berberis vulgaris, 199
Bombyx mori, 77, 144
Brachelytron, 41
Branta canadensis, 10
Brochymena, 40

Caltha palustris, 103
Camponotus, 159
Caracia, 40
Cary a ovata, 199

Centris, 51
Cephaloleia, 111
Ceratophyllum demersum, 56
Cerceris dichroa, 172
Chelone glabra, 91
Cheumatopsyche, 56
Cochliomyia macellaria, 161
Coleomegilla maculata, 102
Conibosa, 123
Copestylum, 115
Comus, 197

Ctenophthalmus pseudagyrates, 10
Cucurbita pepo, 43
Culex, 12, 115
Culicoides, 6
hollensis, 12
Culiseta, 2

Danaus plexippus, 98, 184
Dasypus novemcinctus, 163
Dermacentor albopictus, 21
variabilis, 6, 16, 23
Dermestes carnivorus, 158
Distingphyses, 188
Dumetella carolinensis, 10
Dytiscus fasciventris, 156
verticalis, 152

Ectatomma, 207
Editha, 202
Elodea canadensis, 56
Epargyreus clams, 202
Eritrachys, 41
Eucerceris, 172
baja, 172
bicolor, 172

334

bitruncata, 177
canaliculata, 172
conata, 178
ferruginosa, 172
lacunosa, 172
lacunosa sabinasae, 172
lapazensis, 172
morula, 172
morula albarenae, 172
nevadensis, 172
nevadensis ferruginosa, 172
nevadensis mojavensis, 172
nevadensis nevadensis, 174
pimarum, 176
superba, 172, 179
zimapanensis, 172
Euphydryas phaeton, 89
Euspilotus aenicollis, 158

Felis domestica, 10
Fidicina, 137
amoena, 123
“coffea,” 123
mannifera, 123
pronoe, 123
semilata, 123
sericans, 133
spinocosta, 123
Fraxinus americana, 199

Geomysaprinus (Priscosaprinus) belioculus,
158

Gillisius, 115

Haliaeetus leucocephalus, 10
Helianthus, 43
annuus, 43
Heliconia, 109
aurea, 121
bihai, 116
caribaea, 116
episcopalis, 117
imbricata, 116
rostrata, 111
wagneriana, 116
Hemilucia segmentaria, 158
Herrichella, 41
Heteranthera dubia, 56
Flister punctiger, 158
Hydropsyche, 56
recurvata, 56
Hyla, 152

Inga, 123

Ips confusus, 65

Ixodes dammini spielman, 9

NEW YORK ENTOMOLOGICAL SOCIETY

Janeirona, 40

Larus argentatus, 10
marinus, 10
Leptothrips mali, 166
Lincus, 41

Macrocheles muscadomesticae, 164
Macropygium, 41
Marghita, 40
Mechanitis isthmia, 211
Meconema thalassinum, 170
Melambyrsus, 41
Melanodermus, 41
Melissodes, 43
agilis, 43
Merosargus, 115
Microtus pennsylvanicus, 9
Minilincus, 41
Miopygium, 41
Moncus, 41
Morpho achillaena, 80
achilles, 80

achilles amazonica, 81
peleides insularis, 80
Musca domestica, 9

Nemognatha, 48
Neoadoxoplatys, 41
Nymphalis antiopa, 98, 184
milberti, 146
milberti milberti, 147
milberti viola, 147

Ochlerus, 41
Oenothera biennis, 59
Orbatina, 41
Orchopeas howardii, 10

Pacarina, 134
Pandion haliaetus, 11
Pantala flavescens, 98, 184
Paragonia pyramidata, 80
Paralincus, 41
Parides, 210

iphidamas, 210
Parochlerus, 41

Parthenocissus quinquefolia, 199
Pephysena, 188
ftiscosa, 188
insignis, 188
levis, 188

longirhynchus , 188
microlevis, 188
picta, 188

Pericopus leucophaea, 204

VOLUME LXXXIX, NUMBER 4

335

Peromyscus lecopus, 9
Phaenicia eximia, 158
Pherecles, 41

Pieris glaucus canadensis, 146
glaucus glaucus, 146
Pithecollobium, 125
Polybia simillima, 204
Popillia japonica, 34
Populus deltoides, 103
tremuloides, 59
Pristiphora erichsonii, 96
Prochyliza azteca, 158
Prunus serotina, 199
virginiana, 59
Pyrus malus, 199

Quesada gigas, 123
Quichuana, 115

Rana, 152

Rhabdocelus obscurus, 65
Rhus toxicodendron, 199
Rubrica, 202

Schaderia, 41
Schaefferella, 41
Solanum, 211
Solidago, 199
Sphex ichneumoneus, 170

Stelopolybia, 161
Stictia, 202

Carolina, 202
Stictiella, 202
Stomoxys calcitrans, 9
Svastra obliqua, 43

Tabanus nigrovittatus, 9
Taraxacum officinale, 59
Tenebrio molitor, 65
Tetrochlerus, 41
Triepeolus, 46
helianthi, 48
Trigona, 161
Trirhabda, 197
borealis, 197
virgata, 197

Vaccinium vacillans, 59
Vemonia patens, 204
Vespula, 199
Viburnum, 199

Wyeomyia, 115

Zammara smaragdina, 129
smaragdula, 123
Zygia longifolia, 123
Zyzzyx, 202

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JOURNAL of the

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NEW YORK ENTOMOLOGICAL SOCIETY

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The JOURNAL of the NEW YORK ENTOMOLOGICAL SOCIETY is
devoted to the advancement and dissemination of knowledge pertaining to
insects and their related forms.

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THE SOCIETY solicits book-length manuscripts in any area of Entomology
to consider for publication. Suitable manuscripts will be submitted to Fair-
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published jointly by the Society and Fairleigh Dickinson University Press.
For further information or to submit manuscripts write to President, N. Y.
Entomological Society, American Museum of Natural History, 79th St. &

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Please make all checks, money-orders, or drafts payable to the NEW YORK

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Manual (3rd edition, A.I.B.S., 1972).

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Biochemical abbreviations: follow rules of the U.I.P.A.C.-I.U.B.

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