■

*

% °15.1 0Q>! 3

JOURNAL.

OF THE

NEW YORK

ENTOMOLOGICAL SOCIETY

Iruotrb to jEttitmtologij in <$*tt*ral

VOLUME LII, 1944

Published Quarterly by the Society
North Queen St. and McGovern Ave. Lancaster, Pa.

New York, N. Y.

THE SCIENCE PRESS PRINTING COMPANY
LANCASTER, PENNSYLVANIA

CONTENTS OF VOLUME LII

Page

Alexander, Charles P.

Records and Descriptions of Neotropical Crane-flies

(Tipulidae, Diptera), XVII 45

Records and Descriptions of Neotropical Crane-flies
(Tipulidae, Diptera), XVIII 369

Arnett, Ross H., Jr,

A Revision of the Nearctic Silphini and Nicrophorini
Based upon the Female Genitalia (Coleoptera, Sil-

phidae) 1

Bacon, Annette L.

Bibliography of Frank E. Lutz 69

Bird, Henry

A Re-Survey of Papaipema Sm. (Lepidoptera) 193

Boyd, William M.

Insect Introductions and War 200

See Weiss, Harry B.

Bromley, Stanley W.

Ephraim Porter Felt 223

Brown, F. Martin

Notes on Mexican Butterflies, II, Pieridae 99

Notes on Mexican Butterflies, III, Danaidae 237

Notes on Mexican Butterflies, IV 343

Caldwell, John S.

PsylliidaB from Tropical and Semitropical America

(Ilomoptera) 335

Clench, Harry K.

Two New Subspecies of Everes Comyntas Godart (Lepi-
doptera, Lycaenidae) 59

Two New Subspecies of Lycaenopsis pseudargiolus Bvd.

& Lee. (Lepidoptera, Lycaenidae) 273

New Neotropical Theclinae (Lepidoptera Lycaenidae) 255

Davis, William T.

The Remarkable Distribution of an American Cicada:

A New Genus and Other Cicada Notes 213

iii

Forbes, William T. M.

Lepidoptera from Western Peru and Ecuador 75

Ginsburg, Joseph M.

Outdoor Protection from Mosquitoes 247

Hemming, Francis

Recent Work by the International Commission on Zo-
ological Nomenclature 211

Huckett, H. C.

A Revision of the North American Genus Eremomy-

ioides Mai loch (Diptera, Muscidae) 361

Huntington, E. Irving

Thecla burdi Kaye, a Synonym 328

Jordan, Karl

The Status and Functions of the International Com-
mission on Zoological Nomenclature and the Present

State of Its Work 385

Linsley, E. Gorton

Hibernation of the Syrphid Fly, Lathyrophthalmus
aeneus Scop. 272

New Species of Neopasites with Notes Concerning Others

(Hymenoptera, Nomadidae) 277

McCoy, E. E.

See Weiss, Harry B.

Members of the New York Entomological Society 389

Michener, Charles D.

Differentiation of Females of Certain Species of Culex

by the Cibarial Armature 263

Miller, Dwight D.

Drosophila melanura, a New Species of the Melanica

Group 85

Milne, Lorus J., and Margery Milne

Notes on the Behavior of Burying Beetles (Nicrophorus

spp.) 311

Milne, Margery

See Milne, Lorus J.

Rapp, William F., Jr.

Catalogue of North American Psychodidse 201

Richards, A. Glenn, Jr.

The Structure of Living Insect Nerves and Nerve
Sheaths as Deduced from the Optical Properties 285

IV

Rupert, Laurence R.

A New Species of Lambdina and Notes on Two Species

of Besma (Lepidoptera, Geometridae, Ennominae) 329

Schneirla, T. C.

The Reproductive Functions of the Army-Ant Queen

as Pace-Makers of the Group Behavior Pattern 153

Weiss, Harry B.

The Death-Feints of Alobates pennsylvanica DeG., and

Alobates barbata Knoch. (Coleoptera) 281

Frank Eugene Lutz 62

Insect Food Habit Ratios of New York State 283

Insect Responses to Colors 267

Notes on the Death-Feint of Bruchus obtectus Say 262

Weiss, Harry B., E. E. McCoy, Jr., and William M. Boyd
Group Motor Responses of Adult and Larval Forms of

Insects to Different Wave-Lengths of Light 27

Wolcott, A. B.

A Generic Review of the Subfamily Phyllobaeinae 121

v

JnsocJS

Vol. LII No. 1

MARCH, 1944

Journal

of the

New York Entomological Society

Devoted to Entomology in General

Publication Committee

HARRY B. WEISS JOHN D. SHERMAN, Jr.

T. C. SCHNEIRLA

Subscription $3.00 per Year

Published Quarterly by the Society
N. QUEEN ST. AND McGOVERN AVE.

LANCASTER, PA.

NEW YORK, N. Y.

—

1944

CONTENTS

A Revision of the Nearctic Silphini and Nicrophorini
Based upon the Female Genitalia (Coleoptera, Silphidae)

By Ross II . Arnett, Jr 1

Group Motor Responses of Adult and Larval Forms of In-
sects to Different Wave-Lengths of Light

By Harry B. AVeiss, E. E. McCoy, Jr,, and William
M. Boyd 27

Records and Descriptions of Neotropical Crane-Flies
(Tipulidae, Diptera), XVII

By Charles P. Alexander 45

Two New Subspecies of Everes Comyntas Godart (Lep-
idoptera, Lyceenidae)

By Harry K. Clench 59

Frank Eugene Lutz

By Harry B. Weiss 62

Bibliography of Frank E. Lutz

By Annette L. Bacon 69

Lepidoptera from Western Peru and Ecuador

By Wm. T. M. Forbes 75

Drosophila Melanura, a New Species of the Melanica
Group

By Dwight D. Miller 85

Notes on Mexican Butterflies, II, Pieridae

By F. Martin Brown 99

NOTICE: Volume LI, Number 4, of the Journal of The
New York Entomological Society was published on
January 3, 1944.

Entered as second class matter July 7, 1925, at the post office at Lancaster, Pa.,
under the Act of August 24, 1912.

Acceptance for mailing at special rate of postage provided for in Section 1103.
Act of October 3, 1917, authorized March 27, 1924.

JOURNAL

OF THE

New York Entomological Society

Vol. LII March, 1944 No. 1

A REVISION OF THE NEARTIC SILPHINI AND
NICROPHORINI BASED UPON THE FEMALE
GENITALIA (COLEOPTERA, SILPHID^E)

By Ross H. Arnett, Jr.

Ithaca, New York

The use of the female genitalia as a basis for identification of
genera and species of Coleoptera has been greatly neglected. In
some groups of beetles, however, the female genitalia apparently
offer more diversity of form among species than do those of the
male. Tanner in 1927 pointed out a growing necessity for a
study of the genitalia of beetles for specific descriptions. The
genitalia are naturally more constant within a species than other
parts and they give the taxonomist a better concept of a species,
and its subspecific forms and categories.

The purpose of this study is to present the comparative mor-
phology of the female genitalia of the Neartic Silphini and Nicro-
phorini. It is hoped that the drawings, the key to the genitalia
and the synopsis will be an aid to identification. No basic con-
clusions have been made as to relationships, although certain
tendencies have been indicated.

Acknowledgments. — I wish to express my sincere appreci-
ation and thanks to Professor J. C. Bradley of Cornell University,
under whose direction this work was done, for his many sugges-
tions and criticisms. I am also greatly indebted to Mr. V. S. L.
Pate of Cornell University for help in preparing the paper itself
and to Mr. J. G. Franclemont for aid in developing a technique
for the study of the genitalia.

APR 10 '44

2

Journal New York Entomological Society

[Vol. LII

I am indebted to Professor M. H. Hatch of the University of
Washington and to the United States National Museum for their
generous loan of material for which I thank them sincerely.

Finally, I wish to thank Mr. E. D. McDonald, Jr. and Mr. Ru-
dolph Schuster for their aid in making the drawings, and to my
wife Mary both for aid in typing the paper and for her continual
encouragement.

Methods. — The methods used in this study are somewhat dif-
ferent from those used by Tanner. The beetle was first relaxed
in hot water and the genitalia removed with a pair of forceps
and boiled in caustic potash. They were then put in water and
the mid-ventral membrane cut, the two lateral plates (the para-
procts) flattened out on each side of the dorsal plate (the proc-
tiger) and the coxite bent out to the side of the valvifer so that
the whole organ was flattened out. It was then mounted in
balsam. This enables projection drawings to be made. It is
essentially the same method as that used in studying the male
genitalia of Lepidoptera.

Sexual Differences. — In the female Silphini, the sutural
angles of the elytra are very acute, whereas in the males they are
generally but not always rounded. The hind femora are greatly
enlarged in the males of Silpha littoralis L. and normal in size in
the female sex. Little sexual differences is evident in Thanatophi-
lus truncata Say.

In the Nicrophorini studied, the males have the eyes situated
well forward on the head, the clypeus large and the fore tarsal
pulvilli expanded, whereas in the females the eyes are placed
well towards the back of the head, the clypeus is appreciably
smaller and the fore tarsal pulvilli are simple.

SPECIES STUDIED

Neartic Species. — A list of the species considered in this paper
follows. Only a few forms of subspecific rank have been studied.
Of those studied, however, some changes have had to be made in
their ranking. Of the others, the opinion of other authors has
been followed in considering their rank. In the case of Nicro-
phorus pulsator Gistel, and Silpha tyrolensis Leach, the evidence
of their existence in the Neartic region is insufficient and they
have not been included here.

Mar., 1944]

Arnett: Silphid.®

3

NEARTIC SILPHINI AND NICROPHORINI

Silphini

Silpha L.
lift oralis L.

form surinamensis Fab.
ab. bizonatus Port.
—disciocollis Brulle
analis Chev.
cequinoctialis Gistel
braziliensis Dej.
cayennensis Berg, (nec
Sturm. )

var. elongata Port,
var. discreta Port.
Thanatophilus Leach

Subgenus Oiceoptoma Leach
americana L.
peltata Catesby
ab. affinis Kby.
terminata Kby.
canadensis Kby.
acc. brunnipenis Hatch
noveboracensis Forst.
marginalis Fab.
marginata Kby.
quadripunctata L.

quadripunctulata Muller
quadrimaculata Samouelle
var. sexpunctata Gerh.
ab. bifasciata Schulze.
incequalis Fab.

subsp. rugulosa Port.
subrugata Cherv. nom.
nud.

acc. bicolorata Hatch
ramosa Say
cervaria Mann.
cenescens Casey

Sub genus Thanatophilus s. str.
lapponica Hbst.
tuberculata Germ,
subsp. caudata Say
calif ornica Mann,
subsp. granigera Cherv.
tritub erculat a Kby.

sagax Mann.
coloradensis Wick.

obalskii Port.
truncata Say

mexicana Cherv. in litt.

Blitophaga Keitt.
opaca L.
hirta Schaff.
villosa Naezen
tomentosa Villers
var. samnitica Fiori
var. binotata Port.
bituberosa Lee.

Nicrophorini

Nicrophorus Fabricius

Subgenus N ecrochar is Portevin
carolinus Fab.
mediatus Fab.
ab. mysticallis Ang.
ab. scapulatus Port,
ab. dolosus Port,
ab. floridee Hatch
ab. krautwurini Hatch
ab. lunulatus Hatch
ab. nebraskee Hatch

Subgenus Nicrophorus s. str..
orbicollis Say
halli Kby.

quadrisignatus Cast.

4

Journal New York Entomological Society

[Vol. LII

humator Fab.
sulactus Fisch.
paratype of grandior
Angell

var. atricornis Meier,
ab. maculosns Meier,
ab. rubroplearalis Delah.
delahoni Schilsky i. lift,
ab. signiceps Delab.
subsp. tenuipes Lewis
sayi Cast.

lumulatus Gistel
lunatus Lee.
luniger G. & H.
marginatus Fab.
requiscator Gistel
montezumce Matt,
ab. cordiger Port,
ab. sanjuance Hatch
ab. engelhardti Hatch
ab. leachi Hatch
vespilloides Hbst.
mortuorum Fab.
fractus Port,
ab. andrewesi Port,
ab. aurora Motch.
hebes Kby.
pygmeeus Kby.
vespilloides Lee. (nec.
Hbst.)

defodiens Mann.
disjunctus Wil.-Ellis
ab. humeralis Hatch
ab. Iristis Port,
ab. steinfeldi Smirnov,
ab. maculatus Wil.-Ellis
ab. altumi Westh.
ab. subfasciatus Port.

ab. sub interrupt us Pic.
var. borealis Port,
var. sylvaticus Reitt
ab. sylvivagus Reitt
ab. ruber Hatch
ab. nearticus Hatch
ab. nicolayi Hatch
ab. oregonesis Hatch
subsp. defodiens Mann.
nunemacheri Hatch (nec.

Port.)

ab. binotoides Hatch
binotatus Hatch (nec.

Port.)

ab. conversator Walk.
defodiens var. b. Mann.
lateralis Port.
pollinctor Lee. (nec.

Mann.)

ab. pacificce Hatch
ab. walkeri Hatch
conversator Port. (nec.

Walk.)

ab. gaigei Hatch
ab. kadjakenis Port,
ab. mannerheimi Port,
ab. binotatus Port.
plagiatus Mots.
vespillo L.
vulgaris Fab.
cadaver inus Gravenh.
curvipes Duftschm.
ab. faureli Fauconnet
ab. varendorffi Westh.
ab. bolsmanni Westh.
ab. cethiops Scheicher
ab. minor Westh.
ab. germani Hatch

Mar., 1944]

Arnett: Silphid^e

5

americanus Oliv.
grandis Fab.
virginicus Frol.
melsheimeri Kby.
hybridus Hatch & Angell
var. minesotianus Hatch
nigritus Mann.

var. ruficornis Mots.
pustulatus Hers.
bicolor Newn.
tardus Mann,
ab. coloradensis Hatch
ab. noveboracensis Hatch
ab. fasciatus Port,
ab. unicolor Port.
investigator Zett.
ruspator Er.
infodiens Mann.
confossor Mots.
micro cephalus Thoms.
pustulatus Horn. (nec.

Hers.)

labiatus Mots.
vestigator Gyll. (nec.

Hers.)

subsp. investigator Zett.
ab. suturalis Mots.
infodiens var. b. Mann,
ab. funeror Reitt.
ab. funerator Fanr.
var. variolosus Port,
ab. intermedins Reitt.
ab. jamezi Hatch
ab. lutescens Port,
subsp. maritimus Guer.
aleuticus Guer.
pollintor Mann.
sibiricus Mots.

infodiens var. c. Mann,
ab. martini Hatch
ab. clarencei Hatch
ab. sitkensis Hatch
ab. massetti Hatch
ab. grahami Hatch
ab. charlottei Hatch
ab. particeps Fisch
ab. japani Hatch
tomentosus Web.
velutinus Fab.
ab. communis Hatch
ab. elongatus Hatch
ab. angustefasciatus Port,
ab. splendens Hatch
ab. brevis Hatch
var. aurigaster Port.
germanicus L.
listerianus Fourer
ab. specious Schultze
ab. bimaculatus Steph.
ab. frontalis Fisch.
ab. fassifer Reitt.
ab. apicalis Kraatz
var. ruthenus Mots.
grandior Ang.
guttula Mots.

subsp. guttula Mots,
ab. ruficornis Mots,
ab. sanfranciscce Hatch
ab. punctatus Hatch
ab. shastce Hatch
ab. hypomerus Hatch
ab. lajollce Hatch
ab. vandykei Ang.
ab. quadriguttatus Ang.
ab. kuschei Hatch

6

Journal New York Entomological Society

[Vol. LII

mexicanus Matt.
hecate Bland
ab. wallisi Hatch
ab. California Hatch
ab. intermedins Hatch
ab. disjunctus Port,
ab. woodgatei Hatch
ab. phoenix Hatch

ab. novamexicce Hatch
ab. rubripennis Port,
ab. rubrissimus Hatch
ab. immacnlosis Hatch
ob scums Kby.
melsheimeri Lee.
ab. discontinus Hatch
ab. ruber Hatch

Exotic Forms and Their Relationship to Neartic Species. —
The following list of species are exotic forms of which the female
genitalia have been studied. The first name in each group is that
of the type for the genus or subgenus, or it is a typical neartic
species of that group. The next names are those studied with
the generic or subgeneric name as used by other authors follow-
ing it, if it differs from the names employed in this paper.

Silpha littoralis L.

Silpha cayennesis Sturn.
Silpha bigutatta Phil.
Necrodes bigutatta Phil.
Paranecrodes biguttata
Phil.

Thanatophilus ( Oiceoptoma )
americana L.
Thanatophilus thoracica

L.

Silpha thoracica L.
Thanatophilus japonica
Mots.

Silpha japonica Mots.
Thanatophilus obscura L.

Silpha obscura L.
Thanatophilus carinata
Hbst.

Silpha carinata Hbst.
Thanatophilus Icerigata
Fab.

Silpha Icerigata Fab.
Thanatophilus perforata

Gbl.

Silpha perforata Gbl.
Thanatophilus atrata L.

Silpha atrata L.
Thanatophilus nigrita
Creutz

Silpha nigrita Creutz
Thanatophilus granulata
Oliv.

Silpha granulata Oliv.
Thanatophilus ( Thanatophi-
lus) truncata Say
Thanatophilus sinuatus
Fab.

Silpha sinuatus Fab.
Thanatophilus terminata

Hum.

Silpha terminata Hum.
Thanatophilus rugosus L.
Silpha rugosus L.
Blitophaga opaca L.

Blitophaga oblong a Kust.

Silpha oblong a Kust.
Blitophaga souverbiei
Fairm.

Mar., 1944]

Arnett: Silphid^e

7

Silpha souveriei Fairm.
Blitophaga orientalis
Brnlle

Silpha orientalis Brnlle
Nicrophorus ( Nicrophorus )
vespillo L.

Nicrophorus prcedator

Reitt.

Nicrophorus rotundicollis
Port.

Nicrophorus didymus
Brulle

Nicrophorus interruptus
Steph.

Synopsis of the Neartic Silphini and Nicrophorini. — The
following synopsis is meant to serve as a means of correlating the
external characters with those of the genitalia.

TEIBES

Antennas apparently of ten segments, the second segment being very short
and more or less hidden in the tip of the first; elytra short and not cover-
ing more than the basal five tergitesl Nicrophorini

Antennae clearly of eleven segments, the second not shortened; elytra, if short,
covering more than the basal five tergites Silphini

Genera Silphini

A. Occipital ridge prominent; eyes usually large and prominent; form
usually elongate; labrum broadly emarginate; prothoracic spiracle

sometimes exposed Silpha L.

AA. Occipital ridge usually not prominent; eyes not large and prominent;
labrum broadly or narrowly emarginate; prothoracic spiracle rarely
exposed.

B. Eyes normal, protruding somewhat from the head; labrum broadly
or somewhat narrowly emarginate, but never very narrowly
emarginate unless the head is elongate; head normal or elon-
gate, not short, round or compact Thanatophilus Leach

BB. Eyes very small, not or only very slightly protruding from the
head; head short, round, and compact; labrum very narrowly
emarginate I Blitophaga Beitt.

SILPHA

Pronotum oval, black; elytra usually with red apical spots forming a bar,

sometimes with red basal markings or sometimes immaculate littoralis L.

Pronotum transverse, yellowish-brown with a black central area; elytra im-
maculate discicollis Brulle

THANATOPHILUS

Sub-genera

Labrum broadly or narrowly emarginate; pronotum rarely tomentose, if
tomentose, then orange with a black central area; head and mouth parts
often elongate Oiceoptoma Leach

8

Journal New York Entomological Society

[Vol. LII

Labrum always very broadly emarginate; pronotum usually tomentose, if
not, then elytra without prominent costse; head and mouth parts never
elongate Thanatophilus s. str.

Sub-genus Oiceoptoma

A. Pronotum orange or yellow with a black central area.

B. Elytra rugose americana L.

BB. Elytra smooth.

C. Costse prominent novaboracensis Frost.

CC. Costae obscure, elytra tan with four black spots and the scu-

tellum black quadripunctata L.

AA. Pronotum black.

D. Elytra smooth incequalis Fab.

DD. Elytra rugose ramosa Say

Sub-genus Thanatophilus

A. Pronotum tomentose; elytral costae prominent.

B. Intervals of the elytral costae tuberculate .lapponica Hbst.

BB. Intervals of the elytral costae flat.

C. Two inner elytral costae subequal throughout.

i tritub erculat a Kby.

CC. Two inner elytral costae nearly obsolete at the base.

coloradensis Wick

AA. Pronotum glabrous, costae obscure ; truncata Say

BLITOPHAGA

Surface pubescent ; form more elongate opaca L.

Surface sparsely pubescent; form more oval bituberosa Lee.

NICROPHORUS

A. Pronotum oboval, without distinct sculpturing and very narrowly mar-
gined (subgenus N ecrocharis) carolinus L.

AA. Pronotum orbicular, transverse or cordate with distinct sculpturing and
widely margined on the sides and back (subgenus Nicrophorus s. str.).

B. Pronotum orbicular, widely margined at the sides and the base.

orbicollis Say

BB. Pronotum not orbicular.

C. Pronotum sinuate at the sides, base nearly as wide as the
apex, sides and base widely margined, not cordate.

D. Metasternal epimeron tomentose.

E. Hind tibia curved.

F. Metatrochanter spine small and divergent;
pronotum disc orange, margin black ; front

orange .:.... americanus Fab.

FF. Metatrochanter spine large and convergent;
pronotum black; front black sayi Lap.

Mar., 1944]

Arnett: Silphid^e

9

EE. Hind tibia straight.

G. Spine of the metatrochanter obscure; elytra

immaculate nigritis Mann.

GG. Spine of the metatrochanter prominent.

H. Spine divergent; elytra with orange

fascae pustulatus Hersch.

HH. Spine convergent; elytra immaculate.

humator Fab.

DD. Metasternal epimeron glabrous.

I. Elytra immaculate; hind tibia© usually curved.

germanicus L.

II. Elytra with orange fasciae; hind tibia straight.

J. Three terminal segments of the antennae

black vespilloides Hbst.

JJ. Three terminal segments of the antennae
orange.

K. Metasternal pubescence brown; abdom-
inal pubescence black.

mexicanus Matth.
KK. Metasternal pubescence yellow; abdom-
inal pubescence brown.

investigator Zett.

CC. Pronotum with base much narrower than the apex, sides
strongly sinuate, cordate.

L. Metasternal epimeron glabrous melsheimeri Kby.

LL. Metasternal epimeron tomentose.

M. Thorax tomentose.

N. Thorax entirely tomentose.

tomentosus Web.

NN. Thorax tomentose apically only.

vespillo L.

MM. Thorax glabrous.

O. Basal segment of the antennal club black.

P. Hind tibia straight; disc of the pro-

notum punctate guttula Mots.

PP. Hind tibia arcuate; disc of the pro-
notum nearly smooth.

obscurus Kby.

00. Basal segment of the antennal club orange.

Q. Hind tibia arcuate marginatus Fab.

QQ. Hind tibia straight.

R. Ventral surface of the posterior
tibia densely yellow tomen-
tose hecate Bland.

RR. Ventral surface of the hind
tibia sparsely black tomen-
tose hybridus Hatch & Ang.

10

Journal New York Entomological Society

[Yol. Lll

General Morphology of the Genitalia. — The same terminol-
ogy as that adopted by Tanner, which seems to be a usable inter-
pretation of the relationship of the parts, has been used here.

The dorsal plate or proctiger (PL I, Fig. 3, p.) forms the
upper surface of the genitalia. It has a terminal process (PI.
II, Fig. 8, pro.) sometimes elongate and spatulate, and may
be bent at various angles. Frequently it has terminal hairs. It
apparently serves as the dorsal guide. The paraprocts (PI.
I, Fig. 3, pp.) are lateral plates forming the sides and bottom
of the organ. They sometimes bear setae. The paraproct bears
the valvifer (PI. I, Fig. 3, vf.) which in turn bears the coxite
(PI. I, Fig. 3, c.). The valvifer is sometimes modified into a
lateral guide (PL II, Fig. 8, l.g.) appearing claw-like and lobed
(Pl. II, Fig. 8, 1.) or it may be unmodified and possess setae. The
coxite is a hollow process which supports the stylus (Pl. I, Fig. 3,
sty.) either terminally or laterally on the margin. The stylus is
of various sizes and length and in some species it is expanded at
the apex.

The proctiger is interpreted as a part of the tenth abdominal
tergite. The paraprocts are probably parts of the ninth tergite
with the styli, coxites and valvifers as appendages of the ninth
segment. In this study only the ninth and tenth segments have
been considered. The eighth segment consists of two more or
less unmodified plates, the tergite and sternite.

Key to the Neartic Silphini and Nicrophorini Based
on the Female Genitalia

1. Valvifer at most only slightly lobed, not developed into a curved proc-

ess; proctiger never extended and lobed, ( Silphini ) (2).

Valvifer with a well developed curved process and the proctiger usually
extended and lobed. ( Nicrophorini one genus Nicrophorus Fab.)

(14).

2. Stylus apical or lateral; coxite without a lateral projection (3).

Stylus lateral; coxite with a lateral projection. ( Blitophaga Beitt.)

(12).

3. Stylus always apical, stout, the diameter nearly that of the coxite; cox-

ite stout and more or less uniform throughout. ( Silpha L.) (4).

Stylus apical or lateral, if apical, then very small, much smaller than
the coxite and the coxite is much wider at the base than at the apex.
( Thanatophilus Leach.) . (5).

Mar., 1944]

Arnett: Silphid^

11

SILPEA

4. Coxite with two lateral connecting ridges; stylus broader at the apex.

littoralis L.

Coxite without ridges; stylus more uniform throughout.

discicollis Brulle

THANATOPHILUS

5. Stylus apical or lateral, if lateral then the coxite beyond the stylus is

not greatly flattened or lobed. Subgenus Oiceoptoma Leach (6).

Stylus always lateral; coxite beyond the stylus greatly flattened and
slightly lobed. Subgenus Thanatophilus s. str. (10).

6. Stylus apical 4-punctata L.

Stylus lateral , (7).

7. Stylus long and angulate, nearly reaching the apical end of the coxite.

americana L.

Stylus short and not angulate, and much shorter than the portion of
the coxite beyond the insertion of the stylus (8).

8. Apical portion of the coxite beyond the base of the stylus twice the

length of the stylus or less (9).

Apical portion of the coxite beyond the base of the stylus much more
than twice the length of the stylus ramosa Say

9. Bridge (PL I, Pig. 3, br.) between the coxite and the valvifer wide;

proctiger broadly rounded apically; stylus round at the apex.

novdboracensis Forst.

Bridge between the coxite and the proctiger narrow; proctiger more

angular apically; stylus angulate at the apex inaequalis Fab.

L0. Stylus greatly enlarged at ‘the apex, more than twice the width of the

basal portion truncata Say.

Stylus enlarged at the apex, but much less than twice the width of the
basal portion (11) .

11. Coxite with a prominent basal spine below the insertion of the stylus.

coloradensis Wick.

Coxite without a prominent basal spine below the insertion of the
stylus (12).

12. Stylus inserted on the ventral surface of the coxite so that there is

apparently a lateral flap covering the base of the stylus.

lapponica Hbst.

Stylus inserted on the lateral surface of the coxite tritub erculat a Kby.

BLITOPHAGA

13. Stylus as long as the lateral lobe of the coxite bituberosa Lee.

Stylus much shorter than the lateral lobe of the coxite opaca L.

NICROPHORUS

14. Coxite with a terminal claw as long or longer than the stylus. (Sub-

genus Neocrocharis Port.) carolinus L.

12

Journal New York Entomological Society

[Vol. Lir

Coxite without a terminal claw, or if apparently present, then always
much less than the length of the stylus. (Subgenus Nicrophorus s.
str.) (15).

15. Proctiger lobe short and broad (16).

Proctiger lobe long and narrow, without an apical spatula.

marginatus Fab.

Proctiger lobe medium in length and width, with or without an apical
spatula (18).

16. Proctiger lobe bifurcate orbicollis Say

Proctiger lobe not bifurcate : (17).

17. Proctiger lobe truncate, without prominent apical ridge humator Oliv.

Proctiger lobe round, with prominent apical ridge. (PI. II, Fig.

8, r.) ...... sayi Lap.

18. Proctiger without an apical spatula vespilloides Hbst.

Proctiger with an apical spatula (19).

19. Coxite with a basal-lateral lobe; lobe of the claw of the valvifer longer

than wide vespillo L.

Coxite without a basal-lateral lobe; lobe of the claw of the valvifer
always broader than long (20).

20. Coxite emarginate on the inner lateral margin americanus L.

Coxite not emarginate \ (21).

21. Lobe of the proctiger sub-truncate (22).

Lobe of the proctiger round (24).

22. Lobe of the proctiger greatly curved dorsally-ventrally (23).

Lobe of the proctiger slightly curved dorsally-ventrally nigritus Mann.

23. Spatula of the proctiger lobe broad melsheimeri Kby.

Spatula of the proctiger lobe narrow .• hybridus Hatch & Ang.

24. Lobe of the valvifer claw obscure . ......... (25).

Lobe of the valvifer claw prominent (27).

25. Coxite very narrow hecate Bland.

Coxite broad , (26).

26. Proctiger lobe greatly curved dorsally-ventrally (28).

Proctiger lobe slightly curved dorsally-ventrally (29).

27. Lobe of the claw of the valvifer with setae, spatula ridged.

germanicus L.

Lobe of the claw of the valvifer without setae, spatula not ridged.

pustulatus Hers.

28. Apical margin of the base of the valvifer concave and nearly parallel

with the basal margin mexicanus Matt.

Apical margin of the base of the valvifer nearly straight and not paral-
lel with the basal margin tomentosus Web.

29. Spatula of the proctiger lobe oval guttula Mots

Spatula of the proctiger lobe round (80).

30. Valvifer, exclusive of the lobe, square obscurus Kby.

Valvifer, exclusive of the lobe, trapizoidal investigator Zett.

Mar., 1944]

Arnett: Silphid^e

13

DISCUSSION OF THE NEARTIC SILPHINI AND NICROPHORINI

General. — The tendencies pointed out here are based only on
the species considered in this study and without comparison with
other beetles.

On the basis of the female genitalia alone, the Silphini appear
to be the more primitive of the two tribes and have been treated
so here. But this conclusion is based on the assumption that
simplicity of form, as seen in the Silphini, indicates primitive-
ness rather than reduction. The Nicrophorini are so closely
linked together that it is difficult to tell anything about their
phylogeny.

Silphini

The proctiger is simple, never lobed, usually with setae; the
paraproct is simple, essentially the same as that of the Nicro-
phorini, but with setae. The valvifer at most is lobed only,
usually with setae; the coxite exhibits the greatest variation of
the organ. It may have a basal lobe, or be uniform throughout.
The stylus is attached to the coxite either terminally or laterally.
The stylus is usually uniform in shape, but in some groups it is
enlarged at the apex.

Silpha L.

Type : Silpha littoralis L., designated by Latreille 1810.

In this genus the stylus is terminal and stout, and is wider at
the apex than at the base. The coxite is stout and uniform
throughout. The proctiger, paraprocts and the valvifers are
unmodified.

Silpha littoralis L.

This species is supposedly European, but the characters used
to separate it from the Neartic surinamensis Fab. do not ade-
quately separate the two. In the collection of the author there
are specimens with immaculate elytra, and the genitalia of the
two forms show no differences. Therefore surinamensis Fab. is
a form of littoralis L. and not a distinct species. The genitalia
of this species has a setigerous proctiger. The coxite is stout and
with two ridges on the outer lateral surface which connect at
the apex.

Silpha discicollis Brulle.

14

Journal New York Entomological Society

[Vol. LII

Proctiger with setae; coxite without lateral ridges; stylus less
enlarged at the apex.

Thanatophilus Leach

Type : Silpha rugosa L.

Coxite blade-like, flattened or uniformly triangular; stylus
terminal or lateral, uniform throughout or enlarged at the apex.

Subgenus Oiceoptoma Leach

Type : Silpha thoracica L.

Coxite uniformly triangular with the stylus terminal, varying
to coxite slightly flattened apically and the stylus lateral ; stylus
uniform throughout. Species as described in the key and
synopsis.

Subgenus Thanatophilus s. str.

Coxite flattened at the apex, appearing blade-like and strongly
curved on the outer side ; stylus lateral and enlarged at the apex.
The characters of the species are as presented in the key and
synopsis.

Blitophaga Eeitt.

Type : Silpha opaca L.

Coxite with basal lobe or tooth, terminal portion narrow and
flattened; stylus small and lateral between the basal lobe and
the apex of the coxite, never longer than the basal lobe. The
characters of the species are as presented in the key and the
synopsis.

Nicrophorini

The greatest difference between this tribe and the Silphini is
in the modification of the proctiger. Here the proctiger is
usually greatly extended and generally spatulate at the apex,
nearly always with setae. Also, the valvifer is quite different in
appearance from that of the Silphini. It is greatly enlarged
and extended, flattened and claw-shaped. The coxite is uniform,
bearing the stylus terminally in all cases except Nicrophorus
carolinus L. which has a lateral stylus. The paraproct is with-
out setae, but at times is ridged. The species of this genus are
very closely related with the exception of Nicrophorus carolinus

Mar., 1944]

Arnett: Silphid.®

15

L. which shows characters differing from the others and is placed
in the subgenus Necrocharis Port.

Nicrophorus Fab.

Characters the same as those of the tribe. Type : Nicrophorus
vespillo L., designated by Latreille, 1810.

Subgenus Necrocharis Port.

Type : Nicrophorus Carolina L., one species only with charac-
ters as in the key and the synopsis.

Subgenus Nicrophorus s. str.

The species of the subgenus are all so closely related that they
cannot be separated into species groups. The characters used
in describing the species are inadequate. Color pattern has
little or no value in separating the majority of the species.
Their relationships depend entirely on what set of characters
are used. Many aberrations have been described, but this is
quite unnecessary and becomes extremely confusing, especially
when they are not illustrated. Because of the great variation in
the color pattern, almost any population can be described as a
new aberration. Some changes have been made in the status of
certain forms. Undoubtedly, when other forms are examined,
more changes will be necessary. The following changes have
been made on the basis of the characters presented in the key and
synopsis :

Nicrophorus melsheimeri Kby. is a distinct species and not a
synonym of investigator Zett.

Nicrophorus nigritus Mann, is a distinct species and not a sub-
species of investigator Zett.

Nicrophorus hecate Bland, is a distinct species and not a sub-
species of guttula Mots.

CONCLUSIONS

1. The female genitalia of Silphini and Nicrophorini present
characters which serve to separate the species of the groups.

2. The two tribes have basically the same type of female geni-
talia, but they are two very distinct groups of genera.

3. Silphini tends to be more primitive than Nicrophorini.

16

Journal New York Entomological Society

[VOL. LII

4. The genera Silpha, Thanatophilus , Blitophaga and Nicro-
phorus are distinct groups of species.

5. The color patterns of Nicrophorus are not good specific
characters and aberrations based on those characters are worth-
less.

6. The form and sculpturing of the elytra of Silphini do not
show relationships between the species.

7. Nicrophorus offers no distinct species groups, and the spe-
cies of the genus are very closely related.

BIBLIOGRAPHY

Hatch, M. H. Studies on the Silphinae. Jour. N. Y. Ent. Soc., XXXV,
331-370, 1927. Coleopterorum Catalogus, Part 95, Junk, Berlin,
1928.

Hatch and Reuter. Coleoptera of Washington, Silphidae. Univ. of Wash.
Publ. in Biol., 1: 147-162, 1934.

Horn, G. H. Synopsis of the Silphidae of the U. S. with reference to the
genera of other countries. Trans. Am. Ent. Soc., 8: 219-322, pi.
V-VII, 1880.

Latreille, P. H. Considerations Generales sur Pordre Naturel des Ani-
maux. Paris, 1810.

Leng, C. W. Catalogue of the Coleoptera of America, North of Mexico and
suppl. 1-4. Sherman, Mount Vernon, N. Y., 1920.
Scegoleva-Barovshaja, T. Les Necrophorini (Coleoptera, Silphidae) de la
faune de PU.R.S.S. Travaux de l’lnstitut Zool. de PAcademei des
sciences de PU.R.S.S., 1: 161-191, 1932.

Semenov-Tian Shanskij, A. P. De Tribu Necrophorini (Coleoptera, Sil-
phidae) classification et de ejus distributione geographica. Travaux
de l’lnst. Zool. de PAcademie des Science de PU.R.S.S., 1: 149-
160, 1932.

Tanner, V. M. A Preliminary Study of the Genitalia of Female Coleop-
tera. Trans. Am. Ent. Soc., 53, 5-50, 1927.

Mar., 1944]

Arnett: Silphid^e

17

Abbreviations on Plates

vM

proctiger

spa

spatula

pp

paraprocts

1

lobe

sty

stylus

r

ridge

c

coxite

br. ...

bridge

vf

valvifer

pro

process

lateral guide

18

Journal New York Entomological Society

[Yol. LI i

PLATE I

Figure la. Silpha L. Elytra of female.

Figure lb. Silpha L. Elytra of male.

Figure 2a. Nicrophorus Fab. Head of male.
Figure 2b. Nicrophorus Fab. Head of female.
Figure 3. Thanatophilus quadripunctata L.
Figure 4. Silpha littoralis L.

Figure 5. Silpha discicollis Brulle.

Figure 6. Thanatophilus americana L.

Figure 7. Thanatophilus novaboracensis Forst.
Figure 8. Thanatophilus incequalis Fab.

Figure 9. Blitophaga bituberosa Lee.

Figure 10. Thanatophilus tritub erculata Kby.

(Jour. N. Y. Ent. Soc.), Vol. LII

(Plate I)

10

20

Journal New York Entomological Society

[Yol. Lll

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

Figure

PLATE II

1. Thanatophilus ramosa Say.

2. Thanatophilus lapponica Hbst.

3. Thanatophilus coloradensis Wick.

4. Thanatophilus truncata Say.

5. Blitophaga opaca L.

6. Thanatophilus thoracica L.

7. Thanatophilus rugosus L.

8. Nicrophorus carolinus L.

9. Nicrophorus orhicollis Say.

10. Nicrophorus vespilloides Hbst.

(Jour. N. Y. Ent. Soc.), Vol. LII

(Plate II)

10

22

Journal New York Entomological Society

[Vol. LII

Figure 1.
Figure 2.
Figure 3.
Figure 4.
Figure 5.
Figure 6.
Figure 7.
Figure 8.

PLATE III

Nicrophorus americana Fab.
Nicrophorus sayi Fab.
Nicrophorus nigritus Mann.
Nicrophorus pustulatus Hersch.
Nicrophorus humator Fab.
Nicrophorus germanicus L.
Nicrophorus mexicanus Matth.
Nicrophorus investigator Zett.

(Jour. N. Y. Ent. Soc.), Vol. LII

(Plate III)

7

24

Journal New York Entomological Society

[Vol. LII

Figure 1,
Figure 2,
Figure 3,
Figure 4,
Figure 5.
Figure 7
Figure 6.
Figure 8,

PLATE IY

Nicrophorus melsheimeri Kby.
Nicrophorus tomentosus Web.
Nicrophorus vespillo L.

Nicrophorus guttula Mots.
Nicrophorus ohscurus Kby.
Nicrophorus hecate Bland.
Nicrophorus marginatus Fab.
Nicrophorus hyhridus Hatch & Ang.

(Plate IY)

(Jour. N. Y. Ent. Soc.), Vol. LII

1 2

7

Mar., 1944]

Weiss et al.: Insect Behavior

27

GROUP MOTOR RESPONSES OF ADULT AND
LARVAL FORMS OF INSECTS TO DIF-
FERENT WAVE-LENGTHS
OF LIGHT

By Harry B. Weiss, E. E. McCoy, Jr., and
William M. Boyd

This, the fifth paper of a series, relating to the group behavior
of insects to colors, is concerned with the responses of seven
species of Coleoptera and of sixteen species of lepidopterous,
hymenopterous and coleopterous larvae to ten wave-length bands
of light of equal physical intensities, in disarray, from 3650 A
to 7400A. The tests were run in the sector type equipment de-
scribed in the third and fourth papers of the series.1 As outlined
in these papers, the insects were placed in an introduction cham-
ber, six feet away from the filter chambers, after the lamps were
on and after all filter chambers were open. After the exposure
period, the filter chambers, the central compartment, introduction
chamber and dark chamber were closed, and counts were then
made.

In all previous tests reported upon in the third and fourth
papers the color filters were arranged in a sequence beginning
with the shorter wave-lengths and extending successively to the
longer wave-lengths, as follows: 3650 A (ultra-violet); 4360 A
(violet-blue) ; 4640 A (blue) ; 4920 A (blue-blue-green) ; 5150 A
(blue-green) ; 5460 A (yellow-green) ; 5750 A (yellow-yellow-
green) ; 6060 A (yellow-orange) ; 6420 A (orange-red) ; and
7200 A (infra-red). Each filter chamber was separated from its
neighbor by a black chamber.

All tests reported in the present paper, with adults and larvae,
were made with the filters in disarray, as follows: 3650 A (ultra-
violet) ; 6060 A (yellow-orange) ; 4640 A (blue) ; 7200 A (infra-
red) ; 5150 A (blue-green) ; 6420 A (orange-red) ; 5750 A (yellow-
yellow-green) ; 4360 A (violet-blue) ; 5460 A (yellow-green) ;
and 4920 A (blue-blue-green). The wave-length figures repre-
sent the peak transmissions of the filters.

i Jour. N. Y. Ent. Soc., 50(1) : 1-35, 1942; 51(2) : 117-131, 1943.

28

Journal New York Entomological Society

[Vol. LII

Owing to the deterioration that occurred in the lamps used in
previous tests, new forty-watt, frosted, Westinghouse Mazda
lamps and a new General Electric Mazda mercury lamp (type
A-H4, 100 watts) were utilized for all tests. The same method,
outlined in our first paper,2 was used for determining the relative
positions of the lamps and various filter combinations so that the
physical intensities were approximately equal. A slight change
in technique, designed to improve the equalization, was recently
made and this resulted in a new set of distance settings, differing
slightly from those given in the first paper. These new distance
settings are shown on page 29.

RESULTS WITH ADULT INSECTS

Table I presents the results of exposing seven species of Cole-
optera to ten wave-length bands of light of equal physical intensi-
ties, in disarray, in the sector type equipment. By consulting
the percentage distribution of those reacting positively to the
various wave-length bands and by an examination of the group
behavior curves in Figures 1 and 2, it may be noted that the peak
response for all species except Popillia japonica took place in
the ultra-violet (3650-3663 A) and that secondary peaks occurred
either in the blue-blue-green (4920 A) or in the blue-green
(5150 A). Smaller numbers, in general, appear to have gone to
4360 A (violet-blue) and larger numbers to 6060 A (yellow-
orange) than in previous tests when the filters were not in disar-
ray. Except for the somewhat reduced attractiveness of 4360 A
and the slightly increased attractiveness of 6060 A, the behavior
patterns, with the filters in disarray, did not differ materially
from previous patterns obtained with the filters in orderly array.

Peterson and Haeussler3 in their work with the Oriental fruit
moth and colored lights found that when a less attractive colored
light was placed at right angles to a more attractive colored one,
more fruit moths went to the less attractive light than when the
less attractive one was opposite the more attractive one. There is
no doubt that the stimulating power of certain wave-lengths is
influenced by their positions with respect to other wave-lengths.
But the fact remains that except for the slight differences noted

2 Jour. N. Y. Ent. Soc., 49(1) : 1-20, 1941.

3 Ann. Ent. Soc. Amer., 21(3) : 353-379, 1928.

Mar.,

1944]

Weiss et al.:

Insect Behavior

29

be

O

®

be

c3

"3

<x>

o

Pi

50

2

O

t>

o

&

a

o

10%

• : rH GO rH rH C3 ^ OO CO

|= : CO cq* rH CO oi cq' o 03

^ :10^(M03 03(M<MH

20%

• COCOCqrHCCI>-aOt>;T— 1
§ oi lO 00 rH CO id cl rH* rH*

30%

• O t> CD x lO io t> o_ ©

r~ t> l> H O (> 05 05 GO* GO* rH

u ^NdHHr 1 i— 1

rH

£

o

rH

• 00 H IO m 05 rH CO rH 05 o

§ 03 CO t>* GO co id t- cd id 00

b CO CQ H I— 1 1 — 1 03

50%

• '00 x w IO N S CO X CO o

s th* 03 th" cd i-I cq" id rH rH cd

*C3 CO rH rH i— 1 !— 1 GO

&

o

co

• oo h in cq cq © co cq cq ©

| o i> n id o h d co d oo

^ CO i — 1 rH i — 1 H t-

70%

• CO O rH Cq © © rH 00 rH ©

§= oo id iH rH 03’ o' co' cq’ cq’ oo’

« cq rH rH rH CO

80%

. HCOH rH 03 03 CD OJ 00 IO

§ cd cd 0 co" in 00" cq’ cq' t-h cq*

^3 cq 1 — It — 1 CO

90%

. MHHOOOOHCOM O

§ rH cq’ 03’ cq' 00 cq' rH 1 h 00'

CJ cq rH 10

100%

• co co co cq cq cq co cq 03 0

§ cq" t-h 00’ cq’ cd rH* t-h* o' cq’

cq rH 10

IO N N S

IO r-H O c$

IO IO IO '

I -f I I .

NOOOOOOO^HH £>CO
05NSMCOCOWHH "GO
COaoasrHrHrHlOlOCO " IO

I I I I I I I I I g I

rHCOlOCOOCqOOOOOO mQO
^^TjH^ioiocODaS ^-fCO
cqNcqcocoMmno i>-

CO

fi

<1

M

W

Eh

O

i— i w

h e

w £

H w
EH

o

Eh

<1
P3
■ H
&H
P

o

H

P

O

O

o

o

ft

&0

s *

■rt 02
■S rP

£ s

<X>

<X> i-H

Jg

«H *
O

3-U99

Y

00 ZL

O O iH rH O O O

o

o o

rH rH

^U99

•tad

Y

0Sb9

HHNCQOHO

03 IO

to rH

^U99

J9d

© © t> t> IO 1> C3

to

^ CO

b- Ttf

Y

0909

rH

^U99

I9d

Y

oezs

03 CO CO "HH tJH 03 03

co

O tJH

b- O

q.U99

I9d

Os rJH tO t>- to CO

to

b~ LO

<M rH

Y

09bS

rH

q.U99

J9d

OS H t- tO IO CO 03

o

co

03 IO

Y

OSIS

H ri i — 1 i — 1 i — 1 l — i

rH

rH

03

^U99

J9d

to to OS IO l> o to

to

io to

CO <M

Y

0S6t

i — 1 03 i — 1 rH 03 rH rH

03

rH

!^U99

J9d

OHl>^M>CO

to

io CO

to 03

Y

0f9f

rH

rH

^U90

I9d

Ttl to IO io to 00

IO

o o

b- 03

Y

098k

03

q.U99

I9d

CO C3 CO t> O Tfi

Tin

CO 03

tO 03

Y

0S98

IO CO CO co CO ^ to

CO

30 CO

rH O0

St[^SlI9'[-9A'BA4. Cq
Supxona.i q.u99 19^

J9qU99 UI ^U99 ig^

igqranqo uorpmp

-OJ^UI UI ^U99 J9<J

igqureqo qouiq

UI ^U99 JL9J

sg^nunu gmsodxg;

P9AJ0AUI S^99S
-UI *0U IBXOJQ

S^S9; *ojq

g OS
<3 -g
h?- c3

CH rg

t3

a

c$

O0 CD 03 CO tH 00 O0
Tti rH CO CO Tti ^ 03

30 5D 1>

r-t 03 co

CO 03 CO tJH 30

«ffi05O!D00N

^HMOJOOCO

os os io

1^ tJH tH

30 03 io

co tH «o ^ Tt< co

CO CO CO CO CO CO CO

”T 'T 'T T T "T 'T

to oo co ^h t— co oo

03 Cvl

I I 1 I I I I

30 30 tO to tO tO t—

I I
to b-

^ £ P~> K

c3 c3 O o3 c3 c3

wwznwznw

© © © © © ©

U u V HH +H *N
© © © © © © o

■S -8 -6 -I -8 -I (2 5 -g,

sllslsg | I

» m © m © » ? ft ©

t5 © O O O O 'S o «*

© © © © © © a e c-

r©r©rer©r©r©| g |

©©©©©©© j5 ^

££££££» 8~ |

S-SSSS-SJiS Set,

©©©©©©©O ftrg o

LJ •<?=» H #«S> *<S> t>s CO /-s Q M

^ IO io io io io to si >■*

m ftftftftftftg^ £ ©
^©©©©©as^Sas ST'r©

o o P

s

30 b-
O GO
00 rH

03 CO CO CO

T TT T T

a

fc.

as

£

as

o

as

Q

§

to

>'§

©

CO

©

©

as

ft

•5 _g ^ o

© ri ft"*-*
H© g © 3

Ah

as

m

Peak intensities of bands.

Mar., 1944]

Weiss et al.: Insect Behavior

31

above the group behavior patterns for the species tested remained
materially unchanged with the filters in disarray. Regardless of
the relative positions of the various wave-length bands, the insects
made approximately the same selections time after time.

RESULTS WITH LARVAL FORMS
Most of the experimental work on the behavior of insects to
colored light has been done with adult insects. Nevertheless
there are a few references in the literature to the behavior of
larval forms and mention will be made of the photopositive ones.

For example, Mayer and Soule found that the larvae of Danais
archippus are photopositive to ultra-violet. Gross,4 in his study
of the reactions of arthropods to monochromatic lights of equal
intensities reported that the larvae of Zeuzera pyrina, a lepidop-
terous wood-borer, and of a noctuid moth Feltia subgothica, are
photopositive to colors, the order of the effectiveness of stimula-
tion being blue (4200-4800 A), green (4900-5500 A), yellow
(5700-622 A), and red (6300-6500 A). Lymantria larvae, ac-
cording to Hundertmark5 appear to prefer blue when different
colors are compared. Gotz,6 in his study of the perception of
color and form in lepidopterous larvae found that an appreciation
of color occurs in the larvae of Vanessa and Pieris. These are
attracted by the green color of leaves or pieces of paper, regard-
less of the color of the surroundings, but more so on a white back-
ground than on a black one. Lammert7 reports that caterpillars
will go toward a source of light after a blacking of their eyes.
And Suffert8 states that many caterpillars colored like their sur-
roundings and feeding in exposed situations, orient themselves so
that the light always falls upon them from a particular angle.
These last two instances indicate the possession of a dermal light
sense.

Our tests as reported in the present paper involved the ex-
posure of the larvae of sixteen species of insects to ten wave-length
bands of light, of equal physical intensities, from 3600 A to 7200
A. These bands were in disarray. From Table II and Figures
4 Jour. Exp. Zool., 14: 467-512, 1913.
s Z. vergl. Physiol., 24 : 563-582, 1936.
e Z. vergl. Physiol., 23 : 429-503, 1936.

7 Z. vergl. Physiol., 3 : 225-278, 1925.

8Z. Morph. Oekol. Tiere., 26: 147-316, 1932.

TABLE II

Behavior of Sixteen Species of Larvje to Ten Wave-length Bands of Light, or Colors, in Disarray

b£)
S *

Q5 "So

05

05

I!
^ £

^U90 J0<J

Y 00 ZL

1J.H00 J9(J

Y 0Zf9

^U00 10^

Y 0909

^J.1100 J9J

Y 02Z2

^H00 J0(J

Y 09t2

^H00 ja^

Y 0212

^U00 J0J

Y 0 Z6f

^U00 J0<J

Y 0f9f

!}H99 .I0<J

Y 098b

^H00 «[0<J

Y 0298

Sq^SuapOAHAl Cq
Sup0R9J ^U00 J0,£

J0^H00 III q.U00

aaquinip noppmp
-oj^ui in ^H00

ts

Pi pT

a

Pi

pj

pT

fH

£ £

bC

O O

0

0

0

O

O

«w

be be

be

be

Pi

be

u

tS)

Pi

be

4^>

3 3

3

3

Ml-#

I

co|^

|

3

HW

ft

1

H|M

A

ft

0

0 0

rH

rH

O

0

0

CO

0 0

ft

05

CO

ft

rH

ft

05 CO

ft

ft

ft

VO

rH

ft

O rH

05

- CO

ft

O

0

rH

ft ft

05

00

10

CO

05

rH

rH

O rH

05

05

CO

co

tH

r— 1

CO <M

CO

05

rH

rH

tH

co

co co

CO

05

05

CO

VO

CM

05

05

05

05

0°

co 10

co

ft

IO

CO

rH

* IO

CO ft

10

CO

OO

ft

ft

rH

05

CO CO

ft

ft

CO

05

GO

<M

ft IO

05

rH

05

ft

IO

ft 00

05

05

vo

O

ft

05

05 co

05

ft

co

05

ft

CM

co 0

co

IO

CO

1 1

00

co

CO CO

ft

ft

co

t-

CO

T— 1

ft 05

CO

rH

rH

05

10

T— 1

05

tH

rH

jDquinqo qaiqq
HI q.H00 J0(J

S03.HH11H 0insodxg;

p0A|OAHI S^09S
-HI "OU XR^OJL

co 10

CO CM
ft L-

O- 00
CO O CO

05 ft CO

S^S0^ *0^

05-2

S 05
ft

Pi

C3

ft

o

£ 05

6.3

05

O

CO

^ co

12 4

e 1

■is co

0

Si kT

1 ^
g®

cj

05

ft

O

Pi

II

ft

00 $

3 t

|S

§■-

lH

ft

o

’ft

05

A

^CO

1:

3 co 05

■ft^g

ft

ci

pH

co

3

s-

>5

03 CO

§ SC)

05

O

||T eT e r 88 It

§ oSS 14

| ^’g^’g g^®

W

Peak intensities of bands.

TABLE II — ( Continued )

£ P
bfi be

o o

bJO So
3 3

Ph Ph

P

o

P So
£

o S
So

o

So

° ?
bfi So

pH

bD

1=1 *

t> rP

g|o

f-l p

5x10a

Y 00 Zl

51X88 I8J

Y 03b9

51X80 JStJ

Y 0909

5X188 J8J

Y 09Z2

5X188 18^

Y 09b2

5X188 J8<5

Y 0212

51X88 I8<J

Y 0S6b

O rH O O

O 03 tH O

03 03 03

C00003OC0C0C003

Ocoi— I CO 03 CO 03 t>-

O lO 03 CO

lO CO t>-

5II88 J8(J

Y 0b9k

5X188 I8(J

Y 098k

5X188 18(5

Y 0298

sq5xfri8p8ABAV 05
Sux58'B8I 5X188 J8(5

J85U88 XIX 5X188 J8J

•xaqxcxBqo xxoppnp

-0J5HX XIX 5X188 18(5

CO rfl CO CO 03 t-

rHOlOlOCJOrHrJH

r^COr^OlOqlOCOiH

10 IQ IO CO CO

laqxxiBqo qoBjq
XXX 5X188 .18(5

S85iiuxxn smsodxg;

mioiooooioo

rtHC0rHC0rJHlO03C0

paAJOAXIX S588S

-hi -oxx 56505,

05 03 05 CO CO 1C LO

OOOOlOOCOOlrHCO
LOrtHCOrH'^i— I03i-H

S5S85 *o^[

£ ©
P

ci

8 I

*3 ^
£

CO 83 si co g co

^ TjH

i'll

■s^

£

7 1

co e

r*

I p.

CO o

‘t

l>- ^ 05

5^ CO |

ax s~

S> O rr| CO

en-H e

Pj I P JH

fel

C3

m

>

8

CO

CO

o r* o rH , ^

a- I M ffi S rjH
CO ^ Tj o I
^ T“H ^ rH *rH *K> Cd

<0 1 S> 1 bjo e 1

--05-01 P

Eh Eh ,p o
m

cot

.55 CO

«S rH

| 05

Cl P

csi P

Ah

Peak intensities of bands.

34

Journal New York Entomological Society

[Vol. Lll

3, 4 and 5 which record their group behavior in percentages and
graphically, it is apparent that the peak response for most species
took place in the ultra-violet (3650 A). In nearly all instances
a peak either equal to the one in ultra-violet or secondary to it
occurred in the blue-blue-green (4900 A), or in the blue-green
(5150 A). The minor peak at 6060 A (yellow-orange) is attrib-
uted to the disarray of the filters which resulted in this wave-
length being next to 3650 A (ultra-violet).

The larvae of Diacrisia virginica were the only ones which ultra-
violet light (3650 A) failed to stimulate appreciably. These
larvae are found crawling upon the ground or feeding upon low
plants. As a whole the group behavior of the larvae, in general,
did not differ from that of numerous adult insects, previously
tested, and their color discrimination, so called, was approxi-
mately the same as that exhibited by adult insects in spite of the
fact that their visual organs are less complex than those of adult
insects.

NOTES

Autoserica castanea. This beetle, being nocturnal, was tested
at 10 : 30 p.m.

Hippodamia convergens Guer. The predaceous larvae of this
coccinellid failed to react at all under the conditions of our tests.
When placed in the introduction chamber they climbed up the
sides and remained there. Apparently their negative geotropic
behavior predominated.

Hyphantria cunea Dru. The \ to f grown larvae of this
species, the fall webworm, made a web in the introduction cham-
ber and stayed there, even though they had been previously
deprived of food for twenty hours.

During the course of our work with larvae it was found that,
as a rule, they were more photopositive after having been de-
prived of food for a half-day or more previous to the tests. The
gregariousness of some of the species, especially of the larvae of
Hadena turbidenta Hbn., and Melalopha inclusa Hbn., appeared
to inhibit somewhat their sensitivity to light.

DISCUSSION

In view of the comparative simplicity of the lateral ocelli of
larvae, the similarity of the group behavior of larvae to that of

Mar., 1944]

Weiss et al.: Insect Behavior

35

adult insects with compound eyes is of considerable interest.
Although variable in structure, lateral ocelli in lepidopterous
larvas consist of a group, each ocellus having a structure not un-
like the single ommatidium of a compound eye. In the larvse of
sawflies and of many Coleoptera, the ocellus, of which there is
only one on each side, is a lens-like, transparent thickening of the
cuticle with underlying epidermis, and retinulse, each made up
of two or three visual cells grouped around a rhabdom. These
visual cells may be pigmented, or there may be separate pigment
cells. Dethier in a recent study9 of the corneal lens in caterpil-
lars states that “the cornea possesses a short focal distance, great
depth of focus, and an extremely low f value permitting the
admittance of much light. ’ ’

Although the king-crab, Limulus polyphemus, is not an insect,
the work of Hartline and Graham on the nerve impulses and
responses of single visual sense cells, to light, in the eye of this
animal is of unusual interest and it is within the realm of possi-
bility that a similar process of photoreception may operate in
insects.

The lateral facetted eye of the king-crab contains about 300
large ommatidia and the optic nerve fibres come directly from the
receptor cells with no intervening neurones. These authors10
studied the nerve impulses and developed a technique by which
was recorded the discharge from a single receptor unit, in the
form of oscillograms, representing the potential changes between
the cut end and an uninjured portion of the nerve, upon stimu-
lation of the eye by light. The electrical activity in the optic
nerve brought about by this stimulation was amplified by a vac-
uum tube and recorded by an oscillograph. Among other things
the stimulation of a single ommatidium resulted in a small strand
of the optic nerve showing a regular sequence of nerve impulses.
‘ 4 The discharge in a single fiber begins after a short latent period
at a high frequency, which has been found to be as high as 130
per second. The frequency falls rapidly at first, and finally
approaches a steady value, which is maintained for the duration
of illumination” (Hartline and Graham).

9 Jour. Cell, and Comp. Physiol., 19(3) : 301-313, 1942.

10 Jour. Cell, and Comp. Physiol., 1(2) : 277-295, 1932.

36

Journal New York Entomological Society

[Vol. LII

In a later paper11 these authors studied the responses of single
visual sense cells to visible light of different wave-lengths. This
was done by means of single fiber preparations from a Limulus
eye. It was found that when the energy of the stimulating light
of different wave-lengths was approximately equal, the response
to green was stronger than the responses to either violet or red.
When the energy was increased in the red and violet their level
of response was raised and when the intensities for the different
wave-lengths were adjusted so that the responses were equal,
there was no effect of wave-length as such, indicating that single
sense cells can gauge brightness but cannot distinguish wave-
length. The relative energies of the various wave-lengths re-
quired to produce the same response after being adjusted in in-
verse ratio to the degree to which they are absorbed yielded a
visibility curve, for a single visual sense cell, that had its maxi-
mum in the green near 5200 A and that declined symmetrically
on each side to low values in the violet near 4400 A and in the red
near 6400 A. According to the interpretation of visibility curves
by Hecht and Williams12 the stimulation of a single visual sense
cell by light depends upon the absorption spectrum of the pri-
mary photosensitive substance. The absorption of light by this
substance varies with wave-length and the production of a given
response needs a certain amount of photochemical change, which
in turn requires the absorption of a constant amount of energy.

Hartline and Graham also found that in the same eye of Limu-
lus there was a differential sensitivity among optic nerve fibers
and their attached sensory cells for different regions of the visible
spectrum and they believe that such specialization of the visual
cells, coupled with integrated action may give rise to color vision.

In considering the tests with insects reported upon in this and
in previous papers,13 the following patterns of behavior prevailed
over and over, when various species were exposed to ten wave-
length bands of equal intensities from 3600 A to 7200 A. In the
composite behavior14 of 5,454 insects of various orders, mostly

11 Jour. Gen. Physiol., 18(6) : 917-931, 1935.

12 Jour. Gen. Physiol., 5: 1, 1922.

13 JOUR. N. Y. Ent. Soc., 49 : 1-20, 149-159, 1941 ; 50 : 1-35, 1942 ; 51 :
117-131, 1943.

ii Ent. News, 54: 152-156, 1943.

Mar., 1944]

Weiss et al. : Insect Behavior

37

coleopterous, the peak response took place at 3650 A (ultra-
violet). From here the response declined gradually to a low
point at 4640 A (blue) ; then it increased to a secondary peak
at 4920 A (blue-blue-green), and then declined gradually to a
low point at 5750 A (yellow-yellow-green) from which point it
levelled off to 6420 A (orange-red). In the cases of individual
species there were deviations from this pattern. Drosophila and
various species of Coleoptera in some tests responded almost en-
tirely to 3650 A alone, dropping to a low level at 4360 A and
levelling off at that wave-length. Sometimes the secondary peak
occurred at 5150 A instead of at 4920 A. Although the peak
responses took place at 3650 A and 4920 A, small percentages of
the test animals went to other wave-lengths. In addition, it was
found that when a second test, using the same insects, succeeded
the first, the same group behavior pattern took place. The peak
response occurred at 3650 A, the secondary one at 4920 A. How-
ever, the individuals that made up the peaks in the second test
were not all the same as those making up the peaks in the first
test. In other words, there was a shifting of the individuals that
went to the different wave-length bands, but no difference in the
final result.

Assuming that the light receptors of insects function in the
same way as those of some other invertebrates and considering
the results obtained from the single visual sense cells of Limulus
as outlined by Hartline and Graham and mentioned above, it is
possible to venture an explanation for the group behavior pattern
of insect response to colors. Starting with the fact that the test
insects responded in varying numbers to all wave-lengths from
3650 A to about 6420 A, it is apparent that the photosensitive
substance of their visual sense cells will function at any of the
wave-lengths between 3650 A and 6420 A, if the physical intensity
of the wave-length is sufficient and constant.

When confronted by ten wave-length bands of equalized inten-
sities which converged upon the insects in the introduction cham-
ber of the apparatus, the primary photosensitive substance of the
visual sense cells of a large number absorbed the energy at 3650 A
to a greater extent than the energy at other wave-lengths. This
resulted in a photochemical reaction accompanied by physical

38

Journal New York Entomological Society

[Vol. LII

changes in nerve fibers, one of which was a change in the electric
potential of the point in the fiber that was actively responding
measured with respect to a nearby, but as yet, inactive point.
This electrical activity in the optic nerve fiber was transmitted
to the muscles where changes in tension occurred resulting in the
insects going to the ultra-violet (3650 A) in larger numbers than
to any other test wave-length. As the absorption of light by the
primary photosensitive substance of the single sense cell varies
with wave-length and as the production of a response requires a
certain amount of photochemical change plus a constant amount
of energy, it seems evident that the energy of the remaining test
wave-lengths although equal, was not sufficient to result in a
response that equalled that of the ultra-violet. Consequently
smaller numbers of individuals responded to the test wave-lengths
other than 3650 A. The question then arises as to why all indi-
viduals did not respond to 3650 A alone. In a group of 100 or
more insects collected in the field, it is not expected that they
would all be in the same physiological state at the same time. In
fact, when tested, only some are photosensitive. Others remain
in the introduction chamber, others get as far as the central com-
partment and others go to the black chamber, all exhibiting dif-
ferent degrees of behavior to light. Among those that are photo-
positive it is reasonable to assume that there exist some varia-
tions by individuals in the sensitivity of their visual receptors.
These variations may be connected with different physiological
states. They may be due to a depletion of the primary photosen-
sitive substance in the visual sense cells through the action of
light, resulting in individuals so affected responding in smaller
numbers to wave-lengths other than ultra-violet. Until restora-
tive processes take place in the visual sense cells of such individ-
uals, their sensitivity to ultra-violet declines.

Frequently various species, when tested, responded almost ex-
clusively to ultra-violet. But many others did not. In the case
of Drosophila which was bred under controlled conditions and
which were of uniform ages, the response to ultra-violet was
unusually high.

It is realized that these deductions are based upon the behavior
of single visual sense cells of the king-crab, to light of different

Mar., 1944]

Weiss et al.: Insect Behavior

39

wave-lengths and no consideration has been given to the fact that
individual cellular units act collectively and not independently,
nor to the fact that Graham and Hartline15 found that although
the visibility curves for single sense cells in the same eye are
approximately identical, they differ by significant amounts. In
addition they report that two sense cells were able to distinguish
violet from red and taking all these facts into consideration they
are of the opinion that such differential sensitivity “may be con-
sidered a peripheral mechanism of color vision.”

Another reason for the deductions as outlined consists of the
behavior of the Japanese beetle, Popillia japonica, which was
made to respond to what were unattractive wave-lengths under
equalized physical intensities, by increasing the intensities of such
wave-lengths. In fact with other species as well it was possible
to vary the behavior pattern by changing the intensities. And
in general, from our work over the past several years, it appears
that the behavior patterns of insects to equalized wave-lengths
are not unlike the behavior pattern of a single sense cell, in Limu-
lus, to equalized wave-lengths. Perhaps the behavior curves in
this and in former papers16 may be interpreted as rough approxi-
mations of the absorption spectrum of the photosensitive sub-
stance in the combined visual sense cells of many insects, as well
as indications of their motor responses to equalized wave-lengths
of light.

A word should be said about the comparatively large per-
centages of test insects which remain in the introduction chamber
and central compartment of our testing equipment. We have
always attributed this mainly to low illumination. At low illu-
minations only the most sensitive ommatidia function, there being
different thresholds of response for different ommatidia.17 By
others, a falling off in intensity discrimination, due to low illu-
mination is attributed to a nervous coupling of groups of omma-
tidia to form new units.18

is Jour. Gen. Physiol., 18: 917-931, 1935.

is Jour. N. Y. Ent. Soc., 49 : 1-20, 149-159, 1941 ; 50 : 1-35, 1942 ; 51 :
117-131, 1943.

i7 Hecht and Wald. Jour. Gen. Physiol., 17 : 517-547, 1934.
is Buddenbrock and Shultz. Zool. Jahrb. Physiol., 52: 513-536, 1933.

40

Journal New York Entomological Society

[Vol. Lll

The foregoing discussion is an attempt to explain the group
behavior patterns, or motor responses of insects to various wave-
lengths of light of equal physical intensities on the basis of the
results obtained by investigators who used single visual sense cells
of other invertebrates. It is realized that the motor response to
light of a complex organism such as an insect cannot be ade-
quately and definitely explained on the basis of the behavior of
single visual sense cells of other animals, nevertheless such work
as has been done with single sense cells furnishes valuable clues
to the phenomena of vision in insects. Until similar and addi-
tional investigations are made on the behavior of photoreceptor
cells and optic nerve fibers of insects, singly and in integrated
action, one has to be satisfied with implications.

PLATE Y

Figure 1. Behavior of six lots of Leptinotarsa decemlineata Say, to 10 wave-
length bands, in disarray, from 3650 a to 7200 A. Physical
intensities equalized.

1. 129 beetles. Three tests.

2. 478 beetles. Four tests.

3. 857 beetles. Six tests.

4. 857 beetles. Four tests.

5. 340 beetles. Four tests.

6. 338 beetles. Four tests.

Figure 2. Behavior of six species of Coleoptera to 10 wave-length bands, in
disarray, from 3650A to 7200 A. Physical intensities equalized.

1. Chrysochus auratus Say. 217 adults. Three tests.

2. Tetraopes tetraophthalmus Forst. 144 adults. Five tests. •

3. Chauliognathus marginatus Fabr. 42 adults. Two tests.

4. Photinus pennsylvanica DeG. 78 adults. Three tests.

5. Popillia japonicg Newm. 295 adults. Three tests.

6. Autoserica castanea Arrow. 338 adults. Three tests.

Tested after 10: 30 p.m.

(Jour. N. Y. Ent. Soc.), Vol. LII (Plate V)

42

Journal New York Entomological Society

[Vol. LII

PLATE VI

Figure 3. Behavior of coleopterous, hymenopterous and lepidopterous larvse
to 10 wave-length bands, in disarray, from 3650 A to 7200 A.
Physical intensities equalized.

1. Leptinotarsa decemlineata Say (Col.). 372 larvse, 1 to f grown.

Four tests.

2. Lophyrus lecontei Fitch (Hymen.). 124 larvse, full grown. Two

tests.

3. Macremphytus sp. (Hymen.). 491 larvse, full grown. Three tests.

Larvse starved.

4. Diacrisia virginica Fab. (Lep.). 92 larvse, full grown. Three tests.

5. Dryocampa rubicunda Fab. (Lep.). 100 larvse, full grown. Three

tests.

6. Anisota senatoria A & S (Lep.). 125 larvse, i to f grown. Three

tests.

Figure 4. Behavior of lepidopterous larvse to 10 wave-length bands, in dis-
array, from 3650 A to 7200 A. Physical intensities equalized.

1. Eudamus tityrus Fab. 125 larvse, full grown, starved. Three tests.

2. Hadena turbulenta Hbn. 263 larvse, f grown. Three tests.

3. Datana integerrima G. & E. 188 larvse, f grown, starved. Two tests.

4. Datana ministra Dru. 303 larvse, f to full grown. Three tests.

5. Melalopha inclusa Hbn. 70 larvse, full grown. Two tests.

6. Hyparpax aurora S. & A. 274 larvse, full grown. Three tests.
Figure 5. Behavior of lepidopterous larvse to 10 wave-length bands, in dis-
array, from 3650 A to 7200 A. Physical intensities equalized.

1. Actias luna Linn. 63 larvse, | grown. Two tests.

2. Telea polyphemus Cram. 141 larvae, f-full grown. Two tests.

3. Telea polyphemus Cram. 136 larvse, full grown, starved. Four tests.

4. Ceratomia catalpce Bdv. 110 larvse, full grown. Three tests.

5. Phlegethontius Carolina Linn. 32 larvse, full grown. Two tests.

(Plate VI)

(Jour. N. Y. Ent. Soc.), Vol. LII

7200 A

Mar., 1944]

Alexander: Crane-Flies

45

RECORDS AND DESCRIPTIONS OF NEOTROPICAL
CRANE-FLIES (TIPULID^, DIPTERA), XVII

By Charles P. Alexander
Amherst, Massachusetts

The previous instalment under this general title was published
in September, 1943 (Journal of the New York Entomological
Society, 51(3): 199-212). The materials here considered are
all from Ecuador where they were taken by Mr. William Clarke-
Macintyre, Mr. David B. Laddey, and Professor F. Martin Brown.
Some of the most interesting of the new species were taken at the
station “Zumbi,” in the Province of Santiago-Zamora, southern
Oriente, by Mr. Laddey. For a brief discussion of this station,
the preceding instalment should be consulted. I am greatly
indebted to the collectors for the privilege of retaining the types
of the novelties in my collection of these flies.

Genus Gnophomyia Osten Sacken

Gnophomyia (G-nophomyia) argutula new species.

General coloration of mesonotal praescutum gray with three dark brown
stripes, the lateral portions orange-yellow; thoracic pleura variegated dark
brown and obscure yellow; knobs of halteres dark brown; femora yellow, the
tips conspicuously blackened; wings yellowish gray, with three darker clouds
or bands, the second at the level of cord; cell 1st M2 widened outwardly,
nearly four times as wide at outer end as at base; cell 2nd A broad; abdom-
inal segments bicolored, the basal portions reddish yellow, the remainder
dark brown ; male hypopygium with the outer dististyle acute at tip ; phallo-
some unusually broad and obtuse.

Male. — Length about 7 mm.; wing 7.5 mm.

Rostrum brown; palpi black. Antennae brownish black; flagellar segments
nearly cylindrical; verticils of outer segments much longer and more con-
spicuous than those of the basal segments. Anterior vertex dull orange, rela-
tively narrow, the eyes correspondingly large and protuberant; posterior
portion of head brownish gray, the anterior orbits obscure orange.

Pronotum obscure brownish yellow above, dark brown on sides ; pretergites
obscure yellow. Mesonotal praescutum with the ground color of interspaces
gray, with three entire dark brown stripes, the median one more reddened at
cephalic end; humeral and lateral portions of sclerite obscure orange yellow;
posterior sclerites of notum dark brown, sparsely pruinose; scutellum paler,

46

Journal New York Entomological Society

[Vol. Lll

with a central, dark brown spot ; dorsal pleurotergite obscure yellow. Pleura
chiefly dark brown, sparsely pruinose, restrictedly but conspicuously varie-
gated with obscure yellow, distributed as follows: Dorsal portion of sterno-
pleurite ; posterior border of pteropleurite, and meral region. Halteres short,
stem yellow at base, the remainder dark brown. Legs with the coxae brown,
sparsely pruinose; trochanters obscure yellow; femora yellow, the tips rather
broadly and conspicuously blackened; tibiae and tarsi yellow, only the ter-
minal segment weakly darkened. Wings with the ground color yellowish
gray, the color greatly restricted by three more or less distinct, slightly
darker clouds or bands, the most conspicuous at the cord and over outer end
of cell 1st M2; slightly less distinct bands at proximal fourth of wing and
as a nearly apical darkening in the cells beyond cord; stigma very long and
narrow, dark brown; veins yellow in the ground areas, brown in the darkened
fields. Venation: Sc± ending about opposite the short transverse R2’, -Ss in
longitudinal alignment with E5, r-m at its fork; cell 1st M2 strongly widened
outwardly, nearly four times as wide at outer end as at base; cell 1st M2
approximately as long as vein M± beyond it; cell 2nd A noticeably shorter
and broader than in duplex.

Basal abdominal segments bicolored, dark brown, the basal rings conspicu-
ously reddish or reddish yellow, the subterminal segments more uniform dark
brown; hypopygium and preceding segment more yellowish; sternal pattern
generally like the tergal. Male hypopygium with the outer dististyle much
less conspicuously flattened than in laticincta, its apex acute; base of style
with only two elongate setae. Inner dististyle about one-half as long as the
outer style, provided with numerous setae, including about three of unusual
length. Phallosome unusually broad and obtuse.

Holotype, J1, Zumbi, Rio Zamora, Santiago-Zamora, altitude
700 meters, November 1, 1941 (Laddey).

The nearest relatives are Gnophomyia ( Gnophomyia ) duplex
Alexander and G. (G.) laticincta Alexander, which have the legs
and wings somewhat similarly patterned. The former species is
still known only from the female sex, differing from the present
fly in the venation and in the details of coloration of body and
wings. The latter species, laticincta , differs conspicuously in
the structure of the male hypopygium, especially of the outer
dististyle.

Gnophomyia (Gnophomyia) bulbibasis new species.

General coloration of mesonotum dark brown, very sparsely pruinose, on
praeseutum forming a discal shield; thoracic pleura striped longitudinally
with dark brown and reddish; halteres darkened; legs pale brown; wings
grayish subhyaline, stigma scarcely differentiated; male hypopygium with
the outer dististyle conspicuously bulbous just beyond base; gonapophyses.
appearing as blackened spines.

Mar., 1944]

Alexander: Crane-Flies

47

Male. — Length about 4.5-5.5 mm. ; wing 5-6.2 mm.

Female. — Length about 5 mm. ; wing 5.5 mm.

Rostrum and palpi dark brown. Antennae dark brown; basal flagellar seg-
ments subcylindrical, the outer ones shorter; verticils subequal in length to
the segments. Head dark gray; eyes large; anterior vertex only a little
wider than the diameter of scape.

Pronotum above yellow, darker on sides. Mesonotum almost uniformly
dark brown, very sparsely pruinose, on prsescutum forming a discal shield
that leaves the humeral and lateral portions yellowish, in cases more obscure
than in others. Pleura reddish, with a conspicuous dark brown dorsal stripe
extending from the propleura to the postnotum, passing above the halteres;
dorsopleural region yellow, confluent with the similarly colored lateral
prsescutal borders; immediately ventrad of the dark pleural stripe a more or
less distinct paler longitudinal line extending from behind the fore coxae
to the base of abdomen. Halteres dusky, the knob still darker. Legs with
coxae obscure yellow to testaceous yellow; trochanters yellow; remainder of
legs pale brown, the femoral bases clearer yellow; outer tarsal segments pass-
ing into darker brown. Wings grayish subhyaline, the extreme base yellow ;
stigmal area very restricted and pale, scarcely differentiated; veins pale
brown, yellow in the prearcular field. Venation: Sc long, S cx ending just
before level of B2, Sc2 some distance from its tip, lying opposite or before
the fork of Bs; r-m before or close to fork of Bs-} m-cu about three-fourths
its own length beyond the fork of M .

Abdominal tergites brownish black, the sternites a trifle more piceous;
hypopygium yellowish brown. Ovipositor with eerci relatively short and
stout, with setae to the tips of the valves. Male hypopygium with the outer
dististyle conspicuously expanded or bulbous just beyond base, thence nar-
rowed to a long straight rod that terminates in an acute point. Inner disti-
style much shorter, obtuse at tip; at base with about four powerful setae,
with other shorter setae on distal half, chiefly near apex. Gonapophyses
appearing as blackened spines.

Holotype, Zumbi, Rio Zamora, Santiago-Zamora, altitude
700 meters, November 4, 1941 (Laddey). Allotopotype, 5? with
the type. Paratopotypes, 2 1 $, October 31-November 2,

1941 (Laddey).

Gnophomyia ( Gnophomyia ) bulbibasis has the structure of
the male hypopygium, especially of the gonapophyses, somewhat
as in G. (G.) oxymera Alexander, from which it differs in other
hypopygial characters, as the bulbous basal enlargement of the
outer dististyle.

Gnophomyia (Gnophomyia) fessa new species.

General coloration of mesonotum and the dorsal pleurites dark brown,
ventral pleurites reddish brown ; rostrum, antennee and halteres black ; femora

48

Journal New York Entomological Society

[Vol. LII

obscure yellow, with a narrow subterminal darker ring; wings subhyaline,
stigma reduced to a narrow seam; male hypopygium with the inner dististyle
bulbous, the apex conspicuously wider than the base; gonapophyses incurved,
appearing as separate blades, blackened and microscopically serrulate at
bases.

Male. — Length about 5 mm.; wing 5 mm.; antenna about 1.5 mm.

Female. — Length about 6.5 mm. ; wing 6 mm.

Rostrum and palpi brownish black. Antennae black throughout; basal
flagellar segments subcylindrical, the outer ones more elongate-oval, with
verticils that exceed the segments in length. Head dark gray; eyes (male)
relatively large, the anterior vertex correspondingly narrowed.

Pronotum brown, the pretergites conspicuously light yellow. Mesonotum
almost uniform dark brown, the surface very sparsely pruinose, the central
portion of praescutum and the scutal lobes slightly darker. Pleurotergite and
dorsal pleura dark brown, contrasting abruptly with the reddish brown
ventral pleurites, the surface sparsely pruinose. Halteres brownish black
throughout. Legs with coxae reddish; trochanters yellow; femora obscure
yellow, more infuscated immediately before their tips to form a narrow,
nearly terminal ring; tibiae and basitarsi obscure yellow, the tips narrowly
infuscated; outer tarsal segments passing into brownish black. Wings sub-
hyaline, with a very faint darker tinge; stigma darker brown but reduced
to a linear area adjoining veins B^ and B1+2, scarcely involving the surround-
ing membrane; prearcular and costal fields a trifle more yellowish; veins
brown, somewhat more brightened in the basal areas. Venation : $<?! ending
opposite -K2; Bs straight, oblique; basal section of B5 lacking, r-m at fork
of Bs; B2 variable in position, before the fork of B2+ 3+4 or nearly its own
length beyond this fork; cell 1st M2 narrow, subequal in length to vein
beyond it; m-cu from one-third to two-thirds its length beyond the fork of M.

Abdominal tergites and hypopygium brownish black; sternites yellow.
Ovipositor with cerci relatively small, only weakly sclerotized, with setae
virtually to their tips. Male hypopygium with the outer dististyle a simple,
curved, relatively narrow, blackened rod, the tip subacute. Inner dististyle
relatively short, bulbous, the apex almost twice as wide as the base, with
several setae, chiefly on the distal third. Phallosome with the gonapophyses
incurved, appearing as separate blades, at apex blackened and microscopically
roughened to serrulate; apex of phallosome narrow.

Holotype, Palmar, Rio Maizito, Manabi, altitude 200 meters,
May 20, 1941 (Laddey). Allotopotype, §, pinned with type.

Gnophomyia ( Gnophomyia ) fessa is most similar to species
such as G. (G.) oxymera Alexander and G. ( G .) nectarea new
species, differing in the details of coloration and, especially, the
structure of the male hypopygium, as the inner dististyle and
the phallosome.

Mar., 1944]

Alexander: Crane-Flies

49

G-nophomyia (G-nophomyia) nectarea new species.

Size small (wing, male, about 5 mm.) ; mesonotum and dorsal pleura
opaque black, the ventral pleurites abruptly reddish; halteres dusky; legs
brownish yellow; wings subhyaline, the stigma and an extensive cloud on
proximal third of wing infuscated; vein R2 very faint to nearly atrophied;
cell 1st M2 long and narrow ; cell 2nd A relatively narrow ; male hypopygium
with the outer dististyle. slender; inner dististyle with a conspicuous basal
lobe that is microscopically corrugated or wrinkled.

Male. — Length about 4-4.8 mm.; wing 4.5-5.5 mm.; antenna about 1.3-
1.6 mm.

Female. — Length about 5 mm. ; wing 5 mm.

Eostrum dark brown; palpi black. Antennae of moderate length, dark
brown; flagellar segments subcylindrical ; longest verticils subequal in length
to the segments. Head brown, the orbits and posterior vertex light gray;
anterior vertex relatively narrow, about one-third the diameter of scape; eyes
(male) correspondingly large.

Pronotum dark brown ; pretergites light yellow. Mesonotum chiefly
blackened, the surface opaque by a sparse pruinosity; central portion of
scutum and posterior border of scutellum slightly more reddened. Dorsal
pleurites covered by a broad black longitudinal stripe, this area also involv-
ing the pleurotergite and surrounding the root of halteres; ventral pleurites
reddish, very sparsely pruinose. Halteres dusky, base of stem restrictedly
brightened. Legs with coxae and trochanters yellow; remainder of legs
brownish yellow, the outer tarsal segments more infuscated. Wings sub-
hyaline, the extreme base restrictedly yellow; stigma oval, darker brown;
a conspicuous dusky cloud or wash on basal third of wing in general vicinity
of vein Cu, involving the bases of cells Cu, 1st A and 2nd A, together with
much of M ; veins brown, yellow in the flavous basal region. Venation:
Vein R2 very faint to nearly atrophied; Rs in direct longitudinal alignment
with _K5; branches of Rs all extending generally parallel to one another;
cell 1st M2 relatively long and narrow, its inner end pointed or strongly nar-
rowed, the outer end more widened, the cell subequal in length to vein M4;
m-cu less than its own length beyond fork of M, at near one-fourth to one-
fifth the length of the cell; cell 2nd A relatively narrow.

Abdominal tergites and the hypopygium brownish black; basal sternites
a trifle paler. Male hypopygium with the outer dististyle unusually slender
throughout, narrowed at apex to a subacute point. Inner style with a con-
spicuous basal lobe or shoulder, its surface and adjoining margin of style
microscopically wrinkled. Phallosome of moderate width, its tip blackened
and slightly narrowed to an obtuse point, the surface microscopically wrinkled.

Holotype, J1, Zumbi, Rio Zamora, Santiago-Zamora, altitude
700 meters, November 2, 1941 (Laddey). Allotopotype, §•
Paratopotypes, 2 jy1, October 30-November 5, 1941 (Laddey).

50

Journal New York Entomological Society

[Vol. LII

Gnophomyia ( Gnophomyia ) nectar ea is entirely distinct from
other small-sized regional members of the genus. From all such,
it differs conspicuously in the peculiar wing pattern and in the
structure of the male hypopygium.

Gnophomyia (Gnophomyia) tuber new species.

General coloration of mesonotum medium brown, with poorly defined pat-
tern; pleura with a broad, medium brown, longitudinal stripe over the dorsal
sclerites; halteres yellow; wings with a buffy tinge, brighter in the basal
portions; abdominal tergites reddish brown, weakly darkened laterally;
male hypopygium with the basistyles short, on mesal face near base with a
conspicuous darkened tubercle, this provided with about three conspicuous
setae; outer dististyle compressed-flattened, its apex obtuse; inner dististyle
with the apex very obtuse to subtruncate.

Male. — Length about 6-6.3 mm.; wing 6.8-7 mm.; antenna about 1.5-
1.6 mm.

Rostrum testaceous brown; palpi brownish black. Antennae with the scape
and pedicel brownish yellow, flagellum brownish black; flagellar segments
long-cylindrical, the terminal segments shorter; verticils considerably exceed-
ing the segments in length. Head brownish gray; anterior vertex relatively
narrow, eyes large.

Pronotum light yellow above, darker on sides; pretergites light yellow.
Mesonotal praescutum medium brown, sparsely pruinose, the median area
more darkened, behind forming more or less distinct stripes; scutal lobes
similarly darkened, the posterior border obscure yellow; scutellum brown,
with a yellow spot on either side of base; postnotum medium brown, more
yellowish on suture between mediotergite and pleurotergite. Pleura with a
broad but relatively inconspicuous, medium brown stripe, more intense in
front, becoming diffuse behind; dorsopleural region yellow; ventral pleurites
obscure yellow, paler yellow behind. Halteres yellow. Legs with the coxae
and trochanters yellow; femora obscure yellow, weakly darkened at or near
tips; tibiae and basitarsi yellow, the tips narrowly and inconspicuously infus-
cated; outer tarsal segments brownish black. Wings relatively broad, with
a sandy or buffy tinge, the prearcular and costal fields clearer yellow ; stigma
very restricted in area, pale brown; veins brown, more brownish yellow in
the brightened portions. Venation: Scx ending about opposite the fork of
E2+3+i, £u2 nearly opposite the fork of Es ; E2+3 more than one-third E2+3+i;
Es in direct longitudinal alignment with E5 ; r-m beyond fork of Es ; cell
1st M2 relatively long, subequal to vein Jf4 beyond it ; m-cu about two-thirds
its length beyond fork of M.

Abdominal tergites reddish brown, weakly darkened laterally; sternites
clearer yellow; hypopygium yellowish brown. Male hypopygium with the
basistyles short, on mesal face near base with a conspicuous darkened tuber-
cle, this provided with about three conspicuous setae. Outer dististyle dark-
ened, conspicuously compressed-flattened, widest at near midlength, its apex

Mar., 1944]

Alexander: Crane-Flies

51

obtuse. Inner dististyle extending about to midlength of the outer style,
dark-colored, its apex very obtuse to subtruncate; rather numerous setae,
including a row of four or five larger ones along the lower or cephalic margin.
Phallosome relatively broad, the apex obtuse.

Holotype, J', Playas de Montalvo, Los Rios, altitude 15 meters,
March 5, 1938 (Macintyre). Paratopotypes, 1 <$, 1 sex?, pinned
with type.

In its general appearance and wing coloration, the present fly
is most like Gnophomyia ( Gnophomyia ) acricula Alexander and
G. ( G .) digitiformis Alexander, but has a very different hypo-
pyginm. The tubercle on the basistyle is not found in any other
species known to me.

Genus N eo gnophomyia Alexander

Neognophomyia spectralis new species.

General coloration of mesonotal praescutum reddish yellow, the posterior
sclerites darker; vertex yellow, with a brown central area; thoracic pleura
with a conspicuous dorsal black stripe; tips of femora, tibiae and basitarsi
broadly blackened; wings with a pale yellow tinge, patterned with brown,
including a subbasal fascia; abdominal tergites conspicuously patterned
with brownish black and yellow; male hypopygium with the tergal spines
blackened, nearly straight; phallosome produced at apex into lateral horns.

Male. — Length about 5-5.5 mm.; wing 6-7 mm.

Rostrum obscure yellow; palpi brown. Antennae with scape and pedicel
dark brown; basal flagellar segments pale brown, the outer ones paling to
brownish yellow; flagellar segments passing from oval through long-oval;
verticils conspicuous. Head yellow, the anterior vertex with a conspicuous
brown area; anterior vertex moderately wide, a little more than twice the
diameter of the scape.

Pronotum yellow medially, brownish black on sides. Mesonotal praescutum
reddish yellow, more yellowish on lateral and humeral portions, in cases more
darkened, especially near suture; scutum yellow, each lobe with two brown
areas; scutellum black, more or less pruinose; postnotum brownish black,
including both the mediotergite and pleurotergite. Pleura reddish yellow
with a broad black longitudinal stripe beginning on sides of pronotum, in-
volving the anepisl;ernum, dorsal pteropleurite and the postnotum, as de-
scribed. Halteres yellow. Legs with the coxae yellow to reddish yellow;
trochanters yellow; femora, tibiae and basitarsi yellow, with broad and
conspicuous black tips; remainder of tarsi black. Wings with a pale yellow
tinge, patterned with brown, including a broad seam from stigma across
anterior cord, and narrower seams on m-cu and outer end of cell 1st M2; a
less conspicuous subbasal band from origin of Rs extending obliquely across
cell M into cells Cu, 1st A and 2nd A ; extreme base of wing blackened; veins

52

Journal New York Entomological Society

[Vol. LII

yellow to brownish yellow, darker brown in the patterned areas. Venation:
B2 at near midlength of petiole of cell _E3; cell 1st M2 strongly narrowed at
proximal end; m-cu from about one-half to approximately its own length
beyond the fork of M.

Abdominal tergites yellow, handsomely patterned with brownish black,
the segments chiefly darkened with a large yellow area at posterior border
of each segment, encroaching on the base of the succeeding segment, the
outer segments more uniformly darkened, greatly restricting the yellow color ;
sternites yellow, the terminal segments more darkened; hypopygium chiefly
obscure brownish yellow. Male hypopygium with the tergal spines appearing
as long, nearly straight, blackened blades, a little expanded at base, thence
gradually narrowed to the acute tips. Outer dististyle relatively slender;
outer margin of distal third with five long setae, additional to the single
terminal bristle. Inner dististyle large and massive, terminating in a broad
blackened beak, the apical border with about seven or eight strong setae;
basal tooth or flange of style strongly blackened, provided with four strong
setae. Phallosome relatively wide, at apex produced into a slender spinous
point that is directed laterad, immediately cephalad of which is a broadly
rounded emargination.

Holotype, J1, Banos, Tungurahua, altitude 2,000 meters, July
14, 1939 (Macintyre). Paratopotypes, 7 altitude 1,700-
2,000 meters, May 11-June 2, 1937 ; paratype, 2 J'J1, Pititi, near
Banos, altitude 1,900 meters, June 14, 1937 (Macintyre).

The only generally similar species is Neognophomyict hirsuta
(Alexander) of eastern Brazil, which has the pattern of the
body, legs and wings somewhat the same but with the male
hypopygium entirely different. This latter species has the sub-
basal darkened wing band much wider and more continuous than
in the present fly.

Neognophomyia interrupta new species.

Allied to hostica; general coloration yellow, the praescutum with the disk
chiefly reddish yellow ; wings grayish yellow, with a narrow brown seam along
cord; abdomen pale, with tergites three, five and six conspicuously dark
brown; male hypopygium with the outer dististyle strongly constricted at
near midlength, the bulbous outer portion terminating in two subequal elon-
gate setae; tergal spines pale, angularly bent beyond midlength.

Male. — Length about 4.5 mm.; wing 5 mm.

Eostrum pale yellow; palpi pale, the outer segments darkened. Antennae
with scape and pedicel light yellow; flagellum broken. Head obscure brown-
ish yellow; eyes (male) relatively large.

Pronotum yellow. Mesonotal praescutum yellow, with three more or less
confluent more reddish stripes that form a nearly continuous discal area;

Mar., 1944]

Alexander: Crane-Flies

53

scutal lobes brownish black, the remainder of scutum more brownish yellow ;
scutellum and mediotergite brownish yellow, the pleurotergite almost covered
by an oval velvety black spot, as is common in the genus. Pleura reddish yel-
low the anepisternum a little infuscated. Halteres uniformly pale yellow.
Legs with the coxae and trochanters yellow ; remainder of legs broken. Wings
grayish yellow, the prearcular and costal fields somewhat brighter yellow;
a narrow but conspicuous brown seam along cord, becoming indistinct or
obliterated at fork of M ; veins brownish yellow, darker in the infuscated
areas. Venation: E2 at about one-third the length of petiole of cell R3;
vein ^3‘ oblique; cell 1st M2 only slightly widened outwardly; m-cu at near
one-third the length of the cell.

Abdomen conspicuously patterned; basal tergites brownish yellow, dark-
ened laterally; tergites three, five and six conspicuously dark brown; tergites
four, seven and eight pale; hypopygium and sternites more uniformly yellow.
Male hypopygium with the outer dististyle strongly constricted at near mid-
length, the base widened, the apex bulbous, its extreme tip broadly obtuse
with two subequal elongate setae. In hostica, the style is unusually slender,
gradually narrowed beyond the basal enlargement, the tip unusually slender
and thus with only a single apical seta. Inner dististyle much as in hostica,
more widened at near midlength, the setae at this point slightly more
numerous, longer and paler. Phallosome narrower. Tergal spines somewhat
as in hostica, the basal section shorter.

Holotype, J', Zumbi, Rio Zamora, Santiago-Zamora, altitude
700 meters, November 5, 1941 (Laddey).

The most similar described species is Neognophomyia hostica
Alexander, of Peru, which differs most evidently in the colora-
tion of the abdomen and in the structure of the male hypopygium,
as compared above.

Genus Gonomyia Meigen

Cronomyia (Progonomyia) acrissima new species.

General coloration gray, the praescutum with three conspicuous brown
stripes; thoracic pleura dark gray with a broad yellow longitudinal stripe;
legs dark brown, the tarsi black; wings with a weak grayish tinge, unpat-
terned except for the very pale brown and inconspicuous stigma; male
hypopygium with the mesal face of basistyle before apex with a longitudinal
group of acute spines, these exceeding 60 in number.

Male. — Length 5.5-5. 6 mm.; wing 6-6.2 mm.; antenna about 1-1.1 mm.

Rostrum black, sparsely pruinose; palpi brownish black. Antennae with
the scape dark brown, pedicel and flagellum black; flagellar segments sub-
cylindrical to long-oval with verticels that are subequal in length to the
segments. Head gray; vertex with a median brown line.

Pronotum blackened, pruinose, the lateral portions, together with the
anterior pretergites, obscure yellow. Mesonotal praescutum gray, with three

54

Journal New York Entomological Society

[Vol. LII

conspicuous brown stripes, the median one not reaching the suture, the lat-
erals crossing the suture onto the scutal lobes; pseudosutural foveae black,
conspicuous; posterior sclerites of notum gray, the posterior border of scu-
tellum more obscure yellow; postnotum gray, the dorsal portion of pleuro-
tergite and adjoining portion of mediotergite obscure yellow. Pleura dark
gray, with a broad and conspicuous yellow stripe extending from behind the
fore coxae across the dorsal sternopleurite, behind including the ventral
pteropleurite, meron and metapleura. Halteres with stem yellowish brown,
knob darker brown. Legs with coxae light brown, sparsely pruinose; tro-
chanters brownish yellow; remainder of legs dark brown, the tarsi passing
into black. Wings with a weak grayish tinge, unpatterned except for the
very pale brown stigma; extreme wing base paler; veins brown, brightened
at base of wing. Venation: Sc relatively short, Sol ending a distance be-
yond origin of Bs about equal to m-cu, Sc2 a slightly shorter distance before
origin; Bo indicated by a very faint element at midlength of stigma; cell
2nd M2 nearly twice its petiole; m-cu from three-fifths to two-thirds its
length before the fork of M.

Abdominal tergites dark grayish brown, sternites slightly paler; eighth
and ninth segments somewhat paler brown, the styli darker. Male hypo-
pygium with the apex of basistyle obtuse, unarmed; mesal face immediately
back from tip with an extensive longitudinal area of acute spines, these
totalling in excess of 60. Outer dististyle a semicylindrical blade, its apex
narrowed into an acute black spine. Intermediate style a long darkened
blade, about one-half longer than the outer style; at near one-third its length
bearing a more slender branch or arm that is approximately one-half as long
as the main apical branch; stem near the branch with numerous setae, these
becoming even longer and more conspicuous on base of lateral branch; main
or axial branch glabrous, gradually narrowed to the subacute cultriform
apex. Inner dististyle broad, obtuse at apex, near margin provided with
several setae, those at and near apex longest. JEdeagus relatively slender,
narrowed outwardly, near apex with small and inconspicuous lateral shoulders.

Holotype, Banos, Tungurahua, altitude 1,450 meters, April
23, 1939 (Macintyre). Paratopotype, 1 J*.

The present fly is readily distinguished from all other gen-
erally similar forms by the structure of the male hypopygium,
especially the spinous area near apex of basistyle. Such arma-
ture is uncommon in the subgenus and in all other cases the num-
ber, arrangement and shape of the spines is different; in
Gonomyia ( Progonomyia ) compact a Alexander there is a rela-
tively large group of spiculose points; in G. (P.) serena Alex-
ander, the spines of the general type of the present fly but very
few in number; in G. (P.) thiosema Alexander, an even larger
group of elongate spinous pegs on mesal face of basistyle but

Mar., 1944]

Alexander: Crane-Flies

55

these with obtusely rounded tips. In all these species, the
structure of the dististyles is likewise distinctive.

G-onomyia (Progonomyia) hyperplatys new species.

General coloration of notum grayish brown; antennae black throughout;
pleura blackened, sparsely pruinose, with a conspicuous yellow longitudinal
stripe; knobs of halteres darkened; wings yellowish, restrictedly patterned
with pale brown; S c± ending shortly beyond origin of Es-, male hypopygium
with the phallosome unusually wide, appearing as a broadly flattened plate,
each outer lateral angle further produced.

Male. — Length about 6 mm.; wing 6.6 mm.

Eostrum and palpi dark brown. Antennae brownish black throughout;
flagellar segments subcylindrical, passing into long-oval, with verticils that
exceed the segments. Head brownish gray on anterior vertex, deepening in
color behind the antennal bases, the broad posterior vertex clear gray.

Pronotum obscure yellow medially, dark brown sublaterally ; pretergites
yellow. Mesonotum dark grayish brown, the humeral and lateral portions
light yellow; scutellum more reddish brown; mediotergite dark brown, with
a conspicuous yellow area on either side of basal half; dorsal portion of
pleurotergite similarly yellow. Pleura blackened, sparsely pruinose, with
a conspicuous yellow longitudinal stripe extending from above the fore coxae
across the dorsal sternopleurite to the ventral pteropleurite and meral region.
Halteres with stem yellow, knob weakly darkened. Legs with the coxae
basally dark brownish gray, the tips restrictedly yellow; trochanters brown;
remainder of legs broken. Wings with a yellowish tinge, restrictedly pat-
terned with rather pale brown, the areas chiefly restricted to the vicinity
of the veins, distributed as follows: Origin of Es ; stigma; cord and base
of cell 2nd M2 ; two small isolated spots in Anal cells, one near outer end
of cell 1st A adjoining vein 2nd A, the other near center of cell 2nd A ;
veins brown. Venation: Sc short, S cx ending shortly beyond origin of Es,
Sc2 about an equal distance before this origin, Es relatively long, square and
spurred at origin; E2 about twice E3+i; m-cu from about one-third to one-
half its length before fork of M.

Abdomen brownish black; ninth segment abruptly yellow, the styli again
blackened. Male hypopygium with the outer dististyle a short, slender,
curved rod that narrows to the acute blackened tip, the outer margin with
delicate microscopic setulae. Intermediate style broadly flattened, entirely
dark-colored, the inner margin at near midlength produced into a sharp spine,
the surface basad of this with conspicuous setae; apical portion of style ap-
pearing as a broad flattened basal flange that narrows into a long arm that
is narrowly but deeply notched at apex. Inner dististyle a dusky lobe with
strong spinous setae along the entire inner face, the terminal one longest.
Phallosome unusually wide for a member of this subgenus, appearing as a
broadly flattened plate subtending the aedeagus, each outer lateral angle pro-
duced further into a flattened lobe or blade, with about seven setae chiefly
distributed along the outer margin near apex.

56

Journal New York Entomological Society

[Vol. LII

Holotype, Banos, Tungurahua, altitude 1,450 meters, April
23, 1939 (Macintyre).

The present fly is entirely different from other described species
of the subgenus, differing especially in the structure of the male
hypopygium and particularly of the phallosome. The most
similar form is Gonomyia ( Progonomyia ) velutina Alexander,
which differs in the wing pattern and in all details of structure
of the male hypopygium.

Genus Cryptolabis Osten Sacken

Cryptolabis (Cryptolabis) alticola new species.

General coloration black, more or less prninose ; lateral pretergites abruptly
yellowish white; legs black; wings with a strong brownish tinge, the pre-
arcular and costal fields yellow; Bs very long; male hypopygium with the
dististyle large and conspicuous, exserted, consisting of an outer flattened
setuliferous blade and a bispinous inner body; tergal plate bearing conspicu-
ous lateral arms that terminate in several strong setae; aedeagus long and
slender.

Male. — Length about 4.5 mm.; wing 5.2 mm.

Rostrum and palpi black. Antennae brownish black; flagellar segments
oval, with conspicuous verticils. Head black, presumably pruinose in fresh
specimens.

Pronotum, mesonotum and pleura of the unique type dull black, the surface
presumably more or less pruinose in fresh specimens; pretergites abruptly
and conspicuously yellowish white. Halteres brown, the apex of knob
brighter. Legs black, with relatively conspicuous suberect setae. Wings
with a strong brown tinge, the prearcular and costal fields yellow; axillary
region restrictedly infuscated; veins and macrotrichia brown. Macrotrichia
of cells relatively abundant, beyond the cord extending from outer end of
cell B2 to cell Mi} in the radial field involving all of the cells with the excep-
tion of the basal fourth to fifth. Venation: Sc relatively short, Sc1 ending
some distance before the end of Bs; Bs very long, exceeding in length vein
Bs; B2+ 3+4 at origin nearly perpendicular to Bs , subequal in length to B2+ 3;
cell M3 deep; m-cu at near midlength of Ms+i; vein 2nd A gently sinuous.

Abdomen brownish black; hypopygium black. Male hypopygium with the
dististyle large and conspicuous, exserted; consisting of an outer fleshy lobe
or blade that may represent a separate style, and the main body of the style
itself; outer blade flattened, densely covered with short setae; inner portion
of style with apex dilated into a truncated portion, the outer margin with
two strong blackened spinous points, the more basal one erect, the outer spine
more appressed. What appears to be the tergite appears as a transverse
plate with the median portion strongly produced into a rounded lobe, the
lateral arms appearing as strong cylindrical lobes that are directed strongly

Mar., 1944]

Alexander: Crane-Flies

57

mesad and then caudad, the tips with about seven elongate setae. .ZEdeagus
unusually long and slender, blackened, transversely corrugated.

Holotype, J1, Hacienda Talahua, Bolivar, altitude 3,100 meters,
April 28, 1939 (Brown).

Cryptolabis ( Cryptolabis ) alticola is entirely different from the
other described species of the genus. It is most similar to species
such as C. (C.) chilotanica Alexander, of southern Chile, yet very
distinct in the structure of the male hypopygium. For a discus-
sion of the type locality, consult Brown (Ann. Ent. Soc. Amer.,
34: 848; 1941).

Mar., 1944]

Clench: Lyc^enhle

59

TWO NEW SUBSPECIES OF EVERES
COMYNTAS GODART (LEPIDOPTERA,
EYC/ENIDJE)

By Harry K. Clench
Cambridge, Mass.

Two races of this wide-ranging species have recently come to
my attention, one from South Dakota and one from Montana.

Everes comyntas valerise, new subspecies

TJpperside :

Male. Both wings slightly purplish blue. Fore wing with a narrow dark
border on the outer margin. Costa and costal veins pencilled with light blue.
Hind wing with an internervural row of small spots on the outer margin.
Costa dark bordered. Cu2 with a short tail. Fringe of fore wing dark
basally, white outwardly; of the hind wing white.

Female. Both wings brown. Base of each dark blue. Hind wing with a
black spot in the CurCu2 interspace basally bordered by a shallow, rather
dull orange lunule. A thin marginal pale bluish line borders the outer mar-
gin, interrupted at the veins, basal to which is a similar but scalloped and
more obscure line. The two outline a series of internervural dark spots.
Occasionally these lines are very faint, and the row of spots consequently
almost indistinguishable.

Underside :

Male. Ground color grayish tan. All spots arranged as in typical comyn-
tas, but with the post-discal series usually rounder and darker — occasionally
very heavy. The orange over the two spots in the M3-Cux-Cu2 interspaces of
the hind wing is usually faint, although stronger in the latter than in the
former. On this same wing, just basal to the marginal compound border,
the ground color is white between the veins.

Female. Similar to the male.

Length of fore wing: Male, 11.5-13.5 mm.; Female, 10-12.5 mm.

Holotype, male, near Lead, South Dakota, June 22, 1939 v(V.
H. and A. C. Frederick).

Allotype, female, Terry Peak, South Dakota, el. 5200+ feet,
June 24, 1939 (V. H. and A. C. Frederick).

Paratypes, 1 male, same data as holotype ; 1 male and 2 females,
same data as allotype; 1 female, Harney Peak, el. 5200+ feet,
June 25, 1939 (A. C. Frederick) ; 31 males, 3 females, Spearfish
Canyon and vicinity, el. 5200+ feet, as follows : 3 males, June 26,

60

Journal New York Entomological Society

[Vol. Li l

27, and July 1, 1939, resp. (V. H. and A. C. Frederick) ; 15 males,
2 females, June 29, 1942 (A. C. Frederick) ; 13 males and 1
female, June 30, 1942 (A. C. Frederick) ; 17 males, Icebox Can-
yon, June 29, 1942 (A. C. Frederick). All localities in the Black
Hills, South Dakota.

Holotype and allotype no. 25934 in the collection of the Museum
of Comparative Zoology. Paratypes in the collection of Mr.
Frederick and of the author.

Remarks. This subspecies appears to be most similar to the
southwestern race herrii Grinnell1 from which it differs in the
following particulars : the black border on the wings above in the
male is slightly thinner. This is most apparent on the hind wing,
where the border in herrii is thick enough to include the inter-
nervural spots, while in valerice these spots are almost always
free. There is no orange lunule on the hind wing above, an
almost constant feature of herrii , and even more prominent in
typical comyntas, but more variable in the latter. The ground
color below in both sexes appears to be a little darker. The
female of valerice is largely blackish brown above, with a rather
dark basal blue shading, while in herrii both wings are rather
extensively blue above. The orange lunules above are smaller in
females of valerice than in those of herrii.

This subspecies appears to be quite variable. One male has a
tiny orange lunule in the Cui-Cu2 interspace above — the only
indication of it in the whole type series. The same specimen and
one or two others have the marginal border on both wings thick-
ened, that on the hind wing including the submarginal spots.
Below, the intensity of the spots varies considerably. One speci-
men has the submarginal series of lunules in the compound border
enlarged and very dark, giving the insect a most peculiar appear-
ance. The post-discal series of spots may be enlarged or reduced.

This subspecies is named for Mrs. A. C. Frederick, who mate-
rially assisted her husband in collecting the type series.

Everes comyntas albrighti, new subspecies

TJpperside :

Male. Uniform violet-blue on both wings. Fore wing with a very narrow
dark marginal border. Hind wing also with this border, and in addition,.

i Can. Ent., 33 : 192, 1901.

Mar., 1944]

Clench : Lyc^enidhs

61

an almost obsolete row of small dashes, the heaviest in the CivCuo inter-
space.

Female. Uniform brown on both wings. Base of fore wing blue. On
the hind wing, in the Cux-Cm, interspace, a faint orange lunule surmounts a
tiny dark spot.

Underside:

Male. Fore wing dirty white, with a broad costal border of dark grayish
scaling. The marginal compound border is almost obsolete, save for a few
dark scales. A post-discal row of black spots, large and distinct, runs from
costa to inner margin, the costal ones usually obsolete. Cell closed by a dark
dash. Hind wing with a brownish gray ground color, fading on the outer
margin. The submarginal border is, as in the fore wing, almost completely
absent. The post-discal series, the cell-end bar, and the basal spots are also
nearly gone, but their positions are shown by whitened areas in the gray.
In the Cuj-Cuo interspace is a small black spot, obscurely scaled with metallic,
and capped by a tiny orange crescent.

Female. The single female examined shows a very similar appearance.
The post-discal series of spots on the fore wing is represented, however, only
by spots from M;> to the inner margin. The costal gray-brown on this wing
extends down to M3.

Length of fore wing: Male, 11-11.5 nun.; female, 11 mm.

Holotype, male, Kings Hill, Montana, July 9, 1939 (from C. C.
Albright).

Allotype, female, same data.

Paratypes, two males, same data.

Holotype and allotype to be deposited in the collection of the
Museum of Comparative Zoology. One paratype in the collec-
tion of Mr. C. F. dos Passos. One paratype in the collection of
the author.

Remarks. This subspecies differs from all other North Ameri-
can Everes thus far known in the grayish costal shading on the
fore wing below, and in the gray ground color of the hind wing
below. This subspecies appears to be closer in appearance to
northern specimens of amyntula Boisd.2 since several of its char-
acters correspond quite closely to that species, such as the narrow
margin above, reduced maculation below (aside from the gray
ground color) and reduced orange in both sexes.

This subspecies is named for Dr. C. C. Albright, of Great
Falls, Montana, from whom the specimens were obtained.

2 Ann. Soc. Ent. France, 10 (2) : 294, 1852.

(Jour. N. Y.

Ent. Soc.), Vol. LIT

(Plate VII)

FRANK EUGENE LUTZ

Mar., 1944]

Weiss: Frank E. Lutz

63

FRANK EUGENE LUTZ
1879-1943

Frank E. Lutz, the son of Martin P. Lutz and Anna Amelia
(Brockway) Lutz, was born in Bloomsburg, Pa., on September 15,
1879. His early education took place in the public schools and
the Bloomsburg State Normal School. From Haverford College
in 1900 he received his A.B. degree. During his first two years
in college he specialized in mathematics, upon the advice of his
father, who, being an insurance agent, was impressed by the
large earnings of life insurance actuaries. However, after two
years the boy decided to go into medicine and so mathematics
was dropped for biology. With college over his biology teacher,
H. S. Pratt, advised him to go into biometry in view of his train-
ing in both mathematics and biology. This he did by going to see
Dr. C. B. Davenport of the University of Chicago, who had charge
of a summer biological laboratory at Cold Spring Harbor. At
this laboratory Frank E. Lutz waited on tables and counted the
grooves on scallop shells, finally publishing his first paper, a very
short one, entitled “A Study in the Variations in the Number of
Grooves upon the Shells of Pecten irradiens (Lam.) ” in Science
in 1900. Although brief, this paper helped him to get a scholar-
ship at the University of Chicago where he obtained his A.M. in
1902. While working, as biologist for the North Shore Improve-
ment Association, in mosquito control on the north shore of Long
Island, he earned enough money to go to London (Eng.) where he
studied under Karl Pearson. From September 1902 to June 1903
he was a student in London and Berlin.

From 1904 to 1909 he was employed as resident investigator at
the Station for Experimental Evolution (Carnegie Institution) at
Cold Spring Harbor, N. Y., where he did research work on hered-
ity. Within this period, or in 1907, he obtained his Ph.D. from
the University of Chicago for his dissertation on “The Variation
and Correlation of Certain Taxonomic Characters of Gryllus,”
and at the beginning of this period, or on December 30, 1904, he
married Martha Ellen Brobson, of Philadelphia, Pa.

64

Journal New York Entomological Society

[Vol. Lll

Dr. Lutz then entered the employ of The American Museum
of Natural History in 1909 as assistant curator in the department
of invertebrate zoology. From 1917 to 1921 he served as asso-
ciate curator and in 1921, when the department of entomology was
created, he was appointed curator. For 22 years, or until his
death at the age of 64 on the morning of November 27, 1943, at
Harkness Pavilion, New York City, after an illness of several
weeks, Dr. Lutz continued as chairman and curator of the depart-
ment of insects and spiders.

After coming to the American Museum of Natural History his
activity in biometrics declined and was replaced by an absorbing
interest in insects, although he never had any college training in
entomology and although several of his early museum papers
dealt with the history of Antarctic explorations and with the
string-figures of Patamana Indians. Under Dr. Lutz’s leader-
ship a large exhibition and study collection of insects was assem-
bled, now numbering approximately 2,000,000 specimens. Many
of these collections were made by Dr. Lutz during the course
of 23 field expeditions to various parts of the United States and
to South and Central America and the West Indies. These
expeditions started in 1908 with a trip to Cuba and Mexico, and
after he entered the employ of the Museum, 23 expeditions were
made. The first took place in 1911 to the West Indies, British
Guiana and Florida, and the last in 1941 to California. Between
these dates, Dr. Lutz collected and made observations in Florida,
Louisiana, Texas, California, Colorado, Wyoming, Utah, Idaho,
Panama, Porto Rico, Cuba and British Guiana. In all five trips
were made to the West Indies, five to Panama, five to Florida and
eight to the western part of the United States. Some of these
western trips were made in a special Museum truck-like auto-
mobile which Dr. Lutz had outfitted with equipment for collecting
and living out-of-doors, making him independent of hotels and
trains. In addition, he took an active part in Museum affairs and
commttees and was chairman of publications and editor of the
Museum’s Bulletin and Memoirs from 1917 to 1929. From 1925
to 1928 he directed the Station for the Study of Insects at Tuxedo,
N. Y., and was one of the nation’s leading exponents for nature
trails and museums.

Mar., 1944]

Weiss: Frank E. Lutz

65

During the summer of 1926 he began the first trailside museum
of its kind at Bear Mountain, N. Y., and was called upon by many
organizations for advice in establishing similar trails in parks and
wild-life areas in various parts of the United States. I distinctly
recall his enthusiasm upon this subject, when in company with
Mr. E. L. Dickerson I paid him a visit at Tuxedo when the sta-
tion there was half completed. Dr. Lutz was so anixous to get
things finished that he would not go with us for lunch, preferring
to dine quickly upon some pieces of bread over which he had
broken a raw egg, a nutritious, if not appetizing mixture.

In planning the arrangement of the insect exhibits at the
Museum, Dr. Lutz attempted not only to supply information
about insects, but to interest the visitor in entomology as well.
About 1915 the plan of the Hall of Insect Life involved exhibits,
with continuity, covering ontogeny, anatomy, physiology, tax-
onomy, phylogeny, life-histories, insect associations, insect
enemies and evolution. And in addition, there were exhibits on
miscellaneous topics such as insect architecture, insects as food,
medicine, social insects, etc., etc. Later as exhibit methods
changed there were originated by Dr. Lutz various habitat groups.
Dr. Lutz tells of the early days of his department in ‘‘Natural
History,’ ’ May-June, 1924, under the title “Amateur Entomolo-
gists and the Museum.” During his administration the insect
collection was increased by gifts, purchases and expeditions, well
over a million specimens.

For many years Dr. Lutz took an active part in the affairs of
the New York Entomological Society, serving as president in 1925
and 1926 and on the Publication Committee for 20 years. For a
long period the meetings were held in Dr. Lutz ’s room on the third
floor of the Museum and there, surrounded by preserved spider-
webs, Dr. Lutz’s zoo of living insects and entomological books and
paraphernalia, many interesting entomological discussions took
place, in which he always participated.

A glance at Dr. Lutz’s published writings indicates that from
1910 on, they were concerned exclusively with insects, and spiders,
principally the former. In both his popular and scientific writ-
ings he covered such topics as geographic distribution, insect
sounds, a study of ultraviolet in relation to flower- visiting habits

66

Journal New York Entomological Society

[Vol. LII

of insects, wind and the direction of insect flight, insect life in
thermal waters, and other subjects involving the biology and
behavior of insects. He was not interested in economic entomol-
ogy and he believed that more intensive work on the biology of
insects was needed in view of the fact that comparatively little
is known even about many of our most common species. Im-
portant contributions were made by Lutz in the field of insect
behavior through his research work on insect reactions to ultra-
violet, on the training of bees to come to certain ultraviolet wave-
lengths and patterns for food, on his recordings and sound motion
pictures of insect sounds and insect behavior under various atmos-
pheric pressures. Such work required not only a thoughtful and
enquiring mind, an awareness of the pitfalls in conclusions, but
ingenuity in inventing mechanical devices needed for the tests.
These requirements Dr. Lutz possessed. In 1923 he was awarded
the Morrison Prize for his essay on ‘ ‘ The Colors of Flowers and
the Vision of Insects with Special Reference to Ultraviolet.” In
addition to his research work he did much to popularize ento-
mology and nature study. Tens of thousands of persons use his
4 ‘Field Book of Insects.” This was first published in 1918. A
second edition was published in 1921 and a third in 1935. Its
royalties put Dr. Lutz ’s four children through college.

In 1941 he wrote his last book entitled, “A Lot of Insects.”
This embodies accounts of the insects that, for the most part, were
the objects of Dr. Lutz’s curiosity, experimentation, and enter-
tainment over a period of many years, and includes his sound and
humorous entomological philosophy — all expressed interestingly
and in a lucid style. Dr. Lutz approached all his problems from
a stiumulating and thought-provoking viewpoint and this makes
for fascinating reading.

Dr. Lutz was a Fellow of the New York Academy of Sciences
and the American Association for the Advancement of Science,
a charter member of the Entomological Society of America and
its president in 1927, a member of the American Society of
Zoologists, the American Society of Naturalists, the Ecological
Society of America, Sigma Xi, Phi Beta Kappa, New York
Zoological Society, and the New York Entomological Society, his
presidency of the latter society having already been noted. He

Mar., 1944]

Weiss: Frank E. Lutz

67

was an advisor to the Buffalo Society of Natural Sciences and in
1937 a lecturer in Columbia University. He also served as chair-
man of the committee on Biological Relations Between Flowers
and Insects of the National Research Council.

Dr. Lutz made his home in New Jersey and funeral services
were held at his residence, 13 North Central Avenue, Ramsey,
New Jersey, on November 29, 1943. He is survived by his widow,
Mrs. Martha Ellen Brobson Lutz and four children, a son, Frank
Brobson Lutz, and three daughters, Anna Lutz, Ensign Laura
Lutz of the WAVES, and Mrs. Boyd Sherman.

These few paragraphs are but an inadequate summary of some
of the things accomplished by Dr. Lutz during his lifetime.
They fail to record many activites of which there are no records,
except in the memories of his friends. They fail to record the
happiness that Dr. Lutz’s chosen life-work brought to him, and
the pleasure that was his in creating problems and then solving
them. And they fail to mention his amiable and quizzical
philosophy of biological theories, flashes of which are apparent in
his writings. In a review of Dr. Lutz’s last book, and speaking of
his “Field Book of Insects” as well, Dr. H. M. Parshley said:
‘ ‘ They mirror a rare and admirable personality, a man of genuine
good will, a humorist, and one of a remarkable generation of
American naturalists.” — Harry B. Weiss.

Mar., 1944]

Bacon: Frank E. Lutz

69

BIBLIOGRAPHY OF FRANK E. LUTZ*

By Annette L. Bacon

1900. A study in the variations in the number of grooves upon the shells of
Pecten irradians (Lam.). Science, 12: 373.

1902. Report of Mr. Frank E. Lutz, Biologist. North Shore Improvement

Association. Reports on plans for the extermination of mosqui-
toes of the north shore of Long Island between Hempstead Harbor
and Cold Spring Harbor, p. 42-56.

The ecology of insect sounds. Can. Ent., 34: 64-66.

Inheritance of color among pointers. Science, 15: 571-572. (With
Eliz. B. Meek.)

1903. Note on the influence of change in sex on the intensity of heredity.

Biometrika, 2: 237-240.

Assortative mating in man, a cooperative study. Biometrika, 2 : 481—
498.

1904. Variation in bees. Bioh Bull., 6: 217-219.

1905. Biometry. Jour. Phil. Psych, and Sci. Meth., 2: 12-15.

Assortative mating in man. Science, 22: 249-250.

1906. Preserving spiders’ webs. Science, 23: 391.

The tegminal position in Gryllus. Can. Ent., 38 : 207-209.

1907. The merits of the fruit fly ( Drosophila ampelophila) . School Science

and Math., 7 : 672-673.

1908. Notes and literature: The effect of environment upon animals. Amer.

Nat., 42: 60-61.

The inheritance of the manner of clasping the hands. Amer. Nat.,
42: 195-196.

Notes on the inheritance of variations in the color pattern of Crioceris
asparagi. Psyche, 15 : 50-52.

The variation and correlations of certain taxonomic characters of
Gryllus. Carnegie Inst. Wash., Pub. No. 101, 63 p.

Combinations of alternative and blending inheritance. Science, 28:
317-318.

1909. Notes and literature: The effect of environment upon animals. Amer.

Nat., 43 : 55-57.

Notes and literature : The effect of environment upon animals. Amer.
Nat., 43: 248-251.

Experimental work with pomace flies. Amer. Mus. Jour., 9: 234-236.

1910. A brief history of antarctic exploration. A.M.N.H. Guide Leaflet 31,

27 p.

The annual scourge of flies and mosquitoes. Amer. Mus. Jour., 10:
183-185.

* Exclusive of book reviews and annual reports.

70

Journal New York Entomological Society

[Vol. LII

1911. Experiments with Drosophila ampelophila concerning evolution. Car-

negie Inst. Wash., Pub. No. 143, 40 p.

Relation between habit and structure in the insect world. Amer. Mus.
Jour., 11: 27-28.

Flea carriers of the plague. Amer. Mus. Jour., 11: 95-98.

Notes and literature: Mimicry. Amer. Nat., 45: 190-192.

Index to the generic names in Volume XXIX, Dr. Petrunkevitch ’s
1 1 Catalogue of Spiders of North, Central and South America and
Adjacent Islands.’ ’ Bull. A.M.N.H., 29:793-809.

1912. Inheritance of abnormal wing-venation in Drosophila. Proc. VII

International Zool. Congress (1907), p. 411-419. (Advance
reprint, 1910.)

String-figures from the Patomana Indians of British Guiana.

Anthrop. Papers A.M.N.H., 12: 1-14.

String-figures from the Upper Potaro. Timehri. Jour. Roy. Agric.

Com. Soc. British Guiana, 2(ser. 3): 117-127.

Do butterflies migrate? Amer. Mus. Jour., 12: 106-108.

The importance of insects. Amer. Mus. Jour., 12: 253—254.

1913. Experiments concerning the sexual difference in the wing length of

Drosophila ampelophila. Jour. Exper. Zool., 14: 267-273.

Eactors in aquatic environments. Jour. N. Y. Ent. Soc., 21: 1-4.

The distribution of Occidental spiders. Science, 37 : 567-568.

The life of the butterfly. House and Garden, August, p. 89-92.

1914. Humidity — a neglected factor in environmental work. Amer. Nat.,

48: 122-128.

Collecting in Cuba. Amer. Mus. Jour., 14: 99-106.

Biological notes concerning Drosophila ampelophila. Jour. N. Y. Ent.
Soc., 22: 134-138.

Our common butterflies. A.M.N.H. Guide Leaflet 38, 25 p. (With
F. E. Watson.) 4th ed., 1923, 31 p.; 5th ed., 1926, 21 p.; 7th
ed., 1941, 21 p.

How to collect and preserve insects. A.M.N.H. Guide Leaflet 39, 21 p.
3rd ed., 1917, 22 p.; 4th ed., 1920, 22 p.; 5th ed., 1924, 27 p.;
8th ed., 1936, 27 p.

1915. List of Greater Antillean spiders with notes on their distribution.

Annals N. Y. Acad. Sci., 26: 71-148.

Fragments of spider lore. Amer. Mus. Jour., 15: 424-426.
Experiments with Drosophila ampelophila concerning natural selec-
tion. Bull. A.M.N.H., 34: 605-624.

Outline plan of the hall of insect life. A.M.N.H. Circular, 4 p.

1916. Heredity and sex ; Mendelism and some of its recent developments.

Amer. Mus. Jour., 16: 228-242.

Faunal dispersal. Amer. Nat., 50: 374-384.

The geographic distribution of Bombidse (Hymenoptera) with notes
on certain species of Boreal America. Bull. A.M.N.H., 35: 501-
521.

Mar., 1944]

Bacon: Frank E. Lutz

71

Insects, an inexhaustible and relatively untouched field for recreation
or research. Amer. Mus. Jour., 16: 525—532.

1917. Heredity and sex. Mendelism and some of its recent developments

Scientific American Supplement, 84(2169) : 56-58.

1918. Field book of insects. G. P. Putnam’s Sons. New York, N. Y.

509 p.

General rules for the preparation of manuscript for the Bulletin and
Memoirs of the American Museum of Natural History. A.M.N.H.
Special Brochure No. 2, 14 p.

Insects and disease. A.M.N.H. Guide Leaflet 48, 73 p. (With C.-E.
A. Winslow.)

The control of insects injurious to gardens. A.M.N.H. Circular, 3 p.

1919. Scientific zoological publications of the American Museum for 1918.

Summary of work on invertebrates, fishes, amphibians, and birds.
Nat. Hist., 19 : 341-346.

Scientific zoological publications of the American Museum. Summary
of work on fossil mammals. Nat. Hist., 19: 731-733.

1920. Scientific zoological publications of the American Museum. Summary

of work on whales. Nat. Hist., 20: 107-109.

An entomologist in Colorado. Nat. Hist., 20: 312-325.

Our most brilliant butterflies. Country Life, 38(4): 27-33. (With
F. E. Watson).

Foreign insects newly come to America. Nat. Hist., 20: 501-502.
Notes on the distribution and bibliography of North American bees
of the families Apidse, Meliponidse, Bombidae, Euglossidae, and
Anthophoridae. Bull. A.M.N.H., 42: 491-641. (With T. D. A.
Cockerell.)

How crickets chirp. The Woodcraft Totem Board, 4(4) : 3.

How some insects walk on water. The Woodcraft Totem Board,
4(5): 3.

How moths and butterflies spend the winter. The Woodcraft Totem
Board, 4(6) : 3.

1921. How do insects make galls. The Woodcraft Totem Board, 4(7): 3.
How bees make honey. The Woodcraft Totem Board, 4(8) : 7.

How whales spout. The Woodcraft Totem Board, 4(9): 3.

How to collect insects. The Woodcraft Totem Board, 4(10) : 3.

How to preserve insects. The Woodcraft Totem Board, 4(11) : 3.
Geographic average, a suggested method for the study of distribution.
Amer. Mus. Novitates, 5, 7 p.

Field book of insects. Second edition. G. P. Putnam’s Sons. New
York, N. Y. 562 p.

1922. The reaction of Drosophila to ultraviolet. Science, 55: 519. (With

F. K. Richtmyer.)

Altitude in Colorado and geographical distribution. Bull. A.M.N.H.,
46: 335-366.

1923. Flowers and their insect visitors. Nat. Hist., 23 : 125-134.

72

Journal New York Entomological Society

[Vol. LII

1924. Apparently non-selective characters and combinations of characters,

including a study of ultraviolet in relation to the flower-visiting
habits of insects. Annals N. Y. Acad. Sci., 29 : 181-283.

Amateur entomologist and the Museum. A survey, from the half-
century mark, of the Department of Entomology, American
Museum. Nat. Hist., 24: 337-346.

Hunting stingless bees where east seems to be west. Nat. Hist., 24:
494-508.

Insect sounds. Bull. A.M.N.H., 50: 333-372.

1925. The new insect groups in the American Museum. Nat. Hist., 25:

126-135.

1926. Nature trails, an experiment in out-door education. A.M.N.H. Misc.

Pub. 21, 36 p. Second edition, 1931.

Taking nature lore to the public. Nat. Hist., 26: 111-123.

The friendly insects. Nat. Hist., 26: 147-151.

Principal orders of insects. Nat. Hist., 26: 164, with chart. (With
A. J. Mutchler.)

Our common butterflies. Nat. Hist., 26 : 165-183. (With E. E. Wat-
son.) (Reprint of Guide Leaflet 38.)

Insect sounds. Nat. Hist., 26: 206-213.

A nature trail in the sky. Nat. Hist., 26: 411-428.

1927. The still-open road. Nat. Hist., 27 : 373-382.

Wind and the direction of insect flight. Amer. Mus. Novitates 291,
4 p.

A much-abused but still cheerful cricket. Jour. N. Y. Ent. Soe., 35:
307-308.

1928. A new species of bacteria and the gall of an aphid. Amer. Mus. Novi-

tates 305, 4 p. (With E. Martin Brown.)

Little “beasts of prey” of the insect world. Nat. Hist., 28: 188-190.
Insects that erect tents. Nat. Hist., 28: 264-268.

1929. Experiments with “wonder creatures.” Nat. Hist., 29: 160-168.
Observations on leaf -cutting ants. Amer. Mus. Novitates 388, 21 p.

1930. An analysis by movietone of a cricket’s chirp ( Gryllus assimilis).

Amer. Mus. Novitates 420, 14 p. (With W. R. Hicks.)
Caddis-fly larvae as masons and builders. Nat. Hist., 30: 276-281.
Aquatic insect pets. Nat. Hist., 30: 389-401.

1931. Light as a factor in controlling the start of daily activity of a wren

and stingless bees. Amer. Mus. Novitates 468, 9 p.

Insects vs. the people. Nat. Hist., 31: 49-57.

In defense of insects. Sci. Mo., 32: 367-369. (Science Service Radio
Talks.) (Reprint of Insects vs. the people.)

Insects vs. the people. Condensed from The Scientific Monthly, April,
1931. Readers Digest, 19 : 359-360, 363.

In defense of insects. Article 28 in Science Today, edited by Watson
Davis, Harcourt Brace and Co., p. 184-188 (reprint).

Mar., 1944]

Bacon: Frank E. Lutz

73

A simple evaporimeter and some data obtained by its use in the Canal
Zone. Ecology, 12: 445-448.

Notes on the animal life of thermal waters in the Yellowstone National
Park. Amer. Mus. Novitates 498, 10 p.

1932. Our ignorance concerning insects. Can. Ent., 64 : 25-29, 49-56 and

73-78.

Experiments with Orthoptera concerning diurnal rhythm. Amer. Mus.
Novitates 550, 24 p.

1933. Experiments with 11 stingless bees” ( Trigona cressoni parastigma )

concerning their ability to distinguish ultraviolet patterns.
Amer. Mus. Novitates 641, 26 p.

li Invisible” colors of flowers and butterflies. Nat. Hist., 33: 565-
576.

1934. The il buckwheat problem” and the behavior of the honey bee.

Amer. Mus. Novitates, 688, 10 p.

Reactions of Drosophila to 2537 A radiation. Amer. Mus. Novitates
706, 14 p. (With E. N. Grisewood.)

World of the bee remains a mystery. New York Times Magazine,
Aug. 19, p. 12 and 15.

From low to high. Grand Canyon Nature Notes, 9 : 327-329.

1935. Field book of insects. Third edition. G. P. Putnam’s Sons. New

York, N. Y. 510 p.

1936. How about the tent caterpillar? Nat. Hist., 37: 149-158.

1937. Culture methods for invertebrate animals. 590 p. By P. S. Galtsoff,

F. E. Lutz, P. S. Welch, J. G. Needham, et al. Comstock Pub.

Co. Ithaca, N. Y. Collecting and rearing terrestrial and fresh-
water invertebrates, by Lutz, Needham, Welch, p. 40-50.

1938. The insect glee club at the microphone. Nat. Hist., 42: 338-345, 378.

1939. Big bees. School Nature League Bull. No. 10 (ser. 9), 3 p.

1941. The truth about termites. Nat. Hist., 48 : 113-115.

A lot of insects. G. P. Putnam’s Sons. New York, N. Y. 304 p.

1942. Insects, ticks and human diseases. A.M.N.I1. Guide Leaflet No. 113,

38 p. (With C. H. Curran.)

Mar., 1944]

Forbes: Lepidoptera

75

LEPIDOPTERA FROM WESTERN PERU AND
ECUADOR

By Wm. T. M. Forbes

Department of Entomology, Cornell University
Ithaca, New York

The occasion of this note is a little lot of Lepidoptera collected
by Mr. and Mrs. D. L. Frizzell in the arid northwest corner of
Pern, and Puna Id., Ecuador. While not many, the striking
character of the fauna is indicated by the presence of two new
Citheroniidae, and the region evidently is worth intensive collect-
ing. Among the normal and widespread things, may be men-
tioned Coea acheronta, Herse convolvuli from the Parinas Valley,
near Negritos, Peru; Pholus labruscce, Celerio annei, Utetheisa
ornatrix from the Parinas Valley; Hymenia fascialis, Eudioptis
hyalinata and Conchylodes arcifera from the Quebrada Mogollon.

The following are worthy of more specific mention. A female
Monarch shows the dull color and heavy black of the Lima
females. Single females of Ascia monuste from Negritos, Peru,
and Puna Id., Ecuador, suggest but hardly prove a racial dif-
ference.

Lycaena ramon Dognin. Parinas Valley, May 7, 1939. This
species was described from near Loja, Ecuador, a high temperate
and semiarid locality in the heart of the Andes1 but is equally
at home at sea level, where I took it commonly at Lima. It also
occurs on the western slope of the Andes at Chosica and Matu-
cana, and we have a specimen from Eten, a little north of Lima —
so it doubtless covers the whole arid area of western Ecuador
and at least northern Peru. We received the Eten specimen as
hanno, and other material may be floating around under that
name, but it is easily recognized by the ocelli on the hind wing
below — two larger between M3 and Cu2, and two only a little
smaller behind Cu2. The following key to the American species
of Lycazna, subgenus Hemiargus will place it more precisely.

1 Brown, Ann. Ent. Soc. Am., 34: 832.

76

Journal New York Entomological Society

[Vol. LII

1. Postmedial spots of fore wing below large and black, contrasting with the

small and fuscous subterminal series isola

-. Pm. spots of fore wing similar to subterminal ones ....; 2

2. Hind wing below with a large ocellus in cell M3 (cell 3 of Herrich-

Schaeffer system), similar to the one behind it 3

Hind wing with no ocellus in cell M3 5

3. Inner half of hind wing below contrastingly darkened, nearly obliterat-

ing the usual marks, which are much enlarged in this area; four small

ocelli . martha

Hind wing with ground all one color, the fuscous spotting generally
uniform :..... ......... 4

4. Two ocelli in anal area (behind Cu2) about two-thirds as large as the

ones in front of it and similar ramon

These ocelli small with a small silver spot only, or dull and similar to the
anterior subterminal markings :.... sachaeina

5. Anal area with a single large ocellus, similar to the one in front of Cu2 ... 6
Anal area with two subequal and inconspicuous spots or ocelli, about as

in zachaeina, or none .....I..:........... 8

6. Gray-brown below with spots all small, subequal and grayish ... bahamensis

-. Pale gray below a......... : :.. ." 7

7. Hind wing below, and above in female, with a very broad white submar-

ginal band ; all spots below small and similar dominica

-. Hind wing below with less conspicuous white submarginal band or none ;
three of the dark spots black and conspicuous ...... ammon ( catilina auct.)

8. Largely gray above; hind wing with slight tail and anal lobe bornoi

— . Mostly blue above; hind wing evenly rounded hanno ( catilina )

Goniurus jethira Butler. Easily recognized by the very large
honey-yellow spots, which are even larger in this specimen than
in the type. (Lep. Exot., p. 65, pi. 25, fig. 4). Parinas Valley,
May 7, 1939. The original locality was merely “Peru” but I
did not take it either at Lima or in the Chanchamayo, and suspect
it is a specialty of the arid Northwest.

Arsenura harrietse, new species

Closely similar to A. richardsoni Druce in major features; the fuscous
ground very lightly dotted with black, most definitely on posterior half of
median area. Antemedial of two widely separated blackish lines, the inner
nearly straight and outer much bowed out below cell, as in richardsoni, but
with heavier blackish filling, and the lines themselves more contrasting ; post-
medial line much further out, nearer to subterminal than to discal lunule,
blackish, more definite and more bowed out opposite lower angle of cell, the
median area conspicuously whitish toward inner margin; the following line
(subterminal in position, but probably morphologically the outer postmedian)
black, defined outwardly by a clay colored line, which is much finer and more

Mar., 1944]

Forbes: Lepidoptera

77

contrasting than the defining pale shade in richardsoni ; course much as in
richardsoni, but closer to margin, especially on costal third, and not nearly
as much extended toward base on inner margin; subterminal area not pale
as in richardsoni, but as dark as ground on fore wing and contrastingly
blackish on hind wing, with much paler yellow-brown terminal area. Sub-
apical black spot shorter than in richardsoni, not distinctly defined with
white; the terminal area below it vaguely shading between dull and red
brown, without the contrasting red-brown wedges of richardsoni. Discal
lunule black and contrasting with central tawny lunule, as in richardsoni,
but without the tawny bar extending from its outer side.

Hind wing generally similar to fore wing, but with only a faint darker
antemedial shade in place of the double line and dark filling, this shade
incorporating the faint discal bar (which is more distinct in richardsoni ) ;
postmedial band as on fore wing, much more conspicuous than in richardsoni ;
the outer pattern differing from the fore wing as noted.^ Wing form rounder
than in richardsoni, the apex of fore wing and angle of hind wing less
extended. Under side much less mottled than richardsoni, with three wavy
outer bands, varying from obsolescent to rather conspicuous, the outer
strongest and middle one weakest. Body plain brown as in richardsoni.

Expanse 92-110 mm., much smaller than richardsoni.

Puna Id., Ecuador; type and two paratypes in collection Cornell Uni-
versity.

This may possibly be a race of richardsoni, but the discontinu-
ous distribution, different wing form and many differences in
pattern suggest rather a good species. The following skeleton
key will place it in the genus :

1. Antemedial line double, of an outwardly oblique inner and a strongly

excurved outer element, the latter sometimes faint; st. space on pos-
terior half of fore wing and hind wing much broader than terminal

area I ...... 2

-. Am. line single, straight and outwardly oblique; both wings with a

sharply defined even slender pale marginal stripe *romulus

-. Am. line single, outcurved or angled, inwardly oblique to inner margin;
st. space narrower, usually much narrower than terminal space, except
sometimes for narrow extensions 4

2. Fore wing roundly falcate and deeply excavate below apex; hind wing

with tooth large, 12 mm. long on anterior side; hind wing with inner
st. line deeply sinuate, passing half way between margin and cell

at Mx ■„ *championi

-. Wings less irregular; st. line of hind wing crossing Mx two-thirds way
out to margin : 3

3. Discal spot of fore wing with a simple orange central lunule; margin of

wings hardly irregular *harrietae

78

Journal New York Entomological Society

t Vol. LII

Discal spot of fore wing with a short extension of middle of outer side
of lunule, forming a Greek e ; tail of hind wing 6 mm. long.

*ricliardsoni

4. Hind wing at least with a series of dark spots or lunules in terminal

area, wholly distinct from the subterminal complex; fore wing with
black markings conspicuous in cells Mx and almost always M2, the
upper usually joining to the apical pattern but conspicuous, the lower

usually free 5

-. Hind wing without this series of markings, though often with somewhat
similar extensions of the st. area; black patches in cells Mx and M2
normally absent, sometimes mere dashes, or lost in general blackish
shading : 11

5. Generally smaller species (female alcmene expanding 140 mm.), head

solid black, contrasting with the paler brown or fuscous thorax 6

-. Larger species (160 mm. and often more) ; head with at least a contrast-
ing pale bar over bases of antennae 8

6. Postmedial line not defined with pale; st. area expanded into a large

patch, occupying two-thirds the area between st. line and margin in

cells B4 and B5 . alcmene

-. Pm. line conspicuously defined by a following dirty white shading; st.
area below apex less extensive .4$;. 7

7. Markings of fore wing corresponding to the admarginal lunules of hind

wing taking the form of two similar large blotches in cells M3 and Cux

(feet not seen) pandora

-. This element of pattern taking the form of a waved diffuse admarginal

line ; tarsi concolorous dark brown *angulata

-. This element obsolete, except at anal angle, where it is not conspicuous;
tarsi cream white, contrasting *xanthopus

8. No black st. patch in cell M2; pm. area of both wings heavily shaded

with black; discal spot of fore wing lunulate; no admarginal spots on

fore wing sylla, hercules

-. Cell M2 heavily marked subterminally with black; discal spot a simple
bar . | : 9

9. No admarginal lunules on posterior part of fore wing; the black spot in

M2 fused with the blotch in cell M* aspasia

-. Spot in cell M2 separate, conspicuous, and followed with dark shades in
cells M3 and Cux similar to those on hind wing , 10

10. Ground with strong yellowish tint ; the two black patches in cells Mx and

M2 similar, very large, separated by hardly more than the light vein,

and scaled heavily with blue *meander

— . Ground with olive tint; the second black patch narrow and ovate.

*biundulata

11. Inner subterminal line nearly even, and marked with contrasting whitish

dots on veins or more irregular whitish patches; antemedial line when
distinct formed of a straight or concave bar across cell and a very
oblique lower portion from lower side of cell to basal angle 12

Mar., 1944]

Forbes: Lepidoptera

79

— . Inner st. line more irregular, normally with two large scallops between
M3 and Cu2, with only limited and irregular white marks; am. line
when distinct with lower portion less set off from portion crossing cell,
frequently in the form of a single excurved band 14

12. Lower segment of am. line conspicuous, in line with and more or less

continuing the black shade subterminally across cell Mx; a conspicuous

pale st. patch just below it in cell M2 ponder osa

— . Lower segment of am. line inconspicuous ; no single pale st. patch 13

13. Outer margin strongly irregular; postmedial area contrasting bright

chestnut brown loatesi

— . Margins less irregular; ground rather even dull light brown.

*crenulata

— . Margins still less irregular; pm. area somewhat contrasting, but light
brown arcaei

14. Discal spot lunulate, with contrasting pale center; dentations of inner

and outer st. lines if present not closely corresponding 15

— . Discal spots simple; the inner and outer st. lines closely parallel over
the dentations at cells M3 and Cux 16

15. Inner st. with distinct and outer with very strong dentations, the white

accompanying shade irregular *cymonia

— . Inner st. line nearly straight, and with even accompanying white shade;
outer st. obsolete thomsoni

16. Outer st. line with very strong, narrow black-filled dentations on both

wings, three or four of them on fore wing similar polyodonta

— . Outer st. line closely fitted to inner, both without strong dentations, save

for two in cells M3 and CUi of fore wing group *armida

— . Intermediate; the two upper teeth on fore wing present, but only half
as large as the two lower, and filled not with special black triangles
but with extensions of the generally blackish contrasting pm. area;
very large, expanding 175 mm * archianassa

In structural characters, the wing-form varies too widely in
obviously closely related species to serve for major subdivision,
but two groups are set apart by having pectinate antennae, sylla
and arcaei with their relatives — alternatives 5 and 12 of the key.
The residue, so far as seen, and including harrietce, have serrate
and fasciculate antennae.

Dysdwmonia species. There was badly broken material of a
very striking undescribed Dysdaemonia, with scalloped wings.
It will be described by Mr. Johnson, who has a better specimen
from the same region.

* Species represented in coll. Cornell University, many of them the gift of
Mr. Frank Johnson.

80

Journal New York Entomological Society

[Vol. LII

Givira tristani Schaus. A rubbed specimen from Puna Id. is
this species or very close.

Euclea copac Schaus. Puna Id. Agrees so far as can be seen,
but not good enough for certainty. The species was described
merely from “Peru.”

S 'eirocastnia elaphebolia Druce. Looks to me like a good spe-
cies. Described from Ecuador.

Monodes convexa, new species

Superficially similar to the North American festivoides group, but with
relatively much smaller body and arched fore wings. Structures of the nor-
mal Monodes, without sex-scaling ; male antennae ciliate, legs unmodified, with
a rough tuft beyond middle of mid tibiae; vestiture normal for Monodes, as
described by Hampson, but with the posterior thoracic tuft extended back,
almost completely covering the basal abdominal tuft, and roundly truncate
behind. Palpi with second joint upturned only a little beyond middle of
front, as also in normal species of Monodes.

Body fuscous, thorax somewhat mottled, with darker lower half of collar,
the upper half of front, vertex and lower half of collar contrasting blackish
in dark specimens; palpi with first and second joints with paler apices, the
outer sides contrasting blackish in dark specimens. Pore wing gray, varying
extremely in tint, light specimens with the terminal third much darker, dark
ones sometimes with the costal area rather darker. Costa with numerous
dark bars in light specimens, dark, cut with the pale gray ante- and post-
medial lines in dark specimens, and with about four small whitish bars
between postmedial and the pale apical shade. Subbasal line of vague paler
powdery scaling, toward costa, toward inner margin represented by an oblique
blackish bar; antemedial obscure, except for the contrasting pale bar at
costa; postmedial represented by dark spots on veins, followed by minute
whitish ones, broadly and evenly excurved on costal two thirds, then oblique
in to inner margin and slightly concave. Subterminal obscure, sometimes
indicated as the irregular boundary between a grayer terminal and browner
subterminal area; orbicular and reniform spots large, the orbicular outlined
with black except above, usually heavily before and behind; reniform very
large, only partly outlined; the area before orb. blackish, the filling between
orb. and ren. blackish or shaded with dark, without the sharp boundaries of
the festivoides group, the space between ren. and postmedial line usually
somewhat darkened, but without a blackish spot. Claviform minute, whitish,
usually contrasting, heavily outlined with black, especially before and beyond.
Apical oblique shade varying from obsolete to conspicuous, cream white to
ash gray, with the basal half shaded with buff in light specimens and slightly
warmer brown in dark ones; starting from pm. line opposite cell, curving up
and widening in a horn shape, and ending on outer tenth of costa; partly
edged behind with black. Terminal blackish dots, obscure in dark speci-

Mar., 1944]

Forbes: Lepidoptera

81

mens, preceded by whitish points, alternating with the pm. ones. Fringe
powdery gray, the outer half rather smoother and less powdery. Hind wing
dirty white, shading into fuscous on outer half; alula cream, clothed with
large scales and conspicuous. Expanse 17-20 mm.

This species in normal specimens will run in Hampson’s key to
the festivoides group, from which it is distinguished by the total
lack of warm coloring, much smaller body and arched fore wings,
also in maculate forms by the less sharply defined blackish about
the orbicular and reniform. It is probably close to bogotana
Felder and aphaidropa Dyar (which are presumably merely color
forms of each other) but is smaller, and none of our series are as
pale as Felder’s figure of bogotana, nor show the reddish shadings
along costa of aphaidropa. It varies enormously, from speci-
mens (males) with the basal two thirds luteous and only the outer
third blackish, much like Felder’s figure, to specimens (females)
that are wholly suffused with smoky gray and coal black, with
all the markings obsolescent. Such specimens will probably key
out to phceopera, but differ from it and its relatives by the rather
even dull gray, with the darker area between orbicular and reni-
form, and the paler apical area at least faintly visible.

The male genitalia of Monodes are extremely varied, and con-
vexa resembles nucicolora, grata or fusimacula more closely than
festivoides, having the valves slender, with a complicated basal
chitinization composed of parts of sacculus, costa, and perhaps
clasper, and sending a spike forward across the costa; weak
clasper at a third way out, crossing costa; juxta slenderly ex-
tended as a complete anellus, and transtilla also bent into a round
arch closely parallel to it. MCdoeagus with two massive spines,
formed of fused cornuti.

Holotype, male from Chosica, Peru, May 25, 1920. Numerous
paratypes of both sexes, from Chosica and Lima, Peru, May
1920; a couple caught by Parish at Lima in 1915; and a pair
from the Amotape Mts., N.W. Peru, collected by the Frizzells;
all in collection Cornell University. I believe other specimens
exist in collections, perhaps labelled bogotana, but the Lima
fauna has been extraordinarily neglected, considering how many
good collectors have passed through it or even used it for a base.

Cydosia phcedra Druee. Puna Id.

Cobubatha numa Druce. Amotape Mts.

82

Journal New York Entomological Society

[Vol. LII

Dichochroma, new genus

Similar to the Pyraustine genus Dichogama. Yestiture of large, smooth
scaling; palpi upturned to middle of front, close-scaled, slightly flattened
against the front; the segments well marked off, third segment larger than
in Dichogama, two-thirds as long as second, continuing the direction of sec-
ond; maxillary palpi rough-scaled, flattened against the face, as in Dicho-
gama. Tongue strong. Legs very short for a Pyraustine, as in Dichogama,
mid tibia about as long as femur without trochanter, and tarsus hardly
longer. Fore wing presumably with the long Arctiid-like frenulum hook of
Dichogama (male not seen) ; R3 and R4 stalked, R5 stalked with M4, well
separated from R4, M2 and M3 stalked, Cu4 parallel to M3, arising well before
end of cell. Hind wing without fringe on Cu; M2 and 3 strongly stalked, as
in D. fernaldi, but unlike the other Dichogamas at hand.

Third A of fore wing is strong and makes a wide loop, but I cannot see
if it runs back into 2d A.

This genus is clearly a development of Dichogama, differing
from it, and from all Pyraustinae known to me in the stalked
R5 and Mi (veins 6 and 7) and from most in the stalking of
M2 and M3 in both wings. It is also far smaller than any Dicho-
gama yet known, and is found on the Mainland, while Dichogama
is essentially Antillean, only D. diffusalis not yet being known
from the Antilles. In Hampson’s key (Proc. Zool. Soc., 1898,
594) it will run to Hymenia or Macaraetera, according to how the
third segment of the palpus is interpreted, but has no real kin-
ship to either. The stalked veins will easily separate it.

By the way there is no reason for marking several species of
Dichogama “incertae sedis” as Klima does in Lep. Cat. 89, p. 122.
We have colotha, fernaldi and gudmanni from Porto Rico and
they are normal Dichogamas, save for the stalked instead of ap-
proximate M2 and M3 in fernaldi ; and amalyilis and bergii show
patterns that would hardly occur elsewhere ; but smithii, unques-
tioned by Klima, is the well known Noctuid, Casandria abseuzalis,
which I found common in Porto Rico.

Dichochroma muralis, new species

Head and thorax mouse gray, dusted with whitish scale-tips; thorax im-
maculate ; shaft of antennae blackish ; palpi with first segment whitish, second
mouse gray, but whitish along the ventral inner keel; third segment blackish
with contrasting dirty white tip. Under side cream white, the front side of
the fore legs fuscous. Abdomen above mouse gray, immaculate, below
nearly white.

Mae., 1944]

Forbes: Lepidoptera

83

Fore wing mouse gray, immaculate but with pale scale-tips especially
toward base, and sparsely overlaid with narrow whitish strap-shaped scales;
fringe whitish. Hind wing translucent white with narrow and broken fus-
cous terminal line. Expanse 16 mm.

Amotape Mts., N. W. Peru, H. & D. L. Frizzell; type one female in coll.
Cornell University.

In sum this little fauna from the north end of the arid coastal
strip of South America is a curious one. While the typical mate-
rial of this strip is present ( e.g ., Celerio annei and Monodes con -
vexa) there is also a definite Central American element (notably
the Arsennra, which is closest to A. richardsoni) , and a few spe-
cies which now appear to be endemic, such as the undescribed
Dysdsemonia, Euclea copac and Dichochroma murina. Plainly
many more interesting things are due to come out of the area.

Mar., 1944]

Miller: Drosophila

85

DROSOPHILA MELANURA, A NEW SPECIES OF THE

MELANICA GROUP

By Dwight D. Miller*

California Institute of Technology, Pasadena, California

During most of the summer of 1941 the author collected Droso-
philas and related forms on the River Campus of the University
of Rochester at Rochester, New York. One of the commonest
types to be found in the fermented banana traps used in the col-
lections was a form identified as Drosophila melanica Sturtevant
(1916). From June 5 through August 25 there were gotten
1263 individuals of this species, 538 of which were females, 725
males. In a recent paper Patterson (1942) has indicated that
D. melanica should be divided into two subspecies, melanica and
paramelanica, differing in distribution, melanica having been
found in the southeastern United States and in Mexico, para-
melanica occurring in the northeastern part of this country. It
is probable, on the basis of locality, that the D. melanica indi-
viduals collected at Rochester belonged to the subspecies para-
melanica. Towards the end of the collecting period, from
August 19 through August 25, there occurred in the traps some
male Drosophilas that seemed to differ from the D. melanica
males only in the unusually dark pigmentation of their genital
region. Altogether there were 10 such males collected. Since
it was suspected that these males represented a new form, a num-
ber of apparently D. melanica females collected at the time were
isolated individually into culture bottles in the hope that some
of them would have offspring, males of which would be of this
new type. Unfortunately, none of the females so isolated bred.
Since it seemed unlikely that the females would have any progeny
if left to themselves, an attempt was made to mate them to some
of the melanica- like males. Culture bottles that had contained
one of the females with two such males yielded some offspring.
The male progeny were all of the new type in that their genital

* This study was begun while the author was holding a teaching assistant-
ship at the University of Rochester, Rochester, New York.

86

Journal New York Entomological Society

t Vol. LII

region was darkly pigmented. These offspring proved to be fer-
tile, and through allowing them to mate among themselves a
strain was derived. Males of succeeding generations have all
persisted in differing from D. melanica males in the dark pig-
mentation of the genital region. On the basis of this constant
character of difference, as well as because of others noted below,
the melanica- like form collected at Rochester is here designated
a separate species, Drosophila melanura. A description of the
new species is given next, with notes on comparison of the new
form with D. melanica following just afterwards.

Description of the Species

Drosophila melanura, sp. nov.

External Characters of Imagines.

$ Arista with about 9 branches. Antennas brownish, pollinose. Front
blackish. Middle orbital length about i that of posterior one. Second oral
length about ^ that of first. Carina broad below, sulcate. Face brownish.
Cheeks pale brownish, about i greatest diameter of eye at their greatest
width. Eyes red, with black pile.

Acrostichal hairs in irregular rows; about six rows at the anterior dorso-
central bristles. Anterior scutellars convergent. Mesonotum dull brown,
with four slightly paler longitudinal stripes, two extending forward from
just inside anterior dorsocentral bristles, two lying just outside the lateral
pairs of dorsocentral bristles. Pleurae brownish. Sterno-index about 0.9.
Legs pale brownish. Two apical bristles on first tibiae, one on second; pre-
apicals on all three.

Broad, brown bands on abdominal segments 2 through 6, with posteriorly
narrowing median interruptions, with but little interruption near the lateral
margins of the tergites. Genital region (genital arch, anal plates, claspers,
penis apparatus) dark brown.

Wings clear, veins brown. Costal index about 2.9; 4th vein index about
1.4; 5x index about 0.9; 4c index about 0.6. Two bristles at apex of first
costal section. Third costal section with bristles on its basal

Body length about 3.3 mm. (alive) ; wings 2.8 mm.

$ Broad, brown bands on abdominal segments 2 through 6, with posteri-
orly narrowing median interruptions, often with wide lateral interruptions
on the 6th segment.

Body length about 3.5 mm. (alive) ; wings 3.0 mm.

Internal Characters of Imagines.

Testis with about 3 inner coils (probably the seminal vesicle) and 3 outer
ones, the coiled portion tending to be orange. Ejaculatory sac with 4 long
diverticula, 2 anterior and 2 posterior.

Spermathecae chitinized. Ventral receptacle with about 30 coils.

Mar., 1944]

Miller: Drosophila

87

Additional notes.

Egg. — With 2 filaments, each about f the length of the egg.

Puparium. — Amber. About 9 branches in each anterior
spiracle. Horn index (length of puparium/length of anterior
spiracle horn) about 12.

Chromosomes. — Female metaphase plate contains : one pair of
large V’s, two pairs of rods, one pair of medium V’s, and one pair
of small V’s. Male metaphase plate has a J in place of one of
the large V ’s.

Key Position. — The following is to be taken as an expansion
of the first line of couplet 50 of the Key to North American Spe-
cies of Drosophila of Sturtevant (1942), to which point an
attempt to classify a specimen of D. melanura should lead one.

50a. Male genital region dark brown melanura

Male genital region not dark brown melanica

Distribution. — This species has been collected only in a small
wood on the campus of the University of Rochester (River
Campus), Rochester, New York.

Type specimens of D. melanura are deposited in the American
Museum of Natural History, New York, N. Y.

Comparison with D. melanica

Drosophila melanura has been compared with individuals
taken from D. melanica strains kept in this laboratory. These
strains have been classified as to subspecies ( melanica or para-
melanica) by Prof. A. H. Sturtevant. The above description of
D. melanura , as well as the notes which follow, is based largely
on individuals of the single existing strain of this species, de-
rived as indicated in the first paragraph.

The impression has been gotten that D. melanura is a larger
form than D. melanica. This has been gotten not so much from
wild individuals as from flies raised in the laboratory (the size
measurements given in the above description were taken from
“average” laboratory individuals). The difference in size may
be partly due to the fact that the new species has not bred as
vigorously in the laboratory as has D. melanica and has usually
had the advantage of relatively uncrowded culture bottles.

88

Journal New York Entomological Society

[Vol. LII

D. melanura has also seemed to have a lighter body color than
D. melanica. This difference is especially noticeable on the dor-
sal surface of the thorax, which may be described as medium
brown in the new form, dark brown or very dark brown in D.
melanica. Accompanying this difference is a greater prominence
of the mesonotal stripes in melanura than in D. melanica. It is
the impression of the author that the D. melanica subspecies tend
to differ from each other in that melanica is somewhat darker
than paramelanica. The thorax color difference between me-
lanura and D. melanica melanica has been quite striking. Since
a body color difference was not appreciated at the time the col-
lections of melanura and D. melanica (probably paramelanica)
were being made, the separation of wild females on the basis of
color was not attempted. It remains to be seen how reliable a
criterion this is for the identification of wild individuals in
general.

The abdominal banding pattern of D. melanura males was
found to be different from that of the D. melanica males exam-
ined. Whereas in melanura males the band on the 6th segment
(as well as the others) was always found to be complete laterally,
this was never seen to be the case in D. melanica males. In a
D. melanica strain ( paramelanica ) from South Amherst, Massa-
chusetts, the males’ abdominal bands were all found to become
indistinct near the lateral margins of the tergites. In a D.
melanica strain ( melanica ) from Walnut Creek, Texas, while the
bands on the 2nd through 5th abdominal segments generally
extended all the way to the edges to the tergites, the band on the
6th segment was interrupted laterally.

Mention has already been made of the darkly pigmented geni-
tal region of males of the new species. This seems to be the best
character whereby it may be recognized as different from D. me-
lanica. In D. melanura males the genital arch, anal plates, and
claspers become dark brown a few days after emergence. The
plates of the penis apparatus also become dark. The result is
that the genital region stands out conspicuously against the light
ventral abdominal wall. In D. melanica the male genital region
seems relatively inconspicuous.

In D. melanura and in D. melanica the penis apparatus appar-
ently consists of two rather broad lateral plates as well as the

Mar., 1944]

Miller: Drosophila

89

structure probably homologous to the chitinous rod called the
penis by some authors (for example, Nonidez, 1920, in D. melano-
g aster). The appearance of the lateral plates was found to

Fig. 1. A. Male genital region of B. melanura. B. Male genital region
of B. melanica paramelanica (South Amherst, Mass.). The labels indicate:
the anal plates (a), claspers (c), genital arch (g), and penis apparatus (p).
These drawings were made from dead specimens, and the penis apparatus is
shown farther forward than its usual position in living, etherized individ-
uals. C. Penis apparatus of B. melanura. D. Penis apparatus of B.

melanica melanica (Walnut Creek, Texas). E. Penis apparatus of B.
melanica paramelanica (South Amherst). C, D, and E are side view draw-
ings made on a somewhat larger scale than A and B. The edges labelled
with a “p” in C and E correspond to the edges so labelled in A and B
respectively. The point labelled with an “x” in D was found to vary some-
what in prominence.

differ between melanura and D. melanica. In melanura they
were deeply incised at the anterior edge (Figure 1, C), whereas

90

Journal New York Entomological Society

[Vol. LII

in D. melanica they were not so much or scarcely at all so incised
(Fig. 1, D & E). The structure of the plates in D. melanica
melanica strains examined (Walnut Creek, Texas; Coffeeville,
Kansas; Taneycomo, Missouri) seemed intermediate (Fig. 1,
D) between that of D. melanura (Fig. 1, C) and that of D. me-
lanica paramelanica strains (Madison, Wisconsin; South Am-
herst, Massachusetts) (Fig. 1, E). In D. melanura the impres-
sion was gotten that the penis apparatus is somewhat larger and
more conspicuous relative to its surroundings than in D. melanica
(Fig. 1,A&B).

The anterior spiracle horns of the puparium were found to be
relatively shorter in D. melanura than in D. melanica. The horn
index (length of puparium/length of horn) was determined to
be about 12 in melanura, whereas melanica estimates were 9 for
the South Amherst strain ( paramelanica ) and 7 for the one from
Walnut Creek {melanica).

The chromosomes of D. melanura were investigated by means
of acetic orcein smear preparations of larval ganglia and of
ovaries and testes taken from pupse and adults (using the 70 per
cent acetic acid stain of LaCour, 1941). Examples of chromo-
some groups are given in Figure 2, A, B, and C. It may be seen
that the smallest chromosome is not dot-shaped, as was reported in
D. melanica by Metz (1916), but, rather, a small V. An investi-
gation of D. melanica chromosomes, both paramelanica (South
Amherst and Madison) and melanica (Walnut Creek), has con-
firmed Metz’ report of dot-like chromosomes in this species
(Figure 2, D, E, and F). Of interest here is the statement of
Griff en (1942) that the melanica group species D. nigromelanica
Patterson and Wheeler (1942) has rod-shaped microchromosomes
rather than dots. The salivary gland chromosomes of D. me-
lanura have not been studied.

As may be seen from the first paragraph of this paper D.
melanura males were gotten in the summer, 1941, collections at
Rochester in a much smaller number than were D. melanica
males. Moreover, D. melanura was only collected past the
middle of summer, late in August, while D. melanica occurred in
the traps by early June. An ecological difference between the
two species is suggested. D. nigromelanica was collected also

Mar., 1944]

Miller: Drosophila

91

and had an occurrence in the traps similar to that of D. me-
lanura; from July 26 to August 25 there were gotten 26 indi-
viduals of this species, 11 of which were males. A record of the
collections of these melanica group species is given in Table 1.

Fig. 2. A and B. D. melanura ovary metaphase figures. C. D. melanura
testis metaphase. D and E. D. melanica paramelanica (South Amherst)
ovary metaphases. In E some of the chromosomes, including the dot-like
ones, are obviously split. F. D. melanica melanica (Walnut Creek) testis
metaphase. The dots are close together. G, H, and I. Metaphase figures
found in the testes of hybrids derived from melanica (Walnut Creek) females
mated to melanura males.

D. melanura has been kept with some difficulty on the Droso-
phila culture media in use at this laboratory. The impression
has been gotten that D. melanica has not been so difficult to main-

92

Journal New York Entomological Society

[VOL. LII

tain. In December, 1941, onr corn meal-molasses-agar medium
was changed in that agar was omitted from it and rolled oats
were added (formula of Dr. R. H. MacKnight). The result has
been a somewhat softer, wetter medium. From soon after the

Table 1. Week-by-week record of melanica group species collections at
Rochester, New York, during the summer of 1941. The number of D.
melanura females gotten is not recorded; these were most probably all classi-
fied as D. melanica females. Since the circumstances of collection (number
of traps, number of collections, etc.) varied from week to week, the variation
in absolute numbers should not be taken to reflect very well the changing
state of the wild population.

Collections for the week beginning: .

June 1 June 8

June 15

June 22

June 29

melanica

$ $

5

18

40

$ $

2

3

3

26

41

nigromelanica

melanura

July 6

July 13

July 20

July 27

Aug. 3

melanica

$ $

171

47

74

134

115

$ 2

111

19

38

125

93

nigromelanica

$

1

$ $

1

1

6

melanura

Aug. 10

Aug. 17

Aug. 24

Total

Total

melanica

$ $

46

57

18

725

$ $

26

35

16

538

1263

nigromelanica

$ $

7

3

11

$ $

5

2

15

26

melanura

$ $

7

3

10

10

change was made until about the middle of May, 1942, the D.
melanura strain refused to breed at all. Following this period
there has been a gradual recovery of ease of culture.

Hybrids with D. melanica

A few attempts were made to cross D. melanica females with
D. melanura males, several individuals of a kind being used in
each mating. Progeny have been gotten both from crosses of
melanica (Walnut Creek) females by melanura males and from
paramelanica (Madison) females by melanura males (Table 2).

Mar., 1944]

Miller: Drosophila

93

A few matings were also made between D. nigromelanica females
and melanura males, using a Harden County, Texas, strain fur-
nished by Prof. J. T. Patterson, but no progeny were gotten
(Table 2).

In general the interspecific cross progeny have been not very
different from the maternal species. The penis apparatus of the

Table 2. Interspecific crosses involving D. melanura. The flies were
put into fresh culture bottles on the dates listed under each mating. The
numbers of progeny are placed opposite the dates appropriate to the bottles
in which they occurred.

Cross

Progeny

$ $ $ $ Totals

D. melanica melanica $ $ xD. melanura $ $

Walnut Creek

$ $ x Rochester $ $

(7 of each)

8- 7-42

8-10-42

8-15-42

Walnut Creek

$ 9 x Rochester $ $

(22 of each)

8- 8-42

8-11-42

15

18

8-15-42

7

7

22

25

47

Walnut Creek

$ $ x Rochester $ $

(15 of each)

8-10-42

8-15-42

20

15

8-19-42

40

35

8-24-42

7

6

8-31-42

46

31

9- 8-42

24

24

137

111

248

D. melanica paramelanica $ $ xfi. melanura $ $
Madison $ $ x Rochester $ $ (5 of each)

8- 8-42
8-11-42
8-15-42

Madison $ $ x Rochester $ $ (7 of each)
8-10-42
8-15-42
8-19-42

94

Journal New York Entomological Society

[VOL. LII

Table 2 — ( Continued )

Cross

D. melanica paramelanica $ $ x D. melanura $ $
(cont.)

Madison 9 $ x Rochester $ $ (15 of each)
8- 7-42
8-10-42
8-15-42
8-24-42

8- 31-42

9- 8-42

D. nigromelanica $ $ xD. melanura $ $

Harden Co. $ $ x Rochester $ $ (8 of each)
9- 8-42
9-12-42

Harden Co. $ $ x Rochester $ $ (4 of each)
9-1-42
9-4-42
9-8-42

Harden Co. $ $ x Rochester $ $ (13 of each)
9- 4-42
9- 8-42
9-10-42

2 $

6

18

5

29

Progeny
$ $ Totals

6

16

4

1

27 56

hybrid males, for example, has been rather similar to that of
males of the form to which the mother belonged. Both sexes
have seemed quite viable, and the sex ratio has been about 1 to
1 (Table 2).

Metaphase plates found in the testes of hybrid males derived
from both the kinds of crosses mentioned above contained one
dot-like chromosome and one small V (Fig. 2, G, H, and I). This
evidence may be offered as one form of proof of the hybrid nature
of these offspring. The salivary gland chromosomes of several
larvae were looked at, and although a very careful study of these
chromosomes was not made, it did seem likely that they were
heterozygous for a number of chromosomal rearrangements,
probably inversions.

Mar., 1944]

Miller: Drosophila

95

Table 3. Backcrosses of hybrid females to B. melanura and to D.
melanica.

Cross

Progeny

$ $ $ $ Totals

( melanica 9 2 x melanura) $ 9 x B. melanura $ $

(W. C. $ $ xRoch.) $ 9 x Rochester $ $ (13
of each)

8- 31-42

9- 3-42
9- 8-42

(W. C. 9 $ xRoch.) 9 $ x Rochester $ $ (7
of each)

9-1-42

9-4-42

9-8-42

( paramelanica 9 2 x melanura ) 9 9 xD.
melanura $ $

(Mad. 9 2 xRoch.) 9 $ x Rochester $ $

(2 of each)

9- 4-42
9- 8-42
9-10-42

( melanica 9 $ x melanura) 9 $ x B. melanica
melanica $ $

(W. C. 9 $ xRoch.) 9 2 X Walnut Creek $ $
(17 of each)

9- 4-42

9- 8-42

9-10-42

45

30

9-18-42

26

15

9-25-42

31

20

102

65

(. paramelanica 9 9 x melanura) 9 9 x para-
melanica $ $

(Mad. 9 2 xRoch.) 9 9 x Madison $ $
(9 of each)

9-30-42

10- 3-42
10- 8-42

96

Journal New York Entomological Society

[Vol. LII

No offspring have been gotten from hybrid males and females
kept together. It seems likely that the males are sterile.
Although hybrid testes were found to contain many sperms, no
motility of these sperms in Kingers’ solution was seen. More-
over, acetic orcein smears have shown these testes to contain a
number of rather large, spindle-shaped, dully-staining bodies,
but no regular sperm heads. It is probable that these elongate
structures are the product of an abnormal spermiogenesis.

Attempts have been made to backcross hybrid females both to
D. melanura and to D. melanica males, and offspring were gotten
in the case of a group mating of hybrid females (Walnut Creek
22 x Rochester to Walnut Creek males ( melanica ) (Table
3). A rather obvious excess of females over males may be seen
in these progeny. Since no mutants were involved in the crosses
that gave rise to them, little could be concluded about the genetic
constitution of the back-cross individuals. Some of the males
were dissected 10 or 11 days after emergence and their testes
examined. Out of 20 such males 2 had testes containing small,
pyknotic bodies of various shapes, but no normal sperm heads,
while the others had normal appearing sperms. Several recently
hatched males were dissected and their testes examined for
chromosome groups. In one of the testes there were found
rather clear figures showing the presence of a dot and small Y
together, and in this testis there were a number of quite normal
appearing sperm heads.

SUMMARY

Drosophila melanura, a new species similar to D. melanica
Sturt., is described here. Notes on comparison of the two forms
are presented. These species differ cytologically in the appear-
ance of the smallest chromosome in the metaphase plate, D. me-
lanura having a small V where D. melanica has a dot. Hybrids
have been obtained from crosses between D. melanura females
and D. melanica males, and the female hybrids have proved to be
fertile.

ACKNOWLEDGMENTS

The author wishes to thank Dr. H. D. Stalker of Washington
University, St. Louis, Missouri, and Prof. A. H. Sturtevant of

Mar., 1944]

Miller: Drosophila

97

the California Institute of Technology for advice and assistance

in connection with this study.

LITERATURE CITED

Griffen, A. B. 1942. Relationships in the Melaniea Species Group. Univ.
of Texas Publ., 4228: 68-73.

LaCour, L. 1941. Acetic-orcein: A New Stain-fixative for Chromosomes.
Stain Techn., 16: 169-174.

Metz, C. W. 1916. Chromosome Studies on the Diptera. III. Additional
Types of Chromosome Groups in the Drosophilidse. Amer. Nat., 50 :
587-599.

Nonidez, J. F. 1920. The Internal Phenomena of Reproduction in Dro-
sophila. Biol. Bull., 39 : 207-230.

Patterson, J. T. 1942. Interspecific Hybridization in the Genus Drosoph-
ila. Univ. of Texas Publ., 4228: 7-15.

and M. R. Wheeler. 1942. Description of New Species of the

Subgenera Hirtodrosophila and Drosophila. Univ. of Texas Publ.,
4213: 67-109.

Sturtevant, A. H. 1916. Notes on North American Drosophilidse with
Descriptions of Twenty-three New Species. Ann. Ent. Soc. Amer.,
9: 323-343.

. 1942. The Classification of the Genus Drosophila, with Descrip-
tions of Nine New Species. Univ. of Texas Publ., 4213 : 5-51.

Mar., 1944]

Brown: Pieridje

99

NOTES ON MEXICAN BUTTERFLIES, II, PIERID^

By F. Martin Brown

The collections made by Hoogstraal’s parties and by Potts con-
tained 33 of the 71 species of Pieridse reported by Hoffmann.
Those lacking are primarily the truly tropical species that enter
Chiapas and the southern lowlands and some of the western
Nearctic things that are found in Sonora and Baja California.
I have added the data from a small collection made by H. D.
Thomas in 1936.

Several localities not mentioned in the first paper on the
Papilionidae (g.u.)# are noted here:

Geographic Data

Acahuato, Michoacan, 19° 20' N., 102° 20' W., 3000 ft.

“On the edge of the plateau above Apatzingan. Open semi-
desert with scattered trees below the town and open pine forest
above it. Transitional between the tropics and temperate area. ’ ’
Hoogstraql.

Cumbres, Vera Cruz, 6000 ft., km. 295 on road to Vera Cruz
from Mexico City.

“North and west of Orizaba, very definitely up and out of the
tropical zone and rain belt, into a dry upland type of country
even though still the bottom of the valley. A lot of straggling
thorn bush, tremendous organ cactus, a lot of Opuntia. Short
grass and rock slopes. Temp, at about 75 or less.” Potts.

El Mante, San Luis Potosi.

“A strange conglomeration in this region of swampy country
with tall palms and very tall thorn bush jungle next to bananas,
corn fields and maguey. Cactus growing in mud ! Temp, proba-
bly 95° F.” Potts.

Hda. Potrero Viejo, nr. Paraje Nuevo, Vera Cruz.

‘ ‘ Cultivated country for centuries, yet plenty of original vege-
tation nearby. I collected only in a small meadow near the
* Jour. N. Y. Ent. Soc., Yol. 51, No. 3, p. 161-178, 1943.

100

Journal New York Entomological Society

[Vol. LII

hacienda itself.” Potts. Between Fortin and Orizaba on the
Mexico City-Vera Cruz highway.

Hda. Santa Engracia, Tamaulipas.

I have no data on this station. F.M.B.

Monte Alban, Oaxaca, 16° 50' N., 96° 20' W., 6000 ft.

“ Hills back of Oaxaca, the only moist country nearby!”
Potts.

Ojo de Agua, Sabinas Hidalgo, Nuevo Leon, 26° 40' N., 100° 15'
W., 1000-1600 ft.

‘ ‘ Collections were made up to 1600 ft., low shrubs and cactus.”
Hoogstraal.

Orizaba, Vera Cruz, 18° 45' N., 96° 50' W., 2200 ft.

“About a kilometer SE. of the town. Just the top end of the
tropics or at least the jungle tropics, along the valley slope.
Impenetrable underbrush, lush, rains some 9 or 10 months of the
year here. ” Potts.

Sta. Lucrecia.

A Thomas station which must be on the west coast, probably in
Michoacan, from the material so labeled. F.M.B.

Rio Balsas, Guerrero, 18° dz S., 98°-102° W.

The great river basin of southern Mexico. ‘ ‘ The river itself is
pretty dry. Collected up a side canyon to the south. Temp.
105-110, muggy. Thorn bush and rocks.” Potts. The collec-
tion was made near Mexcala (18° 0' S., 99° 30' W., 2400 ft.).

Tuxpango, Vera Cruz, ? 1500 ft.

“Top of a big wide barranca SE. of Orizaba. County simi-
lar to ‘Orizaba.’ ” Potts.

Dismorphiinae

I am surprised that these two collectors, Hoogstraal and Potts,
were so unsuccessful in finding members of this sub-family. At
least nine species are known from the areas they visited. Only
Hoogstraal collected any and his material represents one species.

Mar., 1944]

Brown: Pierid^:

101

61. Enantia jethys Boisduval.

G. & S. (1), 2: p. 182-183, 727, pi. 61, f. 56; pi. 74, f. 27,

28 (as albania).

R. (2), p. 100.

H. (3), p. 654-5.

Tancitaro, Michoacan, 6600 ft. 3 J'.J' 2 52 vii.20-viii. 14.40
(H.H.).

Hoffmann, p. 654U5, recognizes three species, citronella Felder
(1861), jethys Bdv. (1836) and mita Reak (1866). I feel that
these represent but one variable species. I have a long series of
specimens from various tropical stations and among them are all
intergrades from jethys to citronella to mita. Whether these are
in turn only varieties of melite Linnaeus or not I am not sure.
The most heavily marked specimen should be referred to f. jethys
Bdv., those without the dark bar along the inner margin of the
forewing to f. citronella Felder, and those with further restric-
tion of the dark apical markings to f . mita.

The specimens noted above are of f. citronella. I suspect that
the degree to which the dark markings are developed is related
to the season or moisture, the darkly marked individual being
1 1 wet season ’ ’ the lightly marked, 1 1 dry season. ’ ’

63. Enantia marion, Godman & Salvin.

G. & S., 2 : 184.

R., p. 100.

H. , p. 655.

El Sabino, Uruapan, Michoacan, 1 vii. 15-30.36
(H.D.T.).

Pierinas

(Euchloini)

72. Hesperocharis costaricensis Bates.

G. & S., 2 : 126, pi. 60, f. 3, 4.

R., p. 79, pi. 23f.

H, p. 656.

Rio Blanco, Yera Cruz, 2200 ft., 1 J1 v.10.41 (R.P.).

El Sabino, Uruapan, Michoacan, 1 2 vii. 15-30.36
(H.D.T.).

102

Journal New York Entomological Society

[Vol. LI1

Apparently this is the first record of the species from the east
coast of Mexico. Hoffmann, p. 656, reports the species from the
west coast of Guerrero, Michoacan, Jalisco and Colima.

(Rhodocerini)

74. Colias eurytheme Boisduval.

G. & S., 2 : 151 (as chrysotheme) .

R., p. 93, pi. 271

H. , p. 656.

Hda. Yista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 2 ?5 vi.17.40 (H.H.).

Galeana, Nuevo Leon, 6500 ft., 1 J1 vii.30.39 (H.H.).
Tancitaro, Michoacan, 6600 ft., 1 £ viii.11.40 (H.H.).
These few specimens do not indicate the abundance of the
species in Mexico. It is probable that the species was ignored in
most localities. The Nuevo Leon material is f. amphidusa Bdv.,
the other is typical of f . eurytheme .

77. Zerene caesonia Stoll.

G. & S., 2 : 153, 727 (as cesonia).

R., p. 94, pi. 261

H. , p. 656.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 4 1 $ vi.16-18.40 (H.H.).

Arroyo del Meco, Tamaulipas, 1320 ft., 1 $ iv.28.41

(R.P.).

Jacala, Hidalgo, 4500 ft., 7 J'J1 1 J vi.25-vii.l.39 (H.H.).
El Banito, Valles, San Luis Potosi, 200 ft., 1 vi. 28.40
(H.H.).

Tancitaro, Michoacan, 6600 ft., 1 J vii.30.40 (H.H.).
Apatzingan, Michoacan, 1050 ft., 1 vii.5.40 (H.H.).

Rio Balsas, Guerrero, 2400 ft., 1 5 v.26.41 (R.P.).
Chilpancingo, Guerrero, 1 $ v.26.41 (R.P.).

The specimens are all in a condition indicating recent emer-
gence.

78. Anteos clorinde Godart.

G. & S., 2 : 148, 726.

R., p. 89, pi. 24g.

Mar., 1944]

Brown: Pierid^:

103

H., p. 656.

Victoria, Tamaulipas, 1 J1 vi.25.35 (H. A. Freeman).

El Pujal, San Luis Potosi, 100 ft., 16 J'J' vii. 17.39
(H.H.).

Arroyo del Calabezas, San Luis Potosi, 250 ft., 2 J'J'

iv. 30.41 (R.P.).

Hda. Potrero Viejo, nr. Parare Nuevo, Vera Cruz., 1500
ft., 1 J1 v.5.41 (R.P.).

Rio Blanco, Vera Cruz, 2200 ft., 1 v.10.41 (R.P.).

Jacala, Hidalgo, 4500 ft., 3 $$ vi.2^-vii.7.39 (H.H.).
Apatzingan, Michoacan, 1 5 viii.2.40 (H.H.).

The Apatzingan 5 is badly rubbed, but appears to lack the
usual large yellow area on the forewing.

79. Anteos maerula Fabricius.

G. & S., 2 : 149, 726.

R., p. 89, pi. 24g.

H. , p. 656.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 2 $$ vi.17-18.40 (H.H.).

Victoria, Tamaulipas, 1 vi.25.35 (H. A. Freeman).

60 mi. S. of Victoria, Tamaulipas, 1 J' vii. 6. 36 (H.D.T.).
El Banito, Valles, San Luis Potosi, 200 ft., 4 J'df 2 55
vi. 26.40 (H.H.).

El Pujal, San Luis Potosi, 100 ft., 12 J'J' vii.17.39
(H.H.).

El Sol, Tamazunchale, San Luis Potosi, 150 ft., 1^15

v. 29.41 (R.P.).

Jacala, Hidalgo, 4500 ft., 3 vii.1-10.39 (H.H.).

Chichen Itza, Yucatan, 1 g iv.7.36 (F.M.B.).

The females from El Banito are f. gueneeana Bdv., the one
from El Sol is the nymotypical form.

80. Phoebis sennae Linnaeus.

G. & S., 2 : 141 (as eubule).

R., p. 85.

H. , p. 657.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 3 <?<? 1 5 vi. 16-21.40 (H.H.).

104

Journal New York Entomological Society

[Vol. LIl

Hda. Santa Engracia, Tamaulipas, 2 J'J' vii.27.39
(H.H.).

El Banito, Valles, San Luis Potosi, 400 ft., 2 J'J' 1 5
vi. 26-28.40 (H.H.).

El Sol, Tamazunchale, San Lnis Potosi, 400 ft., 1^1$
v.29.41 (R.P.).

Jacala, Hidalgo, 4500 ft., 3 vi.29.39 (H.H.).

Chichen Itza, Yucatan, 1 § iv.8.36 (P.M.B.) ; 1 §
viii.30.36 (H.D.T.).

Sta. Lucrecia, 1 $ ix. 24.39 (H.D.T.).

Apatzingan, Michoacan, 1200 ft., 2 1 5 viii. 3-5.40

(H.H.).

El Sabino, Uruapan, Michoacan, 1 ^ 1 J vii.15-30.36
(H.D.T.).

Tancitaro, Michoacan, 4500 ft. 1 <£ viii.11.41 (H.H.).

The males from Nuevo Leon are fairly typical of race eubule L.
the others tend toward marcellina Cr., but none of them is as
strongly marked as the typical males from tropical South
America. It is a pity that no one bothers to collect this common
species in sufficient numbers to settle the question of what race
the Mexican material represents. I would designate all of the
above males eubule f. drya Fabricius. I fully realize that else-
where (Amer. Mus. Nov. 368, p. 7, 1929) I stated that Mexican
material was referable to marcellina Cr. I also noted (p. 8) that
the tropical race might well represent two races. The more
material that I see from Mexico the more I feel that these state-
ments need revising, but I have not the material to do so. It is
quite probable that in Mexico the two races so intergrade that we
have a continuous cline from eubule to marcellina.

The El Banito female is f . pallida Cockerell, the El Sol, yamana
Reakirt. The Hda. Vista Hermosa female is a good marcellina
Cr. The Apatzingan female is badly rubbed but is probably
f. yamana. F.M.B.’s from Chichen Itza is pallida while H.D.T.’s
is marcellina.

81. Phoebis philea Linnaeus.

G. & S., 2 : 140, 726.

R., p. 86, pi. 25c.

H. , p. 657.

Mar., 1944]

Brown: Pierim:

105

El Pujal, San Luis P.otosi, 100 ft., 2 vii.17.39 (H.H.).

(H.H.).

Apatzingan, Michoacan, 1200 ft., 1 viii.5.40 (H.H.).

83. Phoebis agarithe maxima Neumoegen.

G. & S., 2: 145, 726 (as agarithe).

R., p. 86 (as argante agaritha).

H. , p .657.

Hda. Vista Hermosa, Nuevo Leon, 1500 ft., 6 3 22

vi.16-21.40 (H.H.).

Sabinas Hidalgo, Nuevo Leon, 960 ft., 2 2

vi. 15-18. 39 (H.H.).

Arroyo del Meco, Tamaulipas, 1320 ft., 2 iv.28.41

(R.P.).

Hda. Santa Engracia, Tamaulipas, 2 J'J' vii.25.39
(H.H.).

Victoria, Tamaulipas, 1 2 vi.28.35 (H. A. Freeman).

60 mi. S. Victoria, Tamaulipas, 1 2 vii.6.36 (H.D.T.).

El Pujal, San Luis Potosi, 100 ft., 3 vii.20-21.39

(H.H.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 £
v.29.41 (R.P.).

Tamazunchale, San Luis Potosi, 500 ft., 1 £ vi.30.40
(H.H.).

Arroyo del Calabezas, San Luis Potosi, 250 ft., 1
iv.30.41 (R.P.).

f vii.22.39

El Banito, San Luis Potosi, 200 ft., 3 J'J' 1 $ < ^2^40

l (H.H.).

El Mante, San Luis Potosi, 1 iv.29.41 (R.P.).

Orizaba, Vera Cruz, 2000 ft., 1 J* v.6.41 (R.P.).

These are all good maxima except the El Sol male which ap-
proaches agarithe. The northern specimens are absolutely typ-
ical, those from the south show some of the brownish markings
on the underside. All the females are f . albarithe Brown.

83c. Phoebis agarithe agarithe Boisduval.

Progresso, Yucatan, 1 2 viii.27.36 (H.D.T.).

Chichen Itza, Yucatan, 1 2 viii.30.36 (H.D.T.).

106

Journal New York Entomological Society

[Vol. LII

These two females are much more like typical South American
agarithe than maxima and I have so designated them. They are
not antiilia Brown, which might be expected on the peninsula.
The Progresso specimen is the yellow form, the Chichen Itza one
the white form. Hoffmann has not included this race in his
check-list.

84. Phoebis intermedia Butler.

G. & S., 2: 143, 726 (as rurina).

R., p. 86.

H. , p. 657.

Tancitaro, Michoacan, 4500 ft., 1 vii. 25.40 (H.H.).

86. Aphrissa statira jada Butler.

G. & S., 2 : 147 (as statira).

R., p. 87 (as statira).

H. , p. 657.

El Banito, Valles, San Luis Potosi, 200 ft., 1 J vi.26.40
(H.H.).

This is a pale lemon yellow female like those of true statira.

87. Kricogonia lyside Godart.

G. & S., 2: 150, 151, 726 (as unicolor).

R., p. 89.

H. , p. 658.

Sabinas Hidalgo, Nuevo Leon, 900 ft., 9 10

vi.15-18.39 (H.H.).

Ojo de Agua, Sabinas Hidalgo, N. L., 1000 ft., 2
vi.14.40 (H.H.).

Villa Santiago, Nuevo Leon, 1500 ft., 2 viii.8.39.

(H.H.).

35 km. W. of Linares, Nuevo Leon, 1 J viii.7.39. (H.H.).
Galeana, Nuevo Leon, 2 J'J' 2 22 vii. 28-viii. 4.39. (H.H.).
Hda. Sta. Engracia, Tamaulipas, 1^1} vii.25-27.39
(H.H.).

El Banito, Valles, San Luis Potosi, 1 § vii.22.39 (H.H.).
nr. Apatzingan, Michoacan, 500 ft., 3 J'.J' 3 22 viii.3.40
(H.H.).

Apatzingan, Michoacan, 1050 ft., 1 $ 2 viii.2-5.40

(H.H.).

Mar., 1944]

Brown: Pierid^e

107

El Sabino, Uruapan, Michoacan, 2 1 J yii.15-30.36

(H.D.T.).

Acahuato, Michoacan, 3000 ft., 1 $ viii.2.40 (H.H.).
Tancitaro, Michoacan, 6600 ft., 1 £ vii.20.40 (H.H.).

z

p 2 cs

It is rather strange that Potts took no specimens of this common
species. The above series contains examples of all of the named
forms that occur in Mexico.

typical $ terrissa $ unicolor $ fantasia

Nuevo Leon 21 5 4 1?

Tamaulipas 1 1

San Luis Potosi 1

Michoacan 7 3 1 4

The specimens noted as fantasia are not typical; they all lack
the costal yellow streak and the ground color of the wings tends
to be greenish rather than yellow. The size of the specimens
varies greatly. This is best shown in the text figure, based upon
the baso-apical radius of the forewings, and the following table
of data.

N. Mean S.D. Critical limits Range

Total

series ... 45 21.37 ± 0.57 mm. 3.66 mm. 14.1 mm., 28.7 mm. 16-28 mm.

Sabinas

Hidalgo 19 22.90 ± 0.34 mm. 3.27 mm. 16.5 mm., 29.4 mm. 16-28 mm.

108

Journal New York Entomological Society

[Vol. LII

88. Eurema albula Cramer.

G. & S., 2: 166.

R., p. 84.

H. p. 658.

K. (4), p. 121.

El Sol, Tamazunchale, San Lnis Potosi, 400 ft., 11
iv.30, v.29.41 (R.P.).

Ojo de Agua, Yera Cruz, 1600 ft., 4 v.12.41 (R.P.).

El Sabino, Urnapan, Michoacan, 1 vii. 15-30. 36

(H.D.T.).

Two of the Ojo de Agua specimens are intermediate to albula
and f. tapeina Bates.

89. Eurema jucunda Boisduval.

G. & S., 2 : 168, pi. 64, f. 9-12.

R., p. 83, pi. 24e.

H. , p. 658.

K., p. 124.

nr. Acapulco, Guerrero, 100 ft., 1 2 v.26.41 (R.P.).
nr. Chilpancingo, Guerrero, 4 jy1 3 2$ v.26.41 (R.P.).
Rio Balsas, Guerrero, 2400 ft., 1 2 v.26.41 (R.P.).

El Sabino, Uruapan, Michoacan, 10 J'J1 9 22 vii.15-30.36
(H.D.T.).

Apatzingan, Michoacan, 3000 ft., 1 $ viii.2.40 (H.H.).
Tancitaro, Michoacan, 6600 ft., 1 y 1 2 vii. 25, viii.11.40
(H.H.).

All the males are of f . sidonia Felder. This is also true of the
females except for two from Chilpancingo which tend toward the
typical in respect to the margin of the hindwings. The general
color of the females varies from white to pale yellow. One of the
Chilpancingo specimens and the one from Rio Balsas have white
forewings and yellow hindwings.

89a. Eurema jucunda lydia Felder.

G. &S., 2:170, pi. 63, f. 20-23.

R., p. 83 (as delia lydia).

H. , p. 658.

K., p. 126 (as palmyra lydia).

Jacala, Hidalgo, 4500 ft., 3 jy 2 2? vi.25-vii.2.39
(H.H.).

Mar., 1944]

Brown: Pierhx®

109

El Pujal, San Lnis Potosi, 100 ft., 5 2 vii.17-

21.39 (H.H.).

Tuxpango, Vera Crnz, 1500 ft., 1 c? 2 5? v-9.41 (R.P.).

Hda. Potrero Viejo, Vera Cruz, 1500 ft., 1 J' 1 J v.5.41
(R.P.).

Orizaba, Vera Cruz, 2000 ft., 4 1 J v.6.41 (R.P.).

Rio Blanco, Vera Cruz, 2200 ft., 2 1 2 v.10.41 (R.P.).

This series of “barred” Euremas from the east coast is quite
variable. All have the pearly white under surfaces characteristic
of jucunda. The females are fairly constant, having a reduced
grey bar on the inner margin of the forewing and an incomplete
black margin on the hindwing. One Jacala specimen is faintly
flushed with yellow on the forewing and the Rio Blanco female
bears this flush on the hindwing. The males are extremely varied
but I hesitate to refer the specimen to anything other than lydia.
The marginal band of black on the hindwings is broad and com-
plete on two (Rio Blanco and Orizaba), narrower but complete
on two (El Pujal) and interrupted on the rest. In no case is the
margin reduced to an apical spot. The hindwings range from
yellow (J1 Jacala) to white with many specimens just faintly
tinged with yellow. Until a lot of life history work is done and
enormous series taken over long periods of time for many locali-
ties we will not be able to unravel the puzzle of the “barred”
Euremas.

91. Eurema boisduvaliana Felder.

G. &. S., 2 : 159, pi. 63, f. 1-4.

R., p. 81 (as mexicana, in part).

H. , p. 658.

K, p. 133.

Galeana, Nuevo Leon, 6500 ft., 1 £ 10.29.41 (R.P.).

Jacala, Hidalgo, 4500 ft., 4 1 $?, vi.29-vii.5.39

(H.H.).

El Pujal, San Luis Potosi, 100 ft., 11 vii.12-21.39

(H.H.).

Arroyo del Calabezas, San Luis Potosi, 1 J' iv.30.41
(R.P.).

El Banito, Valles, San Luis Potosi, 1^1$ vi. 26-27. 40
(H.H.).

110

Journal New York Entomological Society

[Yol. LII

El Sol, Tamazunchale, San Lnis Potosi, 2 v.29.41

(R.P.).

Fortin, Yera Cruz, 1600 ft., 2 J'J' v.4.41 (R.P.).
Orizaba, Vera Cruz, 2000 ft., 1 v.4.41 (R.P.).

Rio Blanco, Yera Cruz, 2200 ft., 8 J'J' v.10.41 (R.P.).
Chichen Itza, Yucatan, 1 <£ viii.30.36 (H.D.T.).

El Sabino, Uruapan, Michoacan, 1 §?, vii.15-30.36
(H.D.T.).

Apatzingan, Michoacan, 3200 ft., 1 5 viii.2.40 (H.H.).
The questioned female from Jacala is aberrant. The apical
patch on the hindwings is reduced to a few scales along the
nervule. The same is true of the El Sabino female.

92. Eurema xanthochlora Kollar.

G. & S., 2 : 161, 727, pi. 63, f. 5-8.

R., p. 81, pi. 24b.

H. , p. 658.

K., p. 134.

El Banito, Yalles, San Luis Potosi, 200 ft., 1 $ vi.28.40
(H.H.).

Tuxpango, Yera Cruz, 1500 ft., 1 £ v.9.41 (R.P.).

93. Eurema mexicana mexicana Boisduval.

G. & S., 2 : 157.

R., p. 81, pi. 24a.

H. , p. 659.

K, p. 134.

Hda. Yista Hermosa, Villa Santiago, Nuevo Leon, 1500-
3000 ft., 6^1$ vi.17-21.40 (H.H.).

Villa Santiago, Nuevo Leon, 1500 ft., 1 J' viii.8.39
(H.H.).

Jacala, Hidalgo, 4500 ft., 37 J'J' 1 5 vi.23-vii.2.39
(H.H.).

El Pujal, San Luis Potosi, 100 ft., 1 <$ vii.17.39 (H.H.).
El Sol, Tamazunchale, 400 ft., 8 J'.J' v.28-29.41 (R.P.) ;
1$ iv.31.41 (R.P.).

Tuxpango, Yera Cruz, 1500 ft., 1 J v.8.41 (R.P.).

Rio Blanco, Yera Cruz, 2200 ft., 5 v.10.41 (R.P.).

Cumbres (km. 295), Yera Cruz, 6000 ft., 1 v.7.41

(R.P.).

Mar., 1944]

Brown: Pierid^e

111

El Sabino, Uruapan, Michoacan, 3 J'J' vii.15-30.36
(H.D.T.).

Tancitaro, Michoacan, 6600 ft., 1 $ viii.14.40 (H.H.).

Cerro Tancitaro, Michoacan, 7000 ft., 2 vii.20.40

(H.H.).

All these specimens are typical mexicana. Hoffmann includes
in his listing of the species both ab. recta Klots and ab. bieder-
manni Ehrmann with question marks. I have a distinct feeling
that these two “aberrations’7 describe the material from the
northern reaches of the race such as the mountains of Colorado
and should be raised to racial status. Ehrmann’s name is avail-
able for this. There is some variation in size among the speci-
mens before me. A male from Villa Santiago measures only 15
mm. along the baso-apical radius of the forewing while the largest
specimen from El Sol measures 27 mm. The black margin of the
hindwing is variable; on some specimens it is a small patch an-
terior to M2 while in others it extends to Cu2. On these latter
it is broadest at the M2-M3 interspaces and has a basad spur in
Ms— Cul

94. Eurema salome Felder.

G. & S., 2 : 160 (as fabiola).

R., p. 81.

H. , p. 659.

K, p. 136.

Jacala, Hidalgo, 4500 ft., 16 vi.23-vii.2.39 (H.H.).

Fortin, Vera Cruz, 1600 ft., 1 $ v.3.41 (R.P.).

These are all f. limoneus Felder. I disagree with Hoffmann
and agree with Klots on the status of this name. I feel that the
species is splitting into two races and that the northern material
from Mexico is sufficiently different from Ecuadorian salome to
require a name in this genus.

95. Pyristia gundlachia Poey.

G. & S., 2 : 156.

R., p. 81, pi. 21a.

H. , p. 659.

K, p. 137.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft.,2$2vi.l7-22.40 (H.H.).

112

Journal New York Entomological Society

[Vol. LII

96. Pyristia proterpia Fabricius.

G. & S., 2: 155.

R., p. 81, pi. 24a.

H. , p. 659.

K., p. 137.

nr. Villagran, Tamaulipas, 1160 ft., 1 J' iv.28.41 (R.P.).
C. Victoria, Tamaulipas, 1 g vi.19.39 (H.H.).

Hda. Sta. Engracia, Tamaulipas, 1 vii.27.39 (H.H.).

El Pujal, San Luis Potosi, 100 ft., 3 1 5 vii. 17-20.39

(H.H.).

Rio Blanco, Vera Cruz, 2200 ft., 2 J'J' v.10.41 (R.P.).
Chichen Itza, Yucatan, 1 g viii.30.36 (H.D.T.).
Acapulco, Guerrero, 100 ft., 1 v.26.41 (R.P.).

Apatzingan, Michoacan, 1050 ft., 5 viii.2-5.40

(H.H.).

El Sabino, Uruapan, Michoacan, 10 J'J' 4 5? vii. 15-30. 36
(H.D.T.).

Tancitaro, Michoacan, 7000 ft., 1^1} vi.25-30.40
(H.H.).

On the northernmost specimen (Villagran) the black markings
are greatly reduced. The Chichen Itza specimen approaches
watsoni from the similar arid area in Ecuador !

97. Pyristia dina westwoodi Boisduval.

G. & S., 2 : 163 (as dina).

R., p. 82, pi. 24b.

H. , p. 659.

K., p. 139.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 7 J'J'
4 22 iv. 30-31, v.28-29.41 (R.P.).

Arroyo del Calabezas, San Luis Potosi, 250 ft., 1 $
iv.30.41 (R.P.).

Chichen Itza, Yucatan, 1 £ viii.30.36 (H.D.T.) ; 1 2
iv.7.36 (F.M.B.).

N. of Iguala, Guerrero, 1 $ v.22.41 (R.P.).

All the females are much lighter than the males and in many
respects, other than size, compare favorably with f . 2 citrina
Poey from the West Indies. The West Coast male is the exact
counterpart of race parvumbra Kaye from Jamaica!

Mar., 1944]

Brown: Pierid^e

113

98. Pyristia lisa euterpe Menetries.

G. & S., 2 : 162.

R., p. 83.

H. , p. 659.

K., p. 138.

Sabinas Hidalgo, Nuevo Leon, 960 ft., 1 £ vi.15.39
(H.H.).

nr. Villagran, Tamanlipas, 1150 ft., 3 iv.28.41

(R.P.).

60 mi. So. of Victoria, Tamaulipas, 2 vii.6.36

(H.D.T.).

Jacala, Hidalgo, 4500 ft., 1 vii.6.39 (H.H.).

El Pujal, San Luis Potosi, 100 ft., 1 5 vi.20.39 (H.H.).
Rio Blanco, Vera Cruz, 2200 ft., 1 v.10.41 (R.P.).

The female is f. centralia Herrick-Schaffer {alba Strecker).

99. Pyristia nise perimede Prittwitz.

G. &S., 2:165, 727 (as tenella).

R., p. 83, pi. 24d (as nise).

H. , p. 659.

K, p. 140.

Sabinas Hidalgo, Nuevo Leon, 960 ft., 1 vi.31.39

(H.H.).

nr. Villagran, Tamaulipas, 1160 ft., 2 J'J' iv.28.41 (R.P.).
C. Victoria, Tamaulipas, 1 J vi.19.39 (H.H.).

60 mi. So. of Victoria, Tamaulipas, 1 $ vii.6.36 (H.D.T.).
Hda. Sta. Engracia, Tamaulipas, 1 J' vii.25.39 (H.H.).
Jacala, Hidalgo, 4500 ft., 5 2 5? vi.23-vii.2.39

(H.H.).

El Pujal, San Luis Potosi, 100 ft., 3 vii.18-21.39

(H.H.).

Arroyo del Calabezas, San Luis Potosi, 250 ft., 1 J*

iv. 30.41 (R.P.). z

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1^1$

v. 28-29.41 (R.P.).

El Banito, Valles, San Luis Potosi, 150 ft., 5 2 5?

vi. 26.40, vii.22.39 (H.H.) ; iv.30.41 (R.P.).

Hda. Potrero Viejo, Paraje Nuevo, Vera Cruz, 1500 ft.,
3 v.5.41 (R.P.).

114

Journal New York Entomological Society

[Vol. LII

Orizaba, Vera Cruz, 2000 ft., 2 v.6.41 (R.P.).

Rio Blanco, Vera Cruz, 2200 ft., 8 v.10.41 (R.P.).

Ojo de Agua, Vera Cruz, 1600 ft., 1 5 v.12.41 (R.P.).
Acapulco, Guerrero, 100 ft., 1 J' v.23-25.41 (R.P.).

Sta. Lucrecia, 1 ix.24.39 (H.D.T.).

Apatzingan, Michoacan, 500 ft., 1 <$ viii.3.40 (H.H.).

El Sabino, Uruapan, Michoacan, 5 2 vii.15-30.36

(H.D.T.).

There is a great deal of variation in this series, none of it re-
lated to geographic distribution. The ground color varies from
pale to deep lemon. The dark margin on the hindwings varies
from complete to absent ; on over half of the specimens it is rep-
resented by only a row of black dots on the nervules. There is
some variation in the extent of the black marking on the fore-
wing. On some the underside is boldly marked; on others it is
almost immaculate.

100. Abaeis nicippe Cramer.

G. & S, 2 : 155, 727.

R., p. 81, pi. 24a.

K., p. 132.

H. , p. 659.

Sabinas Hidalgo, Nuevo Leon, 960 ft., 5 J'J' vi.15-16.39
(H.H.).

Ojo de Agua, Sabinas Hidalgo, Nuevo Leon, 1300 ft.,
1 vi.14.40 (H.H.).

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 1 <$ 2 ?? vi.16-21.40 (H.H.).

Galeana, Nuevo Leon, 6500 ft., 2 jy1 vii.30-viii.3.39
(H.H.).

Jacala, Hidalgo, 4500 ft., 11 J'J' vi.23-vii.2.39 (H.H.).
El Pujal, San Luis Potosi, 100 ft., 1 J1 n.d. (H.H.).
Hda. Potrero Viejo, nr. Paraje Nuevo, Vera Cruz, 1500
ft., 1 $ vi.5.41 (R.P.).

Monte Alban, Oaxaca, 6000 ft., 1 y v.18.41 (R.P.).

The Monte Alban specimen is notable for the reduction of the
dark apical markings on it.

101. Nathalis iole Boisduval.

G. & S., 2 : 172, pi. 64, ff. 15-18.

Mar., 1944]

Brown: Pierid^

115

R., p. 95, pi. 27h (as jole).

H., p. 659.

bet. Adjuntas & Sabinas Hidalgo, Nuevo Leon, 3000
ft., 1 c? vi.21.40 (H.H.).

Sabinas Hidalgo, Nuevo Leon, 1500 ft., 1 1 J vi.17.40

(H.H.).

Ojo de Agua, Sabinas Hidalgo, Nuevo Leon, 1000 ft.,
1 g vi. 15.40 (H.H.).

Galeana, Nuevo Leon, 6500 ft., 1 vii. 30.39 (H.H.).

nr. Villagran, Tamaulipas, 1160 ft., 1 iv.28.41 (R.P.).

C. Victoria, Tamaulipas, 1 § vi.19.39 (H.H.).

60 mi. So. of Victoria, Tamaulipas, 1 2 vii.6.36
(H.D.T.).

Jacala, Hidalgo, 4500 ft., 10 1 5 vi.23-vii.l.39.

(H.H.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 $
v.29.41 (R.P.).

El Banito, Valles, San Luis Potosi, 150 ft., 2 ££ iv.30.41
(R.P.).

El Pujal, San Luis Potosi, 100 ft., 1^1} vii.l7.39J
(H.H.).

Tuxpango, Vera Cruz, 1500 ft., 2 ££ v.9.41 (R.P.).

Orizaba, Vera Cruz, 2000 ft., 1 g 1 2 v.6.41 (R.P.).

Cumbres (km. 295), Vera Cruz, 8000 ft., 3 ££ v.7.41
(R.P.).

Monte Alban, Oaxaca, 6000 ft., 1 2 v.18.41 (R.P.).

The females from all localities other than Jacala and Monte
Alban are f. irene Pitch. One of the Jalaca males has the fore-
wing bar reduced to a submarginal spot.

(Pierini)

103. Neophasia terlootii Behr.

R., p. 55, pi. 18a.

H., p. 660.

Cerro Tancitaro, Michoacan, 5500 ft., 1 $ viii.17.40
(H.H.).

This capture of Hoogstraal’s extends the range of this species-
over 300 miles southward. I suspect that it will be recovered still

116

Journal New York Entomological Society

[Vol. LII

further south, possibly in Central America in the pine-oak forests
of Guatemala and Honduras.

104. Catasticta flisa Herrick-Schaffer.

G. & S., 2 : 117.

R., p. 73, pi. 22e.

H. , p. 660.

Jaeala, Hidalgo, 4500 ft., 1 vi.29.39 (H.H.).

This extends the range of the species northward from Vera
Cruz (see H., p. 660).

105. Catasticta nimbice nimbice Boisduval.

G. & S., 2 : 118.

R., p. 70, pi. 22b.

H. , p. 660.

Jaeala, Hidalgo, 4500 ft., 2 vi.2L-28.39 (H.H.).

Tancitaro, Michoacan, 6600 ft., 1 J vii.30.40 (H.H.).
The female is freshly emerged, the males are worn.

110. Appius ilaire Godart.

G. & S., 2 : 135 (as margarita).

R., p. 59, pi. 19c.

H. , p. 661.

El Banito, Valles, San Luis Potosi, 200 ft., 1 £ vi. 28.40
(H.H.).

113. Pieris protodice Boisduval & Le Conte.

G. & S., 2 : 130, 724.

R., p. 59.

H. , p. 661.

Sabinas Hidalgo, Nuevo Leon, 960 ft., 1 5 vi.15.39
(H.H.).

Hda. Vista Hermosa, nr. Villa Santiago, Nuevo Leon,
1600 ft., 1 3 vi.16.40 (H.H.).

Galeana, Nuevo Leon, 6500 ft., 1 J viii.1.39 (H.H.).
Monterrey, Nuevo Leon, 1600 ft., 1 § iv.27.41 (R.P.).
nr. Villagran, Tamaulipas, 1160 ft., 1^1} iv.28.41
(R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 2 J'J' vi.28.40
(H.H.).

Mar., 1944]

Brown: Pierid^e

117

114. Pieris rapae Linnaeus.

R., p. 58, pi. 19b.

H., p. 661.

Galeana, Nuevo Leon, 6500 ft., 2 <£<§ viii.4.39 (H.H.).
This species seems to have reached Mexico since the publication
of the Biologia (since 1900?).

116. Leptophobia aripa Boisduval.

G. & S., 2 : 136, 726 (as elodia).

R., p. 60, pi. 19d.

H. , p. 661.

nr. Villagran, Tamaulipas, 1160 ft., 1 5 iv.28.41 (R.P.).
Jacala, Hidalgo, 4500 ft., 2 J'J' vi.28.39 (H.H.).

Fortin, Vera Cruz, 1600 ft., 1 J1 v.3.41 (R.P.).

Ojo de Agua, Vera Cruz, 1600 ft., 1 v.12.41 (R.P.).

Orizaba, Vera Cruz, 200 ft., 1 £ v.6.41 (R.P.).

117. Itaballia demophile calydonia Boisduval.

G. & S., 2 : 131.

R., p. 63.

H. , p. 661.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 £
v.29.41 (R.P.).

The subapical cross bar on the upper sides of the forewings is
totally lacking, on the underside it is reduced to little more than
a broad line. This specimen extends the range northward from
southern Vera Cruz (see H., p. 661).

119. Pieriballia viardi viardi Boisduval.

G. & S., 2 : 133, 725, pi. 62, f. 1-4.

R., p. 62.

H. , p. 661.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 $
iv.31.41 (R.P.).

El Pujal, San Luis Potosi, 100 ft., 1 J* 1 J vii.20-21.39
(H.H.).

121. Ascia monuste cleomes Boisduval.

G. & S., 2 : 132, 725 (as monuste).

R., p. 57.

H. , p. 662.

118

Journal New York Entomological Society

[Vol. LII

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1600
ft., 1 ? vi.17.40 (H.H.).

Galeana, Nuevo Leon, 6500 ft., 1 5 iv.29.41 (R.P.).

nr. Villagran, Tamaulipas, 1160 ft., 1 § iv.28.41 (R.P.).

Arroyo del Meco, Tamaulipas, 1 y iv.28.41 (R.P.).

C. Victoria, Tamaulipas, 2 §f vi.27.35 (H. A. Free-
man) ; 2 yy vi.19.39 (H.H.).

60 mi. So. of Victoria, Tamaulipas, 3 yy vii.6.36
(H.D.T.).

Jacala, Hidalgo, 5400 ft., 1 y 1 ? vii.1.39 (H.H.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 2 yy

v. 29.41 (R.P.).

Arroyo del Calabezas, San Luis Potosi, 250 ft., 1 y
10.30.41 (R.P.).

El Baiiito, Valles, San Luis Potosi, 200 ft., 2 yy 2

vi. 26.40, vii.19.39 (H.H.).

El Pujal, San Luis Potosi, 100 ft., 3 yy vii. 17-19. 39
(H.H.).

Tuxpango, Vera Cruz, 1500 ft., 1 y v.9.41 (R.P.).

Hda. Potrero Viejo, Paraje Nuevo, Vera Cruz, 1500 ft.,
1 ? v.5.41 (R.P.).

Rio Blanco, Vera Cruz, 2200 ft., 1 y v.10.41 (R.P.).

Apatzingan, Michoacan, 1200 ft., 2 yy 1 J viii.5.40
(H.H.).

The males are rather uniform throughout. The marginal dark
spots on the hindwing are variable, being represented by either
a few scales or a small triangular patch. This does not seem to
be related to the origin of the specimens. One male from 60
miles South of Victoria is almost immaculate on both upper and
lower surfaces. The females fall into two clearly defined groups.
Those from Nuevo Leon, Tamaulipas and Michoacan have a pink-
ish iridescence on the upper surface and one (from Victoria)
approaches phileta Fabricius. Those from San Luis Potosi,
Hidalgo and Vera Cruz are a dull buff -yellow on the hindwings
and the same with a slightly pinkish tone on the forewings.

123. Melete isandra Boisduval.

G. & S., 2 : 139, 726, pi. 61, ff. 13, 14.

R, p. 77, pi. 23b.

Mar., 1944]

Brown: Pierid^

119

H., p. 662.

El Banito, Valles, San Lnis Potosi, 200 ft., 1 $ 1 J
vii.19.39 (H.H.).

El Pnjal, San Lnis Potosi, 100 ft., 3 yii. 17-20.39
(H.H.).

BIBLIOGRAPHY

(1) . Godman & Salvin, Biologia Centrali-Americana, Rhopalocera, 2: 113-

188. 1889-1890.

(2) . Roeber, in Seitz’ Macrolepidoptera of the World, vol. 5, pp. 53-111.

1909.

(3) . Hoffmann, Anales del Instituto de Biologia, 11: 639-739. 1940.

(4) . Klots, Entomologica Americana, 9: 99-171. 1929.

No. 2

Vol. LII

JUNE, 1944

Journal

of the

New York Entomological Society

Devoted to Entomology in General

Edited by HARRY B. WEISS

Publication Committee

HARRY B. WEISS JOHN D. SHERMAN, Jr.

T. C. SCHNEIRLA

Subscription $3.00 per Year

Published Quarterly by the Society
N. QUEEN ST. AND McGOVERN AVE.

LANCASTER, PA.

NEW YORK, N. Y.

1944

CONTENTS

A Generic Review of the Subfamily Phyllobaeinae

By A. B. Wolcott 121

The Reproductive Functions of the Army-Ant Queen as
Pace-Makers of the Group Behavior Pattern

By T. C. Schneirla 153

A Re-Survey of Papaipema Sm. (Lepidoptera)

By Henry Bird 193

Insect Introductions and War

By William M. Boyd 200

Catalogue of North American Psychodidse

By William F. Rapp, Jr 201

Recent Work by the International Commission on Zoologi-
cal Nomenclature

By Francis Hemming 211

NOTICE: Volume LII, Number 1, of the Journal of the
New York Entomological Society was published on
April 6, 1944.

Entered as second class matter July 7, 1925, at the post office at Lancaster, I’a..
under the Act of August 24, 1912.

Acceptance for mailing at special rate of postage provided for in Section 1103.
Act of October 3, 1917, authorized March 27, 1924.

JOURNAL

OF THE

New York Entomological Society

Vol. LII June, 1944 No. 2

A GENERIC REVIEW OF THE SUBFAMILY PHYLLO-
BiENINiE (OLIM HYDNOCERIN^E) (COL.)

By A. B. Wolcott
Downers Grove, III.

It has become increasingly evident that the genera of this sub-
family are in a very chaotic condition, and that they are neither
clearly defined nor readily understood. This state of confused
conditions has been produced by various causes, such as the inclu-
sion within the group of extraneous elements, which should have
been assigned elsewhere ; the omission of certain genera ; disre-
gard of the rules of priority and nomenclature; the all too fre-
quent totally inadequate diagnoses and the lack of an ample and
accurate synonymy, all of which have been contributing factors
tending to create the utmost disorder.

In the present paper the writer will endeavor to eliminate the
alien elements, to tabulate the several genera which are compo-
nents of this subfamily, to give where needful, or at all possible
an amended diagnosis, to give an ample synonymy, to indicate
the type of each genus and record their geographic range.

Phyllob^enin.® Subfamily nom. nov.

Subfamily Characters

Cleridae; maxillary palpi with terminal segment cylindrical,
usually much shorter than that of the labial palpi, which are
large, dilated and securiform. Head with the eyes included
broader than the prothorax; antennae short; eleven-segmented
(with the exception of one or two genera), usually with a two-

122

Journal New York Entomological Society

[Vol. LIT

to five-segmented clava (rarely feebly developed). Thorax with
the sides more or less dilated at, or before the middle. Elytra
usually elongate, completely covering the abdomen, or more or
less abbreviated, elytral sculpture variable. Terminal segments
of abdomen, especially in the male, with sexual modifications.
Legs long, slender, tarsi five-segmented, basal segment very small,
atrophied or wanting, when present covered by the second seg-
ment, or elongate and distinct, not at all hidden from view from
above.

The following key to the genera, studied in connection with the
generic diagnosis which follow, will, it is believed, render possible
a ready recognition of the several genera.

KEY TO THE GENERA

3.

4.

5.

6.

8.

10.

11.

Antennae ten-segmented, clava uni-segmented Lasiocallimerus Corp.

Antennae eleven-segmented 2.

Antennal clava indistinct 18.

Antennal clava distinct; basal segment of hind tarsi small, atrophied or

absent, usually covered by the second segment 3.

Clava two-segmented 4.

Clava three- to five-segmented ......: 8.

Terminal segment of clava larger than the tenth Parmius Sharp.

Terminal segment of clava subequal in size to tenth, sometimes connate,

causing the antennae to appear ten-segmented Neohydnus Gorh.

Terminal segment of clava minute or very small 5.

Tarsal ungues with a broad basal tooth Phyllobcenus Dep**

Tarsal ungues simple or at most only slightly thickened at base 6.

Prothorax much longer than broad; third segment of antennae about

twice as long as broad, cylindrical ! Isohydnocera Chpn.

Prothorax subequilateral , , 7.

Third segment of antennae stout, only slightly longer than broad.

Emmepus Motsch.

Third segment of antennae as broad or broader than long, trapezoidal.

Wolcottia Chpn.

Clava three-segmented; basal segment of hind tarsi short 9.

Clava three- to five-segmented; basal segment of hind tarsi long, not

covered by the second segment 15.

Clava rather indistinct; protliorax one-half longer than broad.

Cephaloclerus Kuw.

Clava distinct .....: - 10.

Body apterous; eyes coarsely granulate Paupris Sharp.

Body alate; eyes finely granulate 11.

Eyes flat or depressed Abrosius Fairm.

Eyes strongly convex : 12.

June, 1944]

Wolcott: Phyllob^nin^e

123

12.

13.

14.

15.

16.

17.

18.

Eyes large, globular, entire Isolemidia Gorh.

Eyes smaller 13.

Tarsi of hind legs nonlamellate Metaxina Broun.

Tarsi of hind legs with at least two segments lamellate 14.

Clava lax, serrate I Eurycranium Blanch.

Clava rather compact; eyes feebly emarginate Lemidia Spin.

Very small (3 mm.), Phyllobcenus- like in form Theano Cast.

Much larger (5 mm. or more) : 16.

Form broad and compact; elytra never with scales; clava five-seg-
mented sub. gen. Brachycallimerus Chpn.

Form elongate or oblong; adorned with white or yellow scales 17.

Head, the eyes included, only slightly broader than the anterior part of
the prothorax sub. gen. Callimerus Gorh.

Head, the eyes included, much broader than the anterior part of the
prothorax; body depressed sub. gen. Cucujocallimerus Pic.

Elytra depressed, much broader than the prothorax at base, elongate.

Evenus Cast.

Elytra convex, elongate, only slightly broader than the prothorax at
base Allelidea Waterh.

Genus Phyllob^inus Dejean

A hundred and twenty years ago Thomas Say (Jonr. Acad.
Phila., Ill, 1823, p. 192), described an insect to which he gave
the name Clerus humeralis, a year later E. F. Germar (Ins. Spec.
Nov., I, 1824, p. 80) described the same insect (which had evi-
dently come into his possession bearing a label with the name
given to the species by Say), as he adopted for his species the
name of Clerus humeralis , thus creating a synonym of Say’s
species. Several years later Count Dejean (Cat. des Col. de la
coll., edition 3, 1837, p. 127) included under Phyllol>cenus Dejean,
humeralis Germ., which is the first and only valid species, con-
tained in the list of ten names, nine of which are given with
“Dej.” as author, but as he did not describe any of these species
they are to be considered as nomina nuda. Dejean’s citation of
humeralis Germ., according to the International Code ( vide
Article 21 and Opinion 1, B) constitutes an indication, and gives
validity to the genus Phyllobcenus Dejean, with P. ( Clerus )
humeralis Say as the genotype.

Edward Newman (1838: 379) erected the genus Hydnocera
based on serrata Newm., which is a synonym of pallipennis Say
(1825: 176). Newman two years later (1840: 362) described

124

Journal New York Entomological Society

[Vol. LII

three more species of this genus, at which time he states that
“The Count Dejean, in his ‘Catalogue des Coleop teres’ has given
to the genus the provisional name of Phyllobaenus : this I learn
by finding one of them so named by Dr. Harris.” He also re-
marks ‘ ‘ One of the species has been described by Germar, in his
‘Insectorum Species,’ under the name of Clems humeralis ; and
the same insect has also been labeled as the Tillus humeralis of
Say, but I have no reference whatever to any description by
the American entomologist.” Newman’s statements show how
deeply he was imbued with the antiquated and obsolete idea that
a generic name unaccompanied by a characterization or diag-
nosis was of no significance. And upon this supposition some
of these names have been freely and rashly employed as names
to designate other genera. An instance of the kind just men-
tioned is that of Phyllohcenus Spinola (Clerites II, 1844, p. 1),
of which he makes the following statements: “My Phyllobenes
are not those of Mr. Dejean, the latter belong to the G. [enus]
Hydnocera of Mr. Newman, a genus of which that author has
published the characters and of which he is the true founder,
the name that he has assigned to it has incontestably the right of
priority. That of Phyllohcenus having become vacant, I like it
better to take possession of it than rack one ’s brains in looking for
a new name to apply to a species that Mr. Dejean has placed in
the G. [enus] Notoxus and which cannot remain there.” The
action of the present writer in assigning Phyllobcenus to its
proper nomenclatural status leaves Phyllohcenus Spinola vacant,
and necessitates a new name in its stead, the name Phlogisto-
sternus (/Aoyiotos, crematus areppov, sternum, is here proposed.

Genotype. — Phyllobcenus dislofatus.

The writer’s thanks are due his good friend Dr. Edward A.
Chapin, Curator Division of Insects, United States National Mu-
seum, who has assisted the author in many ways; also it was he
who first called the author’s attention to the fact that Hydnocera
Newman is a synonym of Phyllobcenus Dejean.

Phyllohcenus Dejean, Cat. des Col., 3rd Edition, 1837, p. 127.

Hydnocera Newman, The Ent. Mag., Lond., V, 1837 (1838), p.
379. — Newman, Mag. Nat. Hist., (2) IV, 1840, p. 362. — Spinola,
Rev. Zool., 1841, p. 75. — Klug, Clerii, 1842, p. 311. — Spinola, Mon.

June, 1944]

Wolcott: Phyllob^enin^e

125

Clerites, II, 1844, p. 39. — Leconte, Ann. Lyc. Nat. Hist. New
York, V, 1849, p. 26. — Lacordaire, Gen. Col., IV, 1857, p. 471. —
Chenu, Encycl. d’Hist. Nat. Col., II, 1860, p. 268. — Leconte,
Classif. Col. N. Amer., I, 1861, p. 196. — Gorham, Trans. Ent. Soc.
Lond., 1877, pt. 3, p. 260. — Gorham, Biol. Centr.-Amer., Ill, pt. 2,
1883, p. 168. — Leconte and Horn, Classif. Col. N. Amer., ed. II,
1883, p. 218. — Wickham, Can. Ent., XXVII, 1895, pp. 245 and
251. — Schenkling, Gen. Ins. (Wytsman) Cler., 1903, p. 92. —
Schenkling, Col. Cat. (Jnnk) Cler., pt. 23, 1910, p. 102. — Gahan,
Ann. Mag. Nat. Hist., (8) V, 1910, p. 65. — Wolcott, Bull. Ind.
Dept. Geol. Nat. Res., I, 1910, p. 855. — Chapin, Bull. Brookl. Ent.
Soc., XII, 1917, p. 83.— Chapin, Proc. Biol. Soc. Wash., XXXV,
1922, p. 55. — Bradley, Man. Gen. Beetles Amer. N. of Mex., 1930,
p. 105. — Chagnon, Le Nat. Canad., LXII, 1935, pp. 174 and 175.

Theano Chevrolat, Ann. Soc. Ent. France, I (2), 1843, p. 33.

Diagnosis of Phyllobcenus Dejean

Head, with the eyes included, broader than the prothorax,
mandibles feebly curved with a strong internal tooth before the
apex. Head very short, front large, vertical, eyes rather large,
prominent, oval, transverse, entire or the front margin with a
feeble vestige of emargination. Antennas very short, eleven-seg-
mented, first segment thick, obconical, second very short, seg-
ments three to nine nearly cylindrical, the ninth segment as
broad as long, tenth segment large, swollen, ovate, terminated
by the scarcely discernible, minute eleventh segment. Maxillary
palpi with terminal segment short, nearly cylindrical or atten-
uate, apex truncate, terminal segment of labial palpi large, trans-
versely securiform. Prothorax usually longer than broad, sides
more or less dilated before the middle. Elytra much broader
than the prothorax at base, usually moderately elongate parallel
or narrowed near the middle, often abbreviated at apex, sculp-
ture variable. Legs long, posterior femora reaching slightly be-
yond the apex of the elytra, tarsi five-segmented, basal segment
short, the three succeeding lamellate, the posterior tarsi with the
lamella of basal segment long and compressed, lamellae of second
and third segments depressed, bifid. Ungues with a broad basal
tooth.

126

Journal New York Entomological Society

[Yol. LII

Newman in his diagnosis of the genus Hydnocera states that
the antennae are 10-articulate, but it is evident that he failed to
observe the very small and closely united true apical segment.
This error was corrected by Lacordaire (1857: 471).

Genotype. — Phyllobcenus ( Clerus ) humeralis Say. Monobasic.

Geographical range, 118 species of North, Central and South
America.

Genus Lemidia Spinola

Lemidia Spinola, Rev. Zool., 1841, p. 75. — Klug, Clerii., 1842,
p. 311. — Spinola, Clerites, II, 1844, p. 32. — Lacordaire, Gen. Col.,
IY, 1857, p. 470.— Chenu, Encycl. d’Hist. Nat. Col., II, 1860, p.
267. — Gorham, Trans. Soc. Ent. Lond., 1877, p. 249. — Schenkling,
Deutsch. Ent. Zeit., 1898, p. 169. — Schenkling, Gen. Ins., (Wyts-
man) Cler., 1903, p. 90. — Schenkling, Col. Cat., (Junk) Cler.,
1910, p. 108. — Lea, Ann. Soc. Ent. Belg., LI, 1907, pp. 331 et 362.

Eumede Pascoe, Ann. Mag. Nat. Hist., (4) XVII, 1876, p. 50. —
Broun, Man. New Zealand Col., I, 1880, p. 334.

Hydnocera Newman, The Entom., 1841, p. 36. — Newman, idem,
1842, p. 365. — White, Cat. Cleridae Brit. Mus., IV, 1849, pp. 61
et 62.

Clerus Newman, Zoologist, 1843, p. 119 (sp. hilaris).

Hoploclerus White, l.c. p. 40. — Westwood, Proc. Zool. Soc.
Lond., XX, 1852, p. 52.

Hydnocerus Westwood, lac cit., 1852, p. 47.

Lemidius Westwood, idem, 1852, p. 47.

Laiomorphus Pic, Mel. Exot.-ent., L, 1937, p. 20.

Diagnosis of Lemidia, revised

Body small, rather elongate, parallel or posteriorly wider.
Head short, with the eyes broader than the prothorax, front
broad between the eyes, which are large, prominent, feebly, shal-
lowly emarginate in front, finely granulated. Maxillary palpi
with the terminal segment cylindrical, somewhat oval, acute at
apex ; that of the labial palpi much larger, depressed, securiform.
Labrum prominent, emarginate. Antennae very short, reaching
the hind margin of the head, eleven-segmented, segment 1 thick
and long, subcylindrical, not rising to the height of the front, seg-
ment 2 very short, swollen at middle and subglobular, segments

June, 1944]

Wolcott: Phyllob^enin^e

127

3 to 8 short, diminishing in length, without becoming sensibly
narrower, segments 9 to 11 thick and depressed, equally dilated,
forming a quite distinct clava of these three segments, the first two
are equal, subtransverse, the last longest and oval. Prothorax
longer than broad, narrower than the head, much narrower at
base, anteriorly with at most a feeble transverse impressed line.
Elytra oblong, surface depressed, entirely covering the abdomen,
irregularly punctured, suture usually closed, spices conjointly
rounded, rarely separately mucronate. Legs long and slender,
the femora extending to or beyond apex of the elytra, tarsi five-
segmented (four segments only visible from above), the first
segment very small or rudimentary but usually perfectly visible
from below, the first three lamellate, the second, third and fourth
segments diminishing in length. Ungues small, simple or at
most with an indistinct tooth at base.

Genotype. — Lemidia ( Hydnocera ) nitens Newman (Spinola).
Monobasic.

Geographical range, sixty-two species of this genus are found
in Australia, one in New Zealand, two in Tonkin, and seven in
Tasmania.

Genus Isohydnocera Chapin

Isohydnocera Chapin, Bull. Brookl. Ent. Soc., XII, 1917, p.
83. — Bradley, Man. Gen. Beetles Amer. N. of Mex., 1930, p. 105.

Diagnosis of Isohydnocera

“Body very elongate; front vertical; eyes prominent, finety
granulate, entire or minutely emarginate near antennal inser-
tion; terminal segment of maxillary palpi cylindro-conical, that
of labial palpi large, triangular; mandibles with a tooth near
apex on the inside. Antennae 11-segmented, first segment thick,
moderately long and somewhat arcuate, second short, globose,
third to ninth cylindrical, longer than broad, tenth very large,
forming with the eleventh a compact club which is much thicker
than the ninth segment. Prothorax distinctly longer than broad,
nearly cylindrical, lateral dilation weak. Elytra broader than
the thorax, shorter than the abdomen, attenuate or parallel and
truncate, lateral margins and usually tips strongly serrate. Legs
long and thin, tarsi with five segments, the first covered by the

128

Journal New York Entomological Society

[Vol. LII

second from above, segments two, three and four bearing lamel-
lae beneath, ungues long and thin, simple” (Chapin).

■ ‘ To this genus are to be assigned the following species : I. curti-
pennis Newman ( longicollis Ziegl.), tabida Lee., schusteri Lee.,
ornata Wole., gerhardi Wole., pusilla Schaeff., aegra Newm.,
brunnea sp. nov. and albocincta Horn” (Chapin).

Genotype. — Isohydnocera ( Hydnocera ) curtipennis Newman,
designated by Chapin.

Geographical range, North, Central and South America.

Genus Wolcottia Chapin

Wolcottia Chapin, Bull. Brookl. Ent. Soc., XII, 1917, p. 84.—
Bradley, Man. Gen. Beetles Amer. N. of Mex., 1930, p. 105.

Diagnosis of Wolcottia

“Body elongate, slightly attenuate posteriorly. Head short,
front flat, lateral margins slightly concave to accommodate eyes,
which are not prominent, finely granulate and minutely cemargi-
nate as in Isohydnocera. Labrum broad, entire. Mandibles
falciform, with a small internal tooth near apex. Terminal seg-
ment of maxillary palpi conical, as long, or slightly longer than
the preceding, that of the labial palpi very large, triangular.
Antennae short and stout, 11-segmented ; first segment slightly
longer than broad, second nearly spherical, third to eighth as
long as broad, trapezoidal, becoming progressively broader, ninth
broader than long, nearly as wide as tenth, tenth and eleventh
forming an oval mass nearly as long as the three preceding seg-
ments. Thorax slightly broader than long, sides feebly dilated,
slightly narrower than the head with eyes. Elytra at humeri
broader than the head across eyes, attenuate posteriorly, tips
rounded separately and with irregular margins. Legs long and
thin, hind femora reaching beyond the tips of the elytra, tarsi
of five segments, the first covered by the second from above,
second, third and fourth bearing lamellae beneath. Ungues long
and thin, simple or but slightly thickened” (Chapin).

Genotype. — W. ( Hydnocera ) pedalis Leconte, designated by
Chapin.

Geographical range, North America.

June, 1944]

Wolcott: Phyllob^enin^e

129

Dr. Chapin states that “The three species assigned to this
genus, W. pedalis Lee., sobrina Fall, and parviceps Schaeffer,
have antennae which seem to be midway between the three-seg-
mented club type of Lemidia Spinola and the two-segmented club
type of Hydnocera Newman.”

Genus Parmius Sharp

Parmius Sharp, Ent. Mo. Mag., XIII, 1877, p. 272. — Broun,
Man. New Zealand Col., I, 1880, p. 331. — Schenkling, Gen. Ins.,
(Wytsman) Cler., 1903, p. 96. — Schenkling, Col. Cat., (Junk)
Cleridae, pars 23, 1910, p. 112.

Diagnosis of Parmius, revised

Elongate, alate, slender. Head with the eyes as broad as the
elytra, broader than the thorax. Terminal segment of the maxil-
lary palpi small, that of the labial palpi very large and broad.
Eyes very prominent, finely granulated, entire or with a more
or less definite emargination, antennae short and rather slender,
11-segmented, the two or three terminal segments forming a short,
broad clava, the tenth being strongly transverse and the eleventh
rather stouter than the tenth and hardly as long as broad.
Thorax subcylindrical but a little dilated at the middle, about as
long as broad, but narrower than the elytra, constricted in front
and behind, very narrow and parallel, the humeri distinct and
rectangular. Legs long, tarsi moderately long and broad, very
distinctly five-segmented, the second segment inserted very near
the tibia. Hind coxae rather widely distant.

“The two species ( debilis Shp., and longipes Shp.,) differ from
Paupris aptera by their finely granulated eyes, by the presence
of wings and the more normal form of the elytra ’ ’ ( Sharp ) .

Genotype. — Parmius longipes Sharp, by present designation.

Geographical range, four described species known, all from
New Zealand.

Genus Neohydnus Gorham

Neohydnus Gorham, Ann. Mus. Genova, (2) XII, 1892, p.
742. — Schenkling, Gen. Ins., (Wytsman) Cler., fasc. 13, 1903,
p. 94. — Schenkling, Col. Cat., (Junk) Cleridae, pars 23, 1910,
p. 107.— Chapin, Phil. Jour. Sci., XXV, No. 2, 1924, p. 180.

130

Journal New York Entomological Society

[Yol. LIJ

Emmepus Motchoulsky, Bull. Imp. Nat. Moscou, XXXIV,
1861, p. 127 (sp. flavipes).

Diagnosis of Neohydnus, revised

Head vertical. Labrnm entire ; mandibles stout, falcate, with
a short internal tooth toward apex. Eyes large, finely granu-
lated, pubescent, entire or minutely emarginate at base of an-
tennae; the head plane between the eyes (very much as in Cal-
limerus). Terminal segment of maxillary palpi cylindro-acumi-
nate, that of labial palpi dilated, elongate-triangular. Antennae
short 11-segmented (often apparently ten-segmented, the tenth
and eleventh segments being anchylosed), first segment short
and thick, slightly bent, second nearly spherical, third to ninth
trapezoidal, becoming successively shorter to eighth or ninth,
ninth sometimes longer than the eighth, tenth and eleventh to-
gether oval and flattened. Thorax with length and breadth sub-
equal, not widened in front (as in Phyllobcenus) , with a con-
stricted line near the front, sides slightly dilated below this con-
struction, lateral foveae deep, oblique above dilations, usually
two or three in number, no basal constriction but margined at
base, anterior coxal cavities widely open behind. Elytra usually
as long as body, confusedly punctate, no trace of striae, suture
slightly dehiscent toward apices, lateral margin and tips usually
strongly serrate. Abdomen with six visible ventral segments.
Sexual modifications of males usually conspicuous. Legs slender,
femora somewhat swollen ; tarsi short, of five segments, first seg-
ment short and ventral to second, ungues with a broad basal
process.

Genotype. — Neohydnus despectus Gorham.

Geographical range, Indo-Malaysia.

Genus Metaxina Broun

Metaxina Broun, Ann. Mag. Nat. Hist., (8) III, 1909, p. 407.

Diagnosis of Metaxina, revised

Head (eyes included) nearly as large as the thorax, forehead
limited between the antennae by a slightly raised suture. Palpi
stout, long, the terminal segment of maxillary palpi quite oblique

June, 1944]

Wolcott: Phyllob^nin^e

131

at apices; that of the labial palpi subtriangular or securiform,
truncate at apices. Eyes prominent, distinctly granulated, ap-
parently rotundate, in reality transversely broadly oviform, en-
tire. Antennae eleven-segmented, inserted not in cavities, but
on slight prominences, or articulations in front of the eyes ; basal
segment stout, pyriform, third segment slightly longer than
second or fourth ; segments four to eight nearly equal in breadth,
each longer than broad, and narrowed at the base ; segments nine
and ten distinctly broader than the preceding, but little longer ;
eleventh rather larger and oviform ;. these organs therefore more
nearly resemble those of Paupris and Parmius than of Phymato-
phcea. Prothorax somewhat cordiform, transverse, apex trun-
cate, base rounded, the lateral margins fine and curved toward
the base, but without forming posterior angles; scutellum elon-
gate. Prosternum truncate in front. Coxae with trochanters;
the anterior prominent, separated only by the prosternal process ;
intermediate rather less prominent, almost contiguous; the pos-
terior small, only moderately separated. Metasternum moder-
ately elongate, its sides distinctly punctured. Elytra parallel-
sided, broader than the thorax, with obtusely prominent humeri,
the punctuation moderately coarse, not close and subseriate, sur-
face subdepressed, apices not quite smooth, elongate. Tarsi five-
segmented, basal segment abbreviated above, its lobes prolonged
below; segments two to four of nearly equal length, their lobes
also prolonged but without perceptible lamellae underneath ; fifth
simple, nearly as long as the preceding four together; basal seg-
ments of the anterior tarsi furnished with long slender, brush-
like setae below. Abdomen composed of six segments, the basal
segment rather larger than the others, segments two to five about
equal ; sixth short, deeply emarginate, with a supplementary
conical segment protruding therefrom.

Genotype. — Metaxina ornata Broun, designated by Broun.

Geographical range, 1 species of New Zealand.

Broun remarks that “the type of this genus is quite unlike
any Australian or New Zealand species known to me, and cer-
tainly an aberrant form, owing to the absence of tarsal lamellae,
the unnotched eyes, &c.” This genus and its genotype has not
been mentioned in the literature since the genus was created by

132

Journal New York Entomological Society

[Vol. Lll

Broun. This is truly an anomalous genus, but this being an
insular insect, and obviously a highly adaptive form, the singu-
larity of its aspect need cause no great doubt as to the propriety
of including this genus in the present subfamily.

Genus Abrosius Fairmaire

Abrosius Fairmaire, Bull. Mus. Hist. Nat., Paris, VIII, 1902,
p. 315. — Schenkling, Gen. Ins., (Wytsman) Cler., 1903, p. 96. —
Schenkling, Col. Cat., (Junk) Cler., 1910, p. 112. — Gahan, Ann.
Mag. Nat. Hist., (8) V, 1910, p. 65.

Diagnosis of Abrosius translated from the original

“This new genus of Cleridae approaches the Hydnocerides by
the tarsi having the first segment indistinct, the eyes entire and
the antennae inserted near their lower border; but the eyes are
rather depressed, the legs of ordinary length and the facies is
very different from that of Evenus, recalling greatly that of
Aulicus. The antennae are slender, of eleven articles, the three
terminal segments forming a small, rather narrow, lax mass ; the
front is rather large, but the eyes moderately convex. Prothorax
scarcely longer than broad, base moderately narrow, sides
rounded, scarcely sinuate at base. Elytra oblong, humeri rather
angulate, surface with basal portion rather strongly seriate-
punctate, posteriorly minutely and confusedly punctate. Legs
slender, rather short, tarsi seen from above four-segmented, the
first segment indistinct, the intermediate segments provided with
rather long lamellae. ’ ’

Genotype. — Abrosius cyaneorufus Fairm., by sole inclusion.

Geographical range, 1 species of Madagascar.

The genus Abrosius is unfortunately unknown in nature to the
writer, otherwise additional characters would be given to enable
recognition of this very poorly characterized genus, the diagnosis
of which fails to mention the chief characters usually employed
for this purpose, such as the structure of palpi, and gives only a
very ambiguous and contradictory account of the eyes and an
inefficient description of the legs and the tarsi.

Genus Cephaloclerus Kuwert

C ephaloclerus Kuwert, Ann. Soc. Ent. Belg., XXXVII, 1893,
p. 486. — Kraatz, Deutsch. Ent. Zeit., 1899, p. 105. — Schenkling,

June, 1944]

Wolcott: Phyllob^nin^e

133

Bull. Mus. d’Hist. Paris, VIII, 1902, p. 330 (in species). —
Schenkling, Gen. Ins., (Wytsman) Cler., 1903, p. 90. — Hintz,
Deutsche. Ent. Zeitschr., 1905, p. 312. — Schenkling, Deutseh. Ent.
Zeit., 1906, heft 2, p. 300. — Gahan, Ann. Mag. Nat. Hist., Lond.,
(8) V, 1910, p. 65. — Schenkling, Col. Cat., (Junk) Cleridae,
1910, p. 111.

Hydnocera Boheman, Ins. Caffr., I, fasc. 2, 1851, p. 511.

Diagnosis of Cephaloclerus, revised

Head large, rather strongly rounded, with the eyes much
broader than the thorax, clypeus strongly emarginate, labrum
emarginate; terminal segment of maxillary palpi short, slender,
cylindrical, acute at apex ; that of labial palpi larger, bell-shaped.
Antennae very short, scarcely longer than the thorax, eleven-seg-
mented, slender, segments four to ten very short, serrate, clava
elongate, but little distinct, three-segmented, terminal segment
rounded, its apex obtuse, within subacuminate, eyes large, ovate,
entire, moderately convex, prominent. Thorax nearly one-half
longer than broad, angles strongly rounded, margins acute, pos-
teriorly bisinuate, apex truncate, narrower than the head, sides
before the middle nearly straight then behind the middle ab-
ruptly narrowed, posteriorly narrower, basal margin truncate,
thinly reflexed, near apex and near base a transverse impressed
line, surface slightly convex, sides at apex and base narrowly
constricted. Elytra oblong-quadrate, narrower than the head,
nearly three times as long as the thorax and twice as broad as
the thorax at base, base slightly broadened, obliquely truncate,
humeri strongly rounded, not elevated, sides thinly margined,
straight (sinuate in C. basalis), apex conjointly rounded, coarsely
and deeply punctate. Legs slender, posterior tarsi viewed from
above evidently four-segmented, segments one, two and three
dilated, penultimate segments lamellate, ungues simple.

Genotype. — Cephaloclerus corynetoides Kuwert. Monobasic.

Geographical range, 23 species, all of Africa.

Genus Eurycranium Blanchard

Eurycranium Blanchard, Hist. Ins., II, 1845, p. 87. — Chenu,
Encycl. d’Hist. Nat. Col., II, 1860, p. 245.

134

Journal New York Entomological Society

[Vol. LII

Eurycranus Blanchard, Hist. Ins., II, 1845, p. 84. — Gemm. et
Harold, Cat. Col., VI, 1869, p. 1745 (pars). — Gahan, Ann. Mag.
Nat. Hist., (8) V, 1910, p. 64 and 65.

Eurymetopum Blanchard, Voy. d’Orb., 1844, p. 92.1 — White,
Nomenc. Col. Ins. Brit. Mus. Cler., IV, 1849, p. 42. — Chevrolat,
Cat. des Cler., Rev. Mag. Zool., 1874, p. 275.

Epiclines Spinola, Gay Hist. Chile, Zool., IV, 1849, p. 386
(pars). — Lacordaire, Gen. Col., IV, 1857, p. 463 (pars). — Gem-
minger et Harold, Cat. Col., VI, 1869, p. 1745 (pars). — Gorham,
Cist. Ent., II, 1876, p. 91 (“nil ad rem”). — Lohde, Ent. Zeit.
Stett., IX, 1900, p. 71. — Schenkling, Gen. Ins., (Wytsman) Cler.,
1903, p. 71 (pars). — Schenkling, Col. Cat., (Junk) Cler., 1910,
p. 99 (pars).

Clerus Spinola, Mon. Cler., II, 1844, p. 158 (in species).

Thanasimus Spinola, Gay Hist. Chile, Zool., IV, 1849, p. 392
(pars). — Fairmaire et Germain, Col. Chilensia, Paris, II, 1861,
p. 3. — Philippi, Stett. Ent. Zeit., XXV, 1864, p. 266 (pars). —
Philippi, Ann. Univ. Chile, Santiago, XXVI, 1865, p. 656 (in
species).

Dereutes Chevrolat, Mem. des Clerites, 1876, p. 29 (pars) (a
group name only!).

Thereutes Harold, Col. Hefte, XVI, 1879, p. 248 (pars)
(inutilis emendat.) .

Diagnosis of Eurycranium, revised

Head short, very wide across the eyes, which are very promi-
nent and nearly entire. The maxillary palpi are short, the ter-
minal segment elongate, conico-cylindrical, that of the labial

i Eurymetopum Blanchard is preoccupied by Eurymetopon Eschscholtz,
Atlas, Zoologique du voyage du capitaine Kotzebue, fascicule 2, p. 8, pi. 18,
figs. 1 and 2, to designate a genus of the family Tenebrionidae. Later
Schonherr (Genera et Species Curculionidum, Paris, VI, 1, 1840, p. 112)
employed the name Eurymetopus to designate a genus of Tetramerous coleop-
tera, of the Brachypterous Curculionides. This author has described but one
species ( E . fallax ) of the great number now known from South America.
Chevrolat says (Mem. Cler., p. 30), “The three names represent the same
root. That [of Dereutes ] which I employ indicates the mode of life of the
greater part of the true derides. Not only do they live on insects in the
larval state, but arriving at the perfect state they run on the bark of trees,
pouncing upon those that they meet devouring them with avidity.”

June, 1944]

Wolcott: Phyllob^enisle

135

palpi securiform, very large, the mandibles are rather small with
a feeble tooth on internal side. Antennae slender, long, eleven-
segmented, nearly as long as head and thorax united, the funicle
very long with the three last segments of antennae forming a
small ovate, serrate clava. Prothorax long, cylindrical and con-
stituting nearly a third of the total length of the insect, sides of
thorax more or less broadly dilated at middle, apical and basal
constrictions and transverse impressions usually quite feeble,
basal margin thinly reflexed. Elytra rather long, entirely cover-
ing the abdomen, flexible, suture closed, apex conjointly rounded,
surface very uneven, puncturing imperceptible. Legs moder-
ately long, slender, tarsi five-segmented, first segment very short
and small, but still visible from above, the second longer than
the first but still less stout, third slightly shorter than the second,
fourth dilated.

Genotype. — E. maculatum Blanchard, by present designation.

Geographical range, South and Central America.

Genus Paupris Sharp

Paupris Sharp, Ent. Monthly Mag., XIII, 1877, p. 271. — Gor-
ham, Trans. Ent. Soc. Lond., 1877, p. 263. — Schenkling, Gen. Ins.,
(Wytsman) Cler., 1903, p. 95. — Broun, Man. New Zealand Col.,
I, 1880, p. 331. — Schenkling, Gen. Ins., (Wytsman) Cler., 1903,
p. 95. — Gahan, Ann. Mag. Nat. Hist., (8) V, 1910, p. 65. — Schenk-
ling, Col. Cat., (Junk) Cler., 1910, p. 112.

Diagnosis of Paupris, revised

Elongate, apterous. Head (the eyes included) rather broader
than the thorax at its broadest part. Eyes rather small, moder-
ately prominent, coarsely granulated, with a very small tri-
angular excision. Maxillary palpi with terminal segment small
and cylindrical, not at all dilated, while that of the labial palpi
is very large, transversely broad, and dilated. Antennse short and
stout, inserted well before the eyes, eleven-segmented, segment
two shorter than the third, segments of the funicle three to eight
all subequal in length, segments nine to eleven each broader than
long, about twice as broad as the preceding segments, the terminal
segment as broad as long, its apex subacuminate, these segments

136

Journal New York Entomological Society

[Yol. LII

forming a rather lax clava. Prothorax elongate and narrow,
longer than broad, sides at middle somewhat dilated, but scarcely
tuberculate. Elytra small and narrow, much abbreviated,
usually nearly three dorsal segments shorter than the abdomen,
base narrow, humeri absent, suture closed, elongate-obovate, very
sparsely covered with short erect coarse hairs, apex conjointly
rounded. Legs long and rather stout, tarsi moderately broad,
five-segmented, first segment very small and short, nearly covered
by the elongate second segment, segments three and four each
shorter than the second, and strongly lamellate, ungues small,
simple.

Genotype. — Paupris aptera Sharp. Monobasic.

Geographical range, one species of New Zealand.

Genus Isolemidia Gorham

Isolemidia Gorham, Trans. Ent. Soc. Lond., 1877, pp. 249,
257. — Gorham, Biol. Centr.-Amer., Ill, pt. 2, 1883, p. 177. —
Schenkling, Gen. Ins., (Wytsman) Cler., 1903, p. 92. — Schenk-
ling, Col. Cat., (Junk) Cler., 1910, p. 107. — Gahan, Ann. Mag.
Nat. Hist., (8) V, 1910, p. 65. — Wolcott, Publ. Field Mus. Nat.
Hist., Zool., VII, 1910, p. 376 (in species). — Chapin, Proc. Ent.
Soc. Wash., XXII, 1920, pp. 51 and 52 (in species).

Diagnosis of Isolemidia , revised

Head with the eyes broader than the thorax, eyes very large,
globular, entire or at most minutely emarginate, maxillary palpi
subfiliform, terminal segment truncate at apex ; that of the labial
palpi securiform. Thorax subcylindrical, but often short and
sometimes quadrate or broader than long, the sides broadly
rounded or tuberculate at middle, strongly constricted near apex
and base, the basal and apical transverse impressed lines very
distinct. Antennae very short, usually little longer than the
head, as in Lemidia, eleven-segmented, those segments succeed-
ing the basal segment a little longer than wide, the three last
transverse, forming a small, short, rather connate, oblong, clava.
Elytra elongate, usually much broader than the thorax, and
broadest behind the middle, the apex conjointly rounded, usually
covering the abdomen, roundly truncate ( pulchella Gorh.), the

June, 1944]

Wolcott: Phyllob^enusle

137

humeri distinct, sometimes with a very small callus (/. cariniceps
Wole., the elytra are much abbreviated, the suture widely dehis-
cent, the apices separately obtusely rounded with a few well
developed teeth). Legs long and slender, posterior femora mod-
erately long, tarsi five-segmented, the basal segment as in Lemidia,
atrophied, second segment covering the first, ungues simple.

Genotype. — I. pulchella Gorham, designated by Gorham.

Geographical range, thirteen species of South and Central
America.

Genus Emmepus Motschoulsky

Emmepus Motschoulsky, Bull. Moscou., XVIII, 1845, p. 41 (sub
Staphylin.). — Lacordaire, Gen. Col., IV, 1857, p. 494. — Chenu,
Encycl. d’Hist. Nat. Col., II, 1860, pp. 82 and 266 (sub Staphy-
lin.).— Gorham, Trans. Ent. Soc. Lond., 1877, p. 263. — Reitter,
Verh. Nat. Ver. Brunn., XXXII, 1893 (1894), p. 39. — Reitter,
Best. Tab., 1894, p. 6.— Reitter, Wien. Ent. Zeit., XV, 1896, p.
283. — Schenkling, Gen. Ins., (Wytsman) Cler., 1903, p. 96. —
Gahan, Ann. Mag. Nat. Hist., (8) V, 1910, p. 65. — Schenkling,
Col. Cat., (Junk) Cler., 1910, p. 112.

Brachyclerus Fairmaire, Compt. rend. Soc. Ent. Belg., XXVII,
1883, p. 157. — Bedel, Ann. Soc. Ent. Fr., (6) VII, 1887, p. 197.

Diagnosis of Emmepus, revised

Body slender, small. Head transversely quadrate, front broad,
mandibles internally with a distinct tooth near apex. Maxillary
palpi with the terminal segment small, nearly cylindrical, at apex
much smaller ; that of labial palpi large, securiform. Eyes large,
prominent, finely granulated, ? entire. Antennae very short,
eleven-segmented, the two first segments large and thick, the
succeeding segments much shorter and more slender, the two
terminal segments forming a ball-like clava, first segment of
clava (the tenth of the antennae) much swollen, partly covering
the very small apical segment. Prothorax as long as broad, the
sides at middle broadly subangulately dilated. Elytra twice as
long as broad, much larger than the thorax, much shorter than
the abdomen, suture strongly dehiscent, scarcely covering the
abdomen behind the middle. The wings long, extending well
beyond the apex of the abdomen which it covers. Legs long and

138

Journal New York Entomological Society

[Vol. LII

slender, tarsi rather long and slender, distinctly five-segmented,
first segment longer than the second, segments three and four
each successively shorter than the preceding segments, segments
one to four with lamellae beneath, segment five elongate-clavate,
much longer than the two preceding segments united, ungues
simple.

Genotype. — Emmepus arundinis Motsch. Monobasic.

Geographical range, three species known from Caucasus, 'Cas-
pian Sea, South Algeria, Turkestan and Ceylon.

Genus Theano Castelnau

Theano Castelnau, Silberm. Rev., IV, 1836, p. 51. — Lacordaire,
Gen. Col, IV, 1857, p. 493.— Chenu, Encycl. d’Hist. Nat. Col, II,
1860, p. 268. — Gorham, Trans. Ent. Soc. Lond, 1877, p. 363. —
Schenkling, Gen. Ins, (Wytsman) Cler, 1903, p. 97. — Gahan,
Ann. Mag. Nat. Hist, (8) V, 1910, p. 65. — Schenkling, Col. Cat,
(Junk) Cler, 1910, p. 113.

Diagnosis of Theano , revised

Body small, rather long. Head with the eyes broader than the
thorax. Labrum entire, its front arcuate, mandibles with an
internal tooth near apex. Terminal segment of labial palpi large,
securiform, acuminate at apex. Eyes moderately large, promi-
nent, entire. Antennae short, slender, eleven-segmented, the first
two segments equal, rather large, the third segment slender, the
succeeding five segments rather slender, trianguler, the last three
segments forming a swollen, ovate or somewhat globular clava.
Thorax much narrower than the head, rounded at the sides.
Elytra somewhat elongate, surface very coarsely punctate. Legs
long and slender, posterior femora extending slightly beyond the
apex of the abdomen, tarsi with the basal segment conical, the
succeeding three segments short, dilated and bifid, provided
with prolonged lamellae beneath.

Genotype. — Theano pusilla Cast. Monobasic.

Geographical range, one species from South America (Colom-
bia).

Subgenus Callimerus Gorham

Callimerus Gorham, Cist. Ent, II, 1876 (1875-1882), p. 65. —
Schenkling, Gen. Ins, (Wytsman) Cler, fasc. 13, 1903, p. 24. —

June, 1944]

Wolcott: Phyllob^enesle

139

Gahan, Ann. Mag. Nat. Hist., (8) V, 1910, p. 65. — Schenkling,
Col. Cat., (Jnnk) Cler., 1910, p. 26. — Chapin, Philipp. Jonr. Sci.,
XXV, No. 2, 1924, pp. 180 and 191. — Pic, Exot.-Ent., fasc. 54,
1929, p. 17 (pars).

Xylobius White, Nomenc. Col. Brit. Mus., Cler., IV, 1849, p. 50
(in species) .—Westwood, Proc. Zool. Soc. Lond., XX, 1852, p. 40
(in species).

Lemidia Thomson, Mus. Scient., II, 1860, p. 61 (in species). —
Pascoe, Jour. Ent., I, 1860, p. 48. — Chevrolat, Rev. Mag. Zool.,
1874, p. 321. — Kuwert, Ann. Soc. Ent. Belg., XXXVII, 1893, p.
484.

Caloclerus Kuwert, Ann. Soc. Ent. Belg., XXXVII, 1893, p.
480. — Schenkling, Ann. Soc. Ent, Belg., VL, 1901, p. 105.

Diagnosis of Callimerus, revised

Form elongate, or oblong, head with eyes slightly broader than
apex of thorax, eyes prominent, finely granulate, very slightly
emarginate at base of antennae. Labrum entire ; mandibles stout,
falcate, with internal tooth near apex ; terminal segment of maxil-
lary palpi twice as long as preceding segment, subulate ; that of
the labial palpi elongate-triangular. Antennae rather short,
slender, distinctly eleven-segmented, first segment stout, bent, at
least twice the length of the second, which is from nearly as long
as broad to equilateral and globular, segments three to eight much
longer than broad, each segment shorter than the preceding, ninth
to eleventh forming a lax, oval clava, the eleventh oval at apex.
Thorax longer than broad, somewhat constricted before and be-
hind, moderately to strongly dilated at or before the middle,
anterior coxal cavities very widely open behind. Elytra entirely
covering the abdomen, long, narrow, sides parallel or gradually
becoming narrower posteriorly, adorned with white or yellow
scales, these scales often forming a, definite pattern, apex some-
times abbreviated and truncate or rarely each mucronate, surface
variously punctate. Abdomen with six visible ventral segments ;
in the male the terminal segments are often profoundly modified ;
in the female the modification, if any, occurs on the penultimate
segment. Legs rather long, posterior tibiae with or without a
notch with comb of hairs near its apex; tarsi moderately long,

140

Journal New York Entomological Society

[Yol. lii

five-segmented, the basal segment the longest. Ungues nearly
simple or with a broad plate-like tooth.

Subgenotype. — Clerus ( Xylobius ) dulcis Westw., designated
by Gorham.

Geographical range, 102 species known from Indo-Malaysia.

Subgenus Cucujocallimerus Pic

Cucujocallimerus s. g. Pic, Mel. Exot.-ent., fasc. LIV, 1929,
pp. 17-18.

Callimerus auct. (pars).

Diagnosis of Cucujocallimerus, slightly revised

Prothorax less than, or little longer than broad, or sometimes
very elongate, posterior margin less narrow, often rather broad,
or sometimes indistinct, femora more or less slender or a little
stouter, sometimes the posterior stouter than the others; the
tarsi the most often slender and long. Form of the body oblongo-
elongate or narrow. Antennas less short. Head with the eyes
much broader than the front part of the prothorax, this very im-
pressive. Body wholly, as a matter of fact, depressed above.

“The sub-genus Cucujocallimerus mihi has the legs principally
the posterior femora much longer than the others, whereas the
greater portion of the species of the s. genus Callimerus has them
little or moderately longer than the others” (Pic).

Subgenotype. — Cucujocallimerus ( Callimerus ) coomani Pic.,
designated by Pic.

Geographical range, 1 species of Tonkin.

Subgenus Brachycallimerus Chapin

Br achy callimerus Chapin, Philipp. Jour. Sci., XXV, No. 2,
1924, pp. 180 et 190. — Corporaal, Eev. Fran^aise d’Ent., IV,
1937, p. 60.

Callimerus auct. (pars), loc. cit., p. 190.

Crassocallimerus Pic, Mel. Exot.-ent., LIV, 1929, pp. 17 et 18.

Diagnosis of Brachycallimerus, only slightly revised

Head broad, eyes very prominent, very slightly emarginate
near antennal socket; labrum entire, mandibles broad and flat,

June, 1944]

Wolcott: Phyllob^enin^:

141

falcate, with a sharp tooth internally near apex; antennae short
and compact, eleven-segmented, first segment short and stont,
second nearly spherical, third to sixth longer than broad, seventh
to eleventh broader than long, each broader than the preceding,
together forming a compact obtriangnlar five-segmented clava.
Thorax broader than long, polished, with a few distinct punc-
tures. Elytra broader at base than the thorax, punctures numer-
ous, not seriate. Entire dorsal surface of insect devoid of scales.
Abdomen with six visible ventral segments, secondary sexual
modifications conspicuous. Legs moderate in length, posterior
tibiae with subapical notch, tarsi rather long and slender, five-
segmented, first segment longer than the second, distinct and not
covered by the second, ungues broadly toothed at base.

Subgenotype. — Callimerus latifrons Gorh., designated by
Chapin.

Geographical range, Indo-Malaysia.

Chapin states that “The insects which I include in this new
genus have heretofore been classified with the species of Calli-
merus Gorh. They differ from those species in their broad and
compact form, short and compact antennae, and the total absence
of scales from the upper surface. In addition to the genotype,
I would include the following species : latesignatus Gorh., rus-
ticus Gorh., pectoralis Schklg., and probably trifasciatus Schklg.”
Corporaal (1937 : 60) includes the last named species without
question and adds pallidus Gorh., and doesburgi Corp., n. sp., as
members of this genus.

Genus Evenus Castelnau

Evenus Castelnau, Silb. Rev., IV, 1836, p. 41. — Spinola, Rev.
ZooL, 1841, p. 75. — Klug, Abhandl. Berl. Akad. 1842, p. 315. —
Spinola, Mon. Clerites, II, 1844, p. 28. — Lacordaire, Gen. Col.,
1857, p. 469. — Chenu, Encycl. d’Hist. Nat. Col., II, 1860, p. 266.
— Gorham, Trans. Ent. Soc. Lond., 1877, p. 249. — Schenkling,
Gen. Ins., (Wytsman) Cler., 1903, p. 89. — Gahan, Ann. Mag. Nat.
Hist., (8) V, 1910, p. 65. — Schenkling, Deutsche Ent. Zeitschr.,
1906, p. 300. — Schenkling, Col. Cat., (Junk) Cler., 1910, p. 111.

Diagnosis of Evenus, revised

Body very narrow, filiform. Head large, constricted behind,
front concave. Maxillary palpi small, terminal segment cylin-

142

Journal New York Entomological Society

[Vol. LII

drical, rounded at apex ; labial palpi long, three times as long as
the maxillary palpi, terminal segment elongate, securiform,
slender at base. Eyes large, very prominent transversely oval,
not visibly emarginate. Antennae very short, inserted between
the eyes and very near their extreme anterior margin, eleven-
segmented, first segment rather stout, long, cylindrical, segments
three to eight obconical, small but distinct, each progressively
broader but without becoming longer, the ninth as broad as the
eighth and forming with the two last segments a narrow, elongate
clava, terminating in an acute point, a little flattened and at their
articulations rather serrate rendering at least very doubtful
the independent mobility of the tenth and eleventh segments.
Thorax narrow and elongate, disk depressed and narrowed in
front, sides feebly dilated, front and hind margins equal in width,
anterior coxal cavities open behind. Elytra depressed, much
broader than base of thorax, very elongate parallel, but not reach-
ing to apex of the abdomen, suture closed, irregularly rounded
at apex, surface rather smooth (in the typical species sometimes
seriately punctate). Legs elongate, thin, the posterior pair
twice as long as the others, their femora reaching beyond the apex
of the elytra, tibiae rather longer than the femora, straight and
cylindrical, . tarsi very long, five-segmented, equally visible, long
and narrow, the posterior having the two first elongate conical,
the first shorter than the second by about one-third, third and
fourth short, bifid, lamellate beneath, the second longer than any
of the others, subequal to segments three and four united, and
somewhat similarly depressed, a little dilated and feebly emargi-
nate at apex, the fifth segment shorter than either of the preced-
ing two, terminated by two large and short simple ungues, all
tarsi pubescent beneath.

Genotype. — Evenus filiformis Cast. Monobasic.

Geographical range, thirteen species all of Madagascar.

Genus Lasiocallimerus Corporaal
Lasiocallimerus Corporaal, Tijds. v. Ent., 82, 1939, p. 194.

Diagnosis of Lasiocallimerus Corporaal
il Tarsorum articulo primo longiore quam secundo. Statura
generi Brachycallimero similis , sed antennce 10-articulatce, arti-

June, 1944]

Wolcott: Phyllob^enin^e

143

culo robusto, articulo secundo ceque crasso, paulo breviore,
articulo tertio subelongato, ariiculis 4-9 transversis, articido
ultimo subelongato, ovato, ad apicem subacuminato. Corpus
longe pilosum et, in thorace, dense squamuloso-villosum. Pedes
quotes in genere Brachycallimero, tibiis sine tuber culo, tar sis
latis, unguiculis latis, ad apicem emarginatis.

“With the first tarsal segment longer than the second. Head
broad; eyes [rather large] very prominent, [finely facetted, with
long, erect, pale hairs] very slightly emarginate near antennal
socket, labrum entire, mandibles broad and flat, falcate. An-
tennae short and compact, ten-segmented ; first segment short and
stout, second of equal thickness, but shorter, third a little longer
than broad, fourth to ninth broader than long, tenth longer than
broad, ovate, a little acuminate. Prothorax of the same form as
in Brachycallimerus, heavily punctate, under the longer pilosity
closely covered with crisp, scale-like hairs. Elytra broader than
prothorax, evenly punctured, but not in rows. Abdomen with
six visible ventral segments. Legs of moderate length, tibiae
without notch, tarsi short and broad, claws broad, emarginate at
tip” (Corporaal).

Genotype. — Lasiocallimerus vestitus Corp., designated by Cor-
poraal.

Geographical range, one species of Java.

Genus Allelidea Waterhouse

Allelidea Waterhouse, Trans. Ent. Soc. Lond., II, 1839, p. 193.
— Lacordaire, Gen. Col., IV, 1857, p. 473. — Chenu, Encycl.
d’Hist. Nat., Col., II, 1860, p. 266. — Gorham, Trans. Ent. Soc.
Lond., 1877, p. 263. — Blackburn, Trans. Roy. Soc. South Austral.,
XIV, 1891, p. 302. — Schenkling, Gen. Ins., (Wytsman) Cler.,
1903, p. 97. — Gahan, Ann. Mag. Nat. Hist., (8) V, 1910, p. 65. —
Schenkling, Col. Cat., (Junk) Cler., 1910, p. 112.

Diagnosis of Allelidea, revised

Body elongate, cylindrical. Head with the eyes as broad or
broader than the thorax, constricted posteriorly, front verticle,
labrum transverse, emarginate. Terminal segment of maxillary
palpi ?, that of labial palpi oblique, securiform. Mandibles in-

144

Journal New York Entomological Society

[Vol. LII

ternally with a small tooth before the apex. Eyes moderately
prominent, entire. Antennae very short, eleven-segmented, ar-
ticulations very distinct, segments very small, gradually growing
larger, the three last segments slightly larger than the others
forming a very lax elongate-oblong clava. Thorax longer than
broad, narrow and elongate, subcylindrical, sides a little before
the middle roundly dilated, at bast strongly narrowed. Elytra
not much broader than thorax at base, seriately punctured, elon-
gate, gradually and very feebly broader posteriorly, apex serrate,
as long as the abdomen (J' ?), or much shorter (5 ?). The three
pairs of legs equal in length, long and slender, posterior femora
slightly shorter than the abdomen, tarsi five-segmented, first seg-
ment not much longer than the succeeding segments together,
segments two to four with bilobed membraneous appendices, fifth
segment moderate. Ungues with membraneous appendices.

This is a small insect with the facies of Phyllohoenus.

Genotype. — Allelidea ctenostomoides Waterh., designated by
Gorham.

Geographical range, seven species of Australia.

Remarks on genera more or less allied to the genera of the
subfamily Phyllobseninae, or transferable to that subfamily.

Genus Brachyptevenus Pic

Brachyptevenus Pic, Mel. Exot.-ent., LXXI, 1939, pp. 24-25.

Diagnosis of Brachyptevenus , revised

Head broader than the thorax, posteriorly long attenuate, not
strangulate. Thorax elongate, sides sinuate, middle impressed
posteriorly. Elytra but little reduced, anteriorly subattenuate,
narrow and elongate, apex slightly broader. Legs slender, ab-
domen distinctly exceeding the elytral apex. This new genus is
near E venus Cast. It is clearly characterized by the form of the
head and the elytra. It is established for the following new
species B. niger Pic.

Genotype. — Brachyptevenus niger Pic. Monobasic.

Geographical range, 1 species of Madagascar.

The writer has been compelled by reason of the brief and un-
satisfactory diagnosis of this genus to omit it from consideration
in the body of this account.

June, 1944]

Wolcott: Phyllob^enin^e

145

Genus Achlamys Waterhouse

Achlamys Waterhouse, Cist. Ent., II, 1875-1882 (1879), p.
530. — -Schenkling, Gen. Ins., (Wytsman) Cler., 1903, p. 16. —
Schenkling, Col. Cat., (Junk) Cler., 1910, p. 17.

Diagnosis of Achlamys , revised

Form cylindrical. Head (with the eyes) broader than the
thorax, the eyes prominent, coarsely granulate, scarcely emargi-
nate in front. Antennae as long as the head and thorax taken
together, the first segment thick, ovate, the second segment small,
the third elongate, subcylindrical, the fourth, fifth and sixth be-
coming gradually stouter and shorter, the seventh as broad as
long, the eighth, ninth, tenth and eleventh segments forming a
distinct four-segmented clava. Apical segment of the maxillary
palpi elongate, a little narrowed before the apex ; apical segment
of the labial palpi securiform. Prothorax very convex, one-third
longer than broad, very slightly constricted before the front
margin, strongly constricted before the base. Elytra parallel,
a little broader than the thorax, each with seven lines of deep
oblong punctures which do not extend quite to the apex, which
is smooth and obtuse. Tarsi with five distinct segments, the
ungues with a single tooth at the base.

“Closely allied to Pallensis, but with prominent eyes and dis-
tinct club to the antennas” (Waterhouse).

The writer believes Achlamys should be removed from Tillinae
to the Phyllobaeninae and there with Paupris Sharp, both genera
having coarsely facetted eyes, a character exceptional in this sub-
family, these two genera may be placed as a distinct group or
tribe designated by the term Pauprini.

Gahan (1910: 65) has transferred the genus Ellipotema
Spinola to the subfamily Corynetinae, and the genera Cleropiestus,
Calendyma Lac., and Epiclines Chevr., to the subfamily Clerinas,
he also briefly differentiates ( loc . cit p. 64) the genus Epiclines
and Eurycranus (= E-ury cranium Blanch.) and calls attention to
the long-standing and absolutely absurd erroneous synonymy
which has been perpetuated ever since the issuance of Lacor-
daire ’s Genera des Coleopteres. Of the species in our Catalogues
under Epiclines only a few, as far as can now be determined,

146

Journal New York Entomological Society

[Vol. LII

really belong to that genus, those that should be so placed are
as follows: gayi Chevr., (nec Spin.), cenea Phil., basalis Blanch.,
puncticollis Spin., ruficollis Phil., similis Schklg., advena Chevr.,
viridis Phil., and probably tristis Spinola. The rest of the listed
species with scarcely an exception should go to the genus Eury-
cranium Blanchard.

Genotype of Epiclines, E. gayi, Chevrolat. Monobasic.

Geographical range, Chile.

The genera of this subfamily may be grouped according to
natural characters as follows:

Group 1

Clava two-segmented; basal segment of hind tarsi short, cov-
ered above by the second segment, prothorax variously propor-
tioned.

Pliyllobcenus Dejean, Lemidia Spinola, Isohydnocera Chapin.

Group 2

Clava indefinitely two- or three-segmented; basal segment of
hind tarsi short, covered above by the second segment, prothorax
with length and breadth subequal.

Wolcottia Chapin, Parmius Sharp, Neohydnus Gorham,
Metaxina Broun.

Group 3

Clava three-segmented ; basal segment of hind tarsi short, cov-
ered above by the second segment, prothorax with length usually
greater than breadth.

Abrosius Fairmaire, C ephaloclerus Kuwert, Eury cranium
Blanchard, Paupris Sharp, Isolemidia Gorham.

Group 4

Clava two-segmented • basal segment of hind tarsi long, not cov-
ered by the second segment, prothorax as long as broad.

Emmepus Motchoulsky.

Group 5

Clava three-segmented; basal segment of hind tarsi long, not
covered by the second segment, prothorax at least as long as
broad.

June, 1944]

Wolcott: Phyllob^nin^

147

Theano Castelnau, sub. gen. Callimerus Gorham, sub. gen.
Cucu jo callimerus Pic.

Group 6

Clava five-segmented ; basal segment of hind tarsi long, not cov-
ered by the second segment, prothorax broader than long.

Sub. gen. Br achy callimerus Chapin.

Group 7

Clava indistinctly four-segmented ; basal segment of hind tarsi
long, not covered by the second segment, prothorax longer than
broad.

Evenus Castelnau.

Group 8

Clava one-segmented ; basal segment of hind tarsi long, not cov-
ered by the second segment, prothorax broader than long.

Lasio callimerus Corporaal.

Group 9

Clava indistinctly three-segmented ; basal segment of hind tarsi
long, prothorax longer than broad.

Allelidea Waterhouse.

BIBLIOGRAPHY

Bedel, L.

1887. Recherches sur les Coleopteres du nord de l’Afrique. Recherches
synonymiques. Ann. Soc. Ent. Fr., (6) VII, p. 195-202.

Blackburn, T.

1889. Further notes on Australian Coleoptera, with descriptions of new
genera and species. Proc. Linn. Soc. New South Wales, (2)
III, p. 1387-1506.

1891. Further notes on Australian Coleoptera, with descriptions of new
genera and species: X. Trans. Roy. Soc. South Australia,
XIV, p. 292-345.

Blanchard, Smile.

1844. Voyage dans l’Amerique meridionale de M. Alcide d’Orbigny,

VI, 1837-1844 (1844). Insectes, p. 1-222. Paris.

1845. Histoire des Insectes, traitant de leurs mceurs metamorphoses en

general et comprenant une nouvelle classification fondee sur
leurs rapports naturels, II. Paris.

1853. Voyage au pole sud et dans l’oceanie sur les corvettes 1 ’Astrolabe
et la Zelee; etc. Zoologie, IV, Description des Insectes, p.
59-65. Paris.

148

Journal New York Entomological Society

[Vol. LII

Boheman, Carl H.

1851. Insecta Caffrariae annis 1838-1845 J. A. Wahlberg collecta amici
auxilis suffultus, Coleoptera, I, fasc. 2, p. 299-626. 3 pis.

Holmiae.

Bradley, J. Chester.

1930. Manual of the genera of beetles of America north of Mexico.
New York.

Broun, Thomas.

1880. Manual of New Zealand Coleoptera, I, p. 330-337. Wellington.
1909. Descriptions of new genera and species of New Zealand Coleop-
tera. Ann. Mag. Nat. Hist., (8) III, p. 385-415.
Castelnau, Comte Francis de Laporte.

1936. Etudes entomologiques ou descriptions d ’insectes nouveaux et

observations sur la synonymie. Silb. Rev. Ent., IY, p. 5-60.
Chagnon, Gustave.

1935. Coleopteres de la province de Quebec. Nat. Canad., LXII, p.
165-175.

Chapin, Edward A.

1917. Studies in the Hydnocerini — The Hydnoceroid Genera. Bull.
Brookl. Ent. Soc., XII, p. 83-85.

1920. New American Cleridae, with note on the synonymy of Microp-
terus Chevr. (Col.). Proc. Ent. Soc. Wash., XXII, p. 50-54.
1922. New North American Hydnocera. Proc. Biol. Soc. Wash.,
XXXV, p. 55-58.

1924. Classification of the Philippine components of the Coleopterous
family CleridEe. Philipp. Jour. Sci., XXV, 2, p. 159-286.
pis. 1-5.

Chenu, Jean Charles.

1860. Encyclopedic d’histoire naturelle ou traiite complet de cette
science d’apres les travaux des naturalistes les plus eminents,
etc. Coleoptera, II, p. 1-312, pis. Paris.

Chbvrolat, Leon Auguste.

1829. Les Insectes. Les Coleopteres. Troiseme famille. — Les Serri-
cornes. Guerin’s Iconographie du Regne Animal de Cuvier.
Insectes, partie VII. 1829-1838. Paris.

1843. Description de vingt-quatre nouvelles especes de Terediles pour
faire suite a la Monographic des Clairones de M. le Docteur
Klug. Ann. Soc. Ent. Feance, (2) I, p. 31-42.

1874. Catalogue des Clerides de la collection de M. A. Chevrolat. Rev.
Mag. Zool., (3) II, p. 252-329.

1876. Memoire sur la Famille des Clerites. pp. 1-51. Paris.
Corporaal, J. B.

1937. A New Brachycallimerus from Java. Rev. Frangaise d’Entom.,

IV, p. 60-62.

1939. Studies in Callimerus and allied genera. (Col.). Tijd. v. Ent.,
LXXXII, p. 182-195.

June, 1944]

Wolcott: Phyllob^nin^e

149

Dejean, Comte Pierre Francois M. A.

1837. Catalogue des Coleopteres de la collection de le baron Dejean.
Edit. 3, corrected and augmented, p. 1-503.

Eschscholtz, Johann F.

1831. Voyage du capitainne Kotzebue. Zoologischer Atlas, IV, fasc.
2, pi.

Fairmaire, Leon.

1883. Emmepus Bonnairei. Compt. rend. Soc. Ent. Belg., XXII, p.
157.

1902. Description des Clerides recuillis par le Dr. Deeorse dans le Sud-
est de Madagascar. Bull. Mus. Hist. Nat., VIII, p. 306-317.
Paris.

Fairmaire, Leon, et Germain, P.

1861. Coleoptera Chilensis, II, p. 1-4. Paris.

Fursov, N. I.

1936. Eine neue Gattung und 5 neue arten aus Fam. Cleridae (Coleop-
tera) von Ost-Sibirien und Mittelasien. Bull. Soc. Nat.
Moscou, Sect. Biol., Nouvelle serie, XLV, 5, p. 351-354.
Gahan, Chas. J.

1902. Observations on some mimetic insects and spiders from Borneo
and Singapore, with appendices containing descriptions of

New Species by and C. J. Gahan. Proc. Zool.

Soc. Lond., II, p. 250-284, pi.

1910. Notes on Cleridae and descriptions of some new Genera and
Species of this Family of Coleoptera. Ann. Mag. Nat. Hist.
Lond., (8) V, p. 55-76.

Gemminger, Max.

1870. Epiclines spinolai. Col. Hefte, VI, p. 127.

Gemminger, Max., et Harold, Edgar von.

1869. Catalogus Coleopterorum Hucusque descriptorum synonymicus et
systematicus, VI, p. 1722-1759.

Germar, Ernst Friedrich.

1824. Insectorum species novae aut minus cognitae, descriptionibus
illustratrates. Volumen primum. Coleoptera, p. 80-81.
Gorham, Henry Stephen.

1876. Notes on the Coleopterous family Cleridae, with descriptions of

new genera and species. Cist. Ent., II (1875-1882), p. 57-
106.

1877. Descriptions of new species of Cleridae. Trans. Ent. Soc. Lond.,

p. 245-263.

1877. Descriptions of new species of Cleridae, with notes on the genera

and corrections of synonymy. Trans. Ent. Soc. Lond., p.
401-426.

1878. Descriptions of new genera and species of Cleridae, with notes on

the genera and corrections of synonymy. Trans. Ent. Soc.
Lond., p. 153-167.

150

Journal New York Entomological Society

[Vol. LII

1882. Malacodermata. Cleridae. Biol. Centr.-Amer., Ill, pt. 2, 1882-

1883, p. 129-193. Suppl., 1886, p. 332-346, 13 pis.

1892. Viaggio di Leonardo Fea in Birmania e regioni vieine, xlviii,

Cleridae. Ann. Mus. Genov., (2) XII, p. 718-746.

Harold, Edgar von.

1879. Genus Thereutes. Col. Hefte, XYI, p. 248-249, nota.

Hintz, E.

1905. Neue Cleriden aus Westafrika. Deutsch. Ent. Zeitsch., II, p.
305-314.

Klug, Johann C. F.

1842. Versuch einer systematischen Bestimmung und Auseinander-
setzung der Gattungen und Arten der Clerii, eine Insecten-
familie aus der Ordnung der Coleopteren. Abh. Konigl.
Akad. Wiss. Berlin, p. 259-397, 2 pis.

Kraatz, Gustav.

1899. Verzeichniss der von Hrn. Conradt in Westafrica (Kamerun,
Togo) gesammelten Cleriden. Deutsch. Ent. Zeitschr., p. 81-
107.

Kuwert, A.

1893. Neue Afrieanische Cleriden. Ann. Soc. Ent. Belg., XXXVII,

p. 486-497.

Lacordaire, Jean Theodore.

1857. Histoire naturelle des Insectes. Genera des Coleopteres, ou
expose methodique de tous les genres proposes jusqu’ici dans
cet ordre d’Insectes, IV, derides, p. 415-496. Paris.

Lea, Arthur M.

1907. Notes on the genus Lemidia with descriptions of new species.
Ann. Soc. Ent. Belg., LI, p. 331-362, 2 pis.

Leconte, John Lawrence.

1849. Synopsis of the Coleopterous insects of the group Cleridae which
inhabit the United States. Ann. Lye. Nat. Hist. N. Y., V,
p. 9-35, pp. [1-27 of reprint].

1861. Classification of the Coleoptera of North America. Pt. 1.

Smiths. Misc. Coll., Ill, p. 1-208 issued 1861, p. 209-278
issued 1862.

Leconte, John L., and Horn, George Henry.

1883. Classification of the Coleoptera of North America. Smiths.

Misc. Coll., XXVI, No. 507, p. i-xxxviii + 1-567. Washing-
ton.

Lesne, Pierre.

1909. Notes sur la nomenclature des Clerides (Col.). Bull. Soc. Ent.
France, p. 206.

Motschoulsky, T. Victor von.

1845. Remarques sur la collection de Coleopteres russes de Victor
Motschoulsky. Bull. Soc. Imp. Nat. Moscou, XVIII, No. 1,
p. 1-127, pi. 3.

June, 1944] Wolcott: Phyllob^nin2E 151

1861. Essai d’un cat. des Ins. de Pile Ceylan. Bull. Soc. Imp. Nat.
Moscou, XXXIV, p. 125-127.

Newman, Edward.

1837. Entomological notes. Entom. Mag. London, p. 372-402.

1840. Descriptions of new species of Coleopterous insects. Mag. Nat.

Hist., (2) IV, p. 362-368.

1841. Entomological notes. The Entomologist, 1840-1842, p. 1-37.

London.

1843. On new species of Coleoptera. The Zoologist, I, 1843-1861, p.
1-384. London.

Pascoe, Francis P.

1860. Notes on little known genera and species of Coleoptera. Jour,
of Entom., I, p. 36-64.

1876. Descriptions of new genera and species of New Zealand Coleop-
tera, Pt. 2. Ann. Mag. Nat. Hist., (4) XVII, p. 48-60.
Philippi, R. A. et Friedr.

1864. Beschreibung einer neuen Chilenischen Kafer. Stett. Ent.

Zeitung, p. 266-284.

Philippi, Rodulfo Amando.

1865. Description de algunos insectos nuevos chilenos por don Rodulfo

Amando Philippi. Ann. Univ. Chila, XXVI, p. 651-660.

Pic, Maurice.

1927. Coleopteres du Globe. Mel. Exot.-ent., L, p. 1-36.

1929. Genera allied to Callimerus. Mel. Exot.-ent., LIV, p. 16-17.
1939. Mutations et nouveautes diverses. Mel. Exot.-ent., LXXI, p.
1-36.

Reitter, Edmund.

1894. Bestimmungs-Tabelle der Coleopteren-Familie der Cleriden, des
palaearctischen Faunengebietes. Verh. Naturf. Ver. Brunn.,
XXXII, 1893 (1894), p. 37-89.

1894. Bestimmungs-Tabelle der Coleopteren-Familie der Cleriden.
Heft XXVIII, p. 1-56.

1896. Hydnoeera — Emmepus. Wien. Ent. Zeit., XV, p. 283.

Say, Thomas.

1823. Descriptions of Coleopterous insects collected in the late Expe-
dition to the Rocky Mountains, performed by order of Mr.
Calhoun, Secretary of War, under the command of Major
Long. Jour. Acad. Nat. Sci. Phila., Ill, p. 139-216.

1825. Descriptions of new Coleopterous insects inhabiting the United
States. Jour. Acad. Nat. Sci. Phila., V, p. 160-204.
SCHENKLING, SlGMUND.

1898. Revision der Cleridengattung Lemidia Spin., nebst Beschreibung
einiger neuer Arten. Deutsche Ent. Zeit., I, p. 169-182.
1901. Neue Cleriden des Koniglichen Museums zu Briissel. Ann. Soc.
Ent. Belg., XLV, p. 104-108.

Cleriden nouveaux du Museum naturelle de Paris. Bull. Mus.
Hist. Nat., VIII, p. 317-332.

1902.

152

Journal New York Entomological Society

[Vol. LII

1903. Genera Insectorum. Coleoptera Malacodermata. Fam. Cleridae.

(Wytsman), fasc. 13, p. 1-124, 5 pis. Bruxelles.

1908. Some Cleridae of the Indian Museum. Rec. Ind. Mus., II, pt. 4,
387-388.

1910. Coleoptera Catalogus. Auspiciis et auxilio W. Junk editus a
S. Schenkling, pars 23, Cleridae. Berlin.

SCHONHERR, CARL J.

1840. Genera et species Curculionidum, YI, pt. 1, p. 1-174. Paris.
Sharp, David.

1877. Description of some new genera and species of New Zealand
Coleoptera. Entom. Month. Mag., XIII, p. 271-272.
Spinola, Marquis Maximilien.

1841. Monographic des Terediles. Tableau synoptique des Clairons.

Rev. Zool. por la Soc. Cuvier., p. 70-76. Paris.

1844. Essai Monographique sur les Clerites insectes coleopteres, I, II
and Suppl., 47 pis. Genes.

1849. in Claudio Gay: Historia fisica y politica de Chile, Zoologia, IY,
Cleroideos, p. 381-414, 1 pi. Paris.

Thomson, James.

1860. Materiaux pour servir a une monographic nouvelle de la famille
des Clerides. Musee scientifique ou recueil d’histoire natur-
elle, p. 46-67.

Waterhouse, Charles Owen.

1879. New species of Cleridae and other Coleoptera from Madagascar.
Cist. Ent., 1875-1882, p. 529-534.

Waterhouse, George Robert.

1839. Descriptions of some new species of exotic insects (Coleopt.,
Hymenopt.). Trans. Ent. Soc. Bond., II, p. 188-196, fig.
Westwood, John Obadiah.

1852. Descriptions of new Cleridae from Asia, Africa and Australia.

Proc. Zool. Soc. Lond., XX, p. 35-55 (Annulosa), pis. 24-27.

White, Adam.

1849. Nomenclature of Coleopterous insects in the collection of the
British Museum, pt. IY, Cleridae, p. 1-68. British Museum,
London.

Wickham, Henry Fredr.

1895. The Coleoptera of Canada, 13. The Cleridae of Ontario and
Quebec. Can. Ent., XXYII, p. 247-253, 4 text figs.
Wolcott, Albert Burk.

1910. The Cleridae of Indiana, in Blatchley, Col. Ind. Bull 1, Dept.

Geol. Nat. Res., p. 846-862.

1910. Reprint of the above title issued separately.

1910. Notes on some Cleridae of Middle and North America with de-
scriptions of new species. Field Mus. Publ. Zool., VII, p.
339-401.

June, 1944]

Schneirla: Insect Behavior

153

THE REPRODUCTIVE FUNCTIONS OF THE ARMY-
ANT QUEEN AS PACE-MAKERS OF THE
GROUP BEHAVIOR PATTERN

T. C. Schneirla1

Although they are among the commonest ants encountered in
the tropical forests of the Old and New Worlds, the dorylines have
long remained among the least known. In Africa and Asia the
“driver ants’ ’ and in tropical America the “legionary” or
“army” ants of this subfamily present many intriguing problems,
especially in the predatory raids and nomadic life of their colonies
and the biological status of their huge wingless fertile females, the
dichthadiigynes. These last individuals are not very well known
to science, since up to a relatively short time ago virtually no pre-
cise information was available concerning their relations to the
colony. Actually we find, beyond propagative functions which
differ rather strikingly in some respects from those of fertile
females in other insects, the properties of the doryline queen make
her a factor of critical importance in the general process of colony
behavior.

This paper is directed toward accounting for the essential onto-
genetic basis of the behavior pattern characteristic in the subgenus
Eciton s. str., an American branch of the subfamily Dorylinae.
Our study specifically concerns these terrestrial army ants, espe-
cially the type species of the subgenus, E. hamatum. The results
may prove eventually to have relevance among the dorylines far
beyond the conditions of the species on which this report is based.2

1 Department of Animal Behavior, American Museum of Natural History,
New York, and Department of Psychology (Washington Square College),
New York University.

2 Acknowledgments : The field work for this study was subsidized by a
grant from the Bache Fund of the National Academy of Sciences. Prepara-
tion of the manuscript was made possible by a grant from the Committee for
Research in Problems of Sex (National Research Council), administered by
Dr. Frank A. Beach. The writer also wishes to express his thanks to Mr.
James Zetek, Custodian of the Barro Colorado Island Biological Reservation,
for his hospitality and his unstinting help with arrangements for work at the
station.

154

Journal New York Entomological Society

[Vol. LII

The chief characteristics of the behavior pattern of E. hamatum
involve the formation of bivouacs, raiding, and bivouac-change
movements which in series may be termed migrations. In this
species the temporary nest or bivouac of a given colony is a more or
less cylindrical mass formed by the clustered bodies of ants alone,
without foreign materials, usually depending from a log or vines,
or hanging against a tree. From his studies in the rainy season
the writer (1933; 1938) has described two distinctive conditions
through which colonies of E. hamatum pass successively in alter-
nation at such times : the nomadic and the statary activity phases.
Roughly, each phase lasts close to 20 days before it is succeeded
by the other. The principal characteristics of the nomadic phase
are 1) highly developed daily raids and 2) a regular change of the
bivouac site at the termination of each day. In rather sharp con-
trast, the statary phase is marked by 1) less developed raids and
2) occupancy of the same bivouac site throughout the interval.
In addition there are other features which characterize each of
these behavior phases. These are essentially intrinsic to the
colony, yet as we shall find they are intimately involved in the
entire system of events in Eciton behavior.

Our essential hypothesis is that a close relationship exists be-
tween behavior outside the bivouac (i.e., raiding and bivouac-
change processes) and conditions within the bivouac, and that this
pattern of events basically depends upon the functions of the
reproductive agent of the colony. Accordingly in this paper at-
tention focusses upon the role of the mother queen, whose proper-
ties apparently are of key importance in the entire mechanism
of the Eciton behavior pattern.

Material for this study was gathered and most of the field ob-
servations were carried out on Barro Colorado Island in the
Panama Canal Zone, in the rainy season months of 1936 and 1938
(May to September). In that locality the rainy period usually
begins in April and ends in early December. Investigations of
the Eciton problem have yet to be conducted in the dry season.
This limitation is important, since it is very probable that rainy-
season conditions are optimal for the appearance of relationships
such as we shall describe. In an indirect manner the results of
this study suggest that with extreme changes in prevalent atmos-

June, 1944]

Schneirla : ' Insect Behavior

155

pheric conditions important variations may occur in the described
behavior pattern. In view of this, it is a fact of some importance
that onr evidence was gathered in a portion of the Caribbean
lower rain-forest zone in which the rainy and dry seasons are dif-
ferentiated rather sharply from year to year. What differences
will be found under more variable meteorological conditions rep-
resent an interesting problem.

PROPERTIES OF THE ECITON QUEEN

From the time Andre (1885) captured the first dichthadiigyne
from the subterranean bivouac of an E. ( Labidus ) coecum colony
the list of captures has grown very slowly, and although fertile
females are at present known from nearly 25 of the more than 100
recorded species of Eciton and all of the eight recorded species of
Eciton s. str. (Bruch, 1934), they are still among the most highly
prized collector ’s items. In view of the great difficulty of captur-
ing them, it is scarcely surprising that these rare insects have been
almost invariably clapped into alcohol when taken, under the
influence of what Creighton has appropriately termed ‘ ‘ collector ’s
itch. ’ ’ This practice of course has not precluded the accumulation
of much valuable information concerning the external morphol-
ogy and the taxonomic affinities of the specimens; yet it has
unfortunately kept in the remote background several important
questions which can find their answers only in study of the living
queen.

Thus it is unfortunate that from the records of more than a
score of collections very little can be learned about the behavior
and biological properties of the living individual. With the ex-
ception of general observations carried out by Wheeler (1900)
with captive colonies of E. schmitti in Texas, no special investi-
gations have been attempted. The present study stems from a
general investigation of army-ant behavior which has revealed the
probability that the dichthadiigyne plays a crucial though in-
direct role in the phenomena of Eciton behavior (Schneirla,
1934; 1938).

Eciton queens may be found in either of two very different con-
ditions, the “normal” or contracted condition and the physo-
gastric or egg-producing condition (see Fig. 1). For the present
study, the dichthadiigyne material comprised 13 queens of E .

156

Journal New York Entomological Society

[Vol. LII

hamatum and one of E. burchelli in the contracted condition, and
one queen of E. hamatum in the physogastric condition. In each
case, field notes on the colony from which a given queen was
taken covered the general situation of the colony and its behavior
(especially its raids and bivouac-change movements) over a period
of days, the status of the brood or broods and the condition of the
queen at the time of capture. Each queen was kept alive in the
laboratory as long as possible after capture, for observation and
test both alone and in relation to the workers of her colony.

There is a notable disparity in our material as to condition of
the queen when captured. Why so many of the cases were in the
contracted condition and only one case physogastric when taken
will become clear as the general circumstances of Eciton life are
disclosed.3 For a number of reasons, physogastric army-ant
queens are exceedingly difficult to capture. There is only one
other case on record in which such an individual was observed
during her short span of life in captivity, that of a physogastric
queen of E. ( Labidus ) coecum taken by Weber (1941) in Trini-
dad. Fortunately we are not forced to depend upon direct evi-
dence concerning the queen at all stages, since an abundance of
other evidence coordinates nicely with results derived from work
on the queen herself.4

A descriptive contrast of queens and workers. — The Eciton
queen (Fig. 1) stands out as strikingly unique among fertile in-
sect individuals and as a highly distinctive member of her colony.
We may use the queen of E. hamatum as example.5 Unlike the
queens of virtually all other ants, she is wingless throughout life.
In color she is almost uniformly ferrugineous mahogany, darker
than the yellowish brown which characterizes the workers, and
lacks the cephalic paleness of major workers. She differs strik-

3 It is a striking fact that among 16 captured queens representing ten
Eciton s. str. species and sub-species, reported by various authors prior to
1942 (see e.g., Wheeler, 1921; Bruch, 1934), all were in the contracted
condition.

4 In the present paper the supporting evidence is reported rather concisely.
Further details and additional evidence will be incorporated in a monograph
on this subject, now in preparation with the collaboration of Dr. Harold R.
Hagan of the Department of Biology, City College of New York.

s The reader is referred to Wheeler Js systematic descriptions of the first
queens of E. hamatum (1925) and E. hurchelli (1921) to be discovered.

June, 1944]

Schneirla: Insect Behavior

157

ingly from the workers in gross size and in anatomical detail.
The greater bulk of the queen is suggested by the fact that her
over-all length (contracted) is close to 17.1 mm. (Av. 6 speci-
mens), whereas the range of body length in the workers is 2.5 mm.
(minor) to 9.5 mm. (major). The polymorphic worker types re-
semble the queen only very broadly, for example both queen and
workers possess well-developed and dentate tarsal claws, near the
inferior occipital corners of the queen’s head there are small
obtuse projections homologous to the acute spines of the worker
major, and the queen possesses strikingly prominent pairs of
epinotal and petiolar horns contrasting with two simple aligned
nodes in the worker. Among many impressive morphological
differences are the large lateral pseudo-ocelli (Werringloer, 1932)
of the queen and the relatively tiny ones of workers, the great
bulk of the queen’s thorax and her well-developed legs as con-
trasted with corresponding slenderness in the worker ’s structures,
and in particular the great size of the queen ’s abdomen contrasted
with the much smaller gaster of the worker. Associated with an
enormous expansion of reproductive functions in the queen and
the probable absence of such functions in the worker, the hamatum
queen’s gaster even when contracted measures near 9.1 mm. in
length and 5.5 mm. in its greatest width, whereas the gaster of the
largest major worker measures only 2.6 mm. in length and 1.8 mm.
in width. The queen’s gaster terminates distinctively in a large
triangular shelf, the hypogynium. Another striking character-
istic of the queen is the marked development of her external
respiratory orifices or stigmata, suggesting a high stage of de-
velopment in the respiratory system generally.

This general morphological comparison may suffice to illustrate
the extent to which the Eciton queen diverges from the workers,
paralleling a functional specialization which correspondingly
differs markedly from that of the workers. The nature and
extent of the dichthadiigyne ’s specialization becomes clear when
we consider some typical characteristics of her behavior and her
place in the life economy of the colony.

Behavior and functional 'properties of the “normal” or con-
tracted queen. — In E. hamatum and probably in Eciton s. str.
generally the reproductive capacity of any given colony is cen-

158

Journal New York Entomological Society

[Vol. LII

tered in a single individual, a fecund dichthadiigyne. A selective
mechanism of some kind evidently is involved which excludes ad-
ditional queens, possibly at a time shortly before or after young
queens are fertilized. Circumstances indicate that this one
fecund individual must pass around nine-tenths of her days in
the rainy season in the resting or contracted condition, which as a
consequence may be termed her “normal” state. Ordinarily she

Fig. 1. Queens of E. liamatum in the contracted (upper) and the physo-
gastric or gravid condition. ( Cf . Fig. 2 and Table I.) Total length of the
contracted queen, 17 mm. (Drawing by Miss Janet Roembild, from photo-
graphs by Dr. Virgil Argo, Dept, of Biology, City College of New York.)

does not leave the confines of the bivouac, except when she is
drawn into a bivouac-change movement, an event that scarcely
ever engages her before nightfall.

Circumstances indicate that the sequestration of the queen may
be due largely to the effect which light exerts upon her. In labo-
ratory tests she reacts specifically to directionalized bright light

June, 1944]

Schneirla: Insect Behavior

159

by turning away from the source and moving energetically about
until in darkness. Whenever she chances into the light, facing
toward the source, the queen volte faces abruptly and runs off.
(Among numerous observations, this test was repeated in one
instance 12 times in 30 minutes with the same queen, — always
with the described result.) When illumination is not intense
the response is more variable, yet even then as a rule the queen
settles down only when in a fairly dark part of the nest.

In contrast to the queen, the workers seem virtually incapable of
specific orienting responses to light. They are aroused to activity
by light after a period of darkness; thus colonies are regularly
stimulated to begin their daily raiding after dawn (Schneirla,
1938; 1940). This however is a mere photokinetic effect ( i.e
a generalized excitation by light), and they appear incapable
of orienting to directionalized illumination except in a very crude
fashion as an outcome of restless movements over a considerable
time. That the basis of this outstanding difference between queen
and workers rests in different optoneural equipment is suggested
by the fact that in E. hamatum the gross size of the queen ’s lateral
pseudo-ocellus is more than twice that of the major worker’s.
We should expect to find correspondingly accentuated differences
in the internal structure of the visual receptor, and in related
mechanisms, underlying the queen’s pronounced normal reac-
tivity to light.6

This recalls the fact that the fertilized queens of ants in general
are repelled by light. Whether the Bciton queen, like any others,
is positive to light prior to fertilization cannot be said at present ;
however, the sharpness of the functional queen’s avoidance re-
sponse is clear. The basis of a highly adaptive circumstance is
thereby provided — that the queen is cloistered in a well-protected
situation except when the colony shifts its home site. The prev-
alence of this condition is attested by the fact that Eciton queens
are never seen at other times, as during raids — its effectiveness is
emphasized by the difficulties one experiences in capturing these
queens.

6 Although no queen material was available for Werringloer *s (1932) valu-
able histological study of Eciton visual mechanisms, species differences were
found in optic histology which appear to parallel typical behavior differences.

160

Journal New York Entomological Society

[Vol. LII

That the queen keeps to the bivouac in the daytime, during
raids, is not attributable to any inability to follow the colony’s
trails. In the laboratory, captive hamatum queens may run for
hours at a time in a column following a circular path established
by workers or, when one is set down near a vacated trail of her
own colony in the forest, she follows the route successfully once
she hits upon it. Evidently it is her “photophobic” proclivities
that mainly account for the queen’s sequestration in the bivouac.

Although for a long time the idea has held sway that the army-
ant queen is moved passively about by the workers when the colony
shifts its site, in Eciton s. str. the facts are contrary to this sup-
position. In laboratory nests the queen readily makes her way
about despite the usual added burden of several workers hitch-hik-
ing and dragging from her legs or gaster. The physical char-
acteristics of her well-developed legs give every indication of great
strength, as does her robust frame in general — an impression
borne out fully by her behavior. The test comes during the long
bivouac-change march in the wild when the queen must make her
way over a route generally much longer than 100 meters, passing
along a narrow and tortuous trail full of hazards such as vine
bridges and sharp turns although she is swarmed under at nearly
every step by a frenzied crowd of workers.

On a number of occasions the hamatum queen’s participation
in the colony movement has been observed, always at night except
in two instances when colonies were greatly oversize and the move-
ments unduly extended. The bivouac-change usually gets under
way before dusk and is completed during the night, with the
queen coming along during the latter part and frequently near the
very end of the trek. The explanation seems to be that she is
normally stationed in the innermost recesses of the cluster, cen-
tered in the brood with a mass of workers minor around her, in a
section of the bivouac which is likely to be almost the last to be
drawn into the evacuation.

Just before the queen appears from the bivouac there is a very
noticeable increase in excitement among the workers on the trail,
agitated variable movement becomes common instead of the
monotonous plodding seen previously, and the column begins to
widen perceptibly from the 2-3 cm. which is characteristic. As

June, 1944]

Schneirla: Insect Behavior

161

the column becomes more crowded it broadens to as much as 15
cm. within a few minutes, then the queen appears in the thickest
part. As the queen moves along she is not only surrounded by
jostling workers of all sizes, but much of the time is literally cov-
ered with them, has them underfoot and hanging to her. The
workers hamper her movements particularly when she slows in
mounting steep grades or in rounding sharp corners ; at such times
they may crowd around so that she is actually stopped for a time.
The remarkable fact is that amid this great excitement and
arduous labor the queen is able to follow the trail in its devious
windings and through its difficult sections.7 Tests show that this
ability depends upon a response to Eciton chemical. For exam-
ple, if the queen is taken up and returned to the route when there
are no workers upon it, she follows it readily. There can be little
doubt about the fact that the queens of Eciton s. str. participate
quite actively in the bivouac-change movements of their colonies
and get to the new site under their own power.

The adaptive significance of the workers’ behavior toward the
queen during the lengthy migratory trek deserves mention. Since
at all times a high pitch of excitement is maintained in the queen ’s
section of the march, any intrusion is immediately responded to
by large numbers of frenzied workers so aroused that they bite
and sting with very little provocation. In the darkness of night,
the feverish activity that prevails in the few meters of column
near the queen contrasts strongly with the lethargic monotonous
lock-step movement that prevails throughout the other sections of
the movement. Only in the entourage of the queen are the work-
ers aroused to a degree of excitement and responsiveness that
resembles their daytime activity in raiding. The outcome is that
the colony’s reproductive specialist is well protected at the time
when exposure is great.

The strength and vitality of the queen are shown impressively
when the colony is etherized for capture. Because of the agility
and speed with which she is capable of moving when disturbed,
the queen generally is able to make her escape from the bivouac
while the principal mass of workers succumbs to the anaesthetic.

7 Reichensperger (1934) observed similar occurrences in a bivouac-change
movement of an E. lucanoides colony in Costa Rica from which he captured
the queen.

162

Journal New York Entomological Society

LVol. LII

Thus she is frequently discovered under a leaf or piece of bark
near the outer edge of the sheet used to imprison the bivouac, still
somewhat active although workers in numbers lie prone around
her, and if the way is not blocked she may make good her escape
along a raiding trail.

Wheeler (1921) has remarked upon the possibility that an ex-
tensive development in the respiratory system of the Eciton queen
is an important adaptation to a sequestered bivouac life. The
pronounced development of this system in general is indicated by
the prominence of the spiracles, the external openings of the
breathing tubes. With a highly efficient mechanism for gaseous
exchange, it is possible for the queen, despite her huge bulk and
relatively great oxygen needs, to remain for long times within the
close air of the bivouac without any apparent detriment to health.
Furthermore, the large size of the main tracheal vessels of the
abdomen and the extensive ramifications of these vessels among
the reproductive and other visceral organs indicates that the queen
is well equipped in this respect to meet the crucial demands of a
greatly increased metabolism during her brief gestational periods.

Although nothing is known concerning the Eciton queen’s nor-
mal span of life with her colony, in all probability it is equal at
least to one rainy season. However, the hamatum queen for some
reason is particularly short-lived after removal from the midst
of her colony. Three of our 13 contracted specimens lived only
two days and only one lived longer than seven days in captivity
after capture (see Table I). As a control, one queen (1938 B)
was permitted to remain with her colony for more than two
weeks in the laboratory, then was removed (without ansesthesis)
in good condition, yet this queen died within four days after she
was segregated with a small group of workers. Removal from the
colony thus seems to introduce some change which makes in-
evitable the early death of captive Eciton queens. Although in
this study various preliminary attempts were made to discover
the nature of the lethal factor, the results were not sufficiently
clear to warrant discussion here.8 Whatever this factor may be,

8 A possibility worth testing is that the Eciton queen when isolated from
the mass of her colony is deprived of some essential food substance. For
instance, Zahl (1939) has found that workers and queens of the tropical
ponerine ant Dinoponera grandis sicken and die after a few weeks in cap-
tivity unless a larval brood is present in the colony.

June, 1944]

Schneirla: Insect Behavior

163

it appears to be specific for queens and not for workers, since in all
cases workers remained alive in captivity for weeks after their
queens had died.

From the above description of workers’ responses to the queen
during the bivouac-change movement, it is apparent that she must
exert a powerfully attractive stimulative effect upon them. The
basis appears to be mainly chemical. Even to the human observer
the queen is distinguished by a delicate, fragrant odor, quite un-
like the heavy, somewhat foetid odor characteristic of Eciton
workers. Workers are especially attracted to pieces of cardboard
formerly in use as floors for the queen’s cell, and will collect in
the one of two compartments in which a queen has rested for a
time. In artificial nests they gather about her, and follow closely
as she runs about. When she comes to rest they pile the brood
around her, and spend much time licking her and stroking her
with antennae. Our queens, placed in small wire cells at the top
of the large cylindrical nests in which their respective colonies
were clustered, were visited by large numbers of minim workers
which penetrated the fine screening and remained to cluster in the
cell. Thus the queen may be considered a factor of some impor-
tance in normal trophallaxis (Wheeler, 1928), the stimulative
interrelationship of individuals which provides the basis for social
organization.9

Characteristics and behavior of the physogastric queen. — At
the time this study began a number of queens of Eciton s. str.
species had been captured in various parts of Central and South
America: notably E. burchelli by Wheeler (1921) in British
Guiana, E. vagans by Gaige in Colombia (Wheeler, 1921), E.
hamatum by Wheeler (1925), and E. lucanoides and E. mattogros-

9 However, it is doubtful that the workers in general are widely sensitive
to the actual presence or absence of the queen as Wheeler (1921) implies they
were in the case of a colony of E. burclielli from which the queen was re-
moved. “I infer,” he says, “that this was the only remaining female in
the colony, for after her removal a perceptible apathy or dejection seemed
to fall on the whole body of ants” (p. 298). Actually, this effect of
1 1 apathy, ’ ’ specifically a rather abrupt drop in general activity, characteristi-
cally ensues after large numbers of the ants have been stirred up and then
permitted to recluster. Once reclustered, they fall into a lethargic condition
whether or not the queen has been removed, hence this behavior cannot be
attributed to a removal of the queen.

164

Journal New York Entomological Society

[Vol. Lll

sense by Reichensperger (1926, 1934) and E. quadriglume and
E. rogeri in the Argentine by correspondents of Carlos Bruch
(1934). Thus by 1934 the queens of nearly all of the species of
Eciton sensu stricto had been taken. It is interesting to note that
in all of these cases the queen’s gaster was contracted, a circum-
stance which led various writers ( e.g Bruch, 1934) to conclude
that the queen in question was “young,” or “virgin.”

The experience of the present writer was similar in capturing
Eciton s. sir. queens in Panama for behavior studies. In 1933 two
queens, E. hamatum and E. burchelli, were taken; in 1936 four
queens, one of E. lucanoides and three of E. hamatum, and in
1938 nine more of the last species, all of them alike in the fully
contracted condition (see Table I) . Prom these and other facts it
becomes apparent that few if any of the queens of Eciton s. str.
reported in the literature could have been virgin when captured,10
and that the dichthadiigyne must be capable of returning to the
contracted state after having delivered eggs. This interpretation
in fact was offered as one alternative by Wheeler (1925) in report-
ing the first capture of the queen of E. hamatum.

Early in the present investigation, evidence concerning periodic
changes in colony behavior, together with the experience of cap-
turing numerous queens singly from colonies with broods in all
stages of development, strongly indicated that the queen must
pass repeatedly from the contracted to the physogastric condition,
remaining in the latter condition only briefly before returning to
the contracted state. Finally it became clear circumstantially
that the eggs of a given brood must be laid during a short period
of a few days near the end of the first week of the statary phase
in colony behavior, when the colony is non-migratory.

Following this conclusion, in 1938 an effort was made to capture
a hamatum colony at the critical time, which on circumstantial
grounds was placed about seven days after the beginning of the
statary period. On July 21 a colony (record number 38 H)
was found which although nomadic at the time was evidently
nearing the statary phase, judging from the fact that its larval
i° Probable exceptions are the two queens of E. burchelli taken by Wheeler
and Emerson at Kartabo, British Guiana, in 1920 (Wheeler, 1921), in which
circumstances point to the recent emergence of at least one and perhaps both
of them from cocoons.

June, 1944]

Schneirla: Insect Behavior

165

brood seemed very advanced and about ready to spin cocoons.
After three further bivouac-change movements on successive
days, when spinning appeared to be mainly finished in the brood,
the colony settled down at a spot near station 4, Barbour trail.11

The statary bivouac was formed in a deep cranny near one end
of a decayed fallen tree-trunk. The cluster formed a long ellip-
tical curtain-mass filling the opening of the niche. On each of the
following days, when the colony was revisited, the ants had only
one raiding-system, a typical sign of the statary condition. The
site was revisited on July 30, i.e., one week after the beginning of
the statary period, with materials for capturing the colony.

The colony had withdrawn about 30 cm. farther into the recess,
very probably in response to frequent spattering by rain. Fortu-
nately it was still fairly accessible. The bivouac was quiet, its
forward wall sprinkled with ants holding cocoons in their mandi-
bles. A dampened sheet was fastened securely across the open-
ing to entrap the ants, then was sprinkled with ether. After
four minutes, when the covering was removed, the major portion
of the colony, a mingled heap of ants and cocoons on the floor
of the cavity, was quickly scooped into a large jar. A few strands
of ants still hanging, together with masses of eggs and clusters of
small workers from the rear of the bivouac, were put into a second
jar. In examining the material promptly after returning to the
laboratory the first jar was found to contain the major part of the
large enclosed pupal brood as well as most of the worker popula-
tion. In the second jar, which contained workers and egg masses
from the rear strands of the bivouac, a phy so gastric queen (Fig.
1) presently was brought to light in the part of the mass where
workers minor and eggs were most numerous. Judging by rela-
tive positions in the jar, this queen must have occupied a position
near the back of the cluster, probably among the hanging strands
which contained most of the workers minor and most of the eggs.
A careful inspection of the material revealed no other queens, no
males, and two broods — a large enclosed pupal brood and an enor-
mous clutch of recently delivered eggs.

Within the following twenty-four hours, which proved to be the

11 The queen was seen in the procession during the final bivouac-change
movement, and was judged to be in the contracted condition at that time.

166

Journal New York Entomological Society

[Vol. LIl

span of life remaining to the 38 H dichthadiigyne, her behavior
was studied as continuously as other duties permitted. —

A few minutes after she was removed from the jar (4: 30 p.m., one hour
after capture) the queen appeared to be recovering from ansesthesis, as indi-
cated by reflex twitchings of tarsi and antennas, although most of the workers
remained immobile. At 4 : 45 p.m. she had recovered sufficiently to gain an
upright posture and had laid 20 eggs. At 5 : 10 p.m. she was running about
the Petri dish in which she had been placed, despite the trammeling effects
of her huge distended gaster which tilted to one side or the other as she
moved. From rough measurements at that time, her gaster was 16.5 mm. in
length.

Until 8 : 15 p.m. the queen remained undisturbed in a darkened Petri dish,,
laying more than 300 eggs. Then for an hour she was exposed at intervals
to brilliant photoflood light when motion pictures were taken. Despite evi-
dent disturbance from the light, marked by shock reactions when light was
introduced and by turning from the source, there was a describable regularity
about the queen ’s behavior during this interval. There were successive
periods of a few minutes each in which the queen stood in place, observably
engaged in egg-laying, with intermittent periods in which she ran about the
dish more or less continuously, dragging or carrying numerous workers on
her gaster and followed closely by others. At no time was she moved by the
workers, although generally a number of them remained close to her whether
or not she was in motion.

The following representative notes were taken during two of the quiescent
intervals, the first lasting 5 minutes after 9: 37 p.m. and the second 2 min-
utes after 9: 44 p.m.:

The queen abruptly breaks pace and suddenly stops in place, headed away
from the photoflood source. She stands firmly on all six legs, with the
ventral surface of the gaster resting on the floor. In oviposition, the eggs
emerge in a wide jet from the vaginal orifice, and spread out radially upon
the hypogynium. The process is marked by slight laterad oscillations of the
the gaster and a shivering of the body and legs, which cease when eggs no
longer emerge. — Eggs were laid during both of these halts.

At 11 : 30 p.m. the queen with 6 workers was placed in dim light for obser-
vation. During the next 90 minutes there were 11 quiescent periods, varying
between 30 sec. and 11 min. in duration (6 of them lasting between 1 and 4
min.). The intervening periods of activity were shorter, most of them
around 3 min. in duration. In each case, locomotion began rather abruptly.
The quiescent periods likewise began abruptly, with the queen usually sprawl-
ing close to the floor at once. Generally she stopped in a position facing
away from the light, which although dim was sufficient to orient her. In
some of the shorter intervals no eggs were laid. In the course of one or two
of the longest stops, the queen shifted position briefly or moved forward a
short distance once during the interval. The workers remained close to her,,
frequently applying their mouth parts to her body at the vaginal orifice and

June, 1944]

• Schneirla: Insect Behavior

167

at the surface of the hypogynium and licking these surfaces. After one of
the stops a drop of viscous greenish-yellow fluid remained where the tip of
the queen’s gaster had rested. The workers soon found this drop and applied
their mouth parts to it, evidently feeding. — Observations were discontinued
at 1: 30 a.m., since no eggs had been laid during the preceding 30 minutes. —
Between 5: 10 p.m. and 12: 00 a.m., 1245 eggs were laid.

In the morning, at 6: 30 a.m., only 358 eggs were found to have been laid
during the night ( i.e ., after 1: 30 A.M.). As before, alternating periods of
quiescence and of activity were observed. Between 8: 15 and 9: 54 a.m. the
queen was shielded by a ruby-glass filter and remained under observation.
During this interval there were sixteen quiescent periods alternating with
intervals of continuous locomotion, virtually all of the phases of quiescence
and of activity falling between 1 and 4 min. in duration. In some of the
stops between 20 and 30 eggs were laid, in others no eggs. The following
notes typify egg-laying behavior:

8 : 23 to 8 : 26 a.m. — The queen stops abruptly, facing away from the weak
light. Promptly there begins a shivering movement of the gaster, increasing
at times to an oscillatory movement ; then the oscillation becomes continuous,
at times reaching 1 mm. in amplitude. These oscillations of the gaster begin
locally and increase in amplitude, meanwhile spreading to include the entire
body with the legs participating. Near the height of such spasms the eggs
begin to appear. A jet of eggs between 12 and 15 units wide is forced slowly
from the vagina directly backward over the hypogynium, with the eggs
fanning radially to each side as they emerge. The eggs in small packets are
either picked off by the workers or drop to the floor when the queen moves
away/ — Finally the queen breaks abruptly into motion, in an interval of loco-
motion which lasts 4 minutes in this case.

At 4 : 15 p.m. the queen seemed less energetic than in the morning, and had
laid only about 60 eggs since 10: 00 a.m. For about 20 minutes she was
exposed to bright sunlight while photographs were taken, and although she
was shaded between exposures the intense light appeared to exert a decidedly
injurious effect upon her. At length she fell upon one side with flexed legs,
unable to regain an upright position. At 6 : 00 p.m. there were unmistakable
signs of approaching death, which in the contracted queen is characterized
by intermittent struggling and by local tarsal and antennal reflexes lasting
a number of hours. — At 6 : 30 p.m. she was chloroformed for fixation in
Carnoy’s solution and eventual preservation in 70% alcohol.

Notwithstanding the fact that disturbances incident to labora-
tory captivity must have disrupted the normal oviposition
mechanism greatly, and although the factors which character-
istically hasten death in captive Eciton queens must have been at
work, certain inferences concerning the normal circumstances of
physogastric function may be drawn from these observations.
Concerning the queen herself, the regular occurrence of short

168

Journal New York Entomological Society

[Yol. LII

intervals of action and of quiescence (and egg-laying) in alterna-
tion indicates that a rhythmic process underlies the delivery of
eggs from the ovarioles. The abrupt onset of each quiescent
period together with the oscillatory movements of the gaster and
at times the entire body during oviposition speak for the opera-
tion of a fairly distinct egg-laying process arising at short inter-
vals. Likewise, the abruptness with which the queen broke into
motion after each quiescent period indicates the regular recur-
rence of a rhythmic change in this process, evidently a phase dur-
ing which more eggs became viscerally available for delivery.

Under the conditions of these observations the queen was at-
tended by only a few workers, and space permitted her to run
about in the intervals between egg-laying stops. It is probable
that under normal conditions in the bivouac, when she is sur-
rounded by masses of workers, free locomotion is out of the ques-
tion for the queen. In that situation her viscerally-stimulated
overt activity must be reduced to stirring in place and to inter-
stimulative relations with workers.

This queen died with a considerable portion of her eggs still
undelivered, a fact which together with other circumstances indi-
cates that in all probability lethal processes setting in with cap-
tivity impaired the egg-production rhythm seriously from the
beginning. Under normal conditions in the bivouac this rhythm
presumably functions smoothly and continuously throughout the
period of approximately four days in which all of a given clutch
of eggs is delivered. Then, as we shall see, circumstances are such
that the queen, contracted once more but in an exhausted condi-
tion, may profit by a long interval of recuperation before the
parturitive ordeal must be repeated.

Our results shed some light upon the behavioral relation of the
physogastric queen to her colony. First of all, the fact that the
queen is quite capable of locomotion during her time of physo-
gastry effectively negates the hypothesis (Muller, 1886) that the
colony must remain in place at such times because of an inability
to drag along the heavy egg-laden queen. She moves readily, and
even drags or carries workers about with her, hence there is reason
to believe that she would be quite capable of joining a bivouac-
change movement if one happened to get under way. She would

June, 1944]

Schneirla: Insect Behavior

169

have to make periodic (viscerally enforced) stops on the trail,
but could make the trek even though it might well be fatal. We
shall find that the sessile (i.e., statary) status of the colony when
the queen is laying her eggs is indeed related to the queen’s
oviposition cycle, but very indirectly, and very differently than
the above hypothesis would suggest.

Our observations indicate that the normal stimulative attrac-
tiveness of the queen for the workers (i.e., her trophallactic rela-
tionship with workers) is considerably increased during the time
of oviposition. The intersegmental membranes of her enormously
swollen gaster and the hypogynium and vaginal regions in par-
ticular are highly effective centers of attraction. The workers are
constantly crawling upon her and licking these regions as well as
the eggs when they appear (cf. Wheeler, 1900) and the (evidently
much relished) small drops of fecal material which issue at times
after egg-depositing episodes.12

There can be little question that this queen produced all of the
eggs taken with the colony, a total of 17,062, which was estimated
to be two or three thousand short of the actual number laid prior
to capture. To this we may add 2,046 eggs laid while the queen
was held captive, and 7,190 unlaid eggs taken from her gaster in
post-mortem study. The total of 26,298 eggs approaches fairly
close to the size of Eciton brood populations for which census
studies have been made in other cases, offering further evidence
that one individual dichthadiigyne is capable of delivering single
broods of that size. In view of circumstances indicating that the
eggs of each huge brood are laid within a short interval of prob-
ably no more than three or four days, the entire performance
seems quite worthy of being termed a stupendous feat.

The great susceptibility to death of the Eciton queen in this
condition is attested by the fact that our specimen survived only
about 30 hours after capture. It is probable of course that her
demise was hastened by the anassthesis and by other shock-effects
( e.g ., from intense light during photography), yet the fact that
contracted queens similarly treated usually remain alive for as
long as a week in captivity speaks for a specific physiological
vulnerability to non-optimal conditions in the gravid Eciton

12 Emerson (1939) has reported a similar observation for queen-worker
relationships during egg-laying in termites.

170

Journal New York Entomological Society

t Vol. LII

queen. It is a fact of great adaptive significance that during this
biologically critical time the dichthadiigyne ordinarily is assured
the essential optimal situation through the operation of intrinsic
factors which render the colony sessile. No less remarkable is the
fact that the queen herself , through her peculiar reproductive
properties , is indirectly the basic controller of this state, of affairs.

EVIDENCE FOR A FUNCTIONAL RELATIONSHIP BETWEEN
BROOD AND COLONY

Significant periodic differences in colony behavior. — As the
first step toward working out relationships between the queen and
her colony, let us examine the general situation of the series of E.
hamatum colonies from which queens were captured for this study
(see Table I).

In their behavior twelve of the hamatum colonies (i.e., colonies
33 A; 36 A, B, D, and G; and 38 C, X, D , E, F, G, and I) from
which queens were taken conform to the pattern previously de-
scribed as “ nomadic’ 9 (Schneirla, 1938). On the other hand
only three (i.e., colonies 38 A, B, and H) conform to the pattern
described as “statary. ” Since all of these colonies were studied
for at least three days prior to capture, and some of them for
longer times, any important deviations should have become ap-
parent ; but in all cases there appeared very clearly the character-
istics of one or the other of the described activity patterns.

In cases showing the nomadic pattern, all of the colonies were
migratory, that is, they had all engaged in bivouac-change move-
ments near the end of each day in the period preceding capture.
Without exception among many observations, the raiding activi-
ties of a given day were terminated by a complete movement of
the colony, generally during the evening and early night, so that
the next day found the colony developing a new raid from a dif-
ferent site generally more than 100 meters removed from the last.

Although their bivouacs were situated in rather different topo-
graphical circumstances, the clusters of the “nomadic” colonies
were not secluded as is typical of statary colonies. The former
almost without exception established themselves beneath logs,
under matted vines, or against the sides of trees between but-
tressed roots, seldom within cavities such as hollow logs or trees.
As a rule the larger portion of the more or less cylindrical mass

Data Relevant to the 15 Queens of E. hamatum Captured in the Present Study, and to the Colonies from

Which These Queens Were Taken ( cf . Fig. 2)

June, 1944]

Schneirla: Insect Behavior

171

O

3 £
© &
O ©
ao

nO nS
od cC

a a
, © ©

.2 fl ©

cS c§ d »

O © 03 O O ©

© ©
^3 ^3

>5

e

■K>

e

$2

v.

e

e

£2

&1

£

<0

OD

© ©
bfi be
oS d

a s I 1

ro • «© • M -oo

>. g 5© go go

Sol

W- £H W g V.M • (.TJ

a i a i a i a i | a

b • oo

b a «

. CO
b t-

g’

bp

b o

S

^ (Tf J XT \T J L'* -^T UNJ w»j ^ L'* 1 1 J W NJJ

f>» r^s ^o'do'ddddHdrioHNMO'd
£ ££ £ > H II II II II II II II II II II II II II II II

be

be

00 ©

« a

o ©

©

© 03
© «M
© Oh £•"»

°V.£

«H O >

'S'S*5

3 ©
o

cc cc cd b c3

d

d

d

rr-y

d

d

d

d

jb

r© '"C nO '©S r©

nO

n©

no

no

no

o

© CO Tji b- S .

CO

CO

o

t-

5©

i o'?

fH 53 U

w

•g

,<0 no no no no

nO

no

nO

no

no

no

nO

^.no 03

0} <D 0) Qp

4^> 4^ 4-3 4-3 4-3 4-3

O O O O O O

c2 o3 oj ct cd

£ £ rt a « rt
o o o o o o
QOOOUO

f-i Fh Fh >H SH

§£ -g t *
oo a g£

j^-g ^
£ 6

p© _, ^ , rrt

£ § £ ^ ©

? o a

© &%<£ a

l « s

o TtH 00 00 cq

0 05 00 050©

<1<1pqpO<J

CO O 00

CO CO CO

Oi os 03

PQ

q H q $ M w Hs

* The growth stage of the respective broods taken in 1938 is represented by data on the range (E) and the mean (M) of
body lengths (samples ranged between 25 and 200 specimens from each colony). The writer wishes to express his thanks
to Miss Ruth Greene, who generously contributed her time to gather these data.

The larvae of various Eciton species have been described by G. C. Wheeler (1943) with a key for their identification.

172

Journal New York Entomological Society

[Vol. LIT

of ants was fairly exposed to view. It is typical that the bivouacs
of nomadic colonies are far more readily approached and observed
than are those of statary colonies.

In their raiding, particularly, colonies in the two behavior con-
ditions exhibited very different characteristics. In the nomadic
condition the Ecitons always staged vigorous daily raids which
began promptly with the first daylight and grew rapidly into the
characteristic maximal raiding pattern of the species. In E.
hamatum this is marked by the growth of two or three (generally
three) principal raiding systems, each a tree-like pattern of
trails with a single principal trail as its line of communication
with the bivouac. Although of course the details of the trail
systems vary considerably according to the situation of the
bivouac and the general topography, invariably multiple trail-
systems are found under nomadic conditions. At such times a
maximal number of ants from the population is drawn into the
daily foray, developing extensive and complexly branched trails
on a widespread front of raiding. With the raid probing out
along a number of principal lines, the usual result is that roughly
three-fourths of the circular zone around the bivouac is invaded,
with the consequence that relatively enormous quantities of booty
(mainly the soft-bodied young of other insects) are gathered in.
While nomadic , a colony remains near its peak of vigor in raiding
and in other activities.

With those colonies in the statary condition when their queens
were captured the case was quite different in all important
respects concerning behavior. Colonies 38 A, B, and H were
all known to have remained in their same bivouacs at least three
days prior to capture. Colony H, for example, after three ob-
served daily bivouac-changes, clustered within a large crevice
in the side of a massive log on July 22, 1938, and was found in
the same place and in almost the same spot on July 30, when
captured.

Other evidence has been offered (Schneirla, 1933, 1940) for
the existence of the statary period as a distinctive 4 1 sessile ’ ’ phase
of Eciton life. Observations on particular colonies extending
over considerable periods have shown that once a hamatum colony
enters a statary phase, it remains bivouacked in the same place

June, 1944]

Schneirla: Insect Behavior

173

and exhibits other characteristic “statary” features of behavior
during a period of approximately 19 days, before a major change
occurs.

The fact will be recalled that of the fourteen colonies from
which queens were captured in this study, only three were taken
in the statary condition. This difference is scarcely a matter
of accident, but is due to the relatively secluded locations taken
by colonies of Eciton s. str. when they are statary. Colony 38 H,
clustered in a deep crevice on the outer wall of a huge log, was
unusually accessible for a statary colony. The virtual impregna-
bility of statary colonies is attributable to two facts in particular.
First, they are commonly clustered within a cavity, generally
in a hollow log or tree. Colony 38 A had to be extracted from a
hollow log, colony 38 B from within the large hollow root of a
standing tree, by drilling a circle of holes with brace and bit
and breaking out an opening through which the ants could be
removed by hand.13 Then, too, if a statary colony happens to
settle where it is somewhat exposed to the elements, disturbance
from wind, rain or sunlight causes the ants to shift their position
so that after a few days capturing them would require the assis-
tance of a wrecking crew. The proneness of colonies to cluster
in hollow logs or trees when entering the statary period stands
as a highly adaptive circumstance for which no specific explana-
tion can be advanced at the present time.

Without exception, each of our three statary colonies from
which queens were obtained had a single raiding system when
captured. This pattern is readily recognized by the presence of
just one principal raiding trail leading from the bivouac, rather
than two or three as in the nomadic phase. Distance reached
from the bivouac in the raiding is not a secure criterion of its
relative vigor, since in the statary phase the chemically-marked
route of a previously used trail may be employed on more than
one day so that such trails often become greatly extended beyond
the distances ordinarily reached in nomadic-phase raiding. As
more certain indications of reduced raiding, the number of prin-
cipal systems is one rather than three, and there is a smaller

13 Dr. Neal Weber kindly assisted in this operation, which was completed
in a torrential rain.

174

Journal New York Entomological Society

[Yol. lit

number of branch trails in that system than is characteristic of
' a raiding system in the nomadic phase. Thus, in the statary
phase, raiding activity falls much below the level of vigor and of
numbers involved during the nomadic phase.

Before considering the relationship of these periodic differences
in Eciton behavior, it is desirable to examine correlated differ-
ences in the internal constitution of the colony.

Intrinsic conditions underlying colony behavior. — A considera-
tion of the internal characteristics of the colonies (see Table I)
shows that the external behavior differences were paralleled by
others no less outstanding. The intrinsic differences may be
represented by a comparison of the two groups of colonies in
terms of their condition and the status of their brood or broods
when the queens were captured.

First of all, it should be emphasized that the broods invariably
were worker broods, and that no batch of larvae or pupae con-
tained individuals departing sufficiently from the others in form
or size to suggest the anlagen of fertile forms. This statement
is of course not based upon our present facts alone, but finds its
main support in an examination of hundreds of Eciton broods
on Barro Colorado Island in rainy months between May 15 and
September 20 in four different years which has disclosed none
but worker forms in them.

Another fact of significance is that while each of the broods
contained tens of thousands of individuals, all members of a given
brood were roughly at the same stage of development. This con-
dition held not only when a single brood in the larval stage was
present, but also for cases in which there were two broods (i.e.,
a pupal brood in addition to a brood in the very early larval
condition). This statement, likewise, is based not only upon the
present data but also upon the invariable finding that in E.
hamatum the entire population of a given mature larval brood
terminates that stage and spins cocoons within a very few days,
and that pupal broods mature and are removed from their cocoons
as callows within a similarly short period of three or four days.

All three of our statary colonies had two broods at well-sepa-
rated points of development, a fact which we shall find character-
istic of the latter part of this activity phase. Colony 38 H con-

June, 1944]

Schneirla: Insect Behavior

175

tained a well-advanced pupal brood as well as a massive batch
of newly laid eggs. Colony 38 A contained a still more advanced
pupal brood, and its younger brood had largely hatched and
passed into the early larval period. The two broods of colony
38 B corresponded in general to those of 38 A. In the early days
of the statary period only one brood (always a pupal brood)
is present.

In contrast, the nomadic colonies each contained only a single
brood, in every case in the larval condition. From the size data
on these broods, represented in Table I by the averages and ranges
of individual body lengths, it is apparent that the broods were
at very different stages of development when the colonies were
captured. If a greater magnitude of body length in the brood
(as indicated by larger averages and greater ranges) may be
taken as valid evidence for an advanced point in growth, the
nomadic colonies captured in 1938 may be placed in the following
sequence : X , C, D , E, F and G. It will be noticed that with the
single exception of colony X, there is a correspondence between
the chronological order of capture and the growth point attained
by the respective broods. The possible significance of this fact
will be considered in a later connection.

When we bring together our facts concerning the external and
internal characteristics of the colony, a significant parallelism is
evident between them for both the nomadic and the statary
phases. In the statary phase 1) the colony does not migrate
but remains in a given place, 2) by virtue of its secluded location
it is well protected from the elements and from animals, 3) it
raids minimally and is comparatively lethargic, and 4) it con-
tains a pupal brood and in its latter part ( e.g ., colonies 38 A,
B, and H) eggs or a young larval brood as well. The most sig-
nificant parallel circumstances are a low condition of colony
activity and a brood {or broods) incapable of overt activity.

In sharp contrast to the above situation, a colony in the no-
madic phase 1) regularly moves to a new bivouacking site at
the end of each raiding day, 2) generally bivouacs in fairly open
places, 3) raids maximally and appears to be always {i.e., in
the daytime) near the peak of activity, and 4) contains a single
brood in a more or less advanced larval phase. In this case, the

176

Journal New York Entomological Society

[Vol. LII

most significant parallel circumstances are a high point in colony
activity and the presence of a larval brood which is capable of
overt activity .

Theoretical explanation of the parallelism between brood con-
dition and colony behavior. — These facts suggest that a causal

DAYS

Fig. 2. Schema to represent concurrent events in colony behavior (top),
condition of the brood, and function of the queen in E. hamatum. (The
sequence of given events in the colony may be traced horizontally, the con-
currence of events, vertically.) A-H, approximate situation of the respec-
tive colonies from which queens were captured in 1938 ; Ph, physogastric
interval of the queen; E, egg mass delivered in given physogastric interval;
L, larval stage of brood; P, pupal stage of brood; £ 2J, workers minor and
major of a callow brood.

relationship exists between the condition of the brood and the
circumstances of general activity in the colony. The conclusion
appears justified that conditions arising from the capacity of the

June, 1944]

Schneirla: Insect Behavior

177

brood for overt activity somehow account for the critical differ-
ence which appears in general colony activity pattern when active
broods rather than “passive” broods are present. This idea
has been advanced in a previous connection (Schneirla, 1938)
and will be carried further presently.

Evidence from field observation and special test supports this
inference of a causal relationship dependent upon condition of
the brood. Batches of eggs and very young larval broods (as in
the statary colonies 38 A, B, and H) are as a rule gathered itito
a single mass or a few masses which are covered and at times per-
meated by minim workers and are only very indirectly in contact
with the adult population of the colony. On the other hand, when
the larvae have developed further (as in colonies 38 C, D, E, F,
G, and X) and their twisting movements become perceptible to
the unaided eye, these masses are broken up and more or less dis-
tributed through the bivouac, so that larval activity and other
stimulative effects may directly reach large numbers of adult
workers. When the larvae mature, and become enclosed in co-
coons, it is apparent that in the ensuing pupal stage (as in col-
onies 38 A, B, and H) they become passive contents of the bivouac.
Heaped in interior pockets of the cluster and held by workers
distributed through its wall, they are presumably neutral in im-
portance or may actually exert a quieting effect upon the work-
ers that clutch them in their mandibles, somewhat as the pressure
of thumb-in-mouth pacifies an infant.

From laboratory observations we may say that through the
presence of an active larval brood the workers are distinctly
stimulated so that the amount and the vigor of their activity in-
creases greatly. When Ecitons are divided into part-colonies of
comparable size, it is the groups with larvae that show the greater
activity both within the confines of their artificial nest and in the
frequency of visits to an adjoining food-place, as compared with
groups that have no brood or have a pupal brood in cocoons.
When we study the relationships of individuals within the nest,
there are clear indications that this difference in activity level
actually depends upon a stimulative effect from the larval brood.
Typically the larval brood is spread over an area, usually around
the queen if one is present. The workers move restlessly around,

178

Journal New York Entomological Society

[Vol. LII

frequently touching the larvge with mouth-parts or licking them,
intermittently touching and stroking them with antennae, often
picking them up and shifting their position slightly or carrying
them bodily from place to place. When pieces of food are car-
ried into the innermost nest area, they are commonly deposited
near the larvae or actually laid upon them, so that in their almost
incessant twisting movements the larvae readily bring their mouth
parts into contact with the food.14 It is a common observation
that stimulation from passing workers may arouse quiescent larvae
to squirming activity, or that chance stimulation from larvae may
bestir workers. It is manifest that the larvae thus in a number
of ways arouse the adults to increased activity, in the mutual
stimulative relationship between adults and brood which Wheeler
(1928) has termed ‘ ‘ trophallaxis. ”15
On the other hand a group without any brood or with pupae in
cocoons is distinctly less lively than when larvae are present.
Even when exposed to light the workers seldom appear very ex-
cited, and most of the time they stand quietly over their heap
of cocoons or near it, or move slowly about in the vicinity. Under
these conditions Ecitons are chronically less feverish in their ac-
tivity, indulge in many fewer excursions from the nest, and hence
they carry in far less food than in the larva-containing situation.
We may say that the group or colony which contains active larva
as in the nomadic period has a special source of colony “drive.”
In contrast, in the statary period, this factor is greatly reduced
when interindividual stimulation is limited to relationships among
adult workers themselves.16

14 Gallardo (1920) has commented upon the great mobility of Eciton
larvae.

is The gastric anatomy of the larvae of B. burchelli and the feeding be-
havior of these larvae have been described by Wheeler and Bailey (1920).
In this species the larva is fed at frequent intervals on boluses composed of
the soft parts of insect prey, laid by the workers upon the ventrum of the
larva or actually thrust into the larval gullet.

i6 From his observations on a colony of B. burchelli, Wm. Muller (1886)
inferred that an increase or decrease in the raiding activities of a colony is
attributable to the relative need of the brood for food.

“Larven brauchen ja im allgemeinen, besonders kurze Zeit vor
der Yerpuppung, bedeutend mehr Nahrung als die fertigen Insekten,
und so scheint nichts natiirlicher, als dass das Nahrungsbediirfnis der

June, 1944]

Schneirla: Insect Behavior

179

However, in a statary colony with advanced pupae there is a
marked change in the general level of activity when the nearly
mature individuals begin to move within their cocoons. When a
close approach to the end-poinjt of pupal development is indicated
by the appearance of a distinct pigmentation of eyes, mandibles,
and anterior sections of the body, activities such as twitchings of
the legs and slight turnings of the body become observable. Such
movements of the pupae may be increased by picking individuals
up with tweezers or by lightly probing the body, just as they are
perceptibly elicited when the cocoons are picked up by workers.
The stirring of mature pupae appears to excite the workers and
evidently leads to the removal of the brood from cocoons ; the en-
velopes readily tearing open as the workers manhandle them
(Schneirla, 1934, pp. 320-321). That the opening of cocoons de-
pends critically upon pupal movements is suggested by the fact
that test groups did not open cocoons in which the advanced pupae
had been killed with a needle (op. cit., 1938, p. 66).

There is a distinct increase of excitement in an Eciton colony
as emergence of its brood gets under way. The excitement rises
in crescendo until, a day or two later when the largest part of the
brood is free and active as callow individuals, the colony shakes
itself from the statary period by staging an exceptionally vigor-
ous raid ending in a bivouac-change movement. Laboratory ob-
servations show that in the appearance of a new pupal brood and
in the strong stimulative effects from callows during their first
few days as “free” individuals in the colony, an Eciton worker
population receives a great lift in internal “drive.” The conclu-
sion seems in order that this internal change is responsible for
setting a colony into the nomadic phase.

Gesellschaft ein geringeres wird und die Tiere entsprechend weniger

auf Beute ausgehen, nachdem alle Larven eingesponnen 7 ’ (p. 87).

While this point undoubtedly has relevance to the 'phylogenetic problem (con-
cerning the evolution of the Eciton behavior pattern) and describes an
important adaptive function of the pattern, it is teleological and misleading
if applied to the ontogenetic problem (concerning the contemporary causa-
tion of the pattern in a given colony). As the present discussion shows, it is
not the brood ’s food-consuming capacity as such, but the presence or absence
of incidental tactuo -chemical stimulation from the brood which critically
governs periodic changes in colony behavior.

180

Journal New York Entomological Society

[Vol. Lll

Our theory thus accounts for the nomadic-statary (rainy-sea-
son) cycle of Eciton colony activity in terms of cyclical changes
in the internal stimulative properties of different broods. In
Figure 2 the relationship is represented schematically in terms
of a diphasic diagram, on which are indicated the positions evi-
dently attained by our colonies at the times their respective
queens and broods were captured. The “trough” phase of the
cycle signifies the statary period, in which the colony is held in
position and is comparatively inactive and the brood or broods
relatively passive in their social roles. A colony enters this
statary phase when the enclosure of its mature larval brood re-
moves the major internal source of special social stimulation.
The appearance of a new brood as eggs laid after about one-third
of the period has elapsed does not materially change general
colony behavior, since with this brood concentrated in masses,
incapable of movement, it can exert only a relatively negligible
stimulative effect upon the general population. As a matter
of fact, the vigor of raiding seems to decrease somewhat toward
the middle of the statary period, and hamatum colonies sometimes
show raidless days then (also E. burchelli, as Muller (1886)
noticed). With social stimulation low throughout the period, the
workers are only weakly stimulated to leave the colony in raiding,
with the result that statary-period raids are relatively feeble as
compared with nomadic-period raids.

We may say that in the statary period a low summation of
colony excitement directly expresses its limitations in the under-
development of raiding, which roughly is only about one-third as
extensive as in the nomadic period. Such raids may be regarded
as underdeveloped not only in their restricted numbers and scope
but also in falling below the threshold of development at which
a bivouac-change movement becomes inevitable.

To speak of a “threshold” of raiding essential for new behavior
requires us to show why underdeveloped raids cannot lead into
a colony movement. This has been done in special studies of the
relationship between raiding and the bivouac-change movement
(Schneirla, 1938, 1944) which bring out the reason why a colony
can withdraw to its original bivouac from an underdeveloped
statary-period raid but must move onward when a maximal

June, 1944]

Schneirla: Insect Behavior

181

nomadic-period raid has been staged. The actual behavior proc-
ess, fairly complex, centers around the fact that in the statary
period the colony is insufficiently rearoused (by regular post-
meridional atmospheric changes) after the midday lull in ac-
tivity, so that no exodus exists capable of blocking the principal
trail to returning raiders. In the nomadic period, on the other
hand, the afternoon rearousal produces a lasting afternoon exodus
from the bivouac which forcibly reorients returning raiders out-
ward in at least one trail system; then a movement from the
bivouac persists on this route, eventually draining the entire
colony to a new bivouacking site. Thus when extrinsic stimu-
lation (from light in particular) drops away at dusk, a colony in
the statary phase loses its principal source of arousal and grad-
ually approaches quiescence; whereas a colony in the nomadic
phase when raiding declines with dusk possesses an adequate and
persistent internal “ drive’ ’ facilitating the vigorous exodus al-
ready in progress. The key to the situation appears to be the
radical difference in the intrinsic properties of colonies in these
two conditions.

The break from the statary phase appears to be accomplished
not through the influence of the new larval brood, which at that
time seems insufficiently developed to be a social-stimulative
factor of any account, but rather through activities centering
around the maturation of the pupal brood and its appearance as
an enormous horde of hyperactive, voracious callows. Invari-
ably, hamatum colonies shift from their statary sites when the
pupal brood has largely emerged; the deserted spot is littered
with empty cocoons, and the first bivouac-change processions are
distinguished by crowds of pale-colored callow workers. Circum-
stances indicate that the stimulative effect from tens of thou-
sands of callows not only serves to bring the colony into nomadic
life, but also persists for a few (apparently about five) days as
a factor of similar influence and importance. Our colony 38 X
was taken at such a time. As a major source of stimulation, the
presence of the callows appears to overlap the involvement of the
new brood of larvae, in that as the callows lose their early social-
stimulative effect and merge into the adult worker population, the
larvae become increasingly functional in this respect. About five

182

Journal New York Entomological Society

[Vol. LIT

days after the nomadic period has begun, typically, the callows
are almost indistinguishable from regular workers and (judging
from laboratory observations) have dropped to the level of the
ordinary workers in their trophallactic properties. Meanwhile,
the larvae have increased notably in size and activity, most of them
are capable of feeding, and at this time they appear to constitute
the new major source of social stimulation or intrinsic colony
‘ ‘ drive. ’ ’ Our colony 38 C was taken at such a time. When it
was captured, the larval brood was distributed rather widely
through the interior of the bivouac, and in the laboratory a pro-
nounced activity was noted in all but the smallest size category.

Colonies 38 C, D, E, F, and G (taken in chronological sequence)
may be placed at successive points in the nomadic period on the
basis of the invariable fact that an increasing development of the
larval brood parallels an advance through this phase of activity
(see Fig. 2). Not only are the trophallactic properties of the
brood instrumental in maintaining the nomadic pattern of be-
havior by supporting daily maximal raids, but as the larvae de-
velop further their influence appears to increase. The apparent
augmentation of their social effect probably is due to increased
chemo-stimulative properties as well as to greater general activity
as they grow. This is suggested by laboratory observations and
by the fact that in the bivouac more advanced broods tend to be
widely distributed through the cluster, single larvae held by indi-
vidual workers much more frequently than with younger broods.
Thus the intrinsic ‘ ‘ drive ’ ’ effect of the brood not only maintains
the nomadic pattern in the colony, so that maximal raids and
bivouac-change movements appear regularly in the daily routine,
but this relationship appears to rise through a crescendo as brood
development nears its climax. Hence, near the end of the no-
madic period, raids are more extensive and more heavily popu-
lated, and colony movements tend to carry over longer distances
than earlier in the period. In particular, this is revealed by
studies of individual colonies throughout the nomadic period
( e.g colony 1936 A, — Schneirla, 1938; also E. burchelli, colony
381, — Schneirla, 1944).

As we have pointed out, in addition to postulating an excitation
to “maximal’ raiding through the drive effect of active broods,

June, 1944]

Schneirla : Insect Behavior

183

the theory postulates a given threshold in raiding which is essen-
tial for the occurrence of a bivouac-change. As the relationship
has been worked out in a previous paper {op. cit., 1938), only in
the nomadic period does raiding attain the pattern {i.e.y three
trail systems in hamatum) and the degree of colony involvement
that is required if it is to eventuate in a colony movement. That
the essential “maximal” pattern of activity in the colony depends
specifically upon a characteristic larval-brood influence is sug-
gested strikingly by the regularity with which colonies lapse into
the statary period on the very day most of the larvae have spun
their cocoons.

It seems clear that since a critical difference in the pattern of
hamatum colony behavior depends upon the condition of the
brood, the ultimate determination of such matters resides in the
individual that produces the broods, i.e., the queen.

THE ECITON QUEEN AS PACEMAKER FOR COLONY BEHAVIOR

It is characteristic of E. hamatum, E. burchelli, and other rep-
resentatively terrestrial species of Eciton s. str. (if not Ecitons
generally), that broods appearing in the rainy season are very
large, and that all individuals in a given brood develop concur-
rently. That is, in each new hamatum brood the eggs are laid
within a span of a few days, pass through the larval period essen-
tially in step, enter the pupal stage within about four days, and
appear as callow workers all within a very short period. Obvi-
ously this state of affairs depends essentially upon the reproduc-
tive properties of the Eciton queen.

Let us first consider some further facts about brood develop-
ment that throw light indirectly upon the capacities of the queen.
Population studies have been made with four broods captured on
Barro Colorado Island, with these results : E. hamatum — a larval
brood, 26,452 individuals (not complete), a pupal brood, 31,379
individuals (fairly complete) ; E. burchelli — one complete larval
brood of 36,888 individuals, and a pupal brood of 31,298 indi-
viduals. Then too, the mass of eggs from physogastric queen
38 H totalled more than 26,000, probably with a few thousand
missing. Since many other broods inspected in the egg, larval, or
pupal condition were comparable in bulk with these populations

184

Journal New York Entomological Society

[Vol. LII

which were actually counted and sized, it seems that the rainy-
season broods of E. hamatum (and of Eciton s. str. broadly) are
typically very large. In all probability the broods in this season
comprise more than 25,000 individuals as a rule. That a single
queen is capable of delivering an egg mass of these proportions
within a few days is an impressive fact.

Beyond the fact that in nearly twenty captures we have never
found more than one queen in a colony, and the fact that the
enormous clutch of eggs in colony 38 H may be attributed safely
to a single queen, our evidence on condition of the broods seems
to exclude any possibility that more than one queen figures in
producing a given brood. First of all, the range and central
tendency of body size in young and advanced larval broods are
rather constant for given stages (see Table I). The range of
individual sizes is at first limited ( e.g colonies 38 H, A, B, and
X) but increases with the general age of the brood (colonies 38 C,
D, E , F, G, and others). That there is a definite unimodality
in the distribution of individual sizes within a given brood was
indicated by study of the broods mentioned above, and has been
verified in all four of the broods surveyed in toto. Furthermore,
all brood distributions are skewed toward the worker-minor ex-
treme. With the population classified into five body-size cate-
gories (in terms of body length), with No. 1 the smallest and No.
5 the largest, the mode falls in category No. 2 as a rule, well below
the median value. For example, in a hamatum pupal brood (of
31,379 individuals in cocoons) which was surveyed, the following
distribution was found through the five size categories, from
smallest (ca. 5 mm. long) to largest (ca. 10.5 mm. long) : 5,209,
16,860, 8,072, 870, and 368, respectively. The prevalence of the
described distribution pattern in Eciton brood populations indi-
cates the involvement of a single reproductive source in each case,
a maternal source having definite and regular properties.

As further evidence we may repeat the fact that enclosure of
the larval brood and the opening of cocoons when a pupal brood
hatches are matters of mass change, both events accompanied
by prominent changes in colony behavior. In view of these con-
siderations together with our invariable discovery of but one
queen to a colony in many captures, the conclusion seems justified

June, 1944]

Schneirla: Insect Behavior

185

that these large worker broods appearing in hamatum (and in
burchelli) colonies are attributable to single queens in the respec-
tive cases.17

Furthermore, the time relations of successive broods in given
colonies are sufficiently predictable to suggest the function of but
one queen in each case. The results from colony 36 A (E. ha-
matum), which was studied during a period of 42 days, from
August 5 to September 15, 1936, bear directly on this point
(Schneirla, 1938). During seven weeks of observation, this
colony passed from a statary period into a nomadic period which
lasted 17 days, then spent 19 days in a new statary period, after
which it entered a further nomadic period. Three successive
broods were observed, all of them relatively immense and entirely
composed of worker forms. The first of these broods terminated
its development and appeared as callow workers when the study
began. The second brood was present as developing larvae
through the complete nomadic period, was enclosed in cocoons
and entered the phase of pupation when the fully observed statary
period began, passed through its pupal development and ap-
peared as callows as a further nomadic period got under way.
The eggs of the third brood were laid at some time between Au-
gust 28 and September 6, and this brood was well started on its
larval development when the colony was captured September 15.
In the end the colony was anaesthetized and thoroughly examined.
Only one queen (in the contracted condition) was found.

This evidence is brought together in the schematic representa-
tion of the hamatum behavior cycle in Figure 2. The condition
of the brood or broods as indicated at times of major change in
colony behavior corresponds to the facts for colony 36 A and
checks with many other cases, as does the representation of a
developing larval brood through the nomadic period and a pupal
brood through the statary period. Our records indicate that in
the area of this study individual hamatum (and burchelli) col-
onies characteristically pass through the cycle of correlated be-
havior changes and worker-brood production a number of times

17 A possible alternative explanation is that the broods are the product of
two or more queens with exactly synchronized visceral rhythms. This seems,
very doubtful, especially because no polygynous colonies have been dis-
covered.

186

Journal New York Entomological Society

[Vol. LII

in regular succession during the first four or five months of the
rainy season.

To describe the characteristic time relations of the principal
changes in brood development for hamatum, it is necessary to
ascertain the time at which the eggs are laid more exactly than is
possible from the record of colony 36 A. From general evidence,
and from the facts concerning queen 38 H, we have inferred that
the process of egg-laying occurs during a short period following
about one week after the beginning of the statary period. The
conclusion that normally a new brood appears as eggs about ten
days before the end of each given statary period is supported
directly and indirectly by numerous facts. In particular, al-
though eggs are never found in colonies examined during the first
few days of the statary period, large masses of eggs are found
during the last week of this period. Queen 38 H evidently had
just passed the peak of an egg-laying process when captured seven
days after the beginning of a statary period. Since in hamatum
oviposition appears to be of short duration, probably no more
than three or four days from beginning to end, if we set 18-20
days as the usual length of the statary period in this species we
may say that the new brood has roughly ten days of early develop-
ment before the statary period ends.

On this basis we may estimate the developmental period of a
given brood. To the first 10 days of embryonic and early larval
growth when the colony is statary, and 17 days to complete larval
development in the nomadic period, we may add 19 days (as a
close approximation) in the pupal stage during the following
statary period. That makes a total of 46 days for the complete
development of a hamatum brood. Other facts plainly indicate
that the interval between the appearance of successive broods in
a given colony is the number of days from the end of one statary
period (when one brood is delivered as callows) to the end of the
next statary period (when the next brood is delivered), i.e., about
35 days.18

is As may be seen in Figure 2, the difference of about 10 days in the above
figures is attributable to the fact that the oviposition and early development
of a given new brood overlaps the period in which the preceding brood is
completing its pupal maturation.

June, 1944]

Schneirla: Insect Behavior

187

On this basis we may arrive at an approximation of the interval
between successive occurrences of the short but strenuous ovi-
position act of the queen. After having deposited a given clutch
of eggs, the queen evidently lays no more until the early part of
the next statary period. If the peak of one process is roughly
placed at seven days after the beginning of a statary period
and its end at nine days, for the ensuing resting phase there are
ten days remaining in the same period, 17 days in the ensuing
nomadic period, and perhaps five days in the following statary
period, — 32 days in all for the interval between successive inter-
vals of actual egg-laying. This checks fairly well with the figure
offered above for the interval between the appearance of suc-
cessive broods of callows, as it obviously should.

To repeat, there is every reason to believe that in a given
hamatum colony the broods appearing at intervals of about 35
days are the progeny of a single queen. Not only the facts con-
cerning the capture of queens, but also the highly predictable
time relations between successive broods, support this conclusion.
The facts as sketched in Figure 2 indicate that the entire set
of events is highly synchronized; particularly in the ability of
the queen to deliver a new batch of eggs about 12 days before
the previous brood joins the worker population of the colony.
The evidence suggests that in our area of study a given hamatum
queen is capable of repeating this process a number of times
during the first months of the rainy season and perhaps even
longer.

Thus in an indirect manner but nevertheless very effectively
the queen is the pace-setter of the cyclic changes which have
been described for the type species. This relationship appears
to hold for other Eciton s. str. species as well ( e.g E. burchelli,
E. vagans), and perhaps also in other Eciton subgenera.

To summarize, in hamatum we seem to have the master pattern,
as it were : 1 ) the queen at regular intervals and in a strikingly
precise manner furnishes a huge mass of eggs which begin their
development almost simultaneously, 2) the given brood does not
materially influence events until the emergence of the previous
brood as callows dynamizes the population into nomadism, 3)
then the new brood (as larvae) takes over as principal source of

188

Journal New York Entomological Society

[Vol. LII

the “social stimulation” which maintains the highly dynamic
activity pattern of the colony for some time ; 4) when the “drive”
effect of the larval brood is removed as it enters the pupal condi-
tion the colony lapses into its minimal activity pattern, from
which 5) the emergence of this brood as callow workers arouses
it into a new dynamic phase — and so on.

The queen is the key of this entire process, but only in an
indirect manner through her physiological properties as the pro-
ducer of successive broods. After a given brood has appeared as
eggs we may say hypothetically that the presence of the queen
is essential neither for the next major change in colony behavior
nor for the maintenance of this change ( i.e ., for nomadism). In
fact without the queen the next sessile phase of the colony-
behavior cycle may also occur and the colony will even begin
its next nomadic period (c/. Fig. 2). However, if no new brood
is forthcoming we should expect the colony to lapse from its new
nomadic phase into an aberrant and mainly sessile type of ex-
istence, i.e., a protracted statary condition. Some evidence
corroborating this prediction is in fact available from the study
of colonies deprived of their broods.

To state the hypothesis in a different way, it is quite likely
that in the dry season when Eciton broods are very small (either
through metabolic insufficiency in the queen or through brood
cannibalism in the workers, or both of these), the colonies lapse
almost completely into statary life. But under optimal condi-
tions in the rainy season, the adequate condition and regular
labors of the queen periodically furnish the colony, in an inci-
dental manner, with the changes in trophallaxis-based social
stimulation which condition the drive (or lack of drive) under-
lying alternate changes in the general behavior cycle.

The exquisite synchronization of the set of relationships in-
volved in the Eciton behavior pattern is emphasized in several
ways when the queen is considered in her role of key individual.
Through her capacity to deliver an entire batch of eggs within
a few days, the queen indirectly contributes the precision with
which the colony shifts from one mode of life to another. This in
turn promotes a further characteristic of great adaptive value.
Due to the spacing of her successive broods, the queen becomes

June, 1944]

Schneirla: Insect Behavior

189

physogastric and lays her eggs only at times when the colony
happens to be statary (see Fig. 2), a combination of events which
is most opportune for survival of colony and species. It is ap-
parent from our discussion of the indirect relations existing
between the cycles of queen and colony functions that a true
convergence of events occurs in this case, which serves to guard
the queen from injury at the one time she is most vulnerable.
When she is gravid, having to make her way over a long bivouac-
change trek undoubtedly would offer a serious risk for the queen,
carrying the constant threat of dangers such as a tumble from
elevated sections of the path on vines or tearing the tightly
stretched intersegmental membranes on rough surfaces. In-
stead, thanks indirectly to her own visceral regularity, the queen
is safely immured through this critical period, and moreover,
there follows a further time of safety during which she may
convalesce from the organic ordeal of large-scale oviposition.
Not least among the factors fitting into the marked adaptivity of
this synchronized pattern of events, the bivouac of the colony in
its statary phase is characteristically more sequestered than at
other times. Altogether, it would be difficult to find a more
effectively adaptive mosaic of various biological events than this
one, based upon the physiological properties of the Eciton queen.

SUMMABY AND CONCLUSIONS

The functions of the Eciton hamatum queen serve indirectly
as a pace-making factor in the colony behavior pattern of the
species.

The determining processes in a given colony center around the
reproductive properties of the single functional queen. Except
when colony bivouac-change movements occur the fertile queen
is confined to the bivouac, evidently to a large extent because of
her pronounced photonegativity. At regular intervals of ap-
proximately 35 days the dichthadiigyne becomes physogastric and
within a few days releases a mass of more than 20,000 eggs. Be-
tween egg-delivery episodes she remains in the contracted or
“resting’’ condition. The ability of the hamatum queen to
mature and deliver an immense number of eggs within a very
limited time means that all members of a given brood develop

190

Journal New York Entomological Society

[Vol. LIT

and mature roughly in synchronization. These facts hold im-
portant consequences for the colony behavior pattern.

In the rainy season of the Caribbean lower rain-forest zone of
Panama, a colony of E. hamatum passes through statary (mini-
mal raiding; absence of migration) and nomadic (maximal daily
raiding; successive daily bivouac-changes) behavior phases at
regular intervals, each period lasting nearly 20 days. Depending
upon the properties of a fertile queen, the alternation of these
behavioral phases is highly predictable.

Significantly different intracolony circumstances parallel the
two phases of the colony behavior cycle. Colonies in the nomadic
condition invariably contain a single brood passing through its
larval stage, and the period ends precisely when this brood has
become mature and is mainly enclosed in cocoons. In statary
colonies a brood in the pupal stage is found, and the period ends
when this brood has matured and has mainly emerged as callow
workers. Approximately seven days after a given statary period
has begun, a new brood of more than 20,000 eggs appears.

The intimate correspondence which exists between Eciton
colony behavior and brood condition is explicable in terms of the
Wheeler trophallaxis concept. Evidence is cited for the involve-
ment of a special social-stimulative effect, a superadded “ drive’ ’
factor, furnished by broods capable of extensive overt activity.
Emanating from a newly emerged lot of callow workers, this effect
arouses a previously sessile colony to the threshold of maximal
raiding and daily bivouac-change, and thus is responsible for
initiating a new nomadic period. Furthermore, due to a rather
precise synchronization of successive broods, before the energiz-
ing function of a new callow brood runs its course it is overlapped
and succeded by the similar function of a sufficiently developed
larval brood. This larval effect maintains the nomadic condition
over a considerable time, but the colony lapses promptly from
nomadism when the larvae mature and spin their cocoons. Thus
the sessile and relatively inactive statary condition which ensues
is attributable to the existence of an inadequate energization of
the colony when the brood (eggs; pupae) is incapable of function-
ing as an effective source of social stimulation.

Periodic changes characterizing Eciton colony behavior thus
basically depend upon a highly regular reproductive cycle in the

June, 1944]

Schneirla: Insect Behavior

191

queen. By furnishing new broods at fixed intervals, the queen’s
function indirectly governs the presence or absence of the intra-
colony dynamizing factor critically responsible for the ebb and
flow of events in the Bciton behavior pattern.

LITERATURE CITED

Andre, E. 1885. Species des Hymenopteres. (Supplement au species des
Formicides d ’Europe et des pays limitrophes.) 8: 838-840.

Bruch, C. 1934. Las formas femeninas de Eciton . . . Descripcion y re-
descripcion de algunas espeeies de la Argentina. Ann. Soc. Cient.
Argent., 118, 2nd sem., 1934: 113-135.

Emerson, A. E. 1939. Social organization and the superorganism. Amer.
Midi. Nat., 21 : 182-209.

Gallardo, A. 1920. Las hormigas de la Republica Argentina — Subfamilia
dorilinas. Anal. Mus. Nac. Hist. Nat. Buenos Aires, 30: 281-410.
Muller, W. 1886. Beobachtungen an Wanderameisen (Eciton hamatum
Fabr.).i9 Kosmos, 18: 81-93.

Reichensperger, A. 1926. Das $ von Eciton mattogrossensis Luederw.
(Hym.) Ent. Mitteil., 15: 401-404.

. 1934. Beitrag zur Kenntnis von Eciton lucanoides Em. Zool. Anz.,

106: 240-245.

Schneirla, T. C. 1933. Studies on army ants in Panama. Jour. Comp.
Psychol., 15 : 267-299.

. 1934. Raiding and other outstanding phenomena in the behavior of

army ants. Proc. Nat. Acad. Sci., 20: 316-321.

. 1938. A theory of army-ant behavior based upon the analysis of

activities in a representative species. Jour. Comp. Psychol., 25:
51-90.

. 1940. Further studies on the army-ant behavior pattern. Mass

organization in the swarm-raiders. Ibid., 29: 401-460.

. 1944. Studies on the army-ant behavior pattern. Nomadism in the

swarm-raider Eciton burchelli. Proc. Amer. Phil. Soc., Phila., 89
(1) (in press).

Weber, N. 1941. The rediscovery of the queen of Eciton ( Labidus ) coecum
Latr. (Hym.: Formicidce) . Amer. Midi. Nat., 26: 325-329.
Werringloer, Anneliese. 1932. Die Sehorgane und Sehzentren der
Dorylinen nebst Untersuchungen iiber die Facettenaugen der For-
miciden. Zeitschr. wiss. Zool., 141 : 432-524.

Wheeler, G. C. 1943. The larvae of the army ants. Ann. Entom. Soc.
Amer., 36: 319-332.

Wheeler, W. M. 1900. The female of Eciton sumachrasti Norton, with
some notes on the habits of Texas Ecitons.20 Amer. Nat., 34 : 563-
574.

19 Redetermined as E. burchelli.

20 Redetermined as E. schmitti.

192

Journal New York Entomological Society

[Vol. LII

. 1910. Ants, their Structure, Development and Behavior. N. Y. :

Columbia University Press, pp. 663.

— -. 1921. Observations on army ants in British Guiana. Proc. Amer.

Acad. Arts Sci., 56 : 291-328.

. 1925. The finding of the queen of the army ant Eciton hamatum

Fabricius. Biol. Bull., 49: 139-149.

. 1928. The social insects, their origin and evolution. N. Y. : Har-

court, Brace.

and I. W. Bailey. 1920. The feeding habits of Pseudomyrminse

and other ants. Trans. Amer. Phil. Soc., 22, N.S., Art. 4: 235-279.

Zahl, P. A. 1939. To the Lost World. N. Y.: Knopf.

June, 1944]

Bird: Papaipema

193

A RE-SURVEY OF PAPAIPEMA SM. (LEPIDOPTERA)

By Henry Bird
Rye, N. Y.

The intricacies of insect life are abysmal and any individual
studies are bound to be incomplete and fragmentary. This is so
axiomatic as to need no argument and may excuse the limited
viewpoint of any single observer. However, as time goes on data
and observable facts accumulate, given problems here and there
gain enlightenment through various channels.

Retrospective deductions on the part of the writer based on
“the sum of evidence” as this slowly evolves apparently offers
some ground work for the serial arrangement of Papaipema
species.

Conceived as an ontogenetic tree with its phylogenetic roots
outcropping from supposedly more ancient genera, it is interest-
ing to invade this vale of surmise.

To what extent these relationships can be shown in a list
arrangement is unsatisfactory but should be undertaken. As
building bricks there are the characters of the adults including
of course the genitalia of both sexes, the gleaning from larval
evidence backed by attending factors. Thus it becomes requisite
to discourse somewhat at length taxonomically.

Also the final disposition of holotypes should be chronicled.

First, as to the generic basis whereupon Prof. J. B. Smith
established Papaipema in 1899. 1

He named no genotype and the gist of his characterizations
featured moths with primaries rather broad and outwardly acute
at apex; the thoracic tuftings decidedly upright and anteriorly
usually broadened, in form like an “adze” behind the collar; the
antennae are simple ; the f rons smooth ; the male genitalia mainly
show a unique pattern “having the harpes more or less forked
with triangular patch of spinulated surface at the tip. The
clasper in almost all cases a long, stout, curved hook, but is unique
in having the outer curve strongly toothed. ’ ’

1 Revision of Hydroecia, Trans. Am. Ent. Soc., Yol. XXVI.

194

Journal New York Entomological Society

[Vol. Lll

As now considered there are nearly fifty species in the genus
with thirty-seven at least following this genitalic pattern closely.
Smith’s simple drawings of a portion of the male genitalia are
misleading however since it is necessary to chronicle the following
specific errors.

Harrisii and pterisii were considered by him as one species;
duovata, arctivorens and merriccata at least were confused under
“rutila”; circumlucens, ochroptena and form humuli were
treated as circumlucens ; he considered cerussata and frigida form
thalictri under cerussata label; treated purpurifascia and lysi-
machice as one; misidentified sciata for “limpida” ; nepheleptena
for appassionata, while his reference to necopina had largely to
do with maritima.

Criticism should not be levied unduly at these presumed mis-
takes since some of Guenee’s types, notably rutila and limpida,
British Museum uniques, have not been satisfactorily associated,
and two of our names as used hereinafter will probably fall
through this lack of perception.

In 1910 Sir George Hampson,2 following his custom of citing
genotypes, selected cerina as genotype of Papaipema, on First
Species Rule because it headed Smith’s enumeration of the genus.
That was an unfortunate usage since the species does not measure
fully to Smith’s definition. Recognizing this discrepancy from
a mere autopic glance, and though bound by the Rules he never-
theless uses the very proper species harrisii in illustrating vena-
tion and the bodily detail, quite representative of the genus.

Begging Sir George ’s pardon, this writer votes that cerina Grt.
be superseded as genotype of Papaipema by harrisii Grt., if a
more elastic rule be forthcoming some day.

The limitations of generic boundaries are subject to varying
personal ideas but ideally their demarcation should suggest evo-
lutionary trends in so far as that might be surmised.

Avoiding theory as much as possible but judging facts as they
appear today, we can find much aid in turning to the larvae in
their earlier stages.

It is generally conceded that early stage larvae reflect the primi-
tive ancestral line at least phylogenetically. Leaning on this

2 Cat. Lep. Phal. Brit. Mus., Vol. IX, p. 80.

June, 1944] Bird: Papaipema 195

deduction, a personal familiarity with forty Papaipema species
seems to help in some measure while details of color pattern aid
specifically. As larvae Papaipema species are unique.

Their early larval pellicle is distinctly colored whereas most
mining larvae are at all stages more or less translucent.

The great majority show a contrasting middle ring of dark
purplish or pinkish brown, in livid hue, at the first four abdomi-
nal segments while elsewhere longitudinal white or yellowish lines
drawn on the darker body color produce striking individuals.

This intensity continues through the instars up to the penulti-
mate, while maturity usually exhibits a faded translucence.

Three pattern types follow; the dark middle girdle may show
an abrupt termination of all lines ; or the dorsal line may cross it
in unbroken continuity ; or both the dorsal and subdorsal may be
entirely unbroken. These features of pattern aid much specifi-
cally.

They are constant with two exceptions — the Pacific Coast spe-
cies angelica and insulidens , where one, or rarely both lines may
be continuous. One is tempted to assume that a progenitor was
not wholly an internal feeder but subsisted within some en-
circling tissues with both extremities exposed and maintaining
there the linear markings. The above category applies to thirty-
eight known individuals. Two other known species are decidedly
different and, again assuming, feature as admirable connecting
links with their Apamea-Gortyna-Hyroecia relatives possessing
a world-wide, north temperate zone dispersal. Because of this
dispersal as against restricted North American Papaipema it
seems rational to consider the latter as a subsequent offshoot.
Their very close relationship bespeaks a comparatively recent
evolution wherein possibly marked choices of food plants, some
of the latter also restricted to America, may have played a part
in influencing specific origin.

The larvae of the two alleged connecting species, frigida and
beeriana have their markings as transverse segmental rings,
features prevailing with Apamea erepta ryensis, Hydrcecia im-
manis, H. micacea and PL. stramentosa , which is as far as famili-
arity goes.

The exotic Xanthoecia flavago larva is cross banded, while
Parapamea buffalcensis and Emboloecia sauzalitee have larvae

196

Journal New York Entomological Society

[Vol. LII

which are longitudinally lined. This latter trio has the frons
distinctly armed but these various genera may all figure as part
of the ‘ ‘ Gortynid series. ’ ’ This term has been used by the writer
as a convenience, and in view of Grote’s arguments,3 Gortyna,
genotype micacea Och., might properly find generic application
somewhere therein.

As structural larval features, the heavy setigerous plates are
noteworthy, with a peculiar development in many species that
have an additional plate known as IVa, on joint ten. Since this
plate bears no setal hair its transitory character may be adduced.

However it is one of the evidences aiding specific distinction.

The rugged genitalia become a prime structural feature with
the adults. In the males such closeness to the conventional pat-
tern prevails as to indicate the very near relationship of the
species. Greatest modification exists with furcata and eryngii,
while frigida, unimoda and appassionato, make a decided break
with the harpes greatly reduced. One might suggest a generic
break here but full evidence points to connecting species.

With the female genitalia the genital plate at the ostium is
of help specifically. It is a heavy, more or less shield-shaped
process, differing in outline and scobinated characteristically.
Dr. F. Heydemann in a praiseworthy treatment of the nictitans
group of Apamea 4 notes the value of the genital plate and figures
the character without other detail upon a single plate for specific
comparison.

The close proximity of Papaipema species blend them together
into a satisfactory whole. Indeed, in a number of instances they
are so close that if no further evidence was at hand than a few
flown specimens much doubt would arise as to their distinction.

Furthermore, variation is rife particularly in the feature
wherein the ordinarily prominent white marked stigmata may
be obsolescent or vice versa. This is productive of such an autopic
difference that erroneously, two distinct species seem to be in-
volved— vide Guenee’s two alleged species, nebris and nitela.

That a varietal name be given to the lesser of such forms seems
practical. This has been done in the more striking instances and

3 Historical Sketch of Gortyna, Proc. Am. Phil. Soc., Vol. XXXIX, No. 162.

4 Die Arten der Eydroecia nictitans .« Sond. Ento. Zeit., XXXXIX u.

xxxxv.

June, 1944]

Bird: Papaipema

197

the future holds possibilities with slighter stigmatal variance
likely to be grasped by some. Some dimorphism is observable,
the ubiquitous cataphracta with its extensive food habits shows
this in its more northern range, while imperspicua represented by
a unique type may well be in that category. Until rearing proves
the fact its specific standing may remain.

Two formerly considered species must merge as one, verona
and astuta with the latter name preserved as varietal. Smith’s
unique type of verona, from Winnipeg, Manitoba, is a dwarfed
pale form of the species, while astuta was applied to a larger more
colorful variant with the terminal space solidly purplish, easily
suggestive of distinctness. Both can occur in the same locality
and though verona is less numerous in the writer’s experience;
it has priority.

That astuta be retained as a distinct color, or dimorphic form
points to future expediency.

As to the placement of Papaipema holotypes, the writer has
prepared a detailed treatment of the genus under title “The
Epic of Papaipema,” a unique copy which is willed to the
American Museum of Natural History, New York, N. Y.

With it will go his collection of the group. Hence the holotypes
and paratypes, with the relevant literature will be at one place
for future students. The collection embraces something over
fifteen hundred specimens, mainly reared. The various types
number ninety-eight examples. The aggregation brings together
not only the adults, but larval and pupal stages, to some extent
the parasites; the foodplant habitations are also shown. The
genitalic slides are not considered in this summary.

The “Epic” consists of three volumes. It assembles the
principal published literature thus dealing with the historic rise
and subsequent departures in the genus, treats monographically,
matters of taxonomic import, features of parasitism and the
gleanings of several decades of field study. Interlarded between
the published papers, the author’s notes and criticisms bring
such up to current review. Particularly some of his earlier
papers were rather puerile and in need of revision. Volumes I
and II total 1127 pages, inclusive of the relevant articles. Vol-
ume III is a portmanteu affair.

198

Journal New York Entomological Society

[Vol. LII

Seventy plates occupying seven containers, are unbound for
easy comparison; four containers hold autographed letters from
important workers in connection with the subject, many of whom
have now passed on. The plates illustrate by line drawings the
infested food plants in some instances, larval features and the
genitalia of both sexes in so far as possible, while the adults are
shown in color. There is also a booklet of remarks and index of
plates.

By what manner a commingling of the species of Papaipema
can be best portrayed with their ontogenetic proximity appar-
ently shown, it is convenient to resort to a popular vegetative
process and erect a fanciful “tree.”5 Then, as nonconventional
genes seem to have effected the protoplasmic stream, branches or
shoots may materialize, either ascending or paralleling the main
trunk according to the line of thought. Admirable as a basis of
ideas, but to transplant this fruitage to the linear order of list
column spoils the conception entirely. The writer can only sug-
gest the following summary.

List order for the species of the genus

PAPAIPEMA Smith.

(Asterisk denotes larva unknown, synonyms in italics)

frigida Smith.

form thalictri Lyman.
terminalis Strand,
unimoda Smith.*
beeriana Bird.

form lacinarise Bird,
appassionata Harvey.
horni Strand.

purpurifascia Grote & Robin-
son.

luteipicta Strand,
lysimachiae Bird.

nec purpurifascia Auct.
stenoscelis Dyar.
speciosissima G. & R.

form regalis Wyatt & Beer,
s EPIC OF PAPAIPEMA, 1940, pp.

inquaesita G. & R.
form wyatti Barnes & Ben-
jamin,
pterisii Bird.

triorthia Dyar.
anargyria Dyar.*
ochroptena Dyar.

form humuli Bird,
arctivorens Hampson.
merriccata Bird,
araliae Bird & Jones,
harrisii Grote.

form mulieris Strand,
sub. sp. rubiginosa Bird,
verona Smith.

form astuta Bird.

553-554, Yol. II.

June, 1944]

Bird: Papaipema

199

rutila Guenee.*
depictata Benjamin.*
nepheleptena Dyar.

moeseri Bird,
impecuniosa Grote.
circumlucens Smith.
baptism Bird,
form ochroptenoides
Benj.*

sub. sp. vaha Benj.*
marginidens Guenee.
birdi Dyar.

nephrasyntheta Dyar.*
furcata Smith,
rigida Grote.
pertincta Dyar.
limata Bird*
insulidens Bird,
angelica Smith,
cataphracta Grote.
form sulphurata Bird,
race fluxa Bird,
imperspicua Bird.*

duovata Bird,
aerata Lyman,
placida Bird.*
cerina Grote.
dribi Benjamin.*
polymniae Bird,
nebris Guenee.

form nitela Guenee.
duplicata Bird.

obsolescens Strand,
silphii Bird,
necopina Grote.
nelita Strecker.
form linda Bird,
form obicularis Strand,
errans Barnes & McDunnough.
engelhardti Bird,
sciata Bird,
limpida Guenee.*
cerussata Grote.
eryngii Bird,
maritima Bird,
eupatorii Lyman.

200

Journal New York Entomological Society

[Vol. LII

INSECT INTRODUCTIONS AND WAR

The recent press releases on the introduction of potential insect
pests in packages sent home by members of the armed forces over-
seas has evidently stirred some interest. The Bureau of Plant
Industry, New Jersey Department of Agriculture, was called to
inspect and fumigate a reed stool received by a woman in Trenton,
New Jersey from her son in India. This was found to be infested
with numerous small bostrichid beetles which were identified by
W. S. Fisher, Bureau of Entomology and Plant Quarantine at
Washington, D. C., as Dinoderus brevis Horn. D. brevis was
originally described from a single specimen taken in Louisiana,
and was believed to be a native species. This species was later
found to be Oriental, especially common in India, and has been
carried, through commerce, to many parts of the world. — William
M. Boyd.

June, 1944]

Rapp: Psychodid^e

201

CATALOGUE OF NORTH AMERICAN PSYCHODID^

By William F. Rapp, Jr.

In 1905 Aldrich1 published the last catalogue of North Amer-
ican Diptera. In the family Psychodidee he listed 34 species,
which were distributed in four genera. Table I is a comparison
of Aldrich’s 1905 catalogue and this catalogue.

TABLE I

Genus Aldrich 1905 Rapp 1943

Flebotomus 3

Pericoma .. .... 11 13

Psychoda 21 41

Sycorax 1

Trichomyia 1 2

No. of species 34 59

Since 1905 several workers have been attracted to this family
with the result that many changes have arisen in nomenclature,
plus the addition of new species. This catalogue has been com-
piled after a careful survey of the entomological literature.
Synonymy is listed wherever it has appeared in the literature.
It is interesting to note that of the 34 species listed by Aldrich
only 6 have been reduced to synonymy. Of the 54 species de-
scribed since 1905, 19 have proven to be synonyms.

The North American Psychodidae, as a whole, are of little eco-
nomic importance. Certain species of Flebotomus are known
vectors of tropical diseases, but none of these occur in the North
American region. Psychoda alternata Say is at times a pest
around sewage filter plants.2

The area covered in this paper includes all of America north
of Mexico, or the area considered as the nearctic region based
upon zoogeographical division.

1 Aldrich, John M. “A Catalogue of North American Diptera,” Smith-
sonian Miscellaneous Collections , Yol. 46 (1905), p. 1-680.

2 Headlee, T. J., and Beckwith, C. S., ‘ ‘ Sprinkling Sewage Ply, Psychoda
alternata Jour. Econ. Ent., Yol. 11 (1918), p. 395-401.

202

Journal New York Entomological Society

[Vol. LII

FLEBOTOMUS* Rondani
Flebotomus diabolicus Hall.

Phlebotomies diabolicus Hall, Proc. Ent. Soc. Wash., Vol. 38
(1936), p. 28.

Texas.

Flebotomus texanus Dampf.

Phlebotomies texanus Dampf, Anales de la Escuela Nacional
de Ciencias Biologicas, Vol. 1 (1938), pp. 119-122.
Texas.

Flebotomus vexator Coquillett.

F. vexator Coquillett, Ent. News, Vol. 18 (1907), p. 102.
Louisiana, Maryland.

PERICOMA Walker
Pericoma bipunctata Kincaid.

P. bipunctata Kincaid, Ent. News, Vol. 10 (1899), p. 34.
California, Washington.

Pericoma californica Kincaid.

P. californica Kincaid, Ent. News, Vol. 12 (1901), p. 195.
California.

Pericoma Carolina Banks.

P. Carolina Banks, Bui. Brooklyn Ent. Soc., Vol. 26 (1931),

p. 228.

North Carolina.

Pericoma furcata Kincaid.

P. furcata Kincaid, Ent. News, Vol. 10 (1899), p. 34.
Washington.

Pericoma longiplata Haseman.

P. longiplata Haseman, Trans. Am. Ent. Soc., Vol. 33 (1907),
p. 308.

Arizona.

Pericoma ocellaria var. americana Kincaid.

P. ocellaria var. americana Kincaid, Ent. News, Vol. 12
(1901), p. 194.

Maine.

Pericoma satellitia Dyar.

P. satellitia Dyar, Proc. Ent. Soc. Wash., Vol. 29 (1927),
p. 163.

Maryland.

* Phlebotmus of authors.

June, 1944]

Rapp: Psychodidjs

203

Pericoma scala Haseman.

P. scala Haseman, Trans. Am. Ent. Soc., Vol. 33 (1907),
p. 307.

Arizona.

Pericoma sitchana Kincaid.

P. sitchana Kincaid, Ent. News, Vol. 10 (1899), p. 33.
Alaska, Oregon.

Pericoma trialbawhorla Haseman.

P. trialbawhorla Haseman, Trans. Am. Ent. Soc., Vol. 33
(1907), p. 306.

Missouri.

Pericoma triloba Kincaid.

P . triloba Kincaid, Ent. News, Vol. 10 (1899), p. 33.
Washington.

Pericoma truncata Kincaid.

P. truncata Kincaid, Ent. News, Vol. 10 (1899), p. 35.
California.

Pericoma variegata Kincaid.

P. variegata Kincaid, Ent. News, Vol. 10 (1899), p. 33.
Washington.

PSYCHODA Latreille
Psychoda alberta Curran.

P. alberta Curran, Can. Ent., Vol. 56 (1924), p. 219.

Alberta.

Psychoda albipunctata Williston.

P. albipunctata Williston, Ent. News, Vol. 5 (1893), p. 113.
Telmatoscopus meridionalis Eaton, Ent. Mo. Mag. (1894), p.
195.

P. snowii Haseman, Trans. Am. Ent. Soc., Vol. 33 (1907), p.
311-312.

P. erect a Curran, Cat. Ins. Jam. Dept. Agr., Jamaica Ent.

Bui. No. 4, pt. 1 & 2 (1926), p. 102.

Florida, Louisiana, South Carolina, Texas.

Psychoda albitarsis Banks.

P. albitarsis Banks, Can. Ent., Vol. 27 (1895), p. 324.
Maryland, New Jersey, New York, North Carolina, Virginia.
Quebec.

204

Journal New York Entomological Society

[Vol. LII

Psychoda alternata Say.

P. alternata Say, Long’s Exp. St. Peter’s River, App. (1824),
p. 358.

Tipula phalcemoides Scop., Ent. Carn., No. 864 (1763), p.
324.

Psychoda sexpunctata Cnrtis, Brit. Ent., Vol. 16 (1839), p.
745.

P. marginepunctata Roser., Corr. Wurt. landro, Ver. 1
(1840), p. 50.

P. schizura Kincaid, Ent. News, 10 (1899), p. 32.

P. floridica Haseman, Trans. Am. Ent. Soc., 33 (1907), p. 316.
P. nocturnala Haseman, Trans. Am. Ent. Soc., 33 (1907), p.
319.

P. bengalensis Brunetti, Rec. Ind. Mus., Vol. 11 (1908), p.
370.

P. albimaculata Welch, Ann. Ent. Soc. Amer., Vol. 5 (1912),
p. 411.

P. dakotensis Dyar, Insec. Inscit. Menst., Vol. 14 (1926), pp.
107-110.

California, Connecticut, District of Columbia, Florida, Illi-
nois, Kansas, Maryland, Missouri, New Hampshire, New
Jersey, New Mexico, New York, Ohio, Oregon, Pennsylvania,
South Dakota, Texas, Virginia, Washington.

Psychoda annulipes Johnson.

P. annulipes Johnson, Bui. Amer. Mus. Nat. History, Vol. 32
(1913), p. 43.

Florida.

Psychoda aterrima Banks.

P. aterrima Banks, Ent. News, Vol. 25 (1914), p. 128.

New York.

Psychoda augusta Curran.

P. augusta Curran, Can. Ent., Vol. 58 (1926), p. 228.

Quebec.

Psychoda autumnalis Banks.

P. autumnalis Banks, Ent. News, Vol. 25 (1914), p. 127.
Pericoma litt oralis Dyar, Insec. Inscit. Menst., Vol. 14
(1926), pp. 107-110.

Pericoma aldrichana Dyar, Insec. Inscit. Menst., Vol. 14
(1926), pp. 107-110.

June, 1944]

Kapp: Psychodid^e

205

California, District of Columbia, Maryland, Virginia.
Alaska.

Psychoda bicolor Banks.

P. bicolor Banks, Can. Ent., Vol. 26 (1894), p. 33.

P. nigra (Banks) Dyar, Proc. Ent. Soc. Wash., Vol. 30
(1928), p. 87.

District of Columbia, Indiana, Maryland, New York.
Psychoda bishoppi Del Rosario.

P. bishoppi Del Rosario, Philippine Jour. Sci., Vol. 59 (1936),
p. 141.

Maryland.

Psychoda cinerea Banks.

P. cinerea Banks, Can. Ent., Vol. 26 (1894), p. 331.

P. elegans Kincaid, Ent. News, Vol. 8 (1897), p. 144.
Threticus compar Eaton, Ent. Mo. Mag. II, Vol. 15 (1904),
p. 57.

P. domestica Haseman, Ent. News, Vol. 19 (1908), p. 285.

P. compar Tonnoir, Ann. Soc. Ent. Belg., Vol. 62 (1922), p.

67.

P. prudens Curran, Can. Ent., Vol. 56 (1924), p. 219.
Connecticut, District of Columbia, Indiana, Maine, Mary-
land, Massachusetts, Missouri, New Jersey, New York, Ore-
gon, Virginia, Washington.

Alberta.

Psychoda criddlei Curran.

P. criddle Curran, Can. Ent., Vol. 56 (1924), p. 218.

Ontario.

Psychoda degenera Walker.

P. degenera Walker, List of the Specimens of Dipterous In-
sects in the Collection of the British Museum, List I
(1848), p. 33.

Ontario.

Psychoda helicis Dyar.

P. helicis Dyar, Proc. Ent. Soc. Wash., Vol. 31 (1929), p. 63.
Maryland.

Psychoda horizontala Haseman.

P. horizontala Haseman, Trans. Am. Ent. Soc., Vol. 33
(1907), p. 313.

Missouri.

206

Journal New York Entomological Society

[Vol. LII

Psychoda interdicta Dyar.

P. interdicta Dyar, Proc. Ent. Soc. Wash., Vol. 30 (1928),

p. 88.

Maryland, New York.

Psychoda interrupta Banks.

P. interrupta Banks, Proc. Ent. Soc. Wash., Vol. 8 (1907),
p. 150.

Maryland.

Psychoda juno Curran.

P. juno Curran, Can. Ent., Vol. 58 (1926), p. 228.

Ontario.

Psychoda longifringa Haseman.

P. longifringa Haseman, Trans. Am. Ent. Soc., Vol. 33
(1907), p. 318.

Florida.

Psychoda marylandana Del Rosario.

P. marylandana , Del Rosario, Philippine Jour. Sci., Vol. 59
(1936), p. 111.

Maryland.

Psychoda megantica Curran.

P. megantica Curran, Can. Ent., Vol. 56 (1924), p. 217.
Quebec.

Psychoda minuta Banks.

P. minuta Banks, Can. Ent., Vol. 26 (1894), p. 331.
Connecticut, New Jersey, New Mexico, New York, Virginia.

Psychoda nigra Banks.

P. nigra Banks, Can. Ent., Vol. 26 (1894), p. 331.

P. marginalis Banks, Can. Ent., Vol. 26 (1894), p. 333.
P.apicalis Banks, Proc. Ent. Soc. Wash., Vol. 8 (1906), p.
148.

P. basalis Banks, Proc. Ent. Soc. Wash., Vol. 8 (1906), p. 149.
Pericoma orillia Curran, Can. Ent., Vol. 56 (1924), p. 218.

P. varitarsis Curran, Can. Ent., Vol. 56 (1924), p. 220.
Maruina nigra (Banks) Dyar, Insec. Inscit. Menst., Vol. 14
(1926), p. 111.

Pericoma apicalis (Banks) Dyar, Insec. Inscit. Menst., Vol.
14 (1926), p. 149.

Connecticut, District of Columbia, Indiana, Maine, Mary-

June, 1944]

Rapp: Psychodid^:

207

land, New Hampshire, New Jersey, New York, Ohio, Vir-
ginia.

Ontario, Quebec.

Psychoda nitida Banks.

P. nitida Banks, Can. Ent., Vol. 33 (1901), p. 275.

District of Columbia, New York.

Psychoda olympia Kincaid.

P. olympia Kincaid, Ent. News, Vol. 8 (1899), p. 144.
Pericoma olympia (Kincaid) Haseman, Trans. Am. Ent.

Soc., Vol. 33 (1907), p. 305.

Washington.

Psychoda opposata Banks.

P. opposata Banks, Can. Ent., Vol. 33 (1901), p. 274.
District of Columbia, Maryland, New York.

Psychoda phalaemoides (Linn.) Tonnoir.

Tipula phalcenoides Linnaeus, Syst. Nat. ed. 10, No. 32
(1758), p. 588.

P . phalcenoides (Linn.) Tonnoir, Ann. Soc. Ent. Belg., Vol.
62 (1922), p. 67.

P. pacifica Kincaid, Ent. News, Vol. 8 (1897), p. 143.

P. tonnoiri Dyar, Insec. Inscit. Menst., Vol. 14 (1926), p. 105.
California, Colorado, District of Columbia, Idaho, Maryland,
New Mexico, Oregon, Washington, Wisconsin.

Alberta, British Columbia.

Alaska.

Psychoda pusilla Tonnoir.

P. pusilla Tonnoir, Ann. Soc. Ent. Belg., Vol. 62 (1922),
p. 83.

Kansas, Maryland.

Psychoda quadripunctata Banks.

P. quadripunctata Banks, Ent. Soc. of Wash., Vol. 8 (1907),
p. 149.

Virginia.

Psychoda scotiae Curran.

P. scoticB Curran, Can. Ent., Vol. 56 (1924), p. 216.

Nova Scotia, Quebec.

Psychoda severini Tonnoir.

P. severini Tonnoir, Ann. Soc. Ent. Belg., Vol. 62 (1922),
p. 78.

208

Journal New York Entomological Society

[Vol. Lll

California, District of Columbia, Maryland, Montana, New
Mexico.

British Columbia.

Psychoda sigma Kincaid.

P. sigma Kincaid, Ent. News, Vol. 10 (1901), p. 31.
P.surcoufi Tonnoir, Ann. Soc. Ent. Belg., Vol. 62 (1922),
p. 74.

Washington.

Psychoda signata Banks.

P. signata Banks, Can. Ent., Vol. 33 (1901), p. 274.

District of Columbia, Maine.

Psychoda slossoni Williston.

P. slossoni Williston, Ent. News, Vol. 4 (1893), p. 114.

Maine, New York.

Psychoda snowhilli Del Rosario.

P. snowhilli Del Rosario, Philippine Jour. Sci., Vol. 59
(1936), p. 140.

Maryland.

Psychoda squamosa Johnson.

P. squamosa Johnson, Bui. Amer. Mus. Nat. Hist., Vol. 32
(1913), p. 43.

Psychoda superba Banks.

P. superba Banks, Can. Ent., Vol. 26 (1894), p. 332.

District of Columbia, Maryland, Michigan, New Jersey, New
York, Virginia.

Psychoda superba var. conspicua Del Rosario.

P. superba var. conspicua Del Rosario, Philippine Jour. Sci.,
Vol. 59 (1936), p. 125.

Maryland, Virginia.

Psychoda tridactila Kincaid.

tridactila Kincaid, Ent. News, Vol. 19 (1899), p. 32.
Washington.

Psychoda uniformata Haseman.

P. uniformata Haseman, Trans. Am. Ent. Soc., Vol. 33
(1907), p. 319.

Missouri.

June, 1944]

Rapp: Psychodid^e

209

Psychoda uniformis Del Rosario.

P. uniformis Del Rosario, Philippine Jonr. Sci., 59 (1936),
p. 113.

Maryland.

TRICHOMYIA Haliday

Trichomyia lanceolata Kincaid.

Sycorax lanceolata Kincaid, Ent. News, Vol. 10 (1899), p. 35.
California, Washington.

Trichomyia unipunctata Haseman.

T. unipunctata Haseman, Trans. Am. Ent. Soc., Vol. 33
(1907), p. 323.

Arizona.

June, 1944]

Heming: International Commission

211

RECENT WORK BY THE INTERNATIONAL COMMIS-
SION ON ZOOLOGICAL NOMENCLATURE

By Francis Hemming

Secretary to the International Commission on Zoological
Nomenclature

The International Commission on Zoological Nomenclature are
now engaged in the publication of decisions taken before the out-
break of war in 1939. These decisions have been embodied in
Opinions 134^183 and Declarations 10-12. Of these, Opinions
134-155 and the 3 Declarations have already been published or
are in the press. The remainder are ready for printing and will
be published as soon as funds are available.

These Opinions are of particular interest to entomologists, since
in addition to 9 Opinions relating to the interpretation of various
aspects of the International Code, no less than 38 of these Opinions
are directly concerned with entomological subjects.

The Opinions relating to the interpretation of the Code are :
Opinion 138 (meaning of phrase “definite bibliographic refer-
ence” in Article 25) ; Opinion 141 (naming of families) ; Opinion
145 (status of names first published in invalid works) ; Opinion
147 (generic names of same origin and meaning as older generic
names) ; Opinion 148 (status relating to names published as
emendations of, or substitutes for, older names) ; Opinion 164
(position as regards types when two or more genera are united) ;
Opinion 168 (supplementing Opinion 65 regarding genera based
upon erroneously determined species) ; Opinion 172 (status of
type — designations of genera in abstracts, etc.) ; and Opinion 183
(interpretation of Article 8 relating to form in which generic
names should be published).

Of the Opinions specially concerned with entomological sub-
jects, 6 deal with the status of particular works or with the dates
of such works : Opinion 136 (Latreille, 1810) affects all Orders
of insects; Opinion 135 (“Erlangen List,” 1801) is of special in-
terest to hymenopterists ; Opinions 134 (Freyer, Neue Beitrage),
138 (Hiibner, Samml. exet. Sckmett 1807 and Fabricius Mag.

212

Journal New York Entomological Society

[You Lll

Insektenk. (Illiger) 1807) and 150 (Hiibner, Verz. bek. Schmett)
are concerned with Lepidoptera and Opinion 152 (Meigen, 1800)
with Diptera. Opinions 140 and 143 deal with certain family
names in insects.

The remaining 30 Opinions deal with particular generic names
in various Orders of insects. These Opinions either fix the types
of these genera or add the names to the Official List of Generic
Names in Zoology, or do both. Three of these Opinions are con-
cerned with Orthoptera; 13 with Hymenoptera and 14 with
Lepidoptera.

Bach Opinion is published separately but Opinions are con-
secutively paged to facilitate the publication of an index on the
completion of the volume concerned.

The International Commission are most anxious to secure that
Opinions are published as rapidly as possible but they are greatly
hampered by lack of funds. The Commission therefore appeal to
scientific institutions and individual scientific workers for dona-
tions to a special fund to be used for the issue of publications.
Full particulars of this Appeal are given in Part 2 of the Com-
mission’s Official Organ, the Bulletin of Zoological Nomenclature
published in 1943.

Contributions, however small, will be warmly welcomed and
will be acknowledged in the Bulletin. Bankers’ drafts, cheques,
and money orders should be made payable to the ‘ ‘ International
Commission on Zoological Nomenclature” and sent to the Com-
mission at their Publications Office, 41 Queen’s Gate, London,
S.W.7. All orders for the Commission’s publications should be
sent to the same address. Inquiries relating to the work of the
Commission should be addressed to me at 83 Fellows Road (Gar-
den Flat), London, N.W.3.

International Commission on

Zoological Nomenclature,

Publications Office,

41 Queen ’s Gate,

London, S.W.7.

24th April, 1944.

VoL LII

No. 3

Edited by HARRY B. WEISS

1AL

Publication Committee

HARRY B,

SEPTEMBER, 1944

Journal

of the

New York Entomological Society

Devoted to Entomology in General

WEISS JOHN D. SHERMAN, Jr.

T. C. SCHNEIRLA

Subscription $3.00 per Year

Published Quarterly by the Society
N. QUEEN ST. AND McGOVERN AVE.
LANCASTER, PA.

NEW YORK, N. Y.

1944

CONTENTS

The Remarkable Distribution of an American Cicada:

A New Genus and Other Cicada Notes

By William T. Davis 213

Ephraim Porter Felt

By Stanley W. Bromley 223

Notes on Mexican Butterflies, III

By F. Martin Brown 237

Outdoor Protection from Mosquitoes

By Joseph M. Ginsburg 247

New Neotropical Theclinae (Lepidoptera, Lycaenidae)

By Harry K. Clench 255

Note on the Death-Feint of Bruchus obtectus Say

By Harry B. Weiss 262

Differentiation of Females of Certain Species of Culex
by the Cibarial Armature

By Charles D. Michener 263

Insect Responses to Colors

By Harry B. Weiss 267

Hibernation of the Syrphid Fly, Lathyrophthalmus seneus
Scop.

By E. Gorton Linsley 272

Two New Subspecies of Lycaenopsis pseudargiolus Bdv.

& Lee. (Lepidoptera, Lycaenidae)

By Harry K. Clench 273

New Species of Neopasites with Notes Concerning
Others (Hymenoptera, Nomadidae)

By E. Gorton Linsley 277

The Death-Feints of Alobates pennsylvanica DeG., and
Alobates barbata Knoch. (Coleoptera)

By Harry B. Weiss 281

Insect Food Habit Ratios of New York State

By Harry B. Weiss 283

NOTICE : Volume LII, Number 2, of the Journal of The
New York Entomological Society was published on
July 5, 1944.

Entered as second class matter July 7, 1925, at the post office at Lancaster, Pa.
under the Act of August 24, 1912

Acceptance for mailing at special fate of postage provided for in Section 1103.
Act of October 3, 1917, authorized March 27, 1924.

JOURNAL

OF THE

New York Entomological Society

Vol. LII September, 1944 No. 3

THE REMARKABLE DISTRIBUTION OF AN
AMERICAN CICADA; A NEW GENUS,

AND OTHER CICADA NOTES1

By William T. Davis
Staten Island, N. Y.

QUESADA GIGAS, A REMARKABLE CICADA
(Plate VIII, Figs. 1, 2)

A number of species of North American cicadas occur from
the Atlantic seaboard to about western Kansas, or to the one
hundredth meridian, and northward from Florida and Southern
Texas to Nova Scotia and the region of the Great Lakes. In
Western North America, where there are a greater number of
' species than in the Eastern States and Canada, the species extend
eastward to about the 100th meridian. Two species of Okana-
gana, namely rimosa and canadensis , in the northern part of their
range, where they follow the belt of evergreen forest extending
from Nova Scotia to the far west, have a greater east and west
distribution than the cicadas occurring more to the south. In:
“A Preliminary Review of the West Coast Cicadidse,” 1915, Mr.
E. P. Van Duzee states regarding rimosa: “This species occurs
across the whole continent from Vancouver Island to Quebec
and as far south on the west coast as Fresno Co., California.”

In Texas there are at least 40 species of cicadas and several
additional named forms or varieties. The most famous species
occurring in Texas, is Quesada gigas Olivier, remarkable on
account of its distribution as well as its loud song, which has been
1 1 am indebted to Howard Cleaves for photographing the specimens.

214

Journal New York Entomological Society

[Vol. LII

likened by many observers to the shrill whistle of a first-rate
locomotive. In: “The Naturalist on the Eiver Amazons,” Bates
states: “Added to these noises were the songs of strange cicadas,
one large kind perched high on the trees around our little haven
setting up a most piercing chirrup ; it began with the usual harsh
jarring tone of its tribe, but this gradually and rapidly became
shriller, until it ended in a long and loud note resembling the
steam-whistle of a locomotive engine. Half-a-dozen of these
wonderful performers made a considerable item in the evening
concert. I had heard the same species before at Para, but it
was there very uncommon. ’ ’

The common name for gigas in some localities is the: “Loco-
motive Cicada,” and it is also called the: “Soupbug,” because
it is at times attracted to light in its evening flights and lands on
the supper table.

It is the only known American cicada that may be found in
the mature or winged form every month in the year in some part
of its extensive north and south distribution of about 4,000 miles,
from southern Texas through Mexico, Central America, South
America into Argentina.

In the writer’s collection there are many specimens from
southern Texas, where it has thus far been recorded from May
to October ; there are many records from Mexico ; also from
Central America, and in South America specimens from Co- '
lombia, Venezuela, Trinidad, Tobago Island recorded in Feb-
ruary and March; British Guiana in September (Dr. Beebe and
John Tee-Van) ; Ecuador; Brazil in September, December, Jan-
uary, May; Peru in March, June, September, October; Bolivia
in February; Paraguay in December and January. Specimens
from Argentina are dated November and December, and it doubt-
less occurs at other dates.

In his: “Synopsis of the Cicadidas of Ecuador,” 1925, Dr.
Frederic W. Goding states regarding the Genus Quesada that:
“One species has been recognized in Ecuador, which is greenish
yellow, with a fuscous spot on the bases of the second and third
apical cells of tegmina ; it is one of our largest species. ’ ’

Dr. Kenneth J. Haywood, Chief of the Department of Ento-
mology, Tucuman, Argentina, has informed me that Quesada

Sept., 1944]

Davis: Cicadas

215

gigas, is, speaking generally, distributed over Argentina north
of a line drawn between Buenos Aires and Mendoza. So far
there are no available records for Chile or Uruguay. The Doctor
states that this fine cicada is called: “ ‘Chichara grande’ (chi-
chara is a common name for the cicada here), ‘Coyoyo,’ or more
commonly 4 Coyuyo, ’ according to what part of this vast country
you find yourself in. ’ ’

Considering the extended distribution of the species, the speci-
mens from the various parts of its wide range are surprisingly
alike, but sometimes differ individually in color. Those from
Brazil, Argentina, etc., are often quite large, with abdomen notice-
ably broad in the males, but Texas specimens may also differ con-
siderably in size.

There is a colored figure of the insect under the name of Tym-
panoterpes gigas in “Biologia Centrali- Americana, ” 1881, with
an interesting account of its song and habits. In : “ Insect Sing-
ers, A Natural History of the Cicadas,” Dr. J. G. Myers devotes
considerable space to an account of this remarkable species, its
song and habits.

In his: “Catalogue of the Cicadidse, ” 1906, Mr. Distant cites a
number of specific names that have been bestowed upon Quesada
gigas, and when more specimens have been collected from the dif-
ferent parts of its remarkable range, and additional field studies
have been made, including time of appearance, it may be dis-
covered that there are some definite geographic races or even an
additional species involved.

In his: “Preliminary Survey of the Cicadidse of the United
States, Antilles and Mexico,” 1892, Uhler stated concerning
gigas : “ I have examined specimens from various parts of Mexico,
and from Guatemala, Guiana, and Matto Grosso, Brazil. Speci-
mens from Tamaulipas, Mexico, differ in no respect from others
living further South, although the species is a very variable one
especially in the amount and form of black marking on the upper
surface of the body. The region of the Bio Grande of Texas is
the most northern limit of this species, while the northern part of
the Argentine Republic seems to be its most southern habitat.”

In: “Notes Del Museo de la Plata,” Tomo V, Buenos Aires,
1940, Prof. Belindo Adolfo Torres described and figured a dark-

216

Journal New York Entomological Society

[Vol. LII

colored female of gig as, but as be could find no structural differ-
ence he considered the variety unworthy of a name.

In Texas Quesada gigas has been recorded from Starr, Hidalgo,
Cameron and Bexar counties by Mr. F. F. Bibby, and no doubt it
will be discovered over a wider range. There are specimens from
Kingsville, Kleberg County, in the collection of Cornell Uni-
versity.

Mr. H. B. Parks, Director of the State Agricultural Research
Laboratory near San Antonio, Bexar County records gigas as a
yearly visitor observed since 1934. The “Popcorn Whistlers”
occur in the live-oaks about the Laboratory, usually in July and
August, and he heard them singing in the evening and sometimes
in the early morning in 1941. He has found dead specimens
beneath the trees. He also reports their abundance in the coastal
city of Corpus Christi, Nueces County, in August, 1940. The
little boys in town found great sport in climbing trees and catch-
ing the cicadas. They had also been observed and collected in
1938 at Corpus Christi, and I received specimens from both Mr.
Parks and Mr. Emmett S. Claunch, Jr., who reported that they :
“whistle instead of buzz — that is they sound as though they
whistle.”

Many collectors have found gigas about Brownsville, Cameron
County, and Dr. James A. G. Rehn, of the Philadelphia Academy,
likened its song to the shrill tin whistle of a peanut roaster.

Dr. Raymond H. Beamer and associates from the University
of Kansas collected in Bee County and Hidalgo County in July,
1928, and in sending specimens the Doctor wrote that they had
70 more if I cared to see them. This, as well as some of the other
facts are mentioned to show what a highly successful species
gigas really is, both in numbers as well as in wide distribution,
for in some part of its range from north to south, a male gigas
is in song every month in the year.

Mr. Paul C. Avery of Mission, Hidalgo County, near the Rio
Grande, has sent me a great many gigas, which sometimes occurs
very plentifully along the river where the soil is more or less
damp. He has found them often on Mesquite, and describes the
song as: “Very loud, continuous and shrill. The loudest of any
species found,” at Mission. He collected many in 1935, and in

Sept., 1944]

Davis : Cicadas

217

1936 sent me as a sample 400 specimens — 247 males and 153
females — collected in July of that year. He observed the first
gigas on June 13, and in his letter of July 5, 1936, stated that
they sang both early and late, and often were heard singing after
dark. He heard the last one on September 21 in 1936. Mr.
Avery has also observed the Cicada killer, Sphecius, with a gigas ,
which : ‘ ‘ sure was crying loud and mournfully. ’ ’ This species like
many other cicadas, is subject to a fungus disease, and a number
of specimens have been received with the terminal segments of the
abdomen missing, as often happens when the Seventeen-year
Cicada is attacked by fungus.

It will be seen from the foregoing, that in Quesada gigas the
United States includes in its fauna one of the most remarkable
of the known cicadas, which species is sure to attract more and
more attention in the years to come.

Cornuplura, new genus.

In the Journal of the New York Entomological Society for June,
1936, Tibicen curvispinosa was described and figured as a remarkable cicada
from western Mexico. The two conspicuous upturned spines in the male pro-
truding backward from segment IX, were described and figured, and it was
stated that : 1 ‘ the uncus in curvispinosa is deeply cleft with the resulting two
claw-like extremities long and curved inward. ’ ’ These characters are also
noted by Smith and Grossbeck in : “ Studies in Certain Cicada Species, ’ ’
Entomological News, April, 1907, and shown in their figures 7 and 8 on
plate 3.

Tibicen nigroalbata was also described in the 1936 paper, and its resem-
blance to curvispinosa noted. Only a single female nigroalbata taken by
Prof. E. D. Ball in Arizona was available in 1936.

In the Journal of the New York Entomological Society for June,
1942, these remarkable cicadas are again considered in the light of addi-
tional specimens from Nayarit, Mexico, and Santa Cruz County, Arizona, and
it was suggested that nigroalbata might be a northern race of the southern
curvispinosa.

The general form of the body except for the curved spines in the males of
both curvispinosa and nigroalbata, including the position of the cross-veins
in the fore wings, and the terminal central spine on the last dorsal segment
in the males, is as in the genus Tibicen, but the deeply cleft and curved uncus
is not as found in the other species of that genus native to North America,
nor as in plebeja Scopli, of Europe, the type of the genus. It is suggestive
of the uncus in some species of Diceroprocta, but in that genus the dorsal
segment in the male terminates in two lateral lobes and the venation is
different.

218

Journal New York Entomological Society

[Vol. LII

It would appear from the above that a new genus should be erected for
these remarkable insects with curvispinosa as a type, to be placed between
Tibicen and Diceroprocta, for which the name Cornuplura is here proposed.
The genus may also include rudis Walker, from Mexico, as a closely related
species.

The student is referred to several of the structural characters illus-
trated in the above-mentioned three papers, as characteristic of the Genus
Cornuplura.

OKANAGANA SYNODICA (SAY), ITS HABITS, DISTRIBUTION,
AND A NEW COLOR FORM

In 1825 Thomas Say described his Cicada synodica, and stated
that: “Dr. James and Mr. Peale observed this species in great
numbers in one locality at the base of the Rocky Mountains but
it did not occur elsewhere.” He described the body as black
above, and also enumerated the extensive testaceous colored lines
and spots that generally give a number of the insects when seen
together a yellow-brown appearance. Say states : “Scutel [meso-
notum] with a lateral marginal line the elevated X and two dorsal
dilated lines testaceous; the dorsal lines are merely emarginate
on the inner side, and do not form the W ; at the tip of each
anterior line of the X is a conspicuous, black impressed puncture,
and behind the X the posterior edge of the scutel [metanotum] is
visible and testaceous : beneath very pale testaceous. . . . Length
to the tip of the hemelytra less than one inch.”

In the Kansas University Science Bulletin, March, 1920, p. 345,
Dr. P. B. Lawson, in: “The Cicadidse of Kansas,” records syn-
odica from the western part of the state only, and describes it
as : “A small black and honey-yellow species, ’ ’ with length of
body 15 to 18.5 millimeters, and expanse of fore-wings 38 to
44 mm.

Mr. Joseph Duncan Putnam in his : “Remarks on the Habits
of Several Western Cicadae,” Pro. Davenport Academy of Nat-
ural Sciences, March, 1881, records that: “ Cicada synodica Say,
was quite common on the grassy plains near Denver and Boulder,
in Colorado, in June, 1872. The male makes a tolerably loud
rattling noise.” In June, 1920, the late Dr. Lutz, of the Amer-
ican Museum of Natural History, collected four male synodica
at Medicine Bow, Wyoming, about 6,600 feet, and recorded that :
“The small brown cicada in grass has a continuous note, but
sometimes continues for only a short time.”

Sept., 1944]

Davis: Cicadas

219

In June and July of 1935 a brood of this species appeared
in Colfax County, New Mexico. In the writer’s collection there
are 135 specimens representing this brood, and they are all of the
typical brownish-colored form.

In his account of: ‘ ‘ Characteristics of Certain Western
Cicadas,” Jour. N. Y. Ento. Soc., June, 1940, Dr. John W.
Sugden records that : “A large brood of Okanagana synodica was
observed in Emery County, Utah (June 7, 1928). At first, the
sound was mistaken for the humming of the carburetor of the
car. In the field, so many were buzzing that it was impossible
to locate the position of any individuals. After examining the
low, sage-like bushes, thousands of the insects were found.
Fifteen or twenty could be easily collected on a bush not over
a foot high. The note similar to the typical Okanagana song, was
not very loud, but shrill and long continued and what the indi-
vidual lacked in volume was made up for by the large numbers.
Very few would fly if disturbed, but would remain on the bushes
and could be collected by hand. Others would become quiet when
disturbed and fall to the ground, where their color closely re-
sembled the buff-colored soil. Many were mating. The exuvia
were on the ground or attached to the stems. ’ ’

The known range of synodica has been greatly extended in
recent years, and specimens have been examined from Alberta,
Montana, North and South Dakota, Wyoming, Nebraska, western
Kansas, Colorado, Utah, western Texas, New Mexico and Arizona.
It should also be found in Oklahoma as some of the known locali-
ties are close to the state line. About 600 specimens have been ex-
amined, and it has been observed as the specimens accumulated
that the broods appearing in Arizona and New Mexico were some-
times composed of darker-colored individuals than the broods
occurring in Colorado and other more eastern localities.

On May 26, 1941, Mr. Frank H. Parker collected a great many
almost entirely black-bodied individuals at Holbrook, Navajo
County, eastern Arizona, and wrote as follows concerning them :
“The Holbrook series was taken on a large, slightly rolling mesa
covered chiefly with bunch grass, Gutierrezia, and a low (1 foot
or less) shrub somewhat resembling Fairies Feather Duster,
among which was to be found an occasional diminutive Opuntia.

220

Journal New York Entomological Society

[Vol. LII

The capacity of my cyanide jars, and time, were the only factors
preventing the capture of many thousands of this species. ’ ’ Mr.
Parker sent 67 males and 33 females from this brood.

In the writer’s collection there are also two females of the dark
form collected at Holbrook, May 22, 1934, and a male and female
of the same form from the White Mountains, Arizona, July, 1935.
Broods of both the light and dark forms occur in New Mexico,
and both dark and light colored specimens have been received
from Utah, collected by Dr. Sugden, June 7, 1928, in Emery
County.

It will be noted that broods of the dark, or almost wholly
black individuals, generally occur in the area drained by the
Colorado, while those of the lighter-colored form described by
Say, are on the more eastern watershed, or streams tributary to
the Rio Grande, Missouri, etc.

Okanagana synodica (Say) variety nigra, new variety (Plate VIII, tig. 3).
Type male and allotype female from Holbrook, Arizona, May 26, 1941
(Frank H. Parker). Davis collection.

In this dark or melanistic form of synodica, the testaceous markings as
described by Say are absent or very much reduced. The head is entirely
black save for two pale spots one above each antenna, and the ruby-colored
ocelli. The pronotum is narrowly edged all around with orange, and has a
short, median pale line extending to the anterior margin. The mesonotum
has a very small orange spot at the base of each fore wing ; the posterior
margin, including the X, pale, and the two torch-shaped orange marks, ex-
tending forward from the anterior limbs of the X generally lack the often
conspicuous tooth-like mark or sinuation on the inner side found in typical
synodica. The metanotum is pale and the abdominal segments are black or
very narrowly margined posteriorly with orange. In the female the seg-
ments at the end of the body are more broadly margined with orange. The
upper surface of the uncus of the male is black, and the valve pale. The
venation is darker than in typical synodica.

OKANAGANA PALLIDULA DAVIS ; ADDITIONAL NOTES
AND OBSERVATIONS

(Plate VIII, Fig. 4)

In the Journal of the New York Entomological Society for
September, 1938, there is an account of this species, its distribu-
tion and color forms, and the variety nigra is described and re-
corded from Yolo and Kern counties, California. Pallidula can
be confused with some of the forms of Okanagana vanduzeei

Sept., 1944]

Davis: Cicadas

221

which, however, are usually conspicuously hairy on the head, ancl
often on the pronotum, as well as on the under side of the ab-
domen. Also pallidula is duller and does not present the shin-
ing appearance usual in vanduzeei and its forms. While there
are dark specimens of pallidula there are also green ones, which
apparently do not occur in vanduzeei and its varieties eonsoibrina,
etc.

Okanagana pallidula has thus far been examined from the
great Central Valley of California, and from the following coun-
ties: Sutter, Yuba, Yolo, Sacramento, Contra Costa, Merced,
Madera, Fresno, Tulare, Kings, Kern and San Luis Obisbo. In
some years it occurs in great numbers.

As reported on page 308 of the September, 1938, paper referred
to above, this small cicada may be found singing from its hole in
the ground. In June, 1941, Mr. and Mrs. J. N. Knull of the
Ohio State University, while in the Santa Maria River Valley,
California also found specimens of pallidula singing from their
burrows where it was believed they had undergone their last trans-
formation. If the males ultimately found mates, they prob-
ably in due time left the burrows, or perhaps the females flew to
them as has been observed in other species. This is a matter for
future observation and record. Dr. Charles D. Michener of the
American Museum of Natural History, while looking over my
collection of cicadas in 1942, assured me that he had discovered
males of pallidula singing from their one-time burrows, thus
adding to the observations made by others.

222

Journal New York Entomological Society

[Yol. lii

Plate YIII

Figure 1. Quesada gigas (Olivier). Male from Texas.
Figure 2. Quesada gigas (Olivier). Female from Argentina.
Figure 3. Olcanagana synodica (Say) variety nigra. Type.
Figure 4. Olcanagana pallidula Davis.

(Jour. N. Y. Ent. Soc.), Vol. LIT

(Plate YIII)

(Jour. N. Y. Ent. Soc.), Vol. LII

(Plate IX)

EPHRAIM. PORTER FELT

Sept., 1944]

Bromley: Felt

223

EPHRAIM PORTER FELT— 1868-1943

The New York Entomological Society as well as the entire
scientific world has lost one of its most respected and outstanding
members in the passing of Dr. E. Porter Felt. He was a leader
in the field of entomology and was recognized as such inter-
nationally. He was widely known as a scientist, writer and lec-
turer. As State Entomologist of New York for thirty years his
valuable publications were looked upon as standard for this type
of work. As Director of the Bartlett Tree Research Laboratories
from 1928 until his death, he became a world authority on the
care of shade trees. Real leaders are few in any walk of life.
Dr. Felt was one of the few but his influence was valued by many.

Ephraim Porter Felt was born at Salem, Massachusetts, Janu-
ary 7, 1868, a son of Charles Wilson Felt and Martha Seeth
(Ropes) Felt. His background was like one of many similar old
New England families. As a youth he had decided upon the
ministry as a career but became interested in insect problems
while attending college and this became the basis of his life work.
He received the degree of B.Sc. from the Massachusetts Agricul-
tural College in 1891 and also a degree of B.Sc. from Boston Uni-
versity the same year. The degree of D.Sc. was bestowed upon
him by Cornell University in 1894. After teaching Natural Sci-
ence at The Clinton Liberal Institute, Fort Plain, New York,
from 1893-1895, he became Assistant to the State Entomologist
at Albany. Upon Dr. Lintner’s death, he became State Ento-
mologist of New York, which post he held for thirty years, until
1928 when he came to the Bartlett Tree Research Laboratories as
Director and Chief Entomologist.

In the field of insect taxonomy he achieved international fame
for his careful and extensive studies on the . gall midges — the
Cecidomyidse or Itonididas as he preferred to call them. In 1913
he gave the annual public address on Gall Insects before the Ento-
mological Society of America at Atlanta, Georgia. In 1914 he
was elected a member of the American National Committee on
Nomenclature of the Second International Congress. I have
learned from Professor C. P. Alexander of Massachusetts State

224

Journal New York Entomological Society

[Vol. LII

College that Dr. Felt had reported to him in 1942 that he had
described 1,060 new species of gall midges and plant mites. With
the late D. B. Young, his assistant at Albany, he had described
a number of mosquitoes which are indicated in the literature
under the joint authorship of Felt and Young.

Dr. Felt became a corresponding member of the New York
Entomological Society on October 20, 1900, and an active member
on February 5, 1907. After coming to Stamford in 1928 he fre-
quently presented papers at the New York Entomological Society
meetings which included subjects of a varied nature from gall
midges to poetry concerning insects, as well as numerous valuable
contributions on shade tree insects.

Dr. Felt’s first entomological work was in Massachusetts in the
early days of the gypsy moth invasion. His experience with this
destructive forest and shade tree pest led him in 1924 in his work
for the New York State Conservation Commission to propose a
barrier zone two hundred miles in length to stem the spread of
the gypsy moth into New York State from New England.

Dr. Felt was a pioneer in airplane collection of insects and
made extensive studies on windborne insects. He made studies
on the wind drift of insects on the top of the State Education
Building, Albany, N. Y., the Empire State Building in New York
City, as well as on other tall city buildings and his work on wind-
borne insects has been very important in ascertaining the direc-
tion of spread of the Dutch elm disease which is carried to a large
extent by the European elm bark beetle.

In the late 90 ’s and early 1900 ’s, Dr. Felt studied the spread
of many diseases by the common house fly and by mosquitoes and
made important contributions to this phase of entomology. In the
February, 1944, issue of the National Geographic there was an
article “Saboteur Mosquitoes” by Mr. Stage, Senior Entomolo-
gist of the United States Bureau of Entomology and Plant
Quarantine, in which is recorded some of the work done on mos-
quitoes years ago under the direction of Dr. Felt.

Dr. Felt was very much interested in and very capable of doing
editorial work and was editor of the Journal of Economic Ento-
mology since its start in 1908. At one time he was editor of the
National Shade Tree Conference Reports.

Sept., 1944]

Bromley: Felt

225

While State Entomologist of New York he published 25 official
reports covering the activities of that office and many bulletins,
which were published by the New York State Museum, as well as
more than 700 timely papers and articles which included a wide
variety of entomological work comprising general entomology as
well as popular articles of an informative nature for the general
public. He was a frequent contributor to entomological journals
and scientific publications, as well as magazines and newspapers.
He found time to give numerous addresses, to publish several
popular books and to speak over the radio on different phases of
entomological and shade tree activities. His published reports
and bulletins were models of systematic arrangement and clarity
of expression.

During his long and useful career, Dr. Felt had been a collabo-
rator of the United States Bureau of Entomology, the entomologi-
cal editor of the Country Gentleman from 1898 to 1911 and a
member of the Supervisory Board of the American Year Book.
He was a past president of the American Association of Economic
Entomologists, a past president of the National Shade Tree Con-
ference, a fellow of the Entomological Society of America, a life
member of the American Entomological Society, a member of the
New York and Washington Entomological Societies and of Sigma
Xi and Phi Kappa Phi. Among the honors accorded him were a
gold, and three silver medals at the Pan-American Exposition,
Buffalo, N. Y., in 1901. He was a member of the Fraternity of
Alpha Sigma Phi.

For the last twelve years, he was the author of a weekly syndi-
cated newspaper article entitled “ Talks on Trees” which was used
by nearly three hundred newspapers throughout the country.
He was the author of a number of well-known books on insects
and trees, his two-volume work “Insects Affecting Park and
Woodland Trees” appearing in 1906 and “Manual of Tree and
Shrub Insects” in 1923. He was co-author with Dr. W. H.
Kankin of “Insects and Diseases of Ornamental Trees and
Shrubs” published in 1932. His “Plant Galls and Gall Makers”
appeared in 1940.

Three outstanding books on shade trees were published more
recently — 1 1 Our Shade Trees, ’ ’ in 1938, a second edition of which

226

Journal New York Entomological Society

[Vol. LII

appeared in 1942; “Pruning Trees and Shrubs,” in 1941, and
“Shelter Trees in War and Peace,” in 1943.

On December 8, 1943, Dr. Pelt attended the national meetings
of the American Association of Economic Entomologists held in
Columbus, Ohio. On his way back he stopped at Utica, N. Y.,
to visit his son. He was back at his office on Monday, December
13. He seemed none the worse for the rigorous trip and was in
excellent spirits. On Tuesday morning, December 14, he told his
secretary that the night before he doubted whether he would be
in that day. He said he felt as though he was coming down with
the flu the night before but felt all right that morning. With a
twinkle in his eye, he said that he had had such symptoms before
and that nothing came of them. “You see,” he said, “I am still
pretty tough.” These were the last words that Miss Burns, his
secretary, heard him speak. I talked with him a few minutes
later when he was going out to the Ferguson Library to look up
some references and he seemed to be all right then. Later he
came back to the building and died very suddenly of a heart
attack.

Surviving him, in his immediate family, are three daughters
and a son, a sister and two brothers, and thirteen grandchildren.

A few days after Dr. Felt had passed away, I received a letter
from Dr. Peairs, present editor of the Journal of Economic Ento-
mology, recounting his conversation with Dr. Felt during the
Columbus meetings. I would like to quote that part of Dr.
Peair’s letter.

“I cannot refrain from telling you a little episode in Columbus ;
I had a chat with Dr. Felt and he brought up a discussion of some
of the members who had recently died ; I casually mentioned the
fact that four ex-presidents had died in the last few months. Dr.
Felt, with his faint smile, remarked, not seriously at all, ‘I am
hereby resigning my position as an ex-president of the Associa-
tion, effective at once. ’ I am sure he thought of it as nothing but
a little joke, as I did. But somehow, I almost wish the subject
had not come up. ’ ’

Dr. Felt was a great scientist, a splendid gentleman and an
invaluable friend. Honest, sincere and upright, he moved
through life with a tranquil demeanor and faith in his fellow

Sept., 1944]

Bromley: Felt

227

men. Trivialities neither upset him nor deterred him from any
worthy purpose or goal.

His prime thoughts were for the younger, newer men. All were
stimulated by his great personality. He aided their efforts with
cheer and lent a helping hand no matter how adverse the diffi-
culty or how uncertain the going.

I first met Dr. Felt in 1920. I have known hinqas a friend, as
a counsellor, as a superior and as a leader. Since 1929, it was
my privilege and honor to work with him side by side on impor-
tant research and I have never ceased to marvel at his deft ability
to size up and conquer a problem and at his unruffled composure
in the face of trying obstacles.

Of him I can use no greater expression than to say that he was
an all around great American. His flexibility of outlook was
manifested by the ability to direct itself in a flash from the deep
abstruse contemplation of science to the broad generalities of
national life. He was intensely interested in the world about him.
His human ties were broad and comprehensive. His love for
children was apparent to many.

He was an avid reader, and enjoyed the radio from the political
speeches of the great to the humor of Charlie McCarthy. He had
a deep and abiding sense of humor and was quick and accurate
at repartee. A remark was once made on his winged collar, so
characteristic of his dress. Dr. Felt immediately replied 1 1 What
could be more fitting to my profession?” pointing to the two-
winged insects he was studying.

He was faithful to the minute to his commitments. Many were
the occasions when he accepted an invitation to talk at some
humble, out-of-the-way garden club or other meeting. If he said
he would be there, he was always present. This custom alone
stamped him as a man of his word in the eyes of many of his great
circle of friends.

His appearance as well as his character was left unchanged by
the surging tide of years and he was until the last the same stead-
fast, striking figure. His carefully trimmed, white hair and Van
Dyke beard, his calm, upright bearing, his impeccable dress, his
gentle but firm voice, all reflected the imprint of his great
character.

228

Journal New York Entomological Society

[Vol. LIT

How frequently and forcefully the realization has come to me
during the past two months that we will never again experience
together the exhilaration of solving an important problem, or of
unravelling one of nature’s intricacies again; that no more will
we talk over together the problems of the present or the plans of
the future ; that no more will I have his steadfast character upon
which to lean or his guiding hand to point the way !

While his passing has been such a personal loss to me, how
much greater has been the loss to the profession of entomology
and to the entire world of science. His contributions will loom
in importance during the passing years even as the memory of his
personality may dim and fade.

The accompanying bibliography, transcribed from the card
index by his secretary, Miss Beatrice Burns, gives his publications
in chronological order. It is an inescapable conclusion that such
a bibliography cannot be complete. Dr. Felt was accredited with
having written more than seven hundred important and timely
papers and articles which include a wide variety of entomological
works.

The following may be considered his outstanding writings :

BIBLIOGRAPHY OF DR. E. P. FELT

1893. Nomophila noctuella. Can. Ent. 25 : 129.

1894. On certain grass-eating insects. Cornell Agric. Exp. Sta. Bull. 64.
1896. The scorpion flies. (10th Bep’t) N. Y. State Ent. for 1894.

1901. Suggestions toward greater uniformity in nursery inspection laws and

rulings. Cony, of the Assoc, of Amer. Agr. Colleges and Ex-
periment Stations, Bui. No. 99.

A 1902. N. Y. State Mus. Bui. 53 (17th Rep’t), 705-30, 741-44.

1902. Crude petroleum as an insecticide. Twenty-Third Meet. Soc. Prom.

Agr. Sc. p. 1.

1903. Literature of American economic entomology. Bureau of Entomology

Bui. 40, new series, 7—22.

1904. (With L. H. Joutel) Monograph of the genus Saperda. N. Y. State

Mus. Bull. 74, 1-80.

1904. Importance of isolated rearings from culicid larvge. Ent. Soc. Wash.
Proc. VI, Vol. XX, 312-13.

1904. The mosquitoes or culicidse of New York State. N. Y. State Mus.
Bull. 79, 241-400, 113 fig., 57 pi.

1904. The cause and control of insect depredations. Twenty-Fifth Ann.
Meet. Soc. for Promotion Agr. Sc., p. 73.

Sept., 1944]

Bromley: Felt

229

X 1905.
1906.

1906.

X 1906.

X 1906.

1906.

1907.

1907.

1907.

1907.

1907.
& 1907.

X 1908.

1908.
1908.

? 1908.
1908.
1908.

1908.

1908.

1910.

1910.

1910.

1910.

1910.

1911.
1911.

1911.
1911.
1911.
1911.
1911.
1911.
XL 1911.

Cecidomyia hirtipes. N. Y. State Mus. Bull. 97 (Rep’t 1904), 410-11.
Diversities among New York mosquitoes. Proc. 2nd Anti-Mosquito
Conv. New York City, 1904.

Mosquito control. N. Y. State Mus. Bull. 104 (21st Rep’t), N. Y.
State Ent., 1905.

Studies in Cecidomyiidae. N. Y. State Mus. Bull. 104 (21st Rep ’t,
1905), 116-32.

Plant galls & gall makers. N. Y. State Mus. Memoir 8, 2: 615-48.
Insects affecting park & woodland trees. N. Y. State Mus. Mem. 8,
vol. 1, 1-332.

Hort. diseases and pests. N. Y. State Ed. Dept. Rev. of Legislation,
119-22.

New species of Cecidomyiidae. N. Y. State Mus. Bull. 110, 97-165.

Separate, 1-53. Lingnan Science Jour., Yol. 7., 469.

Gall gnats or Cecidomyiidae. Can. Ent., 39: 143-44.

Cecidomyiidae : A statement. Can. Ent., 39 : 197-98.

Cecidomyia acarivora n. sp. Ent. News, 17 : 242.

New species of Cecidomyiidae II, p. 1-23. Same in N. Y. State Mus.
Bull. 124, 286-304,

Cecidomyia johnsonii Sling. Jour. Econ. Ent., 1: 243.

Contarinia gossypii n. sp. Ent. News, 19 : 210-11.

Insect control in its larger aspects. Fruit Growers Proceedings, p. 1.
Some problems in nomenclature. Ann. Ent. Soc. of America, p. 102.
Observations on the genus Contarinia . Jour. Econ. Ent., 1: 225-28.
Observations on the biology and food habits of the Cecidomyiidae.
Jour. Econ. Ent., 1: 18-21.

Studies in Cecidomyiidae II. N. Y. State Mus. Bull. 124 (23d Rep’t),
286-304.

New species of Cecidomyiidae II. N. Y. State Mus. Bull. 124, 307-510.
Schizomyia ipomoece n. sp. Ent. News, 21: 160-61.

Gall midges of Aster, Carya, Quercus and Salix. Jour. Econ. Ent.,
3: 347-56.

West Indian Cecidomyiidae. Ent. News, 21: 268-70.

Two new Cecidomyiidae. Ent. News, 21: 10-12.

Observations on the house fly. Jour. Econ. Ent., 3: 24-26.

A generic synopsis of the Itonidae. Jour. N. Y. Ent. Soc., 19: 31-62.
Summary of food habits of American gall midges. Ann. Ent. Soc.
America, 4: 55-62.

Endaphis Kieff., in the Americas (Dipt.). Ent. News, 22: 128-29.
Endaphis hirta n. sp. (Dipt.). Ent. News, 22: 224.

Two new gall midges. Can. Ent., 43: 194-96, June.

Four new gall midges (Dipt.). Ent. News, 22: 301-5.

Hosts and galls of American gall midges. Jour. Econ. Ent., 4: 451-75.
New species of gall midges. Jour. Econ. Ent., 4: 476—84; 546-59.
Miastor americana Felt, an account of pedogenesis, N. Y. State Mus.
Bull. 147 (26th Rep’t), 82-104.

230

Journal New York Entomological Society

[Yol. lii

1911. A new species of Lasioptera with observations on certain homologies.
Psyche, 18: 84-86.

¥• 1911. New Indian gall midges. Mem. Dept. Agr. India Ent. Ser. VII, No.
4, 23-28.

1911. A new Lestodiplosis. Ent. News, 22 : 10-11.

1911. Two new gall midges (Dipt.) Ent. News, 22: 109-11.

1911. Three new gall midges. Jour. N. Y. Ent. Soc., 19: 190-3.

1911. Rhopalomyia grossularicB. Jour. Econ. Ent., 4: 347.

1912. New W. Indian gall midges (Dipt.). Ent. News, 23: 173-7.

1912. Observations on Uleella Rubs. (Dipt.). Ent. News, 23: 353-54.

1912. Lasiopteryx manihot n. sp. Can. Ent. 44: 144.

1912. Studies in Itonididse. Jour. N. Y. Ent. Soc., 20: 236-48.

1912. Observations on the identity of the wheat midge. Jour. Econ. Ent.,
5: 286-89.

1912. The gall midge fauna of western North America. Pomona Coll. Jour.
Ent., 4: 753-57.

1912. The identity of the better known midge galls. Ottawa Nat., 25:
65-67, 181-88; separate, 1-11.

1912. Biology of Miastor and Oligarces. Science, 33 : 278-80.

1912. Diarthronomyia calif ornica n. sp. Pomona Coll. Jour, of Ent., 4: 752.
1912. Petroleum & petroleum products as insecticides. N. Y. State Ed.
Dept., Sc. Div. 2 p.

1912. Itonida inopis. Jour. Econ. Ent., 5: 368—69.

1912. Control of insect pests in institutions. Jour, of Home Econ., Yol.
IY, No. 1, 16-26.

1912. New Itonidse. Jour. N. Y. Ent. Soc., 20: 102-7.

1912. New gall midges or Itonidse (Dipt.). Jour. N. Y. Ent. Soc., 20: 102-7.
1912. New gall midges or Itonidse (Dipt.). Jour. N. Y. Ent. Soc., 20:
146-56.

1912. Arthrocnodax occidentalis. Jour. Econ. Ent., 5: 402.

1912. Observations on the identity of the wheat midge. Jour. Econ. Ent. 5:

286-9.

1913. Cystodiplosis eugenice n. sp. Ent. News, 24: 175-76.

1913. Adaptation in the gall midges. Can. Ent. 45: 371-79; also Ent. Soc.

Ontario (44 Rep’t), 1913, 1914, 76-82.

1913. The gall midge fauna of New England. Psyche, 20: 133-47.

1913. A study of gall midges. N. Y. State Mus. Bull. 165 (28th Rep’t),
State Ent., p. 127-227.

1913. Three new gall midges (Dipt.). Can. Ent. 45: 304-8.

7C 1913. A study of gall midges. N. Y. State Mus. Bull. 175 (29th Rep’t),
79-214.

1913. Gall midges in an aquatic or semiaquatic environment. Jour. N. Y.
Ent. Soc., 21: 62-63.

1913. Table of hickory leaf midge galls. Bui. Bklyn. Ent. Soc., 8 : 98-99.
1913. Two new Canadian gall midges. Can. Ent., 40 : 417-18.

1913. Didactylomyia capitata n. sp. Psyche, 20: 174.

Sept., 1944]

Bromley: Felt

231

1913. Descriptions of gall midges (Dipt.). Jour. N. Y. Ent. Soc., 21:
213-19.

1913. Itonida anthici n. sp. Jour. Econ. Ent., 6: 278-79.

1913. Gouty pine midge. Jour. Econ. Ent., 6: 278-79.
x 1913. The goldenrod and its gall flies. Guide to Nature, 6: 149-51.

1913. Arthrocnodax Carolina. Jour. Econ. Ent., 6: 488-89.

1914. Cecidomyiidae by J. J. Kieffer. Fasc. 152 in Genera Insectorum.

Ent. News, 25: 185-88.

1914. Additions to the gall midge fauna of New England. Psyche, 20:
109-14.

1914. Acaroletes pseudococci. Jour. Econ! Ent., 7: 148-49.

1914. Diadiplosis coccidivora. Entomologist, 47 : 86.

1914. Aplonyx sarcobati n. sp. Pomona Jour. Ent. & Zool., 6: 93—94.
1914. Gall midges as forest insects. Ottawa Nat., 28: 76-79.

1914. List of zoophagous Itonididse. Jour. Econ. Ent., 7 : 458-59.

X 1914. Cactus midge, Itonida opuntice. Prickly Pear Trav. Comm. Rep’t,
77-78.

¥ 1914. Notes on forest insects. Jour. Econ. Ent., 7: 373-75.

1914. Additions to the gall midge fauna of New England. Psyche, 20:
109-14.

1914. Descriptions of gall midges. Jour. N. Y. Ent. Soc., 22: 124-34.

1914. New gall midges (Itonidae). Insec. Inscit. Menstr., 2: 117-23.

1914. Hormomyia bulla n. sp. Can. Ent., 46: 286-87.

1914. Conical grape gall Cecidomyia viticola. Jour. Econ. Ent., 7 : 339.

1914. Arthrocnodax constricta. Jour. Econ. Ent., 7: 481.

1915. New genera and species of gall midges. U. S. Nat. Mus. Proc., 48:

195-211.

1915. New South American gall midges. Psyche, 22: No. 5.

1915. The gall midges of the pine. Bull. Bklyn. Ent. Soc., 10: 74-76.

X 1915. A study of gall midges II. N. Y. State Mus. Bull. 175 (29th Rep’t),
79-213.

i. 1915. A study of gall midges III. N. Y. State Mus. Bull. 180 (30th Rep’t),
127-289.

1915. New Asian gall midges. Jour. N. Y. Ent. Soc., 23: 73-84.

1915. Fumigation for the box leaf miner. Jour. Econ. Ent., 8: 94-95.

1915. Mycodiplosis macregori n. sp. Jour. Econ. Ent., 8: 149.

* 1915. A new chrysanthemum pest. Am. Florist, 44 : 162.

1915. New North American gall midges. Can. Ent. 47 : 236-42.

X 1915. New gall midges. Jour. Econ. Ent., 8: 405-9.

■K 1916. A study of gall midges III. N. Y. State Mus. Bull. 180 (30th Rept’t),
127-288.

A 1916. A study of gall midges IV-. N. Y. State Mus. Bull. 186 (31st Rep’t),
101-73.

1916. New western gall midges. Jour. N. Y. Ent. Soc., 24: 175-96.

1916. Gall midges of certain Chenopodiaceae (Dipt.). Ent. News, 27: 201-3.

1916. New North American gall midges. Ent. News, 17 : 412-17.

232

Journal New York Entomological Society

[Vol. LII

1916.
H 1916.

1916.
■V 1916.
* 1916.

1917.
1917.
1917.
1917.

^ 1917.

1917.
X 1918.

1918.

X1918.
f 1918.

A 1918.
-V 1918.

/ 1918.

1919.
X 1919.
X 1919.

-f 1919.

1919.

1920.
4. / 1920.

1920.

1920.

1920.
* 1921.

x( 1921.

X 1921.

X 1921.

New Indian gall midges. Can. Ent., 48 : 400-6.

Distribution of gall midges. Natl. Acad. Sci. Proc., 3 : 349-54.

New gall midges. Can. Ent., 48 : 29-34.

Lasioptera fructuaria. Maine Agr. Expt. Sta. Bull. 244, 268-69.

American insect galls. Ottawa Nat. 30 : 37-39.

Indian gall midges (Cecidomyiidse, Dipt.). Ent. News, 28: 73-76.

Entomological research and utility. Scientific Monthly, p. 551.

New gall midges. Jour. N. Y. Ent. Soc., 25: 193-96.

Distribution of gall midges. Natl. Acad. Sci. Proc., 3: 349-54.

Little known gall midges of certain composites. Ottawa Nat., 31:
13-14.

Asphondylia websteri n. sp. Jour. Econ. Ent., 10: 562.

A study of gall midges Y. N. Y. State Mus. Bull. 198 (32d Rep’t),
101-252.

Notes and descriptions of Itonididse in the collections of the American
Mus. of Natural History. Am. Mus. Nat. Hist., 38 (Oct. 6) :
179-82.

Gall insects and their relation to plants. Sci. Mon., 6 : 509-25.

New Philippine gall midges, with a key to the Itonididse. Phil. Jour.
Sci., 13: 281-325.

Key to American insect galls. N. Y. State Mus. Bull. 200, 1-310.

A study of gall midges YI. N. Y. State Mus. Bull. 202 (33d Rep’t),
76-205.

New gall midges. Jour. Econ. Ent., 11: 380-84.

Five non-gall making midges. Ent. News, 30 : 219-23.

New Philippine gall midges. Jour. Sci., 14: 287—94.

The plant galls collected by the Canadian Arctic Expedition 1913-18.
Kept. Can. Arctic Exped., Y. 3.

Insect galls & gall insects. Ottawa Nat., 32: 127-31.

The European corn borer. Cornell Ext. Bull. 31, 35-48.

The European corn borer problem. Jour. Econ. Ent., 13: 59-74.

Memoirs Department of Agriculture in India. Yol. YII — No. 1, for
Imperial Department of Agr. in India.

Four new African gall midges. Annals Natl. Mus., Yol. IY (part 2),
p. 491-96.

New Indian gall midges. Mem. Dept. Agr. India. Ent. Ser. YII,
No. 1, p. 1-11.

New Philippine gall midges. Phil. Jour. Sci., 17 : 231-34.

Three new sub-tropical gall midges (Itonidae Dipt.). Ent. News, 22:
141-43.

Observations on J ohnsonomyia Felt with a description of a new spe-
cies. Can. Ent., 53 : 96.

New species of reared gall midges (Itonididse). Jour. N. Y. Ent.
Soc., 29: No. 2, 165.

A study of gall midges YII. N. Y. State Mus. Bull. (34th Rep’t),
81-241.

Sept., 1944]

Bromley: Felt

233

1 1921.

\ 1921.
/ 1921.
X 1921.

X 1921.

1921.

1921.

1922.
1922.

*1922.
\ 1922.

1924.

1925.

V 1925‘

1925.

1925.

* 1926.

* 1926.
^ 1926.

1927.
X 1927.

1928.

1929.

1929.
#1929.

1930.
1930.

1930.

1930.

1931.

Adaptations among insects of field and forest. Scientific Monthly,
13: No. 2, 165.

Lasioptera apocyni, new species (Diptera). Can. Ent., 53: 149.

NeAv Javanese gall midges. Extrait De Treubia, Yol. II, p. 89-92.
The number of antennal segments in gall midges and a new species.
Bull. Bklyn. Ent. Soc., 16: 93-95.

Indian grass gall midges. Mem. Dept. Agr. India, Pusa Ent. Ser.
VI, No. 3, p. 15-22.

Javanese gall midges, Treubia, 1: 139-51.

A new Javanese gall midge. Treubia, 1: 270-71.

A new Diadiplosis. Zoologica, Vol. Ill, No. 8.

The possibilities of exterminating insects. Scientific Monthly, 15:
No. 1, 35-41.

A new and remarkable fig midge. The Ent. (Summer Number), p.
5-6.

New cecidomyiid parasite of the whitefly. U. S. Natl. Mus. Proc.,
61: 1-2.

Manual of tree & shrub insects. The Macmillan Co., N. Y., 382 p.
School guide to insects and about insects. Univ. of State of N. Y.
Press.

Key to gall midges. (A Besume of Studies I— VII, Itonididae). N. Y.

State Mus. Bull. No. 257, 1-239.

Dispersal of butterflies and other insects. Nature, p. 1.

Insects and human welfare. Scientific Monthly, 21: 649-53.

A new spruce gall midge (Itonididae). Can. Ent., 58: 229.

New non-gall making Itonididae (Dipt.). Can. Ent., 58: 265-66.

New species of Indian gall midges (Itonididae). Memoirs of the Dept,
of Agr. in India, 9 : 241-45.

Four new Indian gall midges. Mem. Dept. Agr. India, 10 : 1-4.

New species of East Indian gall midges. Treubia, 9: 385-89.

A new African gall midge. Jour. N. Y. Ent. Soc., 36: (2), 123-24.
The need for investigations on shade tree insects. Jour. Econ. Ent.,
22: No. 2, 417.

Hickory leaf gall midges Caryomyia species. Jour. Econ. Ent., 22:
422.

Gall midges or gall gnats of the Orient (Itonidae or Cecidomyidae)
Lingnan Sc. Jour., Vol. 7, 413-74.

The Norway maple Nepticula (Lepidoptera) . Proc. Ent. Soc. Wash.,
32: 8, 146-9.

Economic importance of shade tree insects. Jour. Econ. Ent., 23:
1, 109-13.

(With S. W. Bromley) Shade tree problems. Proc. of the Ann.

Meeting of the Natl. Shade Tree Conf., p. 13.

Scientific names. Science, 21 : No. 1834, 215-17.

(With S. W. Bromley) Developing resistance or tolerance to insect
attack. Jour. Econ. Ent., 24: 437.

234

Journal New York Entomological Society

[Yol. lii

1931.

1931.

1931.

y

1931.

1932.

1932.

1932.

1932.

1932.

1932.

1932.

1932.

1932.

1933.

1933.

1934.

1934.

1934.

1934.

1934.

1935.

n 1935.

1935.

1935.

1935.

1935.

1935.

1935.

(^With S. W. Bromley) Insecticide investigations during 1930. Jour.
Econ. Ent., 24: 232.

The effect of certain important insects on trees. Natl. Shade Tree
Conf., 66-73.

(With S. W. Bromley) Observations on shade tree insects. Jour.
Econ. Ent., 24: 157-62.

(With S. W. Bromley) Tests with nicotine activators. Jour. Econ.
Ent., 24: 105.

(With W. Howard Rankin) Insects and diseases of ornamental trees
and shrubs. The Macmillan Co., N. Y., 507 p.

A new citrus cambium miner from Puerto Rico. Jour. Dept, of Agr.

of Puerto Rico, Yol. XYI, No. 2, 117-18.

A new predaceous gall midge for California. Pan-Pacific Ent., 8:
No. 4, 167-68.

A new Japanese gall midge. Bull. Bklyn. Ent. Soc., 27 : No. 2, 124.
Tree insurance. Proc. Ann. Meet. Natl. Shade Tree Conf., p. 15.
Shade tree insects. Proc. Ann. Meet. Natl. Shade Tree Conf., p. 77.
Shade trees threatened by insect pests. Scientific Monthly, 35 : 59-62.
(With S. W. Bromley) Insecticides on shade trees and ornamentals.
Jour. Econ. Ent., 25: 298.

(With S. W. Bromley) Observations on shade tree insects. Jour.
Econ. Ent., 25 : 39.

A hibiscus bud midge new to Hawaii. Proc. Haw. Ent. Soc., VIII,
No. 2, 247-48.

A new enemy of the pineapple mealybug and a list of gall midge
enemies of mealybugs. Jour. N. Y. Ent. Soc., 41: Mar .-June,
187-89.

New gall midges. Bull. Bklyn. Ent. Soc., 29: No. 2, 77-78.
Fundamentals in the control of insect pests. Proc. 10th Natl. Shade
Tree Conf., 37-43.

Classifying symbols for insects. Jour. N. Y. Ent. Soc., 42: 373-92.

A new gall midge on fig. (Dipt.: Itonididse). Ent. News, 45: 131-33.
Shade tree insects in 1933. Jour. Econ. Ent., 27 : 195.

Bark beetles and the Dutch elm disease. Jour. Econ. Ent., 28: No. 1,
231.

(With K. F. Chamberlain) Occurrence of insects at some height in
air, especially on the roofs of high buildings. N. Y. State Mus.
Circ. 17, 70 p.

Trisopsis in the United States (Dipt.: Itonididae or Cecidomyiidae),
Ent. News, 46: 75.

A new melon gall midge. Bull. Bklyn. Ent. Soc., 30: No. 2, 79-80.

A new gall midge. Jour. N. Y. Ent. Soc., 43: 47.

A gall midge on pine cones. Jour. N. Y. Ent. Soc., 43: 48.

Bark beetles and the Dutch elm disease. Jour. Econ. Ent., 28: 231.
The importance of shade tree insects in 1934. Jour. Econ. Ent., 28:
390.

Sept., 1944]

Bromley: Felt

235

1936.

1936.

1936.

1936.

1936.

1936.

1937.

1937.

1938.

1938.

1938.

1938.

1938.

1938.

1939.

1939.

1939.

1940.

X 1940.
1940.
1940.

1940.

1940.

1940.

1940.
’*1941.

1941.
)C 1941.

1941.

New midges on pine and grass. Jour. N. Y. Ent. Soc., 44: 7.

(With S. W. Bromley) Oil effects on shade trees. Jour. Econ. Ent.,
29: (2), 357-60.

Two new cockle burr midges (Dipt.: Cecidomyiidae) . Ent. News, 47:
231.

(With S. W. Bromley) Shade tree insect developments. Jour. Econ.
Ent. 29: 490.

The biology of mayflies, with a systematic account of North American
species. Jour. Econ. Ent., 29: 223.

Forest insects. Jour. Econ. Ent., 29: 1031.

(With S. W. Bromley) Observations on shade tree insects and their
control. Jour. Econ. Ent., 30: (1), 71-75.

Dissemination of insects by air currents. Jour. Econ. Ent., 30: 458.
A note on Lasioptera murtfeldtiana Felt. Jour. N. Y. Ent. Soc.,
46: 44.

(With S. W. Bromley) Shade tree insects and sprays, 1937. Jour.
Econ. Ent., 31: No. 2, 173-6.

Why do insects become pests? Scientific Monthly, 46: 437-40.

A new species of gall midge predaceous on mealybugs. Proc. Haw.
Ent. Soc., No. 1.

Wind drift & dissemination of insects. Excerpt, Can. Ent., 221.

Our shade trees. Orange Judd Publishing Company, N. Y., 187 p.
(With S. W. Bromley) Hurricane of Sept. 21, 1938. Scientific Tree
Topics No. 1, 1939, Yol. 1.

A new juniper midge (Diptera: Cecidomyiidae). Ent. News, 50:
159-60.

A new gall midge on rhododendron. Jour. N. Y. Ent. Soc., 47 : 41-42.
(With S. W. Bromley) New insecticides and spreaders on shade trees.
Jour. Econ. Ent., 33: 244-49.

Rhododendron midge. Scientific Tree Topics 4, Yol. 1, p. 30.
Insecticides and their use. Scientific Tree Topics 4, Yol. 1, p. 26.
European elm bark beetle and Dutch elm disease control. Jour. Econ.
Ent., 33: (3), 556.

Danger to trees in an oversupply of quickly available nitrogen. Scien-
tific Tree Topics 2, Yol. 1, p. 10.

Pine root weevil. Scientific Tree Topics 2, Yol. 1, p. 13.

Factors affecting Dutch elm disease spread. Scientific Tree Topics
3, Vol. 1. p. 18.

Training trees for beauty. Scientific Tree Topics 5, Yol. 1, p. 34.
(With S. W. Bromley) New and unusual shade tree pests. Jour.
Econ. Ent., 34 : (3), 383.

With S. W. Bromley) Major shade tree insects, 1940. Jour. Econ.
Ent., 34: (2), 180.

The minute gall insect of white oaks. Scientific Tree Topics 7, Yol.
1, p. 55.

Weather and winter drying. Scientific Tree Topics 7, Yol. 1, p. 50.

236

Journal New York Entomological Society

[VOL. LII

1941. Pruning trees and shrubs. Orange Judd Pub. Co., Inc., N. Y., 237 p.

1942. (With S. W. Bromley) The increasing importance of Coleopterous

borers in shade trees. Jour. Econ. Ent., 35: (2), 169.

1942. Our shade trees. Eevised edition. Orange Judd Pub. Co., Inc., N. Y.,

316 p.

1943. How may the arboricultural profession serve a nation at war? 18th

Natl. Shade Tree Conf., p. 29.

1943. Dutch elm disease control. Scientific Tree Topics 8, Yol. 1, p. 58.
1943. Shelter trees in war and peace. Orange Judd Pub. Co., N. Y., 320 p.

Stanley W. Bromley.

Sept., 1944]

Brown: Butterflies

237

NOTES ON MEXICAN BUTTERFLIES, III, DANAIDiE

By F. Martin Brown

Danainae

124. Anosia berenice Cramer.

G. & S. (1), 1: 3; 2: 638.

Ha. (2), p. 114, pi. 31a.

Ho. (5), p. 662.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 4 ££ 1 2 vi. 16-18. 40 (H.H.).

Galeana, Nuevo Leon, 6500 ft., 2 J'J' 2 52 vii.29-viii.-
1.39 (H.H.).

nr. Villagran, Tamanlipas, 1160 ft., 1J iv.28.41 (R.P.).

Hda. Sta. Engracia, Tamanlipas, 1 2 vii.27.39 (H.H.).

60 mi. So. of Victoria, Tamaulipas, 1 £ vii.5.36
(H.D.T.).

Jacala, Hidalgo, 4500 ft., 4 J'J' 6 52 vi.25-vii.3.39
(H.H.).

El Sol, Tamazunchale, Sau Luis Potosi, 400 ft., 2 25
v.28-29.41 (R.P.).

El Pujal, San Luis Potosi, 100 ft., 1 £ vii.20.39 (H.H.).

nr. Chilpancingo, Guerrero, 1 2 vi.26.41 (R.P.).

El Sabino, nr. Uruapan, Michoacan, 5 J'.J' 1 2 vii.16-
30.36 (H.D.T.).

Chichen Itza, Yucatan, 1 J1 viii.30.36 (H.D.T.).

All this material is probably referable to race strigosa Bates.
A few of the specimens (Villagran, Galeana, Jacala) have the
greyish scales along the nervules of the upper side of the hind
wings so reduced in number as to be difficult to see. There is
some difference in ground color which I am inclined to believe is
related to the freshness of the specimens. Those that are freshly
emerged are brighter fulvous than those that have flown for some
time which are rather dull dilute reddish brown. There is con-
siderable variation in the development of the white spots in the
dark margin of the hind wings. In two cases (a Jacala 2 and an
El Sol 2) these spots are absent, in all of the others the outer row

238

Journal New York Entomological Society

[Vol. LII

is at least indicated and in some the second row is almost com-
plete. The baso-apical radius of the fore wing varies from 31 to
47 mm. in the males and from 29 to 44 mm. among the females.
There may be a tendency for the more northern specimens to be
brighter and larger, and more clearly strigosa.

125. Anosia cleothera Godart.

G. & S., 1 : 3 ; 2 : 638.

Ha., p. 114, pi. 31b.

Ho., p. 663.

Galeana, Nuevo Leon, 6500 ft., 1 J1 viii.1.39 (H.H.).

60 mi. So. of Victoria, Tamaulipas, 1 £ vii.6.36
(H.D.T.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 J
v. 29.41 (R.P.).

Orizaba, Vera Cruz, 2000 ft., 1 J v.6.41 (R.P.).

Rio Blanco, Vera Cruz, 2200 ft., 1 J' v.10.41 (R.P.).
The Galeana record is rather far north for this species; how-
ever, it has been taken in Texas. In McDunnough’s check list it
is termed eresimus Cramer. I am not sure that cleothera is really
valid for the Central American material, it was described from
‘ ‘ Timor ” ( ! ) and the original description does not apply too well
to our material. Cramer’s species from the Guianas fits it no
better so until a specialist settles the question it is best to let
cleothera stand.

126. Danaus plexippus Linnaeus.

G. & S., 1: 1; 2: 638.

Ha., p. 113 (as archippus F.), Vol. 1, pi. 28c.

Ho., p. 663 (as curassavicce F.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft.. 1 J
v.29.41 (R.P.).

Tancitaro, Michoacan, 6600 ft., 1 2 §5 vii.30-viii.ll.40

(H.H.).

The El Sol specimen and one of the Tancitaro females are f.
fumosa Hulst. Hoffmann lists this species as Diogas curassavicce
Fabricius. Apparently he overlooked Riley’s paper (Tr. Ent.
Soc. Lond., 76: 451. 1929) which definitely linked the Linnean
name with our “Monarch.”

Sept., 1944]

Brown: Butterflies

239

Ithomiinae

My specimens of this subfamily were determined by R. M. Fox,
of the Reading Public Museum. These represent six species of
which Fox says, “All are common members of the Mexican tropi-
cal fauna.’7 All of them were collected by Robert Potts. Of
these six species five are among the twenty-eight Ithomiinae in
Hoffmann’s list. The other has probably been confused by Hoff-
mann with Ithomia patilla Hewitson. I am following Hoffmann’s
order in this listing.

139. Dircenna klugii Hiibner.

G. & S., 1 : 28 ; 2 : 644.

Ha., p. 138, pi. 36e.

Ho., p. 664.

Tuxpango, Yera Cruz, 1500 ft., 1 £ v.8.41.

Fortin, Yera Cruz, 1600 ft., 1 2 v.3-4.41.

Orizaba, Yera Cruz, 2000 ft., 1 2 v.6.41.

141A. Ithomia hippocrenis Bates.

G. & S., 1 : 51, pi. 5, f. 5.

Ha., p. 142.

Ho., p. 665 (I. patilla in part).

Ojo de Agua, Yera Cruz, 1600 ft., 3 3 22 v.12.41.

A pair of these are deposited at the Reading Public Museum.
This species is not listed by Hoffmann.

145. Oleria paula Weymer.

G. & S., 2 : 645 ( L . victoria 1 : 38 in part) .

Ha., p. 150.

Ho., p. 665.

Ojo de Agua, Yera Cruz, 1600 ft., 1 , 1 2 v.12.41.
Hoffmann et al. use the genus Leucothryis for this species.

149. Pteronymia cotytto Guerin.

G. & S., 1: 44; 2: 647.

Ha., p. 155, pi. 40c.

Ho., p. 665.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 3 22
v.28-29.41.

240

Journal New York Entomological Society

[VOL. LII

Tuxpango, Vera Cruz, 1500 ft., 1 2 v.8.41.

Ojo de Agua, Vera Cruz, 1600 ft., 1 J' 2 22 y.12.41.

A female from El Sol and one from Ojo de Agua are deposited
at the Reading Public Museum.

153. Greta oto, Hewitson.

G. & S., 1 : 56 ; 2 : 648.

Ha., p. 164, pi. 41e.

Ho., p. 666.

Ojo de Agua, Vera Cruz, 1600 ft., 2 jy 3 22 v.12.41.
Hoffmann et al. place this and the following species in Hy-
menitis. A pair of oto is deposited at the Reading Public Museum.

154. Greta nero Hewitson.

G. & S., 1: 57; 2: 648.

Ha., p. 163, pi. 41e.

Ho., p. 666.

Fortin, Vera Cruz, 1600 ft., 1 y v.3.41.

SATYRIDiE

As is so frequent with collections made by inexperienced col-
lectors in the tropics the catch of Satyridse brought back by both
Hoogstraal and Potts is small in comparison with what it might
have been. The bulk of the specimens taken are Euptychia.
Most of the jungle-loving species were not captured.

159. Tisiphone maculata Hopffer.

G. & S., 1 : 72, 651.

W. (3), p. 184, pi. 44b.

Ho., p. 666.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 2 jy
1 2 v.28-29.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 2 22 vi.26-
28.40 (H.H.).

The El Sol female is fresh ; all of the other specimens are worn.

163. Taygetis virgilia Cramer.

G. & S., 1 : 97, 659.

W., p. 187, pi. 45a.

Sept., 1944]

Brown: Butterflies

241

Ho., p. 666.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 2 jy
v.28-29.41 (R.P.).

167. Taygetis inconspicua Draudt.

Ho., p. 667.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 7 jy
iv.30, vi. 28-29. 41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 1 y vi. 28.40

(H.H.).

El Pujal, San Luis Potosi, 100 ft., 2 yy vi. 17-18. 39
(H.II.).

The El Banito male and several of the May specimens from
El Sol are worn ; the others are reasonably fresh. I suspect that
this is nothing more than a form of virgilia. The genus will be
treated in monographic form as a member of my series of papers
devoted to Ecuadorian butterflies.

172. Neonympha gemma Hiibner.

G. & S., 1 : 92, 657, pi. 8, f. 12.

W., p. 223 (as Cornelius F.), pi. 49f.

Ho, p. 667.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft, 2 jy 1 ? vi. 18-20.40 (H.H.).

60 mi. So. of Victoria, Tamaulipas, 2 yy vii.6.36
(H.D.T.).

Jacala, Hidalgo, 4500 ft, 3 jy vi.24-29.39 (H.H.).
Fortin, Vera Cruz, 1600 ft, 1 y v.4.41 (R.P.).

179. Megisto hesione Sulzer.

G. & S, 1 : 75, 651 (as ocirrhoe).

W, p. 194, pi. 46d.

Ho, p. 667.

El Sol, Tamazunchale, San Luis Potosi, 400 ft, 7 yy
iv. 30-31, v.28-29.41 (R.P.).

Ojo cle Agua, Vera Cruz, 1600 ft, 2 55 v.12.41 (R.P.).

181. Megisto mollina Ilewitson.

G. & S, 1: 76, 652, pi. 8, ff. 13, 14 (fig. as westwoodi).

242

Journal New York Entomological Society

[Vol. LII

W., p. 195, pi. 46e.

Ho., p. 668.

El Sol, Tamazunchale, San Lnis Potosi, 400 ft., 5 J'J'
v.28-29.41 (R.P.).

Ojo de Agua, Vera Cruz, 1600 ft., 3 J'.J' 3 55 v.12.41
(R.P.).

The females and one male from Ojo de Agua are f. westwoodi
Butler.

182. Megisto fetna Butler.

G. & S., 1 : 77, 652, pi. 8, ff. 15, 16.

W., p. 195.

Ho., p. 668.

Chilpancingo, Guerrero, 1 vi. 26.41 (R.P.).

184. Megisto terrestris Butler.

G. & S., 1 : 79.

W., p. 200.

Ho., p. 668.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 1 S vi.19.40 (H.H.).

This specimen is probably not true terrestris which hails from
eastern South America but may be the same form regarded as
terrestris by Hoffmann. It is far north of Hoffmann’s record
“Selva virgen de Campeche (Tierra caliente).” It compares
well with Weymer’s figure noted above.

188. Megisto rubricata Edwards.

G. & S., 1 : 82, 655, pi. 8, f . 5.

W, p. 202.

Ho., p. 668.

Ojo de Agua, Sabinas Hidalgo, Nuevo Leon, 1000 ft.,
1<? vi.14.40 (H.H.).

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1600
ft., 1 vi.19.40 (H.H.).

Jacala, Hidalgo, 4500-5100 ft., 3 J'J' 2 5? vi.24.39

(H.H.). ^

Sept., 1944]

Brown: Butterflies

243

190. Megisto renata disaffecta Butler.

G. & S., 1 : 82, 655, pi. 8, f. 5 (as renata).

W, p. 204, pi. 47d.

Ho., p. 668.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 y 1 2
iv.30-31.41 (R.P.) .

Fortin, Vera Cruz, 1600 ft., 1 y v.3.41 (R.P.).

Tuxpango, Vera Cruz, 1500 ft., 1 y v.9.41 (R.P.).

Ojo de Agua, Vera Cruz, 1600 ft., 3 yy v. 12.41 (R.P.).

El Sabino, nr. Uruapan, Michoacan, 1 ^ 1 J vii.15-
30.36 (H.D.T.).

Chichen Itza, Yucatan, 2 yy viii. 30.36 (H.D.T.).

Determination tentative.

191. Megisto hermes Fabricius.

G. & S., 1 : 86, 656, pi. 8, ff. 6, 7 (as camerta).

W., p. 207, pi. 48a.

Ho., p. 668.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon,
1600 ft., 3 yy vi. 17-18.40 (H.H.).

Monterrey, Nuevo Leon, 1800 ft., 2 yy 1 2 iv.27.41
(R.P.).

nr. Villagran, Tamaulipas, 1160 ft., 1 y iv.28.41 (R.P.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 2 yy
2 52 v.28-29.41 (R.P.) .

El Banito, Valles, San Luis Potosi, 200 ft., 3 yy
vi. 27.41 (H.H.) ; 1 y iv.30.41 (R.P.).

El Pujal, San Luis Potosi, 100 ft., 2 yy 5 22 vii.17-
21.39 (H.H.).

Hda. Potrero Viejo, Vera Cruz, 1500 ft., 1 2 v.5.41
(R.P.).

Tuxpango, Vera Cruz, 1500 ft., 1 2 v.9.41 (R.P.).

Fortin, Vera Cruz, 1600 ft., 2 52 v.4.41 (R.P.).

Ojo de Agua, Vera Cruz, 2000 ft., 2 yy 1 2 v. 12.41
(R.P.).

Orizaba, Vera Cruz, 2000 ft., 4 yy 2 22 v.6.41 (R.P.).

60 mi. So. of Victoria, Tamaulipas, 1 y 1 2 vii.6.36
(H.D.T.).

244

Journal New York Entomological Society

[Yol. lii

El Sabino, nr. Uruapan, Michoacan, 5 4 52 yii.15-

30.36 (H.D.T.).

Chichen Itza, Yucatan, 2 55 viii.30.36 (H.D.T.).

An extremely varied series. The species “kermes” may be
composite.

197. Megisto libye Linnaeus.

G. & S., 1 : 83, 655.

W., p. 212, pi. 48e.

Ho., p. 669.

Ojo de Agua, Vera Cruz, 1600 ft., 1 5 v.12.41 (R.P.).

198. Megisto glaucina Bates.

G. & 1 : 90, pi. 8, if. 18, 19.

W, p. 216, pi. 49b.

Ho., p. 669.

Ojo de Agua, Vera Cruz, 1600 ft., 1 (J 1 5 v.12.41
(R.P.).

Megisto sp.

Fortin, Vera Cruz, 1600 ft., 1 v.4.41 (R.P.).

Ojo de Agua, Vera Cruz, 2000 ft., 1 v.6.41 (R.P.).

A species near nebulosa Butler. Differing from that South
American species in having five ocelli on the under side of the
fore wing.

200. Pindis squamistriga Felder.

G. & S., 1 : 80, 654, pi. 8, f. 24 (fig. as zabdi).

W., p. 224, pi. 50a.

Ho., p. 669.

Jacala, Hidalgo, 4500 ft., 2 vi.23-29.39 (H.H.).

El Sabino, nr. Uruapan, Michoacan, 1 (5^^.15-30.36

(H.D.T.).

Sta. Lucretia, 2 <5<5 ix.4.39 (H.D.T.).

The two Jacala specimens are badly battered; fortunately one
entire fore wing is intact on one specimen, sufficient to recognize
the genus and species. This is the first record of the species from
so far northeast in Mexico.

Sept., 1944]

Brown: Butterflies

245

202. Paramecera xicaque Reakirt.

G. & S., 1 : 101, 660.

W., p. 225, pi. 50a.

Ho., p. 669.

Tancitaro, Michoacan, 6600 ft., 1 J1 vii.25.40 (H.H.).

A battered specimen.

BRASSOLIDiE

216. Opsiphanes cassina fabricii Boisduval.

G. & S., 1 : 127, 664 (as cassice).

F. (4), p. 302, pi. 51d.

Ho., p. 671.

El Banito, Valles, San Luis Potosi, 200 ft., 3 1 5

vi.28.40 (H.H.).

Hoffmann reports this species from both coasts as far north as
Sinaloa and Tamanlipas in contrast to Fruhstorfer’s report of it
only as far north as Tepic, Nayarit on the west coast and no men-
tion of it in eastern Mexico.

218. Eryphanis aesacus aesacus Herrick-Schaffer.

G. & S„ 1 : 137, 666.

F. , p. 313, pi. 64b.

Ho., p. 671.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 2 J'J'
1 2 v.28-29.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 1 $ v.28.40
(H.H.).

El Pujal, San Luis Potosi, 100 ft., 2 vi.18.39

(H.H.).

These records extend the range of the species a little further
north than that given by Hoffmann. Fruhstorfer says of the
species, 1 1 Everywhere rare. ’ ’

MORPHID.ZE

225. Morpho peleides montezuma Guenee

G. & S., 1 : 119, 663 (as peleides).

F., p. 344.

Ho., p. 672.

246

Journal New York Entomological Society

[Vol. LII

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 5 J'J'
y.29.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 2 J'J'
vii.22.39, vi. 26.40 (H.H.).

Ojo de Agua, Vera Cruz, 1600 ft., 3 J'J' 1 § v.12.41
(R.P.).

The Ojo de Agua material is in very poor condition, especially
the female. The other specimens are in good condition. The
records for San Luis Potosi extend the range of the species a
little to the north.

BIBLIOGRAPHY

1. Godman & Salvin, Biologia Centrali- Americana, Rhopalocera, 1 : 1-139.

1879-1881.

2. Haensch, in Seitz’ Macrolepidoptera of the World, vol. 5, p. 113-171.

1909- 1910.

3. Weymer, in Seitz’ Macrolepidoptera of the World, vol. 5, p. 173-283.

1910- 1912.

4. Fruhstorfer, in Seitz’ Macrolepidoptera of the World, vol. 5, p. 285-356.

1912-1913.

5. Hoffmann, Anales del Instituto de Biologia (Mexico), 11: 639-739.

1940.

Sept., 1944]

Ginsburg: Mosquitoes

247

OUTDOOR PROTECTION FROM MOSQUITOES*

Joseph M. Ginsburg

Biochemist in Entomology, New Jersey Agricultural
Experiment Station

Even in areas where mosquito control work has been systemati-
cally conducted for many years it has not been possible, with our
present means and methods, completely to eradicate the mosquito.
Under conditions favorable for its rapid development, such as
during summers of abundant rainfall, it becomes at times a seri-
ous interference with human comfort.

When a female mosquito bites or pierces the skin to suck our
blood she injects into the wound a small amount of a poisonous
substance, the chemical composition of which has not been defi-
nitely established. The effect of this injected material varies
with different species of mosquitoes as well as with the suscepti-
bility of the bitten individual. While a small number of people
appear to possess a certain degree of immunity, either natural or
acquired, against this poison, the great majority of us, especially
children, suffer from its effects. Immediately following the bite,
an itching sensation is felt which may be followed by considerable
swelling. The scratching induced by the irritation may cause a
secondary blood infection, especially among children, leading to
serious results. The irritation may be relieved by washing the
swelling with any one of the following solutions : soft soap, alco-
hol, glycerin, iodine or borax.

Complete elimination of mosquitoes indoors can be readily
accomplished by proper screening, spraying, or fumigating. On
the other hand, protection from mosquito annoyance outdoors
constitutes a difficult problem, the solution of which cannot
always be successfully attained.

Mosquito repellents. — -Various chemicals, possessing repelling
properties against mosquitoes, have been recommended and com-
mercialized. They are applied in form of lotions, ointments or
powders to the exposed parts of the body. The duration of the

* Journal Series paper of the New Jersey Agricultural Experiment Station,
Rutgers University, Department of Entomology.

248

Journal New York Entomological Society

[Vol. LII

protection may vary from a short time to several hours, depend-
ing on the inherent repellent properties of the chemical, thorough-
ness of application, species and density of the mosquito popula-
tion, degree of attractiveness possessed by the individual, and
atmospheric factors.

Citronella is one of the oldest mosquito repellents used either
in its natural liquid state or in combination with various other
compounds. One of the early formulas recommended by Dr.
L. 0. Howard (1923) contains the following ingredients: oil of
citronella 2 ounces, spirits of camphor 2 ounces, and oil of cedar
1 ounce. This preparation can also be made in form of a salve,
if desired, by mixing in enough petrolatum or similar semisolids.
One thorough application may remain effective for 1 or 2 hours.
Dr. Herms (1939) finds the following formula successful: citron-
ella 3 parts, kerosene 2 parts, and coconut oil 4 parts. To this
mixture is added 1 per cent carbolic acid. Within the last few
years longer lasting and more efficient repellents have been syn-
thesized, as a result of research conducted at the Federal Bureau
of Entomology in Washington and at Rutgers University (Gran-
ett 1940), in cooperation with various commercial concerns.
Three of these are at present widely used, namely, dimethyl
phthalate, Indalone, and formula No. 612.

Though mosquito repellents are contributing a great deal
toward relief from mosquito annoyance to individuals, such as
night watchmen, military pickets, mosquito workers and fisher-
men, compelled to remain exposed for considerable lengths of
time in mosquito-infested areas, their frequent application en-
counters many drawbacks. First, not all the repellent mixtures
thus far developed repel mosquitoes from a distance. In virtu-
ally every case the mosquito has to alight, or at least come very
close to the treated surface, before the chemical acts on it. Com-
plete and thorough coverage is, therefore, necessary, otherwise
untreated spots may be bitten. Second, virtually all of the repel-
lents cause sharp smarting on delicate parts of the skin such as
eyelids, face and forehead. This irritation is apt to become
rather pungent and often unbearable to many sensitive indi-
viduals under warm, humid atmospheres and other conditions
when copious perspiration occurs. Third, because of the solvent

Sept., 1944]

Ginsburg: Mosquitoes

249

properties of the repellents, a treated individual must avoid
coming in contact with varnished and painted surfaces. Either
paint or varnish will be partially removed, resulting in staining
of clothes or any other objects incidentally touched. Fourth, the
repellent does not protect a group, much less a large outdoor
audience, unless each individual is treated. Fifth, continuous
feel of a greasy, somewhat irritating solvent all over one’s skin
becomes rather unpleasant. Unless compelled by military duty
or night work, few individuals will willingly submit themselves
to this treatment night after night. Instead they would rather
stay indoors or in a well-screened porch.

Mosquito sprays. — In view of these objectionable features, the
writer became interested in developing a mosquito spray which
should, with no injury to man, animals, and plants, completely
free an area from adult mosquitoes where groups of people could
spend a summer evening with no mosquito annoyance and with
perfect body comfort. This effort coincided with another prob-
lem, that of developing a mosquito larvicide which should kill
mosquito larvae and pupae with no injury to fish, water fowl, and
aquatic plants, offering at the same time no fire hazard. Such
a product was called for in residential sections, ornamental ponds,
game conservation parks, and similar places where mosquito oil
was objectionable. By 1931 the New Jersey Py rethrum Mosquito
Larvicide was developed and came into wide use as a substitute
for oil wherever the latter meets with objections, Ginsburg
(1930). The larvicide is essentially an emulsion consisting of
66 per cent kerosene, 0.5 per cent sodium lauryl sulfate as emulsi-
fier, 0.07 per cent pyrethrins, and about 34 per cent water. The
concentrated stock emulsion is mixed just before spraying with
about 10 parts of clear water, which may be taken directly from
• the area to be sprayed.

Preliminary tests conducted during 1934 (Ginsburg 1935) indi-
cated that spraying with this diluted larvicide might also prove
effective in ridding a given area from adult mosquitoes. Further
extensive experiments carried out during 1935-36 (Ginsburg
1936, 1937) by the writer in cooperation with the various County
Mosquito Commissions in New Jersey have substantiated this
assumption and have established a definite method of procedure
for spraying.

250

Journal New York Entomological Society

[Yol. lii

The experiments during these three years have given us some
fundamental knowledge as well as authentic information concern-
ing the possibility and practicability of eliminating mosquito
annoyance from outdoor public gatherings without in any way
interfering with human comfort and activities. We have learned
how to accomplish this relief without appreciable injury to plant
life. These results have also thrown some light on the physical
and chemical mechanism involved as well as the role that each
ingredient of the larvicide plays in this process. From the experi-
ence thus far gained the following principles may be evolved as
guides in this work.

In order successfully to eliminate adult mosquitoes from a
given area outdoors, two objectives must be attained. First, all
female mosquitoes resting in the grass, shrubs, and throughout
the area to be treated must be killed. This is accomplished by
thoroughly spraying the entire grounds before the audience
gathers. Second, the influx of mosquitoes from the surroundings
into the protected area must be prevented. For this purpose the
spray is directed upward, as high as the pressure permits, so as
to saturate the atmosphere with a fine mist of larvicide. This
air-fogging may have to be repeated two or three times during the
affair, depending on the species and density of mosquitoes, the
flight intensity, the direction and velocity of wind, and other
atmospheric factors. It should be emphasized that, for complete
protection, it is just as important to kill or incapacitate all the
female mosquitoes hiding in vegetation as it is to bar those in the
surroundings from flying in. The spray should be applied in
highly dispersed form, as a fog or mist. The finer the liquid
particles, the less will be the danger of injury to plants and the
longer will it remain floating in the air. The diluted larvicide
must be kept thoroughly mixed during spraying.

The mechanism involved in this process is twofold, repellency
and contact killing. The female mosquito when in contact with
the larvicide is either killed or paralyzed to such a degree that,
for an indefinite period, she can neither fly nor bite. These symp-
toms are brought about primarily by the pyrethrins incorporated
in the larvicide. The petroleum oil acts as a carrier for the
pyrethrum. It should be mentioned here that neither an aqueous

Sept., 1944]

Ginsburg: Mosquitoes

251

spray of pyrethrum extract nor a kerosene emulsion when applied
separately gave satisfactory protection. Evidently the combi-
nation of both ingredients is necessary for adequate protection.
The water in the larvicide merely serves as the outer phase in
which the two toxic ingredients are dispersed by the aid of the
emulsifier.

The question has been asked : What happens to the mosquitoes
in the sprayed area? The following observations may offer an
explanation. When tents were sprayed inside with the larvicide,
some mosquitoes were found on the ground apparently dead while
others remained on the canvas in a quiescent condition. When
disturbed they attempted to fly but soon again attached them-
selves to the canvas. They remained for some time in this partly
paralyzed condition and finally either dropped to the ground or
flew away. These symptoms closely resemble those observed in
other insects treated with pyrethrum insecticides and can, there-
fore, be attributed directly to the pyrethrins incorporated in the
larvicide. Again, while checking the mosquito density the writer
observed on several occasions that when mosquitoes reach the
treated area they abruptly change their course of flight, turning
back or at a right angle. This behavior suggests repellency.
Evidently mosquitoes find the oil, the pyrethrum, or both com-
bined so disagreeable that they keep away from the treated area.

DIRECTIONS FOR SPRAYING

The concentrated larvicide is first well shaken or stirred. It is
then mixed with 12 parts of water (1 quart to 3 gallons) in the
sprayer. This diluted mixture is ready for spraying. During
the spraying operation it should be frequently shaken or stirred
in order to insure uniform distribution of the larvicide.

Procedure. — Before the affair starts, the entire area, including
grass, shrubs, bleachers, sheds, benches, or any other place where
mosquitoes may rest during the day, is thoroughly sprayed with
the diluted larvicide. This should kill all mosquitoes as well as
many other insects hiding in the grass, shrubs, and other places.
The next object is to prevent mosquitoes from the surroundings
from coming into the protected area. For this purpose the spray
is directed upward so as to saturate the atmosphere with a fine
mist or fog of the larvicide. If the spray is applied against the

252

Journal New York Entomological Society

[Vol. LII

wind the fine mist or fog will drift with the wind throughout the
area. This entire operation should be finished before the gather-
ing takes place. Just about dusk or when the mosquitoes from
the outside begin to fly in, another fogging is necessary. For this
purpose the spray is applied as high as the sprayer permits and
primarily on the side from which the wind is blowing. This mist
in the air wards off mosquitoes from outside the treated area.
If no noticeable wind prevails it may be necessary to fog all
around the area, directing the spray upward so as to keep the
inflying mosquitoes away. A thorough fogging about the time
when the mosquitoes start coming in should be sufficient for the
rest of the evening. Under very heavy infestations, where the
mosquitoes are coming in large numbers, a second fogging about
9 : 30 may be necessary.

Apparatus Required. — (1) on small areas, such as backyards,
porches, and private lawns, a garden sprayer, knapsack sprayer,
bucket pump, or electric sprayer capable of producing a fine
spray, about 10 or 15 feet high, may be used with success.

( 2 ) For spraying large areas, such as picnic grounds, stadiums,
and open air theaters, a power sprayer capable of developing a
pressure of 200 pounds or more per square inch and equipped
with a spray gun is necessary. The larvicide has been found
most efficient in protecting outdoor audiences on comparatively
large areas where power sprayers can be used.

Precautions. — Before attempting to treat an area, one should
make a thorough survey in order to ascertain the following
points: sources of mosquitoes; possible mosquito resting places;
direction of wind; type of vegetation present; water supply for
mixing the larvicide ; kind of spraying outfit necessary ; nature of
gatherings; and any difficulties that may be anticipated during
the spraying operation. This information should enable the
operator to plan his method of procedure.

PRESENT STATUS OF THE NEW JERSEY PYRETHRUM
MOSQUITO LARVICIDE

Since this spray was introduced, numerous outdoor evening
concerts, carnivals, church parties, community gatherings, and
lawn parties have been fully protected from mosquitoes either
directly by the County Mosquito Commissions at a nominal cost,

Sept., 1944]

Ginsburg: Mosquitoes

253

or indirectly through their assistance, guidance, and advice. The
number of persons present on these occasions varied from small
groups to many thousands. Records show that in several in-
stances some 20,000# persons in one gathering have enjoyed an
open air evening concert with no mosquito discomfort as a result
of spraying the stadium where the affair was held. When the
larvicide was sprayed as directed, no injury has resulted to grass,
shrubs, trees, ornamental plants or aquatic plants. Up to and
including 1942, its use increased with each successive season.

Prior to 1934 most of the pyrethrum used in this country came
from Japan, and small amounts from Dalmatia and the Kenya
Colony in British East Africa. By 1939 the picture was re-
versed; most of our pyrethrum was imported from Kenya and
very little from Japan. When World War II broke out, cultiva-
tion of pyrethrum in the British possessions, for various reasons,
decreased. Shipping shortage and submarine losses still further
curtailed the flow of pyrethrum flowers to this country. After
Pearl Harbor, practically all the available and potential supplies
of pyrethrum were taken over by the United States Government
for army use. At present the War Production Board does not
allow the use of pyrethrum for preparing the larvicide. Thus,
the excellent record of this newly developed method for tempo-
rary relief from biting female mosquitoes has been suddenly
terminated for the duration or until enough pyrethrum becomes
available to supply a surplus, above the amount necessary for our
armed forces.

Intensive testing is now being conducted with various other
chemicals as possible substitutes for pyrethrum. At this date,
however, none has proved equal to pyrethrum from the stand-
points of effectiveness against the mosquito and of safety to man,
animals, and plants.

LITERATURE CITED

Ginsburg, J. M. 1930. Studies of pyrethrum as a mosquito larvicide. Proc.
17th Ann. Meet. N. J. Mosq. Ext. Assoc., p. 57.

. 1935. Larvicides and method for temporary protection from adult

mosquitoes in limited areas. Proc. 22nd Ann. Meet. N. J. Mosq.
Ext. Assoc., p. 147.

* Essex County Symphony Society, at the Newark School Stadium.

254

Journal New York Entomological Society

[Yol. lii

. 1936. Protection of outdoor gatherings from the mosquito pest.

Proc. 23rd Ann. Meet. N. J. Mosq. Ext. Assoc., p. 166.

. 1937. Principles underlying the protection of outdoor meetings

from the mosquito pest and methods of application. Proc. 24th
Ann. Meet. N. J. Mosq. Ext. Assoc., p. 5.

Granett, P. 1940. Studies of mosquito repellents. Jour. Econ. Ent.,
33(3) : 563-71.

Herms, W. B. 1939. Medical Entomology. MacMillan, New York.
Howard, L. O. 1923. Remedies and preventives against mosquitoes.
U. S. D. A. Farmers’ Bull. No. 444.

Vannote, R. L. 1937. Methods of applying the larvicide as a repellant.
Proc. 24th Ann. Meet. N. J. Mosq. Ext. Assoc., p. 11.

Sept., 1944]

Clench: Lyoenhle

255

NEW NEOTROPICAL THECLINZE (LEPIDOPTERA,
LYC^NID^E)

By Harry K. Clench
Cambridge, Mass.

The following new Theclinge have come to my attention recently,
two of them during the arranging of the Neotropical Lycsenidse
in the collection of the Museum of Comparative Zoology (M.C.Z.)
and one in a loan of study material from the American Museum
of Natural History.

X

Thecla caramba, new species

Eyes ringed with pale, green. Frons metallic green. Collar above obscure,
laterally brown. Palpi rusty gray, terminal sediment dark. Antennce
black above, white annulate below. Thorax above black-brown (scaled with
metallic blue?), covered anteriorly, laterally and posteriorly with long, back-
directed hairs, bluish in color; below with moderately long, red-brown hair.
Abdomen above metallic blue, becoming gray towards the anal extremity;
below pale yellow, or rich cream. Legs largely black, ringed narrowly and
infrequently by white.

Upperside :

Male. Both wings dark, lustrous blue, with a purple luster. Fore wing
with a narrow outer marginal border, thickening slightly at the apex. Find
wing with costa gray-brown, and inner margin gray. Outer margin very
narrowly black. Anal lobe rusty. Outer margin very slightly produced at the
vein-ends from Mx to the anal angle, but no indication of any tails is present.
Fringe of both wings brown, on hind Aving paler outwardly between the veins.

Underside:

Male. Both wings bright pea-green. Fore wing Avith a gray area from Cu2
to the inner margin. The green along this gray area is bluish, basad. On
the outer margin the green extends down to 2A. Find wing with a white
spot on the costa, near outer angle, basally edged with dark red. A similar,
but post-discal, spot in M2-M3 and one in Cuj-Cua. In Cu2-2A is a white
dash, basally edged with red, and in 2A-inner margin is a similar, but heavier
dash. Anal lobe red, extending on outer margin to Cu2. In Cm-Cua is a
submarginal red dash. Fringe of both wings rusty-red, outwardly paler on
the hind Aving betAveen the veins.

Length of fore wing : Male, 15 mm.

256

Journal New York Entomological Society

[Vol. LII

Holotype, male, Massaranduba-Blumenau, Brasil (ex coll. E. I.
Huntington (no. 1009)); in the American Museum of Natural
History.

Remarks. Evidently closely allied to amyntor Cramer,1 of
which, unfortunately, no males have been examined. The holo-
type of caramba was compared with a female (in M.C.Z., no data,
ex coll. A. G. Weeks) of amyntor , the description and figure of
Draudt (1919, in Seitz, Macrolep. World, 5, p. 762, pi. 153 i),
and Cramer’s descriptions and figures of amyntor ( loc . cit.) and
menalcas (1782, Pap. Exot., 3, p. 117, pi. 259, figs. A and B).

Above, caramba differs from amyntor (of Draudt) in the nar-
rower, better defined marginal border. Below, from the female
at hand, the present species seems to have more of an anal lobe
and less prominent white markings. The absence of a tail is
probably very significant, and would seem to connect amyntor
to the acaste group, particularly to those members of that group
with a green frons. It certainly destroys Draudt ’s character for
splitting the “ amyntor -group” into two sections ( i.e ., the tailed
section, comprising the true amyntor group, to which caramba
obviously belongs, and the tailless section, more aptly called the
acaste group, and to which caramba bears little (relative)
resemblance.

Thecla punona, new species

Eyes ringed with white. Frons with moderately profuse, long, erect pale
gray hairs. Collar above with brown and gray hairs intermingled; on the
sides, white. Antennce largely white below, black above, but with thin an-
nulations of each extending into the other; club black. Thorax black above,
with sparsely strewn bronzy-brown hairs; below with pallid gray hair, quite
profuse. Abdomen gray-tan above, gray below.

Upperside :

Female?. Both wings slightly lustrous brown. Hind wing with outer
margin from M3 to the anal angle scalloped, and with a narrow whitish
marginal thread-line from Cu2 to 2A. Fringe of both wings dirty white,
basally darker ; that of hind wing obscurely dark at the vein-ends.

Underside :

Female?. Fore wing tan. A post-discal, slightly curved, dark brown line,
outwardly white, proceeds from costa to Cu2. Immediately without this line
at the apex is a hoary area, extending from costa roughly to M3, A sub-

1 1779, Pap. Exot., etc., 1, p. 76, pi. 48, fig. E.

Sept., 1944]

Clench: Lyc^enid^e

257

marginal, frequently indefinite line crosses faintly through this, increasing in
visibility below it until its disappearance at 2A. Kind wing hoary gray,
darker on the outer margin. A submarginal row of faint yellowish or pallid-
fulvous lunules parallels entirely the outer margin. A curved, irregular, dis-
cal, brown line, tan within and white without, crosses from costa to inner
margin, and a post-basal, similar one also, though necessarily shorter. The
two close off an area, slightly darker than the surrounding ground, which is
just basad of the Center. Occasionally the ground color is so disposed as to
give the appearance of a row of pale gray lunules capping the yellowish ones
on the outer margin. Fringe similar to the upperside but rather darker.

Length of fore wing : Female?, 9.5-11 mm.

Holotype, female (f), Puno, Peru, 12,500 feet, Nov. 1, 1898
(Wm. J. Gerhard).

Paratypes, two females ( ?), same data as holotype.

Holotype and one paratype, no. 26258 in the M.C.Z. One para-
type in the collection of the author.

Remarks. Rather closely allied to oribata Weymer2 and pos-
sibly only subspecifically distinct from it. The principal differ-
encse are : the less apically produced fore wing and less anally
produced hind wing (if Weymer ’s figure is correct); the less
distinct outer row of spots on the fore wing below, and the less
apparent submarginal line on the hind wing below; the more
prominent basal dashes ; the less distinctly black-checked fringe,
particularly on the hind wing. A single specimen from Chulu-
mani, Bolivia (Nov. 28, 1898, W. J. Gerhard), not made part of
the type series, approaches oribata in several of these respects.

Weymer ’s figure shows the lines on the underside to be quite
dark, almost black, while in punona they are brown. It is prob-
able, however, that the figure errs somewhat in this respect.

Draudt (1919, in Seitz, Macrolep. World, 5, p. 760, pi. 153 h)
described a new species under the name of anosma, which he later
(Seitz, add. and corr., p. 823) synonymized to oribata. In his
description he mentioned “incoherent dark lunular spots” and
“longer small tail on the lower median vein” neither of which is
particularly clear. The latter particularly would seem to exclude
it entirely from the culminicola-gr owp. It is presumed that by
“median ’ ’ he meant 1 1 cubital. ’ ’

2 1890, Lepidopteren gesammelt . . . von A. Stiibel (Rhopaloeera by Wey-
mer), p. 123, pi. 4, fig. 8.

258

Journal New York Entomological Society

[Yol. LII

Thecla kalikimaka, new species

Thecla hemon: Hoffman, 1940, Anales Inst. Biol. Mex., 11, p. 704 (no. 586)
( nec hemon Hiibner). _ Ciu. i w >

Eye ringed obscurely with white. Frons brown. Collar brown. Antennce
black-brown, club dull fulvous ; below, white annulate, club brighter fulvous.
Palpi brown, slightly hoary within. Thorax metallic green-blue above,
slightly hairy; below, covered with uniform gray-brown hair. Abdomen above
green-blue, below brown. Legs brown-black, tarsal segments white annulate.

Upperside :

Male. Both wings somewhat bronzy green. Fore wing with a compound
scent-pad consisting of a dark dash across the cell-end and a rectangular
brown patch just beyond. Outer margin moderately heavily edged with
brown, thickening considerably towards the apex. The basal limit of this
brown is very tenuous and indefinite. Hind wing with a small anal area of
brown, and two tails, one each at Cux and Cu2, the latter longer and emitted
tangentially. Anal lobe deeply incised, as is usual in this group.

Underside :

Both wings brown. Fore wing with a faint, dull, metallic green iridescence.
Paler on inner margin. A discal dark brown line crosses from costa to Cu2,
quite straight. Mind wing with a straight discal line from Rs opposite the
outer angle to just beyond Cu2 in the center of the wing. From there it be-
comes metallic green, and proceeds to 2A, where it angles sharply basad and
terminates at the inner margin. A submarginal line starts at about M2 and
proceeds parallel to the outer margin, reaching just beyond Cu2 where it angles
basad, to terminate at the inner margin, edged inwardly from Cu2 with a few
metallic scales. A pale, hardly noticeable marginal line runs from outer
angle to apex, becoming very obscurely greenish from Cu± on. Anal region
restrictedly irrorated with pale scales. A faint line, post-basal, starts on
costa and proceeds to lower cell-end. Fringe of both wings brown.

Length of fore wing : Male, 16 mm.

Holotype, male, Jalapa, Mexico, date and collector unknown,;
ex coll. A. G. Weeks, Jr.

Paratype, male, same locality, Sept. 8, 1884 (“Morrison”),
ex coll. F. A. Eddy.

Holotype, M.C.Z. 26257. Paratype in the author’s collection.

Remarks. Apparently most closely allied to mavors and
triquetra (references and authorships of these names below).
From both it differs in the brown color of the underside, which
in these species is bright metallic green or blue. The lines below
are less apparent than in either of the two species (except thaf
in kalikimaka there is a discal line on the fore wing, lacking in

Sept., 1944]

Clench: Lyc-enid^e

259

mavors). Above it differs from triquetra in the green color (blue
in that species), and from mavors in the lack of a pale, almost
white, anal line. The apical and marginal border of the fore
wing appears slightly heavier than in either mavors or triquetra.
It would seem closest to mavors, since it has the two tails of that
species, as opposed to the single one of triquetra.

A brief discussion of the species of this group (as limited by
Drauclt, p. 750) might not be amiss.

Thecla mavors Hiibner (1818, Zutr. zur Samml. Exot. Schmett.,
1st. Hund., p. 31 (no. 95), figs. 189, 190). Quite variable, but
whether racially or not cannot be determined from the material
at hand. The male is metallic green above, with a compound
scent-pad, and is bordered narrowly and indistinctly with black-
brown on the fore wing. Hind wing at the anal angle edged
with very pale green, almost white. Below green, fore wing
brown on inner margin, otherwise practically immaculate. Hind
wing with a discal and submarginal line, each angled basad at
Cu2-2A. Outward of the latter is a band of brownish maroon
from Mi to the anal angle. Anal lobe similarly colored. Two
tails, the larger at Cu2 tangentially emitted.

Female, brown above and below. The latter surface crossed
on both wings by thin brown lines; two on fore wings, parallel
to the outer margin, and a short one crossing the cell-end. An-
other pair on hind wing, also parallel to the outer margin, but
angling basad at Cu2-2A. A post-basal line commences on costa,
crosses cell-end and stops.

Specimens in the M.C.Z. from Suapure, Venezuela; Cusilluni,
Bolivia; Cumato Arepo, Savanna, Trinidad; Rio Tapajoz, Brasil.

A Suapure female has the two lines on the hind wing abnor-
mally close together, but whether or not this is an individual
variant cannot be stated without additional material.

Thecla triquetra Hewitson (1862-1869, 111. Diurn. Lep. Lycas-
nidas, p. 76, pi. 28, figs. 17, 18, 19). Differs from mavors in the
male by being blue above, having but one tail (also tangentially
emitted), the bluer color below, and the presence of a discal line
on the fore wing, and heavier ones on the hind wing. Females
differ from mavors females chiefly in the lack of a Cui tail.

260

Journal New York Entomological Society

[Vol. LII

Otherwise they appear very similar. There is, perhaps, a slight
difference in the shape of the hind wing, but more specimens are
needed to confirm it.

In the M.C.Z. from Blumenau and Rio de Janeiro, Brasil.

Theda pawper a Felder (1865, Reise Novara, Lep. 2, p. 246,
pi. 31, fig. 15). May be distinguished from the above in the
male by the dark green color and the lack of a scent-pad on the
fore wing. Differs below in that the inner of the two longest
lines touches the outer at the anal angle.

Apparently not in the M.C.Z. (see under ella).

Theda drucei Lathy (1926, Ann. Mag. Nat. Hist., (9) 17, p.
41). This species, also not in the M.C.Z., is figured in the Trans.
Ent. Soc. London, 78, pi. 9, fig. 9 (plate accompanying paper
that begins on p. 133). It apparently belongs near pauper a and
harrietta, but differs, in the male, by the possession of a scent-pad.
There are other differences as well.

Theda ella Draudt (1919, in Seitz, Macrolep. World, 5, p. 750,
pi. 148 b as mavors). Differs from pauper a in that the inner
line below on the hind wing is w-shaped at the anal angle. This
is. the difference brought out by Draudt. He noted the fact that
paupera was rare, and very likely based this difference on the
figure of Felder, which does show no “W” at the anal angle
of the hind wing below. This character, however, may be very
faint (as in the single specimen here provisionally assigned to
ella ), and consequently overlooked when Felder’s specimen was
figured.

A single specimen in the M.C.Z. (Bogota, Colombia?) ex coll.
A. G. Weeks, may possibly be this species. It has, however, much
heavier borders above than Draudt ’s illustration shows, and
narrower lines below. It might also be paupera , but again the
markings below are all thinner than Draudt ’s and Felder’s
illustrations show.

Theda harrietta Weeks (1901, Can. Ent., 33, p. 294). This
is a perfectly good species, allied to paupera and ella. Besides
the greatly differing color above, the lines below are rather dif-

Sept., 1944]

Clench: Lyozenhle

261

ferent, the central apex of the “W” being rather shorter than
in the specimen cited above, and in Draudt’s figure of ella. The
discal line on the fore wing extends into the Cu2-2A interspace,
which it apparently does not in the other species. In other
Theclinae, however, this character has been found to be individu-
ally variable.

One specimen in the M.C.Z. (Type, M.C.Z. no. 16673) from
Coroico, Bolivia, April 20, 1899 (Wm. J. Gerhard).

262

Journal New York Entomological Society

[Vol. LII

NOTE ON THE DEATH-FEINT OF BRUCHUS
OBTECTUS (SAY)

The common bean weevil, Bruchus obtectus, exhibits a wide
variation in the duration of its comparatively brief death-feint.
Out of 283 freshly emerged beetles, only 72 could be induced to
feign death. Although dropped from a height of 12 inches, hav-
ing their thoraxes pinched, being rolled between the thumb and
forefinger, 211 of the weevils refused to perform in spite of all
the handling. The 72 that actually went into death-feints, fre-
quently did so, with comparatively little trouble. A slight dis-
turbance such as touching them with the tip of a pencil, or causing
them to fall one inch, or gently pressing the sides of the thorax
brought about the desired reaction.

The temperature during these tests was 70° F. For the 72
beetles that reacted, the duration of the feint varied in length
from 1 to 300 seconds. The duration of the weighted, arithmetic
average death-feint was 33.5 seconds, and the standard deviation
was 74.8. The following table shows the duration of the death-
feint in the 72 different beetles that reacted :

No.

beetles

Length of
death-
feint

No.

beetles

Length of
death-
feint

No.

beetles

Length of
death-
feint

No.

beetles

Length of
death-
feint

Seconds

Seconds

Seconds

Seconds

2

1

1

12

5

30

1

75

4

2

2

14

1

32

1

78

3

3

6

15

1

33

1

80

1

4

3

17

1

35

1

100

5

5

1

18

1

40

1

105

1

6

5

20

1

50

1

250

2

8

1

22

1

53

1

290

2

9

1

23

1

57

1

300

3

10

3

25

1

60

2

11

1

29

2

70

Harry B. Weiss.

Sept., 1944]

Michener: Culex

263

DIFFERENTIATION OF FEMALES OF CERTAIN
SPECIES OF CULEX BY THE
CIBARIAL ARMATURE

By First Lieutenant Charles D. Michener1
Sanitary Corps, Army of the United States

The purpose of this paper is to describe characters of the
cibarial (or ‘ ‘pharyngeal”) armature of the Culex found in the
southeastern United States. These characters support the usual
subgeneric classification and serve in addition to distinguish
females, previously considered indistinguishable, of certain of
the species commonly placed in the subgenus Melanoconion (e.g.,
by King, Bradley, McNeel, 1942).

The cibarial armature has been used by a number of authors
in separating the subgenera and certain Old World species of
Anopheles. Christophers (1933) gives a good account of the
structures involved in Anopheles.

The armature lies on the posterior margin of the anterior pump
of alimentary canal. This pump has usually been called the
pharynx, but Snodgrass (1943) has pointed out that it is in
reality the cibarium. The second pump is the true pharynx.
The ventral surface of the cibarium is sclerotic. This sclerotized
area ends posteriorly in a broadly concave margin lying between
the two cibarial cornua, which are apodemal muscle attachments
at the posterior end of the cibarium. The cibarial armature,
which is present only in females, consists of a series of sclerotized
projections or teeth along the margin between the cornua.

With a little practice these structures can be studied about as
easily as the male genitalia. The technique used is as follows:
The head is placed in 10 per cent potassium hydroxide and heated
for a short time. Then it is placed in water on a slide and under
a binocular microscope the entire outer wall of the head is broken
away in large pieces by means of fine needles. The pharynx and
cibarium, attached to the hypopharynx, may now be seen within

1 Acknowledgment for both helpful advice and specimens for dissection is
made to Major Stanley J. Carpenter and Captain Woodrow W. Middlekauff.

264

Journal New York Entomological Society

[Vol. LII

and are transferred, with or without separation from the other
mouthparts, to a drop of chloral hydrate medium (Berlese’s
medium) on a slide. Here the pharynx is pulled away from the
cibarium. The cibarial armature will probably still not be
clearly visible because of the imbricated sclerotized teeth in the
membrane of the dorsal surface of the cibarium between the
cibarial cornua. Therefore, this membrane with its imbricated
teeth is dissected away from the dorsal surface of the cibarium in
order to expose clearly the armature on the ventral surface. This
membrane should be preserved, however, since its imbricated
teeth offer characters of importance. The mount is completed
with a coverglass. The separation of individual teeth as recom-
mended by Christophers (1933) for Anopheles has not proved
to be possible with Culex.

The following key separates the species of Culex found in the
southeastern United States into groups on the basis of female
cibarial armatures. Culex atratus Theobald and C. ~bahamensis
Dyar and Knab, found in this area only on the Florida Keys,
have been omitted because of the lack of specimens for study.

1. Cibarial armature consisting of 25 to 35 small, slender teeth; imbricated

teeth of the membrane of dorsal surface of cibarium between cornua

small, lightly sclerotized, brownish 2

-. Cibarial armature consisting of 3 to 8 large, blunt teeth; imbricated teeth
of membrane of dorsal surface of cibarium between cornua large,
heavily sclerotic, blackish (Subgenus Melanoconion ) 3

2. Cibarial teeth not longer than width of a cibarial cornu.

(Subgenus Culex s. str.) pipiens Linnaeus, quinquef asciatus Say,
tarsalis Coquillett, salinarius Coquillett, nigripalpus Theobald
-. Cibarial teeth much longer than width of a cibarial cornu.

(Subgenus Neoculex) apicalis Adams

3. Cibarium with three teeth pilosus (Dyar and Knab)

-. Cibarium with seven or eight teeth.

erraticus Dyar and Knab, peccator Dyar and Knab

These characters are shown in greater detail in the figures.
Except to strengthen the recognized subgeneric classification, the
cibarial characters contribute little to our knowledge of the sub-
genera Culex s. str. and Neoculex.

The three remaining species, however, are externally indis-
tinguishable in the females except that most specimens of errati-

Sept., 1944]

Michener: Culex

265

cus have some coarse, golden, mesoscntal scales and the broad,
appressed, occipital scales are limited to a band along the eye
margin, while in the other two species the mesoscntal scales are
usually dark and the occiput is usually mostly covered with
broad, appressed scales. These characters do not hold for every
specimen. The cibarial characters thus make possible the identi-
fication of female pilosus.

It is interesting to note that Edwards (1932), on the basis of
larval characters, placed pilosus in the subgenus Mochlostryax,

Fig. 1. Cibarial armatures of: 1, Culex {Culex) restuans; 2, Culex {Neo-
culex) apicalis Adams; 3, Culex ( Melanoconion ) pilosus (Dyar and Knab) ;
4 and 5, Culex ( Melanoconion ) erraticus Dyar and Knab.

and erraticus and peccator in Melanoconion. The cibarial arma-
tures of the three species here discussed support this classifica-
tion, but until these characters have been studied for the numer-
ous tropical species of this group, it is not advisable to consider
the cibarial armature as a subgeneric character.

The cibarial armatures of about twenty specimens of C. pilosus
have been examined and found quite uniform in structure. Sev-
eral of those studied were from a series reared from pilosus larvae
by Mr. Wm. V. Reed. The armatures of forty other female

266

Journal New York Entomological Society

[Vol. LII

specimens of Melanoconion were examined. All were seven- or
eight-toothed. Most of them, on the basis of the characters of
the vestiture already mentioned, were presumed to be C. erraticus
and two were from a series reared from erraticus larvas by Lt.
Basil G. Markos. Among the six specimens studied having seven
or eight cibarial teeth but vestiture as in pilosus and peccator,
two from different localities were collected with males of peccator
and are, no doubt, females of that species. Both of these had
but seven teeth.

LITERATURE CITED

Christophers, S. R. 1933. The fauna of British India, Diptera, Vol. IV,
Family Culicidae, Tribe Anophelini, vi + 371 pp., illus.

Edwards, F. W. 1932. Diptera, Fam. Culicidas, in Wytsmann’s Genera
Insectorum, fasc. 194, 258 pp., illus.

King, W. V., G. IT. Bradley, and T. E. McNeel. 1942. The mosquitoes of
the southeastern states, U. S. Dept. Agr., Misc. Publ. 336, 96 pp., illus.
Snodgrass, R. E. 1943. The feeding apparatus of biting and disease carry-
ing flies: a wartime contribution to medical entomology, Smithsonian
Misc. Colls., vol. 104, no. 1, pp. 1-51, illus.

Sept., 1944]

Weiss: Insect Behavior

267

INSECT RESPONSES TO COLORS

By Harry B. Weiss

The purpose of this article is to call attention to the approxi-
mately similar qualitative results obtained by various workers
who used widely different methods of approach and technique in
studying the behavior of insects to different wave lengths of light.
No attempt will be made here to describe the techniques, as they
are already matters of record. In fact they differ as widely as
the methods of approach.

Bertholf (1, 2) exposed the honey bee, and the fruit fly
Drosophila to two translucent glass plates of equal size, one
illuminated with white light and the other with monochromatic
light obtained by means of a quartz prism. The intensity of the
white light was changed until its effect on the photopositive re-
sponse of the insects was equal to that of monochromatic wave
lengths in different portions of the spectrum. He found that for
the honey bee, the stimulating efficiency increased from zero at
6450 A to a maximum of 100 arbitrary units at 5500 A, then
decreased to 10 at 4350 A after which it rose rapidly to a second
maximum of 450 at 3650 A and then rapidly declined to zero
at about 2800 A.

As for Drosophila, Bertholf (2) stated . . starting with
the longer wave lengths the efficiency is very low until it starts
to rise at about 5750 A; from here it rises to a maximum in the
so-called visible spectrum at 4870 A . . . ; from this wave
length it decreases again at 4250 A ; then it rises suddenly and
attains a maximum value at 3650 A . . . ; from here it de-
creases rapidly to zero at 2540 A. ’ ’ Bertholf worked with 30
wave lengths from 2300 A to 7000 A.

Weiss et al. (6, 7, 8) exposed approximately 15,000 insects,
both adult and larval forms, mostly diurnal, but some nocturnal,
involving 40 species in various orders to 10 wave-length bands of
light of equal physical intensities from 3650 A to 7400 A. Each
test lasted from 15 to 30 minutes and a group of 100 or more of
each species was used for from one to three successive tests. The

268

Journal New York Entomological Society

[Vol.LII

composite group behavior pattern, both for larvas and adult
insects indicated that the stimulating efficiency increased only
slightly from zero at 7200 A to 5750 A. From here it rose to a
maximum at 4920 A. It then declined to a comparatively low
level at 4640 A from which point it ascended to its peak maxi-
mum level at 3650 A.

These authors found that, regardless of the relative positions
of the wave-length bands, the insects made approximately the
same selections time after time, also that when a second and third
test followed the first, there was a shifting of individuals that
went to the different colors, but no change in the final result.
In addition some species such as the Japanese beetle and the Colo-
rado potato beetle responded to what were unattractive wave
lengths under equalized physical intensities, when the intensities
of such unattractive wave lengths were increased. In other
words, it was possible to vary the behavior pattern by varying
the intensities.

Crescitelli and Jahn (3), approached the problem from the
standpoint of the electrical responses of the dark-adapted grass-
hopper eye. “Leads were taken with silver-silver chloride elec-
trodes from fluid-filled chambers about each eye. The entire
surface of one eye was illuminated, and the other eye was kept
in darkness. Records were obtained by means of a cathode ray
oscillograph. For the experiments on colored light Corning color
filters were placed between the light source and the eye.” Six
wave-length bands were employed, extending from about 4000 A
to 7000 A. “The relative intensity transmitted through each
of these six filter combinations was determined by means of a
thermopile and galvanometer. The infra-red radiations were
completely removed from the stimulating light by using 5 cm.
of water and a Corning (AKLO) heat absorbing filter.”

These authors studied the change in form of the electrograms
of the grasshopper eye under variations in intensity of the stimu-
lating light and also the quantitative aspects of the response in
relation to the quality of the stimulating light. They found that
there was apparently no specific effect of wave length on the
electrical response of the whole dark-adapted grasshopper eye.
At equalized intensities there were decided differences in wave

Sept., 1944]

Weiss: Insect Behavior

269

form with the six different spectral bands, but these disappeared
and the color responses were exactly matched when the intensities
of the different spectral regions were properly adjusted. Quoting
again from their paper : ‘ ‘ The form of the electrical response of
the dark-adapted grasshopper eye to brief stimulation by white
or colored light varies according to the intensity of the light.
At very low intensities the response is diphasic, the initial posi-
tive phase of which resembles the a- wave of the vertebrate electro-
retinogram. As the intensity is increased the positive phase
decreases and changes its position while the negative phase be-
comes increasingly prominent. Eventually the positive phase
is completely eliminated and the electrogram takes the form of
the typical high-intensity response. The order of effectiveness
of the different colors in causing this change in wave form is :
green, blue, violet, orange-red, red.”

The curve relating the magnitude of the potential to the wave
length had a peak in the green region of the spectrum, and de-
clined sharply toward the red and less sharply toward the violet.
The magnitude of the electrical response was found to be defi-
nitely related to the quality of the stimulating light and the form
of the response to be influenced by the intensity of the stimu-
lating light, either white or colored.

Jahn and Crescitelli (5), also studied, in the same manner, the
electrical responses of the compound eye of the moth Samia
cecropia, in relation to the quality and intensity of the stimu-
lating light. Part of their conclusions are quoted as follows :
“The electrical responses of the moth and grasshopper eyes to
wave length are surprisingly similar. For both animals the same
type of graph is obtained when the relative magnitude of the
potential is plotted against wave length. This graph has a gen-
eral similarity to the absorption curve of visual purple. Another
aspect of the electrical response to wave length concerns the fact
that no specific effects of wave length on the electrograms are
discernible. By properly adjusting the intensity, the responses
to one color may be exactly matched with the response to any
other color, indicating that the differences in the responses to
different colors of equal intensity are caused merely by differ-
ences in sensitivity and are not effects of wave length per se.”

270

Journal New York Entomological Society

[Vol. LII

In the case of the moth eye the maximum response was obtained
with the green band. The responses dropped sharply toward the
red band and less sharply toward the violet.

Graham and Hartline (4) studying the responses of single
visual sense cells of Limulus to visible light of different wave
lengths found that when the energy of the stimulating light of
different wave lengths was approximately equal, the response to
green was stronger than the responses to either violet or red.
When the energy was increased in the red and violet their level
of response was raised and when the intensities of the different
wave lengths were adjusted so that the responses were equal,
there was no effect of wave length as such, indicating that single
sense cells can gauge brightness, but cannot distinguish wave
length. The relative energies of the various wave lengths re-
quired to produce the same response, after being adjusted in
inverse ratio to the degree of their absorption yielded a visibility
curve for a single visual sense cell that had its maximum in the
green near 5200 A and that declined symmetrically on each side
to low values in the violet near 4400 A and in the red near
6400 A.

Thus the visibility curves of a single visual sense cell of Limu-
lus, although not an insect, of the eye of a grasshopper, a diurnal
insect, and of the eye of a Cecropia moth, a nocturnal insect,
are qualitatively similar to the curve of the relative stimulating
efficiency of different wave lengths of light for Drosophila, as
reported by Bertholf and to the behavior curves for the numerous
adult and larval forms of diurnal and some nocturnal insects as
reported by Weiss et al. These curves are not identical because
of the different methods of approach and technique but they are
all strikingly similar for the visible portion of the spectrum. All
were obtained under wave lengths of equalized physical intensi-
ties. Hartline and Graham and Crescitelli and Jahn by properly
adjusting the intensity were able to match the response to one
color with the response to any other color and Weiss et al., in their
behavior studies found that insects responded to what were un-
attractive colors under equalized intensities, when the intensities
of these colors were increased.

Crescitelli and Jahn (3) report that other authors who worked
with pigeon eyes and the eyes of certain vertebrates also found

Sept., 1944]

Weiss: Insect Behavior

271

that wave form difference are simply intensity differences and
that the electrical response to different wave bands could be
duplicated by adjusting the intensity of the different bands.

Thus it appears that both the electrical responses of the insect
eye and the motor responses of the insect itself to different colors
of equal intensity are due to differences in sensitivity, or to the
absorption of light, which varies with wave length, by the pri-
mary photosensitive substance of the visual sense cells, and are
not the effects of wave length by itself.

REFERENCES

1. Bertholf, L. M. The distribution of stimulative efficiency in the ultra-

violet spectrum for the honey bee. Jour. Agr. Res. 43 (8) : 703-
713. 1931.

2. Bertholf, L. M. The extent of the spectrum for Drosophila and the

distribution of the stimulative efficiency in it. Zeitsch. Yergl.
Physiol. 18: 32-64. 1932.

3. Crescitelli, Frederick, and Theodore Louis Jahn. The electrical

response of the dark-adapted grasshopper eye to various intensities
of illumination and to different qualities of light. Jour. Cellular
and Comp. Physiol. 13 : 105-112. 1939.

4. Graham, C. H., and H. K. Hartline. The response of single visual

sense cells to lights of different wave lengths. Jour. Gen. Physiol.
18 (6) : 917-931. 1935.

5. Jahn, Theodore Louis, and Frederick Crescitelli. ' The electrical

responses of the Cecropia moth eye. Jour. Cellular and Comp.
Physiol. 13 : 113-119. 1939.

6. Weiss, Harry B., Frank A. Soraci, and E. E. McCoy, Jr. Insect be-

havior to various wave lengths of light. Jour. N. Y. Ent. Soc.
49: 1-20; 149-159. 1941. 50; 1-35. 1942. 51:117-131. 1943.

7. Weiss, Harry B., E. E. McCoy, Jr., and William M. Boyd. Group

motor responses of adult and larval forms of insects to different
wave lengths of light. Jour. N. Y. Ent. Soc. 52: 27-43. 1944.

8. Weiss, Harry B. The group behavior of 14,000 insects to colors. Ent.

News 54 : 152-156. 1943.

272

Journal New York Entomological Society

[Yol. LII

HIBERNATION OF THE SYRPHID FLY,
LATHYROPHTHALMUS iENEUS SCOP.

Lathyrophthalmus ceneus Scopoli has the interesting habit of
hibernating gregariously in old nests and burrows of Aculeate
Hymenoptera. On several occasions in the last few winters the
writer has encountered the species in open tunnels of various
anthophorid bees situated in vertical banks or cliffs. In Febru-
ary, 1944, however, several hundred specimens were found hiber-
nating in empty nests of the black and yellow mud-dauber,
Sceliphron servillei (Saussure), in various localities in the San
Joaquin Valley, California. The localities included Merced in
Merced County, Chowchilla and Berenda in Madera County,
Tulare and Tipton in Tulare County, and Kingsburg and Fowler
in Fresno County. In all cases the nests utilized for hibernation
were situated under bridges or culverts. Usually five or six
individuals, more rarely eight or nine, occupied a single empty
mud cell. Prof. F. M. Hull, who very kindly made the identifi-
cation, reported that both sexes were present in a series of ap-
proximately thirty specimens submitted to him, with the females
slightly predominating.

The only other hibernating insect using these nests and ap-
proaching the syrphids in abundance was the elm leaf beetle,
Gallerucella xanthomelcena (Schrank). These, however, were
hibernating elsewhere in the vicinity under a variety of different
conditions, whereas the syrphids appeared to exhibit a marked
preference for this particular habitat. — E. Gorton Linsley.

Sept., 1944]

Clench: Lyc^enid.®

273

TWO NEW SUBSPECIES OF LYC^NOPSIS
PSEUD ARGIOLUS BDV. & LEG. (LEPI-
DOPTERA, LYC/ENIDiE)

By Harry K. Clench
Cambridge, Mass.

The following new subspecies have recently come to my atten-
tion and appear to be worthy of names. As is well known, the
interrelationships of the various named entities of North Ameri-
can Lycamopsis are not as yet satisfactorily determined. The
following two have, therefore, been placed under pseudargiolus
as simple races. It is possible that later on they will be classified
differently.

Lycsenopsis pseudargiolus sidara, new subspecies

Upperside :

Male. Both wings uniform, slightly violet blue, with a thin blackish border
on each. Fringe white, dark toward the apex of the fore wing and occasion-
ally faintly at the vein-ends of both wings.

Female. Both wings shining blue, with a violet tinge. Fore wing with
a broad, dark, marginal border, thickest at the apex. Costa with the blue
whiter and the marginal brown border slightly extended basad. Kind wing:
Costa shaded with brown. Outer margin with a thin black line, basal to
which is a row of spots, one to each interspace. Fringe of both wings similar
to that of the male.

Underside:

Male. Both wings white, grayish in the basal half. Fore wing with the
cell closed by a faint dark line. A thin marginal line runs from costa to
inner margin, basal to which is a row of small spots, each to an interspace.
Basal to these is a scalloped line, sometimes appearing as a row of angled
bars, vertices inward. A post-discal series of heavy short bars runs from
costa to inner margin. The lower (inner marginal) one is very faint, occa-
sionally almost non-existent, but the others are very heavy, and each is set
on the bias, costal end inward, and more heavily so costad than elsewhere.
The costal dash is basally dislocated. Hind wing : The marginal line, series
of dots, scalloped line and cell-end bar as in the fore wing. The post-discal
row of spots is heavy as in the fore wing, but quite dislocated. The two
costal spots are placed inward, the next two outward, and the next four
alternately inward and outward. The base of the wing is scaled with black-
ish. Three basal spots, one on the costa, one in the cell, and one on the

274

Journal New York Entomological Society

[Vol. LII

inner margin, roughly parallel to the body line. These spots, the cell-end
bar and the post-discal series are all outlined faintly in white.

Female. Purer white, lacking the gray basal shading; otherwise similar
to the male.

Holotype, male, Manitou, Colorado, June 5—11, 1882 (ex S. H.
Scudder Coll.).

Allotype, female, same data as holotype.

Paratypes, three males, Starr Ranch, El Paso County, Colorado
(6500 ft.); May 12, 1934; one male, Rock Creek, El Paso Co.,
Colorado (6800 ft.), May 9, 1934; one male, “Col.” (ex S. H.
Scudder coll.).

Holotype, allotype, and four paratypes in the collection of the
Museum of Comparative Zoology. One paratype in the author’s
collection.

Remarks. This subspecies bears a rather close resemblance to
the form named violacea 1 by Edwards. Violacea is represented
in part in the M.C.Z., by a pair of topotypes and a series of speci-
mens labelled “Coalb.,” with various dates around 1883, all re-
ceived through the Scudder collection, they having been received
from Edwards. Sidara may be separated from these specimens
by the closer proximity of the post-discal series of dashes (prin-
cipally on the fore wing below) to the scalloped line. Also, in
sidara the spots of the post-discal series of the hind wing tend to
be more rounded, while in violacea they are either more linear
(usually) or are more quadrate. In sidara the ground color
below tends toward gray, while in violacea it is nearly pure white,
making the white ringing of the discal and basal spots of sidara
absent in violacea. A single male from Cloudcroft, New Mexico,
is apparently of this subspecies, but it is less heavily marked
below, and lacks the basal graying, and the subsequent white-
ringing of the spots.

Lycaenopsis pseudargiolus bakeri, new subspecies

Upperside :

Male. Purplish blue, nearly violet, the costal veins pencilled in whitish
blue. Costa and inner margin of hind wing whitish. Both wings very nar-
rowly bordered by a thin black line. Apical half of the outer margin of the

1 1866, Proc. Ent. Soc., Philadelphia, 6, p. 201. Type loc. : Kanawha River,
West Virginia.

Sept., 1944]

Clench: Lyc^enid^e

275

fore wing narrowly black. Fringe of fore wing white, becoming black to-
wards the apex and at the vein-ends. That of the hind wing white, occa-
sionally faintly dark at the vein-ends.

Female. Fore wing dark brown, with a central area of blue, of varying
extent. A small streak closes the cell. Hind wing also brown, and with a
marginal series of pale blue lunules enclosing small brown spots. Fringe as
in the male.

Underside:

Male. Both wings white. Markings arranged as in other pseudargiolus
forms (see sidara above, for example), the marginal compound border of
both wings rather faint, the post-discal, discal and basal spots very well
marked, dark, and not tending to confluence. Base of hind wing tinged with
greenish.

Female. As in the male.

Holotype, male, Baker, Oregon, April 20, 1941 (J. H. Baker).

Allotype, female, Durkee, Oregon, May 11, 1941 (J. H. Baker).

Paratypes , nos. 1 to 5, male, same data as holotype; nos. 6 to
10, male, Pine Creek, Baker, Oregon, May 26, 1941 ( J. H. Baker) ;
nos. 11, 12, male, Kane Creek, Oregon, March 16, 1934 (F. W.
Lawrence) ; nos. 13, 14, male and female resp., same locality and
collector as allotype, May 19, 1941.

Holotype and allotype in the Museum of Comparative Zoology.
Two male paratypes to be returned to Mr. Baker. Remaining
paratypes in the author’s collection.

Remarks. Both sexes differ from the Californian race echo2 in
the much darker, and therefore more prominent, discal and basal
spots of both wings below. The female above has slightly re-
duced blue areas. Baker i apparently connects, to a greater or
lesser degree, echo with nigrescens Fletcher,3 from which latter
it may be told in the greater amount of blue in the females and
in the constancy of the underside pattern. Quoting Fletcher,
*‘The underside of this variety ( nigrescens ) is remarkable and
specimens of both sexes may be found which, if the underside
alone were seen, might be referred to neglect a, violacea, lucia or
marginata, and some even combine characters of all these. One
beautiful form which frequently occurs, has an irregular, discal,
dark blotch of confluent spots on the secondaries beneath as in

2 1864, Proc. Ent. Soe. Philadelphia, 2, p. 506. Type loc. : California.

3 1903, Trans. Eoy. Soe. Canada, (2) 9, p. 213, fig’d. Type loc.: Kaslo,
B.C.

276

Journal New York Entomological Society

[Vol. LII

lucia, and the clear marginal spots of violacea. This form Mr.
Cockle, who has collected this butterfly for several years and has
been much interested in it, considers to be most typical of the
variety.” In all the 16 specimens (14 of them males) of bakeri
examined, there was but one specimen (paratype no. 8) that ex-
hibited the slightest tendency towards the lucia- like confluence
of spots on the underside of the hind wing, and this specimen
was obviously slightly aberrant. The only indication of cliscal
confluence was the basal thickening of the Cui-Cu2, spot of the
post-discal series. The further indication of aberrance was given
by the thickened scalloped line of the submarginal compound
border.

This subspecies is named for Mr. James H. Baker, of Baker,
Oregon, from whom the majority of the specimens were received.

Sept., 1944]

Linsley: Neopasites

277

NEW SPECIES OF NEOPASITES WITH NOTES
CONCERNING OTHERS (HYMEN-
OPTERA, NOMADIDiE)

By E. Gorton Linsley
University of California, Berkeley

Since the publication of a revision of this genus,1 I have had
the privilege of studying additional material through the kind-
ness of Mr. J. N. Knull, Mr. P. H. Timberlake, and Prof. 0. A.
Stevens. The present paper includes the results of this study.

Neopasites elegans Linsley, new species

Female : Form very robust ; color black ; abdomen deep red throughout ;
integument densely, coarsely punctate, moderately dull. Head very densely
punctate, with a large patch of dense appressed white hairs around the
antennal bases, a large white patch behind each eye, the two connected
posteriorly along the posterior margin of the head and the latter by a narrow
median line from the vertex, through the median ocellus to the antennal white
patch; antennas very dark brown, first flagellar segment about as long as
following two together; upper frons and vertex coarsely, contiguously and
subcontiguously punctate, punctures of ocell-ocular area at least as large as
those of disk of mesoscutum ; clypeus densely but more finely punctate ;
labrum clothed with long, dense white hair at base, surface finely punctate,
indistinctly longitudinally carinate for more than two-thirds of its length ;
mandibles black, apices reddish. Thorax coarsely punctate; pronotal collar
densely white pubescent, tubercles margined with white; tegulae very dark
piceous; mesoscutum coarsely densely punctate, margins and anterior three-
fourths of median line, densely clothed with appressed white hairs; meso-
scutellum coarsely, densely punctate, lateral margins and median line densely
white pubescent, the mid-line expanded at posterior margin into a subtri-
angular white patch; metanotum with a white patch on each side; mesepi-
sterna coarsely, very closely, somewhat rugosely punctate, vertical face
broadly margined with dense appressed white hairs; mesosternum with disk
white pubescent. Wings lightly infuscated, veins and stigma brown. Legs
dark brown; anterior and intermediate femora white pubescent beneath at
apex, intermediate and posterior tibiae with posterior face densely, outer face
more thinly, white pubescent. Abdomen deep red, without any dark cloud-
ing; first tergite with a large, suboval white patch on each side bordering

1 Linsley, E. G. 1943. A revision of the genus Neopasites. Trans. Amer.
Ent. Soc., 69: 119-140, fig. 1.

278

Journal New York Entomological Society

[Vol. LII

lateral margin of basal concavity, the patches separated by nearly twice
their width, basal concavity without white pubescence, tergites two to five
with a large oval white patch on basal margin, the pairs separated by about
their own diameters and becoming successively larger on succeeding seg-
ments except those of the fifth tergite which are smaller, tergites two and
three with a latero-basal white patch on each side those of second tergite
much smaller than the submedian basal patch, those of third tergite scarcely
evident, tergites one to four with a lateral white patch on apical margin,
that of fourth tergite small; fifth sternite bilobate, densely punctate, pubes-
cent, apical margin broadly and shallowly emarginate.

Length 5 mm.

Holotype female (collection of Ohio State University) and one
paratype, female, from El Paso County, Texas, August 30, 1940,
collected by D. J. and J. N. Knu.ll. Two additional paratypes,
female, were taken by Mr. and Mrs. Knull in Culberson County,
Texas, August 30, 1940. One paratype is deposited in the collec-
tion of Mr. P. H. Timberlake, another in the collection of the
writer, the third in the collection of Ohio State University.

In my recent key to Neopasites, this species would run near A.
calliopsidis Linsley but it is larger, more robust, with the integu-
ment duller and more coarsely and densely punctate, the fifth
abdominal sternite more lobate on each side, the disk more densely
punctate and hairy, ‘ and the apical margin broadly, shallowly
emarginate. It further differs in the more extensive white pubes-
cent patches of the head and thorax, the base of the labrum
clothed with white hair, a slightly different pubescent pattern on
the abdominal tergites, and the absence of white pubescence from
the basal abdominal declivity. It is possibly the most beautifully
spotted of the known species.

Neopasites knulli Linsley, new species

Male: Form slender; color black, antennas, mouthparts, legs, and tegulae
brownish or piceous, abdomen red; integument densely, coarsely punctate,
moderately dull. Head very densely punctate, with a large patch of ap-
pressed white hairs around the antennal bases which is denser along the mid-
line where it extends upward nearly to median ocellus, and a large white
patch behind each eye, the two connected posteriorly along posterior margin
of head; antennae brown, distal two-thirds of flagellum reddish, first flagellar
segment about as long as following two together; upper frons and vertex
coarsely, contiguously and subcontiguously punctate, the punctures of ocell-
ocular area a little larger but not quite so dense as those of disk of meso-

Sei>t., 1944]

Linsley: Neopasites

279

scutum; elypeus densely but more finely punctate; labrum with a few long,
erect white hairs at base, surface finely punctate, densely so at base, longi-
tudinally carinate for about one-half its length; mandibles reddish, base and
apex broadly piceous; ventral surface of head shining, area on each side of
gular cavity shining, irregularly punctate, punctures averaging more than
one puncture width apart. Thorax coarsely, very closely punctate; pronotal
collar densely white pubescent, tubercles margined with white; tegulae pice-
ous, margins rufo-testaeeous ; mesose-utum densely, contiguously and more
or less rugosely punctate, anterior and lateral margins and anterior half of
median line clothed with traces of white hairs along lateral and posterior
margins ; metanotum with a white patch on each side ; mesepisterna coarsely,
closely, somewhat rugosely punctate, vertical face broadly margined with
white. Wings very lightly infuscated, veins and stigma light brown. Legs
piceous ; anterior and intermediate femora apically white pubescent beneath ;
anterior and intermediate tibiae thinly, posterior tibiae more densely, white
pubescent externally. Abdomen red; first tergite with a patch of white on
each side bordering lateral margin of basal concavity, the patches separated
by nearly twice their width, basal concavity without white pubescence, ter-
gites two to five with a transverse band of white along basal margin extend-
ing from each side of middle to lateral margin, the pair on tergite two sepa-
rated by about half the distance separating the patches on tergite ohe, those
of each succeeding segment separated by about half the distance separating
those of the preceding tergite, tergites two to four with a small lateral white
patch on apical margin, fifth tergite with an apical fringe of white pubes-
cence, disk faintly clouded with piceous, sixth tergite more distinctly clouded
with piceous; pygidial plate ligulate, at least twice as long as broad; ster-
nites coarsely punctate, apical margins fringed with white pubescence, last
sternite with apical process truncate.

Length 5 mm.

Holotype male (collection of Ohio State University) from Cul-
berson County, Texas, August 30, 1940, collected by D. J. and
J. N. Knull.

This species is superficially very much like Neopasites ( Odon -
topasites) arizonicus Linsley in size, form, and coloration, but
may be distinguished by the naked eyes, less densely punctate
ventral surface of the head, especially on each side of gular
cavity, the short labral carina, and the darker legs and mouth-
parts. It appears to be more closely related to N. (N. ) minimus
Linsley, but the latter species is much smaller (3.5 mm.), with
only a few scattered punctures on the under side of the head,
and the labrum is not carinate.

280

Journal New York Entomological Society

[Vol. LII

Neopasites robertsoni Crawford

Among material recently submitted by Prof. 0. A. Stevens for
study were examples of this species from Hatton, North Dakota,
August 3 (Stevens).

Neopasites stevensi (Crawford)

This species was also taken by Prof. Stevens at Hatton, North
Dakota, on August 3, visiting flowers of Grindelia squarrosa.

Neopasites heliopsis (Robertson)

N. heliopsis was recently recorded1 from Winnecock, North
Dakota. The locality should have read Winnecook, Montana.
The record was based on a specimen in the collection of P. H.
Timberlake.

Neopasites calliopsidis Linsley

This species has been previously recorded from Iowa, Kansas
and Montana. Mr. Timberlake has submitted specimens from
the following localities in Colorado : Boulder County, July 6,
1925 (C. H. Hicks) ; Boulder, June 26 and 28, 1939, on Chry-
sopsis (Timberlake) ; Boulder, 2 miles north, June 25, 1939, on
Chrysopsis (Timberlake) ; and Cuchara Camps, Spanish Peaks,
elevation 8000 ft., July 4, 1939, on Aster (Timberlake). In these
specimens the average extent of the black areas on the abdomen
is greater than in the type series (Iowa), and some males also
have the basal white spots of the tergites confluent on each side.

Sept., 1944]

Weiss: Death-Feints

281

THE DEATH-FEINTS OF ALOBATES PENN-
SYLVANIA DeG., AND ALOBATES
BARBATA ENOCH.

By Harry B. Weiss

Hibernating specimens of both species of darkling beetles were
tested during the last week of April, 1944, in order to determine
the duration of their death-feints. All specimens were kept at a
room temperature of 72° F., several days before testing in order
to fully bring them out of hibernation. Death-feints occurred
when they were picked up or when they were pressed gently,
ventrally or when dropped through a distance of six or twelve
inches. Some beetles required a lot of handling and others very
little in order to bring on the death-feint. The ventral surface
of the thorax appeared to be the most sensitive area. When the
stimulus was applied to the dorsal surface the death-feint did not
occur. It was impossible to apply the stimulus with equal force
each time when it was done by hand, and it is not known if there
is any connection between the force of the stimulus and the dura-
tion of the reaction. However, it does not seem likely that there
is, in view of the fact that a gentle stimulus was just as liable to
promote a long or short death-feint as a hard stimulus. Although
different degrees of pressure and different amounts of handling
were required to initiate the death-feints, both the sensitivity of
the individual and the duration of its reaction probably depend
upon the variable organization of its nervous and motor
mechanism.

Alo~bates pennsylvanica DeG.

Ten specimens of this species were induced to feign death at
Fahrenheit temperatures of 72° and 82°. The durations of the
death-feints are shown in the following table. Some beetles
rested dorsally and others ventrally during the death-feint, but
neither position appeared to affect the duration of the reaction.

Successive death-feints in the same beetle became progressively,
irregularly shorter. One beetle refused to react after the eight-

282

Journal New York Entomological Society

[Vol. LII

Duration of Death-feint

Beetle

At 72° F. April 19

At 82° F. April 26

Seconds

Seconds

A

778

394

B

266

105

C

62

24

D

138

262

E

247

520

F

43

230

Gr

129

173

H

1,095

160

I

35

620

J

1,851

380

Average

464.4

268.8

eenth successive death-feint; another after the sixth, and many
of them after the third or fourth.

Alobates barbata Knoch.

The durations of the death-feints of ten specimens of this
species at Fahrenheit temperatures of 72° and 84° are shown
as follows :

Duration of Death-feint

Beetle At 72° F. April 20 At 72° F. April 25 At 84° F. April 25

Seconds

Seconds

Seconds

A

163

25

48

B

165

306

35

C

6

406

86

D

30

40

180

E

227

213

15

F

414

152

265

G

340

85

154

H

97

41

92

I

76

56

7

J

315

475

20

Average

183.6

179.9

90.0

Successive death-feints in the same beetle of this species were
generally similar to those reported for Alobates pennsylvanica.

Both species reacted alike except for the fact that the death-
feints of A. pennsylvanica endured longer than those of A.
barbata. When the temperature was increased 10 or 12 degrees
the average duration of the reaction declined approximately
one-half, for both species.

Sept., 1944]

Weiss: Death-Feints

283

Apparently there is a variation in the sensitivity of different
individuals, to the external stimuli initiating the death-feint and
also in the period of recovery. These variations are probably due
to differences in the quality of the labile compounds in the
receptive and conductive parts of the nervous system and in the
contractile muscle tissue. The destructive chemical action in-
volved in the reaction to the stimulus and the restoration of the
discomposed substances require different periods of time for dif-
ferent individuals. A high temperature apparently hastens
recovery. It would be of interest to know if internal stimuli
play any part in the restorative processes.

INSECT FOOD HABIT RATIOS OF NEW YORK STATE

By Harry B. Weiss

At various times during the past 15 years I have wondered if
a food habit classification of the species- of insects recorded from
New York would show ratios that differed materially from those
of other sections such as New Jersey, Connecticut, etc. In order
to satisfy my curiosity, I finally classified according to their
family food habits 15,343 of the 15,449 species recorded in “A
List of the Insects of New York,” M. D. Leonard, Editor-in-Chief,
that was published January, 1928, as Memoir 101 of the Cornell
University Agricultural Experiment Station. The Anoplura,
Mallophaga and Siphonaptera were omitted because of their non-
relation to vegetation or to other insects, likewise a few other
species, difficult to classify. Considering the large number of
species involved, these omissions are relatively unimportant.

The difficulty of classifying families of insects in accordance
with the food habits of their members is fully appreciated and
the weaknesses of such a classification have been admitted in a
former paper.1 The terms saprophagous, phytophagous, etc., are
used in their broadest sense and I am aware that such conclusions
as may be drawn from food habit classifications are broad general-
izations.

1 Insect Food Habit Ratios of North Carolina and Mount Desert Island,
Maine. Jour. N. Y. Ent. Soc., vol. 47, p. 155-157, June 1939.

284

Journal New York Entomological Society

[Vol. LII

The following table shows the distribution ratios of food habit
types for New York and, for comparative purposes, the ratios for
five other large areas are included. Five of the six areas are
large and embrace a variety of vegetation. Under such condi-
tions one would not expect the distribution ratios of the types of
food habits to vary widely and it will be noted that the New York
ratios are not unlike those for other areas.

No.

Phyto-

Sapro-

Harpacto- Para-

Pollen

feeders,

etc.

species

phagous

phagous

phagous sitic

Per cent Per cent Per cent Per cent Per cent

Western Arctic
Coast, N. A.

New Jersey

Connecticut

North Carolina
Mount Desert
Island, Me. ...

New York

400

47

27

14

10

2

10,500

49

19

16

12

4

6,781

52

19

16

10

3

9,249

46

17

22

11

4

5,177

52

17

14

15

2

15,343

48

23

17

10

2

In former papers it was brought out that in relatively small
areas, each with a uniform type of vegetation, the ratios of the
types of food habits, based on the species present, varied in
accordance with the type of vegetation, if the numerical ratios
between the species and the factors tending to reduce their num-
bers are considered as constant. In large areas such as are indi-
cated in the table, the ratios appear to vary but little.

Although the food classifications in the table are broad general-
izations of ‘‘apparent” food, comparatively little has been added
to our knowledge of “actual” food, since B. P. Uvarov called
attention in 1928 (“Insect Nutrition and Metabolism,” Trans.
Ent. Soc. London, Dec. 31, 1928), to the limited number of cases
where the true food of insects was actually known.

r.

Vol. LII No. 4

DECEMBER, 1944

Journal

of the

New York Entomological Society

Devoted to Entomology in General

Edited by HARRY B, WEISS

Publication Committee

HARRY B. WEISS JOHN D. SHERMAN, Jr.

T. C. SCHNEIRLA

Subscription $3.00 per Year

Published Quarterly by the Society
N. QUEEN ST. AND McGOVERN AVE.

LANCASTER, PA.

NEW YORK, N. Y.

1944

CONTENTS

The Structure of Living Insect Nerves and Nerve Sheaths
as Deduced from the Optical Properties

By A. Glenn Richards, Jr 285

Notes on the Behavior of Burying Beetles (Nicrophorus
spp.)

By Lorus J. Milne and Margery J. Milne 311

Thecla Burdi Kaye, A Synonym

By E. Irving Huntington 328

A New Species of Lambdina and Notes on Two Species
of Besma (Lepidoptera, Geometridae, Ennominae)

By Laurence R. Rupert 329

Psylliidae from Tropical and Semitropical America
(Homoptera)

By John S. Caldwell 335

Notes on Mexican Butterflies, IV

By F. Martin Brown 343

A Revision of the North American Genus Eremomyioides
Malloch (Diptera, Muscidae)

By II. C. I Iuckett 361

Records and Descriptions of Neotropical Crane-Flies
(Tipulidae, Diptera), XVIII

By Charles P. Alexander 369

A Correction 384

The Status and Functions of the International Commis-
sion on Zoological Nomenclature and the Present State
of Its Work

By Karl Jordan 385

Members of the New York Entomological Society 389

NOTICE: Volume LII, Number 3, of the Journal of the

New York Entomological Society was published on

September 11, 1944.

Entered as second class matter July 7, 1925, at the post office at Lancaster, Pa.,
under the Act of August 24, 1912.

Acceptance for mailing at special rate of postage provided for in Section 1103.

Act of October 3, 1917, authorized March 27, 1924.

JOURNAL

OF THE

New York Entomological Society

Vol. LII December, 1944 No. 4

THE STRUCTURE OF LIVING INSECT NERVES AND
NERVE SHEATHS AS DEDUCED FROM
THE OPTICAL PROPERTIES* 1

By A. Glenn Richards, Jr.

Zoological Laboratory, University of Pennsylvania

In a previous issue of this Journal relatively indirect histo-
chemical data were presented and were interpreted as indicating
the presence of lipid nerve sheaths in insects (Richards, 1943).
These sheaths, like those of so-called non-myelinated nerve fibers
of vertebrates, are extremely thin and are very labile under the
influence of fixation techniques. Optical studies with polarized
light permit the study of living nerves and so obviate the diffi-
culties attendant upon histological fixation. They also give far
greater sensitivity. The present paper presents data on the
ultrastructure of normal living insect nerves and nerve cords as
deduced from optical data.2 The presence of lipo-protein nerve
sheaths in insects is confirmed and amplified.

The studies reported herein were performed on the nervous
system of adult American cockroaches, Periplaneta americana,
and on larvae of the house mosquito, Culex pipiens. Most of the
studies were made on intact nerve cords and peripheral nerves

1 The work described in this paper was done under a contract, recom-
mended by the Committee on Medical Research, between the Office of Scien-
tific Research and Development and the University of Pennsylvania.

2 Thanks are due to Dr. R. S. Bear and Dr. F. O. Schmitt, of the Massa-
chusetts Institute of Technology, for assisting the author with the use and
interpretation of polarized light data, and to Dr. H. B. Steinbach, of Wash-
ington University, for the loan of a Kohler compensator.

286

Journal New York Entomological Society

[VOL. LII

but the data so obtained agree with studies on teased, single-fiber
preparations of the larger nerves of cockroaches. Observations
were also made on single specimens of an assortment of other
arthropods to see if similar optical properties occur for nerves
throughout the phylum Arthropoda.

THE USE OF POLARIZED LIGHT

From the viewpoint of optics a nerve is composed of optically
inactive components and several types of optically active com-
ponents. In studies on nerves polarized light is used to deter-
mine (in living or fixed nerves) the presence and status of the
oriented optically active components. One can in a sense frac-
tionate an intact nerve, and with a knowledge of the optical
activities of the extractable components determine where the
various components are located and what their orientation is in
a living nerve. In experimental studies polarized light can be
used to look for degradations of either orientation or chemical
structure. One is dealing, then, with a kind of cytological in-
spection but with a method of inspection that is extremely sensi-
tive when used for nerve studies.

No detailed treatment of the use and interpretation of polarized
light data can be given here. Some of the terms to be used in
this paper may be mentioned in relatively non-technical language
but for an understanding of the complex phenomena involved
reference must be made to the extensive treatises listed below.3

An ordinary beam of light is considered as a series of wave
motions in which the light rays vibrate in all planes 'perpendicular
to the direction of propagation. Plane polarized light is light in
which the vibrations are all in the same plane4 (Fig. 1). As

3 Descriptions of the instruments and general phenomena can be found in
Rogers & Kerr (1933) and other textbooks on mineralogy and optics, espe-
cially Ambronn & Frey (1926). Excellent diagrams of the optical phe-
nomena involved are given by Myers (1938). The most lucid elementary
presentation for beginners is perhaps that of Cheshire (1932). For the
interpretation of biological data see W. J. Schmidt (1937) and Frey-
Wyssling (1938). A more recent and comprehensive analysis of the optical
phenomena shown by nerve sheaths is given by F. O. Schmitt & R. S. Bear
(1939).

4 Actually a light wave consists of electrical and magnetic vectors which
vibrate in mutually perpendicular planes perpendicular to the direction of

Dec., 1944]

Richards: Insect Nerves

287

such, plane polarized light is analogous to parallel waves on water,
where the waves vibrate np and down in one plane while traveling
forward. With polarized light one studies the birefringence
(= double refraction) of a material. This is accomplished by
observing the specimen between crossed Nicol prisms or other

Fig. 1. Diagram of the effect of polarizing blocks set with the polarizing
planes at a 90° angle. The axes of the polarizing blocks is indicated diagram-
matically by parallel lines. The planes in which the light rays vibrate are
sketched in for clarity of three-dimensional visualization.

A light beam traveling along the dotted line in the direction a — > b is
composed of waves vibrating in all axes perpendicular to the direction of
propagation (four of these planes are drawn in this diagram). On passing
through the polarizer A only waves vibrating in the plane of the paper are
transmitted. On reaching the polarizer B, set at a 90° angle, these are just
the waves that are completely extinguished, and therefore no light is trans-
mitted beyond B*

polarizing materials,* * 5 i.e., one prism is below the object and trans-
mits only light that is polarized in one particular plane, the other
prism is above the object and being set at a right angle to the

propagation, but in polarized light studies it is customary to designate the

vibration plane of the electrical vector as the plane of polarization.

5 For a good diagram of how Nicol prisms function see Myers (1938),
figure 84. The same result is obtained by a different principle in ‘ ‘ Polaroid ’ ’
sheets, etc.

* For clarity of diagrammatic presentation the author has taken some
liberties with optics in Figures 1 and 2. Actually, four waves traveling in
phase along one line would combine into a single wave of intermediate posi-
tion and the sum of the four in height. For illustrating the point in question
the method used here seems clearest even though technically different vibra-
tion planes should be drawn along different lines parallel to a-b.

288

Journal New York Entomological Society

[Vol. LII

first eliminates all of the light except that which has had its plane
of polarization changed by passing through the specimen (Figs.
1-2). The ability to change the plane of polarization at certain
orientations is a property of birefringent materials. Substances
which show birefringence are said to be anisotropic, substances
which do not are called isotropic. Even with anisotropic sub-
stances, however, there is one direction along which light (ordi-
nary or polarized) is unaltered; this is termed the optic axis of
the substance. The optic axis may be called isotropic, all other
axes are anisotropic. The anisotropic axes have the property of
changing the plane of polarization by splitting any transmitted
beam of light into two rays6 which vibrate in mutually perpen-
dicular planes (Fig. 2) and travel with different velocities.7
The optical effects can be completely described diagrammatically
in terms of a direction of vibration for faster (or slower) trans-
mission (see below) and two refractive indices, since the refrac-
tive index is an inverse statement of the relative speed of light.
Substances with only one refractive index are called isotropic,
substances with two different refractive indices are called uni-
axially anisotropic, and substances with different refractive in-
dices for all three vibration planes of rays perpendicular to the
optic axis are called biaxially anisotropic. Most biological aniso-
tropic substances, including those dealt with in this paper, are
uniaxial.

Several kinds of birefringence are shown by biological mate-
rials. In addition to uniaxial and biaxial anisotropy, birefrin-
gence may be either positive or negative with respect to a particu-
lar direction. These are arbitrary terms applied to differentiate
between materials in which the index of refraction is greater

6 Commonly called the ordinary and extraordinary rays.

7 As Cheshire (1932) points out, birefringent substances include fibrous
and crystalline structures where the orientations of the components are such
that the structure is different in the longitudinal and transverse directions.
In a certain sense the structure is analogous to the grain of wood, and, as
Tyndall showed, the speed of heat and sound waves is different for waves
traveling along the long axis of a piece of wood and for waves traveling
perpendicular to this. With birefringent materials and polarized light we
have similar differences in speeds in different directions with the additional
complication that both the ray direction and the vibration direction are con-
cerned.

Dec., 1944]

Richards: Insect Nerves

289

for the vibration in the stated direction than for the other vibra-
tion perpendicular to it ( positive birefringence) , and materials
in which the relative velocities are the reverse ( negative birefrin-
gence). The direction for faster (or slower) vibration is deter-
mined by comparing the effect with a known standard. Com-
monly one takes advantage of interference effects for this purpose
and uses a gypsum plate which makes the entire field appear red.
Any effect of the specimen must be either additive or subtractive,
and reference to Newton’s series of colors shows which it is.
Knowing the orientation of the various parts of the system one

Fig. 2. Same as Figure 1 but with a sheet of birefringent material (the
specimen, C ) inserted between the polarizers with its axis at a 45° angle to
each. The description is the same as for Figure 1 except that the specimen,
C, oriented at a 45° angle splits the plane polarized beam into two mutually
perpendicular waves vibrating at a 45° angle to the beam from A to C, and
therefore at a 45° angle to polarizer B. The waves do not reach B at a 90°
angle to the vibration direction, and therefore light is transmitted. Adding
to these figures the parts of a compound microscope will give a crude diagram
of the optics of a polarizing microscope of the orthoscope type.

can usually tell by clear-cut color changes (from red to blue or
yellow) whether the substance is positively or negatively bire-
fringent.

The terms positive and negative birefringence, however, can
be used in either of two ways: 1) When the orientation of the
optic axes of the anisotropic substance is known, then the material
may itself be called either positive or negative depending on
whether the index for vibrations parallel to the optic axis is
greater or less, respectively, than the index for vibrations per-
pendicular to this axis. 2) Whether the orientation of the optic

290

Journal New York Entomological Society

[Vol. LII

axes is known or not, one can refer to an anatomical structure as
positive or negative with respect to some distinguishing direction.
In many cases in biology the optic axis of the oriented birefrin-
gent material coincides with the long axis of the gross structure
(e.g., muscle, tendon, axis cylinder of nerves, etc.). Thus a
muscle fiber is positively uniaxial in relation to both its long axis
and the optic axes of the anisotropic micelles. In the case of the
nerve sheath, however, the anisotropic micelles are oriented with
their optic axes radial (perpendicular to the long axis of the
nerve). Accordingly, the nerve sheath can be called either
positive or negative depending on whether the reference direction
is the optic axes of the anisotropic particles or the long axis of
the nerve. This can be confusing unless one is careful to note
the reference direction, especially as whichever reference direc-
tion is chosen some nerve sheaths are positive and some negative
in relation to it. In biological literature reference has usually
been made to the anatomically long dimension, and this custom
is followed in the present paper; however in the recent review
by Schmitt & Bear (1939) these authors have shifted to employ-
ing the optic axes as reference directions.

Birefringence may be produced by either the internal structure
of a molecule or its asymmetrical shape. The first is called
intrinsic birefringence, the second is called form birefringence.
Distinguishing between these two types is quite useful in analyz-
ing the source of the observed birefringence. Fortunately this is
readily accomplished. Form birefringence is due to the presence
of oriented submicroscopic rodlets or plates called micelles • im-
mersion of these micelles in a medium of the same refractive
index abolishes their birefringence but has no effect on any in-
trinsic birefringence that may be present. By immersion in
media of various refractive indices one can then determine not
only whether the material possesses any intrinsic birefringence
but more importantly whether there is present a second birefrin-
gent material that is normally masked by the form birefringence
of the first material. Thus, in a so-called non-myelinated nerve
one first observes a birefringence that is positive in relation to
the long axis of the nerve. If such a nerve is immersed in a salt
solution containing glycerine, the sign of birefringence is re-
versed. But if the nerve is first extracted with alcohol the sign

Dec., 1944]

Richards: Insect Nerves

291

cannot be reversed. This is interpreted as due to the positive
form birefringence of the oriented protein normally masking the
negative intrinsic birefringence of the lipids (Schmitt & Bear,
1937).

In treating birefringence quantitatively, certain values can be
measured and others calculated from these measurements. Prop-
erly oriented anisotropic substances split a beam into two rays
which, vibrating in different planes, travel through the specimen
at different velocities. Obviously, if one of the component rays
travels more slowly, it will lag behind and emerge from the
specimen a certain finite distance behind the faster ray.8 This
difference is called the retardation (= phase difference, = ampli-
tude, 4>). With a Kohler compensator the retardation is calcu-
lated from the angle between the compensator plate and the
specimen necessary to compensate for (abolish) the retardation
of the specimen :

where m is the known maximum retardation of the compensator
plate, A the wave length of light (551 mq is taken as the “center
of gravity” of white light), and 3i and (32 the measured angles
using the Kohler method and the Bear & Schmitt modification
respectively (Bear & Schmitt, 1936b). The sign of <p will depend
on whether 3i and j32 are additive or subtractive angles and will
agree with the qualitative determination of the birefringence as
positive or negative in character.

Obviously, the amount the slower ray is behind the faster ray
will depend on both the difference in velocity and the thickness
of the specimen being measured. The difference between the two
velocities, called the magnitude of birefringence, is the funda-
mental property we are interested in obtaining. With sheets of
known composition and orientation the magnitude of birefrin-
gence is readily calculated from :

8 This actually transforms the plane polarized beam of light into an ellip-
tically or circularly polarized beam. However, for our purposes we can con-
tinue to treat it as though we were dealing with two plane polarized beams
vibrating perpendicular to one another.

<p = - sin 2(3i, or
<J> = - 2mA sin 232,

(1)

(2)

(3)

292

Journal New York Entomological Society

[Vol. LII

where nt and n2 are the two refractive indices, <p the retardation,
and d the thickness of the specimen.

In nerve studies it is relatively easy to determine the retarda-
tion (<j>) but sometimes difficult to obtain the true effective dis-
tance ( d ). This is partly due to the heterogeneous nature of
nerves, and in the case of nerve sheaths partly to the complica-
tion introduced by the optic axes of the sheath micelles being
oriented radially. Schmitt & Bear (1937) have derived the fol-
lowing equation to allow for the radial orientation of the sheath
micelles :

where n 1 and n2 are the two refractive indices, (32 the compensa-
tion angle determined by the Bear & Schmitt method, dx the axon
diameter or its equivalent, and d2 the axis cylinder diameter or
its equivalent.

For entire insect nerve cords none of these methods for deter-
mining the magnitude of birefringence seems satisfactory. Due
to the low amplitude it has not been found feasible to attempt
quantitative measurements on teased single nerve fibers of in-
sects except for extracted nerve sheaths and the axis cylinder of
stretched nerves. Equation (1) has been used for the points
plotted in Figure 39 but equations (3) and (4) have been applied
to estimate the more fundamental birefringence values for single
fiber preparations and for the neural lamella surrounding the
nerve cord.

For the work reported in this paper a Leitz polarizing micro-
scope was used, employing orthoscopic methods, white light,
various magnifications and several types of compensators. Quan-
titative measurements were made with a Kohler A/20 rotating
mica plate compensator (Leitz Wetzlar No. 2335) using both the
Kohler and the Bear & Schmitt methods (Bear & Schmitt, 1936b).

THE OPTICAL EFFECTS SHOWN BY INSECT NERVES
1. General

Until recently insect nerves have been considered non-myelin-

9 So far as graphs such as figure 3 are concerned, the same curve would be
obtained from using the angle ((3), the retardation (0) and the magnitude
of birefringence ( nx-n2 ).

9.46 sin 2(32

Dec., 1944]

Richards: Insect Nerves

293

ated (Richards, 1943). In many insects ( e.g ., mosqnito larvae)
single nerves are extremely small (less than 2 q in diameter)
but in the large cockroaches some single fibers attain a diameter
of 10 (j and a few are 20-30 q in diameter. The thicknesses of
the nerve sheaths are apparently not more than a few per cent
of the fiber diameter. One can recognize the sheath and estimate
its thickness around single nerve fibers of the cockroach, but the
sheath around nerves of mosquito larvae is beyond resolution with
ordinary microscopical methods.

It is relatively easy to tease out and identify for study single
fibers from a cockroach nerve cord. Many observations can be
made satisfactorily, however, on intact nerve cords containing
many nerves in one bundle. The intact interganglionic abdomi-
nal connectives in the adult American cockroach have a diameter
of 175-280 jj, while those of fourth (last) instar mosquito larvae
are only 13-24 q in diameter.

Around the outside of the intact nerve cord there is a secreted
sheet of material which Scharrer (1939) has termed the neural
lamella. This sheet is also birefringent but does not intefere
with optical analyses of the included nerves. It will be discussed
in a separate section.

In analyzing nerves, they are first examined in a salt solution,
with or without previous fixation, then they may be examined in
glycerine or other media of high refractive index to mask the
form birefringence and so determine what intrinsic birefringence
is present. In insect nerves such immersion normally results in
a reversal of the sign of birefringence, indicating that in a normal
nerve we are dealing with a balance between birefringent mate-
rials of opposite sign (curve 2, Fig. 3). The lipids may be re-
moved by appropriate solvents {e.g., alcohol) ; comparison of the
optical properties before and after extraction permit both show-
ing and localizing the presence of the optically active lipids.
Since the form birefringence of the proteins normally predomi-
nates in non-myelinated nerves, it is ordinarily sufficient to ex-
amine a preparation in salt solution and then in a dilute solution
of glycerine in saline solution. For some special purposes and
for evaluating the method other media, solvents and variations
of technique may be used.

294

Journal New York Entomological Society

[Yol. LII

2. Analysis of the Birefringent Properties of the
Axis Cylinder

Intact, living ventral nerve cords and larger peripheral nerves
of American cockroach adults and mosquito larvae immersed in
saline solutions10 show a slight birefringence which is usually
positive in relation to the long axis of the nerve fibers (Fig. 14) d1
The birefringence of the axis cylinder12 of living cockroach
nerves in salt solution or of fixed cockroach nerves after lipid
extraction is relatively diffuse, is positive in relation to the nerve
axis and seems to be fairly homogeneous throughout single giant
nerve fibers. In the normal relaxed condition the amplitude
of birefringence (4)) due to the axis cylinder is very small but
it can be increased greatly by tension. Cockroach nerve cords
fixed in formol-saline solutions in a relaxed condition, then teased
to give single-fiber preparations, extracted with alcohol (to re-
move masking lipids) and examined in water, show bright posi-
tively birefringent lines for the sheaths but only faint positive
birefringence for the axis cylinder (Fig. 5). Similar results
are obtained for the axis cylinder with living cockroach nerve
cords teased and examined in salt solutions. In contrast, cock-
roach nerve cords fixed in formol-saline solutions in a stretched
condition (1-J times the relaxed length) and then teased and
treated as above show strong positive birefringence throughout
single fiber preparations (Figs. 8-10). Measurements of the am-
plitude of birefringence show that the birefringence of the
sheaths is affected relatively little (perhaps not at all) by the
stretching but that the birefringence of the axis cylinder is
greatly increased. Immersion in glycerine practically abolishes
the birefringence of the axis cylinder showing that this is due
mainly tb anisodiametric micelles exhibiting form birefringence.

10 For cockroach adults the salt solution used contained NaCl 10.93, KC1
1.57, CaCl2 0.85, MgCl2 0.17 and NaHC03 0.17 grams per liter; for mosquito
larvae NaCl 7.8, KC1 0.62, CaCl2 0.40 and NaHCOs 0.17 grams per liter.

11 Occasionally a normal relaxed nerve cord is found to be negative in
saline. Such nerve cords can be reversed to positive by stretching. Natu-
rally, immersion in glycerine-saline does not reverse the sign of birefringence
of such a nerve cord; it only increases the negativity. See curve 3, Figure 3.

12 The axis cylinder is the nerve axon exclusive of its surrounding mem-
brane and sheath.

Dec., 1944]

Richards: Insect Nerves

295

Whatever the nature of the tension effect may be,13 the fact re-
mains that the axis cylinder of insect nerves contains anisotropic
micelles, presumably protein, with at least a predominant orienta-
tion in the direction of the long axis of the nerve (Fig. 4).

The studies on single-fiber preparations from cockroaches are
in agreement with observations on intact nerve cords and periph-
eral nerves. In intact nerve cords it is easy to measure the total
birefringence but the superposition of many fibers makes it diffi-
cult to determine what components of the birefringence are due
to the axis cylinder and what to the nerve sheaths. In general, a
longitudinally striated appearance to the birefringence of intact
nerve cords can be interpreted as indicating nerve sheath com-
ponents whereas the more homogeneous birefringence seen in
stretched nerve cords can be interpreted as being due to the sum-
mation of sheaths and stretched axis cylinder effects.

An analysis of the form factor involved in this birefringence
is graphed in Figure 3. Since the measurements were all made
with intact nerve cords these curves, of necessity, show a sum-
mation of axis cylinder and sheath effects. Curve 1 shows the
change in amplitude with change in refractive index of the im-
mersion media for relaxed nerve cords from which the lipids have
been removed by extraction with alcohol and ether. This curve
shows that most of the birefringence of lipid-extracted nerves is
due to micellar form, the micelles having a refractive index be-
tween 1.56 and 1.60 when determined in this manner. A small
residue of intrinsic birefringence is probably also present since
the value never falls quite to zero.14 Comparison of these data

13 There are several conceivable explanations of the increase of birefrin-
gence of the axis cylinder on stretching. Mihalik (1937) has made a similar
report for vertebrate nerves. The phenomenon is being studied further. It
is presented here only because the birefringence of the axis cylinder of
insect nerves can be seen most clearly in stretched nerves.

14 This is the interpretation accepted by Schmitt and Bear. The Ambronn
immersion technique commonly encounters difficulties with biological mate-
rials. Theoretically curve 1 should be symmetrical. Correction for the
shrinkage that occurs in media of higher refractive index is not sufficient to
make these curves symmetrical. Such slight asymmetry is seen in various
published curves ( e.g ., Chinn & Schmitt, 1937, Fischer, 1944). Different
methods of fixation give roughly parallel curves of considerably different
height (Bear, Schmitt & Young, 1937b). Curve 1 and 2 of the present paper

296

Journal New York Entomological Society

[Yol. LII

with the data from single-fiber preparations suggests that the
form birefringence plotted in curve 1 is due to both the axis
cylinders and the nerve sheaths but how much is contributed by
each of these components has not yet been determined. Curve 2
shows the change in amplitude of a formalin-fixed nerve cord
immersed in media of increasing refractive index with reversal
of sign at approximately 1.343. Curve 2 is typical for normal
cockroach nerves. Theoretically one would expect this curve to
parallel curve 1 if a series of non-lipid-solvent immersion media
of higher refractive index were available. Curve 3 gives data
from a similar immersion series for one of the occasional nerve
cords which is already negative in saline. Great variation, such
as is shown by the difference between curves 2 and 3, occurs be-
tween different preparations. This seems, at least in part, due to
tension, and will be discussed in a subsequent paper.

It does not seem feasible to attempt estimating the true mag-
nitude of birefringence (nx — n2) from measurements on entire
nerve cords. Estimates from formalin-fixed, alcohol-extracted,
single-fiber preparations of the cockroach using equations (1) and
(3) indicate that the magnitude of birefringence of the axis

and similar curves by Chinn (1938) though roughly parallel are not so far
apart as one would expect. More serious is the great variation of curves for
some materials depending on the immersion series employed. Castle (1936)
has treated in some detail the radically different types of curves shown by
chitin in different series of immersion media where imbibition and possibly
adsorption alter the picture and complicate interpretation. The use of dif-
ferent series of immersion media has given only slight differences for nerve
sheaths (Chinn & Schmitt, 1937) but a much lower value for the refractive
index is assumed from glycerine -immersion series in the recent paper by
Werndle & Taylor (1943). The complex phenomena involved in these unex-
pected variations are not understood. Castle (1936) feels that the situation
is so complex that it defies explanation at present, and that the immersion
method is not valid for the determination of the refractive index of certain
materials such as chitin. The curves from nerves and nerve sheaths by
various methods and media are, however, sufficiently consistent to be reason-
ably certain that '(1) insect nerves are similar to other nerves, and (2) that
the protein component shows largely a form birefringence and that it is of
opposite sign from the lipid birefringence. Under the circumstances it
seems that the best proof of the lipo-protein nature of the nerve sheaths comes
from the results of extraction experiments rather than from the immersion
curves (metatropic effect).

Dec., 1944]

Richards: Insect Nerves

297

•cylinder ranges from nearly zero (immeasurably low) in relaxed
nerves to approximately 0.0018 for stretched nerves.12 The sev-
eral possible reasons for low magnitudes in axis cylinders are dis-
cussed by Bear, Schmitt & Young (1937b) and Richards, Stein-
bach & Anderson (1943).

Fig. 3. Data from immersion experiments with cockroach nerve cords.
Entire interganglionic connectives were used; the measurements accordingly
represent the summation of effects shown by a large bundle of nerves. Curve
1 gives the averages from measurements on six nerve cords fixed in 95%
ethyl alcohol and subsequently measured in media ranging from 1.331 to 1.659
in refractive index,* with corrections being made for differences in size and
for shrinkage in some of the media. Curve 2 gives the readings from a
formalin-fixed nerve cord immersed in distilled water and a series of increas-
ing concentrations of glycerine, range 1.333 to 1.435 in refractive index.
This curve closely approximates the typical picture for fresh nerves in saline,
with reversal of sign occurring at a glycerine concentration between 10 and
15%. Curve 3 gives measurements from one of the few formalin-fixed nerves
which was found to be negative in water.

* The immersion media used were: methyl alcohol, ethyl alcohol, 1-nitro-
propane, methylcyclohexane, chloroform, carbon tetrachloride, benzene, di-
methyl phthalate, methyl salicylate, anethole, aniline, bromoform, carbon
bisulfide and alpha-bromo -naphthalene.

298

Journal New York Entomological Society

[Vol. LII

The nerve cords of mosquito larvae are much smaller than those
of cockroaches, and the included nerves are all very small ( < 1 to
2 (j ) . All studies on mosquito nerves were made with intact nerve
cords since satisfactory single-fiber preparations were not ob-
tained. The intact living nerve cord in appropriate salt solution,
like that of the cockroach, is positively birefringent in relation to
its long axis. In the relaxed condition (Fig. 14) it shows a more
or less striated birefringence which, as noted above, is interpreted
as indicating the presence of a relatively strongly birefringent
sheath component. In the stretched condition the birefringence
is stronger and nearly homogeneous. By comparison with cock-
roach nerves, this is interpreted as indicating an increase of the
axis cylinder component of birefringence. The data from mos-
quito nerve cords are consistent with the idea that they have
qualitatively the same configuration as is found in cockroach
nerves, but they are very small and the amplitude of birefrin-
gence is very low.15

There is no reason to think that the ultrastructure of the axis
cylinder of insect nerves is different from that of nerves of other
animals. The amplitude of birefringence of relaxed insect nerves
is unusually low but this may possibly be reconciled by further
study. Otherwise, the picture recorded above agrees well with
reports by various workers on various nerves (see W. J. Schmidt,,
1937). The birefringence of the axis cylinder is due to aniso-
tropic protein micelles showing chiefly a form birefringence
(anisodiametric particles). These micelles are oriented or at
least predominantly oriented with their optic axes parallel to
the long axis of the nerve since the axis cylinder is isotropic in
cross section. Their birefringence is positive both with respect
to their optic axes and to the nerve axis (Fig. 4). As already
reported with other nerves, there is a slight reduction in the
amplitude of birefringence on histological fixation. There is a
rapid decay or loss of birefringence of the axis cylinder follow-

15 The phase retardation (0) of an entire, relaxed, 18 (a, interganglionic
connective of a mosquito larva in saline as calculated from equation (1) is
of the order of only 2-3 mp. In a stretched nerve cord this may rise to
6-8 mp. These values compare favorably with values given for cultured
Corethra nerves by Pfeiffer (1943).

Dec., 1944]

Richards: Insect Nerves

299

ing mechanical injury, the action of certain venoms and insecti-
cides, and attending post-mortem degeneration.16

3. Analysis of the Birefringent Properties of the
Nerve Sheaths

Insect nerve sheaths are relatively thin. In the largest nerves
of the cockroach they attain a thickness of only a little more than
one micron, i.e., they are only a few per cent of the fiber diameter.
In smaller nerves they seem to be proportionately thinner, and in
the smallest cockroach nerves and in all nerves of mosquito larvae
they are too thin for direct observation (submicroscopic thick-
ness) . Due to the thinness of these sheaths it is difficult to deter-
mine for certain whether or not they are produced by distinct
sheath cells. However, in longitudinal sections of insect nerve
cords one can find a few nuclei between the nerve fibers.17 Bear,
Schmitt & Young (1937a) concluded that nuclei found around
squid giant nerve fibers probably represented cells analogous to
the Schwann cells of vertebrate nerves. Possibly the nuclei seen
in insect nerve cords likewise represent sheath cells analogous
to Schwann cells.

These thin nerve sheaths are metatropic, i.e., they exhibit a
weak positive birefringence (in relation to the nerve axis) which
is readily reversed to negative by immersion in media of slightly
higher refractive index (curve 2, Fig. 3). These data can be
interpreted in the same manner that similar data from other ani-
mal groups have been interpreted by Schmitt & Bear (1937,
1939). The individual nerve sheaths are composed of bound
layers of protein and lipid, both of which are birefringent but
of opposite sign. The protein normally dominates slightly with
its form birefringence. Masking the form birefringence of the
protein with glycerine permits the intrinsic lipid birefringence to
be seen. Removal of the lipids with appropriate solvents ( e.g .,
alcohol) increases the positive birefringence and makes it impos-
sible to reverse the sign of birefringence with glycerine. Since
two oppositely birefringent compounds can be thus demonstrated

16 A presentation of some of the degenerative effects due to venoms and
insecticides will be given in subsequent papers.

17 Other than those associated with tracheae.

300

Journal New York Entomological Society

[Vol. LII

and identified (Figs. 5 vs. 11, 14 vs. 15), it follows that normally
both are present in the sheath, and that the normal sheath bire-
fringence is a picture of how much one component (in this case
protein) predominates in birefringence over the other.

The axis cylinder viewed from the side is birefringent through-
out. The nerve sheaths, however, show birefringence only at or
very near to the edges of the nerve. This is due to the manner
of orientation of the optic axes of the anisotropic micelles. In all
birefringent materials and structures there is' one axis, the optic

Fig. 4. A diagrammatic representation of the orientation of the birefrin-
gent micelles in a nerve and its sheath. The long axes of the rodlets in
these diagrams is used to indicate the orientation of the optic axes of the
micelles. On the left is a hemi-section of a nerve showing the micelles of
the axis cylinder oriented longitudinally while those of the sheath are
oriented radially. On the right is an enlarged sketch showing the arrange-
ment of bound lipid and protein micelles in concentric layers as suggested by
Schmitt, Bear & Palmer (1941).

axis, along which the structure is isotropic. In the nerve sheaths
both the protein and lipid molecules must be oriented with their
optic axes arranged radially (Fig. 4). With such an arrange-
ment, the central part of the sheath should appear isotropic, as
it does, since the optic axes of the micelles lie in the axis of the
microscope, whereas the edges should be strongly birefringent, as
they are. This is confirmed by observation of negative polariza-

Dec., 1944]

Richards: Insect Nerves

301

tion crosses observed in cross sections of alcohol-extracted giant
cockroach nerves (compare Chinn & Schmitt, 1937).

The magnitude of birefringence (Wx-Wa) for the sheath can
be calculated approximately from measurements on single giant
fibers of cockroaches. Fresh nerves in saline show only extremely
faint sheath birefringence. This indicates that the balance of
birefringence from the protein and lipid (opposite signs) is
nearly equal, and the magnitude of birefringence of the combina-
tion is therefore nearly zero. Alcohol-extracted nerves (Figs.
5-7) should show only the protein component. The average of
measurements on eleven alcohol-extracted nerves ranging from
6 to 10 p in diameter using equation (4) is the extremely low
value of nt - n2 = 0.0016. This represents the value for sheath
protein ; the lipid value must be similar but slightly lower.

The nerve cords of mosquito larvae show the metatropic reversal
as well as cockroach nerves, but the small size of the fibers makes
it difficult to show positively that the reversal is in the sheath.
Presumably the situation is the same as in cockroaches except for
the small size and low amplitude (Figs. 14—15).

The protein component of the individual nerve sheaths of in-
sects may be collagenous as in other animals (see e.g., Schmitt,
Hall & Jakus, 1942). Direct proof of this was not obtained but
alcohol-extracted single-fiber preparations from cockroaches ob-
served during treatment with dilute acetic acid showed a reduc-
tion and eventual loss of sheath birefringence. This would be
expected for collagenous sheaths.

The lipid component of the individual nerve sheaths of insects
seems almost certainly to include phosphatids as in other animals
(Schmitt, Bear & Palmer, 1941). The solubilities of the sheath
components are consistent with the idea that they include phos-
phatids. Also it is possible to extract considerable quantities of
phosphatids and other myelin constituents from bee brains (un-
published data by Dumm, Patterson & Kichards).

Metatropic nerve sheaths are found on axons throughout the
insect. They are best studied from interganglionic connectives
and large peripheral nerves {e.g., cereal nerve) but they can be
demonstrated throughout ganglia and in small distal peripheral
nerves. Likely these sheaths extend over the nerve cell bodies

302

Journal New York Entomological Society

[Yol. LII

themselves but this was not studied. Chinn (1938) reported
that a sheath continuous with and similar to the axon sheath
extends over the nerve cells of lobsters, crayfish and leeches.

The metatropic nerve sheaths of cockroaches and mosquito
larvae exhibit numerous of the properties already described for
other nerves. Birefringence is quickly lost following mechanical
injury. The metatropic effect is lost on air-drying. Absolute
ethyl alcohol removes the lipid quickly but weak alcohol removes
it slowly with the production of birefringent particles that are
stainable with Sudan dyes (Richards, 1943) . Other lipid solvents
except acetone also quickly block the metatropic effect (phospha-
tids are relatively insoluble in accetone) . The metatropic reversal
is most readily demonstrated by immersion in glycerine solutions
but it can also be produced by other solutions of high refractive
index (e.g., sucrose) and is independent of shrinkage. The
amplitude is somewhat reduced by fixation with formalin, etc.,
but after fixation with non-lipid solvents the lipids are more
strongly bound and difficult to remove (see Mezzino, 1931).
Likely there is also a relation between fiber diameter and ampli-
tude of birefringence ; certainly the larger nerves of cockroaches
show more birefringence than the small nerves (see Schmitt &
Bear, 1939, Fig. 1).

Cockroach nerve cords degenerating in saline show first a
decrease in amplitude and then loss of the birefringence of the
axis cylinder, then a gradual decrease and loss of the lipid com-
ponent of birefringence. Nerve cords degenerating in glycerine-
saline solutions, however, give rise to sphaerites which gradually
decrease in size and eventually disappear (Fig. 16). These bire-
fringent sphaerites originate from the sheath lipids, they can be
produced by weak alcohol or degeneration in glycerine but have
not been found to occur in cockroach nerve cords degenerating
in saline (compare Baldi, 1929). The lipid component can also
be abolished by the action of cobra venom which is known to con-
tain phosphatase enzymes. They are also affected by certain
insecticides ; these data will be treated in subsequent papers.

Insect nerve sheaths differ quantitatively from other nerves
in the low order of birefringence and especially in the very close
balance of positive and negative components. The balance of

Dec., 1944]

Richards: Insect Nerves

303

these components seems closer in insect nerve sheaths than in
other recorded metatropic nerves. Only 10-15 per cent glycerine
is needed to effect reversal of sign in the insects studied, whereas
30 per cent glycerine is required for Crustacea, 35-40 per cent
for squid giant fibers, 50 per cent for non-myelinated spinal
ganglion cells of the frog, and 70 per cent for leech nerves. In
all of these the refractive index of the anisotropic micelles is
reported to lie in the range 1.56 to 1.60 ; accordingly this varia-
tion in necessary refractive index of the immersion media likely
indicates roughly the ratio of protein birefringence to lipid
birefringence in the various nerves.

METATROPIC NERVE SHEATHS IN OTHER ARTHROPODS

A hasty survey was made covering a number of scattered
arthropods to see if the type of nerve sheath studied in the cock-
roach and mosquito is to be found throughout the Annelid-
Arthropod complex. All species examined exhibited metatropic
nerves. Species examined by the author were an unidentified
spider, a centipede ( Scutigera sp.), a millipede ( Fontaria sp.),
a caterpillar ( Lymantria dispar ), a beetle ( Scarites subterranus)
the honey bee (Apis mellifica) and also a marine polychaete worm
(Nereis virens). Other authors have already reported on the
earthworm (Lumbricus sp.), the medicinal leech (Hirudo medi-
cinalis), Limulus and various Crustacea (prawn, shrimp, crayfish,
crab, lobster).

With all the major groups of the Annelid- Arthropod complex
except the Onychophora represented in the above list, it certainly
seems probable that Schmitt & Bear were correct in suggesting
that metatropic nerve sheaths would be found throughout these
phyla.

THE NEURAL LAMELLA

Around the outside of the nerve cord and of peripheral nerves
is a homogeneous sheet secreted by an underlying layer of non-
nervous cells. Scharrer (1939) studied this rather extensively
in cockroaches and termed the cell layer the “ perineurium’ ’ and
the secreted sheet the 4 * neural lamella.” She pointed out that
the neural lamella is optically homogeneous and stains with dyes
like the connective tissue of vertebrates. It is several microns

304

Journal New York Entomological Society

[Vol. LII

thick in cockroaches but is too thin to measure in mosquito larvae.
It seems to be present in all insects but is commonly very thin
and inconspicuous.

The neural lamella is strongly birefringent. It shows as a
bright line along the edge of all nerve cords and is positively
birefringent in relation to the long axis of the nerve cord (Fig.
12). 18 In cross sections it shows a strong negative polarization
cross. Immersion experiments show that its birefringence is
almost entirely due to form birefringence, but attempts to deter-
mine the refractive index by immersion methods encountered the
same sort of difficulty recorded by Castle (1936) for chitin. Im-
bibition or oriented imbedding in some cases gives quite different
measurements for media of the same refractive index. The re-
fractive index seems most likely in the neighborhood of 1.47, and
clearly is different from both chitin and the sheaths (collagen ?)
around individual nerves. Although the neural lamella shows as
a bright line due to its depth, the true magnitude of its birefrin-
gence is quite low. Using equation (4), nx-n2 is found to be
approximately 0.00022.

The neural lamella is unaffected by extraction with lipid sol-
vents. It is completely and readily dissolved by strong alkali,
and accordingly is not chitinous. It gives a strong protein reac-
tion (xanthoproteic test) but seems not to be collagen since it
does not swell, dissolve or even lose its birefringence in dilute
acetic acid (3 days) and since immersion experiments give differ-
ent results for the neural lamella and the presumably collagenous
sheaths around individual nerves. Serial sections show that the
neural lamella is composed of concentric thin layers. Attempts
to obtain electron micrographs showed only that the neural
lamella fractures in an irregular manner suggesting a non-fibrous
structure. Stretching experiments show that the neural lamella
is elastic and possesses strong, photoelastic properties.

is The neural lamella does not interfere seriously with a study of the
birefringence of the included nerves. It shows only as a bright line along
the edge, and can be ignored. Proof that it does not interfere with readings
made on the included nerve bundle comes from experiments in which the
same reading was obtained for the nerve bundle before and after manual
removal of the neural lamella.

Dec., 1944]

Richards: Insect Nerves

305

The above data suggest that the neural lamella of cockroaches
is a series of elastic, homogeneous, concentric sheets composed of
anisodiametrie protein micelles exhibiting form birefringence and
arranged with their optic axes perpendicular to the surface of
the sheet. The negatively uniaxial protein micelles are arranged
with their optic axes at right angles to the nerve axis, and accord-
ingly the neural lamella appears positively birefringent in rela-
tion to the axis of the nerve cord.

No attempt has yet been made to study accurately the per-
meability of the neural lamella but obviously it must be per-
meable. Exchanges with the blood must take place through it,
fixing fluids used in histology penetrate rapidly, and even the
rather large aggregates of solubilized detergents — Black Sudan
B (a polyazo dye) penetrate the neural lamella more rapidly than
they will dialyze through a collodion membrane.

The birefringence of the neural lamella of a wasp and a beetle
has already been described and figured by W. J. Schmidt (1937,
p. 273, Fig. 70), but seemingly Schmidt erroneously thought that
this sheet compared with the sheaths of single vertebrate nerves.

DISCUSSION

The data presented in this paper supplement those given pre-
viously (Richards, 1943), and show that the structure of insect
nerves and nerve sheaths is closely similar to the structure re-
ported for other animals (see Schmitt & Bear’s review, 1939).
This structure is diagrammed in Figure 4. Around the outside
of insect nerves and nerve cords is a secreted sheet, the neural
lamella, which while serving the same protective purpose is dif-
ferent from the protective coatings around vertebrate nerves and
nerve cords.

Few observations on the birefringence of insect nerves are to
be found in the literature. Bruno (1931) reported that insect
nerves (various species) are isotropic. This bare statement can-
not be evaluated in the absence of any information on the method
of preparation and the type of compensator employed. Perhaps
the negative report is simply an expression of failure to recognize
the extremely low amplitude of relaxed insect nerves. W. J.
Schmidt (1937) has already refuted Bruno’s report. He pub-

306

Journal New York Entomological Society

[Vol. LII

lished a figure showing the birefringence of a fresh peripheral
nerve from the head of a wasp (Fig. 70, p. 273) and described
similar results obtained with the ventral nerve cord of a beetle.
From his brief comments, however, it is not certain how clearly
he distinguished the various components of the birefringence.
Certainly the sheath to which he refers is the neural lamella ; he
seems not to have seen the sheaths of the individual nerves or to
have determined their lipo-protein character. Recently Pfeiffer
(1943) has studied the growth of single nerves from Corethra
larvse (Diptera) in tissue culture. He records a positive uni-
axial birefringence for normal Corethra nerves, with an ampli-
tude of birefringence comparable to that described herein for
mosquito larvae. Pfeiffer was interested in studying nerve
growth ; he does not mention nerve sheaths and seems not to have
distinguished between sheath and axis cylinder effects or to have
recognized the presence of metatropic sheaths.19

Numerous references can be found in entomological literature
stating that insect nerves are non-myelinated. These statements
are readily understandable in view of the extreme thinness of the
sheaths (commonly submicroscopic) and the correspondingly low
lipid content.

SUMMARY

1. Insect nerves and nerve cords in saline solutions show a
positive uniaxial birefringence in relation to the length of the
fiber. This is reversed to negative by immersion in media of
higher refractive index, i.e., the nerves are metatropic. Qualita-
tively they seem to agree well with the structure of other so-called
non-myelinated nerves as recorded by Schmitt, Bear and others.
A diagram presenting the orientation of the optically active
micelles is given in Figure 4.

2. The axis cylinder of insect nerves shows largely form bire-
fringence which is positive in relation to both the nerve axis and
the optic axes of the micelles. Most of this birefringence is due

T9 The extremely small size of these nerves may have caused Pfeiffer to
overlook the sheath. In nerves of comparable size from mosquito larvse the
sheaths are Of submicroscopic thickness. Another possibility is that insect
nerves growing in tissue culture may possibly differ from nerves growing in
an animal.

Dec., 1944]

Richards : Insect Nerves

307

to micellar form but a small residue of intrinsic birefringence
seems to be present. The amplitude of birefringence of the axis
cylinder increases greatly on being stretched.

3. Each insect nerve is surrounded by a discrete sheath of lipo-
protein. In thickness these sheaths are at most only a few per
cent of the fiber diameter, and accordingly in the case of small
nerves are too thin to be observed directly (submicroscopic).
This sheath is responsible for the metatropic reversal effect.
Both the lipid and protein components are oriented with the
optic axes of their micelles arranged radially. They are of
opposite sign. Normally the nerve sheaths appear positively
birefringent in relation to the nerve axis because the positive
component due to the proteins slightly overbalances the negative
component due to the lipids. Removal of the intrinsically bire-
fringent lipid increases the positive birefringence and prevents
metatropic reversal. Masking the form birefringence of the
protein permits the intrinsic birefringence of the lipids to be
seen (metatropic reversal).

The protein component of the sheath may be collagen. Its
micelles are negatively birefringent in relation to their optic
axes but being arranged radially make the nerve appear posi-
tively birefringent in relation to its length. Its birefringence
is mostly due to micellar shape but a small residue of intrinsic
birefringence seems to be present.

The lipid component of the sheath is probably a mixture of
phosphatids with perhaps other “myelin” components. The
lipid micelles are positively birefringent in relation to their optic
axes but being arranged radially make the nerve appear nega-
tively birefringent in relation to its length. The lipid birefrin-
gence is intrinsic since it is unaffected by the refractve index of
immersion media.

4. Insect nerves seem to differ from the non-myelinated nerves
of other animals chiefly in the low order of amplitude of birefrin-
gence and the extreme thinness of the nerve sheaths. These
seeming differences may possibly be illusory since ordinarily only
large nerves are studied whereas insect nerves are commonly very
small.

5. Metatropic nerve sheaths have been reported to date in an
earthworm, sandworm and leech (Annelida), a spider and Limit-

308

Journal New York Entomological Society

[Vol. LII

lus (Arachnida), various shrimps, crabs, etc. (Crustacea), a
millipede (Diplopoda), a centipede (Chilopoda) and five orders
of pterybote insects (cockroach, mosquito, moth, bee, beetle). It
certainly seems probable that at least for the Annelid- Arthropod
complex Schmitt & Bear were correct in suggesting that meta-
tropic nerve sheaths will be found throughout the invertebrates.

6. Around the nerve cord and peripheral nerves is a secreted
permeable elastic layer, the neural lamella, which is structurally
and chemically different from the protective coatings around verte-
brate nerves. The neural lamella is or at least contains a protein
which shows form birefringence and photoelastic properties.
This protein seems to be different from collagen and different
from the protein of the sheaths around individual nerves. Im-
mersion experiments are complicated by the oriented imbedding
of imbibed media. The neural lamella is composed of thin con-
centric layers with the micelles arranged perpendicular to the
surface (radial in relation to the nerve). These micelles, like
sheath proteins, are negatively uniaxial in relation to their optic
axes; being arranged radially they make the sheet appear as a
positively birefringent line (in relation to the nerve axis) along
the edge of the nerve.

LITERATURE CITED

Baldi, F. 1929. Richerche sulla degenerazione Walleriana. Prime in-
dagini a luce polarizzata.- Riv. Neurol., 2 : 56-63.

Bear, R. S. and F. O. Schmitt. 1936. The optics of nerve myelin. Jour.
Optical Soc. Amer., 26: 206-212.

. 1936. The measurement of small retardations with the polarizing

microscope. Jour. Optical Soc. Amer., 26: 363-364.

Bear, R. S., F. O. Schmitt and J. Z. Young. 1937. The sheath components
of the giant nerve fibers of the squid. Proc. Roy. Soc. London, B,
123: 496-504.

. 1937. The ultrastructure of nerve axoplasm. Ibid., 123 : 505-519.

. 1937. Investigation of the protein constituents of nerve axoplasm.

Ibid., 123 : 520-529.

Bruno, G. 1931. Sulla struttura dei nervi di aleuni insetti. Monit. Zool.
Ital., 42: 8-15.

Castle, E. S. 1936. The double refraction of chitin. Jour. Gen. Phys., 19:
797-805.

Cheshire, F. J. 1932. The polarization of light. Jour. Television Soc.,
ser. 2, 1: 201-207.

Dec., 1944]

Richards: Insect Nerves

309

Chinn, P. 1938. Poralization optical studies of the structure of nerve cells.
Jour. Cell. Comp. Phys., 12 : 1-21.

and F. O. Schmitt. 1937. On the birefringence of nerve sheaths

as studied in cross section. Jour. Cell. Comp. Phys., 9: 289-296.
Fischer, E. 1944. The birefringence of striated and smooth mammalian
muscles. Jour. Cell. Comp. Phys., 23 : 113-130.

Mezzino, L. 1931. Rifiessioni ed osservazioni istologiche sulle fibre nervose
midollate. Riv. Biol., 13 : 31-56.

Mihalik, P. 1934. Mechanisch-experimentelle Untersuchungen fiber die
Doppelbrechung der Markhaltigen Nervenfaser. Zts. Zellforsch.
mikr. Anat., 21: 653-656.

Myers, L. M. 1938. Television Optics. 2nd Ed. 362 pp. Pitman & Sons,
London.

Pfeiffer, H. H. 1943. Polarizationsmikroskopische Messungen an gerichtet
gespannten Mutterstficken von Ganglienkulturen in vitro. Natur-
wiss., 31 : 47-48.

Richards, A. G. Jr. 1943. Lipid nerve sheaths in insects and their probable
relation to insecticide action. Jour. N. Y. Ent. Soc., 51: 55-69.

, H. B. Steinbach and T. F. Anderson. 1943. Electron microscope

studies of squid giant nerve axoplasm. Jour. Cell. Comp. Phys.,
21: 129-143.

Scharrer, B. 1939. The differentiation between neuroglia and connective
tissue sheath in the cockroach (Periplaneta americana). Jour.
Comp. Neurol., 70: 77-88.

Schmidt, W. J. 1937. Die Doppelbrechung von Karyoplasma, Zytoplasma
und Metaplasma. Protoplasma Monographien, 11: 1-388.

Schmitt, F. O. and R. S. Bear. 1937. The optical properties of vertebrate
nerve axons as related to fiber size. J our. Cell. Comp. Phys., 9 :
261-273.

. 1939. The ultrastructure of the nerve axon sheath. Biol. Rev., 14:

27-80.

and K. J. Palmer. 1941. X-ray diffraction studies on the nerve

myelin sheath. Jour. Cell. Comp. Phys., 18: 31-42.

, C. E. Hall and M. A. Jakus. 1942. Electron microscope investi-
gations of the structure of collagen. Jour. Cell. Comp. Phys., 20:
11-33.

Werndle, L. and G. W. Taylor. 1943. Sheath birefringence as related to
fiber size and conduction velocity of catfish Mauthner, Mfiller and
peripheral fibers. Jour. Cell. Comp. Phys., 21: 281-293.

PLATE X

In all figures the cross-hairs indicate the orientation of the Nicol prisms
of the microscope. Extra contrast and compensation obtained with a Kohler
V20 rotating mica plate compensator. Photographs, of course, do not dis-
tinguish between positive and negative birefringence; this is determined by
compensators and stated in text and legends.

310

Journal New York Entomological Society

[Vol. LII

Fig. 5. Teased single large nerve fiber from cockroach nerve cord fixed
in 95% ethyl alcohol in relaxed condition. Lipids extracted by alcohol. In
distilled water; birefringence positive in relation to nerve axis. The parallel
light lines represent the protein component (collagen ?) of the nerve sheath.
Diameter of this fiber 16 p,. Magnification 300 X.

Fig. 6. A group of medium-sized nerves from the same preparation as
Figure 5. Diameters of these fibers 5-6 p. Magnification 300 x.

Fig. 7. A single small fiber, 3 p in diameter, from same preparation. Note
how faint the sheath is. Magnification 300 x.

Fig. 8. A single teased cockroach nerve fiber which was fixed in 95%
alcohol while stretched to approximately li x its relaxed length. Note that
the sheath and axis cylinder are of the same general intensity and so cannot
be distinguished. Specimen in distilled water ; birefringence positive in rela-
tion to nerve axis. Diameter of this fiber after stretching 10 p. Compen-
sator set for maximum brightness of nerve. Magnification 300 X.

Fig. 9. Same but with compensator plate rotated for extinction of nerve
retardation. Such brightening and extinguishing effects prove we are deal-
ing with birefringent properties.

Fig. 10. Another nerve from the same preparation. Diameter 12 p. Mag-
nification 300 x.

Fig. 11. Large group of nerve fibers from partly teased cockroach nerve
cord in saline containing 15% glycerine (2 hrs.). The form birefringence
of the protein is here masked sufficiently to give reversal of sign (metatropic
effect) ; the sign of birefringence here is then negative in relation to the
nerve axis. The many parallel light lines are due to the lipid components of
a number of nerve sheaths. Nerves of various diameters (3-8 p on plate
but somewhat shrunken by the glycerine medium). Magnification 500 X.

Fig. 12. A thoracic interganglionic connective of a cockroach in saline
with 15% glycerine (15 min.). The bright lines along the sides are pro-
duced by the neural lamella. The bright interior is produced by the super-
imposed effects of the lipid sheaths of many nerves. The neural lamella
is positive in relation to the nerve cord axis; the included bundle of nerves
is negative. Compensator set for maximum brightness. Magnification 45 X.

Fig. 13. Same but with compensator plate rotated for extinction of nerve
retardation. In this photomicrograph the positive birefringence of the
neural lamella is lost against the white background.

Fig. 14. Abdominal ganglion and interganglionic connectives of a mos-
quito larva. Fresh preparation in saline; birefringence positive in relation
to nerve axes. Compensator set for maximum brightness. Magnification
200 X.

Fig. 15. Same fresh preparation after 5 min. in saline containing 15%
glycerine. The sign has reversed and the nerve cord shrunken slightly.
Magnification 200 x.

Fig. 16. Portions of two interganglionic connectives of a cockroach after
18^ hours degeneration in saline containing 15% glycerine. Shows the break-
down of the sheath lipids into birefringent spheerites. The two bright lines
running diagonally through the center are produced by the neural lamellae
of the two connectives. The wavy bright line in the upper right quadrant
is produced by an air-filled trachea. Magnification 200 X.

(Jour. N. Y. Ent. Soc.), Vol. LII

(Plate X)

Dec., 1944]

Milne: Behavior

311

NOTES ON THE BEHAVIOR OF BURYING BEETLES
(NICROPHORUS SPP.)

By Lorus J. Milne and Margery J. Milne
University of Pennsylvania and Beaver College

For several summers, ending with the season of 1928, the senior
author had observed species of Nicrophorus ( Coleoptera, Staphy-
linidae, Silphinae) burying small carcasses at Irondale (Halibur-
ton County), Ontario, Canada, in July. Some of his observa-
tions were published (Milne, 1928). Much more extensive and
detailed studies of burying beetles were made by Pukowski (1933)
in Europe and Leech (1935) in British Columbia. These authors
followed the life history from the time of burial, while the adults
cared for the young, through the three instars of larval life, the
prepupal and pupal periods, and made observations on emergence
and feeding of the adults. Difficulty was reported in observing
the burying behavior because of its nocturnal nature and because
Nicrophorus was not abundant enough to justify the risks of dis-
turbing beetles at work. Since the burying activities had been
watched repeatedly in daylight at Irondale, the present authors
made a return trip there in the summer of 1944, to prepare a
Kodachrome motion picture of the burying beetles in their work
above ground. Although the time available during the brief
vacation from war research greatly restricted the experimental
studies, it was possible to make further observations which extend
and clarify the behavior of these insects.

1. SPECIES INVOLVED AT IRONDALE

Six species of Nicrophorus have been collected at carrion in
this region by the authors, namely N. sayi Lap., N. orloicollis
Say, N. marginatus Fab., N. pustulatus Hersch., N. vespilloides
Hbst., and N. tomentosus Web. All these are easily distin-
guished in the field. Only tomentosus has the pronotum covered
with yellow, appressed hair (which often becomes abraded on the
two convexities of the disc). Only sayi and marginatus have
distinctly curved hind tibiae. Only vespilloides and tomentosus

312

Journal New York Entomological Society

[Vol. LII

have the antennal clubs entirely black, the other species having
at least part of the club orange red ; these two species are also
considerably smaller in average size. Sayi and orbicollis are
very similar in markings, with the elytra mostly black, but with
orange red in a sub-basal, irregular crossbar and a subapical spot.
Marginatus, vespilloides and tomentosus are similar in markings,
the elytra being mostly orange red, the bands of color continuing
across both elytra, sometimes confluent also along the outer mar-
gins. Pustulatus is unique among these six species in having
the pronotum transversely oval, the others having a circular disc ;
it also has the sub-basal elytral spot small or absent, the subapical
bar often divided in two (sometimes lacking), the insect thus
having an even blacker facies than sayi or orbicollis. Of these,
only tomentosus and orbicollis * were numerous in late July of
1944, and on them most of the following notes are based.

2. METHODS OF STUDY

Irondale is a hilly region from which the original timber was
removed perhaps fifty years ago. Those areas which could be
freed of glacial erratics and which were reasonably level, have
been farmed (chiefly for grain). Other areas have been cleared
for pasture and kept available for sheep and cattle. Less level
land has been allowed to grow up again. The vegetation is typi-
cal Canadian zone, with hemlock, spruce, pine, paper birch, pop-
lar, spiraea, sweet. fern, sweet gale, mountain laurel, club mosses,
bracken, and an abundant lichen flora on exposed rocks and tree
trunks. Among the larger fauna are loons, whip-poor-wills, por-
cupines, groundhogs, skunks, red squirrels and chipmunks.
Most of the glacial lakes contribute to the Burnt River system,
deriving the name from the dark color of the water, due to leach-
ing of iron ore from rock substratum and to solution of decaying
coniferous and other xerophytic debris. Sphagnum is common
and pitcher plants and sundew reach large size.

To attract Nicrophorus, small dead animals were placed at
selected positions in a variety of ecological habitats. Each car-
cass was secured to a nearby stake by a two-foot length of fine steel
wire, to make recovery easy (c/. Milne, 1928). The animals used

* Referred to as N. pollinctor in Milne, 1928.

Dec., 1944]

Milne: Behavior

313

were chiefly deer mice (Peromyscus) , with some house mice
(Mus), shrews (Blarina), birds (robins, thrushes, sparrows,
wrens, domestic turkeys, etc., as found dead) and snakes. In
previous years larger carrion was tried, including groundhog
(Marmota) , skunk, dog, etc., but Nicrophorus was found to show
little interest in carcasses too large for them to bury. For spe-
cies observed at Irondale, a body the size of a robin is perhaps
the limit.

In open fields, such as pastures or where hay had been cut,
N. tomentosus and marginatus arrived to bury mice and snakes.
Competition with ants was frequent and the carcasses deterio-
rated considerably before the beetles could get them buried. In
leaf litter from birch and poplar, second growth woodland, N.
tomentosus and orbicollis were quick to bury mice and birds. In
coniferous duff the same species were somewhat slower in finding
carrion. No activity was observed on mice placed in low, wet
positions such as sedgy swales. Mice placed on particularly
hard ground or on bare rock, were transported by the beetles to
places where the soil was less packed and burial hence easier.
Desiccation of carcasses by the sun did not have any noticeable
effect on the interest in them shown by the beetles.

3. BEHAVIOK OF THE BEETLES

Nicrophorus exhibits a number of interesting behavior pat-
terns. The beetles fly to the general vicinity of the carrion, ap-
parently by smell (c/. Abbott, 1927a & 1927b; Milne, 1928).
Tomentosus is particularly accurate in locating the body before
alighting, buzzing through the bushes like a bumblebee. The
resemblance to Bombus is enhanced by the golden body hair, the
yellow inner surfaces of the elytra (which are held back to back
over the midline) and the creamy cast to the flying wings. All
Nicrophorus run about briskly, forcing their way through or
under tangled vegetation. If disturbed, all but tomentosus are
likely to either feign death, or run away a few feet to hide in
grass roots. Tomentosus takes to its wings and may alight in a
nearby bush, often standing on a slanting stem rather than the
broader surface of a leaf. Pukowski (1933) describes a similar
habit as part of the behavior of a lone Nicrophorus attracting a

314

Journal New York Entomological Society

[Yol. lii

mate, the beetle climbing a stone or plant, elevating the abdomen
obliquely and extending it so much that the coriae are visible.
No indication of such was observed by the present authors, a lone
Nicrophorus regularly getting to work on the burial task and con-
tinuing so engaged until a mate arrived.

After a beetle has arrived at a dead animal, it characteristi-
cally examines the body with palpi and antennae and tests the size
of the carcass by trying to move it. The “carrying” action is
one of the most typical behavior patterns shown loy Nicrophorus.
Another activity closely related is that of “exploring” the sur-
rounding soil for a suitable spot in which the body may be in-
terred. A third procedure is to test the looseness of the soil by
“plowing” it. All of these behaviors are energetic, and are
shown by even a single Nicrophorus.

The carrying action of Nicrophorus demonstrates the strength
of the beetles and the vigor with which they proceed wtih their
task. To move a carcass, say forward in terms of a mouse’s body,
the beetle crawls under the head of the mouse, turns over on its
back, and lifts the mouse bodily over itself. The mouse inches
forward little by little, while the beetle slides slowly on its smooth
back under the mouse until it reaches the posterior end. There
it either emerges and runs around to the anterior end of the
mouse again, or turns over on its feet to crawl under the mouse,
the smooth dorsum of the beetle not disturbing the body. Such
effort is very strenuous, and a rest period or an exploring inter-
lude usually follows a few minutes of moving the carrion. A
single beetle rests more than when a mate has arrived. A slight
tug at the carcass usually stimulates it to renewed activity.
When two beetles are operating one or the other is usually carry-
ing the body while the mate explores or plows. No sexual differ-
ence in degree of activity could be noticed (c/. Wood, 1873;
Furneaux, 1893; Pukowski, 1933). Both members of a pair
were highly industrious under most circumstances. On level
ground devoid of major obstacles, a pair of beetles may transport
a full-grown mouse or shrew as rapidly as three or four feet per
hour, and keep up this pace for as much as two and a half hours,
the limit in every case observed being the distance necessary to
reach sufficiently soft ground for burial use.

Dec., 1944]

Milne: Behavior

315

The exploring behavior was very distressing to the observers.
After a shorter or longer period of work on a carcass, one or
both members of a pair will suddenly leave the body and run
away from it at the usual brisk pace. The beetles may go only
a few feet (seldom less), or as much as a few yards, and in some
cases ( tomentosus ) took to flight, only to return in a matter of
minutes. A specimen of N. tomentosus with a broken elytra! tip
was observed to fly off and back again four times within an hour.
Each time it flew out of sight. During the hour perhaps fifteen
minutes work was done in carrying the mouse and in plowing
in nearby soil, the remainder being either exploring the whole
surrounding area (perhaps fifteen feet in diameter), or resting
with head under a clump of grass, or absent altogether while
away on a flight. This procedure persisted even after the speci-
men was joined by agnate, continuing until the carrion had been
transported (mostly by the mate, in this instance), almost to the
area selected for burial use.

The plowing procedure loosens the earth. The beetle uses its
head as a plowshare or bulldozer, pressing into the earth perhaps
the depth of its stout body below the surface, then forcing its
way forward in an arc while maintaining its depth. The earth
is forced upward and crumbles. Any roots encountered are
either forced aside, or chewed through, but if numerous a new
burial site is sought out. Before a final area is considered satis-
factory, as many as twenty possible sites may be plowed and
found unsuitable. The final area may be many feet away from
the carrion, and the beetle (s) will alternately work on the car-
cass and run to the burial site to do another stretch of plowing.
The route taken between carrion and burial site is usually fairly
direct, and the body is carried along this line. How both mem-
bers of a pair can agree on a site was not determined, nor was
it at all clear how the beetles were able to keep the carcass moving
so uniformly (few exceptions) in one direction. The contrast
was very marked between the cooperation of Nicrophorus and the
great wastage of effort among ants trying to carry a large food
particle.

In a few instances, the beetle bait was placed on soft ground
and the Nicrophorus buried it where it lay. In all instances,

316 Journal New York Entomological Society [Vol. lii

however, the burial procedure was a localization of the plowing
action. The beetles plowed under the carrion, entering at one
side (or end) and emerging at the other, only to turn about and
complete the shuttling action. On each emergence at the side or
end of the carcass, a sizeable amount of earth was forced up, to
accumulate in a loose pile all around the body. Gradually the
earth from below the carrion was displaced to the side and the car-
cass settled into the ground. A continuation of the same activity
gradually drops the body below the surface of the ground, and the
procedure is continued until as much as two inches of earth are
over the carcass. The body is usually let into the ground at a
steep angle. Not only the earth under the carrion, but also that
within a few inches of it is plowed. Closer to the body the
ground is plowed to a greater depth. In its final site, a chamber
is cleared of earth below and to one side from the body. In the
chamber Nicrophorus remains after burial. The chamber may
be as much as two and a half inches long, an inch wide and deep,
and the walls are left packed in the original condition of the
earth, so that cave-ins are unlikely. Burial is usually completed
in five to eight hours, although the beetles will continue for days
if obstructions slow their work. Nicrophorus usually remain
with the carrion at least for many days (c/. Hatch, 1927b ; Milne,,
1928; Pukowski, 1933; Leech, 1935).

There seems to be a relationship between the size of the car-
rion, the size of the Nicrophorus species and the number of pairs
which can work in burying it. A full sized robin (in juvenile
plumage) was handled by a pair each of N. orbicollis and tomen-
tosus. A thirty-inch garter snake was buried by two pairs of
N. tomentosus and one of marginatus. In one instance a full-
grown deer mouse was interred by two pairs of A. tomentosus.
In all other instances only a single pair of a single species did
the work. Additional specimens of orbicollis arriving at a piece
of carrion where a pair were at work, usually crawled under the
body, only to emerge (promptly!) pursued by one of the original
pair. Several rough and tumble fights were witnessed in which
the newcomer resisted briefly the attack of the original 4 ‘ owner. ’ ’
Orbicollis coming to carrion where tomentosus was at work
usually drove the smaller species away. Tomentosus coming to

Dec., 1944]

Milne: Behavior

317

a carcass being carried or buried by orbicollis usually left with-
out any indication of expulsion. Tomentosus coming where
tomentosus were already at work sometimes helped awhile before
leaving, in only one instance (see above) remaining until the
body was completely interred. Thus tomentosus would seem to
make up in small size, greater numbers (see below) and more
sociable reactions for its inability to compete physically with the
larger species. Orbicollis becomes quite excited over expelling a
competitor and stridulates very audibly by rubbing the upper
surface of the abdomen against the under surface of the elytra
(cf. Morley, 1902). This same stridulation is observed when a
pair mate (3 to 4 seconds; cf. Pukowski, 1933), or when an ob-
struction continues to bar the way in moving a carcass. These
were the only occasions and the only species in which such noises
were noted. Pukowski (1933) indicates that pairs working on a
carcass keep in contact with each other by alternately emitting
“zirping” sounds.

Obstructions may be of a variety of types. Naturally encoun-
tered are situations where roots, stems, leaf petioles, etc., are suffi-
ciently anchored at both ends as to restrict an opening through
which the beetle is attempting to move the carrion. The beetle
discovering such an obstacle usually tries first to force it out of
the way, getting head and pronotum under the restraining strand,
feet in the vestiture of the carrion, and crawling ahead. Many
obstructions can be stretched sufficiently by this method to open
a suitable passage for further progress of the carcass. When
force proves inadequate, the beetle may settle down on the re-
straining piece and laboriously chew it through. Often periods
of chewing are alternated with episodes of drawing and pushing
at the carcass or strand, the chewing frequently having weakened
the obstruction sufficiently to let it give. When the obstacle,
proves too resistant to such treatment, soil is plowed from under
the carcass and the needed space obtained in this way.

Obstructions provided experimentally were usually very irri-
tating to the beetles but in no case did they leave permanently.
String tried to the leg or tail of a mouse was the commonest form
of restraint, and one which could be overcome by gnawing the
string until it broke under strain. Fine steel wire was an in-

318

Journal New York Entomological Society

[Vol. LII

superable difficulty. A carefully placed piece of rock, supported
partly on the ground and partly on the body of a mouse, proved
less of a problem. Although the rock weighed several pounds,
the pair of beetles working together were able to lift the rock
where it rested on the mouse sufficiently to push the carrion free
of its pressure and restraint.

One experimental situation gave a good demonstration of the
beetles’ behavior. A mouse laid out on fairly soft ground had a
string tied to one hind leg, the opposite end of the string being
fastened to a stake close to the mouse but in such a way that the
string was almost vertical and held the leg clear of the ground.
A pair of N. orbicollis proceeded to bury the mouse. They did a
fine job, the head of the mouse finally hanging almost vertically
downward into the hole the beetles had excavated. The hind leg
continued to be supported in its original position. The beetles
cleared away the earth until there was none below the head and
shoulders of the mouse for a distance equal to the thickness of a
beetle, and a space was also clear all around the mouse so that it
hung by its one leg over a sizeable, cup-shaped hole. With much
stridulation and rotation of the mouse around and around over
the hole, the beetles failed to go vertically upward to the support-
ing string. Every move in this direction ended in a trip out
the length of the mouse’s tail, to see that it was free. It was,
but the traffic became so heavy and the mouse so “ripe” that the
skin rolled off the tail distally like a glove finger. Finally the
beetles gnawed the tail off the mouse, severing it at the root.
Almost at once they found the supporting string, and after a
few abortive attempts to free the foot, one of the Nicrophorus
settled down to the task of gnawing it through. When the string
gave way, the mouse collapsed in a heap into the bottom of the
hole prepared for it, and burial was completed uneventfully
within half an hour.

Pukowski (1933) indicates that the beetles prepare a conical
hole below the carrion, always smaller than the body, and fold
the carcass as they draw it into the pit. The episode just de-
scribed would seem to indicate that Nicrophorus continue to
enlarge the hole for the carcass until it has been sunk to their
satisfaction, and that a conical hole and consequent folding may

Dec., 1944]

Milne: Behavior

319

be merely economy of effort. When carrion has considerable
length, as for example a snake, no folding was observed by the
present authors. The snake was let into the ground to a depth
of nearly two inches all along its length (except for the tail),
so that it was buried horizontally, in approximately the attitude
occupied previously on top of the earth. It is true that in the
case of snakes, burial advanced rapidly in the region from head
to anus, the tail being left out in the air for nearly twenty-four
hours after the remainder of the carcass was well under the sur-
face. The tail, with its smaller supply of food materials, was
obviously of much less interest to the beetles, forming chiefly an
obstruction to satisfactory burial. In several instances the last
few inches of a snake’s tail was chewed off, and the postanal
remainder pulled into the ground after putrefaction had rendered
it more plastic.

Heavy rain obstructed burying beetles much less than had been
expected. On several occasions Nicrophorus had begun to bury
bait placed on level, hard ground, when torrential downpours
drove the observers to nearby shelter. From the cabin the bait
could be seen almost or quite covered by water, sometimes to a
depth of an inch. No sign of the beetles was noted, yet within
fifteen minutes after the rain slackened enough for the ground
to drain off, the insects were busy in the wet earth, excavating,
plowing and tugging the carcass into their cavity.

The observers’ concern over the seeming desertion of the bait
whenever Nicrophorus went into an exploring episode, suggested
another experiment. As indicated above, the beetles frequently
explore a very sizeable area befor returning to the carcass. There
is no indication, however, that there is ever any difficulty in find-
ing the carcass again, since the beetles often return to it in an
almost straight line from a distance of a yard or two. On several
occasions, duplicate baits were tried, being mice of approximately
equal size and state of preservation, placed a foot apart on even
terrain. A Nicrophorus would come to one, test it for size, then
explore the surrounding ground for a suitable burial site. Dur-
ing the exploration the second mouse was usually discovered.
Almost without exception, the beetle examined the duplicate bait
without attempting to carry it, then hurried back to the mouse

320

Journal New York Entomological Society

[Vol. LII

previously discovered. In no case observed did the beetle (s)
desert the first bait in favor of the second. The same type of
experiment was tried on pairs which were busy burying a rela-
tively fresh mouse. A similar (or more odoriferous) mouse was
placed where they would surely find it during exploration trips.
In no case did the beetles desert the first-found carcass to more
than examine the second. It was quite obvious that the extra
carrion so nearby was a source of great distraction, but the recog-
nition of one body as distinct from another was most marked.

In only a few instances did Nicrophorus show any indication
of feeding on the carrion (cf. Furneaux, 1893 ; Lutz, 1921 ; Steele,
1927 ; Milne, 1928). Usually the beetles seemed to be in a hurry
to get the carcass interred. During daylight the need for rapid
burial was great, since blowflies came in considerable numbers,
laying living larvae if unmolested for a few minutes. The brisk
activities of the beetles and the frequent jerky movements of the
carcass have a deterrent effect on flies of some types (including
blowflies) but where obstacles prevented beetles from quickly get-
ting the carrion under ground during daylight hours, it was
obvious that little of the carcass would be available for other
than dipterous larvae. In many instances the beetles seemed to
realize this, and failed to complete burial. In some cases
Nicrophorus left fly-ridden carcasses sunk below the level of the
ground, covered by perhaps a quarter of an inch of loose earth.
A few days later such carrion was a squirming mass of fat fly
larvae. Rapidly buried bodies, on the other hand, are remark-
ably free of dipterous contamination. This may well be due to
the preference shown by beetles for operations in twilight, at
night or on cloudy (even rainy) days ( cf . Abbott, 1927b) or an
active eating of young maggots by the beetles (noted on a few
occasions; cf. Steele, 1927; Leech, 1935).

In one instance of Nicrophorus feeding, one member of a pair
took time off to investigate a small hole in the abdomen of a
“ripe” mouse, while the mate was busy excavating under the
carcass. For perhaps five minutes the feeding beetle worked
into the hole, until head and pronotum were inside the abdominal
wall. The viscera were explored rather superficially since the
position of the head could be discerned at all times by the moving

Dec., 1944]

Milne: Behavior

321

elevation it produced in the mouse’s skin. It was the observers’
opinion that the beetle was drinking rather than eating. Prior
to this feeding action, small flies had found the hole in the mouse ’s
abdominal wall a very interesting region, and considerable mois-
ture seemed to be present. After the beetle left the hole, there
was no liquid visible and no flies were attracted to the area.
During feeding, several blowflies ran against the posterior end
of the beetle, and were kicked away by violent movement of the
posterior legs. This kicking of molesting flies and ants seems a
common reaction in Nicrophorus.

Nicrophorus apparently discovers carrion entirely by smell,
while ants frequently locate freshly killed mice, seemingly as
part of routine foraging operations. Often ants had removed
the lips and nosetip of the rodent (the first part to be attacked
in all instances observed) before Nicrophorus arrived, but on
warm days (or nights) the beetles began to arrive within an hour
or less. Typical of the speed with which Nicrophorus gather
at a mouse is the following record made between six and ten
o ’clock one warm evening :

5 : 30 P.M.

Freshly killed mouse laid out in birch leaf litter.

6:05

J N. tomentosus.

6:07

N . tomentosus.

6:19

$ N. tomentosus.

6:38

J' N. orbicollis.

6:40

$ Silpha americana.

7:33

2 N. tomentosus.

8:02

2 N. orbicollis.

8:13

$ N. orbicollis.

8:36

<$ S. noveboracensis.

8:41

2 S. americana.

8:47

2 N. orbicollis.

9:00

& 2 8. noveboracensis.

9:40

carabid beetle.

9:50

<£ N . tomentosus.

10:00

Took in mouse for the night.

Thus in four hours, nine Nicrophorus arrived, of which approxi-
mately half were males. The sex ratio is remarkably close to 1 : 1.
Of twenty-two specimens of N. orbicollis collected in the sequence

322

Journal New York Entomological Society

[Vol. Lil

in which they arrived (no selection), ten were females. Of forty-
one N. foment osus collected in the same way (and during the same
time limits), twenty were males. Other observers confirm the
sex ratio (c/. Wood, 1873; Leech, 1935). The ratio of twenty-
two orbicollis to forty-one tomentosus is a very good value to
indicate the relative abundance of the two species in deciduous
woodland. In the sample catch cited above in chronological
order, the proportion of N. tomentosus is unusually low. Some
may have been driven off by N. orbicollis between arrival of
orbicollis and the frequent visits of the observers. Mosquitos
made more constant supervision too uncomfortable.

Another indication of the frequency with which Nicrophorus
come to carrion was afforded by an unintentional experiment.
Usually the observers placed all bait which was to be saved for
the morrow in a glass jar to be kept overnight on ice. On one
occasion a relatively fresh mouse was tossed casually into a but-
terfly net, the net folded on itself a few times, and left standing
outside the cabin over night. In the morning two N. tomentosus
were collected on the outside of the net just over the mouse inside.
Holes had been chewed through the net thicknesses to allow entry
of five other Nicrophorus, two orbicollis which had reached the
mouse, one orbicollis nearby in a fold of the net, and two tomen-
tosus in still other folds. Concern over the damage done to the
net precluded observations on how the beetles might have gone
about burying a carcass under such conditions.

The authors have been unable to identify the sex of Nicro-
phorus in the field without examining the genitalia — a procedure
which is accomplished more satisfactorily on an angesthetized
specimen. Kecords for N. orbicollis were kept, however, to deter-
mine if there were any clue to sex in the size of the specimens.
Ten male and eleven female orbicollis, measured freshly killed
and extended, form the basis of study. Since the telescoping of
the abdomen provided an independent variable, measurements
were made from the most anterior part of the head capsule to the
elytral apex. For the ten males there was a variation in this
measurement from 14 to 22 mm., mean 19.6 mm., with a standard
error of 2.7 mm. or approximately 14%. For the eleven females
the variation was from 16 to 22 mm., mean 18.5 mm., with a

Dec., 1944]

Milne: Behavior

323

standard error of 2.2 mm., or nearly 12%. Since the difference
between the means is only 1.1 mm. (about 6%), the authors were
unable to use size for sex recognition. A small male and a large
female, or vice versa, were encountered more frequently than two
large or two small specimens.

Due to the preference shown by Nicrophorus for work in the
shade or at twilight or night, difficulties were experienced in ob-
taining photographic records. So as to have beetles available to
photograph when the light was good, attempts were made to im-
prison photogenic specimens in glass jars with a little earth. If
the earth were moist, isolated specimens burrowed into it and
were active the following day, showing little agitation over their
confinement and taking on the burial of any mouse provided them
as soon as liberated quietly beside it. However, the beetles be-
came very hungry when kept over night, and unless maintained
in solitude, resorted to cannibalism. Specimens killed by their
fellows were ripped apart most ruthlessly, head from pronotum
from remainder of thorax from abdomen, and each part cleaned
of viscera. The victors frequently lacked tibiae, antennae, some-
times whole legs and elytra, demonstrating the ferocity of the
battles. Onthophagus, histerids and larvae of Silpha seemed able
to escape such attacks, but adult Silpha and Nicrophorus, as well
as other staphylinids and scarabaeids ( e.g Geotrupes) were de-
stroyed by hungry N. orbicollis and tomentosus. When speci-
mens were confined without food for more than a day, they be-
came sluggish and often died. Pukowski (1933) mentions N.
germanicus attacking adult Geotrupes silvaticus, capturing them
at horse dung, seizing them in the legs and mandibles and devour-
ing the viscera. Nicrophorus rolls over on its back or one side
while eating such prey.

4. RELATED OBSERVATIONS

Among other species frequenting, but not burying, small car-
casses, the following beetles were most common :

Staphylinidae : Silpha surinamensis Fab., S. lapponica Hbst.,
S. inaequalis Fab., S. novehoracensis Forst. and S. americana L.,
Staphylinus fossator Grav., Ontholestes cingulatus Grav. and
Creophilus villosus Grav.

324

Journal New York Entomological Society

[Vol. lii

Histeridae: unidentified — at least three species, probably dif-
ferent genera.

Scarabasidas : Geotrupes blackburnei Fab., Onthophagus hecate
Panz., 0. orpheus Panz., 0. nuchicornis L.

Of these Silpha noveboracensis and americana and Ontho-
phagus hecate were most numerous, coming both night and day,
particularly to carrion which had deteriorated considerably.
Staphylinus fossator, Ontholestes cingulatus and Creophilus vil-
losus came to similar carcasses but chiefly in daylight hours. All
of these species chewed at the bait. No evidence was obtained as
to the food of the histerid species. None of these beetles was
driven off by Nicrophorus, but only the histerids . and Ontho-
phagus remained if the carcass was buried.

Larvae of Silpha usually come in small numbers to deteriorated
carrion, and can be found many feet away heading toward the
carcass with remarkable accuracy. The larvae and adults of
Silpha walk with a rapid, jerky movement. In the adults the
jerks are more pronounced, possibly because the legs are longer.
Silpha larvae roll up like terrestrial isopod crustaceans (“sow
bugs, pill bugs”) and feign death, but the adults run away if
disturbed.

Most of the carrion beetles (and dung beetles) carry a number
of mites. Nicrophorus usually have less than twenty (c/. Leech,
1935). On arrival at carrion, many of the mites leave the beetles
and may be found running about on the carcass and nearby
ground. They hurry around on the body of the beetles and ap-
pear to share any agitation shown by the insect. Thus when the
beetle is disturbed, the mites move much more extensively and
leave the beetle much more frequently. The arrival of another
beetle or irritation shown over some obstacle to burial is enough
to greatly excite the mite population. The beetles were never
seen to show any reaction to the presence or position of the mites,
but it was noted also that the mites did not crawl out on the
antennas of the insects, although they ran over the mouthparts,
dorsum, venter and leg bases. Schaupp’s (1881) notes are inter-
esting in suggesting a relationship between mites and the death
of Nicrophorus pupas.

Pukowski (1933) and Leech (1935) have followed in great

Dec., 1944]

Milne: Behavior

325

detail the later phases of the life history, subsequent to burial of
the carcass. The present writers have not had opportunity to
repeat many of these observations. They did note, however, that
carcasses were cleaned fairly well of hair or feathers and worked
into a compact ball, kept free of collected moisture, the skin re-
maining in fair condition while the tissues became a slate-gray,
pasty mass, of a consistency similar to rotting dung. To see some
of the later stages in the life cycle, two mice were laid on the top
of four inches of earth and forest litter packed into a granite dish.
Within a day, both mice were buried by pairs of N. tomentosus,
and the dishful of carrion and insects was screened and carried
home to Pennsylvania. Perhaps due to the agitation of the trip,
the Nicrophorus came out of the ground and were observed run-
ning about over the soil, fluttering their wings and seeking an
exit. Two of the four beetles (a pair) were caught and removed,
and the remaining pair left to minister to any young they might
have on the way. Soil moisture was maintained by occasional
watering. The two beetles were seen running around a few times
more, but on each occasion they returned into the earth. After
two weeks one beetle was noticed on top of the ground, dead and
eviscerated. The pan of carrion and earth was turned out on a
paper. One mouse had dried to a hard mass. The second was
in much better condition, and on it were two fat yellow larvae of
N. tomentosus, so distended with food that their intersegmental
membranes were more conspicuous than the brown sclerites, and
almost helpless to roll over and crawl away. The other parent
(J) was found dead among the earth. About two dozen fly
puparia were among the soil particles, probably from the dried
mouse. No further observations were made and the specimens
were preserved for reference.

The foregoing observations were made during and between
shots with the 16 mm. motion picture camera. A visual record
in Kodachrome was obtained, somewhat over 600 feet in length,
showing the carrying and plowing behaviors, the burial of several
mice, the exhumation of mice and a snake, close-ups of N. tomen-
tosus adults and larvae, of N. orbicollis, including some footage
made at night to the hum of misquitoes while N. orbicollis chewed
through the string which held up the leg of a mouse. The present

326

Journal New York Entomological Society

[Yol. Lll

notes on the burial behavior of Nicrophorus should fill in gaps left
in the excellent work of Pukowski (1933) and Leech (1935) and
with the film, allow entomologists more widely to become ac-
quainted with the activities of this interesting genus.

ANNOTATED BIBLIOGRAPHY

Abbott, C. E. 1927a. Experimental data on the olfactory sense of Coleop-
tera, with special reference to the Necrophori. Annals Ent. Soc.
Amer. 20 : 207-216. Indicates olfactory sense located all over body
of Nicrophorus ; studied N. americcmus, orbicollis and tomentosus
from Md. in June.

. 1927b. Further observations on the olfactory powers of the

Necrophori. Ibid.: 550-553. Same in orbicollis in Wis. (July)

. 1937. The necrophilous habit of Coleoptera. Bull. Brooklyn Ent.

Soc., 32: 202-204. Indicates an obvious ecological succession of
beetles, the kind of carrion determining the numbers and species
involved.

Balduf, W. Y. 1935. The bionomics of entomophagous Coleoptera. (J.

S. Swift Co., Inc.) Pp. 68-75. An excellent summary of the
literature, particularly the work of Pukowski.

Bell, Prof. 1873. A glimpse of insect life. Can. Ent. 5: 94-95. Ob-
served adult Nicrophorus removing maggots from a dead dog.
Clark, C. U. 1895. On the food habits of certain dung and carrion beetles.

Jour. N. Y. Ent. Soc. 3: 61. Mentions Nicrophorus feeding on
maggots.

Davis, W. T. 1915. Silpha surinamensis and Creophilus villosus as pre-
daceous insects. Jour. N. Y. Ent. Soc. 25 : 150. Mentions Ni-
crophorus feeding on maggots from woodchucks and squirrels.
Duncan, C. D. and G. Pickwell. 1937. The world of insects. (McGraw-
Hill Book Co.) Pp. 194. Indicates that Nicrophorus larvae de-
velop in a carcass before bacteria cause serious decomposition.
Fabre, J. H. 1899. Souvenirs entomologiques. YI. Pp. 113-153.
Fowler, W. 1912. Coleoptera — General Introduction, Cicindelidae and
Paussidae. Fauna Brit. India 1912: 12, 24. Discusses olfactory
pits on antennae, and indicates Nicrophorus as typical of temperate
climates.

Furneaux, W. 1893. The outdoor world. (Longmans, Green & Co.)

Pp. 141-143. Indicates that female Nicrophorus is first to settle
on carcass, that male flies about, then both gorge selves on carrion,
after which male buries carcass without help of female.

Hatch, M. H. 1927a. Studies on the Silphinae. Jour. N. Y. Ent. Soc.

35 : 331-371. Primarily a taxonomic paper, indicating Nicrophorus
as the spelling used by Fabricius in erecting the genus.

. 1927b. Studies of carrion beetles of Minnesota, including new

species. Univ. Minn. Agric. Exp. Sta. Tech. Bull. 48: 3-7.

Dec., 1944]

Milne: Behavior

327

. 1940. Observations on Silphinse, with a note on intraspeeific varia-
tions and their designation. Jour. N. Y. Ent. Soe. 48: 233-244.

Jaques, H. E. 1915. The fish feeding Coleoptera of Cedar Point. Ohio
Nat. 15: 525-528. Indicates Nicrophorus feeding on fish washed
ashore.

Leech, H. B. 1935. The family history of Necrophorus conversator Wlk.

Proc. Ent. Soc. B. C. 31: 36-40. The most complete life history
study of a North American species.

Lutz, F. E. 1921. Fieldbook of insects. (G. P. Putnams Sons.) Pp.

295-296. Cites N. americanus as preferring reptiles, but indicates
no burying beetles came to any bait set out under experimental
conditions.

Milne, L. J. 1928. Notes on Silphidae in Haliburton County, Ontario.

Can. Field Nat. 42: 43. Earlier observatiohs on burying behavior.

Morley, R. L. 1902. Notes on stridulation. Ent. Mo. Mag. 38: 249-250.

Indicated that abdomen was rubbed on elytra to produce noise, and
coxae play no part in N. moratorium.

Motter, M. G. 1898. A contribution to the study of the fauna of the
grave. Jour. N. Y. Ent. Soc. 6: 201-231. Indicates no Nicrophorus
on disinterred corpses ( cf . Wood, 1873).

Pukowski, Erna. 1933. Oekolokische Untersuchungen an Necrophorus F.

Zeits. Morph. Okol. Tiere 27 : 518-586, 25 figs., refs. The most
complete account of the life history of European species of Ni-
crophus, including references to the European literature.

Schaupp, F. G. 1881. Description of the larva of Necrophorus tomentosus
Web. Bull. Brooklyn Ent. Soc. 4: 37-38. Indicated that mites
killed 50 per cent of Nicrophorus pupae by sucking them dry.

Selous, C. F. 1911. A preliminary note on the so-called carrion-feeding
Coleoptera. Ent. Mo. Mag. 47 : 86-89. Beetles flew into wind,
walking last few feet to carrion, dragged carcasses to softer earth.
Not convinced that Nicrophorus are feeders on carrion, probably
predators.

Steele, B. F. 1927. Notes on the feeding habits of carrion beetles. Jour.

N. Y. Ent. Soc. 35: 77-81. Indicated N. orhicollis and tomentosus
ate maggots from a woodchuck, and preferred maggots to carrion.

Thomson, Sir J. A. 1935. Biology for everyman. (E. P. Dutton.) Vol.
2 : 866-867. Brief summary of life history.

Wood, J. G. 1873. Insects at home. (Wm. Rutter.) Pp. 86-93. Indi-
cates Nicrophorus attack poorly interred human bodies in Russia.
Found sex ratio about 1:1. Other observations correspond to those
of Furneaux, but more extended.

328

Journal New York Entomological Society

[Yol. lii

THECLA BURDI KAYE, A SYNONYM

By E. Irving Huntington

W. J. Kaye described Thecla burdi from St. Vincent, B.W.I.,
in “The Entomologist, ’ ’ vol. 56, p. 277, 1923. In the collections
of the American Museum of Natural History there is a large
series of Theda angerona Godman and Salvin from Canefields,
Dominica, B.W.I., collected by Mr. L. E. Chadwick between Octo-
ber 24, 1933, and J anuary 16, 1934, as well as three females from
Roseau, Dominica, taken on October 19-20, . 1933, by the Same
collector ; in addition to this series there are one male and two
females taken on the type locality island of St. Vincent by Mr.
E. B. Isaacs, and one male taken at Brimstone Hill, St. Kitts,
B.W.I., March 16, 1929, collector E. I. Huntington. This series
has been compared with Kaye’s original description and there can
be no doubt that the insect described by Kaye as Theda burdi
is the same as that described by Godman and Salvin as Theda
angerona (Proc. Zool. Soc., London, p. 516, 1896).

Through an oversight, this synonomy was not referred to in
“Lycaenidas of the Antilles,” Comstock and Huntington (Ann.
New York Acad. Sci., vol. 45, p. 76).

Dec., 1944]

Rupert: Geometrid^e

329

A NEW SPECIES OF LAMBDINA, AND NOTES ON
TWO SPECIES OF BESMA (LEPIDOPTERA,
GEOMETRIDiE, ENNOMIN7E)

By Laurence R. Rupert
Horseheads, N. Y.

Recent study of various species of the group of the Geometridce,
until recently known as the genus Ellopia, has disclosed an ap-
parently undescribed species of Lambdina Capps, and sufficient
difference between Besma quercivoraria Gn. and B. endropiaria
G. & R., to warrant retention of both these names at specific rank.
A description of the new species and notes on the other two follow.

Lambdina canitiaria new species

Similar to athasaria Wlk., in shape, color, and maculation of wings; head
dark gray, showing no trace of the yellow that is so conspicuous in athasaria ;
thorax and abdomen dark gray, much less yellow than in athasaria ; male
antennae more narrowly pectinate than in athasaria. (The longest pectina-
tions are about 1 mm. long, while in athasaria they average lf-2 mm. long.)
Male genitalia similar to those of athasaria but with slight differences which
may not prove constant when a longer series of canitiaria is available. I
have examined the genitalia of two males of canitiaria, and of twelve males
of athasaria. The canitiaria genitalia both differ from any athasaria ex-
amined and from the figures shown by Capps (Proc. U. S. N. M., Yol. 93,
Plate 3), in having (1) finer and shorter spinules on the furca, and fewer
of them along the basal third; (2) the terminal part of the furca wider and
more rounded; and (3) the auger-like process at the end of the aedeagus less
prominent.

Wing expanse 1-11 inches, averaging a little smaller than athasaria.

At Horseheads, N. Y., the only locality from which it is known,
canitiaria is the earliest Lambdina to appear in the spring. It
is on the wing about three weeks earlier than athasaria and the
periods of flight of the two species have not been observed to
overlap in the same season.

Holotype. — J*, Horseheads, N. Y. — May 22, 1940. (In
Franclemont collection.)

Allotype. — J, Horseheads, N. Y. — May 4, 1938. (In Rupert
coll.)

330

Journal New York Entomological Society

[Vol. LII

Paratypes. — 3 Horseheads, N. Y. — May 22. 1940, and

May 9, 1943. (In Rupert coll.)

Besma quercivoraria Gn. — Two females were taken at Horse-
heads in May 1943. Eggs were obtained from, both, and larvae
from both lots were raised to maturity. No differences of note
were observed among the eggs, larvae, and pupae of the two lots.

The eggs were elliptical, with the surface finely and evenly
pitted, at first translucent, almost colorless, but with a faint
greenish tinge, which soon became darker and more distinctly
green.

The young larvae were pale yellow green, very slender and
very active. They accepted as food several species of oak, but
refused everything else offered including maple. Beech was.
not readily available, and was not offered at this time. On June
22, when most of the larvae were in the last stage, I left Horse-
heads to spend several weeks at Sardinia, N. Y., where oak is
difficult to find. The larvae then accepted beech readily, but
still refused maple.

The mature larvae were dull light yellow-green, with head
somewhat mottled with brown; second thoracic segment with
two prominent brown lateral warts but no conspicuous dorsal
adornment; first, second, third, fifth, and sixth abdominal seg-
ments with inconspicuous lateral warts; third and sixth ab-
dominal segments each bearing in addition to the lateral warts
two prominent subdorsal warts, those on segment 3 somewhat
fused with ehch other, and with the lateral warts of this seg-
ment; other segments without special prominences.

The larvae pupated in late June and early July, forming pale
brown pupae, with wing cases streaked, and abdomen heavily
speckled with dark brown. All of the pupae produced moths
the same season, mostly between July 9 and 25, but with a few
stragglers in August, September, and October.

Besma endropiaria G. & R. — A female of this species was
taken at Sardinia, N. Y., on June 27, 1943, but only fifteen eggs
were obtained. These were similar in size, shape, and surface
texture to those of quercivoraria, but lacked the green tinge, and
showed no color change except the normal darkening just before
hatching. They hatched on July 9, the same day that produced
the first adult of quercivoraria as noted above.

Dec., 1944]

Rupert: Geometries

331

The young larvae were similar in appearance to those of querci-
voraria, but the only food they would accept was maple. They
refused beech, but were not offered oak since it was not readily
available. This preference for maple was not unexpected, for
several years ago a single larva which I found upon maple pro-
duced a male of this species, which I still have in my collection.

The mature larvae were either green or brown, with head more
uniform brown and less mottled than in quercivoraria ; second
thoracic segment with lateral and subdorsal warts so fused as to
form a conspicuous ridge extending completely from one lateral
wart to the other; first and second abdominal segments with
warts similar to those of quercivoraria, and in addition a ventral
prominence on the second segment; third abdominal segment
with warts similarly placed to those of quercivoraria, and simi-
larly fused, but larger; fifth abdominal segment with two well-
developed subdorsal warts, somewhat fused with each other, but
not with the lateral ones; sixth abdominal segment with warts
similar to those of the fifth segment.

Of the thirteen larvae reared, seven were green with brown
warts, similar in color to quercivoraria larvae, while the rest were
dull brown with darker brown warts. It is reasonable to sup-
pose that a brown form of the larva of quercivoraria may occur.

The larvae pupated in late August, producing pupae much
darker than those of quercivoraria. This color difference may
not be constant in large series, however, for it has been noted
that among certain other Geometridce the pupae as well as the
larvae exhibit two or more color phases. These pupae produced
no moths until the following season. The failure of endro-
piaria to produce two generations a season is in accordance with
the results of field collecting at Ithaca, Horseheads, and Sar-
dinia, N. Y. (Franclemont and Rupert), and at Chicago, 111.
(Wyatt). At Ithaca, Horseheads, and Chicago both species
occur, with endropiaria flying between the two broods of querci-
voraria. At Sardinia endropiaria flies in June and querci-
voraria has never been taken.

In addition to the differences noted above between these two
species, there appears to be a constant difference in the male
antennae. This was first called to my attention by Dr. Forbes,

332

Journal New York Entomological Society

[Yol. lii

and I have since confirmed his observation by examining all the
males of both species in my collection. In quercivoraria the
length of the pectinations shows a very gradual increase from
each segment to the next from the base of the antenna, while in
endropiaria this increase is more abrupt. Likewise the decrease
in length of pectinations near the tip is correspondingly gradual
in quercivoraria and abrupt in endropiaria. In general the
pectinations are slightly shorter even at the middle of the antennae
in quercivoraria.

Considering altogether the differences found between these two
species in egg, larval structure and food, period of flight and
number of generations a year, and structure of male antennae,
along with the well-known differences in the appearance of the
adults, it seems quite reasonable to consider them distinct species,
even though the genitalia show no obvious differences. It is true,
as Mr. Capps points out (Proc. U. S. N. M., Yol. 93, p. 142), that
apparent intergrades occur. However, if only one species is
involved, it would seem that such intergrades should be more
commonly found than they are in localities where the typical
forms are both abundant. Intermediate specimens have never
been taken at Horscheads, nor, so far as I can discover, at Ithaca,
where intensive collecting over a period of many years has pro-
duced large series of both regular forms.

PLATE XI

Figure 1.
Figure 2.
Figure 3.
Figure 4.
Figure 5.

Figure 6.

Besma quercivoraria Gn. Larva.

Besma endropiaria G. & E. Larva.

Lambdina canitiaria new species. Furca of male genitalia.
Lambdina athasaria Wlk. Furca of male genitalia.

Besma quercivoraria. Base of male antenna, showing only the-
first five pectinations, and only those on one side.

Besma endropiaria. Similar view of part of male antenna.

(Jour. N. Y. Ent. Soc.), Vol. LII

(Plate XI)

Dec., 1944]

Caldwell: Psylliid^e

335

PSYLLIIDiE FROM TROPICAL AND SEMITROPICAL
AMERICA (HOMOPTERA)*

By John S. Caldwell

ClRCLEVILLE, OHIO

Much of this material consists of species set aside and studied
over a period of time because many are not generically distinct,
that is they possess characters common to more than one genus
according to the present interpretations. Rather than attempt
to revise several genera which would be no more than my per-
sonal interpretation and quite artificial, I believe the true nature
of the generic situation may be represented by a series of fre-
quency curves with the generic types and closely related species
near the crown, and the less related species farther down the
curves. This way it seems possible that two species belonging in
separate genera may be very closely related, which is the true
situation at present between Rhinopsylla and Kuwayama.

The writer takes this opportunity to express his appreciation
to Miss Louise M. Russell of the U. S. Department of Agriculture
for comparing much of this material with specimens in the D. L.
Crawford collection. Dr. Leonard Tuthill has kindly examined
some of this material and expressed his opinion concerning its
validity. Unless stated to the contrary all types are in the
writer’s collection.

Calophya arcuata new species, (Fig. 5)

Length 2 mm., forewing 1.7 mm. Deep orange over all with black eyes
and genae and yellow legs.

Head broad. Vertex smooth, impressed discally making posterior ocelli
greatly elevated ; cephalic half rounded downward and forward. Genae
widely separated, acute, one-third as long as vertex. Thorax scarcely arched ;
pronotum as long as vertex, deflexed, appearing tricarinate. Membrane of
forewing minutely rugose; pterostigma long, open at base; Rs long; M highly
arched around large cubital cell.

* Zoologically speaking, Chermes alni L., 1758, is the same insect now
known as Prociphilus tessellatus (Fitch) [Data from Pehr Kalm, 1756] ;
hence Kirkaldy’s Psyllia, type pyri L., is the type genus of the family
Psylliidae.

336

Journal New York Entomological Society

[Yol. LII

Female genital segment as long as rest of abdomen; dorsal valve bulbose
in basal half, apical half deflexed, extreme apex acute; ventral valve sub-
equal in length to dorsal.

Holotype: female from Bonefish Key, Florida, 2-24—40 (Cald-
well).

Kuwayama striata new species, (Fig. 6)

Length 2.5 mm., forewing 2.1 mm. Greyish-yellow species with four promi-
nent red stripes on mesoscutum.

Vertex rather small, flat, somewhat rounded in front. Genae subspherical.
Prescutum as long as broad, longer than scutum, rounded eephalad. Fore-
wings three times as long as broad, Es reaching as far as furcation of M.

Female genital segment three-fourths as long as rest of abdomen; dorsal
valve straight dorsad, suddenly blunted apically; ventral valve somewhat
stylate in apical fourth.

Holotype: female from Saltillo, Coahuila, 9-23-41 (DeLong,
Good, & Caldwell).

Kuwayama hyalina new species, (Fig. 2)

Length 2.9 mm., forewing 2.3 mm. Yellow over all with black eyes.
Wings very milky white, hyaline.

Vertex scarcely deflexed, short, emarginate caudad; foveae shallow. Genae
roundly swollen. Clypeus visible from front but not prominent. Pronotum
nearly vertical; prescutum flat dorsad, acute eephalad. Forewings two and
a half times as long as broad, not especially acute apically; Es reaching to
furcation of M.

Female genital segment abruptly styliform in apical half; anal opening
located well caudad on dorsal valve.

Female holotype and paratype from Tasquillo, Hidalgo, 10-
24-41, Km. 172 (DeLong & Good).

Kuwayama mexicana new species, (Fig. 1)

Length 4 mm., forewing 3 mm. Vertex cream excepting elongate foveae.
Pronotum cream; thoracic dorsum red with light cream median stripe. An-
tennae, legs and abdomen black.

Eobust species. Vertex relative horizontal, flat. Eyes prominent. Genae
produced into minute cones, blunt. Antennae twice as long as width of head.
Pronotum small, depressed to level of vertex ; prescutum high, acute eephalad ;
scutum short. Forewings large, over twice as long as broad, not acute. Hind
wings small, not reaching to furcation of M in forewings. Pubescence promi-
nent on legs.

Forceps of male simple, slender, as long as proctiger.

Dec., 1944]

Caldwell: Psyllihle

337

Holotype : male from Mexico, D. F., west 18 Kms., 9-1-39
(DeLong).

Kuwayama lateralis new species, (Fig. 3)

Length 3.5 mm., forewing 2.7-3 mm. General color grey with faint red
laterally on prescutum and scutum. Venter of head and thorax black to
dusky. Abdomen black dorsad, dusky ventrad with light lateral stripe on
either side for full length.

Vertex deflexed, foveae deep, ocelli raised. Antennae twice as long as width
of head. Genae swollen. Thorax rather flat, not robust. Forewings almost
three times as long as broad.

Proctiger of male longer than forceps. Forceps produced on cephalic
margins at midlength. Female genital segment almost as long as rest of
abdomen; dorsal valve straight, somewhat styliform apically; ventral valve
abruptly styliform in apical third.

Male holotype, female allotype, and paratypes from Mexico,
D. F., west Km. 20, 11-24-38 (Caldwell).

Trioza rhinosa new species, (Fig. 4)

Length 4.5 mm., forewing 3.5 mm. Shining black over all with white genae.

Head broad; eyes prominent; postocular areas large. Vertex sloped in-
ward toward median line, rolled roundedly forward ; medial ocellus prominent.
Genal cones acute, divergent, one-fourth as long as vertex. Pronotum ver-
tical; rest of thorax scarcely arched. Femora prominent, metatibiae with
apical spur ratio of 3—1. Fore wings twice as long as broad, rounded; cubital
cell smaller than medial; Rs scarcely separated from and paralleling M in
basal fourth.

Forceps of male slender, incurved in caudal aspect. Proctiger long, broad
in lateral aspect.

Holotype: male from Tehuacan, Puebla, 10-17-41 (DeLong,
Good, Caldwell, & Plummer).

The peculiar formation of the head and forewings places this
species close to the Rhinopsylla. In general appearance it is close
to T. diospyri Ashm.

Metatrioza neotriozella new species

Length 2.5-2. 7 mm., forewing 2-2.2 mm. Head and genal cones black.
Mesonotum with red center and two black stripes on either lateral margin.
Costal margin of forewing black basally.

Head as broad as thorax. Vertex strongly concave between eyes, median
suture prominent. Genal cones slender, as long as vertex, closely appressed.
Antennae scarcely as long as width of head. Thorax scarcely arched. Fore-
wings almost three times as long as broad. Hind tibiae with three closely ap-
pressed spurs at apex.

338

Journal New York Entomological Society

[Vol. LII

Forceps of male of even width throughout, truncate apically, evenly arcuate
in caudal aspect.

Dorsal valve of female genital segment styliform in apical half, inflated in
basal half ; ventral valve somewhat styliform.

Male holotype, female allotype, and paratypes from Tncson,
Arizona, 8-16-40, (D. J. & J. N. Knull) are in the Ohio State
University collection at Columbus, Ohio.

Except for the unique form of the vertex this species would
belong in N eotriozella Crawford.

Optomopsylla new genus

Head including eyes much broader than pro and mesonotum, as broad as
metathorax. Vertex smooth except for median groove, vertical, rounded
gently caudad. Posterior ocelli projecting laterad, almost touching the com-
pound eyes. Pronotum much sunken below level of mesonotum and vertex.
Propleurites much compressed, somewhat transversely concave. Forewings
with Cu branched from main stem before R and M.

Related to Ceropsylla but differentiated by the structure and
form of head and thorax.

Type: Optomopsylla formiciformis n. sp.

Optomopsylla formiciformis new species, (Figs. 7, 7-A & 7-B)

Length 3.5 mm., forewing 3.1 mm. Black with the exception of basal four-
fifth of antennse, lateral and ventral portion of pro and mesothoracic femora,
all of metathoracic legs, venter of abdomen and genital segment, most of
mesothorax, scutum of mesothorax and dorsum of metathorax whitish. Fore-
wings clear with R+M+Cu and R heavily darkened.

Genal cones heavy, blunt, divergent, almost as long as vertex. Head ver-
tical. Thorax scarcely arched, flat in profile; pronotum greatly depressed.
Metatibise with apical spur ratio of 3—1. Forewings three times as long
as broad, acute; cubital cell long, flat.

Female holotype from Zamora, Michoacan, 10-2-41 (DeLong,
Good, Caldwell, & Plummer), on willow.

The form combined with the deceptive markings gives this
psyllid the appearance of a black ant in dorsal or lateral aspect.
The genae appear as mandibles, the fore part of the thorax is
much narrowed and the color on the last thoracic segment and
base of the abdomen form the optical illusion of a narrow waist.
The metathoracic legs are white and scarcely visible but the
heavily embrowned vein in the forewings completes the illusion
of a walking leg. This specimen was swept from willow along

Dec., 1944]

Caldwell: Psylliid^e

339

with a net full of ants about the same size. Whether accidental
or not, to me this is a remarkable example of mimicry.

Euphalerus dubius new species, (Fig. 9)

Length 4 mm., forewing 3 mm. General color green variegated with gray.

Head as broad as thorax, almost perpendicular. Vertex twice as broad as
long, flat, median suture very smooth. Genal cones scarcely differentiated
from vertex, short, blunt, contiguous on basal third. Antennae one and a half
times as long as width of head. Thorax strongly arched, very smooth, sutures
very fine between thoracic segments and between head and pronotum.
Pleurites of prothorax subequal. Forewings long, somewhat rhomoboidal;
pterostigma long and broad.

Female genital segment as long as rest of abdomen.

Female holotype from Davis Mts., Texas, 7-2-40 (D. J. & J. N.
Knull). Type in Ohio State University collection at Columbus,
Ohio.

Psyllia martorelli new species, (Figs. 8 & 8-A)

Length 2.5-3. 5 mm., forewing 2-2.5 mm. Specimens in preservative color
unknown. Mesoscutum with broad light stripes.

Head broader than thorax; eyes somewhat stalked; posterior ocelli greatly
elevated. Vertex scarcely deflexed, rolled somewhat roundedly forward.
Genae scarcely swollen; frons much sunken but not covered by genae. An-
tennae almost as long as entire insect. Thorax scarcely arched. Forewings
little over twice as long as broad ; apical margins almost flat ; pterostigma not
apparent ; costal margins pubescent.

Apices of male forceps slightly bifurcate. Female genital segment as long
as rest of abdomen; both valves very slender, stylate in apical half.

Holotype male, allotype female, and paratypes from Villalba,
Puerto Rico, May 1940, on “Inga Inga” (L. F. Martorell).

This species shows some relationship to the Pauropsyllince.

The writer dedicates this outstanding species to his friend Dr.
Luis F. Martorell.

Psyllia berryi new species, (Figs. 11 & 11-A)

Length 5.4 mm., forewing 4.4 mm. Greenish-yellow over all.

Vertex very small, cephalic margin compressed between genae. Genae
greatly developed, larger than vertex, inner margins contiguous, apices blunt.
Antennal insertion in front of ventral margins of eyes. Eyes very small;
postocular area large. Pronotum strongly descending, prominent; prescutum
rounded, longer than scutum. Forewings almost three times as long as
broad; pterostigma very narrow, elongate; cubital cell twice as large as
medial.

Female genital segment short; dorsal valve rounded, somewhat bulbose
apically; anal opening with serrate margins; ventral valve short, thick.

340

Journal New York Entomological Society

[Vol. lii

Female holotype from Santaram, Para, Brazil, October 1942
(L. A. Berry).

The gigantic development of the gense sets this species apart
from any psyllid known to me; however, the structure of the
entire insect is true to the genus. Too many of the present
genera of Psylliidaa have been established on gradational char-
acters for me to add another when there are no fundamental
differences on which to base a decision.

The writer takes great pleasure in naming this unique species
in honor of his friend Lawrence A. Beery, Jr.

Psyllia cedusa new species, (Fig. 10)

Length 2.5 mm., forewing 2.1 mm. General color orange-yellow. Fore-
wings with four black marginal spots.

Vertex twice as broad as long; fovese sharp, deep; posterior ocelli greatly
elevated. Genal cones three-fourths as long as vertex, divergent. Whole
head deflexed, as broad as thorax. Thorax not especially arched yet pro-
notum is nearly vertical. Hind tibiae with small basal spur. Forewings little
over twice as long as broad; cubital cell very highly arched; pterostigma,
small, equilaterally triangular.

Female genital segment as long as rest of abdomen; dorsal valve straight,
stylate in apical third with apex turned up ; ventral valve narrowed in apical
half, curved dorsad.

Female holotype from Jesus Carranza, Veracruz, 10-14-41
(DeLong, Good, Caldwell, & Plummer).

PLATE XII

Figure 1. Kuwayama mexicana. Lateral view of male genitalia.

Figure 2. Kuwayama hyalina. Lateral view of female genitalia.

Figure 3. Kuwayama lateralis. Lateral view of male genitalia.

Figure 4. Trioza rhinosa. Lateral view of male genitalia.

Figure 5. CalopTiya arcuata. Lateral view of female genitalia.

Figure 6. Kuwayama striata. Lateral view of female genitalia.

Figure 7. Optomopsylla formiciformis. Lateral view of female genitalia.
Figure 7-A. Profile of head and thorax.

Figure 7-B. Dorsal view of head and thorax.

Figure 8. Psyllia martorelli. Lateral view of female genitalia.

Figure 8-A. Caudal view of male forceps.

Figure 9. Euphalerus dubius. Profile of head and thorax.

Figure 10. Psyllia cedusa. Lateral view of female genitalia.

Figure 11. Psyllia beeryi. Lateral view of female genitalia.

Figure 11-A. Dorsal view of circum-anal ring.

(Plate XII)

3

Dec., 1944]

Brown: Butterflies

343

NOTES ON MEXICAN BUTTERFLIES, IV

By F. Martin Brown
NYMPHALIDiE — I

Heliconiinae

230. Heliconius ismenius telchinia Doubleday.

G. & S. (1), 1: 149, 667.

S. (2), p. 380, pi. 72b.

H. (3), p. 672.

Ojo de Agua, Vera Cruz, 1600 ft., 1 v.12.41 (R.P.).

237. Heliconius petiverana Doubleday & Hewitson.

G. & S., 1 : 153, 668.

S., p. 392, pi. 78b.

H. , p. 673.

El Banito Valles, San Luis Potosi, 200 ft., 3 J'J' vii.22.39 ;
1^1$ iv. 28-29. 40 (H.H.).

El Pujal, San Luis Potosi, 100 ft., 2 J'.J' 2 J? vii. 18-20. 39

(H.H.).

Ojo de Agua, Vera Cruz, 1600 ft., 1 J* v.12.41 (R.P.).
Rio Blanco, Vera Cruz, 2200 ft., 1 $ 1 2 v.10.41 (R.P.).
El Sabino, Uruapan, Michoacan, 1 J* vii. 15-30. 36
(H.D.T.).

The broad red bar on the forewings of the Vera Cruz specimens
is more rosy red than on the San Luis Potosi specimen. On the
latter the bar is tomato red. Is this a real difference or one due
to the age of the specimen ? In each of the above series there are
fresh and flown specimens ; females seem to be smaller and more
worn than the males.

238. Heliconius charithonia Linnaeus.

G. & S., 1 : 151.

S., p. 394, pi. 79a.

H. , p. 673.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1600
ft., 3 1 ? vi. 16-20.40 (H.H.).

344

Journal New York Entomological Society

[Yol. LII

nr. Villagran, Tamaulipas, 1 , g 1 5 iv.28.41 (R.P.).

60 mi. So. of Victoria, Tamaulipas, 1 J* vii.6.36 (H.D.T.).
Jacala, Hidalgo, 4500 ft., 1 J1 vi.29.39 (H.H.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 4 J'J1
6 ?? v.28-29.41 (R.P.).

El Mante, San Luis Potosi, 1 iv.29.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 1 £ 1 2
yii.22.39, 2 2 ?? vi.26-27.40 (H.H.) ; 1 ?

iv. 31.41 (R.P.).

El Pujal, San Luis Potosi, 100 ft., 3 J'J' 1 J vii. 17-31. 39
(H.H.).

Tuxpango, Vera Cruz, 1500 ft., 1 5 v.9.41 (R.P.).

Fortin, Vera .Cruz, 1600 ft., 4 v.3.41 (R.P.).

Ojo de Agua, Vera Cruz, 1600 ft,, 1 <$ 4 55 v.12.41
(R.P.).

Orizaba, Vera Cruz, 2000 ft., 1 1 5 v.6.41 (R.P.).

Rio Blanco, Vera Cruz, 2200 ft., 1 v.10.41 (R.P.).

Acahuato, Michoacan, 3000 ft,, 1 viii.2.40 (H.H.).

All of the females show a distinct rusty flush over the yellow
bars, it is most intense toward the margins. Among these speci-
mens the females seem to be more worn than the males.

243. Eueides aliphera gracilis Stichel.

G. & S., 1 : 163, 669 (as aliphera).

S, p. 399.

H. , p. 673.

Tuxpango, Vera Cruz, 1500 ft., 1 5 v.9.41 (R.P.).

Ojo de Agua, Vera Cruz, 1600 ft., 1 5 v.12.41 (R.P.)

244. Eueides cleobaea zorcaon Reakirt.

G. & S., 1 : 165, 670.

S, p. 368.

H. , p. 674.

Galeana, Nuevo Leon, 6500 ft., 2 1 5 iv.29.41 (R.P.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 4 J'J'

v. 29.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 2 J'J' 3 §§
vii. 22. 39 ; 1 $ 1 ? vi.28.40 (H.H.).

El Pujal, San Luis Potosi, 100 ft., 1 3 vii. 12-21. 39

(H.H.).

Dec., 1944]

Brown: Butterflies

345

Fortin, Vera Cruz, 1500 ft., 1 v.3.41 (R.P.).

Rio Blanco, Vera Cruz, 2200 ft., 3 3 v.10.41

(R.P.).

There is considerable variation in this species. The ground
color varies from rich fulvous brown to faun. The apical light
spots are either concolorous with the rest of the wing or lighter.
Seitz ( l.c .) states that in the females these spots are “rather pale
yellow.’ ’ They do tend to be lighter in the females but yellow
apical spots are not restricted to females nor do all females have
yellow spots.

There is a peculiar variation in the color of the antennae. The
females have antennae that are almost wholly yellowish. The
males have dark antennae that are yellow tipped.

Dioninae

246. Dryas julia delila Fabricius.

G. & S., 1 : 168, 670.

S, p. 400.

H. , p. 674.

60 mi. So. of Victoria, Tamaulipas, 1 2 vii.6.36 (H.D.T.).
El Sol, Tamazunchale, San Luis Potosi, 400 ft., 15 J'J'
v.28-29.41 (R.P.).

Arroyo del Calabezas, San Luis Potosi, 250 ft., 1
iv.30.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 3 J'J' 1 $
vii.22.39 (H.H.) ; 1 ^ iv.30.41 (R.P.).

El Pujal, San Luis Potosi, 100 ft., 3 J'.J' vii.17.39 (H.H.).
Fortin, Vera Cruz, 1600 ft., 1 v.3.41 (R.P.).

Ojo de Agua, Vera Cruz, 1600 ft., 1 J v.12.41 (R.P.).
Rio Blanco, Vera Cruz, 2200 ft., 6 J'J' v.10.41 (R.P.).
El Sabino, Uruapan, Michoacan, 1 vii. 15-30.36

(H.D.T.).

The black apical streak, characteristic of julia is present in a
reduced fashion on males from El Sol, El Banito, El Pujal and
Rio Blanco, and on both females.

248. Dione juno huscama Reakirt.

G. & S., 1 : 170, 670 (as juno).

S., p. 401, pi. 84e.

346

Journal New York Entomological Society

[Vol. Lit

H., p. 674.

Galeana, Nuevo Leon, 6500 ft., 2 J'J1 iv.29.41 (R.P.).
Jacala, Hidalgo, 4500 ft., 1 $ vi.21.39 (H.H.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 6 J'J'1
1 2 v.28-29.41 (R.P.).

El Sabino, Uruapan, Michoacan, 1 J' vii.15-30.36
(H.D.T.).

Tancitaro, Michoacan, 6000 ft., 1 1 2 vii.20.40 (H.H.).

In general all of these are more boldly marked with black than
Seitz’ figured specimen but not quite so boldly as his figure of
juno juno. The El Sol series is reasonably fresh; the others,
are worn.

249. Dione vanillae Linnaeus.

G. & S., 1: 171, 671.

S., p. 401, pi. 84f.

H. , p. 674 (as v. insular is May).

Jacala, Hidalgo, 4500 ft., 4 3 2? vi.l5-vii.3.39

(H.H.)

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 £
v.29.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 2 J'J' vii.22.39

(H.H.).

Tuxpango, Vera Cruz, 1500 ft., 1 $ v.8.41 (R.P.).

Rio Balsas, Guerrero, 2400 ft., 1 v.26.41 (R.P.).

Apatzingan, Michoacan, 1050 ft., 1 2 viii.5.40 (H.H.).
El Sabino, Uruapan, Michoacan, 1 2 vii.15-30.36
(H.D.T.).

None of these specimens is insularis ; all have fully developed
markings along the margins of the hindwings. The males of the
Jacala series lack the small black dot between M3 and Cui close
to the cell on the hindwings. This dot is present on all of the
other males and on all of the females.

NYMPHALINiE
Tribe Argynnidi

251. Euptoieta claudia Cramer.

G. & S., 1 : 174, 671.

Dec., 1944]

Brown: Butterflies

347

S., p. 403, pi. 85a.

H., p. 674.

Ojo de Agua, Sabinas Hidalgo, Nuevo Leon, 1000 ft., 1 $
vi. 14.40 (H.H.).

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1600
ft., 2 1 ? vi. 16-18.40 (H.H.).

60 mi. So. of Victoria, Tamaulipas, 1 £ vii.6.36 (H.D.T.).
Galeana, Nuevo Leon, 6500 ft., 1 £ viii.3.39 (H.H.).
Jacala, Hidalgo, 4500 ft., 2 vi.25.39 (H.H.).

El Banito, Valles, San Luis Potosi, 200 ft., 2 1 J

vi. 27-28. 40 (H.H.).

Tuxpango, Vera Cruz, 1500 ft., 1 J v.9.41 (R.P.).
Chichen Itza, Yucatan, 1 J viii.30.36 (H.D.T.).

Rio Balsas, Guerrero, 2400 ft., 1 J v.26.41 (R.P.).

El Sabino, Uruapan, Michoacan, 1 5 vii. 15-30. 36
(H.D.T.).

Tancitaro, Michoacan, 6600 ft., 1 J viii. 14.40 (H.H.).

This is probably one of the most adaptable of American butter-
flies. I have found it breeding from sea-level in the tropics to
11,800 ft., in the lower margin of the alpine-arctic zone in Colo-
rado. In that state it is not uncommon far above tree line, at
13,000 ft. or more. I feel that this species is a true archaic an-
cestor of the genus Argynnis. I know of no structural differences
between Euptoieta and Argynnis that are of generic value.

The species claudia is also found in the South Temperate Zone.
I can find no reliable character to separate material from the
Argentine ( hortensia Blanchard) from the material before me.
In this respect the two forms are analogous to Phoebis eubule and
amphitrite.

252. Euptoieta hegesia Cramer.

G. & S., 1 : 175, 671.

S., p. 404, pi. 85a.

H. , p. 674.

Sabinas Hidalgo, Nuevo Leon., 900 ft., 1 J' vi.15.39
(H.H.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 J'
iv.31.41 ; 2 1 ? v.29.41 (R.P.).

348

Journal New York Entomological Society

[Vol. Lll

El Banito, Valles, San Luis Potosi, 200 ft., 1 2 vii.21.39 ;

3^1? vi.28.40 (H.H.).

Fortin, Vera Cruz, 1600 ft., 1 $ v.4.41 (R.P.).

Orizaba, Vera Cruz, 2000 ft., 2 ££ v.6.41 (R.P.).

Rio Blanco, Vera Cruz, 2200 ft., 1^1? v.10.41 (R.P.).
Chichen Itza, Yucatan, 2 $$ 1 2 viii.30.36 (H.D.T.).
Rio Blanco, Guerrero, 2400 ft., 1 5 v.26.41 (R.P.).

Sta. Lucrecia, 2 J'J1 1 2 iv.24.39 (H.D.T.).

Acahuato, Michoacan, 3000 ft., 1 J' viii.2.40 (H.H.).

El Sabino, Uruapan Michoacan, 1 $ 1 § vii.15-30.36
(H.D.T.).

266. Melitaea definita Aaron ( ?).

G. & S., 2: 676, pi. 108, ff. 11, 12 (as schausi).

S., p. 433 (p. 434, pi. 88h, as schausi).

H. , p. 676.

Cuidad Victoria, Tamaulipas, 1 vi.19.39 (H.H.).

Jacala, Hidalgo, 4500 ft., 1 J1 vi.24.39 (H.H.).

These two specimens come fairly close to Texan definita but are
by no means typical. The upper side gives the impression that
they are Euphydryas and the underside continues the impression.
I suspect that this is a paleonearctic species and may be closely
related to the archaic species from which our North American
Euphydryas have sprung. The Jacala specimen extends the
known range of the species.

270. Melitaea theona f. theona Menetries.

G. & S., 1 : 192, 2 : 677 (as Phyciodes) .

S., p. 438, pi. 89g (as Phyciodes) .

Ha. (4), p. 26 (as Phyciodes) .

H. , p. 676.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 2 vi. 15-16.40 (H.H.).

Galeana, Nuevo Leon, 6500 ft., 1 J' iv.29.41 (R.P.).
Jacala, Hidalgo, 4500 ft., 2 $$ 1 2 vi.l9-vii.3.39 (H.H.).
El Sol, Tamazunchale, San Luis Potosi, 400 ft., 4 J'J'
3 $? iv.31; v.29.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 1 2 vii.22.39 ;
, 1 2 vi.26.40. (H.H.) ; 1 J' iv.30.41 (R.P.).

Dec., 1944]

Brown: Butterflies

349

270a. Melitaea theona f. thekla Edwards.

G. &S.,2: 677 (as theona).

S., p. 433, pi. 88f.

Ha., p. 26 (as Phyciodes theona in part).

H. , p. 676.

Ojo de Agua, Sabinas Hidalgo, Nuevo Leon, 1500 ft.,
lcfvi.14.40 (H.H.).

Las Ad juntas, Nuevo Leon, 3000 ft., 1 vi. 21.40 (H.H.).
Arroyo del Meco, Tamaulipas, 1520 ft., 1 2 iv.28.41
(R.P.).

. Melitaea sp.

Two battered specimens of the gabbi group were taken by Potts,
Cumbres, Yera Cruz, 8000 ft., v.17.41, and Rio Balsas, Guerrero,
2400 ft., v.26.41.

272. Phyciodes elada elada Hewitson.

G. & S., 1 : 196, 2 : 679, pi. 21, ff. 6, 7.

S., p. 437, pi. 89f.

Ha., p. 19.

H. , p. 676 (as Melitcea).

Cumbres, Vera Cruz, 8000 ft. (nr. Km. 295) 1 v.7.41

(R.P.).

Apatzingan, Michoacan, 1050 ft., 2 viii. 3-5.40

(H.H.).

Acahuato, Michoacan, 3000 ft., 2 ££ viii. 2.40 (H.H.).
The two Acahuato specimens are much larger than the two
from Apatzingan. The baso-apical radii of the forewings are

15.0, 14.2 and 12.5, 12.0 mm. respectively. The Cumbres specimen
may not be properly placed here, Hoffmann records the species
only from the west coast.

272a. Phyciodes elada callina Boisduval.

Ha., p. 20.

Sabinas Hidalgo, Nuevo Leon, 960 ft., 1 J' vi.15.39
(H.H.).

Jacala, Hidalgo, 4500 ft., 10 3 2? vi.23-vii.6.39

(H.H.).

This may be a mixed series but I cannot bring myself to split-
ting it between two names. Hoffmann does not include the race.
I suspect that his Ulrica is really callina.

350

Journal New York Entomological Society

[Yol. lii

274. Phyciodes liriope guatemalena Bates.

G. & S., 1 : 198, pi. 21, f. 23 (as fragilis).

R. (5), p. 435, pi. 89b (as guatemala) .

Ha., p. 67.

H. , p. 676.

So. of El Mante, San Luis Potosi, 1 £ iv.29.41 (R.P.).

El Sol, Tamazunchale, San Luis Potosi, 2 J'J' 10.31.41
(R.P.).

Hda. Potrero Viejo, Paraje Nuevo, Vera Cruz, 1500 ft.,
1 cfl ? v.5.41 (R.P.).

Ojo de Agua, Vera Cruz, 1600 ft., 2 v.12.41 (R.P.).

Of these specimens only the El Mante specimen is as heavily
marked as the figures given by Roeber (Z.c.). Is Phycoides
mylitta mexicana Hall (p. 44) synonymous with guatemalena f

275. Phyciodes phaon phaon Edwards.

G. &S.,2: 677.

R., p. 436, pi. 89c.

Ha., p. 40.

H. , p. 677.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 2 2? vi.16-17.40 (H.H.).

Monterrey, Nuevo Leon, 1800 ft., 3 22 iv.27.41 (R.P.).

60 mi.*So. of Victoria, Tamaulipas, 1 2 vii.6.36 (H.D.T.).
Jacala, Hidalgo, 4500 ft., 1 vii.2.39 (H.H.).

The Jacala specimen and two of the Monterrey specimens are
typical phaon , the others are f. cestiva Edwards. The known
range is extended by the Jacala specimen.

276. Phyciodes picta pallescens Felder.

G. & S., 1 : 195, 2 : 678, pi. 21, ff. 18, 19.

R., p. 437, pi. 89e.

Ha., p. 50.

H. , p. 676.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 2 v.28.41
(R.P.).

El Sabino, Uruapan, Michoacan, 1 2 vii. 15-30.36
(H.D.T.).

The El Sol specimen establishes a new northern record for this
form on the east coast of Mexico, extending the range given by

Dec., 1944]

Brown: Butterflies

351

Hoffmann. Hoffmann lists pallescens (#276) as a distinct spe-
cies, separate from picta (#281).

277. Phyciodes tharos Drnry.

G. & S., 1 : 193, 2 : 436.

R., p. 436, pi. 89f.

Ha., p. 35.

H. , p. 676.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 1 2 $2 vi.17-18.40. (H.H.).

Galeana, Nuevo Leon, 6500 ft., 1 viii.2.39 (H.H.).

El Sol Tamazunchale, San Luis Potosi, 1 1 J v.28 &

iv. 31.41 (R.P.).

None of these is f. marcia Edwards.

280. Phyciodes vesta vesta Edwards.

G. & S., 1 : 195, 2 : 678.

R., p. 436, pi. 89c.

Ha., p. 50.

H. , p. 677.

Sabinas Hidalgo, Nuevo Leon, 960 ft., 3 1 5 vi.15.39

(H.H.).

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 1 J vi. 18.40 (H.H.).

Hda. Sta. Engracia, Nuevo Leon, 1 vii.25.39 (H.H.).

nr. Villagran, Tamaulipas, 2 55 iv.28.41 (R.P.).

60 mi. So. of Victoria, Tamaulipas, 1 5 vii.6.36 (H.D.T.).
El Mante, San Luis Potosi, 1 <$ iv.29.41 (R.P.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 1 5

v. 28.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 2 J'J1 vii.22.39 ;
1 vi. 27.40 (H.H.).

El Pujal, San Luis Potosi, 100 ft., 1 5 vii.21.39 (H.H.).
The San Luis Potosi specimens tend toward f. loucardi G. & S.
The Nuevo Leon specimens are almost typical vesta.

282. Phyciodes texana texana Edwards.

G. & S., 1 : 200.

R., p. 442, pi. 90c.

Ha., p. 96.

352

Journal New York Entomological Society

[Yol. lii

H., p. 677 (as Athanassa) .

Sabinas Hidalgo, Nuevo Leon, 690 ft., 1 vi.15.39

(H.H.).

Ojo de Agua, Sabinas Hidalgo, Nuevo Leon, 1000 ft., 1 g
vi. 14.40 (H.H.).

Hda. Vista Hermosa, Villa Santiago, Nnevo Leon, 1500
ft., 5 2 $? vi.16-18.40 (H.H.).

Monterrey, Nnevo Leon, 1600 ft., 2 25 iv.27.41 (R.P.).

Galeana, Nuevo Leon, 6500 ft., 1 5 vii. 30.39 (H.H.) ;
1 J1 iv.29.41 (R.P.).

nr. Villagran, Tamaulipas, 1160 ft., 1 1 2 iv.28.41

(R.P.).

60 mi. So. of Victoria, Tamaulipas, 1 § vii.6.36 (H.D.T.).

Jacala, Hidalgo, 4500 ft., 5 J'.J' 1 2 vi.23-vii.2.39 (H.H.).

This is the only member of this particular section of the
Phyciodes that is easily recognized. The broad orange-brown
area at the base of the forewing beneath is the characteristic
which at once identifies it.

285. Phyciodes ptolyca f . ptolyca Bates.

G. & S., 1 : 201 (in part), pi. 21, If. 32, 33.

Ha., p. 91.

H. , p. 677.

Galeana, Nuevo Leon, 6500 ft., 9 1 2 iv.29.41 (R.P.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 2 J'J'
3 $5 v. 29.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 1 $ vii.22.39 ;
1<? vi.26.40 (H.H.).

Hda. Potrero Viejo, Paraje Nuevo, Vera Cruz, 1500 ft.,
1 v.5.41 (R.P.).

Ojo de Agua, Vera Cruz, 1600 ft., 2 ££ v.12.41 (R.P.).

285A. Phyciodes ptolyca f. amator Hall.

Ha., p. 92.

60 mi. So. of Victoria, Tamaulipas, 1 vii.6.36 (H.D.T.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 5
iv.31.41 ; 1<? v.28.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 2 ££ 2 52
vi. 26-27.40 (H.H.).

Dec., 1944]

Brown: Butterflies

353

Hcla. Potrero Viejo, Paraje Nuevo, Vera Cruz, 1500 ft.,
1 J1 1 2 v.5.41 (R.P.).

Fortin, Vera Cruz, 1600 ft., 1 2 v.6.41 (R.P.).

Orizaba, Vera Cruz, 2000 ft., 1 2 v.6.41 (R.P.).

El Capirie, Michoacan, 500 ft., 1 2 viii.3.40 (H.H.).

El Sabino, Uruapan, Michoacan, 1 J1 vii.15-30.36
(H.D.T.).

Hall considers this the western race of ptolyca. I believe it is
only a pale form. Roeber, in Seitz’, confused this species, ptolyca ,
with a form of drusilla, telex. In general ptolyca is smaller than
either ardys or drusilla alethes with which it is easily confused.
I know of no good character for separating this species, but in
series there are several intangible differences.

285a. Phyciodes cortez Hall. ?

Ha., p. 93.

Ojo de Agua, Vera Cruz, 1600 ft., 1 2 v. 12.41 (R.P.).

I believe that I have correctly placed this species. The type
hails from Cuautla (4000 ft.) in S. W. Mexico. It may be a
white form of tulcis. This species, is not included by Hoffmann
in his list.

287. Phyciodes ardys Hewitson.

G. & S., 1 : 204, 2 : 681, pi. 22, ff. 4, 5.

R., p. 442, pi. 90d.

Ha., p. 90.

H. , p. 677.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1 g
12 vi.18.40 (H.H.).

Tuxpango, Vera Cruz, 1500 ft., 2 J'J1 v.9.41 (R.P.).
Fortin, Vera Cruz, 1600 ft., 1 1 2 v.4.41 (R.P.).

Orizaba, Vera Cruz, 2000 ft., 3 % 22 v.6.41 (R.P.).

One pair in copida.

Cumbres, (Km. 295) Vera Cruz, 8000 ft., 1 2 v.7.41
(R.P.).

El Sabino, nr. Uruapan, Michoacan, 1 J* vii.15-30.36

(H.D.T.).

This species may be separated from the following by the band
on the upperside of the hindwing which is broken into spots on
ardys. The Michoacan record is an extension of the known range.

354

Journal New York Entomological Society

[Vol. LII

287A. Phyciodes drusilla alethes Bates.

G. & S., 1 : 201 (as ptolyca in part).

R., p. 443, 8c as lelex on pi. 90, f. d.

Ha., p. 88.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 J
v.28.41 (R.P.).

El Sabino, Uruapan, Michoacan, 4 J'J' 2 vii. 15-30.36

(H.D.T.).

This race of drusilla closely resembles ptolyca. It differs from
that species in the following respects: the light markings on the
upper side are duller; the submarginal markings on the upper
side of the forewings are more frequently complete ; the underside
of the hindwing is less contrasty. Hall states (p. 92) that the
most important character is 1 1 the enlargement of the submarginal
spot at the anterior angle of hindwing above’ ’ on ptolyca. I find
this to be true in about 80% of my specimens.

This species is not included by Hoffmann in his list. He may
have confused it with ptolyca.

290. Phyciodes atronia atronia Bates.

G. & S., 1 : 202, 207, 2 : 681, pi. 22, ff. 19-23 (as atronia,

argentea, cassiopeia) .

R., p. 441, pi. 90a (as obscurata & cassiopeia ) p. 443, pi.

88i (as atronia & argentea).

Ha., p. 108.

H. , p. 677 (as Athanassa) .

Ojo de Agua, Yera Cruz, 1600 ft., 3 v.12.41 (R.P.).

I have followed Hall ( l.c .) in respect to this very variable spe-
cies. My specimens are what Roeber (l.c.) called cassiopeia.

293. Phyciodes myia myia Hewitson.

G. & S., 1: 188,2: 674.

R., p. 444, pi. 90e.

Ha., p. 136.

H. , p. 678 (as Eresia).

Tuxpango, Yera Cruz, 1500 ft., 2 v.9.41 (R.P.).

Fortin, Vera Cruz, 1600 ft., 2 1 ? v.4.41 (R.P.).

Ojo de Agua, Yera Cruz, 1600 ft., 7 $$ 5 5? v.12.41

(R.P.).

Dec., 1944]

Brown: Butterflies

355

El Sabino, Uruapan, Michoacan, 1 2 vii.15-30.36
(H.D.T.).

These are all typical myia. Hall records a single specimen of
the slightly different race griseobasalis Bates from Western
Mexico, Sierra Madre de Tepee. My single female from Michoa-
can does not show any indication of being that race.

297. Phyciodes eranites Hewitson.

G. &S., 1:185, 2:673.

R., p. 446, pi. 92c & d (as evanites).

Ha., p. 164.

H, p. 678.

Fortin, Vera Cruz, 1600 ft., 1 J' v.4.41 (B.P.).

Ojo de Agua, Vera Cruz, 1600 ft., 1 § v.12.41 (R.P.).

298. Phyciodes phillyra Hewitson.

G. & S., 1 : 184, 2 : 674.

R., p. 448, pi. 92c.

Ha., p. 167.

H. , p. 678.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 2
iv.31.41 (R.P.).

Tuxpango, Vera Cruz, 1500 ft., 1 J' 2 22 v.9.41 (R.P.).
Ojo de Agua, Vera Cruz, 1600 ft., 1 £ v.24.41 (R.P.).
Orizaba, Vera Cruz, 2000 ft., 2 v.6.41 (R.P.).

The Tuxpango females are much larger and the coloring less
intense than on the El Sol female. Not previously reported from
San Luis Potosi.

299. Chlosyne janais Drury.

G. &S., 1:178,2:671.

R., p. 451, pi. 91f.

H. , p. 678.

Ojo de Agua, Sabinas Hidalgo, Nuevo Leon, 1000 ft.,
1 J1 vi.15.40 (H.H.).

Sabinas Hidalgo, Nuevo Leon, 960 ft., 1 2 vi. 15.40
(H.H.).

Hda. Vista Hermosa, Nuevo Leon, 1500 ft., 1 2 vi. 17.40
(H.H.).

356

Journal New York Entomological Society

[Vol. Lil

Galeana, Nuevo Leon, 6500 ft., 6 J'J' iv.29.41 (R.P.).

Nr. Villagran, Tamaulipas, 1160 ft., 1 J iv.28.41 (R.P.). '
Arroyo del Meco, Tamaulipas, 1320 ft., 3 J'J' iv.28.41
(R.P.).

Jacala, Hidalgo, 4500 ft., 1 J' 1 ? vi.24.39 (H.H.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 J'
v.28.41 (R.P.).

Ojo de Agua, Vera Cruz, 1600 ft., 2 J'J' 4 v. 12.41
(R.P.).

300. Chlosyne lacinia Geyer.

The multitude of forms of this species are so hopelessly con-
fused and the names proposed for them so numerous that I am
unwilling to put names to any without much more material and
study. Therefore I shall merely list the specimens under brief
descriptions.

a) rufous markings on the upper side of both wings, those of the
forewing forming a more or less continuous band as in G. & S.
pi. 19, f. 8 & 10 : the underside of the hindwings as in f. 7 : under-
side of forewings as in f. 13 with added basal spots. This form
I believe is best called adjutrix Scudder.

Sabinas Hidalgo, Nuevo Leon, 960 ft., 1 J vi. 15.39
(H.H.).

Monterrey, Nuevo Leon, 1800 ft,, 3 J'J1 1 2 iv.27.41
(R.P.).

Jacala, Hidalgo, 4500 ft., 1 vii.2.39 (H.H.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 2 J'J1
iv.31 & 3 22 v.28.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 2 vi.26-

27.41 (H.H.).

El Sabino, Uruapan, Miehoacan, 1 £ vii. 15-30.36
(H.D.T.).

The Miehoacan specimen is atypical to this extent : the basal
spots are absent and the transverse bands are chestnut, narrow
and ill-defined on the upper surface.

5) similar to a) but lacking the yellow marginal lunules on the
underside of the hindwings.

Jacala, Hidalgo, 4500 ft., 1 2 vii.2.39 (H.H.).

Dec., 1944]

Brown: Butterflies

357

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 2

iv. 31.41 (R.P.).

c) similar to a) on the upper side but lacking the brown mark-
ings on the forewing. Underside of hindwing with orange
brown spots outside of yellowish buff transverse band — see Seitz’
pi. 91d — mediatrix.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 $

v. 28.41 (R.P.).

Fortin, Vera Cruz, 1600 ft., 2 v.4.41 (R.P.).

Orizaba, Vera Cruz, 2000 ft., 1 $ v.6.41 (R.P.).

Rio Blanco, Vera Cruz, 2200 ft., 1 £ v.10.41 (R.P.).

d) similar to c) but lacking the series of orange brown spots on
the underside of the hindwings.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 4
2 5$ v.28-29.41 (R.P.).

Tuxpango, Vera Cruz, 1500 ft., 1 £ v.9.41 (R.P.).
Fortin, Vera Cruz, 1600 ft., 1 g v.4.41 (R.P.).

Orizaba, Vera Cruz, 1600 ft., 1 <£ v.6.41 (R.P.).

e) similar to c) but brown area on hindwing above diffuse and
reduced.

El Sabino, Uruapan, Michoacan, 1 2 vii. 15-30. 36

(H.D.T.).

/) similar to d) and varying from that as e) does from c).

El Sabino, Uruapan, Michoacan, 1 2 vii.15-30.36

(H.D.T.).

g) similar to c) on the upper side but brown band reduced to a
series of small spots ; beneath lacking the yellow transverse band
on the hindwing.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 2 22
v.28.41 (R.P.).

Fortin, Vera Cruz, 1600 ft., 1 2 v.4.41 (R.P.).

h) upper surface lacks all trace of brown : underside with yellow
submarginal lunules, transverse band and basal spots on the
hindwing ( quehtala Reakirt?).

358

Journal New York Entomological Society

[Vol. Lll

Iguala, Guerrero, 1 J1 v.27.41 (R.P.).

El Sabino, Uruapan, Michoacan, 1 $ 1 2 vii.15-30.36
(H.D.T.).

i) similar to h) but lacking the yellow on the underside of the
hind wings ( adelina Stdgr. f).

Rio Balsas, Guerrero, 2400 ft., 2 v.26.41 (R.P.).

Apatzingan, Michoacan, 1050 ft., 1 ^ 1 J in copula
viii.2.40 (H.H.).

El Capirie, Michoacan, 500 ft., 1 g viii.3.40 (H.H.).

j) white spotted forewing, hindwing with a light area extended
at least half way from the base to the outer margin which it
parallels. The area is not solidly colored but varies from buff
to orange brown. Under side like the upper side ( lacinia
Hiibner?).

Galeana, Nuevo Leon, 6500 ft., 1 J* iv.29.41 (R.P.).

60 mi. So. of Victoria, Tamaulipas, 1 J' 1 J vii.6.36
(H.D.T.).

El Banito, Valles, San Luis Potosi, 200 ft., 2 J'J' 1 J
vi. 26-28. 40 (H.H.).

303. Chlosyne hippodrome Geyer.

(figured as hyperia by Seitz ’ pi. 91f.).

H., p. 679.

Ojo de Agua, Vera Cruz, 1600 ft., 6 <£$ 5 22 v.12.41
(R.P.).

Chichen Itza, Yucatan, 2 viii.30.36 (H.D.T.).

Acahuato, Michoacan, 1 J1 viii.2.40 (H.H.).

El Sabino, Uruapan, Michoacan, 1 2 vii.15-30.36
(H.D.T.).

There is no difference between east and west coast specimens.
Hoffmann does not record this species from Michoacan.

305. Chlosyne erodyle Bates.

G. & S, 1 : 180, 2 : 672, pi. 20, ff. 3, 4.

R, p. 452, pi. 91h.

H, p. 679.

Rio Blanco, Vera Cruz, 2200 ft., 1 <$ v.10.41 (R.P.).

Dec., 1944]

Brown: Butterflies

359

307. Chlosyne dryope Godman & Salvin.

G. & S., 2 : 672, pi. 108, ff. 3, 4.

H. , p. 679.

Iguala, Guerrero, 1 v.22.41 (R.P.).

Rio Balsas, Guerrero, 2400 ft., 1 2 v.26.41 (R.P.).
Hoffmann records this species only from Jalisco.

311. Chlosyne endeis Godman & Salvin.

G. & S., 2 : 673, pi. 108, ff. 5, 6.

H. , p. 679.

Jacala, Hidalgo, 4500 ft., 3 1 2 vi.23-vii.6.39 (H.H.).

The transverse discal band on a fresh specimen is fulvous. On
the other three which are flown it appears white. This seems to
be due to a great extent to the loss of the fulvous scales which
seem to be very loosely attached.

This is the first record of this species on the eastern slope of
Mexico.

314. Microtia elva Bates.

G. & S., 1 : 120, 2 : 682, pi. 20, f. 23.

R., p. 453, pi. 88h.

H. , p. 669-70.

Sabinas Hidalgo, Nuevo Leon, 960 ft,, 1 £ vi.18.39
(H.H.).

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 2 $$ vi.17-20.40 (H.H.).
nr. Villagran, Tamaulipas, 1160 ft., 1 J* 1 $ iv.28.41
(R.P.).

Arroyo del Meco, Tamaulipas, 1320 ft., 3 <$<$ iv.28.41
(R.P.).

Jacala, Hidalgo, 4500 ft., 6 5 22 vi.23-29.39 (H.H.).

So. of El Mante, San Luis Potosi, 1 5 iv.29.41 (R.P.).
El Banito, Valles, San Luis Potosi, 200 ft., 1 2 vii.22.39 ;
2 <$<$ 1 2 vi. 26-27. 40 (H.H.) ; 2 1 2 iv.30.41

(R.P.).

Acapulco, Guerrero, 100 ft., 1 2 v.26.41 (R.P.).

Iguala, Guerrero, 1^12 v.22.41 (R.P.).

Acahuato, Michoacan, 3000 ft., 1 2 viii.2.40 (H.H.).

360

Journal New York Entomological Society

[Vol. Lll

Apatzingan, Michoacan, 1050 ft., 1 J viii.2.40 (H.H.).
El Capirie, Michoacan, 500 ft,, 1 J1 viii.3.40 (H.H.).
One of the males from El Banito, iv.30.41, and the Sabinas
Hidalgo male may be termed f. horni Rebel. As a matter of fact
these two specimens are closer to f. draudti Rober which I con-
sider to be merely an intermediate form lying between elva and
f. horni.

The west coast females tend to be generally larger than those
from the east coast, 18 mm. vs. 16 mm. baso-apical radius of
the forewing. The males are about the same size from both
coasts.

315. Morpheis ehrenbergii Huebner.

G. & S., 1 : 211, 2 : 682.

R., p. 454, pi. 93a.

H. , p. 670.

Jacala, Hidalgo, 4500 ft., 3 jy* 10 vi.23-vii. 24.39
(H.H.).

Tancitaro, Michoacan, 6586 ft., 2 1 J vii.20-30.40

(H.H.).

Hoffmann carefully listed the States from which this local
species has been reported. Neither of the two here noted occurs
in his list.

BIBLIOGRAPHY

1. Godman & Salvin, Biologia Centrali-americana, 1: 143-211, 1881-1882;

2: 667-682, 1901.

2. Seitz, Macrolepidoptera of the World, 5: 357-433, 1911-1913.

3. Hoffmann, Anales del Instituto de Biologia, 11: 639-738, 1940.

4. Hall, Supplement to Bull. Hill Museum, 2-4: 206 pp., 1928-1930.

5. Roeber, in Seitz’ Macrolepidoptera of the World, 5: 434—454, 1913-1914.

Dec., 1944]

Huckett: MuscidvE

361

A REVISION OF THE NORTH AMERICAN GENUS
EREMOMYIOIDES MALLOCH (DIPTERA,
MUSCIDZE)

By H. C. Huckett
Riverhead, N. Y.

The native genus Eremomyioides was established by Malloch1
for the reception of three species, Pegomyia setosa Stein,
Eremomyia cylindrica Stein and a new form Eremomyioides
parkeri. The second species, cylindrica , was chosen as the geno-
type. Later Malloch2, 3 recorded two new species belonging to
the genus, E. fuscipes and E. similis, and published a key to the
known forms. There is now before me a sixth species which has
not been recognized, and also the male of parkeri.

The distinguishing characters of Eremomyioides are well
marked and distinctive. All the species possess setulae on all
pleural and sternal sclerites of thorax, including propleura,
pteropleura, hypopleura, prosternum and metasternum. In ad-
dition, all species lack cruciate setae on interf rontalia ; they possess
a bristle near middle of anteroventral surface of middle femora
and middle tibiae, and bear numerous stout short setulae on vi-
brissal angle. In the male the structure of hypopygium and
copulatory appendages is typical of all species, being more robust
in parkeri and setosa. The prebasal sclerite of hypopygium
(tegrum 6 of Crampton)4 has numerous bristles. The females
have two or more of the fore tarsal segments broadened. The
genus in my opinion finds its closest relationships in the major-
group of the genus Pegomyia ,5 and in Eremomyia as delimited
by Ringdahl.6

1 Malloch, J. R. 1918. Notes and descriptions of some anthomyid genera.
Proc. Biol. Soc. Wash., 31: 67-68.

2 Malloch, J. R. 1920. Descriptions of new North American Anthomyiidse
(Diptera). Trans. Amer. Ent. Soc., 46 : 182-183.

3 Malloch, J. R. 1921. Synopses of some North American Anthomyiinae
(Diptera). Can. Ent., 53: 76-77.

4 Crampton, G. C. 1941. The terminal abdominal structures of male
Diptera. Psyche, 48 : 94, fig. 20.

s Huckett, H. C. 1941. A revision of the North American species belong-

362

Journal New York Entomological Society

[Vol. LII

The adults of the species are commonly collected in the Spring
in the neighborhood of woodland terrain. From available infor-
mation it would seem that the species cylindrica is widely dis-
tributed across the continent from Alaska to Quebec. A single
specimen of similis has been taken at Boulder, Colorado, and of
fuscipes in the foothills of the Appalachian Mountains in Georgia.

Genus Eremomyioides Malloch

Eremomyioides Malloch, 1918, Proc. Biol. Soc. Wash., 31 : 67.
Curran, 1934, Fam. Gen. N. A. Dipt., p. 391. Seguy, 1937,
Gen. Insect., Fasc. 205, p. 122.

Eremomyoides Malloch, 1921, Can. Ent., 53: 76. Huckett, 1924,
N. Y. (Cornell) Agr. Exp. Sta., Mem. 77 (1923), p. 12.
Genotype: Eremomyia cylindrica Stein.

KEYS TO SPECIES
Males

1. Eyes separated at narrowest by a distance greater than twice that be-

tween posterior ocelli inclusive ; paraf rontals bristled to vertex, vertical
and paraorbital bristles robust; hypopygium prominently protruded
caudad, anal sclerite (tergum 9) longer than length of process on fifth

abdominal sternum 2

Eyes separated at narrowest by a distance not greater than twice that
between posterior ocelli inclusive; paraf rontals bare caudad, verticals
slender, paraorbital bristles lacking; hypopygium inconspicuous, not
protruding caudad, anal sclerite not longer than process of fifth ab-
dominal sternum ; 3

2. Hind femur with long, stiff bristles on proximal region of antero- and

posteroventral surfaces which are directed apicad, stouter than those

on distal half of anteroventral surface parkeri Mall.

Hind femur with bristles on proximal region of antero- and posteroventral
surfaces directed ventrad and not stouter than those on distal half of
anteroventral surface setosa (Stein)

3. Eyes separated by a distance not greater than that between posterior

ocelli inclusive; bristles of middle pair of presutural acrosticals sepa-
rated by a distance greater than that between eyes; hind tibia largely
reddish yellow, apical anterior bristle weak cylindrica (Stein)

ing to the genus Pegomyia. (Hiptera: Muscidse) Mem. Amer. Ent. Soc.,
No. 10, p. 14.

6 Ringdahl, O. 1933. Oversikt av i Sverige funna Hylemyia-arter. Ent.
Tidskr., Arg. 54, haft 1, p. 30.

Dec., 1944]

Huckett: Muscid^e

363

Eyes separated by a distance greater than that between posterior ocelli
inclusive; bristles of middle pair of presutural acrosticals separated by
a distance less than that between eyes 4

4. Hind tibia with 3 or 4 anterodorsal and 3 or 4 posterodorsal bristles.

conscripta n. sp.

Hind tibia with fewer bristles on one or other of dorsal surfaces, usually
with 2 anterodorsal and 2 posterodorsal bristles 5

5. Costal setulse scarcely as long as width of costa, serially conforming to

costal hairs ; thorax with 3 pairs of presutural acrostical bristles ; inner
margin of second antennal segment extending obliquely across base of
third segment and angularly projected on the lower half ...fuscipes Mall.
Costal setulse proximad of costal thorn slightly longer than width of costa,
more widely spaced apart than costal hairs ; thorax with two pairs of
presutural acrosticals; inner margin of second antennal segment ex-
tending transversely at right angles across base of third segment, and
not forming an angular prominence on lower half similis Mall.

Females

1. Fifth abdominal tergum obtusely rounded and swollen at apex, curved

ventrad, armed with a loose tuftlike series of bristles; opening to ovi-
positor situated on ventral surface of abdomen and basad of apex.

parlceri Mall.

Fifth abdominal tergum not rounded nor swollen on caudal region, armed
with a transverse series of marginal bristles; opening to ovipositor
situated at apex of abdomen 2

2. Mid and hind femora reddish yellow cylindrica (Stein)

Mid and hind femora largely blackish 3

3. Hind tibia with 3 or 4 anterodorsal and 3 or 4 posterodorsal bristles;

hind tibia largely reddish yellow 4

Hind tibia with fewer bristles on one or other of dorsal surfaces, usually
with 2 anterodorsal and 2 posterodorsal bristles; hind tibia largely
blackish tinged 5

4. Costal setulse robust and in a prominent series distad and proximad of

costal thorn, longest setulse about equal in length to humeral cross vein
and most of the setulse nearly twice as long as width of costa.

setosa (Stein)

Costal setulse weak, inconspicuous, scarcely longer than width of costa.

conscripta n. sp.

5. Inner margin of second antennal segment extending obliquely across base

of third segment and angularly projected on lower half; costal setulse

scarcely longer than width of costa | fuscipes Mall.

Inner margin of second antennal segment extending transversely at right
angles across base of third segment, and not forming an angular
prominence on lower half ; costal setulse longer than width of costa.

similis Mall.

364

Journal New York Entomological Society

[Vol. LII

Eremomyioides parkeri Malloch

Eremomyioides parkeri Malloch, 1918, Proc. Biol. Soc. Wash., 31 :
67. Seguy, 1937, Gen. Insect., Fasc. 205, p. 122.
Eremomyoides parkeri Malloch, 1921, Can. Ent., 53: 76 Strick-
land, 1938, Can. Jonr. Res., Sect. D, 16: 209.

The male resembles setosa, having eyes widely separated and
frons bristled to vertex. It may be distinguished from the latter
species by the bristles on proximal half of antero- and postero-
ventral surfaces of hind femur, which are coarser and stouter and
are directed apicad. The male has a stout apical bristle on an-
terior surface of hind tibia, but the bristle at middle of anterior
surface is very weak in the specimens before me, as is also the
lower posthumeral bristle. The fore femora lack the usual series
of longish setulae on median plane of anterior surface.

Alberta: J1, Medicine Hat, April 2, 1926 (F. S. Carr), allotype
[C.N.C.] . ?, Elk Island, May 16, 1937 (E. H. Strickland) [Univ.
Alberta]. §, Calgary, May 1, 1939 (W. S. McLeod).

Saskatchewan: Oxbow, May 11, 1907 (Fredk. Knab)

[U.S.N.M.] . Ogema, June 16, 1916 (N. Criddle) [C.N.C.].

Eremomyioides setosa (Stein)

Pegomyia setosa Stein, 1898, Berl. Ent. Zeitschr., (1897) 42, heft
3 & 4, p. 247. Aldrich, 1905, Misc. Coll. Smithsn. Inst., 46 :
558.

Eremomyioides setosa Malloch, 1918, Proc. Biol. Soc. Wash.,
31 : 67.

Eremomyia setosa Stein, 1919, Arch. f. Naturgesch., (1917) 83,
Abt. A, heft 1, p. 153. Seguy, 1937, Gen. Insect., Fasc. 205,

p. 122.

Eremomyoides setosa Malloch, 1921, Can. Ent., 53: 76, 77.

In both setosa and parkeri the hypopygium is much larger than
in other species belonging to the genus. The female of setosa has
all femora largely blackish, costal setulae robust, and hind tibiae
armed with 3 or 4 anterodorsal and 3 or 4 posterodorsal bristles.
Alaska: 2 ^, Katmai, June, 1917 (J. S. Hine).

Idaho: $, Moscow, cotype [Chicago Nat. Hist. Mus.].*
Juliaetta, May 11, 1902 [N.S.N.M.].

* Formerly the Field Museum of Natural History.

Dec., 1944]

Huckett: Muscid^e

365

Oregon: J, Meacham, May 8, 1927, 3680 ft. alt. (H. A. Scul-

len) [Ore. State Coll.].

Washington: J, Davenport, April 4, 1935 (J. Wilcox).

Eremomyioides cylindrica (Stein)

Eremomyia cylindrica Stein, 1898, Berl. Ent. Zeitschr., (1897)
42, heft 3 & 4, p. 226. Aldrich, 1905, Misc. Coll. Smithsn.
Inst., 46 : 554. Smith, 1910, Ann. Rept. N. J. State Museum
1909, p. 791. Johnson, 1913, Bull. Amer. Mus. Nat. Hist., 32,
Art. 3, p. 78. Stein, 1919, Arch. f. Naturgesch., (1917) 83,
Abt. A, heft 1, p. 153. Stein, 1920, Arch. f. Naturgesch.,
(1918) 84, Abt. A, heft 9, p. 73, 74.

Eremomyioides cylindrica Malloch, 1918, Proc. Biol. Soc. Wash.,
31 : 67. Seguy, 1937, Gen. Insect., Fasc. 205, p. 122.
Eremomyoides cylindrica Malloch, 1921, Can. Ent., 53: 76.
Huckett, 1924, N. Y. (Cornell) Agr. Exp. Sta., Mem. 77
(1923), p. 12. Johnson, 1925, Occas. Pap. Boston Soc. Nat.
Hist., 7 : 233. Johnson, 1925, Proc. Boston Soc. Nat. Hist.,
38 : 93. Leonard, 1928, N. Y. (Cornell) Agr. Exp. Sta., Mem.
101 (1926), p. 837. Strickland, 1938, Can. Jour. Res., Sect.
D, 16 : 209.

The male of cylindrica may invariably be distinguished by the
combination of characters given in the key. In this sex the second
antennal segment and tibiae range in color from yellowish or red-
dish to blackish. In the female the middle and hind femora and
all tibiae are reddish yellow, and the costal setulge, like those of the
male, are short and arranged in an inconspicuous series.

Alberta: §, Wabamun, April 24, 1939, J1, Edmonton, May 1,
1937, 3 5, Fawcett, May 8, 1934 (E. H. Strickland).

Massachusetts: J*, New Bedford, April 29, 1896 (Hough), co-
type [Chicago Nat. Hist. Mus.]. J, Wellesley, May 15, 1919 (J.
D. Tothill).

Michigan: 2 2, Ann Arbor, May 18, 1923 (J. S. Hine) [Ohio
State Mus. ] .

Minnesota: J, St. Anthony Park (Lugger), cotype [Chicago
Nat. Hist. Mus.].

New York: <^, Ithaca, May 15, 1914 [C.U.]. Patterson,
April 24, 1935, 5, Brewster, April 30, 1936 (H. Dietrich) [C.U.].
J1, $, Riverhead, Long Island, May 8, 1927.

366

Journal New York Entomological Society

[Vol. LII

Ohio: J', West Liberty, April 12, 1931 (R. B. Gordon). $,
Neotoma, Hocking County, March 28, 1932 (E. S. Thomas) [Ohio
State Mus.] .

Ontario : Strathroy, April 22, 1927 (H. F. Hudson). 5, Low
Bush, Lake Abitibi, June 8, 1925 (N. K. Bigelow) [C.N.C.]. J,
Jordan, June 18, 1926 (G. S. Walley). J, Fenelm Falls, May 27,
1927 (F. P. Ide).

Pennsylvania : §, Broomall, May 6, 1934 [Brigham Young
Univ.].

Quebec : ?, Aylmer, May 14, 1925 (G. S. Walley) . ?, Hull, May
12, 1925 (C. H. Curran) [C.N.C.].

Eremomyioides conscripta new species

Male, blackish, resembling fuscipes in habitus, second antennal segment
reddish along distal margin, parafacial pruinescence brownish, mesonotum
subshining, with a dorsocentral stripe and trace of sublaterals caudad of
transverse suture, abdomen grayish white pruinescent, dorsocentral vitta
gradually becoming broader caudad, legs blackish, hind tibiae with trace of
reddish tinge, calyptrae whitish, halteres purplish tinged.

Eyes separated by about width of third antennal segment, inner margin
of second antennal segment squarely transverse across base of third segment,
two pairs of presutural acrostical bristles, apical bristles of processes of fifth
abdominal sternum fine and slender, costal setulae not longer than width of
costa, inconspicuous ; fore tibia with 2 posteroventral bristles, mid tibia with
1 anteroventral, 1 anterior, 1 anterodorsal, 1 posterodorsal, 3 posterior bris-
tles, hind tibia with 2 or 3 anteroventral, 4 anterodorsal and 4 posterodorsal
bristles, 1 or 2 median anterior bristles, apical anterior bristle well developed.
Length 8 mm.

Female, as male, second antennal segment more broadly reddish, abdomi-
nal stripe less distinct, cross veins slightly infuscated, hind tibise largely
reddish, costal setulae fine, slightly longer than width of costa, tibiae bristled
as in male, fore tarsal segments 2, 3, 4 and 5 narrowly broadened, when com-
pared to those of mid tarsus, fourth fore tarsal segment fully twice as long
as wide. Length 7.5 mm.

Holotype and allotype : J1, §, Copper Mountain, British Colum-
bia, April 8, 1928, Betula occidentalis sap (G. Stace Smith)
[C.N.C.].

The species conscripta comes closest to fuscipes and similis,
from both of which it may usually be distinguished by the greater
number of bristles on hind tibia. In conscripta the costal setulae
are weak, the inner margin of second antennal segment does not

Dec., 1944]

Huckett: Muscidje

367

proceed obliquely across base of third segment, and there are
only two pairs of presutural acrostical bristles. The female of
conscripta has paler hind tibiae than in fuscipes and similis, and
the costal setulae are much weaker than those of setosa.

Eremomyioides fuscipes Malloch

Eremomyioides fuscipes Malloch, 1920, Trans. Amer. Ent. Soc.,
46 : 182. Frison, 1927, Bull. 111. Nat. Hist. Surv., 16, Art. 4,
p. 198. Seguy, 1937, Gen. Insect., Fasc. 205, p. 122.
Eremomyoides fuscipes Malloch, 1921, Can. Ent., 53 : 76.
Huckett, 1924, N. Y. (Cornell) Agr. Exp. Sta., Mem. 77
(1923), p. 12. Leonard, 1928, N. Y. (Cornell) Agr. Exp.
Sta., Mem. 101 (1926), p. 837.

The speces fuscipes and similis are closely allied, both having
the tibiae largely blackish in both sexes. However the hind tibiae,
and to a less extent the middle tibiae, do exhibit a more or less
obscure reddish tinge in certain specimens. In fuscipes the hind
tibia may, or as is more general, may not have a bristle at middle
of anterior surface. The costal setulae in both sexes are weak, and
the inner distal margin of second antennal segment is oblique in
its course across base of third segment, thereby forming an angu-
lar prominence or outline on lower half. In similis the costal
setulae are longer than width of costa, and inner margin of second
antennal segment is more nearly rectangular across base of third
segment.

Alberta: Edmonton, May 13, 1937 (E. H. Strickland).

Connecticut: J', South Meriden, March 15, 1915 (H. L. John-
son).

Georgia: J, Clayton, April 15-22, 1940 (H. C. Huckett).

Illinois: Urbana, March 18, 1918 (Frison and Malloch),

paratype [111. Nat. Hist. Surv.].

New York: J', Ithaca, March 25, 1917 (R. C. Shannon). J',
Fall Creek, Ithaca, April 24, 1922 (L. S. West). J', Cooper Ceme-
tery, Staten Island, March 17, 1918.

Ohio : 2, Columbus, March 27, 1907.

Pennsylvania: 2 J1, Hawley, April 20, 1936 (H. Dietrich).

South Carolina: 2, Clemson College, February 22, 1936 (D.
Dunavan).

368

Journal New York Entomological Society

[Vol. lii

Eremomyioides similis Malloch

Eremomyioides similis Malloch, 1920, Trans. Amer. Ent. Soc., 46 :
183. Frison, 1927, Bull. 111. Nat. Hist. Surv., 16, Art. 4, p.
198. Seguy, 1937, Gen. Insect., Fasc. 205, p. 122.
Eremomyoides similis Malloch, 1921, Can. Ent., 53: 76, 77.

The distinguishing characters and relationships of this species
have already been discussed in notes concerning conscripta and
fuscipes. The species has in error been recorded by me as occur-
ring in New York from specimens which I now regard as belong-
ing to fuscipes.

Alberta : J', Edmonton, May 13, 1937 (E. H. Strickland).
Colorado: J1, Campus, University of Colorado (Cockerell)
[N.S.N.M.] .

Illinois: 5, Tuscola, March 29, 1918, paratype [C.N.C.].
Urbana, Brownfield Woods, April 23, 1919, paratype [111. Nat.
His. Surv.].

South Dakota : Brookings, April 29, 1919 [Ohio State Mus.].

Wisconsin: J', Dane County, April 10, 1900 (F. M. Snyder).

Dec., 1944]

Alexander: Crane-Flies

369

RECORDS AND DESCRIPTIONS OF NEOTROPICAL
CRANE-FLIES (TIPULIDZE, DIPTERA), XVIII

By Charles P. Alexander
Amherst, Massachusetts

The preceding part under this title was published in March,
1944 (Journal of the New York Entomological Society, 52:
45-57). The species discussed at this time are chiefly from Costa
Rica, Panama and Bolivia, derived from sources that are men-
tioned under the individual species. The types of the novelties
are preserved in my private collection of these flies, except where
stated to the contrary.

Genus Tanypremna Osten Sacken

Tanypremna (Tanypremna) saltatrix new species.

Size large (length, male, over 30 mm.) ; mesonotal praescutum with disk
almost covered by four reddish brown stripes, the usual interspaces darkened
and impressed; thoracic pleura yellow, with a narrow interrupted transverse
girdle extending from the praescutum to the sternopleurite ; tibiae with a
conspicuous white ring about its own length before tip ; basal four tarsal
segments black with broad white tips; wings subhyaline, the veins seamed
with pale brown; cell Sc2 small and narrow, tending to become closed at
margin; male hypopygium with the dististyle provided with 13 or 14 small
black spines arranged in a single row.

Male. — Length about 33-34 mm. ; wing 18.5-19 mm. ; abdomen alone about
28-29 mm.

Frontal prolongation of head yellow; palpi greenish yellow. Antennas
with scape and pedicel yellow ; flagellar segments pale brown, the outer ones
darker; segments cylindrical, the verticils longer than the segments. Head
chiefly yellow sericeous, darker behind.

Pronotum conspicuously blackened above, paling to yellow on sides. Mes-
onotal praescutum with the disk almost covered by four reddish brown stripes
that are very narrowly separated by capillary black lines, the ones delimiting
the usual interspaces impressed ; a dark transverse line on praescutum extend-
ing from cephalic end of lateral stripe to the lateral margin, interrupting the
broad yellow lateral and humeral border ; scutal lobes reddish brown, divided
into two unequal areas by narrowly blackened borders and lines ; scutellum
testaceous brown ; mediotergite dark brown, the central portion behind some-
what paler; lateral border of mediotergite and dorsal portion of katapleuro-
tergite pale, the remainder of pleurotergite dark brown. Pleura yellowish
white, with a more or less interrupted transverse girdle extending from the

370

Journal New York Entomological Society

[Vol. LII

lateral praescutal border, as above described, crossing the dorsopleural mem-
brane, on the anepisternum and sternopleurite being broken into linear spots.
Halteres with stem obscure yellow, knob dark brown. Legs with coxae yel-
low, outer face of each with a brownish black stripe, broadest on posterior
coxae; trochanters yellow; fore and middle legs with femora yellow basally,
passing into dark brown, most intense at outer ends but preceded by a very
vague paler subterminal ring; tibiae black, with a relatively narrow but con-
spicuous white ring about its own length from tip; fore tibiae with extreme
bases vaguely whitened; basitarsi black with the tips white, on fore legs
involving about the distal fourth, on the middle legs a little less ; succeeding
three tarsal segments black with their tips broadly white; terminal segment
uniformly brownish black; posterior legs similar but with the white rings
on tibiae and basitarsi even wider, the latter including almost the outer third
of segment. Wings subhyaline, the small stigma dark brown; cell Sc and
seams along most of veins paler brown, these seams wider and more diffuse
on the posterior and caudal veins. Venation: Cell Sc2 very small and narrow,
in holotype closed at margin by approximation of veins Sc2 and -B1 + 2; Us
straight and oblique, in alignment with jS4+5; cell 1st M2 about one-half wider
than long, narrower at outer end; cell about twice its petiole; m-cu
subequal to basal section of M3+ 4; cell 2nd A relatively wide.

Abdomen elongate; tergites brown, on more proximal segments narrowly
ringed with obscure yellow ; outer segments more uniformly darkened ; pleural
membrane dark; sternites yellow, their posterior borders darkened; eighth
and ninth segments dark, the tips of basitarsi pale. Male hypopygium with
tergal lobes broadly and obtusely rounded. Basistyle elongate; mesal face
at near two-thirds the length with a conspcuious lobe that is provided with
about a dozen long pale setae. Dististyle provided with about 13 or 14 short
blackened spines arranged in a single row, with more than half placed on
the distal third of row ; inner branch of style obtuse, provided with abundant
short setae.

Habitat. — Costa Rica.

Holotype, Rivas, altitude 2,875 feet, January 1939 (Dean
Rounds). Paratype, Pedregoso, altitude 2,075 feet, January
1939 (Dean Rounds).

The nearest relatives of the present fly are the Brazilian
Tanypremna ( Tanypremna ) longissima (Enderlein) and T. (T.)
manicata Osten Sacken, both of which differ conspicuously in the
relative proportions and in details of coloration of body and legs.
These species have been keyed and described in an earlier paper
by the writer (Jour. N. Y. Ent. Soc., 22: 205-218, 1914).

Tanypremna (Tanypremna) clotho new species.

Mesonotum very high and gibbous, dark brown, the sides broadly and
abruptly yellow; pronotum narrowly darkened medially; pleura pale yellow;

Dec., 1944]

Alexander : Crane-Flies

371

mesosternum with a conspicuous black spot; legs black, tibiae with a broad
white subterminal ring ; tarsal segments one and two black basally, their tips
broadly white; segments three and four white; wings pale brown, the costal
region slightly darker; stigma and a cloud over cord darker brown; E2+3
sinuously bent ; cell 1st M2 large, square at base, narrowed outwardly ; petiole
of cell M1 about twice m ; cell 2nd A unusually narrow; abdomen elongate,
tergites chiefly dark brown, restrictedly patterned with obscure yellow; cerci
black.

Female. — Length about 41 mm. ; wing 20 mm. ; abdomen alone 36 mm.

Frontal prolongation of head short, almost white; nasus distinct; palpi
pale, tinged with greenish, the proximal end of the terminal segment darker.
Antennae with scape and pedicel pale, the basal flagellar segments greenish,
outer segments dark brown; flagellar segments elongate, with conspicuous
verticils. Front and anterior vertex silvery white ; occiput and the transverse
adjoining portions of vertex almost black, the remainder of vertex paling to
brown ; genae obscure yellow.

Pronotum conspicuously dark brown medially, the sides abruptly yellow.
Mesonotal praescutum almost uniform dark brown, the area strongly narrowed
in front to leave a very broad humeral and lateral area on either side; the
four praescutal stripes differentiated by impressed lines at the interspaces;
scutal lobes similarly dark brown, the median region sparsely yellow polli-
nose; scutellum testaceous brown, the parascutella darker; mediotergite
chiefly dark brown, the lateral and posterior borders pale to form a U-shaped
border; median region at base more reddened; pleurotergite dark brown, the
posterior angle above the halteres abruptly yellow. Mesonotal praescutum
high and gibbous. Pleura uniformly pale yellow; sternum pale except for a
very evident oval black area on mesosternum. Halteres relatively long, stem
brown, yellow at base, knob slightly brightened. Legs with coxae pale
greenish yellow, the posterior pair with a brown spot on caudal aspect;
trochanters greenish yellow, the middle pair blackened on posterior face;
fore and hind femora brown, paler basally, darker brown at and before tips,
middle femora uniform brownish black; tibiae brownish black, with a broad
white subterminal ring, narrowest on fore legs where it is only a little more
than twice the blackened apex, broad on middle and posterior tibiae, especially
the latter, where the white is approximately four times the blackened apex;
basitarsi black on proximal portion, the tip broadly white, involving about
the distal two-thirds of segment, slightly less extensive on middle legs ; tarsal
segment two black at base, the tip white, broadest on fore legs where about
the distal three-fourths to four-fifths of segment is included, narrower on
middle and hind legs where more than half the segment is whitened, more
extensively so on middle legs ; tarsal segments three and four greenish white ;
terminal segment pale brown; claws (female) simple. Wings with a pale
brownish tinge, the broad costal cell slightly darker, cell Sc even darker
brown; stigma small, ill-defined, darker brown; a restricted broAvn cloud on
cord and adjoining portions of outer radial veins; veins brownish black.
Venation: Es straight, subequal to E2+ 3 which is sinuously bent, narrowing

372

Journal New York Entomological Society

[Vol. Lll

cell Bx at its outer end; vein B1+2 weak and pale, lying close to the free tip
of Sc2, which, in turn, is about the same distance from the spur of Scx; cell
R3 narrowed at about two-thirds its length; basal section of Bi+S almost
lacking to punctiform, r-m likewise reduced; cell 1st M2 large, square at
base, narrowed outwardly; petiole of cell Mx about twice m; m-cu at near
two-thirds the length of 2kf3+4; cell 2nd A unusually narrow, its greatest width
(opposite anal angle) about as long as petiole of cell Mx or somewhat shorter
than m-cu.

Abdomen elongate, as shown by the measurements; basal tergites obscure
yellow, narrowly darkened laterally, more broadly so dorsally, the amount of
dark color increasing on outer segments ; vague paler rings at near midlength
of tergites two to five, inclusive ; posterior borders of segments slightly dark-
ened, the adjoining base of the succeeding segment restrictedly pale at the
sides; basal sternites greenish yellow, the outer segments darker; genital
shield brownish black. Ovipositor with long straight black cerci, their tips
narrowly rounded.

Habitat. — Brazil.

Holotype, J, Palmeiras, Estado Kio de Janeiro, November 6,
1940 (Lopes).

The most similar species is Tanypremna ( Tanypremna ) longis-
sima (Enderlein), likewise from southeastern Brazil. This dif-
fers in the larger size (Female, length 55 mm. ; wing 22.5 mm. ;
abdomen alone 48.5 mm.), more especially the longer abdomen in
proportion to the wing length. The latter fly is still not known
to me except from the original description. From this it is indi-
cated that the praescutum is directed more strongly forward over
the head, completely concealing the latter when viewed from
above. There are various discrepancies in color between the two
flies, including the ovipositor, legs and wings, as the distinctly
seamed veins of the outer fourth of the wing in longissima.

It is now very evident that there are numerous species of the
subgenus in Tropical America, with an apparent concentration of
forms in southeastern Brazil. The venation and the basic plan of
structure of the male hypopygium indicates that the supposedly
very distinct generic groups, Ozodicera Macquart, Longurio Loew
and Tanypremna Osten Sacken are, in reality, all closely related
and may well be found to pertain to a single major generic group.
A very comparable case is found in Limonia Meigen, which rather
curiously parallels the present condition. Thus, the conspicu-
ously branched antennas of Ozodicera- are suggested by one of the
subgeneric groups of Tanypremna, Tanypremnodes Alexander,

Dec., 1944]

Alexander : Crane-Flies

373

where the flagellar pectinations while short are perfectly distinct.
The comparable condition in Limonia is to be found in the sub-
genera Bhipidia Meigen, Idioglochina Alexander, and Zelando-
glochina Alexander. Furthermore, the delicate, ethereal build
of species of Tanypremna, as compared to the stouter and more
massive condition in Ozodicera and Longurio, is closely paralleled
by various subgeneric groups of Limonia, as Thrypticomyia Skuse,
Doaneomyia Alexander, and Euglochina Alexander. It is of in-
terest to note that the three typical Tipuline groups listed above
have been placed in various tribes and subtribes within the
Tipulinse. The entire subject of genera and subgenera in the
more primitive Tipuline crane-flies requires much further con-
sideration and probably will result in a further reduction in the
number of supposedly valid generic groups.

Genus Holorusia Loew

Holorusia (Holorusia) furcifera new species.

General coloration of mesonotal praescutum testaceous brown, with four
more reddish brown stripes that are narrowly bordered with darker, especially
the intermediate pair; antennae (male) relatively long, about one-third the
length of body or wing; basal flagellar segments bicolored; thoracic pleura
uniformly pale yellow; femora yellow, the tips narroAvly brownish black;
wings weakly infuscated, with a more whitish central streak as far as the
cord; costal border, outer radial field, m-cu and a spot in cell R brown; cell
1st M2 small, irregularly hexagonal; abdomen yellow, with a broad sub-
terminal, dark brown ring; male hypopygium with the outer dististyle nar-
row, at near midlength bearing a slender lateral branch.

Male.— Length about 14 mm.; wing 15 mm.; antenna about 5 mm.

Frontal prolongation of head obscure yellow, darker on ventral half ; nasus
long and conspicuous; palpi black, the terminal segment paling to brownish
yellow. Antennae (male) about one-third the length of body; scape, pedicel
and first flagellar segment light yellow; succeeding segments bicolored, the
proximal end darkened, with about the distal two-thirds of the more basal
segments yellow, the latter color decreasing in amount on the outer segments,
on the eighth and remaining segments becoming uniformly brown; flagellar
segments cylindrical. Head medium brown, the front and very narrow orbits
yellowish gray ; a narrow but conspicuous dark brown median stripe on vertex.

Pronotum obscure brownish yellow. Mesonotal praescutum testaceous
brown, with four more reddish yellow stripes, the intermediate pair more
distinct, margined by narrow, slightly darker brown lines; humeral region
slightly darkened, especially internally; scutal lobes chiefly reddish yellow,
more obscure laterally; a narrow, darker brown, median vitta that extends
onto the extreme cephalic portion of scutellum, the latter pale yellow; para-

374:

Journal New York Entomological Society

[Vol. Lll

scutella more infuscated ; postnotum light yellow, the lateral border of
mediotergite and the dorsal and posterior borders of pleurotergite somewhat
darker. Pleura and most of pleurotergite clear light yellow. Halteres dark
brown, the extreme base of stem narrowly yellow. Legs with coxae and
trochanters clear light yellow; femora and tibiae yellow, their tips narrowly
brownish black, the amount subequal on all legs; tarsi passing into black.
Wings with the ground color weakly infuscated, variegated with conspicuous
darker brown and restricted whitish subhyaline areas; the darker pattern
includes especially the costal border, involving the prearcular field; outer
radial field as far caudad as vein -E4+5; arcular areas; origin of Bs ; m-cu,
and a conspicuous spot in cell B beyond midlength of vein M; a whitened
area in center of wing as far as cord, especially conspicuous across bases of
anal and cubital cells, M and outer end of cell B, crossing the base of cell
1st M2; veins brown. Posterior border of wing with scattered black setae
interspersed with the marginal fringe, as common in genus. Venation: Bs
a little less than twice m-cu, angulated at origin; vein Bi+5 strongly arcuated
at near midlength, constricting cell _B3; cell 1st M2 small, irregularly hexag-
onal in outline; m-cu close to midlength to vein Ms+4; cell Mx subequal in
length to its petiole.

Abdomen yellow, the three subterminal segments brownish black; hypo-
pygium yellow. Male hypopygium with the tergite narrowed outwardly, the
lateral angles produced caudad into relatively narrow lobes, the mesal faces
of which bear numerous black spinous setae; median area of tergite notched,
at base of emargination produced into a shorter lobe. Outer dististyle nar-
row, at near midlength bearing a conspicuous slender lateral branch. Inner
dististyle with crest high, obtusely rounded ; beak broad and obtuse.

Habitat. — Costa Rica.

Holotype, J', Pedregoso, altitude 2,075 feet, January 1939
(Dean Rounds).

The present fly is most similar to species such as Holorusia
( Holorusia ) strangalia Alexander, differing in the small size,
details of coloration, and structure of the male hypopygium. The
conformation of the outer dististyle is noteworthy.

Genus Tipula Linnaeus

Tipula (Eumicrotipula) votiva new species.

General coloration of mesonotum brownish gray, the prsescutum with three
brown stripes ; pleura yellow dorsally, dark brown beneath ; frontal prolonga-
tion of head yellow above, brownish black beneath ; antennal flagellum black ;
femora with a narrow yellow subterminal ring; wings strongly and almost
uniformly infuscated, restrictedly patterned with dark brown and obscure
yellow.

Male. — Length about 8 mm.; wing 8.7-9 mm.; antenna about 3. 8-3.9 mm.

Frontal prolongation of head yellow above, abruptly brownish black on

Dec., '1944]

Alexander: Crane-Flies

375

sides and beneath; nasus distinct, yellow; palpi dark brown, the incisures
pale. Antennae (male) of moderate length; scape blackened on basal third
to half, the remainder yellow ; pedicel and flagellum black ; flagellar segments
scarcely incised, subcylindrical ; verticils shorter than the segments. Head
brown, more pruinose on front.

Pronotum yellow, the extreme lateral portions dark brown. Mesonotal
praescutum brownish gray with three brown stripes that are best-delimited
on posterior half of sclerite, especially the broad median stripe; lateral por-
tions of praescutum darkened, including a velvety black spot on margin be-
hind pseudosutural foveae; scutum dark brown; scutellum and postnotum
light brown, sparsely pruinose. Pleura dark brown ventrally, the color in-
cluding the propleura, almost all of sternopleurite, and the meral region;
dorsal pleurites, including the dorsopleural membrane, conspicuously yellow;
dorsal anepisternum, posterior portion of sternopleurite and virtually all of
the pteropleurite yellow; pleurotergite blackened, especially posteriorly.
Halteres with stem yellow, knob brownish black. Legs with coxae brownish
black, sparsely pruinose ; trochanters yellow ; femora brownish black, brighter
basally, with a narrow yellow subterminal ring that is placed more than its
own length before the black tip; tibiae and tarsi black. Wings with the
ground very extensively and almost uniformly infuscated, sparsely patterned
with obscure yellow and darker brown areas; cell C brown, slightly bright-
ened subbasally; cell Sc yellow, with four major dark areas that are more
extensive than the pale interspaces; more or less distinct creamy areas in
prearcular field, beyond arculus in proximal ends of basal cells and as a very
faintly indicated area beyond stigma in outer radial field; veins brown, yel-
low in the flavous portions. No macrotrichia in cells of wing. Venation:
K1+2 entirely atrophied; R2 oblique, in direct alignment with beyond the
free tip of Sc2’, petiole of cell M1 exceeding m; cell 2nd A narrow; venation
of medial field very different in the two available specimens; in holotype
normal for the subgenus, that is, with M3+4 present and with m^cu beneath
cell 1st M2) in the paratype, the venation is almost exactly as in the genus
Nephrotoma, with m-cu uniting with vein M4 just before the base of cell
1st M2.

Abdomen with basal tergites obscure yellow, beyond the first with the
lateral borders conspicuously blackened, greatly restricting the amount of
yellow; outer segments, including hypopygium, uniformly brownish black.
Male hypopygium with the caudal margin of tergite nearly transverse, with
a small U-shaped median notch, the lateral portions of the latter slightly
produced into small lobes; lateral portions of tergal plate with long abun-
dant setae.

Habitat. — Bolivia.

Holotype, Buenavista, Santa Crnz (J. Steinbach). Para-
topotype, J1.

The most similar described species is Tipula ( Eumicrotipula )
fatidica Alexander, of Ecuador, which differs conspicuously in

376

Journal New York Entomological Society

[Yol. lii

the larger size, coloration of the body and wings, the short an-
tennae, and in the details of venation, as the much wider cell
2nd A .

Tipula (Micro tipula) epione new species.

Allied to smithi; mesonotal praescutum and scutum almost uniformly red-
dish brown; antennae (male) elongate, yellow, the flagellar segments weakly
bicolored ; halteres inf uscated ; legs yellow ; wings weakly infuseated, the
ground color of outer half of wing more whitish hyaline, heavily patterned
with brown, including the broad costal border and outer radial field; B1+s
entire; male hypopygium with a single dististyle, its inner angle bearing two
or three strong black spinous setae; lobe of ninth sternite a depressed-flat-
tened quadrate plate, with its apex truncated; eighth sternite with posterior
border evenly rounded.

Male. — Length about 12 mm. ; wing 12 mm. ; antenna about 7.5 mm.

Frontal prolongation of head brown; nasus stout; palpi brown, the outer
segments a trifle darker. Antennae (male) elongate, exceeding one-half the
length of wing ; basal three segments yellow ; succeeding segments very
weakly bicolored, brownish yellow, the small basal enlargements a little
darker brown; segments almost cylindrical; verticils much shorter than the
segments. Head brownish gray; anterior vertex at widest point a little
exceeding twice the diameter of scape, more narrowed behind between the
large eyes.

Praescutum and scutum almost uniformly reddish brown, without markings ;
scutellum and postnotum more testaceous yellow, parascutella darker. Pleura
obscure yellow. Halteres infuseated. Legs with the coxae and trochanters
yellow; remainder of legs yellow, the outer tarsal segments infuseated; claws
somewhat weakly toothed. Wings with the ground color weakly infuseated,
more whitish hyaline in outer half, including vicinity of cord and outer medial
field ; a very heavy and conspicuous brown pattern, including the broad costal
border which involves the prearcular region, cells C and Sc, all of B except-
ing the outer end, and virtually the whole outer radial field; conspicuous
brown seams on m-cu and narrower ones at outer end of cell 1st M2 and fork
of M1+2; veins pale, darker in the patterned areas. Squama with setae.
Venation: B1 + 2 entire; Bs relatively long, about one-third longer than m-cu,
the latter at near midlength of M3+4; basal section of Bi+5 very long, greatly
reducing r-m; cell 1st M2 relatively long; cells Jfi and 2nd A wide.

Basal abdominal segments obscure yellow, beyond the second passing into
brown; subterminal segments blackened, forming a narrow ring; hypo-
pygium yellow. Male hypopygium with the ninth tergite long, narrowed
posteriorly, the apex terminating in a broad flattened lobe that is gently
emarginate, to produce two lobules, both set with conspicuous black setae;
ventral tergal appendage (tegmen) pale and fleshy. Dististyle single, ap-
pearing as a subquadrate plate, the outer angle produced into a conspicuous
flattened obtuse lobe, its apical border narrowly blackened; inner angle less

Dec., 1944]

Alexander: Crane-Flies

377

produced, at apex bearing two or three long black spinous setae, the largest
of which is about two-thirds the total length of the style itself. Lobe of
ninth sternite a depressed-flattened subquadrate plate, slightly widened out-
wardly, its apex truncated or with vague indications of a median notch, the
surface densely setiferous. .ZEdeagus stout, terminating in a decurved hook-
like projection. Eighth sternite extensive, the posterior border broadly
rounded, provided with numerous long setae.

Habitat. — Bolivia.

Holotype, <$, Buenavista, Santa Cruz (J. Steinbach).

The present fly is most similar to Tipula (Microtipula) in-
temperata Alexander, T. (M.) smithi Alexander, and T. ( M .)
temperata Alexander, differing from all in the structure of the
male hypopygium. The wing pattern is generally similar in all
of these species yet they differ among themselves in the presence
or absence of major setae on the squamae. In other subgeneric
groups of Tipula the squamal character has been deemed to be of
primary importance in differentiating groups.

Tipula (Microtipula) nicoya new species.

Allied to perangustula ; size small (wing, male, about 10 mm.) ; mesonotal
praescutum obscure brownish yellow with three black stripes; thoracic pleura
yellow, with a narrow, transverse, black girdle occupying the mesepisternum ;
knob of halteres blackened; claws of male simple; wings with cells basad of
cord brownish yellow, the prearcular and costal fields, together with the outer
radial cells, strongly darkened; male hypopygium with the lateral lobes of
both the ninth tergite and the eighth sternite broadly and obtusely rounded ;
outer dististyle a simple cylindrical blackened rod; aedeagus not subtended
by a pair of spines.

Male. — Length about 9 mm.; wing 10 mm.; antenna about 4 mm.

Frontal prolongation of head yellow ; nasus short and stout ; palpi brown.
Antennae relatively long, as shown by the measurements; scape and pedicel
yellow; first flagellar segment yellowish brown; succeeding segments black,
the incisures of the more proximal ones vaguely brightened; segments only
weakly incised; verticils shorter than the segments. Head light brown;
anterior vertex wide, approximately four times the diameter of scape, without
tubercle.

Pronotum obscure yellow laterally, infuscated on central portion. Meso-
notal praescutum with the restricted ground obscure brownish yellow, the sur-
face polished, with three black stripes, the median one relatively narrow ; pos-
terior sclerites of notum almost uniformly blackened, the parascutella paler;
pleurotergite yellow. Pleura yellow, with a conspicuous transverse brown
girdle, extending from the cephalic end of the praescutal stripe which bends
laterad to the margin, crossing the dorsopleural membrane and covering most
of the mesepisternum. Halteres with stem yellow, knob blackened. Legs

378

Journal New York Entomological Society

[Vol. Lir

with coxae and trochanters yellow; femora obscure yellow, the tips rather con-
spicuously blackened ; tibiae obscure yellow, the tips narrowly darkened ; tarsi
yellowish brown to obscure yellow; claws simple. Wings with the cells basad
of cord brownish yellow; prearcular field, cells C and Sc, stigma, and cells
beyond anterior cord conspicuously infuscated, the last slightly less intense
than the others; proximal portions of cells distad of posterior cord and very
indistinct markings in bases of cells and B5 brighter in color. Venation:
B1+2 entire but the distal end faint and without trichia; Bs relatively short,
arcuated at origin, subequal to m-cu; cell 2nd A very narrow, about as in
perangustula.

Abdominal tergites obscure yellow on their basal portions, the caudal mar-
gins broadly black, especially on segments two and three, on the succeeding
tergites the amount of black somewhat more restricted ; lateral tergal borders
darkened; a subterminal black ring; sternites chiefly yellow; hypopygium
yellow. Male hypopygium in its general features much as in perangustula
but differing in all details. Ninth tergite with the lateral lobes broadly and
obtusely rounded, with a conspicuous median brush of setae, in addition to
the usual paired ventral tufts; these setae are much more elongate and less
spinous than in allied forms. Outer dististyle a simple, slender, cylindrical,
blackened rod, its tip obtuse; setae relatively short and inconspicuous. In
perangustula, the style on distal half expanded into an elongate-triangular
head, with long coarse setae. iEdeagus without a pair of subtending slender
spines, as in perangustula. Eighth sternite with the caudal margin con-
spicuously bilobed, each lobe obtusely rounded and provided with numerous
very long pale setae.

Habitat. — Costa Rica.

Holotype, J', Pedregoso, altitude 2,300 feet, January 1939
(Dean Rounds).

The specific name is that of an Amerind tribe inhabiting Costa
Rica. The most similar described species is Tipula ( Microti-
pula ) perangustula Alexander, of Colombia and Venezuela, which
differs especially in the hypopygial characters, as compared
above.

Genus Limonia Meigen

Limonia (Limonia) curraniana new species.

General coloration brown; anterior vertex silvery; antennal scape yellow,
the remainder of organ black; mesonotum brown, with a conspicuous orange
dorsomedian stripe; legs dark brown, the tips of the tibiae and all tarsi with
the exception of the terminal segment and proximal portions of basitarsi of
fore and middle legs, white ; wings strongly tinged with brown ; ventral disti-
style of male hypopygium without rostral spines.

Male. — Length about 5 mm. ; wing 6-6.2 mm.

Dec., 1944]

Alexander : Crane-Flies

379

Mouthparts very reduced; palpi small, black. Antennae with the scape
pale yellow, the remainder of organ black; flagellar segments gradually nar-
rowed and more slender toward end, the terminal segment about one-half
longer than the penultimate segment. Anterior vertex silvery, concave, the
anterior vertex several times as wide as diameter of scape ; posterior portions
of head light brown.

Mesonotum brown, with a conspicuous orange dorsomedian stripe, extend-
ing from the anterior portion of the praescutum to the base of the medioter-
gite. Pleura yellow. Halteres dark brown. Legs with the coxae and tro-
chanters yellow; femora dark brown; tibiae dark brown, the tips broadly
white; fore and middle legs with the proximal half of basitarsi darkened,
the remaining segments, with the exception of the last, white; posterior tarsi
white. Wings strongly tinged with brown; stigma not indicated; costal
region and barely perceptible seams along cord and over outer end of cell
1st M2 darker brown; wing apex insensibly darkened; veins dark brown.
Venation: Sg1 ending just beyond midlength of Bs, Sc2 at its tip; free tip of
Sc2 in transverse alignment with B2 ; cell 1st M2 short and nearly square;
m-cu at the fork of M, longer than the distal section of Cu

Abdominal tergites dark brown, the basal sternites pale yellow; hypo-
pygium dark. Male hypopygium with the ninth tergite transverse, its caudal
border very gently emarginate. Basistyle with the ventromesal lobe exten-
sive, very low and broad. Dorsal dististyle nearly straight, a little expanded
before apex, thence narrowed into an acute beak. Ventral dististyle fleshy,
the rostral prolongation elongate, gently curved, without evident rostral
spines. Gonapophyses appearing as broadly flattened plates, the mesal-
apical angle produced into a straight point.

Habitat. — Panama (Canal Zone).

Holotype, J1, Barro Colorado, January 9, 1929 (C. H. Curran) ;
type in American Museum of Natural History.

I take great pleasure in naming this fly in honor of Dr. C.
Howard Curran, distinguished Dipterologist. The species is most
nearly related to Limonia ( Limonia ) capnora Alexander, L. ( L .)
lutzi (Alexander) and other species but is readily distinguished
by the coloration of the legs and wings and by the details of
venation.

Limonia (Limonia) mesotricha new species.

General coloration brown, the mesonotal praescutum and scutum variegated
with brownish orange; rostrum small, yellow, palpi reduced in size; an-
tennae black throughout; front silvery; halteres elongate, black; legs black;
wings narrowed at bases, strongly tinged with brown; macrotrichia in distal
cells; B2 shortened, the free tip of Sc2 correspondingly lengthened; male
hypopygium with the rostral prolongation of ventral dististyle unarmed with
spines.

380

Journal New York Entomological Society

[Vol. LII

Male. — Length about 5 mm.; wing 6 mm.

Rostrum small, yellow; palpi black, very reduced, apparently only 1-seg-
mented. Antennae black throughout; basal flagellar segments short-oval, the
outer ones more elongate; terminal segment one-half longer than the pen-
ultimate ; verticils elongate. Front silvery ; posterior portion of head brown-
ish fulvous.

Mesonotal praescutum with the three usual stripes obscure brownish orange,
narrowly margined with brown; scutum with median area pale, the lobes
obscure brownish orange, encircled by brown; scutellum dark brown, the
median line a little paler; mediotergite with cephalic half brown, the caudal
portion brightening to obscure yellow. Pleura obscure yellow. Halteres
elongate, black throughout. Legs with the coxae and trochanters yellowish
testaceous; remainder of legs black, the extreme femoral bases restrictedly
brightened. Wings strongly tinged with brown, the darker stigmal region
restricted to a vague seam over the free tip of Sc2 ; vague darker seams over
the veins, the centers of the cells paler; veins brown. Wings narrowed
basally, long-petiolate ; relatively sparse macrotrichia in apical cells from
the stigma to cell Mi} inclusive, in most cases being restricted to rows in the
centers of the cells. Venation: Sc long, Sc2 ending shortly before the fork
of Es, Sc± shorter than Sc2; Es long, angulated at origin; vein E± bent
strongly caudad, at E2 approaching vein E2+ 3, strongly reducing E2, the free
tip of Sc2 correspondingly lengthened; m-cu near the fork of M, longer than
the distal section of

Abdominal tergites dark brown; sternites obscure brownish yellow; hypo-
pygium chiefly brown, the subterminal segments paler. Male hypopygium
with the lobes of tergite somewhat obliquely truncated. Basistyle with ven-
tromesal lobe unusually extensive, occupying almost the entire mesal face of
style. Dorsal dististyle straight, slender, the apex a little enlarged and
curved into a short point. Ventral dististyle relatively small, fleshy, its
rostral prolongation flattened, without evident spines. Gonapophyses pale
throughout, the mesal-apical lobe slender.

Habitat. — Panama (Canal Zone).

Holotype, J', Barro Colorado, December 21, 1928 (C. H. Cur-
ran) ; type in American Museum of Natural History.

Limonia ( Limonia ) mesotricha is very different from all other
regional members of the subgenus, particularly in the apically
hairy wings. The course of vein Rlt with the accompanying
shortening of vein R2 suggests the even more accentuated condi-
tions found in L. (L.) insularis (Williston) and allies.

Genus Gnophomyia Osten Sacken

Gnophomyia (Gnophomyia) podacantha new species.

General coloration black, the head polished, the mesonotum more opaque;
anterior lateral pretergites light yellow; thoracic pleura with a broad black

Dec., 1944]

Alexander : Crane-Flies

381

dorsal stripe, the ventral pleurites paler, pruinose; metapleura and posterior
coxae yellow; halteres uniformly blackened; wings with a blackish tinge, the
cord and vein Cu even more darkened; R2+a+i only slightly elevated; cell
1st M2 long and narrow, very little widened outwardly; male hypopygium
with the caudal border of tergite truncate, provided with a chiefly double
row of approximately one hundred spinous setae; cephalic end of mesal face
of basistyle produced into a long-cylindrical lobe that terminates in two con-
spicuous black spines; both dististyles narrowed and pointed at their tips.

Male. — Length about 5 mm.; wing 5. 5-5. 6 mm.

Female. — Length about 5.5 mm.; wing 5.6— 5.8 mm.

Rostrum and palpi black. Antennae black throughout, relatively long;
flagellar segments elongate, with verticils that are subequal to or slightly
longer than the segments. Head black, more or less polished ; anterior vertex
broad, sparsely pruinose, provided with a low tubercle.

Pronotum black. Anterior pretergites light yellow, the posterior preter-
gites more obscure. Mesonotum black, the surface opaque by a very sparse
pruinosity. Pleura with a broad black dorsal stripe, the ventral pleurites
paler, pruinose, on the metapleura and meron paling to yellow. Halteres
blackened. Legs with the coxae dark brown, the posterior pair paling to
yellow; trochanters testaceous brown; remainder of legs black, the femoral
bases paler. Wings with a blackish tinge, the basal third slightly more
brightened; stigma narrow, long-oval, dark brown; a slightly indicated in-
fuscation along cord and vein Cu; veins dark brown, paler in the more basal
fields. Venation: Scx ending nearly opposite the fork of £2+3 + 4; Rs short
and oblique, in direct alignment with R5; R2+ 3 + 4 only slightly elevated; R2+ 3
about twice R2; branches of Rs extending generally parallel to one another
for virtually all their lengths; cell 1st M2 long and narrow, only slightly
widened outwardly, with m-cu at about one-third to two-fifths its length.

Abdomen, including hypopygium, black. Ovipositor with elongate cerci,
these blackened basally, the distal half obscure yellow; hypovalvae short,
black. Male hypopygium with the tergite large, the caudal margin truncate,
provided with a chiefly double row of long spinous setae, there being approxi-
mately 100 or more such setae; lateral tergal angles produced into lobes, with
spinous setae to their summits. Basistyle at cephalic end of mesal face pro-
duced into a long cylindrical lobe that bears two conspicuous black spines at
summit, these slightly unequal in length and diameter. Outer dististyle a
powerful, nearly straight blackened horn or spike that narrows gradually to
the acute tip. Inner dististyle much smaller, broad at base, thence similarly
narrowed to the nearly acute apex, before this with a few erect setae.

Habitat. — Costa Rica.

Holotype, J', San Juan Miramar, Pacific slope, in cloud forest,
altitude 3,500 feet, January 1939 (Dean Rounds). Allotopotype,
5, pinned with type. Paratopotypes, 1 J', 3 $$.

From other members of the subgenus that center around
maestitia Alexander and allies, the present fly is readily told by

382

Journal New York Entomological Society

[Vol. Lll

the very different male hypopygium, especially the armature of
the basistyle which has suggested the specific name.

Genus Neognophomyia Alexander

Neognophomyia productissima new species.

General coloration of mesonotum reddish yellow; thoracic pleura with a
broad interrupted black dorsal stripe, the ventral pleurites pale; wings gray-
ish yellow, with a brown seam on anterior cord; male hypopygium with the
tergal spines unusually long and slender, entirely pale, at near midlength
narrowed and produced into a long curved needle-like point; phallosome
relatively narrow, its apex broadly obtuse to subtruncate.

Male. — Length about 4.5-4. 7 mm.; wing 4. 6-4. 8 mm.

Eostrum yellowish brown; palpi brown. Antennae brown, the pedicel
slightly brightened ; flagellar segments subcylindrical, the outer ones passing
into elongate-oval, with very long verticils. Head dark brown, more reddish
brown on central portion of vertex; eyes large.

Pronotum obscure yellow above, darker on sides. Mesonotal praescutum
shiny yellow, with poorly indicated, more reddish brown stripes, the lateral
pair especially distinct, continued backward across the suture onto the scutal
lobes; central portion of mesonotum, including the posterior portion of prae-
scutum, scutum and most of scutellum and mediotergite more yellowish.
Pleura and pleurotergite on dorsal portion with four disconnected black areas
that form a broad broken stripe, these areas located on propleura, anepis-
ternum, dorsal pteropleurite and pleurotergite; ventral pleurites more red-
dish yellow. Halteres with stem pale, knob weakly darkened. Legs with
the coxae and trochanters yellow, femora, tibiae and proximal two tarsal seg-
ments yellow, the tips very faintly and vaguely darker; outer tarsal seg-
ments brownish black. Wings with the ground grayish yellow, the pre-
arcular and costal fields clearer yellow; a relatively broad and conspicuous
brown seam extending from the stigma across anterior cord to fork of M;
much narrower seams on posterior cord and outer end of cell 1st M2; veins
brown, more yellowish in the brightened fields. Yenation: E2 placed shortly
beyond one-third the length of petiole of cell Es; vein ending close to
wing tip; cell 1st M2 rather conspicuously widened outwardly, its proximal
end very narrow; m-cu at near one-third the length of cell.

Basal abdominal tergites broadly yellow medially, conspicuously brownish
black on sides; fourth tergite paler than the adjoining ones; tergites five and
six conspicuously blackened, seven and eight again pale; sternites chiefly
yellow, the outer segments more darkened; hypopygium dark brown. Male
hypopygium with the tergal spines unusually long and slender, entirely pale,
at near midlength abruptly narrowed into a long curved needle-like point
that is subequal in length to the basal section. Outer dististyle relatively
short and stout, with about four long setae at outer end, one apical in posi-
tion. Inner dististyle short and stout, curved, at apex narrowed into a more
sclerotized beak or point. Phallosome relatively narrow, its apex broadly
obtuse to subtruncate.

Dec., 1944]

Alexander : Crane-Flies

383

Habitat. — Costa Rica.

Holotype, J1, Pedregoso, altitude 2,500 feet, January 1939
(Dean Rounds). Paratopotypes, 1 J', pinned with type; 4 J'J',
altitude 2,075-2,300 feet, January 1939 (Dean Rounds).

The most similar described species are Neognophomyia pana-
mensis Alexander and N. trinitatis Alexander, which have the
structure of the male hypopygium somewhat the same. The
present fly is readily distinguished by the very unusual elonga-
tion of the so-called tergal spines.

Genus Erioptera Meigen

Erioptera (Symplecta) macroptera argentina new subspecies.

Male. — Length about 3.5 mm.; wing 3.8-4 mm.

As in typical macroptera (Philippi), differing in the details of structure
of the male hypopygium, especially the gonapophyses. Hypopygium with
the outer dististyle only moderately expanded at apex, the latter with about
a dozen teeth. Inner dististyle slightly longer, appearing as a flattened lobe
that narrows gradually to the obtuse tip. Gonapophyses appearing as flat-
tened blades that have about the same size and somewhat the same shape as
the inner dististyle, the outer margin with numerous teeth, those toward the
base longer and more conspicuous; upper surface of apophysis with smaller
denticles. In typical macroptera (Chile), the apophyses appear as slender
blades, the apical third glabrous and only about one-half as wide as the inner
dististyle, the more basal portions of the apophysis with relatively few, very
coarse teeth.

Habitat. — Argentina.

Holotype, J1, Hornadita, Province of Jujuy, altitude 3,400
meters, November 30, 1919 (Vladimir Weiser). Paratypes, 1 J1,
1 $, Bolson, Province of Catamarca, altitude 2,700-2,750 meters,
March 7-9, 1924 (Weiser) ; 1 Province of Buenos Aires (C.
Bruch).

The late Doctor Bruch (Physis, 17 : 47 ; 1939) records the spe-
cies macroptera from La Plata, Delta of the Parana, Rio Negro
and Mendoza. It is presumed that these further records, at least
in part, pertain to the present subspecies. In an earlier paper,
Bruch (l.c., 17 : 26—28 ; 1939) provides a detailed account, with
figures, of the immature stages of this same fly.

384:

Journal New York Entomological Society

[Vol. LII

A CORRECTION

In the issue of this Journal, Vol. LII, June, 1944, pages 193-
199, under the title “A Re-Survey of Papaipema,’ ’ there was
given a contemplated list arrangement of the Papaipema species.

It has developed that in this list a form name, “ ochr opt enoides
Benj.,” there given as a variety of P. circumlucens Sm., was
never validated and should be eliminated.

Ample proof of the matter has been furnished by Dr. Carl
Heinrich, and Dr. J. McDunnough called attention to the prob-
able error concerning “ ochr opt enoides.”

It may be further remarked that a synonym of Papaipema
frigida Sm., perobsoleta Lyman, was omitted, also the synonym
imperturbata Bird, a dwarf form of P. necopina Grt., was not
listed. Both these synonyms were shortly given their correct
status in American literature. — Henry Bird.

Dec., 1944]

Jordan: Nomenclature

385

THE STATUS AND FUNCTIONS OF THE INTERNA-
TIONAL COMMISSION ON ZOOLOGICAL
NOMENCLATURE AND THE PRESENT
STATE OF ITS WORK

By Karl Jordan, Ph.D., F.R.S.

President of the International Commission

The International Commission on Zoological Nomenclature was
established by the Third International Congress of Zoology at its
meeting held at Leyden in 1895. The functions then assigned to
the International Commission were to study existing unofficial
Codes and to make proposals to a later meeting of the Interna-
tional Congress for the establishment of an International Code
of Zoological Nomenclature.

After six years’ work, the International Commission submitted
a draft International Code to the Fifth International Congress of
Zoology, by whom the proposed Code was adopted, with certain
amendments, at its meeting held at Berlin in 1901. The prepara-
tion of the authoritative French text of the Code and of the official
English and German translations thereof was completed in 1904
and the three texts were published in 1905.

At Boston in 1907 the Seventh International Congress of
Zoology authorised the International Commission to render
Opinions on questions of interpretation arising from the prac-
tical application of the Code. At Monaco in 1913, the functions
of the International Commission were extended in two important
directions by the decision of the Ninth International Congress of
Zoology (i) to confer upon the International Commission the
power to suspend the rules in cases where the strict application
of the rules would clearly result in greater confusion than uni-
formity and (ii) to entrust the International Commission with
the duty of compiling the Official List of Generic Names in
Zoology.

As originally constituted, the International Commission con-
sisted of five Commissioners, but in 1898 this number was in-
creased to fifteen. Later it was decided to increase the number

386

Journal New York Entomological Society

[Yol. Lll

of Commissioners to eighteen, the present number. Members of
the International Commission are elected by the International
Congress of Zoology, one-third of the Commission {i.e., six Com-
missioners) retiring at each meeting of the Congress. Retiring
Commissioners are eligible for immediate re-election. Vacancies
occurring between meetings of the International Congress may be
filled by the International Commission itself.

The International Commission have always attached great im-
portance to securing that the principal countries should be repre-
sented on the Commission and that, subject to this consideration,
the membership of the Commission should include one or more
specialists in each of the main divisions of the Animal Kingdom.
At the outbreak of war in 1939, the distribution of seats on the
International Commission, as between the various countries, was
as follows: the United States (4 Commissioners); the United
Kingdom (3 Commissioners) ; Germany (2 Commissioners) ; and
Argentina, Australia, Brazil, Canada, France, Hungary, Italy,
Japan and Poland (1 Commissioner each).

The present state of war makes it impossible for the Secretary
of the International Commission to communicate with certain of
the members of the International Commission, and in consequence
it will not be possible for the International Commission to make
use of their plenary powers to suspend the rules in any given
case until after the war is over, since unanimity in the Commis-
sion is required for the use of these powers. In all other matters,
the International Commission is empowered to act by a majority
and, therefore, can take — and is taking — decisions, notwithstand-
ing the present state of war.

Shortly before the outbreak of the present war, the late Dr.
C. W. Stiles resigned the Secretaryship (though not his member-
ship) of the International Commission on grounds of ill-health
and on the election of his successor, Mr. Francis Hemming, the
present Secretary, the headquarters of the International Com-
mission were transferred from Washington to London. This
change rendered it necessary to make new administrative ar-
rangements, including the transfer of the records of the Com-
mission, the raising of funds for the printing of the Commission ’s
publications, etc. At the outbreak of war in 1939, for a time

Dec., 1944]

Jordan: Nomenclature

387

operations had to be suspended, but in 1942 arrangements were
made to resume activities and during the two years that have
since elapsed a record number of publications has been issued by
the International Commission.

The publications of the International Commission consist of (i)
the Opinions and Declarations of the Commission, of which to-
gether 167 have so far been published, and (ii) the Bulletin of
Zoological Nomenclature. Thirty-seven Opinions and Declara-
tions have been published by the International Commission under
the new arrangements and many other are awaiting publication
and will be issued as soon as possible. In addition, a start is
being made in the publication of an annotated edition of the older
Opinions , the original issue of which has long been exhausted.
The Bulletin of Zoological Nomenclature was founded in 1943
as the Official Organ of the International Commission to serve
as a medium for the publication of cases submitted to the Com-
mission for deliberation and decision, thereby affording zoologists
of all countries an opportunity of commenting on any such pro-
posal before any decision is taken on it by the International Com-
mission.

In addition, the International Commission have made arrange-
ments to publish as soon as possible both the Official List of
Generic Names in Zoology and an up-to-date edition of the Inter-
national Code of Zoological Nomenclature. The first of these
has never previously been published in book form, while of the
latter there does not exist any edition containing the latest amend-
ments approved by the International Congress of Zoology.

The publications of the International Commission are obtain-
able from the Commission’s Publications Office, 41, Queen’s Gate.
London, S.W. 7.

Dec., 1944]

Members

389

MEMBERS OF THE NEW YORK ENTOMOLOGICAL

SOCIETY

(July 1, 1944)

Alexander, Charles P., Massachusetts State College, Am-
herst, Mass.

Angell, J. W., 330 W. 88th St., New York City 24, N. Y.

Bacon, Annette L., Amer. Mus. Nat. Hist., New York City
24, N. Y.

Ballou, Charles H., Est. Exper. Agric., El Valle, D. F.,
Venezuela

H.# Banks, Nathan, Museum of Comparative Zoology, Cam-
bridge, Mass.

Barber, George W., 274 Dwight St., New Haven, Conn.

Barber, H. G., 143 E. Third Ave., Roselle, N. J.

Becker, George G., Inspection House, 209 River St., Ho-
boken, N. J.

Bell, Ernest L., 150-17 Roosevelt Ave., Flushing, L. I., N. Y.

L.* Bequaert, Joseph C., Harvard Medical School, 25 Shattuck
St., Boston, Mass.

Bigelow, David, 345 E. 68th St., New York City 21, N. Y.

L. Bird, Henry, 600 Milton Road, Rye, N. Y.

Blackwelder, Richard E., U. S. Natl. Mus., Washington 25,
D. C.

Blanton, Franklin S., U. S. Ent. Lab., P. O. Box 786, Baby-
lon, L. I., N. Y.

Boyd, H. P., Boy Scouts Amer., 9337 210th St., Belair,
Queens, N. Y.

Boyd, William M., 170 Millerick Ave., Trenton, N. J.

Bromley, Stanley W., Bartlett Tree Res., Lab., P. O. Box
1337, Stamford, Conn.

Brown, F. Martin, Fountain Valley School, Colorado
Springs, Colo.

t Casselberry, R. C., 55 Edgemont Road, Scarsdale, N. Y.

Cazier, Mont A., Amer. Mus. Nat. Hist., New York City 24,
N. Y.

* H. = honorary member ; L. = life member,
t Elected after July 1, 1944.

390

Journal New York Entomological Society

[Vol. LII

Church, Frederic E., 655 Park Ave., New York City 21,
N. Y.

Clausen, Lucy W., Amer. Mus. Nat. Hist., New York City
24, N. Y.

H. Cockerell, T. D. A., 908 Tenth St., Boulder, Colo.

Collins, Donald L., Whitemarsh Res. Lab., Chestnut Hill,
Philadelphia, Pa.

Comstock, William P., Amer. Mus. Nat. Hist., New York
City 24, N. Y.

Connola, Donald P., Boyce Thompson Inst., 1086 N. Broad-
way, Yonkers, N. Y.

Crawford, J. C., Bureau of Entomology, Washington 25,
D. C.

Creighton, William S., C.C.N.Y., 139th and Convent Ave.,
New York City 31, N. Y.

Dalmat, Herbert T., Dept. Ent., Cornell Univ., Ithaca, N. Y.

H. Davis, William T., 146 Stuyvesant Place, New Brighton,
Staten Island, N. Y.

Dethier, Vincent G., 15 Beech St., Norwood, Mass.

L. Detgen, Gustav, 303 W. 42nd St., New York City 17, N. Y.

Dietrich, Henry, Dept. Ent., Cornell Univ., Ithaca, N. Y.

Engelhardt, Mrs. Elizabeth S., 28 Club Way, Hartsdale,
N. Y.

Farrelly, James T., 1507 Popham Ave., Bronx, New York
City, N. Y.

Fenton, Alfred, Texaco Sulphur Co., 2nd Natl. Bank Bldg.,
Houston, Texas

Fleming, Henry S., Dept. Tropical Res., Bronx Park, New
York City 60, N. Y.

Forbes, James, 2986 Marion Ave., New York City 58, N. Y.

Forbes, W. T. M., Dept. Ent., Cornell Univ., Ithaca, N. Y.

Fox, Henry, R.F.D. No. 1, Cape May Court House, N. J.

Funkhouser, William D., Univ. Kentucky, Lexington, Ky.

Garman, Dr. Philip H., Conn. Agric. Exper. Sta., P. O. Box
1106, New Haven, Conn.

Gertsch, Willis J., Amer. Mus. Nat. Hist., New York City
24, N. Y.

Goelbert, Raymond, 71-15 65th St., Glendale, L. I., N. Y.

Dec., 1944]

Members

391

f Goodnight, Clarence, State Teachers College, Jersey City,

N. J.

Granek, Irving, 1364 Sterling Place, Brooklyn, N. Y.

Gray, Alice, Amer. Mus. Nat. Hist., New York City 24,
N. Y.

Gny, Margaret L., Amer. Mus. Nat. Hist., New York City
24, N. Y.

Hallock, H. C., U. S. D. A, Twin Falls, Idaho
Hagan, Harold R., Coll, of City of N. Y., 139th and Convent
Ave., New York City 31, N. Y.

Harriot, Samuel C., 200 W. 58th St., New York City 19,
N. Y.

Hartzell, Albert, Boyce Thompson Inst., 1086 N. Broadway,
Yonkers, N. Y.

Haskins, Caryl P., 1188 Avon Rd., Schenectady, N. Y.
Heineman, Bernard, 175 W. 72nd St., New York City 23,
N. Y.

Hessel, Sidney A., 8 Woodmere Blvd., Woodmere, L. I.,
N. Y.

Hood, J. Douglas, Dept. Ent., Cornell Univ., Ithaca, N. Y.
Horsfall, J. L., Amer. Cyanamid Co., 30 Rockefeller Plaza,
New York City 20, N. Y.

H. Howard, L. O., 45 Pondfield Rd., W., Bronxville, N. Y.

Huckett, H. C., R.F.D., Riverhead, L. I., N. Y.

L. Hunter, R. J., 68 Broad St., Freehold, N. J.

Huntington, E. I., 115 E. 90th St., New York City 28, N. Y.
Janvrin, E. R. P., 38 E. 85th St., New York City 28, N. Y.
L. Johnson, Frank, Griffin, Georgia

Johnston, James W. [address unknown]

Jones, Frank M., 2000 Riverview Ave., Wilmington, Del.
King, James C., Red Oaks, White Plains, N. Y.

Kisliuk, Max, 150-20 87th Rd., Jamaica, L. I., N. Y.

Klots, Alexander B., 170 Elm Ave., Mount Vernon, N. Y.
Lacey, Lionel, 485 Pelham Rd., New Rochelle, N. Y.

H. McDunnough, J., Dept. Agric., Ottawa, Canada
McKenna, George F., Box 147, Adrian, Mich.

H. Melander, A. L., 4670 Rubidoux Dr., Riverside, Calif.

Michener, Charles D., American Mus. Nat. Hist., New York
City 24, N. Y.

392

Journal New York Entomological Society

[Yol. lii

Mulvaney, Gardiner, 3 Patchin Place, New York City 11,
N. Y.

L. Mutchler, Andrew J., Box 55, South Harwich, Mass.

Nadeau, Raoul, 610 W. 145th St., New York City 31, N. Y.

H. Needham, James G., Dept. Ent., Cornell Univ., Ithaca, N. Y.
Nicolay, Alan S., 18 Duryea Rd., Upper Montclair, N. J.
Noaks, John W., 500 Washington Ave., Brooklyn, N. Y.

L. Notman, Howard, Circle Rd., Dongan Hills, Staten Island,
N. Y.

Oakley, Dr. R. G., Inspection House, 209 River St., Ho-
boken, N. J.

Olsen, Chris E., Amer. Mus. Nat. Hist., New York City 24,
N. Y.

H. Osborn, Herbert, Ohio State Univ., Columbus 10, Ohio.

Osburn, Raymond C., Ohio State Univ., Columbus 10, Ohio
Pallister, John C., 3754 Independence Rd., Cleveland, Ohio
Parsons, Carl T., 205 W. 57th St., New York City 19, N. Y.
dos Passos, Cyril F., Mendham, N. J. (Deceased)

Payne, Nellie M., Amer. Cyanamid Co., Boston Post Rd.,
Stamford, Conn.

Petrunkevitch, Alexander, Peabody Museum, Yale Univ.,
New Haven, Conn.

Procter, William, Bar Harbor, Maine
Rau, George, 6023 Harrison PL, West New York, N. J.
Readio, Philip A., Dept. Ent., Cornell Univ., Ithaca, N. Y.
Rex, Edgar G., Alumni House, New Brunswick, N. J.
Richards, A. Glenn, Jr., Dept. Zool., Univ. Penn., Philadel-
phia 4, Pa.

Riehl, Louis A. [address unknown]

Rosenblum, Joseph [address unknown]

Roth, Louis M., 351 E. 45th St., Brooklyn, N. Y.

Ruckes, Herbert, 167-11 33rd Ave., Flushing, L. I., N. Y.
Rumpp, Norman L., 701 El Centro St., South Pasadena,
Calif.

Sanford, Leonard J., Amer. Mus. Nat. Hist., New York
City 24, N. Y.

Sartor, M. H., Inspection House, 209 River St., Hoboken,
N. J.

Dec., 1944]

Members

393

Satterthwait, A. F., 806 Ohio St., Urbana, 111.

Schiller, Walter, Cook County Hospital, Chicago, 111.

Schmitt, August, 389 E. 151st St., Bronx, New York City,
N. Y.

Schneirla, T. C., Amer. Mus. Nat. Hist., New York City 24,
N. Y.

Schott, Fred M., 267 E. Clinton Ave., Bergenfield, N. J.

Schwarz, Herbert F., Amer. Mus. Nat. Hist., New York
City 24, N. Y.

Schwarz, Marjorie, 1111 Park Ave., New York City 28, N. Y.

Scotland, Minnie B., N. Y. State Teachers Coll., Albany,
N. Y.

Shannon, H. J., 88-60 162nd St., Jamaica, L. I., N. Y.

Sherman, John D., Jr., 132 Primrose Ave., Mount Vernon,
N. Y.

Silliman, Fred B., 505 W. Taft Ave., Bridgeport, Conn.

H. Snodgrass, Robert E., Bureau of Entomology, Washington
25, D. C.

Soraci, Frank A., N. J. Dept. Agric., Trenton 8, N. J.

Sordillo, Lina, Amer. Mus. Nat. Hist., New York City 24,
N. Y.

Spieth, Herman T., Amer. Mus. Nat. Hist., New York City
24, N. Y.

Stone, F. Courtney, 1349 Lexington Ave., New York City
28, N. Y.

Sturtevant, A. H., Calif. Inst. Tech., Pasadena, Calif.

Swift, F. R., Fleischmann Lab., 158th and Grand Con-
course, Bronx, New York City, N. Y.

Teale, Edwin W., 93 Park Ave., Baldwin, L. I., N. Y.

Thomas, Charles A., Penn. State College Lab., Kennett
Square, Pa.

Dietz, Harrison M., 233 E. Fairmont Ave., State College,
Pa.

L. Watson, Frank E., 137-58 Geranium Ave., Flushing, L. I.,
N. Y.

Weiss, Harry B., 19 North 7th Ave., Highland Park, N. J.

Zerkowitz, Albert, 127 W. 79th St., New York City 24, N. Y,

Ziegler, J. B. [address unknown]

INDEX TO NAMES OF INSECTS AND PLANTS
IN VOLUME LII

Generic names begin with capital letters. New genera, subgenera, species,
subspecies, varieties and new names are printed in italics.

Abaeis

nicippe, 114
Alobates

barbata, death-feint of, 281
pennsylvanica, death-feint of,
281

Anosia

berenice, 237
cleothera, 238
Anteos

clorinde, 102
maerula, 103
Aphrissa
statira

jada, 106

Appius

ilaire, 116

Army-ant, behavior of, 153
Arsenura

harrietce, 76
Ascia

monuste

cleomes, 117

Behavior of

army-ant, 153
burying beetles, 311
insects to colors, 267
Besma

endropiaria, 330
quercivoraria, 330
Bruchus

obtectus, death-feint of, 262
Burying beetles, behavior of, 311
Butterflies, Mexican, 99
list of, 343
notes on, 237, 343

Calophya

arcuata, 335

Catastica

flisa, 116
nimbice

nimbice, 116

Colias

eury theme, 102
Colors

behavior of insects to, 27, 267
Cornuplura, 217

Cibarial armature of Culex, 263
Crane-flies, descriptions of, 45, 369
Cryptolabis

alticola, 56
Culex

differentiation by cibarial arma-
ture, 263

pipiens, nerve structure of, 285

Danaeus

plexippus, 238
Death-feints of
Alobates, 281
Bruchus obtectus, 262
Edchochroma, 82
muralis, 82
Dinoderus

brevis, 200
Dircenna

klugii, 239
Drosophila

melanura , 86

Eciton

hamatum, 153
Enantia

jethys, 101
marion, 101
Eremomyioides

conscript a, 365
cylindrica, 365

395

396

Journal New York Entomological Society

[Vol. Lll

fuscipes, 367
parkeri, 364
revision of, 361
setosa, 364
similis, 368
Erioptera

macroptera

argentina, 383
Eryphanis
aesacus

aesacus, 245

Everes

comyntas

albrighti, 60
valerice, 59
Euphalerus

dubius, 339
Eurema

albula, 108
boisduvaliana, 109
jacunda, 108
lydia, 108
mexicana

mexicana, 110
salome, 111
xanthochlora, 110

Felt, Ephraim Porter,
bibliography of, 228
biography, 223
portrait, 223

Food habit ratios of insects of New
York, 283

Gnomyia

acrissima, 53
hyperplatys, 55
Gnophomyia

argutula, 45
bulbibasis, 46
fessa, 47
nectarea, 49
podacantha, 380
tuber, 50
Greta

nero, 240
oto, 240

Hesperocharis

costaricensis, 101
Holorusia

furcifera, 373

Insect

behavior to colors, 27
food habit ratios of New York,
283

responses to colors, 267
International Commision on Zoologi-
cal Nomenclature, 385
recent work by, 211
Itaballia

demophile

calydonia, 117

Ithomia

hippocrenis, 239
Key to

genera of Phyllobaeninae, 122
species of

Arsenura, 76
Blitophaga, 11
Eremomyioides, 362
Nicrophorus, 11
Silpha, 11
Thanatophilus, 11
Kricogonia

lyside, 106
Kuwayama

Jiyalina, 336
lateralis, 337
mexicana, 336
striata, 336

Lambdina

canitiaria, 329

Larvae, behavior to colors, 27
Lathyrophthalmus

seneus, hibernation of, 272
Leptophobia
aripa, 117
Limonia

curraniana, 378
mesotricha, 379

Index

397

Dec., 1944]

Lutz, Frank E.

bibliography of, 69
obituary of, 63
portrait of, 62
Lycsena

ramon, 75
Lyesenopsis

pseudargiolus
balceri, 274
sidara, 273

Megisto

fetna, 242
glaucina, 244
hermes, 243
hesione, 241
libye, 244
renata

disaffecta, 243
rubricata, 242
terrestris, 242
Melete

isandra, 118
Members of New York Entomologi-
cal Society, 389
Metatrioza

neotriozella, 337
Mexican butterflies, 99
notes on, 343
Monodes

convexa, 80
Morpho

peleides

montezuma, 245

Mosquitoes, outdoor protection from,
247

Nathalis

iole, 114
Neognophomyia
interrupta, 52
productissima, 382
spectralis, 51
Neonympha

gemma, 241

Neopasites

calliopSidis, 280
elegans, 277
heliopsis, 280
Tcnulli, 278
robertsoni, 280

Neophasia

terlootii, 115

Nerve sheaths, living, structure of,
285

Nerves, living, structure of, 285

Nicrophorini
list of, 3

nearctic, revision of, 1

Nicrophorus

marginatus, 311
orbicollis, 311
pustulatus, 311
spp., behavior of, 311
tomentosus, 311
vespiloides, 311
sayi, 311

Nomenclature, recent work of Inter-
national Commission on Zoologi-
cal, 211

Okanagana

pallidula, 220
synodica, 218
nigra, 220

Oleria

paula, 239

Optomopsylla, 338

formiciformis, 338

Papaipema

list of species, 198
re-survey of, 193

Paramecera

xicaque, 245

Periplaneta

americana, nerve structure of,
285

Phoebis

agarithe

ararithe, 105

398

Journal New York Entomological Society

[Yol. Li I

maxima, 105
intermedia, 106
philea, 104
sennae, 103
Phyllobaeninas

generic review of, 121
Pieriballia
viardi

viardi, 117

Pieris

protodice, 116
rapas, 117
Pindus

squamistriga, 244
Psychodidae

catalogue of North American,
201

Psyllia

berryi, 339
cedusa, 340
martorelli, 339
Pteronymia

cotytto, 239
Pyristia
dina

westwoodi, 112
gundlachia, 111
nise

perimede, 113
proterpia, 112

Silphini

list of, 3

nearctic, revision of, 1

Tanypremna
clotho, 370
saltatrix, 369
Taygetis

inconspicua, 241
virgilia, 240
Thecla

angerona, 328
burdi, 328
caramba , 255
drucei, 260
ella, 260
harrietta, 250
kalikimaka, 258
mavors, 259
paupera, 260
pwnona, 256
triquentra, 259
Tipula

epione, 376
Tisiphone

maculata, 240
nicoya, 377
votiva, 374
Trioza

rhino sa, 337

Quesada

gigas, 213

Zerene

eaesonia, 102

The

New York Entomological Society

Organized June 29, 1892 — Incorporated June 7, 1893
Reincorporated February 17, 1943

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

Annual dues for Active Members, $3.00; including subscription to the Journal, $4.50.
Members of the Society will please remit their annual dues, payable in January, to
the treasurer.

Honorary President, WILLIAM T. DAVIS

i

Officers for the Year 1944

President, EDWIN WAY TEALE Baldwin, Long Island, N. Y.

Vice-President, GEORGE G. BECKER U. S. Bureau Ent. & Plant Quar., Hoboken, N. J.

Secretary, ANNETTE L. BACON American Museum of Natural History

Assistant Secretary, LINA SORDILLO American Museum of Natural History

Treasurer, DR. WILLIS J. GERTSCH American Museum of Natural History

Assistant Treasurer, MARGARET L. GUY American Museum of Natural History

TRUST EES

Wm. T. Davis

Edwin W. Teale

Dr. S. W. Bromley

Wm. P. Comstock H. F. Schwarz

PUBLICATION COMMITTEE

Harry B. Weiss

T. C. SCHNEIRLA
PROGRAM COMMITTEE

John D. Sherman, Jr.

A. S. Nicolay

Wm. P. Comstock
FIELD COMMITTEE

H. S. Fleming

Lucy W. Clausen

Chris. E. Olsen

DELEGATE TO THE N. Y. ACADEMY OF SCIENCES
William T. Davis

JOURNAL

of the

NEW YORK ENTOMOLOGICAL SOCIETY

Published quarterly by the Society at N. Queen St., and Mc-
Govern Ave., Lancaster, Pa. All communications relating to
manuscript for the Journal should be sent to the Editor, Harry B.
Weiss, 19 N. 7th Ave., Highland Park, New Jersey; all subscrip-
tions to the Treasurer, Dr. Willis J. Gertsch, American Museum
of Natural History, New York, N. Y. Orders for back issues
should be sent to the Treasurer, Willis J. Gertsch, American
Museum of Natural History, 79th St., & Central Park W., New
York, N. Y. The society has a complete file of back issues in stock.
The Society will not be responsible for lost Journals if not noti-
fied immediately of change of address. We do not exchange
publications.

Terms for subscription, $3.00 per year, strictly in advance.

Please make all checks , money-orders , or drafts payable to
New York Entomological Society.

Twenty-five reprints without covers are furnished free to
authors. Additional copies may be purchased at the following

rates :

4 pp. 8 pp. 12 pp. 16 pp. 24 pp. 32 pp.

25 copies .... $2.40 $5.22 $5.58 $5.58 $9.00 $9.60

Additional 100’s 60 1.44 1.92 1.92 3.00 3.00

Covers 50 copies, $2.75; additional 100’s, $1.65,

Half-tone prints 1% cents for each half-tone print.

Authors whose papers are illustrated with text figures or
full page plates will be required to supply the electroplates or
pay the cost of making the same by the Journal and also to
pay the cost of printing full page plates on coated paper, when
advisable.

JOURNAL

OF THE

NEW YORK

ENTOMOLOGICAL SOCIETY

Iruot^ to lEntomology in General

VOLUME LIII, 1945

Published Quarterly by the Society
North Queen St. and McGovern Ave.

New York, N. Y.

Lancaster, Pa.

THE SCIENCE PRESS PRINTING COMPANY
LANCASTER, PENNSYLVANIA

CONTENTS OF VOLUME LIII

Page

Abbott, Cyril E.

The Mechanism of Oviposition in Phaenicia (Lncillia)

Sericata Meig. (Diptera) 227

Alexander, Charles P.

Records and Descriptions of Neotropical Crane-Flies

(Tipulidae, Diptera) XIX 49

Records and Descriptions of Neotropical Crane-Flies

(Tipulidae, Diptera ) XX 279

Book Notice 89, 218

Boyd, William M.

Notes on the Potato Tuber Moth 68

Bromley, S. W.

Insect Enemies of the House-Fly, Musca domestica L 145

Brown, F. Martin

Notes on Mexican Butterflies 31

Comstock, William P.

Viola Harriet Dos Passos and Her North American

Moths 47

Cooper, Janet L.

See Rapp, William F., Jr.

Corrections to List of Members 88

Cutkomp, Lawrence K.

See Richards, A. Glenn, Jr.

Dobbins, T. N.

See Hawley, I. M.

Dos Passos, Cyril F.

Some Collections of Lepidoptera 62

Enzmann, Ernst V.

See Haskins, Caryl P.

Forbes, Wm. T. M.

The Eunnomid Pupa (Lepidoptera, Geometridae) 177

Frost, S. W.

Insects Feeding or Breeding on Indigo, Baptisia 219

Goodnight, Clarence J., and Marie L. Goodnight

Phalangida from the United States 239

iii

/

Haskins, Caryl P., and Ernst V. Enzmann

On the Occurrence of Impaternate Females in the

Formiciche 263

Hawley, I. M., and T. N. Dobbins

The Distribution and Abundance of the Japanese Beetle
from 1935 Through 1943, with a Discussion of Some of

the Known Factors that Influence Its Behavior 1

Malkin, Borys

A Supplement to the New York State List of Coleoptera,

No. 6, Additions and Corrections 91

McCoy, E. E.

A Laboratory Aid in the Isolation of Entomogenous

Fungi 238

Michener, Charles D.

Seasonal Variations in Certain Species of Mosquitoes

(Culicidae, Diptera) 293

Needham, James G.

A Pedicel Gall on Trichostema 259

Proceedings of the Society 69, 169

Rapp, William F., Jr.

Check-List of Psychodidae of South and Central Amer-
ica 21

A New Name for Pericoma Unicolor Abreu 262

New Psychodidae from Barro Colorado Island 309

Rapp, William F., Jr., and Janet L. Cooper

Check-List of African Psychodidae 247

Check-List of Psychodidae of Asia and Australia 211

Check-List of the Psychodidae of Europe 117

Richards, A. Glenn, Jr., and Jane L. Weygandt

The Selective Penetration of Fat Solvents Into the Ner-
vous System of Mosquito Larvae 153

Richards, A. Glenn, Jr., and Lawrence K. Cutkomp

Neuropathology in Insects 313

Sanford, Leonard J.

Anteos Maerula, a New Butterfly Recorded from Florida 136
Schiff, Howard

Notes on Thysania zenobia 88

Schwarz, Herbert F.

Review of “Notas sob re Insectos Daninos Observados en
Venezuela 1938-1943” by Charles N. Ballou 218

IV

The Wax of Stingless Bees (Meliponidae) and the Uses

to Which It Has Been Put 137

Wang, Yushi Moltze

A Preliminary Report on Chilopoda at Isham, Kwangsi

and Meitan 63

Weiss, Harry B.

Ancient Remedies Involving Insects 246

C. V. Riley ’s Pirated Work 226

Early Entomological Manuscripts 258

* Insect Food Habit Ratios of the Lloyd-Cornell Reser-
vation 167

The Longevity of Entomologists 231

Review of “A Bibliography of the Homoptera” by Zeno

Payne Metcalf 89

Some Early Entomological Ideas and Practices in

America 301

William Thompson Davis — 1862-1945 127

Weygandt, Jane L.

See Richards, A. Glenn, Jr.

v

■

VoL LIII

No. 1

MARCH, 1945

Journal

of the

New York Entomological Society

Devoted to Entomology in General

HARRY B. WEISS JOHN D. SHERMAN, Jr.

T. C. SCHNEIRLA

Subscription $3.00 per Year

Published Quarterly by the Society
N. QUEEN ST. AND McGOVERN AVE.
LANCASTER, PA.

NEW YORK, N. Y.

1945

CONTENTS

The Distribution and Abundance of the Japanese Beetle
from 1935 Through 1943, with a Discussion of Some of
the Known Factors That Influence Its Behavior

By I. M. Hawley and T. N. Dobbins 1

Check-List of Psychodidae of South and Central America

By William F. Rapp, Jr 21

Notes on Mexican Butterflies

By F. Martin Brown 31

Viola Harriet Dos Passos and Her North American Moths

By William P. Comstock 47

Records and Descriptions of Neotropical Crane-Flies
(Tipulidae, Diptera), XIX

By Charles P. Alexander 49

Some Collections of Lepidoptera

By Cyril F. dos Passos 62

A Preliminary Report on Chilopoda at Ishan, Kwangsi
and Meitan

By Yuhsi Moltze Wang 63

Notes on the Potato Tuber Moth

By William M. Boyd 68

Proceedings of the New York Entomological Society 69

Corrections to List of Members 88

Notes on Thysania zenobia

By Howard Schiff 88

Book Notice 89

NOTICE: Volume LII, Number 4, of the Journal of the
New York Entomological Society was published on
December 19, 1944.

Entered as second class matter July 7, 1925, at the post office at Lancaster, Pa.,
under the Act of August 24, 1912.

Acceptance for mailing at special rate of postage provided for in Section 1103.
Act of October 3, 1917, authorized March 27, 1924.

JOURNAL

OF THE

New York Entomological Society

Vol. LIII March, 1945 No. 1

THE DISTRIBUTION AND ABUNDANCE OF THE
JAPANESE BEETLE FROM 1935 THROUGH
1943, WITH A DISCUSSION OF SOME
OF THE KNOWN FACTORS
THAT INFLUENCE ITS
BEHAVIOR

By I. M. Hawley and T. N. Dobbins
Bureau of Entomology and Plant Quarantine, Agricultural Research
Administration, United States Department of Agriculture

It has been 10 years since a paper treating of the distribution
and abundance of the Japanese beetle ( Popillia japonica Newm.)
has been published. During this 10-year period the insect has
been subjected to a variety of climatic conditions and has en-
countered a wide range of physiographic types that have influ-
enced its behavior. The reaction of the beetle to the conditions
encountered in its spread, the variations in its abundance, and
the factors responsible for changes in numbers are discussed in
this paper.

CHANGES IN THE GENERALLY INFESTED AREA AND
FACTORS INFLUENCING THE RATE OF SPREAD

The progressive dispersal of the Japanese beetle in the United
States prior to 1934 has been treated in several papers by Henry
Fox (1, 2, 3),1 who also carried on adult-beetle surveys within the
generally infested area in 1934 and 1935. As pointed out by Fox
(3), the total range of the Japanese beetle in the United States
falls naturally into two subdivisions, the smaller of these consist-

1 Numbers in parentheses refer to Literature Cited.

2

Journal New York Entomological Society

[Vol. LIII

ing of an area, known as the area of general distribution, in which
the beetle is present at nearly all points with suitable environ-
mental conditions; whereas in the larger subdivision, or periph-
erally infested zone lying beyond this generally infested area,
beetles occur only in localized colonies of various sizes separated
by extensive areas free from the pest. Fox designated this large
peripheral zone as the area of discontinuous infestation, but in
recent years it has been more generally referred to as the outer

Fig. 1. Dots indicate all known points in the outer zone at which the
Japanese beetle has been found prior to and during the summer of 1943.

zone. These two areas, as found in 1943, are shown graphically
in Figure 1. Surveys of adult-beetle abundance were carried on
in the area of general distribution each year from 1925 through
1939 by the staff of the Moorestown, N. J., laboratory of the
Bureau of Entomology and Plant Quarantine, United States De-
partment of Agriculture. In 1940 there was no scouting of this
type, and since 1941 surveys have been more restricted and less
thorough because of personnel and travel limitations; their con-
tinuance has been possible only because of the active cooperation

Mar., 1945]

Hawley & Dobbins: Japanese Beetle

3

of entomological agencies in the states involved. Information
relative to conditions in the outer zone has been obtained largely
from the trapping activities of the Division of Japanese Beetle
Control, of the Bureau.

The area of general distribution has gradually developed to its
present extent by the spread of the beetle, largely by flight, from

Fig. 2. The area of general distribution of the Japanese beetle in the
summer of 1943.

Note. In figures 2, 4, 5, 6, and 7 the relative abundance of the beetle is
shown by the closeness of the dots in stippled areas. The single dots beyond
the limits of the area of general distribution in figures 5 and 6 mark the
locations of isolated beetle infestations.

the original point of introduction in this country near Riverton,
in west-central New Jersey. This dispersal has taken place in all
directions over a period of more than 25 years, until at the close
of the 1943 summer season the pest had covered an area of
roughly 29,200 square miles and had invaded eight states and the
District of Columbia (Fig. 2). The growth of the area of gen-

4

Journal New York Entomological Society

[Vol. LIII

eral distribution is shown graphically in Figure 3, and the pro-
gressive increase in the size of the infested area is shown in
table 1.

In recent years one of the most significant factors involved in
the progressive increase of the area of general distribution has
been the absorption by this area in its outward spread of a num-

Fig. 3. Progressive changes in the outer limit of the area of general
distribution of the Japanese beetle from 1925 through the summer of 1943.

ber of extensive secondary centers of dispersal, which have de-
veloped independently of the primary infestation. The largest
of these secondary centers have evolved, not from single isolated
infestations, but from numbers of such infestations which have
united to form extensive tracts of generally infested territory.

The largest secondary center of dispersal has been that in the
parts of Maryland and Virginia lying east of Chesapeake Bay
(Figs. 2 and 4). The infestation there originally consisted of a

Mar., 1945]

Hawley & Dobbins: Japanese Beetle

5

number of local colonies which had fused to such an extent that,
when this area merged with the area of general distribution in
1942, all the Eastern Shore of Maryland and Virginia except one
relatively small tract was involved. The same type of situation
existed when the fringe of the generally infested, area, which had
been moving north in Connecticut above New Haven, met and
united with a strong local colony that had existed around Hart-
ford for a long time, and when this latter colony, in turn, joined
one spreading south from Springfield, Mass. (Figs. 2 and 4).

TABLE 1

Increase in Size of Area of General Distribution of the Japanese
Beetle, by States, from 1935 through 1943

Estimated infested area (square miles)

State 7

1935 1937 1939 1941 1943

New Jersey t 6,460 6,980 7,250 7,431 8,224

Pennsylvania 3,100 4,358 5,013 6,114 , 7,169

Delaware 670 946 1,064 1,550 1,965

Maryland 480 664 1,546 3,016 * 5,887

New York 690 858 1,141 1,722 2,418

Connecticut 45 286 620 2,200

District of Columbia 62 62

Virginia 85 1,085

Massachusetts 190

Total infested area 11,400 13,851 16,300 20,600 29,200

Increase in area 2,451 2,449 4,300 8,600

These large additions were largely responsible for the sizeable
increase in the area of general distribution between 1941 and
1943, as shown in Table 1. There have be.en other instances of
this kind in earlier years, when the area centering about Harris-
burg, Pa., was added in 1937 (Fig. 5) and when the spread of the
area of general distribution to the southwest below Baltimore
reached and united with a local infestation about Washington,
D. C., in 1941 (Fig. 4). In many instances, however, these iso-
lated colonies were so small that their addition to the area of
general distribution would not have been recognized had not the
area involved been scouted just before the map was prepared.

6

Journal New York Entomological Society

[Vol. LIII

Topography, or rather a complex of factors conditioned by
topography, is becoming an increasingly important influence on
the natural dispersal of the Japanese beetle. The rate of spread
of the insect was fairly uniform in the earlier years, owing largely
to the fact that the dispersal at that time was through fairly uni-

Fig. 4. The area of general distribution of the Japanese beetle in the
summer of 1941. The extent of certain large isolated centers of beetle dis-
persal in the outer zone is shown by stippling.

form terrain. As the spread has progressed inland to the north
and west, however, the beetle has reached the eastern rim of the
Appalachian Mountains, which extend in a northeastern-south-
western direction across Pennsylvania, northern New Jersey, and
southeastern New York. By 1943 the outer limit of dispersal had
either reached or had begun to penetrate this mountain system in

Mar., 1945]

Hawley & Dobbins: Japanese Beetle

7

all three states (Fig. 2). Observations here and at other points
have shown that such physical barriers, although not permanent
obstacles, do markedly retard the rate of beetle dispersal.

The physiography of the easternmost part of the Appalachian
Mountains in Pennsylvania is characterized by a series of ap-
proximately parallel, heavily wooded ridges separated by narrow
valleys, the ridges rarely rising more than 600 to 800 feet above

Fig. 5. The area of general distribution of the Japanese beetle in the
summer of 1937.

the immediate lowlands. Occasionally these ridges are bisected
by river valleys of various sizes and by natural breaks known as
gaps. It is becoming increasingly evident that the dispersal of
the Japanese beetle has been more rapid up these valleys and
through the gaps than in the rougher, higher terrain. In 1943
the infestation extending up the Susquehanna River in Pennsyl-
vania could be considered continuous as far as Sunbury, a pene-
tration of roughly 50 miles into the Appalachian Mountain sys-

8

Journal New York Entomological Society

[Yol. liii

tem, while in the lower Hudson River Valley in New York the
beetle had made its way through the bordering semimountainous
country to a point some distance above Poughkeepsie (Fig. 2).
The spread to the north through Connecticut has largely followed
the Connecticut River Valley. Thus the outline of the area of
general distribution, which in earlier years was roughly arcuate
in shape, is becoming progressively irregular, being character-
ized by prominent arms or streamers that extend outward
through the river valleys.

In 1935 it was estimated by Fox (4) that the area of general
distribution covered 11,400 square miles. It included the north-
ern half of Delaware, the extreme northeastern part of Maryland
around the head of Chesapeake Bay, the greater portion of the
open, rolling farming country lying east of the lower reaches of
the Susquehanna River and southeast of the Appalachian Moun-
tains in Pennsylvania, and all of New Jersey except the extreme
northwestern part (Fig. 6). In New York State the generally
infested area included Staten Island and the metropolitan area
of New York City,, and extended some distance up the Hudson
River Valley; beetles were also present over the western fourth
of Long Island.

By 1937 beetles had spread over much of the northern two-
thirds of Delaware, and there had been some dispersal to the west
and south in extreme northeastern Maryland (Fig, 5). There
had been a movement up the Susquehanna River in southeastern
Pennsylvania that had reached and joined a group of isolated
infestations centering about Harrisburg. In Pennsylvania to the
east of this area there had been only a moderate dispersal beyond
the outer limits noted in 1935. Similarly, in New Jersey the dis-
persal had been rather slow in the semimountainous area in the
northwestern part of the State. There had been a slight spread
up the Hudson River Valley in New York and to the east on Long
Island. The State of Connecticut had been invaded for the first
time when the area of general distribution moved into the south-
west corner.

On the maps for 1935 and 1937 (Figs. 6 and 5, respectively)
the larger isolated colonies that lie in the outer zone just beyond
the area of general distribution are designated by single dots ; on

Mar., 1945]

Hawley & Dobbins: Japanese Beetle

9

later maps only a few of the more important colonies are shown,
and the extent of these is indicated by the limits of the stippled
areas.

By 1939 there had been a moderate dispersal to the south in
Delaware and eastern Maryland; to the west in Maryland the

Fig. 6. The area of general distribution or the area generally infested
by the Japanese beetle in the summer of 1935.

spread had carried the beetle beyond the city of Baltimore ; and
in Pennsylvania the Susquehanna River had been crossed from
the Maryland State line to a point well above Harrisburg (Fig.
7 ) . As in previous years, there had been only a slight movement
to the northwest in the Appalachian Mountains in Pennsylvania
and New Jersey, and only the normal spread had occurred up the

10

Journal New York Entomological Society

[Vol. LIII

Hudson River Valley and to the east on Long Island, in New York
State. There had been a pronounced eastward dispersal in Con-
necticut along Long Island Sound, resulting from the inclusion
of several local infestations in the area of general distribution.
The presence of a large isolated center of dispersal on the Eastern

Fig. 7. The area of general distribution of the Japanese beetle in the
summer of 1939.

Shore of Virginia and in southeastern Maryland is indicated on
the map for 1939 (Fig. 7). There were other local colonies in this
area, especially to the southward in Virginia, but, as only a par-
tial survey of this area was made in 1939, their extent at that time
was uncertain.

By 1941 all but the extreme southern part of Delaware had

Mar., 1945]

Hawley & Dobbins: Japanese Beetle

11

become infested by the Japanese beetle, and the isolated center
of dispersal on the Eastern Shore of Maryland and Virginia had
increased in size until it had almost reached the lower fringe of
the area of general distribution just to the north (Fig. 4). The
main infested area in Maryland had spread sonthwestward be-
yond Baltimore and had united with an isolated colony centering
about Washington, D. C., by a slender strip that followed the
main Baltimore-to-Washington highway. There had been con-
siderable dispersal to the westward in Pennsylvania through the
open country west of the Susquehanna River and a slight though
gradual spread northwestward in most of the Appalachian Moun-
tain region. At one point near the New Jersey State line there
had been a penetration through the Delaware Water Gap into the
higher country beyond. Up the Hudson River Valley in New
York State the beetle had spread beyond Newburgh and it had
covered roughly half of Long Island in its sweep to the east. In
Connecticut the dispersal to the east had carried the insect well
beyond New Haven, and north of there a strong local colony was
developing and spreading south from Hartford.

In 1943 all of Delaware was in the area of general distribution
and the spread of the beetle to the south had met and joined the
previously large isolated center of dispersal occupying the East-
ern Shore of Maryland. This new area was found to be continu-
ous to the extreme tip of the Eastern Shore of Virginia (Fig. 2).
There was only one small section of Maryland east of Chesapeake
Bay that the beetle had not reached ; on the western side of the
bay the insect could be found everywhere well below the latitude
of Washington, D. C. There is some question as to the exact loca-
tion of the western border of the area of general distribution in
Maryland in 1943, as this part of the state was not carefully
scouted, and, as may be seen from the map, there is a large iso-
lated center of dispersal to the west that may possibly have be-
come joined to the generally infested zone at some point. The
spread to the west in southeastern Pennsylvania west of the
Susquehanna River had continued and it will probably move
rapidly in the future, as this open, fertile farming country is well
fitted to support a heavy beetle population. Beetles were found
along both banks of the Susquehanna River as far as Sunbury,

12

Journal New York Entomological Society

[Vol. LIII

where a local colony has existed for some years. By 1943 all the
open farming country in southeastern Pennsylvania south and
east of the Appalachian Mountains was generally infested, and at
several points the insect had begun to penetrate this mountain
system. In 1943 all of New Jersey was placed in the area of
general distribution for the first time. In New York State the
Hudson River Valley became infested beyond Poughkeepsie and
beetles may now be found in the hilly terrain near the Connecti-
cut State line beyond Pawling. There had been a steady dis-
persal to the east on Long Island, slower in the central part than
in the shore areas because this inland section is unfavorable for
beetle spread and development. As already noted, there had been
a marked increase in the infested area in Connecticut since 1941,
for, in addition to a considerable dispersal to the east, there had
been a spread up the Connecticut River Valley in which the area
of general distribution moving north from New Haven had met
and united with the strong isolated infestation that had been
moving down from Hartford for several years. North of Hart-
ford there had been fusion with a dispersal center that had been
moving south from Springfield, Mass.

By the end of the 1943 beetle season the area of general distri-
bution was estimated at 29,200 square miles. This is more than
twice the size of the continuous area in 1937, when 13,851 square
miles were infested. It required from 1916 to 1937, a period of
21 years, to cover a smaller area than has been covered in the
6 years from 1937 through 1943. As previously noted, this strik-
ing increase came about largely as a result of the inclusion in the
area of general distribution of a number of large isolated centers
of dispersal. It is of interest that the extreme outer limits of
general dispersal to the south, Cape Charles, Va., and to the
north, Springfield, Mass., are each approximately 200 miles from
the seat of the original infestation in this country near Riverton,
N. J., while some of the more remote points of dispersal to the
northwestward into the mountainous section of Pennsylvania are
only about 75 miles away.

It will be noted that the distribution maps which accompany
this paper are not all on the same scale and that there are differ-
ences in their structural make-up. This is because these maps

Mar., 1945]

Hawley & Dobbins: Japanese Beetle

13

have been drawn by different persons and because the size of each
map had to be adjusted to cover the limits of the area of general
distribution as found at the time.

BEETLE ABUNDANCE IN THE AREA OF GENERAL DISTRI-
BUTION AND CLIMATIC AND BIOTIC FACTORS THAT
CAUSE CHANGES IN NUMBERS

The abundance of the Japanese beetle at any point is depen-
dent on many factors, among the more important of which are the
age of the infestation, the type of habitat, which includes the
availability of favored food plants and suitable places for ovi-
position, the amount of summer rainfall, and the presence or
absence of predaceous and parasitic insects and pathogenic micro-
organisms. It is not within the scope of this paper to go into all
these subjects in detail, as they have been treated in other publi-
cations (4, 5, 8, 9), but influences known to have been especially
active in the years 1935 through 1943 will be briefly discussed.

When the Japanese beetle first spreads into new territory, so
few are to be found that it is necessary to search the more favored
food plants carefully to locate them. This would be the condition
along the .outer fringe of the area of general distribution. From
this small start the population will gradually increase for a num-
ber of years until, if conditions are favorable, beetles will be pres-
ent in highly destructive numbers ; eventually the population will
decline until a more or less stable condition at a lower level is
reached. The abundance of the insect at any place, therefore,
will depend to some extent on the age of the infestation or the
position of the given place in the above cycle.

The Japanese beetle has been found to thrive in suburban resi-
dential areas where there are plenty of garden and shade-tree
hosts and an abundance of thrifty turf for oviposition; beetle
colonies also flourish in agricultural areas having good loamy soil,
an abundance of pasture land, and plenty of food plants in the
form of fruit trees, cultivated crops, or favored weed hosts.
Beetles do not usually develop to great numbers in densely
wooded regions, in neglected land overgrown with plants that
are unattractive as food, or in places with a very light type of
soil. Beetle concentrations will depend to some extent, therefore,
on the environmental conditions encountered as the insect moves

14

Journal New York Entomological Society

[Vol. Lin

into new areas (5, 8). It should be noted, however, that strong
beetle colonies have been found in some locations that would
appear to be unfavorable for their development, showing that the
beetle has great adaptability.

The amount of rainfall during the summer months, when eggs
and newly hatched larvae predominate in the soil, has an impor-
tant bearing on the size of the beetle population the following
year, because the eggs need an abundance of moisture to complete
their embryological development (4, 6). The rainfall at any
point varies greatly from year to year and, as precipitation in
summer is often in the form of local showers, there may be plenty
of moisture in the soil at one point and a deficiency at places
nearby. In the Philadelphia area the critical period for eggs in
the soil is roughly from July 1 to August 10, as soil surveys have
shown that nearly 95 per cent of the eggs in an average season
are found during this time ; at points south of Philadelphia this
period occurs earlier, and it is slightly later farther north. Vari-
ations in rainfall can therefore cause marked fluctuations in the
beetle population.

Insect parasites of the different stadia have, at least locally,
an influence on the size of the beetle populations. Under varying,
limited environmental conditions parasitic nematodes, fungus
diseases, and an undetermined number of bacterial diseases may
also become highly important factors in reducing the soil popula-
tion. In restricted areas predators, such as birds, skunks, mice,
and moles, are often active in destroying various stages of the
beetle (5).

Of the various factors effecting a measure of biotic control of
the Japanese beetle, however, probably the most widespread and
generally effective is a group of bacterial pathogens that produce
the condition known as milky disease of beetle larvae. The infec-
tive stage of this group in the soil is a bacterial spore which is
well adapted to survive under a wide variety of environmental
conditions, because it is highly resistant to desiccation and can
maintain its viability and infectiousness over a period of several
years and then be able to transmit the disease to other larvae that
ingest the spores along with soil particles in feeding. When dis-
eased larvae die, the infective spores that fill the body cavity are

Mar., 1945]

Hawley & Dobbins: Japanese Beetle

15

left in the soil and, in places with high larval populations, the
spore concentration increases so rapidly that milky diseases be-
come an increasingly important factor in keeping the population
of the insect at a low level.

Following several years with favorable climatic conditions and
in the absence of a strong concentration of milky disease spores
in the soil, the Japanese beetle by 1935 had become abundant over
much of the infested area in New Jersey and eastern Pennsyl-
vania, as is evident from the extent of the closely stippled and
dark areas on the distribution map for that year (Fig. 6). In the
more heavily stippled areas obvious foliage injury would be of
general occurrence and locally there would be extremely severe
damage. From this destructive condition there would be a grada-
tion to one of only slight feeding in the lightly stippled areas
along the lower Atlantic coast, where beetles have rarely been
present in destructive numbers. The feeding habits and food
plants of the beetle are treated in a circular by Hawley and
Metzger (8).

In the interval between the summer of 1935 and that of 1937
climatic conditions had a marked reducing effect on beetle popula-
tions (Fig. 5). The summer of 1936 was warmer than usual and
also deficient in rainfall during July in the section of New Jersey
north of Trenton and the contiguous part of Pennsylvania to the
west. Elsewhere in the area of general distribution precipitation
was rather uneven in 1936, and, as a result, infestations in 1937
tended to vary greatly in intensity. In January and February,
1936, there was an extended period when low temperatures com-
bined with abnormal soil moisture brought about the most wide-
spread mortality of hibernating larvae that has occurred since the
beetle first became established in this country. As noted in a
paper by Hawley and Dobbins (7), this winter-killing occurred
largely in the southern half of New Jersey, in southeastern Penn-
sylvania, and at certain points in Delaware and Maryland. The
combination of these unfavorable weather conditions resulted in
a marked drop in the beetle population in much of the infested
area by the summer of 1937.

Rainfall was far below normal in July and August, 1937, in the
southern half of New Jersey, and the already depleted beetle

16

Journal New York Entomological Society

[Vol. Lin

population was still further reduced. Elsewhere in 1937 and
everywhere in 1938 rainfall was adequate for the favorable devel-
opment of the insect, and by the summer of 1939 an increase in
beetles at most points was clearly evident. The highest concen-
trations continued to exist in southeastern Pennsylvania, north-
eastern Maryland, and northern Delaware (Fig. 7).

In 1939 and 1940 summer rainfall was below normal in parts
of northern New Jersey and in all of southern New Jersey, and by
1941 the infestation in the southwestern part of the State, where
a few years before beetles had been more destructive than else-
where in the country, had markedly decreased from that observed
in former years. Numbers were also reduced in Pennsylvania,
except in the densely infested zone in the southeastern corner of
the State. This same high concentration of the insect was appar-
ently also maintained in northern Delaware and northeastern
Maryland. In the isolated infested area in southeastern Mary-
land beetles were also present in considerable numbers and there
were increases at some points in southeastern New York (Fig. 4).

In both 1941 and 1942 there was a return to a condition of at
least normal summer rainfall, and this was accompanied by in-
creases in the beetle population throughout most of the enlarged
area of general distribution. The infestation in southwestern
New Jersey showed a remarkable increase in intensity, and there
were larger areas of high beetle concentration in northern New
Jersey, in parts of Connecticut, and in Pennsylvania, Delaware,
and Maryland, as well as on Long Island (Fig. 2). The current
trend appears to be toward the development of several large
heavily infested tracts separated by more lightly infested zones.
In any of the darker areas on the more recent maps severe foliage
injury would be general.

In discussing changes in beetle abundance from 1935 through
1943 the influence of summer rainfall has been stressed, not be-
cause it is the only factor involved but because it is the one best
understood. At the beginning of this period the organisms caus-
ing milky disease were present locally in the oldest infested area
about Philadelphia, but diseased larvge were found rarely, if at
all, in the more remote parts of the area of general distribution.
There has been some natural dispersal of milky disease infection

Mar., 1945]

Hawley & Dobbins: Japanese Beetle

17

during the period from 1935 to date which has carried the organ-
ism into new areas, but this spread has not kept pace with the
natural dispersion of the beetle. For this reason the bacterial
pathogens that cause the most prevalent type A disease have been
introduced at many points throughout the beetle-infested states
in an extensive colonizing program carried out by the Moores-
town, N. J., laboratory in cooperation with entomological agencies
in the states involved (13, 14). As shown by soil surveys dis-
cussed in papers by White (10, 11) and by White and Dutky
(12), the type A disease has become so well established at many
points that it is now an important factor in reducing the soil
population of the Japanese beetle. The widespread distribution
of this disease was not started until 1939 and, as it is possible to
treat only a relatively small percentage of the land in any given
area with the limited spore material available, it is still too soon
to expect large reductions in beetle populations at the more recent
points of introduction. Soil surveys have shown, however, that
there is such a high incidence of disease at certain points in Con-
necticut, New York, Delaware, and Maryland, where treatments
were applied early in the distribution program, that recent reduc-
tions in beetles in these areas are undoubtedly due to this cause.
As already noted, spore concentrations of milky disease in the soil
increase rapidly in the presence of high larval populations ; there-
fore this disease should become an even more effective agent as
time goes on.

The situation in southwestern New Jersey deserves especial
consideration. It has been possible to trace the yearly changes
in the soil population in this area by surveys made by the per-
sonnel of the Moorestown laboratory. Several years ago, when
beetles were plentiful in this area, spores of the milky disease
built up to a high concentration; later, when a marked drop in
the larval population occurred, the disease still persisted, although
a smaller number of the infected larvae were recovered. This was
the condition in 1940, but, following two summers with favorable
rainfall, the larval population had greatly increased by 1943 and
the disease incidence had become so high that this factor alone
undoubtedly would have brought the soil population down to a
much lower level. However, the summer of 1943 was again ex-

18

Journal New York Entomological Society

[VOL. LIII

tremely dry, and this will also tend to reduce the beetle concen-
tration in this area in 1944. Apparently, therefore, there are two
main agencies operating to bring about changes in the soil popu-
lation, one of which, the milky disease, when once established,
continues to build up and increase in effectiveness as a control
measure; whereas the other, summer rainfall, is an uncertain,
fluctuating factor that may bring about either an increase or a
decrease in numbers. These two factors working together are
probably responsible for most of the larger changes in population
density. As already noted, other agencies that go to make up the
biotic complex are operative, but the area in which they occur is
usually more restricted. There is some evidence that at certain
places, in the outer zone to the north, unrecognized factors are at
work, therefore the picture of population changes, as we know it
now, is still far from complete.

STATUS OF THE BEETLE IN THE ISOLATED COLONIES
OF THE OUTER ZONE

In the outer zone, beyond the limits of the area of general dis-
tribution, the Japanese beetle occurs in isolated colonies of vari-
ous sizes with uninfested areas in between. These colonies exist
because at some time in the past beetles were carried to these
points by automobile, train, airplane, or in the transportation of
plants or other materials (5, 9). Every known point where
beetles have been found in the outer zone is indicated by a dot on
the map in Figure 1. In some places, as in Bratenahl, a section
of Cleveland, Ohio, and at Providence, R. I., Richmond, Va., and
Asheville, N. C., these colonies have increased in size and strength
over a period of years until beetles are now present in destruc-
tive numbers. At many points shown on the map only a few
beetles were originally found and in some places it was impossible
to find any beetles when these locations were checked by trapping
or scouting. For example, 1 beetle was found at Fort Madison,
Iowa, the most western point, in the summer of 1937 and none has
been taken since. At 3 locations in Florida that are shown on the
map a total of 10 beetles have been taken in 2 trapping seasons
and there is no evidence that permanent colonies now exist in
this state. A combined scouting and suppression program is
carried on in the outer zone each year by the Division of Japanese

Mar., 1945]

Hawley & Dobbins: Japanese Beetle

19

Beetle Control of the Bureau of Entomology and Plant Quaran-
tine, in cooperation with state regulatory agencies. As a part of
this program traps are operated at key locations in many states
and information is thus obtained as to the presence or absence of
the insect at these remote points. The presence of beetles at most
of the locations shown on the map (Fig. 1) was discovered in these
trapping operations. Beetles have been taken at only a few of
the many places that have been trapped. At many points in the
outer zone an effort is being made to eradicate these local infesta-
tions by treating the soil with lead arsenate to destroy the larvae
as they feed. Trapping in these treated places has shown that a
good control has been obtained and few beetles will now be found
at most such places.

It is not within the scope of this paper to consider how far the
Japanese beetle will eventually spread in this country or what its
status as a pest will be in its future range. This subject has been
ably treated in a paper by Fox (4) published in 1939, and most
of the evidence obtained since this time indicates that the insect
will be able to maintain itself in most of the Eastern States, as
Fox predicted, and that in much of its new range it could build
up to destructive numbers. It is pointed out by Hawley (6) that
under the colder climatic conditions in the northern part of the
outer zone the seasonal cycle of the insect is so modified that it
will probably not become so numerous or destructive there as it
has farther south. Some uncertainty still exists as to how the
beetle will react in the coastal sections of the extreme Southern
States with their lighter types of soil, the higher temperatures,
the prolonged periods of scanty rainfall, and the differences -in
vegetation. Sufficient evidence is available, however, to show
that the Japanese beetle can eventually disperse into the outer
zone well beyond the present limits of the area of general distri-
bution before encountering any effective barrier to its successful
establishment.

LITERATURE CITED

(1) Fox, H. 1927. The present range of the Japanese beetle, Popillia

japonica Newm., in America and some factors influencing its

spread. Jour. Econ. Ent., 20: 383-391, illus.

(2) . 1932. The distribution of the Japanese beetle in 1930 and

1931, with special reference to the area of continuous infestation.

Jour. Econ. Ent., 25: 396-407, illus.

20

Journal New York Entomological Society

[Vol. LIII

(3) . 1934. The known distribution of the Japanese beetle in 1932

and 1933. Jour. Econ. Ent., 27: 461-473, illus.

(4) . 1939. The probable future distribution of the Japanese

beetle in North America. Jour. N. Y. Ent. Soc., 47: (105)-123.

(5) Hadley, C. H. and Hawley, I. M. 1934. General information about

the Japanese beetle in the United States. U. S. Dept. Agr. Cir.
332, 23 pp., illus.

(6) Hawley, I. M. 1944. Notes on the biology of the Japanese beetle.

U. S. Bur. Ent. and Plant Quar. E 615, 18 pp., illus. [Processed.]

(7) and Dobbins, T. N. 1941. Mortality among hibernating

larvae of the Japanese beetle with special reference to conditions in
the winter of 1935-36. Jour. N. Y. Ent. Soc., 49: 47-56, illus.

(8) and Metzger, F. W. 1940. Feeding habits of the adult Japa-

nese beetle. U. S. Dept. Agr. Cir. 547, 31 pp., illus.

(9) Smith, L. B. and Hadley, C. H. 1926. The Japanese beetle, U. S.

Dept. Agr. Cir. 363, 67 pp., illus.

(10) White, R. T. 1940. Survival of type A milky disease of Japanese

beetle larvae under adverse field conditions. Jour. Econ. Ent., 33:
303-306.

(11) . 1941. Development of milky disease on Japanese beetle

larvae under field conditions. Jour. Econ. Ent., 34: 213-215.

(12) and Dutky, S. R. 1940. Effect of the introduction of milky

diseases on populations of Japanese beetle larvae. Jour. Econ. Ent.,
33: 306-309.

(13) and Dutky, S. R. 1942. Cooperative distribution of organ-

isms causing milky disease of Japanese beetle grubs. Jour. Econ.
Ent., 35: 679-682.

(14) and McCabe, P. J. 1943. Colonization of the organism caus-

ing milky disease of Japanese beetle larvae. U. S. Bur. Ent. and
Plant Quar. E 605, 6 pp. [Processed.]

Mar., 1945]

Rapp : Psychodhee

21

CHECK-LIST OF PSYCHODID^ OF SOUTH
AND CENTRAL AMERICA1

By William F. Rapp, Jr.

This check-list, which has been compiled after a careful study
of the entomological literature, includes all species described up
to December 31, 1943. It covers Central America south of the
United States, all of South America and the West Indies. Based
upon zoogeographical division, this area would be considered the
Neotropical Realm.

I wish to thank Miss Janet L. Cooper for the help she has given
me in the preparation of this paper.

ALEPIA Enderlein

scripta Enderlein, Dtsch. ent, Z., 1936, p. 95.

Bolivia.

BRU CHOMYIA Alexander

argentina Alexander, Ann. Ent. Soc. Amer., Vol. 13 (1920), p.
405.

Argentina.

brasiliensis Alexander, Rev. Ent. Rio d*e J., 11 (1940), p. 795.
Brazil.

peruviana Alexander, Proc. U. S. Nat. Mus., Yol. 75 (1929) Art.
7, p. 6.

Peru.

shannoni Alexander, Proc. U. S. Nat. Mus., Vol. 75 (1929), Art.
7, p. 4.

Peru.

CHIROLEPIA Enderlein
albicollara Enderlein, Dtsch. ent. Z., 1936, p. 104.

Bolivia.

maculipennis Enderlein, Dtsch. ent. Z., 1936, p. 103.

Bolivia.

DICTYOCAMPSA Enderlein
guttata Enderlein, Dtsch. ent. Z., 1936, p. 100.

Brazil.

1 Second Contribution to a Check-list of Psychodidae of the World.

22

Journal New York Entomological Society

[Vol. LUX

FLEBOTOMUS2 Rondani

alphabeticus da Fonseca, Rev. Ent. Rio de J., Vol. 6, p. 327.
Brazil.

ulmazani .0alliard, Ann. Parasit. hum. comp., Vol. 12, p. 193.

, Mexico.

umarali Barretto and Coutinho, Fac. Med. S. Paulo Univ., Vol.
16 (1940), p. 134.

Brazil.

amazonensis Root, Amer. Jour. Hygiene, Vol. 20 (1934), p. 244.
Peru.

antunesi Coutinho, Bol. Bio. (n.s.), Vol. 4 (1939), p. 181.

Bolivia.

aragaoi de Costa Lima, Mem. Inst. Oswaldo Cruz, Vol. 26, p. 48.
Brazil.

araozi Paterson and Shannon, La Semana Med., Vol. 33 (1926),
p. 236.

Argentina.

arthuri da Fonseca, Rev. Ent. Rio de J., Vol. 6, p. 324.

Brazil.

atroclavatus Knab, Ins. Ins. Mens., Vol. 1 (1913), p. 135. ( F .

tejereoe Larrousse, Etud. Syst. et Med. des Phleb., 1921,
P- 71.)

Trinidad, Venezuela.

avellari da Costa Lima, Mem. Inst. Oswaldo Cruz, Vol. 26, p. 48.
Brazil.

ayrozai Barretto and Coutinho, Fac. Med. S. Paulo Univ., Vol. 16
(1940), p. 131.

Brazil.

braziliensis da Costa Lima, Mem. Inst. Oswaldo Cruz, Vol. 26,
p. 48.

Brazil.

brumpti Larr., Bull. Soc. Path. Exot., Vol. 13 (1920), p. 659.
Brazil.

cardosoi Hereim*- Barretto and Coutinho, Papeis avalsos Dep.
Zool. Sao Paulo, Vol. 1 (1941), p. 148.

Brazil.

2 Phlebotomus of authors.

Mar., 1945]

Rapp: Psychodid^e

23

cavernicolus da Costa Lima, Mem. Inst. Oswaldo Crnz, Vol. 26,
p. 49.

Brazil.

castroi -Pereira7 Barretto and Coutinho, Papeis avnlsos Dep. Zool.

Sao Paulo, Vol. 1 (1941), p. 180.

^ Brazil.

chagasi da Costa Lima, Acta Medica Bio de J., Vol. 7, p. 3.

Brazil.

eolombianus Bistorcelli and Day Van Ty, Parasit. hum. comp.,
Vol. 18 (1941), pp. 73, 263.

Columbia.

cortelezzii Brethes, La Semana Medica, Vol. 30 (1923), p. 361.
Argentina.

cruciatus Coq., Ent. News, Vol. 18 (1907), p. 102.

Guatemala, Panama, Trinidad.
cruzi Mangabeira, Mem. Inst. Oswaldo Cruz, Vol. 33, p. 351.
Brazil.

davisi Root, Amer. Jour. Hygiene, Vol. 20 (1934), p. 242.

Brazil.

evandroi Costa Lima and Antunes, Brazil Medico, Vol. 50 (1930),
p. 419.

Brazil.

fischeri Pinto, Sciencia Med., Vol. 4 (1926), p. 373.

Brazil.

fonsecai da Costa Lima, Mem. Inst. Oswaldo Cruz, Vol. 26, p. 49.
Brazil.

gaminarai Cordero, Vogelsang and Cossio, Cuarta Reun. Soc.

Arg. de Pat. Reg. del. Norte, 1928, p. 649. (Bol. Inst.
Clin. Quir., No. 25-31, 1928.)

Uruguay.

guimaraesi Coutinho, -Pereir^- and Barretto, Rev. Biol. Hyg. S.
Paulo, Vol. 11 (1941),

Brazil.

intermedins Lutz and Neiva, Memorias do. Instituto Oswaldo
Cruz, Vol. 5, p. 92.

Brazil.

lanei 'Re*ieir&r Barretto and Coutinho, Papeis avulsos Dep. Zool.
Sao Paulo, Vol. 1 (1944), p. 184.

Brazil.

24

Journal New York Entomological Society

[Vol. LIII

h

I

lentfl Mangabeira, Mem. Inst. Oswaldo Cruz, Vol. 33, p. 350.
Brazil.

limai Barretto and Coutinho, Fac. Med. S. Paulo Univ., Vol. 16
(1940), p. 127.

Brazil.

lloydi Antunes, Rev. Biol, hyg., Vol. 8, p. 24.

Brazil.

longipalpis Lutz and Neiva, Memorias do Instituto Oswaldo Cruz,
Vol. 5 (1912), p. 90.

Brazil, Venezuela.

lutzianus da Costa Lima, Mem. Inst. Oswaldo Cruz, Vol. 26, p. 48.
Brazil.

mangabeirai ¥e rmm^Barretto and Coutinho, Papeis avulsos Dep.
Zool. Sao Paulo, Vol. 1 (1941), p. 151.

Brazil.

mazzai Paterson^ Bol. del. Inst, de Clin. Zuir, Vol. 67 (1926).
Argentina.

maracayensis Nunez Tovar, Mosq. y Fleb. de Venez, 43 (1924) ;

Dyar and Nunez Tovar, Ins. Ins. Mens., Vol. 14 (1927),
p. 155.

Venezuela.

migonei Franca, Bull. Soc. Part. Sci. Nat., Vol. 8 (1920), p. 230.
Paraguay.

monticolus da Costa Lima, Mem. Inst. Oswaldo Cruz, Vol. 26,
p. 50.

Brazil.

neivai Pinto 1926, Sciencia Med. (Rio), Vol. 4 (1926), p. 371.
Brazil.

nitzulescui da Costa Lima, Mem. Inst. Oswaldo Cruz, Vol. 26,
p. 46.

Brazil.

noguchii Shannon, Science (n.s.), Nov. 23, Vol. 68 (1928), p. 494.
Peru.

oliverioi 'Pereira** Barretto and Coutinho, Papeis avulsos Dep.
Zool. Sao Paulo, Vol. 1 (1941), p. 223.

Brazil.

osornoi Ristorcelli and Dao Van Ty, Ann Parasit. hum. comp.,
Vol. 18 (1941), pp. 72, 260.

Columbia.

Mar., 1945]

Rapp: Psychodhle

25

panamensis Shannon, Jour. Wash. Acad. Sci., Vol. 16 (1926),

p. 192.

Panama, Venezuela.

paraensis da Costa Lima, Acto Medico Rio de J., 7, p. 7.
Brazil.

peruensis Shannon, Science (n.s.), Nov. 9, Vol. 68 (1928), p. 494.
Peru.

pestanai Pweira? Barretto and Coutinho, Papeis avulsos Dep.
Zool. Sao Paulo, Vol. 1 (1941), p. 144.

Brazil.

pessoai Coutinho, Ferreira- and Barretto, Rev. Biol. Hyg. S.

Brazil.

pintoi da Costa Lima, Mem. Inst. Oswaldo Cruz, Vol. 26, p. 48.
Brazil.

quinquefer Dyar, Am. Jour. Hygiene, Vol. 10 (1929), p. 114.
Argentina.

rostrans Summers, Bui. Ent. Res., Vol. 3 (1912), p. 209.

Brazil.

rickardi da Costa Lima, Rev. med. Cirurg., Vol. 44 (1936), p. 288.
Brazil.

sallesi Ayrosa-Galvao and Coutinho, Ann. Fac. Med. S. Paulo,
Vol. 15 (1939), p. 125.

Brazil.

shannoni Dyar, Aiper. Jour. Hygiene, Vol. 10 (1929), p. 121.
Panama.

singularis da Costa Lima, Mem. Inst. Oswaldo Cruz, Vol. 26, p. 50.
Brazil.

trinidadensis Newstead, Ann. Trop. Med. & Parasit., Vol. 18,

Trinidad.

verrucarum Towns. Ins. Ins. Mens., Vol. 1 (1914), p. 107.

Peru.

walkeri Newstead, Bui. Ent. Res., 5 (1914), p. 190.

Found on boundary between Brazil and Bolivia.
whitmani Antumes and Coutinho, Bol. Biol, (n.s.), Vol. 4 (1939),

p. 448.

yucatanensis Galliard, Ann. Parasit. hum. comp., Vol. 12, p. 1.

Paulo, Vol. 10 (1940), pp. 89-103.

p. 47.

Mexico.

26

Journal New York Entomological Society

[Vol. LIII

LEPIDIELLA Enderlein
lanuginosa Enderlein, Dtsch. ent. Z., 1936, p. 89.

Peru.

LEPRIA Enderlein

squamosa Enderlein, Dtsch. ent. Z., 1936, p. 112.

Costa Rica.

MARUINA F. Muller

hirta Johannsen, J. Agric. Univ. P. Rico, Vol. 22, p. 224.

Porto Rico.

indica Feuerborn, Arch. Hydrobiol. Suppl., Vol. 11 (1932), p. 58.
Brazil.

pilosella F. Muller, Tr. Ent. Soc. London, 1895, p. 483.

Brazil.

spinosa F. Muller, Tr. Ent. Soc. London, 1895, p. 483.

Brazil.

Ursula F. Muller, Tr. Ent. Soc. London, 1895, p. 483.

Brazil.

MECYSMIA Enderlein

schoenemanni Enderlein, Dtsch, ent. Z., 1936, p. 108.

Chile.

NEMONEURA Tonnoir

dealbata Tonnoir, Dipt. Patagonia and S. Chile, pt. 2, p. 29.
Chile.

punctata (Phil.) ( Psychoda punctata Philippi), Verh. zool. bot.
Ges. Wien, p. 631.

Chile.

NEMOPALUS Alexander

A

dampfianus Alexander, Rev. Ent. Rio de J., Vol. 11 (1940), p. 706.
Brazil.

pallipes Shannon and del Pone, Rev. Inst. Bact., Buenos Aires,
Yol. 4, pp. 733-734. A
Argentina.

pilipes , Tonnoir, Ann. Soc. Ent. Belgique, Vol. 62 (1922), pp.
130-134.

Argentina.

sziladyi Tonnoir, 6 Congr. Int. Ent., Madrid.

Costa Rica.

Mar., 1945]

Rapp: Psychodid^e

27

vexans Alexander, Rev. Ent. Rio de J., Vol. 11 (1940), p. 798.
Brazil.

PERICOMA Walker

albitarsis Williston, Trans. Ent. Soc. Bond., 1896, p. 284.

St. Vincent, West Indies.
busckana Dyar, Inc. Ins. Mens., Vol. 14, p. 107.

Martinique.

chilensis Tonnoir, Dipt. Patagonia and S. Chile, pt. 2, p. 14.
Chile.

complicate/, Tonnoir, Dipt. Patagonia and S. Chile, pt. 2, p. 15.
Chile.

contigua Tonnoir, Dipt. Patagonia and S. Chile, pt. 2, p. 24.
Chile.

fenestrata Tonnoir, Dipt. Patagonia and S. Chile, pt. 2, p. 20.

Chile.

edwardsi Tonnoir, Dipt. Patagonia & S. Chile, pt. 2, p. 20.

Chile.

griseata Tonnoir, Dipt. Patagonia and S. Chile, pt. 2, p. 22.
Chile.

inornata Tonnoir, Dipt. Patagonia and S. Chile, pt. 2, p. 23.
Chile.

nigropimetata Schiner, Reise der Novara, Dipt., 1868, p. 29.
Brazil.

niveopunctata Tonnoir, Dipt. Patagonia and S. Chile, pt. 2, p. 18.
Chile.

pallidula Tonnoir, Dipt. Patagonia and S. Chile, pt. 2, p. 16.
Chile.

pictipennis Tonnoir, Ann. Soc. Ent. Belgique, Vol. 60, p. 155.
Jamaica.

simplex Tonnoir, Dipt. Patagonia and S. Chile, pt. 2, p. 25.
Chile.

speciosa Tonnoir, Dipt. Patagonia and S. Chile, pt. 2, p. 11.
Chile.

sqaamita^sis Tonnoir, Dipt. Patagonia and S. Chile, pt. 2, p. 21.
Chile.

viduata Tonnoir, Dipt. Patagonia and S. Chile, pt, 2, p. 26.

Chile.

28

Journal New York Entomological Society

[Vol. Lin

PLATYPLASTINX Enderlein

solox Enderlein, Dtsch. ent, Z., 1936, p. 107.

Costa Rica.

sor dellii Shannon and Del Ponte, Rev. del Inst. Bact., Vol. 4
(1927), p. 730.

Argentina.

squamiventris Lntz and Neiva, Memorias do Instituto Oswaldo
Cruz, Vol. 5 (1912), p. 89.

Brazil.

tejeraoe Larr., Travail Lab. Parasitologie Fac. Med. Paris, 1921,
p. 71.

Venezuela.

PSYCHODA Latreille

alternata Say ( Tipula phalcenoides var. Scop.) ( Trichoptera
phalcenoides Meig.) (P. phalcenoides Meig.) (P. sex-
punctata Curtis) (P. marginepunctata Roser) (P.
schizura Kincaid) (P. floridica Haseman) (P. noctur-
nala .Haseman) (P. bengalensis Brunetti) (P. albi-
maculata Welch) (P. dakotensis Dyar), Long’s Exp.
St. Peter’s River, App., 1824.

West Indies: St. Vincent.

amplipenna Knab, Proc. U. S. Nat. Mus., Vol. 46, p. 103.

Cuba.

angustipennis Williston, Dipt. St. Vincent, p. 284.

West Indies : St. Vincent.

antennalis Williston, Dipt. St. Vincent, p. 283.

West Indies : St. Vincent.

dubia Bigot, Miss. Sci. Cap. Horn, Vol. 6 (1888), p. 4.

Cape Horn.

duripuncta Curran, Dept, Sci. and Agr. Jam. Ent. Bui. No. 4,
pt. 3 (1928), App., p. 30.

Jamaica.

erecta Curran, Dept. Agr. Jam. Ent. Bui. No. 4, pt. 1, 2 (1926),
App.

Jamaica.

fimbriatissima Blanchard, Gay hist, fisica de Chili, 1852, p. 468.

Chile.

Mar., 1945]

Rapp: Psychodid^

29

fumata Knab, Proc. U. S. Nat. Mus., Vol. 46, p. 104.

Mexico.,

grisescens Tonnoir, Ann. Soc. Ent. Belg., 62 (1922), pp. 87, 88.
Jamaica.

guianica Curran, Bui. Am. Mus. Nat. Hist., Vol. 66 (1934), p. 309.
British Guiana.

helicis Dyar, Proc. Ent. Soc. Wash., Vol. 31 (1929), p. 63.

Cuba.

hirtiventris, Tonnoir, Ann. Soc. Ent. Belgique, Vol. 60 (1920),

p. 151.

Brazil.

hyalinata Blanchard, Gay hist, fisica de Chili, 1852, p. 468.

Chile.

incomplete Knab, Proc. U. S. Nat. Mus., Vol. 46, p. 105.

Panama.

interdicta Dyar, Proc. Ent. Soc. Wash., Vol. 30 (1928), p. 88.

West Indies: Santa Lucia, Gastries.
notata Blanchard, Gay hist, fisica'de Chili, 1852, p. 468.

Chile.

pallens Williston, Dipt. St. Vincent, p. 283.

West Indies: St. Vincent.

pseudocompar Tonnoir, Dipt. Patagonia and S. Chile, pt. 2, p. 7.
Chile.

pulla Rondani, Arch. Canestr., Vol. 3 (1865), p. 90.

Chile.

punctata Philippi, Verh. zool. bot. Gesellsch., Vol. 15 (1865),
p. 631.

Chile.

punctatella Townsend, Ann. and Mag. Nat. Hist., Vol. 19, p. 19.
Vera Cruz.

septempunctata Philippi, Verh. zool. bot. Gesellsch., Vol. 15
(1865), p. 631.

Chile.

simillima Tonnoir, Dipt. Patagonia and S. Chile, pt. 2, p. 8.
Bariloche.

stellulata Loew, Neue Beitr., Vol. 2 (1854).

Brazil.

subimmaculata Tonnoir, Dipt. Patagonia and S. Chile, pt. 2, p. 6.
Chile.

30

Journal New York Entomological Society

[Vol. LIII

tenella Philippi, Verh. zool. bot. Gesellsch., Vol. 15, p. 631.
Chile.

tricolor Knab, Proc. U. S. Nat. Mus., Yol. 46, p. 105.
Panama.

venusta Weyembergh, Tijdschr. Ent., Yol. 29 (1886), p. 132.
Argentina.

SYCORAX Curtis

ckilensis Tonnoir, Dipt, Patagonia and S. Chile, pt. 2, p. 31.
Chile.

SYNTOMOZA Enderlein
niveitarsis Enderlein, Dtsch. ent. Z., 1936, p. 89.

Peru.

TELMATOSCOPUS Eaton

albipunctatus (Willist.) (Psyckoda albipunctatus Williston) (T.

meridionalis Eaton) ( Psyckoda snowii Haseman), Ent.
News, Yol. 5 (1893), p. 113.

Cosmopolitan: — Bahamas, Bermuda, Panama, Porto Rico,
Haiti, Mexico, Costa Rica, British Guiana, Jamaica,
Cuba, West Indies: St. Domingo, Tabago Islands, Trini-
dad.

TERMITADELPHOS Holmgren
silverstrii Holmgren, Zool. Ang., Yol. 29 (1905), p. 530.

Peru.

wasmanni Holmgren, Zool. Anz., Yol. 29 (1905), p. 533.

Peru.

TONNOIRA Enderlein

pelliticornis Enderlein, Dtsch. ent. Z., 1936, p. 106.

Peru.

TRICHOMYIA Curtis

cirrata Coquillett, Jour. N. Y. Ent. Soe., Vol. 10, p. 137.

Mexico.

edwardsi Tonnoir, Dipt. Patagonia and S. Chile, pt. 2, p. 29.
Chile.

Mar., 1945]

Brown: Butterflies

31

NOTES ON MEXICAN BUTTERFLIES, V

F. Martin Brown

NYMPHALIDiE, II
Tribe Vanessidi

323. Nymphalis antiope Linnaeus.

G. & S., 1: 215; 2: 682 (as Vanessa).

S., p. 457, pi. 93f.

H. , p. 680.

Galeana, Nuevo Leon, 6500 ft., 1 $ vii.30.39 (H.H.).

El Sabino, Uruapan, Michoacan, 1 2 vii. 15-30.36
(H.D.T.).

The Galeana specimen is battered, the other fresh.

324. Vanessa atalanta Linnaeus.

G. & S., 1: 219 (as Pyrameis) .

S., p. 458, pi. 94a.

H. , p. 680.

Galeana, Nuevo Leon, 6500 ft., 1 J' viii.4.39 (H.H.).
Las Adjuntas, Nuevo Leon, 2500-3000 ft., 1 $ vi.21.40
(H.H.).

Orizaba, Vera Cruz, 2000 ft., 1 2 v.6.41 (R.P.).
Tancitaro, Michoacan, 6586 ft., 1 2 viii.11.40 (H.H.).
The Galeana and Orizaba specimens are fresh.

325. Vanessa virginiensis Drury.

S., p. 459, pi. 94b.

H, p. 680.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 12 vi.18.40 (H.H.).

Tancitaro, Michoacan, 6586 ft., 1 2 vii. 25.40 (H.H.).

326. Vanessa cardui Linnaeus.

G. & S., 1: 217 (as Pyrameis).

S., p. 458.

32

Journal New York Entomological Society

[Vol. Lin

Tancitaro, Michoacan, 6586 ft., 1 2 52 vii. 20-25.40

(H.H.).

These are worn specimens but they seem a little paler than
those we take at the same altitude in Colorado.

328-9. Junonia lavinia Cramer.

Hoffmann separates this species into two, coenia Hubner (328)
and genoveva Cramer (329). I believe these to be nothing more
than forms of lavinia Cramer, which name has several years
priority over genoveva.

G. & S. 1 : 220 ; 2 : 684 (as coenia) .

S., p. 461-2, pi. 94d & e.

H. , p. 681.

Sabinas Hidalgo, Nuevo Leon, 960 ft., 1 § vi.25.39
(H.H.) (f. coenia).

Galeana, Nuevo Leon, 6500 ft., 1 £ vii. 1.39 (H.H.) (f.
lavinia).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 $
v.29.41 (R.P.) (f. lavinia).

El Banito, Valles, San Luis Potosi, 200 ft., 1 5 vi.28.40
(H.H.) (f. genoveva) .

Chichen Itza, Yucatan, 2 viii. 30.36 (H.D.T.) (f.
lavinia) .

Sta. Lucrecia, 2 ix. 24.39 (H.D.T.) (f. genoveva).

El Sabino, Uruapan, Michoacan, 3 1 J vii.15-30.36

(H.D.T.) (f. coenia).

330. Anartia jatrophe Johannes.

G. & S., 1: 221 (as iatrophe).

S., p. 462, pi. 94e.

H. , p. 681.

Sabinas Hidalgo, Nuevo Leon, 960 ft., 1 5 vi. 15.39
(H.H.).

Monterrey, Nuevo Leon, 1 § vi.15.35 (H. A. Freeman).

60 mi. So. of Victoria, Tamaulipas, 3 1 5 vii. 6. 36

(H.D.T.).

Jacala, Hidalgo, 4500 ft., 1 vii. 11. 39 (H.H.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft. 1 £ 3 §5
v.28,41 (R.P.).

Mar., 1945]

Brown: Butterflies

33

El Banito, Valles, San Lnis Potosi, 200 ft., 4
vi. 27-28.40 (H.H.) ; 2 1 $ iv.30.41 (R.P.).

El Pujal, San Lnis Potosi, 100 ft., 3 £<$ vii.17-21.39
(H.H.).

Hda. Potrero Viejo, Paraje Nuevo, Vera Cruz, 1500 ft.,
1 v.5.41 (R.P.).

Rio Blanco, Vera Cruz, 2200 ft., 1 2 v. 10.41 (R.P.).

Sta. Lucrecia, 1 g 1 $ ix. 24.39 (H.D.T.).

El Capirie, Michoacan, 500 ft., 1^1$ viii.3.40 (H.H.).

El Sabino, Uruapan, Michoacan, 3 J'J' 1 2 vii. 15-30.36
(H.D.T.).

331. Anartia fatima Fabricius.

G. &S., 1:222.

S, p. 463, pi. 94f.

H, p. 681. .

Galeana, Nuevo Leon, 6500 ft., 3 c?c? iv.29.41 (R.P.)
(f. albifaciata) .

nr. Villagran, Tamaulipas, 1160 ft., 1 vi.28.41 (R.P.)

(f. albifaciata) .

C. Victoria, Tamaulipas, 1 2 vi.19.39 (H. A. Freeman)
(f. albifaciata) .

60 mi. So. of Victoria, Tamaulipas, 2 1 2 vii. 6. 36

(H.D.T.) (f .fatima).

Jacala, Hidalgo, 4500 ft., 4 J'J' vii.1-11.39 (H.H.) (f.
fatima) .

El Sol, Tamazunchale, San Luis Potosi, 400 ft. 6
1 2 v.28-29.41 ; 2 iv.30-31.41 (R.P.) (If.

fatima & albifaciata) .

Arroyo del Calabezas, San Luis Potosi, 250 ft., 1 J'
iv.30.41 (R. P.) (f. albifaciata) .

El Banito, Valles, San Luis Potosi, 200 ft., 4 <$■<$ 1 2
vi. 26-27.40 (H.H.) ; 4 <?<? 3 ?? iv.30.41 (R.P.) (ff.
fatima & albifaciata) .

El Pujal, San Luis Potosi, 100 ft., 8 J'J' 2 22 vii.17-21.39
(H.H.) (ff. fatima & albifaciata) .

Tuxpango, Vera Cruz, 1500 ft., 4 v.9.41 (R.P.) (f.

fatima) .

34

Journal New York Entomological Society

[VOL. LIII

Fortin, Vera Crnz, 1600 ft., 1 y v.4.41 (R.P.) (f. albi-
faciata).

Ojo de Agua, Vera Cruz, 1600 ft., 2 yy v.12.41 (R.P.)
(f . fatima) .

Orizaba, Vera Cruz, 2000 ft., 3 yy 1 $ v.6.41 (R.P.)
(if. albifaciata & fatima).

Chichen Itza, Yucatan, 2 yy 1 5 viii.30.36 (H.D.T.)
(ff. fatima & albifaciata) .

Sta. Lucrecia, 2 yy ix.24.39 (H.D.T.) (if. fatima &
venusta) .

El Capirie, Michoacan, 500 ft., 1 y viii.3.40 (H.H.) (f.
venusta).

Apatzingan, Michoacan, 1050 ft., 1 y viii.5.40 (H.H.)
(f.venusta) .

El Sabino, Uruapan, Michoacan, 5 yy 1 § vii.15-30.36
(H.D.T.) (ff. fatima , venusta & albifaciata) .

The distribution of the various forms of this species is interest-
ingly told by the above list of specimens. I cannot agree with
Holfmann as to the status of venusta. I believe it to be a west
coast form and he considers it to be a west coast race.

333. Victoria stelenes biplagiata Fruhstorfer.

G. & S., 1: 280.

S., p. 463, pi. 95a (as steneles).

H. , p. 681 (as steneles) .

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 9 yy

v. 28-29.41 (R.P.).

El Bahito, Valles, San Luis Potosi, 200 ft., 2 5?

vi. 27-28.40 (H.H.) ; 1 y iv.30.41 (R.P.).

El Pujal, San Luis Potosi, 100 ft., 1 y 2 5? vii.17-18.39
(H.H.).

Chichen Itza, Yucatan, 1 y 1 $ viii. 30.36 (H.D.T.)
(y = f. pallida Fruh.).

El Sabino, Uruapan, Michoacan, 1 y 4 $$ vii.15-30.36
(H.D.T.).

334. Victorina epaphus Latrielle.

S., p. 464, pi. 95b.

H., p. 681.

Mar., 1945]

Brown: Butterflies

35

Tuxpango, Vera Cruz, 1500 ft., 1 <$ v.8.41 (R.P.).

Ojo de Agua, Vera Cruz, 1600 ft., 1 v.12.41 (R.P.)-

336. Hypanartia lethe Fabricius.

G. & S., 1: 212; 2: 682 (as Eurema).

5., p. 460, pi. 94b.

H. , p. 682.

Tuxpango, Vera Cruz, 1500 ft., 1 2 v.9.41 (R.P.).
Fortin, Vera Cruz, 1600 ft., 2 2 22 v-4.41 (R.P.).

Orizaba, Vera Cruz, 2000 ft., 2 <■?<? v.6.41 (R.P.).

337. Hypanartia godmani Bates.

G. & S., 1: 213 (as Eurema).

5., p. 460, pi. 94c.

H. , p. 682.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 <$

iv. 31’41 (R.P.).

Orizaba, Vera Cruz, 2000 ft., 1 2 v.6.41 (R.P.).

Tribe Didonidi

340. Didonis aganisa Boisduval.

G. & S., 1: 276; 2: 690.

S., p. 465, pi. 94f.

H. , p. 682.

Arroyo del Meco, Tamaulipas, 1320 ft., 1 iv.28.41

(R.P.).

C. Victoria, Tamaulipas, 1 J1 vi.25.35 (H. A. Freeman).
Hda. Sta. Engracia, Tamaulipas, 2 22 vii.24.39 (H.H.).
Jacala, Hidalgo, 4500 ft., 1 £ vii.2.39 (H.H.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1^12

v. 28-29.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 1^22?

vi. 27-29.40 (H.H.).

El Pujal, San Luis Potosi, 100 ft., 1 $ vii. 20.39 (H.H.).
Ojo de Agua, Vera Cruz, 2200 ft., 1 J' v.12.41 (R.P.).
Rio Blanco, Vera Cruz, 2200 ft., 1 v.10.41 (R.P.).

Bet. Tule and Mitla, Oaxaca, 1 2 v.18.41 (R.P.).

36

Journal New York Entomological Society

[Vol. LIII

341. Mestra amymone Menetries.

G. & S., 1: 278; 2: 690 (as Cystineura) .

S., p. 467, pi. 115e.

H. , p. 682.

Sabinas Hidalgo, Nuevo Leon, 960 ft., 3 yy vi.15.39
(H.H.).

Villa Santiago, Nuevo Leon, 1600 ft., 1 y viii.8.39
(H.H.).

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 1 y vi.18.40 (H.H.).

Monterrey, Nuevo Leon, 1800 ft., 1 y 1 § iv.27.41
(R.P.).

nr. Villagran, Tamaulipas, 1160 ft., 1 y 3 $? iv.28.41
(R.P.).

Jacala, Hidalgo, 4500 ft., 4 yy 2 ?? vii.2-3.39 (H.H.).
El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 y

v. 28.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 5 yy

vi. 26-27.40 (H.H.).

Rio Balsas, Guerrero, 2400 ft., 2 yy 1 5 v.26.41 (R.P.).
Acahuato, Michoacan, 3000 ft., 1 y 1 § viii.2.40 (H.H.).
Apatzingan, Michoacan, 6586 ft., 1 y viii.5.40 (H.H.).
The west coast specimens tend to be more strongly marked
than the east coast material. However, the difference is slight
and no one would fail to recognize the material from the two
coasts as being the same species.

Tribe Epicaliidi

344. Pyrrhogyra otolais neis Felder.

G. & S., 1: 293.

S., p. 475.

H, p. 682.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 2 yy
1 ? iv. 30-31.41 ; 1 y 1 ? v.28.41 (R.P.).

Ojo de Agua, Vera Cruz, 1600 ft., 1 y v.12.41 (R.P.).

345. Pseudonica flavilla canthara Doubleday.

S., p. 476, pi. 97b.

H., p. 682.

Mar., 1945]

Brown: Butterflies

37

Ojo de Agua, Vera Cruz, 1600 ft., 1 ^ 1 J v.12.41
(R.P.).

These specimens are even more lightly marked than the one
figured by Seitz ( l.c .). The submarginal band on the hindwings
is reduced to a dot at the anal angle.

346. Temenis laothoe liberia Fabricius.

S., p. 476, pi. 97c.

H., p. 683.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 4' J'J'
v. 28.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 1 2 vi.28.40
(H.H.); 1 ^iv.30.41 (R.P.).

348. Epiphile adrasta bandusia Fruhstorfer.

G. & S., 1: 234; 2: 685.

S., p. 479.

H. , p. 683.

Galeana Canyon, Tamaulipas, 1 J1 iv.29.41 (R.P.).

El Sol, Tamazunchale, San Luis Potosi, 2 J'J' 1 2
v. 28-29.41 (R.P.).

El Banito, Valles, San Luis Potosi, 1 vi.26.40 (H.H.).

349. Catonephele nyctimus Westwood.

S., p. 480, pi. 98c.

H, p. 683.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 J' 1 2
v.28-29.41 (R.P.).

Tuxpango, Vera Cruz, 1500 ft., 1 v.8.41 (R.P.).

352. Mycelia cyaniris Doubleday.

G. &S., 1:230.

R., p. 483, pi. 99a (as ethusa J*).

H. , p. 683.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1
iv.30.41 ; 2 JV? v.29.41 (R.P.).

El Pujal, San Luis Potosi, 100 ft., 1 ^ vii.20.39 (H.H.).
Ojo de Agua, Vera Cruz, 1600 ft., 1 2 v.12.41 (R.P.).
This species may easily be confused with the females of the
following species when the upper surfaces are compared. It may

38

Journal New York Entomological Society

[Vol. LIII

always be recognized by the more strongly falcate forewings and
the presence of a single white band running the length of the
cell. The underside of cyaniris is much lighter and uniformly
colored than that of ethusa. The two species fly together. The
ratio of their occurrence in this collection is 1:6! This reverses
the usually accepted notion of the rarity of the two species.
(See G. & S., l.c.).

353. Mycelia ethusa Boisduval.

G. & S., 1 : 232.

R., p. 483 (pi. 99a J as cyaniris ?).

H. , p. 683.

El Mante, San Luis Potosi, 1 2 iv.29.41 (R.P.) (? §
ethusa or aberrant 2 rogenhoferi ?)

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 8
v.28-29.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 2
vii.22.39 ; 10 iv.26-29.40 (H.H.) ; 5 $$ iv.30.41

(R.P.).

El Pujal, San Luis Potosi, 100 ft., 7 vii.17-20.39

H.H.).

See the note following 353A, Mycelia rogenhoferi Felder.

353A. Mycelia rogenhoferi Felder.

G. & S., 1: 231, pi. 24, ff. 3, 4.

R., p. 483, pi. 99a.

H. , p. 683 (as syn. of ethusa).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 J* 1 2
v.29.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 2 22
vii.22.39; 1 ? vi.26.40 (H.H.)

El Pujal, San Luis Potosi, 100 ft., 3 ££ 2 22 vii.17-21.39
(H.H.).

I am puzzled about this material that I call rogenhoferi. Hoff-
mann (l.c.) lists it as a synonym of the female of ethusa. Yet I
have males and females that are identically marked. The mark-
ings in the cell of the forewing on the upper side corresponds
exactly in the two sexes with the figure given by Godman &
Salvin and copied by Roeber. This is quite different from the

Mar., 1945]

Brown: Butterflies

39

markings found on ethusa — Roeber’s ethusa is cyaniris (pi. 99a)
and in the same line his cyaniris $ may be ethusa J but is more
likely a rogenhoferi. The female that I have listed under ethusa
from El Mante may be a slightly aberrant rogenhoferi ; it is like
Roeber’s figure called cyaniris but has four distinct bluish
bands on the hindwing while the figure shows only three plus
some lunules at the margin. I am bothered by the lack of any
clear-cut females of ethusa in such a long series. I may be in
error about the males noted above and Holfmann may be correct.

354. Mycelia cyananthe Felder.

R. , p. 483.

H., p. 683.

Rio Balsas, Guerrero, 2400 ft., 1 J' 1 J v.26.41 (R.P.).
The specimens are badly battered but recognizable.

355. Eunica tatila Herrick-Schaffer.

G. & S., 1: 229; 2: 685.

S. , p. 484, pi. lOOAa.

H. , p. 683.

Galeana Canyon, Tamaulipas, 12 iv.29.41 (R.P.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 17 J'J'
v.28-29.41 (R.P.).

The light spots on the forewing are very faint on the Galeana
Canyon material. However the wings have a distinct purple
reflection.

355A. Eunica caerula Godman & Salvin.

G. & S., 1: 229.

S., p. 484, pi. lOOAa.

H. , p. 683.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 £
v.28.41 (R.P.).

Hoffmann considers this a synonym of tatila H.^S. I do not.

356. Eunica monima Cramer.

G. & S., 1: 228.

S., p. 485, pi. lOOAc.

H. , p. 684.

Galeana Canyon, Tamaulipas, 2 $5 iv.29.41 (R.P.).

40

Journal New York Entomological Society

[Vol. LIII

El Sol, Tamazunchale, San Luis Potosi, 1 g 1 § v.28-
29.41 (R.P.).

Tribe Catagrammidi
369. Callicore anna Guerin.

G. & S., 1: 255; 2: 688.

R., p. 501, pi. 102Bc.

H, p. 685.

Tamazunchale, San Luis Potosi, 1 5 vii.1.36 (H. A.
Freeman).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 £ 3 55
iv.31.41 (R.P.).

Tuxpango, Vera Cruz, 1500 ft., 2 <$$ v.9.41 (R.P.).
Fortin, Yera Cruz, 1600 ft., 1 1 5 v.3.41 (R.P.).

374. Dynamine mylitta Cramer.

R, p. 507, pi. lOlAh.

H., p. 685.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 15
5 5$ v.28-29.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 1 g vi.28.40 ;
1 5 vii.22.39 (H.H.).

El Pujal, San Luis Potosi, 100 ft., 2' J'J' 1 5 vii.17-20.39
(H.H.).

Ojo de Agua, Yera Cruz, 1600 ft., 2 55 v.12.41 (R.P.).

376. Dynamine dyonis Geyer.

G. & S., 1: 250.

R., p. 508, pi. lOlAh, i, lOlBa.

H. , p. 685.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 $
iv.30.41; 7 v. 28-29.41 (R.P.).

El Banito, Yalles, San Luis Potosi, 200 ft., 1 J' vi. 27.40
(H.H.) ; 1 iv.30.41 (R.P.).

El Pujal, San Luis Potosi, 100 ft., 1 5 vii.17.39 (H.H.).

Tribe Ageroniidi
379. Ageronia februa Huebner.

F., p. 539, pi. 105d.

Mar., 1945]

Brown: Butterflies

41

H, p. 685.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 8 J'J'
6 2? v.28-29.41 (R.P.).

El Banito, Yalles, San Luis Potosi, 200 ft., 8 J'J' vi.26-
29.40 (H.H.) ; 2 A? iv.30.41 (R.P.).

El Pujal, San Luis Potosi, 100 ft., 2 1 2 vii. 17-18.40

(H.H.).

Bio Blanco, Yera Cruz, 2200 ft., 1 J' v.10.41 (R.P.).
Acapulco, Guerrero, 100 ft., 1 <$ v.26.41 (R.P.).

I agree with Hoffmann that Fruhstorf er ’s Mexican forms are
untenable.

381. Ageronia ferox fictitia Fruhstorfer.

F. , p. 541, pi. 105e.

H, p. 686.

60 mi. So. of Victoria, Tamaulipas, 2 c?c? vii.6.36
(H.D.T.).

El Pujal, San Luis Potosi, 100 ft., 1 J* vii.17.39 (H.H.).
Of this Hoffmann says, “No se conoce su distribucion geo-
grafica. ’ ’ My two records place the form on the Gulf Coast.

382. Ageronia atlantis Bates.

G. & S., 1: 269; 2: 689.

H. , p. 686.

Acapulco, Guerrero, 100 ft., 1 v.23-25.41 (R.P.).

383. Ageronia feronia farinulenta Fruhstorfer.

F., p. 542.

H, p. 686.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 4 J'J'
1 2 v.29.41 (R.P.).

El Pujal, San Luis Potosi, 100 ft., 1 $ vii.19.39 (H.H.).

384. Ageronia guatemalena marmarice Fruhstorfer.

F., p. 542.

H., p. 686.

60 mi. So. of Victoria, Tamaulipas, 2 J'J' vii.6.36
(H.D.T.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 J' 1 2
v.28-29.41 (R.P.).

42

Journal New York Entomological Society

[Vol. Lin

El Pujal, San Luis Potosi, 100 ft., 1 vii.18.39 (H.H.).

The species feronia and guatemalena can be readily separated
by observing the ocelli on the upper side of the hindwing. These
in guatemalena contain a narrow ring of blue scales around the
greyish or white pupil.

385. Ageronia iphthime Bates.

G. & S., 1: 274.

F., p. 543, pi. 105a.

H. , p. 686.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 6 <£$
3 55 v. 28-29. 41 (B.P.).

Ojo de Agua, Yera Cruz, 1600 ft., 2 v.12.41 (R.P.).

386. Ageronia fornax fornacula Fruhstorfer.

F., p. 543.

H., p. 686.

Tuxpango, Yera Cruz, 1500 ft., 1 J' v.9.41 (R.P.).

387. Ageronia amphinome mexicana Lucas.

F. , p. 543.

H., p. 686.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 £
v.29.41 (R.P.).

Tribe Marpesiidi
390. Timetes chiron Fabricius.

G. & S., 1: 288; 2: 691.

S., p. 468, pi. 96a.

H. , p. 687.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 4 J'J'

1 $ v. 28-29. 41 (R.P.).

Arroyo del Calabezas, San Luis Potosi, 250 ft., 5 J'J'
iv.30.41 (R.P.).

El Banito, Yalles, San Luis Potosi, 200 ft., 1 vii.22.39 ;

2 cf -c? 1 ? vi. 26-28.40 (H.H.).

393. Athena petreus Cramer.

G. & S., 1: 283; 2: 690 (as Timetes peleus).

S., p. 469, pi. 96e (as Megalura peleus).

Mar., 1945]

Brown: Butterflies

43

H., p. 687.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 3
1 ? v.28-29.41 (R.P.).

Arroyo del Calabezas, San Luis Potosi, 250 ft., 4 J'J'

iv. 30.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 1 J' iv.30.41
(R.P.).

El Pujal, San Luis Potosi, 100 ft., 1 <$ vii.17.39 (H.H.).
Tribe Limenitidi

The material from the genus Adelpha was purchased by Mr.
Frank Johnson and will be published by him.

414. Heterochroa bredowi bredowi Geyer.

G. & S., 1: 310.

S., p. 533 (as Limenitis).

H. , p. 688.

Hda. Vista Hermosa, Villa Santiago, Nuevo Leon, 1500
ft., 2 <?<? 1 ? vi.19-21.40 (H.H.).

415. Basilarchia archippus Cramer.

S., p. 534.

H., p. 689.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 J'

v. 29.41 (R.P.).

This is f. obsoleta Edw. recorded by Hoffmann from the north-
eastern part of Mexico.

416. Basilarchia astyanax arizonensis Edwards.

G. & S., 1 : 311 ; 2 : 693 (as Limenitis).

S., p. 534.

H. , p. 689.

Galeana, Nuevo Leon, 6500 ft., 1 $ viii.4.39 (H.H.).
Monterrey, Nuevo Leon, 1 J vi.15.35 (H. A. Freeman).
Arroyo del Meco, Tamaulipas, 1320 ft., 1 J' iv.28.41
(R.P.).

Tribe Apaturidi

419. Chlorippe pavon Latrielle.

G. & S., 1: 315; 2: 693 (as pavonii).

44

Journal New York Entomological Society

[Vol. LIII

R., p. 546, pi. llOBc.

H., p. 689.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 3 J'J'
1 5 iv.30-v.29.41 (R.P.).

Arroyo del Calabezas, San Luis Potosi, 250 ft., 1 $

iv. 30.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 2 vi. 27.40
(H.H.); 2 iv.30.41 (R.P.).

El Pujal, San Luis Potosi, 1 J1 vii.17.39 (H.H.).

424. Asterocampa leilia Edwards.

G. & S., 1: 319; 2: 694 (as Doxocopa).

R. , p. 550, pi. HOAc.

H, p. 689.

Sabinas Hidalgo, Nuevo Leon, 690 ft., 1 <$ 2 22 vi.15-
16.39 (H.H.).

Ojo de Agua, Sabinas Hidalgo, Nuevo Leon, 1000 ft.,
1 vi. 14.40 (H.H.).

424A. Asterocampa lycaon flora Edwards.

R., p. 549, pi. llOAc.

Sabinas Hidalgo, Nuevo Leon, 960 ft., 2 J'J' vi.17-18.39
(H.H.).

Villa Santiago, Nuevo Leon, 1500 ft., 1 2 vi. 17.40
(H.H.).

Galeana, Nuevo Leon, 6500 ft., 1 2 viii.4.39 (H.H.).
Hda. Sta. Engracia, Tamaulipas, 1 2 vii.25.39 (H.H.).
This species is not listed by Hoffmann.

Tribe Gynaeciidi

428. Smyrna blomfildia datis Fruhstorfer.

G. & S., 1: 329; 2: 696.

S. , p. 471, pi. 104d.

H. , p. 690.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 2

v. 29.41 (R.P.).

El Pujal, San Luis Potosi, 100 ft., 3 J'J' vii. 18-20.39
(H.H.).

Mar., 1945]

Brown: Butterflies

45

Ojo de Agua, Vera Cruz, 1600 ft., 3 $5 v.12.41 (R.P.).
The Ojo de Agua females are in tatters but recognizable.

430. Gynaecia dirce Linnaens.

G. & S., 1: 265; 2: 689.

S., p. 473, pi. 97a.

H. , p. 690.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 2 ■£$
y.29.41 (R.P.).

Chichen Itza, Yucatan, 1 J viii. 30.36 (H.D.T.).

Tribe Charaxidi

431. Prepona demophon centralis Fruhstorfer.

G. & S., 1: 320; 2: 694 (as demophon) .

F, p. 554, pi. 111a.

H, p. 690.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1
v.28.41 (R.P.).

El Pujal, San Luis Potosi, 100 ft., 2 1 $ vii. 19-20.39

(H.H.).

433. Prepona antimache gulina Fruhstorfer.

F. , p. 555.

H., p. 690.

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 1 2
v.29.41 (R.P.).

443. Protogonius hippona cecrops Doubleday & Hewitson.

G. & S., 1: 357; 2: 699.

R., p. 580, pi. 117a.

H. , p. 691.

El Pujal, San Luis Potosi, 100 ft., 1 $ vii. 21. 39 (H.H.).
Ojo de Agua, Vera Cruz, 1600 ft., 1 $ v.12.41 (R.P.).
Ancea: — Mr. Frank Johnson purchased these and will publish
upon them.

LIB Y THE ID ^

464. Libythea carinenta Cramer.

G. & S., 1: 360.

46

Journal New York Entomological Society

[Vol. LIII

S., p. 623, pi. 120De.

H., p. 693.

Sabinas Hidalgo, Nuevo Leon, 960 ft., 4 52 vi.15.39
(H.H.).

Arroyo del Meco, Tamaulipas, 1320 ft., 2 iv.28.41

(R.P.).

El Sol, Tamazunchale, San Luis Potosi, 400 ft., 2
v.29.41 (R.P.).

Arroyo del Calabezas, San Luis Potosi, 250 ft., 1
iv.30.41 (R.P.).

So. of El Mante, San Luis Potosi, 1 iv.27.41 (R.P.).

El Banito, Valles, San Luis Potosi, 200 ft., 1 2 vii. 22.39 ;
2 vi.26.40 (H.H.).

Riodinidae: — These are being studied by Mr. Wm. P. Comstock.

Lycaenidae: — These are being studied by Mr. E. I. Huntington.

Hesperioidea : — These are being studied by Mr. Ernest L. Bell.

BIBLIOGRAPHY

1. Godman & Salvin, Biologia Centrali-Americana, 1: 215-360; 2: 682-699.

2. Seitz, Macrolepidoptera of the World, 5: 457-623.

3. Hoffmann, Anales del Instituto de Biologia, 11: 639-738. 1940.

(S = Seitz, R = Roeber, F = Fruhstorf er, in Seitz ’ 1 1 Macrolepidoptera. ”

Mar., 1945]

Comstock: Violet Dos Passos

47

VIOLA HARRIET DOS PASSOS AND HER
NORTH AMERICAN MOTHS

There is romance in the making of a collection. As Milton said
in Lycidas, ‘ ‘ To-morrow to fresh woods, and pastures new. ’ ’ The
joy of a fine day in the field, a discovery, an observation, always
carried in precious memory, such, I think, was the pleasure of
Mrs. dos Passos in her collection. Much of it was of her own
collecting, and her orderly cabinets testify to her care, skill and
patience in the preparation and arrangement of her specimens.
The collection has personality, her personality reflected in it.

In the spring of 1931, at Rangeley, Maine, the collection had its
beginning, a joint undertaking, for her husband Cyril F. dos
Passos, was equally interested. Their first reference literature
was “The Butterfly Book” and “The Moth Book” by Dr. Hol-
land. It was soon decided that Mrs. dos Passos would make the
North American moths her specialty and that Mr. dos Passos
would devote his energies to the collection and study of the North
American butterflies.

Collecting at Rangeley, season after season, was productive of
many rare and interesting species. Collecting methods were im-
proved and a very large light trap was devised and constructed.
Added to this, through a beautiful white birch grove, was a trap
line, with about a dozen “Rummel type” bait traps, always pro-
viding an exhilarating morning walk, filled with surprises as each
trap revealed its content of Catocala, lesser noctuids, perhaps a
fine sphingid or two and many smaller beauties of the woods.
Then there was the sugaring trail for an exciting evening and
it is easy to understand why Rangeley Camp meant so much to
Viola and Cyril dos Passos.

Mrs. dos Passos spent some winters in Florida where she col-
lected many butterflies and some moths, as at Key West where
she captured a fine Cocytius antaeus resting on a wall. In
Florida, she made the acquaintance of many local collectors,
among them Marguerite S. Forsyth from whom she obtained a
number of Florida’s rarities.

Early in the winter of 1931 the dos Passos moved to Mendham,

48 Journal New York Entomological Society [Vol. liii

New Jersey, and there one summer was spent collecting with Dr.
and Mrs. George H. H. Tate who lived nearby. A sugaring trail,
along the wooded skirts of a swamp, gave a good sample of the
x local fauna and the variety was augmented by contributions from
the Tate collection.

The home at Mendham, in ample acres, built on a southerly
sloping hillside overlooking a brook which is the headwaters of
the Passaic River, is ideally located for those who enjoy the
beauties of nature. Virginia deer browse at evening in the upper
reaches of the swamp, birds of passage and of residence are at
home there, even the purple martin, and Viola dos Passos knew
and loved the birds. The west wing of the house provided a con-
servatory with cages for tropical birds and, at the end of the
second story, a study for the butterfly and moth collections and
library. As the collections grew, this space was entirely given
over to the butterflies, Cyril dos Passos’ preserve, and another
large room in the central wing of the house gave ample space for
Mrs. dos Passos ’ collection of moths.

Two extensive collections of Lepidoptera, one of considerable
historic interest, were acquired by Mr. dos Passos and the moths
from these collections were placed in the collection of Mrs. dos
Passos. One was the collection of Max Rothke, made for the most
part at Scranton, Pennsylvania, and the other, that of Thomas E.
Bean, containing material taken at Galena, Illinois, and the rich
collection made at Laggan, Alberta, in the seventies and eighties
of the last century.

Mrs. dos Passos suffered a heart attack, early in the year 1939,
which restricted her physical activities. This affliction resulted
in her death at Rangeley, Maine, on August 29, 1944. She was
the youngest daughter of Anthony H. Van Hise and Harriet
Louise Acker and was born at Newark, New Jersey, on November
24, 1891. She is survived by her huband, Cyril F. dos Passos,
and their son, Manuel dos Passos.

The collection of North American moths, consisting of over
12,000 specimens, and the library relating to the Heterocera were
presented to The American Museum of Natural History by Mr.
dos Passos late in the year 1944. — William P. Comstock.

Mar., 1945]

Alexander : Cranflies

49

RECORDS AND DESCRIPTIONS OF NEOTROPICAL
CRANE-FLIES (TIPULID^, DIPTERA), XIX

By Charles P. Alexander
Amherst, Massachusetts

The previous instalment under this general title was published
in 1944 (Journal of the New York Entomological Society, 52 :
369-383). Most of the materials discussed at this time are from
Costa Rica and Panama, with fewer species from the Greater
Antilles and South America. I am greatly indebted to the vari-
ous collectors for their interest in saving these flies. The types
of the novelties are preserved in my personal collection except in
the cases of three species from the United States National Mu-
seum, sent to me for examination by Dr. Alan Stone, and one fur-
ther species received from Professor J. Speed Rogers.

Genus Brachypremna Osten Sacken

Brachypremna geijskesi new species.

Size large (wing, male, over 20 mm.) ; mesonotum, especially the praescutum,
more or less inflated ; praescutum reddish brown, patterned with black, these
areas especially evident near the suture where they are separated by equally
distinct, obscure yellow spots ; legs black, the femoral tips, narrow tibial bases
and tibial tips whitened, the last broadest on hind legs ; basitarsi of fore and
middle legs more or less infuscated on proximal third ; wings pale brown, pat-
terned with darker brown and cream-colored areas ; stigma with pale center ;
tip of vein B3 curved at about a right angle into the margin ; veins Mx and M 2
not strongly decurved ; male hypopygium with the dististyle expanded at apex
into a subcircular head, this further produced into a cultriform blade.

Male. — Length about 15-17 mm.; wing 20-21 mm.

Head broken. Mesonotum, especially the praescutum, more or less inflated,
somewhat as in quasimodo but not so accentuated. Praescutum, with the
cephalic portion reddish brown, the posterior half with four blackened stripes,
these latter near the suture separated by conspicuous, obscure yellow spots
that become infuscated in front; further yellow brightenings in humeral
region and on the sides of sclerite before the suture, separated by a reddish
brown extension to the lateral border; scutum chiefly dark brown, narrowly
bordered by more brownish black, the central portion very narrowly testa-
ceous; scutellum obscure yellow, with a brown central marking; parascutella
reddish brown, margined in front by blackish; mediotergite with three dark
brown longitudinal stripes, the more restricted intervening lines obscure

50

Journal New York Entomological Society

[Vol. Lin

brownish yellow. Pleura obscure brownish yellow, spotted with dark brown.
Halteres with stem reddish brown, narrowly yellow at base, the knob black-
ened. Legs with coxae testaceous brown, the outer face, especially of the
posterior pair, marked with dark brown; trochanters obscure yellow; femora
brownish black, the bases very restrictedly brightened, the tips rather broadly
and abruptly whitened; tibiae black, the base and apex whitened, the former
about one-half as extensive as the femoral tip ; the amount of white at tibial
apex is narrowest (about distal tenth) on fore legs, somewhat more extensive
on middle legs, occupying about the distal fourth on posterior legs ; fore and
middle basitarsi just beyond their bases weakly infuscated for about one-third
their length, the posterior pair, with the remainder of tarsi, yellowish white.
Wings weakly suffused with brown, patterned with darker brown and cream-
colored areas, arranged as is common in the genus; stigma with pale center,
lying behind vein Ex; tips of veins Ei+ 5 and Mx broadly seamed with brown,
alternating with creamy spots in the centers of the cells; vein Cu, cord and
the veins beyond it, particularly the outer medial field, conspicuously seamed
with brown; veins brown. Venation: Es relatively long; E3 very strongly
curved to margin, forming a right angle ,or virtually so; veins Mx and M2 not
strongly decurved, as in quasimodo ; m-cu shortly beyond fork of M.

Abdomen with somewhat the usual pattern of the genus; basal rings yel-
low, the posterior portions of the tergites extensively infuscated, including
the eighth segment; intermediate sternites with the dark central areas long-
oval, more pointed outwardly, on outer sternites becoming much shorter;
hypopygium with proximal ends of basistyle darkened, the outer third
abruptly pale; dististyles infuscated. Male hypopygium with the dististyle
of distinctive conformation; outer spine acute at tip which is directed
strongly distad; apex of style dilated into a subcircular head that is further
produced into a cultriform blade, the apex very obtuse; before the rounded
apex on disk of blade with about 25 circular pores, each with a very small
conical peg ; in other similar species, including quasimodo, the pores and their
spines are much larger and placed closer to outer margin of style.

Holotype, J*, Brownsberg, Surinam, altitude 400 meters, Sep-
tember 19, 1938 (Geijskes). Paratype, <$, Litanie, Surinam,
July 12, 1939 (Geijskes).

I take great pleasure in naming this interesting Brachypremna
for the collector, Dr. D. C. Geijskes, who has added so materially
to our knowledge of the insects of Dutch Guiana. The fly is most
similar to species such as B. diversipes Alexander, differing in the
details of coloration of the body and legs, in the venation, and in
the somewhat inflated prsescutum. This latter character slightly
suggests the more accentuated condition to be found in the other-
wise entirely distinct B. quasimodo Alexander. The male hy-
popygium is different from all other species of the genus in which
this sex is known.

Mar., 1945]

Alexander: Cranflies

51

Genus Tanypremna Osten Sacken

Tanypremna (Tanypremna) elegantior new species.

Size large (wing 15 mm.) ; thorax uniformly orange-yellow, polished, with-
out pattern; head above orange-yellow with a major brownish black area on
the vertex; legs dark brown, the tarsi black; wings with the basal two-thirds
bright yellow, particularly the prearcular and costal fields, the distal third
more obscured, at the apex distinctly darkened; veins glabrous; Rs longer
than R2+s; cell 1st M2 large and subquadrate, nearly as wide as long; cell
2nd A wide; abdomen yellow, the tergites extensively infuscated medially.

Sex? — Wing 15 mm.; antenna about 1.3 mm. Abdomen broken beyond the
sixth segment, the total length probably about 19-20 mm.

Frontal prolongation of head yellow, very short, a little longer in the region
of the stout, broad nasus; opposite the lateral portions, the length much
shorter; dorsum of prolongation with abundant long black setae; mouthparts
black; palpi brownish black. Antennae short, 13-segmented; basal flagellar
segment oval, narrowed at base, the outer segments becoming long-cylindrical,
with long conspicuous verticils. Head above orange-yellow with a major
brownish black area on vertex adjoining the margin of eye, chiefly on the
posterior vertex, very narrowly interrupted at the midline, prolonged back-
ward on the orbits ; head beneath and on front paler yellow.

Thorax uniform orange-yellow, without markings, the surface glabrous,
polished. Halteres with stem yellow, knob infuscated. Legs with the coxae
and trochanters yellow; femora dark brown, narrowly yellow at base; tibiae
dark brown, the tips very narrowly blackened; tarsi very elongate, basitarsi
dark brown, the remainder of tarsi passing into black; claws simple. Wings
with basal two-thirds or to the general level of the cord bright yellow, par-
ticularly in the prearcular and costal fields; beyond the cord the yellow
becomes more obscured, at the wing-tip passing into dark brown, this includ-
ing the outer ends of cells R2 to M1} inclusive, these latter enclosing incon-
spicuous whitish central streaks ; stigma brownish yellow, narrowly margined
with brown; very narrow and little-evident brown seams along vein R2+ 3 and
over the anterior cord ; veins dark brown, yellow in the more intensely flavous
prearcular and costal fields. Veins unusually glabrous, including all veins
beyond cord; Rs with one or two trichia near base; the only veins with abun-
dant trichia are C, R, Rx, and Sc2 + Rx as far distad as the free tip of the latter.
Venation: Distance on C between Scx and the free tip of Sc2 a little longer
than that between Sc2 and R1+2’, Rs long, gently arcuated, exceeding R2+ 3, the
latter strongly arched at base; cell Mx long, widened subbasally; m about
one-half longer than the petiole of cell Mx; cell 1st M2 large and subquadrate,
the basal section of Mx+2 one-half longer than m; m-cu at near three-fifths
the length of M3+4:) subequal to the distal section of Cu a; cell 2nd A wide.

Abdomen broken at end of sixth segment ; tergites orange, brightest at base
and on outer segments ; dorsum with a broad, dark brown stripe, on outer seg-
ments more brightened on sides of basal rings, the latter delimited behind on
either side by more impressed areas ; sternites yellow.

52

Journal New York Entomological Society

[Vol. Lin

Holotype, Sex? Probably from Colombia, exact locality un-
known (Fred Walker No. 93) ; University of Michigan, through
Professor J. Speed Rogers. The exact data pertaining to this
Walker number is not available at this time.

The most similar described species are Tanypremna ( Tany -
premna ) carbonipes Alexander and T. ( T .) hodgei Alexander,
both of which differ conspicuously in all details of coloration of
the body and wings and in the venation. From the simple claws,
it might be assumed that the present specimen was a female ; how-
ever, the male sex in carbonipes has toothed claws whereas in
hodgei these are simple in both sexes, so nothing can be affirmed
on this basis.

Genus Limonia Meigen

Limonia (Rhipidia) ingenua new species.

Allied to multifida; mesonotal praescutum chiefly brownish yellow, with
three incomplete stripes behind ; posterior sclerites of notum and pleura dark
brown^ the anterior pleurites yellow; legs yellow, the outer tarsal segments
darkened; fore coxae yellow, the remaining pairs brownish black; wings
brownish yellow with a restricted brown pattern; Sc long, m^-cu before the
fork of M-; male hypopygium with the tergite strongly emarginate, the lobes
fringed with powerful flattened setae ; ventromesal lobe of basistyle and apical
lobes of aedeagus small; rostral prolongation of ventral dististyle with two
unequal spines, the outer one strong and slightly curved.

Male. — Length about 5 mm.; wing 5.7 mm.

Rostrum and palpi black. Antennae broken beyond the third segment;
scape black; pedicel and the simple first flagellar segment dark brown (from
its obvious relationship with multifida and sejugata, it may be assumed that
the male antennae will be found to be long-bipectinate). Head dark brown;
anterior vertex reduced to a linear strip that is about as wide as a single row
of ommatidia.

Pronotum dark brown, the scutellum and pretergites paler. Mesonotal
praescutum strongly arched but not pointed, the color chiefly brownish yellow,
clearer and somewhat whitish pruinose on sides; three medium brown stripes
are indicated on the posterior portion of the sclerite, the median one longer
and broader; posterior sclerites of notum dark brown to brownish black, the
parascutella paler ; sides of mediotergite and dorsal portions of pleurotergite
more reddish brown. Pleura chiefly yellow, the pteropleurite and metapleura
abruptly brownish black. Halteres yellow. Legs with the fore coxae and
trochanters yellow, middle and posterior coxae and trochanters brownish black ;
remainder of legs yellow, the terminal two tarsi segments infuscated; claws
with a single erect basal spine. Wings brownish yellow, the prearcular and
costal portions slightly clearer yellow; a very restricted brown pattern that
consists of small spots that are limited to the vicinity of the veins, as follows :

Mar., 1945]

Alexander: Cranflies

53

Origin of Bs ; fork of Sc; cord and outer end of cell 1st M2 ; small darken,-
ings at ends of all longitudinal veins, largest over the tip of 1st A; stigma
bicolored, chiefly yellow, the outer end, over B2 and free tip of Sc2, darker
brown, conspicuous ; proximal end of stigma and a subapical washing in outer
radial field paler brown; veins yellow, infuscated in the clouded portions.
Venation: Sc relatively long, Scx ending about opposite two-thirds the length
of Bs, Sc2 at its tip; Bs nearly perpendicular at origin but not angulatfed;
m-cu about one-third its length before fork of M ; cell 1st M2 a trifle shorter
than vein M1+2 beyond it.

Abdomen bicolored, obscure yellow, the caudal borders of the segments
narrowly infuscated, more extensively so on the more proximal segments; a
darkened pleural spot on the intermediate segments; hypopygium yellow.
Male hypopygium with the caudal margin of tergite with a deep V-shaped
notch, the conspicuous lateral lobes fringed with setae, smallest near the mid-
line, becoming very large and flattened at apex of the narrowly rounded lobes ;
remainder of tergite virtually glabrous. Basistyle with the ventromesal lobe
low and rounded, with about three setae on the lower or cephalic portion very
long and conspicuous, the others short and normal; face of style at base of
lobe with a pair of long setae. Dorsal dististyle a strong, powerful sickle,
its tip acute and slightly upcurved. Ventral dististyle fleshy, the rostral pro-
longation slender, with two very unequal spines ; outer spine placed more than
its own length before apex of prolongation, short and powerful; second spine
slightly more basal, long and straight, about one-third longer than the outer.
Gonapophyses with mesal-apical lobe slender, blackened. ^Edeagus with
apical lobes small and inconspicuous.

Holotype, J*, Pedregoso, Costa Rica, altitude 2,500 feet, Jan-
uary 1939 (Dean Rounds).

The most nearly allied species are the Mexican Limonia ( Rhi -
pidia) multi fida Alexander and L. (R.) sejugata Alexander.
The former has a distinctive coloration of the body and wings,
and a very different structure of the hypopygium, including the
tergite, basistyles and aedeagus; the latter is much more similar
in its general appearance, including the wings and legs, but dif-
fers in the structure of the male hypopygium, involving the ter-
gite, basistyles, ventral dististyles, gonapophyses and aedeagus,
becoming most evident in the single rostral spine of the ventral
dististyle and in the quite different gonapophyses.

Limonia (Peripheroptera) cochabambse new species.

Size large (wing, male, over 11 mm.) ; mesonotum yellow pollinose, with
three conspicuous brownish black stripes ; knobs of halteres blackened ; femora
obscure yellow, the tips brownish black; wings pale yellow, with a restricted
brown pattern; cord in transverse alignment; free tip of Sc2 lying far before
B2; cell 1st M2 subequal in length to vein M1+2 beyond it.

Male. — Length about 8 mm.; wing 11.5 mm.

54

Journal New York Entomological Society

[Vol. LIII

Rostrum brownish black ; palpi black. Antennae black throughout ; flagellar
segments suboval, the outer ones becoming more slender and cylindrical; ver-
ticles conspicuous. Head yellowish gray, somewhat clearer gray on front; a
conspicuous brown area on vertex, becoming a little wider on the posterior
vertex.

Pronotum dark brown above, broadly obscure yellow on sides. Mesonotal
praescutum with the ground yellow pollinose, with three conspicuous brownish
black stripes that are more or less obscured by a faint pollen, the median
stripe unusually broad and conspicuous ; scutum broadly yellow pollinose
medially, the lobes more brownish yellow, darkest on their antero-lateral por-
tions ; scutellum and postnotum brownish yellow, more or less yellow pollinose.
Pleura brownish yellow, heavily yellow pollinose; cephalic portion of anepi-
sternum and ventral sternopleurite more infuscated. Halteres with stem
yellow, knob conspicuously blackened. Legs with the coxae castaneous; tro-
chanters more yellowed, rimmed with black at apices ; femora obscure yellow,
the tips rather narrowly brownish black, on the posterior legs involving about
the outer tenth, subequal in amount on all legs; tibiae reddish brown; tarsi
passing into black; claws with a single long tooth. Wings pale yellow, the
prearcular field, costal border and outer radial field more saturated yellow ;
stigmal region a trifle more infuscated than the remainder of costal border ;
a very restricted dark brown pattern, including narrow seams over arculus,
Sc2, cord, outer end of cell 1st M2, free tip of Sc2 and R2; veins brown. Vena-
tion: Prearcular region relatively extensive; first section of vein R a little
longer than the second section ( Sc2 + Rx ) ; free tip of Sc2 far before R2, vein R±
more than twice R2; cord transverse; basal section of R4+ 5 straight; cell 1st M2
long, gently widened outwardly, subequal to vein M1+2 beyond it ; m-cu oblique
and weakly sinuous, about one-third its length beyond the fork of M ; cell
2nd A narrow, the greatest width about twice that of the constricted basal
portion.

Abdomen dark brown, more or less yellow pollinose, especially on the
sternites; hypopygium brownish black. Male hypopygium with the ventral
dististyle large and fleshy, its area about three times that of the basistyle;
rostral prolongation relatively small; spines placed close together, straight,
subequal to the length of the prolongation beyond their bases. Gonapophyses
with mesal-apical lobe slender and gently curved.

Holotype, J1, Chapare, Cochabamba, Bolivia, November 1934
•(Franz Steinbach).

The most similar described species is Limonm ( Peripheroptera )
trimelcenia Alexander, of Peru, which differs especially in the
venation and pattern of the wings.

Genus Helius St. Fargeau

Helius (Helius) schildi new species.

Belongs to the mirabilis group, allied to miranda ; mesonotum in front
broadly yellow, praescutum blackened sublaterally, this color crossing the

Mar., 1945]

Alexander: Cranflies

55

suture onto the outer portion of the scutal lobes, the disk of praescutum and
scutum obscure brownish yellow; pleura yellow; knobs of halteres orange-
yellow; wings yellowish subhyaline, the prearcular and costal portions light
yellow; two conspicuous brown crossbands, one postarcular, the second at
cord; cell 1st M2 rectangular, m-cu only a short distance beyond fork of M ;
abdomen yellow, the posterior portions of the segments brownish black.

Female. — Length about 8 mm. ; wing 6 mm.

Rostrum a little exceeding the remainder of head, brown; palpi more
brownish black. Antennae of moderate length; scape and pedicel brown,
flagellum yellow, especially the outer segments; flagellar segments passing
through oval to elongate-cylindrical, provided with unusually long verticils,
especially on the outer segments. Head dark brown, gray pruinose, especially
on front and narrow anterior vertex.

Pronotum and the broad cephalic and lateral portions of praescutum yellow,
the latter obscure brownish yellow on posterior half, blackened sublaterally,
this color crossing the suture onto the lateral portions of the scutal lobes;
central portion of scutum obscure brownish yellow; scutellum dark brown:
postnotum, including pleurotergite, blackened, the latter restrictedly pruinose
on its more dorsal portion. Pleura yellow, sparsely pruinose ventrally. Hal-
teres orange-yellow, especially the knobs. Legs with the coxae obscure yellow ;
remainder of legs broken, excepting the basal fourth of a single femur which
is clear yellow. Wings yellowish subhyaline, conspicuously crossbanded with
brown; prearcular field and costal border conspicuously light yellow; basal
dark band post-arcular, slightly more widened at vein Cu; outer band at cord,
completely traversing the wing from stigma to the posterior margin at CuL,
narrowest at and above the fork of M ; cell 1st M2 chiefly pale; veins yellow,
darker in the banded portions. Venation: Branches of Es on their basal half
generally parallel to one another ; basal section of Ei+5 in longitudinal align-
ment with Es, about twice r-m; m-cu only a short distance beyond the fork
of M, this distance not exceeding r-m ; cell 1st M2 rectangular.

Abdominal tergites conspicuously banded with yellow and brownish black,
the bases of the segments yellow, the remainder darkened; on the outer seg-
ments the pale color slightly exceeds the dark; second tergite with an addi-
tional darkened basal ring; intermediate sternites more uniformly pale, the
outer segments banded as are the tergites ; valves of ovipositor very long and
slender, yellowish horn color.

Holotype, §, Higuito, San Mateo, Costa Rica (Pablo Schild) ;
United States National Museum.

I am pleased to name this interesting species for the collector,
Mr. Pablo Schild (Paul Schild), who added materially to our
knowledge of Costa Rican insects. The fly is most similar to
Helius ( Helius ) miranda (Alexander), of Amazonian Brazil and
Pern, differing in the pattern of the body and wings, especially
the broad and conspicuous basal wing band and the handsomely
patterned mesonotum.

56

Journal New York Entomological Society

[Vol. Lin

Helius (Helius) phasmatis new species.

Belongs to the albitarsis group ; thorax yellow, the central portion of
scutum and mediotergite, together with the scutellum, a very little infus-
cated; lialteres uniformly pale yellow; legs pale yellow, the tarsi extensively
snowy white ; wings subhyaline, stigma very long and narrow, brown ; r-m
obliterated by fusion of veins Bi+5 and M1+2’, abdominal tergites brown,
hypopygium obscure yellow.

Male. — Length about 7 mm. ; wing 7.5 mm.

Rostrum obscure brownish yellow, about as long as the remainder of head.
Antennae relatively short, subequal in length to rostrum, yellow throughout ;
flagellar segments passing through suboval to elongate-cylindrical, the longest
verticils considerably exceeding the segments in length. Head dark brown,
sparsely pruinose; eyes large, the anterior vertex reduced to a capillary line
that is scarcely wider than a single row of ommatidia.

Cervical region, pronotum and mesonotum yellow, the scutellum, central
region of scutum and the mediotergite a very little infuscated; pleura and
broad lateral borders of the praescutum and scutal lobes still paler yellow.
Halteres uniformly pale yellow. Legs with the coxae and trochanters pale
yellow; femora, tibiae and most of basitarsi pale yellow, the apex of the last
and remainder of tarsi snowy white. Wings subhyaline, cells C and Sc a trifle
darker; stigma very long and narrow, still darker brown; veins brown. Vena-
tion: Crossvein r-m obliterated by the short fusion of Bi+5 on M1+2, the fusion
shorter than the basal section of Bi+5-, cell 1st M2 large, irregularly hexagonal,
the longest faces being the caudal, proximal and cephalic; m about one-third
the basal section of M3; all veins beyond 1st M2 much exceeding the cell; cell
2nd A relatively narrow.

Abdominal tergites brown, the sternites a little paler ; hypopygium abruptly
obscure yellow.

Holotype, J*, Santo Domingo, without exact geographical data,
March 1928 (P. A. Ricart) ; United States National Museum.

Helius ( Helius ) phasmatis differs conspicuously from all other
described species of the albitarsis group in the yellow coloration
of the thorax, as well as in the yellow legs and halteres.

Genus Oxydiscus de Meijere

Oxydiscus (Oxydiscus) destitutus new species.

General coloration of thoracic notum chestnut brown, the praescutum with a
darker brown median stripe; wings with a pale brown suffusion; macrotrichia
of cells restricted in number; Bs strongly arcuated; B2+3+4: elevated, in direct
longitudinal alignment with vein B3, the fork of cell B3 asymmetrical ; vein B2
close to fork of B2+3+i ; male hypopygium with the ventral spines of the
aedeagus apparently lacking.

Male. — Length about 3.6 mm. ; wing 4.4 mm.

Rostrum chestnut brown; palpi broken. Antennae broken. Head dark
brown; anterior vertex moderately wide, approximately equal to the trans-
verse diameter of eye as viewed from above.

Mar., 1945]

Alexander: Cranflies

57

Pronotum dark brown. Mesonotal praescutum chiefly chestnut brown, the
median region darkened to produce a poorly delimited median stripe; pos-
terior sclerites of notum obscure yellow, the scutal lobes weakly darkened;
pleurotergite yellow. Pleura brownish yellow. Halteres pale, the knobs
weakly darkened. Legs with the coxae and trochanters testaceous yellow;
femora brown, the tibiae and tarsi a little paler brown; tibial spurs con-
spicuous; claws elongate, simple. Wings not conspicuously widened, with a
pale brown suffusion ; prearcular and costal regions somewhat more yellowed ;
stigma pale brown and very inconspicuous, lying entirely beyond the level of
vein E2; veins brown. Macrotrichia of cells restricted to sparse series in
central portions of outer ends of cells E3 to 2nd M2, inclusive. Venation: As
compared with costaricensis ; Scx subequal to vein Ex between the origin of Es
and Sc2; Es approximately of the same length but much more arcuated;
J?2+3+4 more elevated, in direct alignment with Ez so the fork of cell Ea is asym-
metrical; ^2+3+4 more than one half Es; vein E2 close to fork of E2+3+i; cell
1st Mo narrower; m-cu about one-half its length beyond the fork of M ; vein
2nd A ending shortly beyond the level of origin of Es, bent rather suddenly
into the margin. In costaricensis, Sc1 is only a little more than one-half Ex;
Es is only slightly arcuated so cell Ex is correspondingly narrow ; E2 about its
own length beyond fork of E2+3+4- fork of cell E3 symmetrical; m-cu nearly
its own length beyond fork of M.

Abdomen almost uniformly dark brown, the incisures a trifle paler. Male
hypopygium with the dististyles about as in the other regional species of the
genus. The paired spines of the ventral plate of the aedeagus, found in the
other species, are here apparently lacking ; in acutissimus and mexicanus, these
appear as elongate spines.

Holotype, J1, Pedregoso, Costa Rica, altitude 2,075 feet, Jan-
uary 1939 (Dean Rounds).

The present fly is very distinct from the only other described
Costa Rican species, Oxy discus (Oxy discus) costaricensis (Alex-
ander), differing particularly in the venational features above
indicated. The details of structure of the male hypopygium in
costaricensis are not known to me. From 0. (0.) acutissimus
Alexander and 0. (0.) mexicanus Alexander, of southern Mexico,
the present fly differs most evidently in the structure of the male
hypopygium, particularly of the ventral plate of the aedeagus.
It is now evident that there are rather numerous species of the
genus in mountainous Middle and northwestern South America.

Genus N eognophomyia Alexander

Neognophomyia heliconise new species.

General coloration yellow, the anepisternum and pleurotergite each with a
conspicuous black spot; wings subhyaline, the prearcular and costal fields a

58

Journal New York Entomological Society

[Vol. LIII

trifle more yellowed ; a relatively narrow but conspicuous brown band at cord ;
abdomen yellow, tergites two and three on sides, five and six almost entirely
blackened ; male hypopygium with tergal spines yellow, very long and slender,
especially their hairlike tips ; both dististyles small and simple, the inner one
narrowly blackened; phallosome a depressed plate, its apex broadly obtuse,
with a small median blackened knob or point.

Male. — Length about 4.5-5 mm.; wing 4.8-5. 2 mm.

Female. — Length about 4.8-5 mm.; wing 5-5.2 mm.

Rostrum yellow; palpi with two basal segments yellow, the outer ones in-
fuscated. Antennae with scape and pedicel yellow, flagellum black; flagellar
segments long-oval, the verticils conspicuous. Head yellow; anterior vertex
narrow, especially so in male where it is about equal to the diameter of the
scape, in female a little wider; posterior vertex with a Y-shaped or triangular
depression between its posterior portion and the anterior vertex.

Pronotum light yellow. Mesonotum yellow, the surface shiny, the scutal
lobes somewhat deeper in color ; pleurotergite with a very large and extensive
blackened area. Pleura yellow, with a second circular black spot that occupies
most of the anepisternum. Halteres yellow, tips of knobs weakly darkened.
Legs yellow; tips of basitarsi and remainder of tarsi brownish black; in
some specimens, the fore tarsi more extensively darkened. Wings subhyaline,
the prearcular and costal fields a trifle more yellowed; a relatively narrow
but conspicuous brown band at cord, widest at costa, narrowed behind, ending
at Cu; outer end of cell 1st M2 very narrowly seamed with brown; veins
brownish yellow, more flavous in the brightened fields, dark brown in the
patterned areas. Venation: E2+ 3+4 varying from about two-thirds to fully
as long as R3+4; vein Es oblique, E± very long, terminating at wing-tip; cell
1st M2 short, widened outwardly ; m-cu from one-fourth to one-half its length
beyond fork of M ; cell 2nd A relatively narrow.

Abdomen yellow, tergites two and three with large and conspicuous black
lateral areas; tergites five and six, in most specimens, entirely brownish
black; in the males with the central portion a trifle paler, forming more or
less distinct yellow triangles; sternites, remainder of tergites and the hypo-
pygium of male yellow; in female, the genital segment weakly infuscated.
Male hypopygium with the tergal spines yellow, very long and slender, the
basal half of each appearing as a narrow blade, the distal half narrowed into
an exceedingly slender hairlike point, somewhat as in productissima. Both
dististyles small and simple, the outer style with about four elongate setae,
one apical in position; inner style relatively narrow, especially on its distal
half, the apex subacute, narrowly blackened, the outer surface of basal por-
tion with several long powerful setae. Phallosome a broad, depressed plate,
its apex produced into a small black median knob, somewhat as in monophora,
but with the lateral shoulders much more conspicuous, the general apex of
the plate being broadly obtuse.

Holotype, J1, Barro Colorado, Canal Zone, April 1940, ex flow-
ers of Heliconia marice (Musaceae) ; (James Zetek, No. 4645) ;

Mar., 1945]

Alexander: Cranflies

59

United States National Museum. Allotopotype, §. Paratopo-
types, 3 10

In the structure of the male hypopygium, the present species is
quite distinct from other related members of the genus. In the
pale, exceedingly produced tergal spines, it is most similar to the
otherwise distinct Neognophomyia productissima Alexander,
while in the nature of the phallosomic plate it somewhat suggests
species such as N. monophora Alexander, but all details of this
plate, as well as other structures of the hypopygium, are quite
distinct in the two flies.

Genus Gnophomyia Osten Sacken

G-nophomyia (Gnophomyia) apicularis new species.

Allied to tuber; general coloration of mesonotum brown, more or less
pruinose, the praescutum with indications of a slightly darker brown stripe;
femora obscure yellow, with a very vague, light brown subterminal ring;
wings light brown, stigma scarcely evident; Bs in alignment with JR5; male
hypopygium with the mesal face of bastistyle produced into a setiferous
tubercle; outer dististyle a relatively short, blackened, paddle-like blade;
inner dististyle blackened, its lower apical angle produced into a slender
spine ; phallosome at apex produced into a diamond-shaped median point.

Male. — Length about 6.5-7 mm. ; wing 6-6.5 x 1.7-1. 8 mm. ; antenna about
1. 8-1.9 mm.

Eostrum and palpi dark brown. Antennae with scape and pedicel brown-
ish yellow to brown, flagellum brown, the first segment narrowly pale at base ;
segments subcylindrical, the outer ones shorter; terminal segment long-oval,
shorter than the penultimate. Head brownish gray, light gray on the orbits ;
eyes large; anterior vertex relatively narrow, about three times the diameter
of the scape.

Pronotum obscure brownish yellow, more infuscated on sides; pretergites
pale yellow. Mesonotum brown, more or less pruinose, the praescutum with
indications of a slightly darker brown median stripe, the laterals undiffer-
entiated ; humeral and lateral regions of praescutum more yellowed ;
pseudosutural foveae reddish brown; posterior border of scutellum narrowly
reddened, more conspicuous on sides ; lateral border of anterior half of
mediotergite and much of dorsal portion of pleurotergite deep obscure yel-
low. Pleura infuscated dorsally, more reddened on the sternopleurite and
meron, the surface distinctly pruinose; metapleura light yellow. Halteres
short, obscure yellow, the base of knob very weakly darkened. Legs with
the coxae and trochanters yellow, the posterior coxae somewhat paler; femora
obscure yellow, with very vague indications of a narrow, light brown, sub-
terminal ring; tibiae and basitarsi obscure yellow, second and third tarsal
segments obscure yellow, tipped with darker; outer segments more uniformly
brownish black; claws simple. Wings moderately wide, tinged with very

60

Journal New York Entomological Society

[Vol. LIU

light brown, the prearcular and costal fields light yellow; stigma reduced to
a scarcely evident seam along vein B1+2, not reaching vein B3 behind; veins
brown, more brownish yellow in the basal portions. Venation: Sc± ending
jnst before level of B2, Sc2 slightly variable in position, from just beyond
level of fork of Bs to about opposite one-third the length of R2+3+4; r-m at
or just beyond fork of Bs, the latter in alignment with B5 ; cell 1st M2 sub-
equal to vein Mi beyond it; m<-cu about its own length beyond fork of M
or nearly opposite two-fifths the length of cell 1st M2; cell 2nd A broad.

Abdominal tergites dark brown, the caudal borders of the sternites more
testaceous; hypopygium brown. Male hypopygium with the basistyle short
and stout, on mesal face near base with a small setiferous tubercle, somewhat
as in tuber. Outer dististyle a relatively short, blackened, paddlelike blade,
its apex obtuse. Inner dististyle less than one-half the length of the outer,
blackened, the lower apical angle produced into a slender beaklike spine, the
apex beyond this point strongly produced. Phallosome with apex emarginate,
further produced into a conspicuous blackened structure that is more or less
diamond-shaped, the apex truncated.

Holotype, .J', Pedrogoso, Costa Rica, altitude 2,100 feet, Janu-
ary 1939 (Dean Rounds). Paratopotype,

The most similar described species is Gnophomyia {Gnopho-
myia) tuber Alexander, of Ecuador, which is approximately alike
in its general appearance, differing conspicuously in the structure
of the male hypopygium, particularly the outer dististyle, inner
dististyle and the phallosome.

Gnophomyia (Gnophomyia) subapicularis new species.

Allied to tuber ; general coloration of mesonotum blackened, more or less
pruinose, especially on the posterior sclerites; pleura variegated with black,
reddish brown and yellow; halteres brownish black; posterior legs brownish
yellow, paler than the fore pair; wings relatively narrow, with a weak dusky
tinge; Scx ending about opposite two-thirds the length of J22+3+4; abdomen
chiefly blackened, hypopygium qnd ovipositor paler; male hypopygium with
a setiferous tubercle on cephalic end of mesal face of basistyle; outer dis-
tistyle relatively slender; inner dististyle with lower apical angle produced
into a chitinized point.

Male. — Length about 5 mm. ; wing 5. 3-5.5 X 1.3-1.4 mm.

Female. — Length about 5.5 mm. ; wing 6 mm.

Rostrum light brown ; palpi darker brown. Antennae brownish black
throughout ; flagellar segments elongate, subcylindrical, shortening very
gradually to the terminal ones; verticils subequal in length to the segments.
Head blackened, heavily gray pruinose ; eyes large and protuberant ; anterior
vertex broad, nearly five times the diameter of scape.

Pronotum yellow medially, dark brown on sides, pretergites even clearer
yellow. Mesonotum almost uniformly black, the humeral region of praescu-

Mar., 1945]

Alexander: Cranflies

61

turn slightly more reddened; pseudosutural fovese black; scutal lobes and
lateral portions of praescutum more polished black; median region of scutum,
scutellum and postnotum similarly blackened but more pruinose. Pleura
with the dorsal sternopleurite and pteropleurite reddish brown, the propleura
and anepisterum extensively blackened ; ventral sternopleurite darkened ;
metapleura paling to light yellow; surface of pleura more or less heavily
pruinose. Halteres brownish black, base of stem restrictedly brightened.
Legs with the coxae yellow, the fore pair more darkened basally; trochanters
yellow; fore legs with femora, tibiae and basitarsi brownish black, on the
other legs paler brown or, on posterior legs, brownish yellow; outer tarsal
segments blackened; claws simple. Wings relatively narrow, with a weak
dusky tinge; stigma elongate, darker brown; veins brownish black. Vena-
tion: S c1 ending about opposite two-thirds the length of E2+3+4, Sc2 nearly
opposite fork of Es; r-^m variable in position but approximately at the fork
of Es; m-cu usually less than its own length beyond the fork of M.

Abdominal tergites black or piceous; sternites usually obscure yellow,
darkened laterally; hypopygium obscure yellow to light brown. Ovipositor
and genital shield brownish yellow, the tips of the cerci narrowly darkened.
Male hypopygium with the basistyle stout, the mesal face near cephalic end
produced into a conspicuous setif erous lobe, approximately as in apicularis
and tuber. Outer dististyle longer and more slender than in apicularis, the
apex narrow. Inner dististyle darkened, the lower apical angle produced
into a toothlike point; apex of style beyond this truncated, not strongly
produced as in apicularis. Phallosome narrowed outwardly, the blackened
apex produced into four points, two of which are more elongate blackened
rods, the remaining pair representing the low outer lateral angles of a ven-
tral median plate.

Holotype, <$, Pedregoso, Costa Rica, altitude 2,100 feet, January
1939 (Dean Rounds). Allotopotype, <j>. Paratopotypes, 7 $ §>
altitude 2,075-2,100 feet, January 1939.

Gnophomyia ( Gnophomyia ) sub apicularis is most similar
to G. ( G .) apicularis new species and G. ( G .) tuber Alexander,
differing especially in the structure of the male hypopygium,
particularly of both dististyles and the phallosome. The tubercle
on the basistyle and the tooth at apex of the inner dististyle pro-
vide noteworthy characters among the host of allied and gener-
ally similar forms.

62 Journal New York Entomological Society [Vol. Lin

SOME COLLECTIONS OF LEPIDOPTERA

It is often of interest and sometimes of importance to ascertain
where a certain collection of Lepidoptera is located. English col-
lectors are fortunate in having a book entitled ‘ ‘ Where is the

collection?” by Charles Davies Sherborn, which gives such infor-
mation, but there is no similar work on American collections.
Even a search of the literature often does not produce results.
For that reason I am publishing this brief note respecting several
collections acquired by me.

Part of the collection made by E. H. Blackmore of Victoria,
British Columbia, was purchased during 1935; the collection,
library and correspondence of Thomas E. Bean, were purchased
in April, 1936. This collection was made, in part, at Galena, Illi-
nois, but mostly at Laggan, Alberta. Bean was a correspondent
of William Henry Edwards and supplied him with much of the
data contained in the third volume of “The Butterflies of North
America.” The collection of Max Rothke of Scranton, Pennsyl-
vania, was purchased in November, 1936 ; that of R. F. Sternitzky
of San Francisco, California, in February, 1938; the collection
of Owen Bryant, including his Arctic material, in January, 1941,
and I also acquired the small local collection of Louis Doerfel of
Newark, New Jersey.

Some of these collections are of considerable historic interest to
lepidopterists and contained a number of types and paratypes.
The types were given to The American Museum of Natural His-
tory. Most of the paratypes remain in my collection. — Cyril F.
dos Passos.

Mar., 1945]

Wang: Chilopoda

63

A PRELIMINARY REPORT ON CHILOPODA
AT ISHAN, KWANGSI AND MEITAN,
KWEICHOW

Yuhsi Moltze Wang

Zoology Laboratory; Biology Department,

National University of Chekiang

The Chilopoda which are described in the present report belong
to the orders Geophilomorpha, Scolopendromorpha, Lithebie-
morpha and Scntigeromorpha and consist of 6 families, 6 genera
and 7 species. They should all be included in the paleoarctic
forms, the oriental forms and subtropical forms. The species and
their characteristics and locations may be stated as follows :

Order GEOPHILOMORPHA
Family Geophilida]

Body long, worm-shaped. Eyes absent. Basal shield as long
as wide. Ventral pores and coxal pores present.

Genus Geophilus Leach
Geophilus duponti Attem

Two specimens examined, measurements in mm.

No. 20 No. 21

Body length .....; 56 58

Body width 2 2

Basal shield length 1 1

Basal shield width 1 1

Neck length 1.5 1.5

Antenna 2 2

Anal leg 2 2

Ocellus absent ; antenna short, 14 joints ; basal shield as long as
wide; terga a longitudinal band; walking legs 73 pairs; sternum
squarius, haired; ventral pores present, crescent shaped at pos-
terior part of each sternum ; the 24th to 36th segment, 2 grooves
present at the anterior edge of each sternum; anal leg short, 20
coxal pores on each coxa.

64

Journal New York Entomological Society

[Vol. LIII

Color pale brown; it occurs at Kwangsi and Kweichow. I
found it in Meitan under soil.

Family Mecistocephalid,®

Basal shield longer than wide ; ventral pores absent.

Genus Mecistocephalus Newport
Mecistocephalus mecistocephalus Newport

One specimen only. Body length 70 mm. Body width 2 mm.,
length of basal shield three times longer than width ; neck 1 mm. ;
antennae 7 mm. ; anal leg 4 mm.

Ocellus absent; antenna 14 joints; labium 2 dentitions; body,
walking legs and antenna haired ; walking legs 65 pairs ; no ven-
tral pores ; coxal present, more than 68 on each anal leg.

Color yellowish brown, head chestnut red. It occurs at
Kwangsi, and feeds upon earthworms.

Order SCOLOPENDROMORPHA
(1) Family Scolopendrid^e

Ocellus present, by the genus Mimops with a white eye spot.
Tarsus always two joints. Sternum with lateral longitudinal fur-
rows or without furrow, rarely with one median furrow, none
with transverse furrow ; each half of the intercalary sternum two
parted, the intercalary terga, for the most part, not greatly de-
veloped, often not visible.

Genus Scolopendra
Scolopendra subspinipes Leach

Three specimens, measurements in mm.

No. 1

No. 2

No. 3

Body length

143

120

114

Width

11

10

9.5

Head length

8

7.5

7

Width

8.5

8

6

Neck width

10.5

10

10

Antennae

28

22

Damaged

Uropods

26

25

21

Sex

Female

Male

Female

Locality

Ishan

Ishan

Meitan

Mar., 1945]

Wang: Chilopoda

65

Head brownish yellow, terga dark green, sterna brownish yel-
low, ocellus 4 on each side of head; labium 5 dentate plates;
antenna 18 joints; terga 21 in number, the twelfth, the broadest;
stigmata, 9 pairs on each pleuron of the third, fifth, eighth, tenth,
twelfth, fourteenth, sixteenth, eighteenth, and twentieth segment.
Walking legs 21 pairs, uropods the longest, anal segment with two
strong spines. The prefemur of uropod with 3 spines on interior
upper edge, 2 spines on exterior lower edge; and 2 spines on in-
terior lower edge; no spine on tarsus. It is found at Kwangsi
and Kweichow, more abundant at Kwangsi. I have found it at
Kiangsu, Chekiang and Jukien.

Scolopendra japonica L. Koch

Two specimens, male, measurements in mm.

No. 4 No. 5

Body length 71 65

Body width 7 6

Head length 5 5

Head width 5.5 5

Neck width 6.5 6

Antenna 14 17

Uropods 16 16

Locality Ishan Meitan

Head and terga reddish brown, terga bordered with black,
sterna brownish yellow. It differs from subspinipes in two char-
acteristics: (1) With 4 clentates on labium; (2) 3 spines on
exterior lower edge of the prefemur of uropods and one spine
on the tarsus of uropods.

Family Cryptopine

Eyes absent. Tarsus of the lst-19th pairs of legs one joint,
only in Trigonocryptes two joints. Sterna with median furrow,
rarely absent, often with transverse furrow, none with 2 longi-
tudinal furrows. Intercalary sterna half, simple, intercalary
terga most strongly developed.

Genus Otocryptops Hasse
Otocryptops rubiginosus (L. Koch)

Four specimens, measurements in mm.

66

Journal New York Entomological Society

[Vol. LIII

No. 6 ' No. 7 No. 8 No. 9

Body length 49 36 31 26

Body width . 4.5 4 4 3

Head length 4 3 3 2.5

Head width 4 3 3 2.5

Neck width 4 3 3 2

Antenna 9 8 8 7

Uropods •. Damaged 9 Damaged 7

Location Ishan Ishan Ishan Meitan

Head without median furrow, antenna 17 joints, haired ; labium
without dentates. Terga 23 in number, stigma 10 pairs, on each
pleuron of the 3rd, 5th, 8th, 10th, 12th, 14th, 16th, 18th, 20th and
22nd segment. Walking legs 23 pairs, uropods with 1 spine each
on upper and lower surface of prefemur. Color orange brown
to reddish brown, head paler. It occurred at Kwangsi and
Kweichow, more abundant at Ishan.

Order LITHOBIOMORPHA

Only one family, Lithobudae, and one genus, Monotarsobius
Verhoeff, has been collected by the laboratory.

Monotarsobius holstu Pocock

Four specimens, measurements in mm.

Body length

No. 10
17

No. 11
16

No. 12
15

No. 13
8

Body width

2.5

2

2.5

1.2

Head length

2

2

3

1

Head width

’ 2.5

2

3.2

1.5

Neck width

2.2

2

3

1

Antenna

8

7

9

3.2

Uropods

Damaged

8

7

3

Locality

Meitan

Color dark brown. Head with 3 ocelli on each side ; labium
with 2 dentates, the base of labium furrowed, antenna 21 joints,
haired ; terga 15 in number, 8 larger, the 5th segment the broad-
est ; stigma 6 pairs, on each pleuron of the 3rd, 5th, 8th, 10th, 12th
and 14th segment. Walking legs 15 pairs, spinous; uropods rela-
tively long, with 0, 1, 3, 1, 0 spines. It feeds on insects and is
found under bark and stone. The younger has only 7 segments.

Mar., 1945]

Wang: Chilopoda

67

It is more abundant in Meitan. I have found it at Kwangsi and
Chekiang.

Order SCUTIGEROMORPHA
Family Scutigerid^e
Genus Thereuonema Verhoeff
Thereuonema tuberculata (Wood)

Six specimens, measurements in mm.

No. 14 No. 15 No. 16 No. 17 No. 18 No. 19

Body length 15 20 17 11 8 4

Body width 3 3 2.5 2 1.5 0.7

Head length 2 2.5 2 1.5 1.2 0.7

Head width 2.5 3.2 2.5 2 1.8 0.8

Neck width MHL. 1.5 , 2 2 1 0.9 0.5

Antenna -.. ,§§ 20 24 22 18 14 5

Uropods :... 27 30 29 20 17 7

Locality Meitan

Ocelli numerous; labium with 4 long dentates on each parti-
tion; antenna long, 4 parted, more than 400 joints, 11 terga vis-
ible, eight larger, the 5th tergum, the longest. Stigma 7 in single
number on posterior border of each larger tergum, walking legs
15 pairs, each with 8 blue rings, tarsus 47 joints; uropods very
long, spine 1, 2, 2, 3, on humerus, prefemur, femur and tibia.

Color dark brown with 3 blue black longitudinal bands, legs
paler.

It is active at night, may be found on the walls and in corners
or under stones and soil of human habitations, and is more
abundant at Kweichow and Kwangsi. I have found it occurring
at Chekiang and Kiangsu.

68

Journal New York Entomological Society

[Vol. LIU

NOTES ON THE POTATO TUBER MOTH—
Gnorimoschema (Phthorimaea) operculella
(Zell) IN NEW JERSEY

During the inspection for the tuber moth in New Jersey the
New Jersey Department of Agriculture learned of a dump of
condemned, southern potatoes on a farm in Burlington County.
Clean up of this dump was postponed in the hope that our north-
ern winter would destroy the infestation by this southern insect.

An inspection was made November 10, 1943, and many moths
were on the wing. A few larva? were present. Many pupae and
prepupae were found. No infested tubers found. Three nights
of freezing to this date. A week later moths were still flying;
many pupae and prepupae; no larvae. Seven nights of freezing
to this date.

No moths were found on February 10, 1944. Many pupae in
various stages of development but the prepupae were dead. Pupae
gathered and caged started emergence four days later. Seventy-
seven days of freezing weather to date.

Healthy pupae were collected March 31, 1944 and started to
emerge three days later. Many pupae subjected to prolonged
moisture had died and were covered with molds. The major
portion of live pupae was found in the ears and folds of the bag-
closures. Total days of freezing weather to date, 112. A start
was made in burning over the dump with Hauck weed burners.

Burning was half finished April 13 and was completed April
19, 1944. Bains and the wet condition of the pile prevented
finishing the burning sooner and in the interval moths had
emerged. The farmer had planted potatoes in the field adjoin-
ing this potato pile and it was necessary to bury and fumigate
the potatoes in the dump. These notes show that the potato
tuber moth can survive our winter, under certain conditions.

No attempt was made to correlate moisture and temperature
during the period of observation. It is assumed that moisture
was at an optimum in the lower layers of the pile, with variation
only in the upper layer. The death of the pupae under the upper
strata shows that excessive moisture destroys the insect.

The progressive death of larvae and prepupae and the disap-
pearance of the moth leads one to assume that survival was only
possible in the pupal stage. — William M. Boyd.

Mar., 1945]

Proceedings of the Society

69

PROCEEDINGS OF THE NEW YORK
ENTOMOLOGICAL SOCIETY

Meeting of January 5, 1943

The Annual Meeting of the New York Entomological Society was held on
January 5, 1943, in the American Museum of Natural History, with Presi-
dent Weiss in the chair; thirteen members and nine visitors present.

The treasurer’s report for 1942 was read by Dr. Spieth. Dr. Gertsch
reported that the Auditing Committee had found the treasurer’s books in
good order. Both these reports were accepted.

The Nominating Committee ’s recommendations for the officers for 1943
were read by Mr. Soraci as follows:

President Wm. P. Comstock

Vice-President .....: : Edwin Way Teale

Secretary !... Annette L. Bacon

Treasurer : Willis J. Gertsch

Librarian and Curator Leonard J. Sanford

Executive Committee ,.,;......S. W. Bromley

Wm. T. Davis
Albert Hartzell
A. B. Klots
F. E. Lutz

Publication Committee ....Ernest L. Bell

Herbert Buckes
Edwin W. Teale
Harry B. Weiss

Delegate to the New York
Academy of Sciences Wm. T. Davis

There being no further nominations from the floor, a motion was adopted
that the secretary cast one ballot and elect the above proposed officers for
1943. Mr. Weiss then turned the meeting over to the new President, Mr.
Comstock.

The President called the attention of the members to the fact that the
Certificate of Incorporation of The New York Entomological Society would
expire according to its terms on or about the Twenty-fifth day of February,
1943, and that it was desirable to extend the existence of the Society pur-
suant to Section 45 of the General Corporataion Law. After a discussion,
upon motion duly made by Mr. dos Passos, seconded by Mr. Sherman, and
unanimously carried, it was:

Resolved, That a special meeting of the members of The New York Ento-
mological Society be held at The American Museum of Natural History,

70

Journal New York Entomological Society

[Vol. Lin

Central Park West, New York City, New York, on the 19th day of January,
1943, at 7 : 45 P.M., Eastern War Time, to vote upon a proposition to extend
the existence of The New York Entomological Society in perpetuity, or for
such length of time as the members may decide at said meeting.

The secretary read the following Notice of the Special Meeting and the
proxy that was sent to all members of the Society.

THE NEW YORK ENTOMOLOGICAL SOCIETY

THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK, WEST
NEW YORK CITY, NEW YORK

NOTICE OF A SPECIAL MEETING OF THE MEMBERS TO
BE HELD ON THE 19th DAY OF JANUARY, 1943

TO THE MEMBERS OF THE NEW YORK ENTOMOLOGICAL SOCIETY:

Please take notice that by resolution of the members of The New York
Entomological Society duly adopted at a regular meeting thereof held on
the 5th day of January, 1943, a special meeting of the members of The New
York Entomological Society will be held at The American Museum of Natural
History, Central Park West, New York City, New York, on the 19th day of
January, 1943, at 7:45 P.M., Eastern War Time, for the purpose of voting
upon a proposition to extend the existence of said Society in perpetuity,
or for such length of time as the members may determine.

Attention is called to the fact that the corporate existence of The New
York Entomological Society will expire on the 25th day of February, 1943,
unless extended as the result of action taken at the aforesaid special meeting.
It is, therefore, very important that you attend said meeting either in person
or be represented by proxy. Please sign and return the enclosed proxy in any
event, so that we may be assured of a quorum.

January 6, 1943

Annette L. Bacon Wm. P. Comstock

Recording Secretary President

Note: No member in arrears in the payment of dues for over six months
shall be entitled to vote at said meeting unless satisfactory explanation is
given to the Executive Committee. (By-Laws: Art. XVI.)

THE NEW YORK ENTOMOLOGICAL SOCIETY

THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST
NEW YORK CITY, NEW YORK

PROXY

For a special meeting of the members of
The New York Entomological Society

Know all men by these presents, that the undersigned, a member of
The New York Entomological Society, hereby makes, constitutes and appoints
Harry B. Weiss, William P. Comstock, and Annette L. Bacon, or any one or

Mar., 1945]

Proceedings of the Society

71

more of them, true and lawful attorney and proxy for and in the name, place
and stead of the undersigned, to attend at the special meeting of the members
of The New York Entomological Society called to be held on the 19th day
of January, 1943, at The American Museum of Natural History, Central Park
West, New York City, New York, and to vote upon a proposition to extend
the existence of said Society in perpetuity or for such length of time as the
members may determine, and for the transaction of such other business as
may come before the meeting or any adjournment thereof, whether inci-
dental to any of the foregoing or otherwise; and at said meeting and any
adjournment or adjournments thereof, to vote in the name of the undersigned
as fully as the undersigned could do if personally present, hereby ratifying
and confirming all that the said attorney and proxy, or substitute duly
appointed, may do by virtue hereof, and hereby revoking any or all proxy
or proxies or powers of attorney in this behalf heretofore made by the
undersigned.

January , 1943. (L.S.)

The following resolutions were then adopted: Resolved, That the form of
notice of said special meeting, presented and read by the Recording Secre-
tary, be and the same hereby is approved; and it was further Resolved, That
the form of proxy to be used at said meeting, presented and read by the Re-
cording Secretary be and the same hereby is approved; and it was further
Resolved, That the Recording Secretary be, and she hereby is instructed and
directed to cause said notice of said special meeting to be given to the mem-
bers of the Society in accordance with the provisions of the corporation laws
of the State of New York.

There were four proposals for membership: Prof. T. C. Schneirla, New
York University, Department of Psychology; Mr. Henry S. Fleming, Depart-
ment of Tropical Research, Bronx Park; Mr. H. P. Boyd, Boy Scouts of
America; Mr. John G. Thorndike, 1075 Park Avenue, New York City.

The resignation of Mr. Adolf Klein was accepted with regret.

To serve on the Program Committee, Mr. Comstock appointed Mr. Becker,
Dr. Michener and Mr. Soraci.

Since the By-Laws of the Society were found to be out-of-date, the Presi-
dent appointed Mr. dos Passos and Miss Bacon members of a committee to
review the present By-Laws and to recommend modifications and corrections
of them sometime during this year.

Dr. Mont A. Cazier, the speaker of the evening, discussed Modern Methods
Applied in Systematic Analyses, using a genus of beetles as an illustration.
After a discussion, mainly by Mr. Weiss and Mr. Comstock, the meeting
was adjourned.

Minutes of a Special Meeting of the New York
Entomological Society

t

A special meeting of the members of The New York Entomological Society
was held at The American Museum of Natural History, Central Park West,
New York City, New York, on the 19th day of January, 1943, at 7:45 p.m.

The meeting was called to order by Mr. William P. Comstock, the President
of the Society, who acted as temporary Chairman.

72 Journal New York Entomological Society [Vol. Lin

Miss Annette L. Bacon, the Becording Secretary of the Society, acted as
Secretary of the meeting.

The Secretary presented a certified alphabetical list of the members of the
Society entitled to vote at the meeting.

The calling of the roll was dispensed with by unanimous consent.

The Secretary reported that the following nine members were present in
person: A. L. Bacon, C. F. dos Passos, J. D. Sherman, Jr., W. P. Comstock,
N. M. Payne, H. T. Spieth, W. T. Davis, L. J. Sanford, H. B. Weiss.

The Secretary reported that 66 members were represented by proxies to
Harry B. Weiss, William P. Comstock and Annette L. Bacon or any one or
more of them.

The proxies were presented and examined and upon motion duly made,
seconded and unanimously carried were ordered to be filed. Thereupon the
temporory Chairman announced that a quorum was in attendance at the meet-
ing and that the election of a permanent chairman was in order.

Upon motion duly made, seconded and unanimously carried, Mr. William
P. Comstock was elected Chairman of the meeting and Miss Annette L. Bacon
was appointed Secretary of the Meeting.

The Secretary presented an affidavit of Annette L. Bacon sworn to the
7th day of January, 1943, proving the service by mail on the 6th day of
January, 1943, of the notice of the special meeting of the members of The
New York Entomological Society to be held on the 19th day of January, 1943,
upon all the members of said Society, a list of whom is thereto attached and
marked exhibit B and the same was ordered to be filed.

After a discussion of the purpose for which the meeting was called, it was :

Resolved, That the duration of the Corporation is to be perpetual and it
was further:

Resolved, That the proper officers of the corporation to wit: William P.
Comstock, President, and Annette L. Bacon, Becording Secretary, be and
they hereby are authorized, directed and empowered to execute, subscribe and
acknowledge a Certificate of the Extension of The New York Entomological
Society and to file the same in the office of the Secretary of State of the
State of New York, pursuant to Section 45 of the General Corporation Law,
and it was further :

Resolved, That the said officers of the corporation be, and they hereby
are authorized, directed and empowered to take such further steps and pro-
ceedings as may be necessary or advisable in order to accomplish the purpose
of this meeting, including the payment of the filing fee of $25.00.

There being no further business the meeting adjourned.

CEBTIFICATE

I," the undersigned, Annette L. Bacon, Becording Secretary of The New
York Entomological Society, do hereby certify that annexted hereto and
marked Exhibit “A,” is a true and complete list of all the Members of The
New York Entomological Society as of January 19, 1943, entitled to vote at
the Special Meeting of the Members of said Society held on said date.
Dated, January 19, 1943

Annette L. Bacon
Recording Secretary

Mar., 1945]

Proceedings op the Society

73

State of New York )

County of New York j ss'

Annette L. Bacon duly sworn, deposes and says that she is the Recording
Secretary of The New York Entomological Society, that she executed the
foregoing Certificate as such and that the same is true to her knowledge.

Sworn to before me this
19th day of January, 1943

Members of The New York Entomological Society
entitled to vote on January 19, 1943

Alexander, C. P.

Gertsch, W. J.

Payne, N. M.

Angell, J. W.

Granek, I.

Petrunkevitch, A.

Bacon, A. L.

Gray, A.

Procter, W.

Barber, G. W.

Groth, C. F.

Rau, G.

Barber, H. G.

Hagan, H. R.

Readio, P. A.

Bell, E. L.

Halloek, H. C.

Rex, E. G.

Bequaert, J.

Harriot, S.

Richards, A. G.

Bird, H.

Hartzell, A.

Riehl, L. A.

Blackwelder, R. E.

Haskins, C. P.

Rosenblum, J. H.

Bromley, S. W.

Heineman, B.

Roth, L.

Brown, F. M.

Hessel, S. A.

Ruckes, H.

Church, F. E.

Hood, J. D.

Rumpp, N. L.

Clausen, L. W.

Horsfall, J. L.

Sanford, L. J.

Collins, D. L.

Huckett, H. C.

Satterthwait, A. F.

Comstock, W. P.

Hunter, R. J.

Schiller, W.

Connola, D. P.

Huntington, E. I.

Schmitt, A.

Crawford, J. C.

Janvrin, E. R. P.

Schott, F. M.

Creighton, W. S.

Johnson, F.

Schwarz, H. F.

Dalmat, H.

Johnston, J. W.

Scotland, M. B.

Davis, W. T.

Jones, F. M.

Shannon, H. J.

Dethier, Y. G.

Kisliuk, M.

Sherman, J. D.

Dietrich, H.

Klots, A. B.

Soraci, F. A.

Engelhardt, E. S.

Lacey, L.

Spieth, H. T.

Felt, E. P.

Lutz, F. E.

Swift, F. R.

Fenton, A. •

Melander, A. L.

Teale, E. W.

Forbes, J.

Michener, C. D.

Thomas, C. A.

Forbes, W. T. M.

Mutchler, A. J.

Watson, F. E.

Fox, H.

Nicolay, A. S.

Weiss, H. B.

Funkhouser, W. D.

Notman, H.

Zerkowitz, A.

Furness, G. C.

Osburn, R. C.

Ziegler, J. B.

Garman, P. H.

dos Passos, C. F.

Meeting of January 19, 1943

President Comstock in the chair; 23 members and visitors present.

The following were elected members of the Society: Mr. H. P. Boyd,
Mr. H. S. Fleming, Dr. T. S. Schneirla and Mr. J. G. Thorndike.

74

Journal New York Entomological Society

[Vol. LIII

Dr. James C. King, Red Oaks, White Plains, New York, was proposed
for membership.

The resignations of Dr. Wm. C. Moore and Mr. Samuel Harriot were
read, and accepted with regret.

Mr. Wm. T. Davis read a letter from the New York Academy of Sciences
stating that it would be necessary for the Society to share the cost of Museum
guards for the evening meetings. After much discussion, it was the con-
sensus of opinion that we oppose the charge.

Dr. H. T. Spieth moved that the members present express their thanks
to Mr. dos Passos for the work and time he spent on the reincorporation of
the Society. This was unanimously approved by a standing vote.

Mr. Comstock exhibited some Ancea butterflies from Chiapas, in southern
Mexico, from the collection of Mr. Frank Johnson, showing interesting sea-
sonal dimorphism.

Mr. Henry S. Fleming of the Department of Tropical Research, New York
Zoological Society, spoke of his experiences on a recent trip to Venezuela.

Meeting of February 2, 1943

President Comstock in the chair; 24 members and visitors present.

Dr. James C. King was elected a member of the Society.

Mr. Comstock reported on a New York Academy of Sciences Council Meet-
ing, which he attended as representative of our Society, regarding the charges
for rooms used during the evening for Society meetings. No action was
taken by the Society.

Because of the uncertainty of the presence of members of the Society,
Mr. Comstock decided not to appoint an Auditing Committee until one was
needed.

Mr. Alan S. Nicolay and Mr. Henry S. Fleming were appointed members
of the Field Committee.

Mr. E. L. Bell reported the death of Mr. John Boyd in the Solomon Islands.
The following resolutions were adopted:

Upon motion duly made seconded and unanimously carried by a standing
vote the following preambles and resolutions were duly adopted:

Whereas the Society has learned with deep regret of the death of John
Boyd of Southern Pines, North Carolina, as the result of wounds received
at Guadalcanal while serving in the armed forces of the United States of
America and

Whereas the deceased, a young and promising entomologist, was well and
favorably known to many members of the Society for his interest and field
work in Lepidoptera be it

Resolved, That the Society hereby records its deep regret at the untimely
passing of John Boyd and expresses its sincere sympathy to his parents,
Mr. and Mrs. Jackson H. Boyd of Southern Pines, North Carolina, and it
is further

Resolved, That the Secretary be, and she hereby is requested to transmit a
certified copy of these preambles and resolutions to Mr. and Mrs. Boyd.

Mr. Thomas D. Mulhern spoke on the Relation of New Jersey Mosquito
Control Work to the War Activity and showed 1600 feet of movie film. He

Mar., 1945]

Proceedings of the Society

75

brought several members of his staff with him. A vigorous discussion fol-
lowed in which many questions were answered.

Meeting of February 16, 1943

President Comstock in the chair; 14 members and visitors present.

A letter from Mr. Mulhern was read saying that the Eastern Association
of Mosquito Control Workers was about to issue a Directory of active mos-
quito workers.

Mr. Alan S. Nicolay spoke on Beetling in the Southern Appalachians with
particular emphasis on the races of beetles found on different ranges and
mountains, especially in Tennessee.

Meeting of March 2, 1943

Vice-President Teale in the chair; 24 members and visitors present.

The Committee on Reincorporation of the Society reported that the Cer-
tificate of Extension of Existence was obtained and that the Society was
extended in perpetuity as of February 17, 1943.

Dr. T. R. Gardner ,of the Foreign Parasite Introduction Laboratory,
Bureau of Entomology and Plant Quarantine, spoke on Unusual and Inter-
esting Habits associated with Oviposition of Parasitic Insects. An informal
discussion followed.

Meeting of March 16, 1943

Former President Weiss in the chair; 30 members and visitors present.

Dr. Robert Cushman Murphy, Chairman of the Department of Birds at the
American Museum, spoke on Insects and Other Arthropods of the Peruvian
Guano Islands, illustrating his talk with colored lantern slides.

Meeting of April 6, 1943

President Comstock in the chair; 17 members and visitors present.

Dr. R. G. Oakley, Bureau of Entomology and Plant Quarantine, Hoboken,
N. J., was proposed for membership.

Dr. Oakley spoke on Observations and Experiences in Guam, discussing the
topography, agriculture, and insect fauna of the island. Of the 1150 species
of insects recorded, many are pests — both of the crops (such as weevils, leaf
miners and corn borers) and of man (such as mosquitoes, flies and roaches).

Meeting of April 20, 1943

President Comstock in the chair; 28 members and visitors present.

Mr. dos Passos read the report of the By-Law Committee stating that a
new set of By-Laws had been drawn up, copies of which were being sent to
members of the Society. The report was accepted as read and the Com-
mittee was discharged.

Dr. R. G. Oakley was elected a member of the Society.

Dr. E. P. Felt spoke on Some Early Work in Entomology. He outlined
the work done by about twenty American economic entomologists who laid

76

Journal New York Entomological Society

[Vol. LIII

the ground work in the nineteenth century for a great deal of modern eco-
nomic and systematic entomology.

An interesting discussion followed, containing many reminiscences about
entomologists known to members of the Society.

Meeting of May 4, 1943

President Comstock in the chair; 54 members and visitors present.

Because of the large number of visitors present, all business was post-
poned until the next regular meeting.

Dr. A. L. Melander showed one of his excellent films of colored motion
pictures entitled 1 1 Animals from A to Z. ’ ’ This film was particularly de-
signed as an educational one to show children the many kinds of animals
found in this country. The members were very glad to see it before Dr.
Melander left for Biverside, California.

Meeting of May 18, 1943

President Comstock in the chair; 14 members and visitors present.

A resolution to give the remainder of the books belonging to the Society
to the Library of the American Museum of Natural History was passed.

Mr. Wm. T. Davis reported on observations of two methods of singing by
cicadas — vibrating membranes and the use of wings. In some species, both
males and females click their wings to produce a crackling sound.

Mr. G. C. Furness told of experiments leading to insect repellents for the
Army. Such repellents provide protection against malaria-bearing mos-
quitoes in the tropics. Different species of these are repelled with greater
effectiveness by different types of repellents.

Mr. Edwin W. Teale showed photographs of emerging Microgaster, the
parasite of sphingid caterpillars; also a picture of a mantid dining on a
short-tailed shrew.

Mr. Wm. P. Comstock showed five cases of mounted butterflies represent-
ing species found in Puerto Bico. Distribution and the effects of different
habitats on several of these species were discussed. The mass movements of
certain species, particularly the sulphurs, were considered as were such
peculiarities as seasonal variation, brood differences and local populations.

Meeting of October 5, 1943

President Comstock in the chair; 19 members and visitors present.

A letter from Miss Hazel Gay, Librarian of the American Museum, was
read in which she thanked the Society for the books given to the Museum
Library.

The scheduled speaker, Mr. C. F. W. Muesebeck of the U. S. National
Museum, was unable to be present so Mr. Comstock called for reports on
summer activities by members of the Society. On the whole not much col-
lecting was done, few members having time or means of transportation to
get out as they had done in the past.

Mr. Comstock showed a collection of butterflies from New Guinea.

Mar., 1945]

Proceedings of the Society

77

Meeting of October 19, 1943

Mr. Becker in the chair; 12 members and visitors present.

Dr. Joseph M. Ginsburg of Butgers University spoke on the Protection of
Outdoor Gatherings from Mosquitoes. In his discussion Dr. Ginsburg made
brief remarks on the more important species of mosquitoes and their habits.
Kepellents of the type applied to the skin, the essential features of long-
range mosquito control projects in New Jersey, and the efficacy of various
larvicides were discussed. He also spoke of the efficacy of the various aero-
sols used in repelling and knocking down adult mosquitoes.

Meeting of November 16, 1943

President Comstock in the chair; 12 members and visitors present.

Miss Lina Sordillo, Department of Insects and Spiders at the American
Museum of Natural History, was proposed for membership.

The amended By-Laws of the Society were read by Mr. Teale. Copies of
these were sent to the members of the Society before being voted upon.

President Comstock appointed as members of the Auditing Committee Dr.
Schneirla (Chairman), Mr. Fleming and Mr. Furness; and of the Nomi-
nating Committee Mr. Schwarz (Chairman), Mr. King, Mr. Nicolay and Mr.
Sherman.

Mr. Sherman reported that Dr. L. O. Howard was now living in Bronxville,
New York, with his daughter.

Dr. Nellie M. Payne of the American Cyanamid Company spoke on The
Life History of the Flat Grain Beetle, Lcemophloeus minutus (Oliv.).

Meeting of December 7, 1943

President Comstock in the chair; 18 members and visitors present.

Miss Lina Sordillo was elected a member of the Society.

Miss Margaret L. Guy, Department of Insects and Spiders in the American
Museum of Natural History, was proposed for membership.

A resolution of thanks, to the Department of Insects and Spiders of the
American Museum of Natural History for the gift of $75.00 to the Society,
was adopted.

The treasurer was authorized to offer a complete set of the Journal of the
Society to the Joint University Libraries, Nashville, Tennessee, at a special
price of $90.00.

It was with deep regret that the Society heard of the death of Dr. Frank
E. Lutz. It was moved and passed that a resolution be drafted by Dr.
Schneirla and sent to the family. A letter from Mrs. Lutz was read thank-
ing the members of the Society for the flowers they had sent.

A brief report of the treasurer was read mentioning the larger receipts
and disbursements for the year, and saying that the state of the finances of
the Society was about the same as a year ago.

Mr. Comstock read a few changes that had been made in the By-Laws, made
necessary by the use of the term “trustees” in the original certificate of

78

Journal New York Entomological Society

[Vol. LIII

Incorporation instead of “ executive committee” as said group has been
commonly called.

Dr. T. C. Schneirla spoke of The Place of Learning in Insect Life, and
illustrated his interesting discussion with lantern slides.

Meeting of December 21, 1943

President Comstock in the chair; 21 members and visitors present.

Miss Margaret L. Guy was elected a member of the Society.

Dr. Schneirla read the following resolution which was adopted, and the
secretary was instructed to send a copy of it to Mrs. Frank E. Lutz.

“We, the members of the New York Entomological Society, realize poig-
nantly that with the death of Dr. Frank E. Lutz on November 27 last there
has passed from the active lists one of the most valued and beloved of our
number. A senior member of the Society, having joined on March 16, 1909,
Dr. Lutz contributed fundamentally to the growth of the organization, serv-
ing it always as he served the interests of science in general, with the best
capacities of his unusual intelligence and genial personality. As a model for
others there stands his fine record of leadership as President of the Society
in 1925-1926 and as a member of the Executive Board for several years.

11 Vividly there remain in his many valuable publications, in the annals
of the Society, and in the memories of his countless friends, ineradicable
signs of his magnificent contributions as a natural scientist and great
teacher. His very human personality gleams constantly through his achieve-
ments, even as a masterful pun used in the title of a book to conceptualize
an important part of his work with ‘ i A Lot of Insects. ’ ’ His life work
shows most impressively the value of studying the small, ordinary and com-
mon things in nature as well as the great moments, an attitude that brought
him to many original and fundamental accomplishments in the study of
nature in general and insect nature in particular. His accomplishments are
indeed a priceless source of guidance and inspiration for the future.

“We resolve upon this statement as a token of the high esteem and honor
accorded Dr. Lutz as scientist and man by his fellow members in this
Society. ’ ’

It was with regret that the Society heard of the deaths of two more of
our old members: Dr. E. P. Felt, a member since 1907; and Mr. Christian
F. Groth, one of the first members of the Society. Mr. Groth became a
member in 1892 and held several offices during the next ten years. Mr. Sher-
man read the following resolution which was adopted.

“Whereas Christian F. Groth, our former President and one of the
earliest members of The New York Entomological Society, died on Decem-
ber 12th, 1943, at the age of 79 years, be it hereby Resolved, That the mem-
bers of this society convey to his son, Edward M. Groth, counsellor to the
U. S. Legation at Pretoria, South Africa, their deepest sympathy together
with an expression of appreciation of his father ’s enthusiastic interest in the
welfare of our Society, especially in its early years, and of the sense of per-
sonal loss felt by our older members in the death of this genial, lovable
friend. ’ ’

A motion was made and passed that the amended By-Laws as read on
December 7 and sent to all members on December 14 be adopted. A copy
of these By-Laws is printed below.

Mar., 1945]

Proceedings of the Society

79

The speaker of the evening, Dr. H. D. Smith of the U. S. Department of
Agriculture, spoke on Some Entomological Observations in Mexico. He told
of our government’s efforts to head off and control a possible invasion of
the citrus black-fly pest by the successful introduction of a chalcid parasite
into the Mexican areas affected.

Annette L. Bacon, Secretary.

CERTIFICATE OF INCORPORATION AND BY-LAWS OF
THE NEW YORK ENTOMOLOGICAL SOCIETY
ORGANIZED JUNE 29, 1892
INCORPORATED FEBRUARY 25, 1893
CERTIFICATE EXTENDED FEBRUARY 17, 1943

CERTIFICATE OF INCORPORATION OF
THE NEW YORK ENTOMOLOGICAL SOCIETY

We, the undersigned, Berthold Neumoegen, Charles Palm, Gustav
Beyer, Roderigues Ottolengui, citizens of and residents within the State
of New York, and George W. J. Angell, a citizen of and resident within
the State of Connecticut, all being of full age and citizens of the United
States, do hereby certify that we desire to form a Society pursuant to the
provisions of an act entitled “An Act for the incorporation of benevolent,
charitable, scientific and missionary Societies, ’ ’ passed April 12th, 1848, and
the several acts amendatory thereof.

First: The name of said Society shall be “The New York Entomological
Society ’ \

Second: The particular business and objects of said Society are the advance-
ment of the science of Entomology in all its branches.

Third: The term of existence of said Society shall be fifty years.

Fourth : The number of trustees who shall manage the affairs of said
Society is five.

Fifth : The names of the trustees .who shall manage the affairs of the Society
for the first year of its existence are, Berthold Neumoegen, Charles Palm,
Gustav Beyer, Roderigues Ottolengui and George W. J. Angell.

In witness whereof, we have hereunto subscribed our names this twenty-
first day of February, 1893.

Berthold Neumoegen.

R. Ottolengui.

Chas. Palm.

G. W. J. Angell.

G. Beyer.

80

Journal New York Entomological Society

[Vol. LIII

State of New York j
County of New York j SS'

On this twenty-first day of February, 1893, before me personally came
Berthold Neumoegen, Charles Palm, Gustav Beyer, Roderigues Otto-
lengui and George W. J. Angell to me personally known and known to
me to be the individuals described in and who executed the foregoing cer-
tificate, and they thereupon severally acknowledged to me that they had exe-
cuted the same.

Herbert F. Andrews

t

Notary Public, N. ¥. Co.

I hereby consent to and approve of the filing of the within certificate of
incorporation. N. Y., February 24th, 1893. Edwd. Patterson, Justice,
Supreme Court.

No. 70

State of New York |

City and County of New Yorkj

I, Henry D. Purroy, Clerk of the said City and County, and Clerk of the
Supreme Court of said State for said County, Do Certify, That I have com-
pared the preceding with the original Certificate of Incorporation of

THE NEW YORK ENTOMOLOGICAL SOCIETY

on file in my office, and that the same is a correct transcript therefrom,
and of the whole of such original.

Endorsed Filed and Recorded 25th February, 1893.

In witness whereof, I have hereunto subscribed my name and affixed my
official seal, this seventh day of June, 1893.

Henry D. Purroy
Cleric

CERTIFICATE OF EXTENSION OF EXISTENCE OF
THE NEW YORK ENTOMOLOGICAL SOCIETY

Pursuant to Section Forty-five of the General Corporation Law.

We, the undersigned, William P. Comstock and Annette L. Bacon, being
respectively the President and the Recording Secretary of The New York
Entomological Society do hereby certify:

1. The name of the corporation is The New York Entomological
Society.

2. The certificate of incorporation of said corporation was filed and re-
corded in the office of the Clerk of the County of New York on the
25th day of February, 1893, and filed in the office of the Secretary of
State on the 25th day of February, 1893.

3. The date on which the term of existence specified in the certificate of
incorporation will expire is the 25th day of February, 1943.

4. The duration of the corporation is to be perpetual.

Mar., 1945]

Proceedings of the Society

81

In witness whereof, we have made, subscribed and acknowledged this
certificate, this 9th day of February, 1943.

Seal

Attest: Annette L. Bacon
Recording Secretary

The New York Entomological Society
By William P. Comstock

President

State of New York |

County of New York j SS‘

On this 16th day of February, 1943, before me personally appeared Wil-
liam P. Comstock to me known and known to me to be the person described
in and who executed the foregoing Certificate of Extension of Existence of
The New York Entomological Society and he duly acknowledged to me that
he executed the same.

Thelma L. Ryan
Notary Public

Notary Public: Putnam County
Certificate Filed In

N. Y. Co. Clerk’s No. 1048-Reg. No. 4R617
Com. expires March 30, 1944

State of New York |

County of New York^SS‘

On this 16th day of February, 1943, before me personally appeared Annette
L. Bacon to me known and known to me to be the person described in and who
executed the foregoing Certificate of Extension of Existence of The New
York Entomological Society and she duly acknowledged to me that she exe-
cuted the same.

Thelma L. Ryan
Notary Public

Notary Public: Putnam County
Certificate Filed In

N. Y. Co. Clerks No. 1048-Reg. No. 4R617
Com. expires March 30, 1944

State of New York j
County of New York j

William P. Comstock, being duly sworn, deposes and says: that he is the
President of The New York Entomological Society; that he has been duly
authorised to execute and file the foregoing certificate by the votes, cast in
person or by proxy, of a majority of the members of record of the corpora-
tion; that such votes were cast at a meeting of the members duly called for
that purpose upon like notice as that required for the annual meeting of the
corporation and the date of such meeting was the 19th day of January, 1943.

William P. Comstock
President

82

Journal New York Entomological Society

[Vol. LIII

Subscribed and sworn to before me
this 9th day of February, 1943.

Charles J. Kerr, Notary Public

Notary Public, Queens County Clerk’s No. 1186

N. Y. Co. Clk’s #426, Beg. #4-K-196 Commission Expires March 30, 1944

State of New York )

County of New York rS’

Annette L. Bacon, being duly sworn, deposes and says: that she is the
Eecording Secretary of The New York Entomological Society; that she
has been duly authorised to execute and file the forgoing certificate by the
votes, cast in person or by proxy, of a majority of the members of record
of the corporation; -that such votes were cast at a meeting of the members
duly called for that purpose upon like notice as that required for the annual
meeting of the corporation and the date of such meeting was the 19th day
of January, 1943.

Subscribed and sworn to before me
this 9th day of February, 1943.

Charles J. Kerr, Notary Public
Notary Public, Queens County Clerk’s No. 1186
N. Y. Co. Clk’s #426, Eeg. #4-K-196
Commission Expires March 30, 1944

I Certify That I have compared the preceding copy with the original
Certificate of Extension of Corporate Existence of

filed in this department on the 17th day of February, 1943, and that such
copy is a correct transcript therefrom and of the whole of such original.

Witness my hand and the official seal of the Department of State at the
City of Albany, this seventeenth day of February, one thousand nine hundred
and forty-three.

State of New York Department of State Filed Feb. 17, 1943 Tax $ None
Filing Fee $25 Thomas J. Curran, Secretary of State By B. Horan,
Asst. Cashier

The original is filed in the office of the County Clerk and Clerk of the
Supreme Court, New York County. File Number 477-1893. Filed February
26, 1943.

Annette L. Bacon
Eecording Secretary

State of New York
Department of State

THE NEW YOEK ENTOMOLOGICAL SOCIETY

Frank S. Sharp
Deputy Secretary of State

Mar., 1945]

Proceedings of the Society

83

BY-LAWS
Article I
Members

The Society shall consist of active, life and honorary members.

1. Active members shall be persons interested in entomology, who shall be
entitled to vote and hold office.

2. Life members shall be active members who shall have paid fifty dollars
($50.00) in lieu of annual dues. They shall be entitled to vote and hold
office.

3. Honorary members shall be eminent entomologists elected in recognition
of their service to science. There shall not be more than twelve (12) honor-
ary members at any one time. They shall not be entitled to vote or hold
office.

Article II

Election of Members

All candidates for membership must be proposed by an active member of
the Society at a regular meeting. They shall be voted upon individually at
the following regular meeting and the affirmative vote of at least two-thirds
of the members present (given by voice, or by ballot if demanded) is re-
quired for election unless these provisions are waived by unanimous consent
of the members present.

Article III

Officers and Committees

1. Officers of the Society shall consist of a President, a Vice-President,
a Secretary, an Assistant Secretary, a Treasurer, an Assistant Treasurer, and
an Editor.

2. Elective committees of the Society shall consist of five trustees (one
of whom shall be the president) who shall constitute the Executive Com-
mittee who are hereinafter referred to as such, and a Publication Committee.
The Executive Committee shall be composed of the President (Chairman),
and four active members, all entitled td vote. The Editor, Secretary, and
Treasurer shall also be members of the Executive Committee but not entitled
to vote. The Publication Committee shall be composed of the Editor (Chair-
man), and two active members.

3. Standing Committees of the Society to be appointed by the President,
shall consist of an Auditing Committee, composed of three active members;
and a Field Committee, composed of two active members.

4. Temporary committees may be appointed by the President at his dis-
cretion to perform special duties which he shall define. The President also
shall appoint a Nominating Committee, consisting of three active members,
to nominate a full slate of officers, trustees and elective committees at the
annual meeting.

84

Journal New York Entomological Society

[Vol. LIII

ARTICLE IV

Election of Officers and Committees

1. Officers, trustees and members of elective committees shall be elected at
the annual meeting of the Society by a majority vote of the members pres-
ent, or voting by proxy. They shall hold office for one year and/or until
their successors shall be elected.

2. Any vacancy that may occur among the officers, or elective committees,
except as hereinafter provided, shall be filled by appointment by the Execu-
tive Committee. The person appointed to fill the vacancy shall hold office
until the next annual meeting and/or until the election of his successor.

ARTICLE V

Duties of Officers and Committees

1. The President shall preside at all meetings. He shall appoint all com-
mittees except the elective committees, and shall be chairman of the Executive
Committee and a member^ ex-officio of all other committees except the publi-
cation committee.

2. The VicerPresident shall assume the duties of the President in case of
the death, resignation, absence or disability of the President. In case both
the President and Vice-President are absent at a meeting a temporary chair-
man may be chosen by the members present to preside at that meeting.

3. The Secretary shall keep the minutes of the meetings of the Society and
of the Executive Committee; give notice of the meetings of the Society when
not otherwise herein provided for; advise members of their election; attend
to all general correspondence; keep all records and files of the Society and
generally perform such services as may be delegated to him by the Society.

4. The Assistant Secretary shall act in case of the death, resignation,
absence or disability of the Secretary and shall assist the Secretary as
need be.

5. The Treasurer shall receive all moneys for the Society and deposit them
in the name of the Society in such banking institutions as the Executive Com-
mittee may direct; he shall pay therefrom by draft or check all bills and
obligations not exceeding Twenty-five Dollars ($25.00), and all others when
approved by the President or the Editor. He shall keep an account of all
monetary transactions and shall exhibit a statement of them when called for
by the President, Editor, Executive Committee or Auditing Committee and
shall make a full report for the preceding calendar year at the annual meet-
ing. He shall notify members respecting the payment of dues within ten
days after their election and thereafter when annual dues become payable,
and shall send out membership cards on receipt of dues. At the expiration
of his term of office, the Treasurer shall deliver to his successor all funds,
papers, books and vouchers belonging to the Society.

6. The Assistant Treasurer shall act in case of the death, resignation,
absence or disability of the Treasurer and shall assist the Treasurer as
need be.

Mar., 1945]

Proceedings of the Society

85

7. The Editor shall have general charge, management and supervision of
the publication known as the Journal of the New York Entomological Society.
The Editor shall be Chairman of the Publication Committee, as hereinbefore
provided.

8. The Executive Committee shall meet at the call of the President, who
shall be Chairman thereof. It is empowered to call for a report from any
of the officers or committees of the Society at its discretion. It shall keep
minutes of its proceedings which shall be submitted at the next succeeding
regular meeting of the Society. It shall have general charge of the funds,
investments and property of the Society. It shall decide on the status of
members in arrears of dues. It shall determine the subscription price of the
Journal and discounts allowed in connection with its sale, as well as the price
of any other publications of the Society. It shall appoint one active member
of the Society as Delegate to meetings of the New York Academy of Sciences.

9. The Publication Committee shall cooperate with the Editor in the publi-
cation of the Journal of the Society. If, at any time, other publications are
undertaken, the Publication Committee shall be in charge of the production
thereof.

10. The Auditing Committee shall examine the accounts and reports of the
Treasurer and shall report to the Society thereon.

11. The Program Committee shall plan and arrange for the programs of
the meetings.

12. The Field Committee shall arrange for and manage the excursions and
outings of the Society.

13. The Society’s Delegate to the New York Academy of Sciences shall
attend meetings thereof and shall report at the next regular meeting of the
Society concerning any action taken which may affect the Society. The Dele-
gate shall have no power to obligate the Society without the previous authori-
zation of the Executive Committee or except by action taken at a meeting
of the Society.

ARTICLE YI
Publication Funds

All funds subscribed or donated for the Journal or other publications of
the Society shall be used for no other purpose than those specified.

ARTICLE YII
Dues

The dues of active members shall be Three Dollars ($3.00), per annum,
payable in advance on the first day of January of each year. New members,
if elected on or after October 1, shall pay no dues for the year of their
election. Honorary members shall be exempt from the payment of any dues.

ARTICLE VIII
Members in Arrears

All members in arrears in the payment of dues for one year, shall lose the
privilege of voting or holding office. Before the annual meeting the Treasurer

86

Journal New York Entomological Society

[Vol. LIII

shall present a list of the members in arrears in the payment of dues to the
Executive Committee, which shall decide upon dropping such members from
the roll of the Society or postponing for good cause the payment of their dues
for a definite period.

ARTICLE IX

Subscription to the Journal

1. The subscription price of the Journal and the price of single numbers to
active members, and discounts allowed to subscription agencies and on sales
of sets shall be determined by the Executive Committee.

2. Subscriptions shall be payable in advance on the first of January of each
year.

3. The Journal shall be sent gratis to all Life and Honorary members.

ARTICLE X
Meetings

1. Regular meetings of the Society shall be held at The American Museum
of Natural History (or at such other place as the membership shall determine)
on the first and third Tuesdays of each month at 8 : 00 p.m. No regular meet-
ings will be held during the months of June, July, August and September or
upon a legal holiday or upon the first Tuesday of January.

2. The annual meeting of the Society shall be held at The American
Museum of Natural History (or at such other place as the membership shall
determine) on the first Tuesday in January in each year at 8: 00 p.m., if not
a legal holiday, otherwise on the third Tuesday.

3. Special meetings of the Society may be called by the Secretary upon a
written request of the President or 10 active members. Such request shall
state the purpose for which the meeting is to be called and the time and place
where it is to be held. No other business except that specified in the call
shall be transacted except by unanimous consent of the members present.

4. Eleven (11) members shall constitute a quorum for the transaction of
business at an annual meeting and seven (7) members shall constitute a
quorum at any regular meeting.

5. At any special meeting, members in good standing may vote or be rep-
resented by proxy.

6. Whenever notice of any meeting is required by these by-laws it shall be
deemed sufficient if published in the Bulletin of the New York Academy of
Sciences or if given by postal card and addressed to each member of the
Society at his last known address at least ten (10) days and not more than
twenty (20) days before the meeting, or if given as required by the General
Corporation Law of the State of New York.

ARTICLE XI
The Order of Business

The order of business of regular meetings shall be as follows:

1. Reading of minutes.

Mar., 1945]

* Proceedings of the Society

87

2. Reports of officers.

3. Reports of committees.

4. Election of members.

5. Proposals for membership.

6. Miscellaneous business.

7. New business.

8. Reading of papers and scientific discussion.

9. Adjournment.

The order of business of the annual meeting shall be as follows :

1. Reading of minutes.

2. Roll call, verification of proxies^

3. Annual reports of officers.

4. Reports of committees.

5. Election of officers, trustees and elective committees for ensuing year.

6. Miscellaneous business.

7. Proposals and elections for membership.

8. Reading of papers and scientific discussion.

9. Adjournment.

The order of business may be changed or suspended at any meeting with the
consent of two-thirds or more of the members present.

ARTICLE XII
Amendments

These by-laws may be amended at any regular meeting or at a special meet-
ing of the Society called for that purpose by the vote of two-thirds or more
of the members present, provided that the proposed amendment or amend-
ments shall have been submitted in writing and presented at a previous meet-
ing of the Society and due notice thereof having been given in conformity
with the provisions of Article X.

88

Journal New York Entomological Society

[Vol. LIII

CORRECTIONS TO LIST OF MEMBERS OF THE
NEW YORK ENTOMOLOGICAL SOCIETY

Mr. F. E. Watson was elected to honorary membership on Oc-
tober 3, 1939, and this fact should have been indicated opposite
Mr. Watson’s name in the list of members appearing in the
December, 1944 issue of this Journal.

An unfortunate error was made in the same list of members,
by printing ‘ ‘deceased” after the name of Cyril F. dos Passos.
The death of Mrs. Cyril F. dos Passos occurred on August 29,
1944 and not that of her husband. See the article entitled “Viola
Harriet dos Passos and Her North American Moths,” by William
P. Comstock in this issue.

NOTES ON THYSANIA ZENOBIA (CRAMER),—
LEPIDOPTERA, (HETEROCERA)

By Howard Schiff

A female specimen of Thysania zenobia (Cramer), common in
the tropics but as yet rare in New York State, was captured in
the village of Montieello, New York, on September 18, 1944. The
weather was fairly warm for the time of the year and a large
variety of different species of Catocala and other Noctuidae were
on the wing. The specimen of Thysania zenobia had alighted
near lights and was quite sluggish and easily caught. Rips on
both forewings and other wing damages seemed to indicate a long,
hard trip to New York from a distant point.

Mar., 1945]

Book Notice

89

BOOK NOTICE

A Bibliography of the Homoptera (Auchenorhyncha). By Zeno
Payne Metcalf, N. C. State College of Agriculture and En-
gineering of the University of North Carolina, n.d. [1945],
n.p. [Raleigh, N. (C.]. 10J x 8J inches, Vol. I, 886 p., Vol.

II, 186 p. Cloth, $7.50 net, postage 24 cents.

Although the accumulated knowledge of ages is to be found in
books and papers, students are lost in an ocean of literature until
the repositories of this knowledge are made known to them by
bibliographies. I am always glad to see a good bibliography,
especially of a subject in which I am interested, and it is always
a relief to know that I have been saved an enormous amount of
preliminary work. Bibliographers never receive all the credit
that is due them and we are apt to use the results of their knowl-
edge and industry, without gratitude.

This new bibliography of the suborder Homoptera, series
Auchenorhyncha which includes the superfamilies Fulgoroidea
and Cicadoidea is an impressive work in two volumes. In the
first volume the authors are listed alphabetically and their papers
are listed chronologically, with titles, sources and dates of pub-
lication. This volume includes approximately 8,000 titles which
appeared in 1,000 separate journals and in over 900 books, all in
various languages. All titles up to June 1, 1942 are included.
Where the title of the paper is not indicative of its contents, a
brief note supplies this deficiency and serves as a guide in deter-
mining its value for a particular purpose. Volume II consists of
a list of the periodicals, their abbreviations and complete titles,
with places of publication, and a highly useful topical index,
which is general, taxonomic and geographic, thereby enabling the
student to find the topic in which he is interested.

In the Introduction, Dr. Metcalf states that the work has been
in preparation during his spare time, for more than 30 years, and
from the magnitude of the finished product, this may be readily
understood. There are some who take the position that a bibli-
ography should be all inclusive and list everything that has been
written, even if it is a repetition of previous publications. This

90

Journal New York Entomological Society

[Vol. LIII

may be a safe method for bibliographers who are not intimately
acquainted with a particular subject, but I am glad to see that in
the present work Dr. Metcalf exercised his discrimination and
eliminated summaries of other publications, popular articles and
works of an ephemeral nature. There is nothing more annoying
than investigating long lists of references that yield exactly noth-
ing. Users of this bibliography can depend upon Dr. Metcalf’s
wisdom and experience in separating the valuable from the
worthless.

For many years Dr. Metcalf has specialized on the Homoptera
and I am sure that his impressive and authoritative bibliography
will receive the praise and recognition that it deserves. — Harry
B. Weiss.

No. 2

Vol. LIII

JUNE, 1945

Journal

of the

New York Entomological Society

Devoted to Entomology in General

HARRY B. WEISS JOHN D. SHERMAN, Jr.

T. C. SCHNEIRLA

Subscription $3.00 per Year

Published Quarterly by the Society
N. QUEEN ST. AND McGOVERN AVE.

LANCASTER, PA.

NEW YORK, N. Y.

1945

CONTENTS

A Supplement to the New York State List of Coleoptera,

No. 6, Additions and Corrections

By Borys Malkin 91

Check-List of the Psychodidae of Europe

By William F. Rapp, Jr., and Janet L. Cooper 117

William Thompson Davis — 1862-1945

By Harry B. Weiss 127

Anteos maerula, a New Butterfly Recorded From Florida

By Leonard J. Sanford 136

The Wax of Stingless Bees (Meliponidae) and the Uses to
Which It Has Been Put

By Herbert F. Schwarz 137

Insect Enemies of the House-Fly, Musca Domestica L.

By S. W. Bromley 145

The Selective Penetration of Fat Solvents Into the
Nervous System of Mosquito Larvae

By A. Glenn Richards, Jr., and Jane L. Weygandt 153

Insect Food Habit Ratios of the Lloyd-Cornell Reservation

By Harry B. Weiss 167

Proceedings of the Society 169

NOTICE: Volume LIII, Number 1, of the Journal of
the New York Entomological Society Was Published
on May 4, 1945.

Entered as second class matter July 7, 1925, at tlie post office at Lancaster, Pa.,
under the Act of August 24, 1912.

Acceptance for mailing at special rate of postage provided for in Section 1103.
Act of October 3, 1917, authorized March 27, 1924.

JOURNAL

OF THE

New York Entomological Society

Vol. LIII June, 1945 No. 2

A SUPPLEMENT TO THE NEW YORK STATE LIST
OF COLEOPTERA, NO. 6, ADDITIONS
AND CORRECTIONS1

By Borys Malkin

Since the publication of Cooper’s2 supplement to the New York
State list of Coleoptera, numerous records have been added to the
known beetle fauna of the state. These the writer has attempted
to gather together in a single paper presented below. As in the
above mentioned pamphlet, it contains not only records new to
the state or those inadequately represented in the list, but also
a list of current literature through which a number of records
and notes on New York beetles have been scattered.

As usual, the double asterisk (**) signifies species new to the
state list. A single asterisk (*) indicates species new to the
so-called “ continental’ ’ New York. A double dagger (J) de-
notes species previously unrecorded from Long Island, while rec-
ords new to Staten Island are preceded by a single dagger ( f ) .

Although the writer’s own collection served as the main source
of material listed in this paper, numerous records came to his at-
tention from elsewhere. Collections of the American Museum of
Natural History, and of Mr. William Spector, Lt. (j.g.) D. G.
Kelley, Sgt. J. H. Kremer, Mr. Chas. Ragot, and Mr. J. W.

1 Cornell University Agr. Exp. Sta. Mem. 101, Aug. 1926 (issued Jan.
1928). (Coleoptera, pp. 203-520.)

2 For a complete list of literature pertaining to New York Coleoptera see:
K. Cooper, “A Supplement to the New York State List of Insects Devoted
to the Coleoptera. Additions, Notes and Corrections. ’ ’ Bull. Brooklyn Ent.
Soc., Vol. XXX, No. 4, pp. 142-159.

92

Journal New York Entomological Society [Vol. liii

Green must be mentioned here. To Sgt. Kremer and to Mr.
Green the writer is indebted for extensive cooperation, not only
in the way of specimens and identifications, but also for help in
compiling necessary data of varied nature.

All records are followed by the collector ’s name .in italics. If
no such name appears it may be assumed that the author was the
collector. The names of the taxonomists are represented by
abbreviations as follows:

H. B. Leech— (L), C. A. Frost— (F), J. W. Green— (G), K.
M. Fender — (KF), R. E. Blackwelder — (REB), C. T. Parsons —
(P), F. T. Scott— (S), W. S. Fisher— (WSF), 0. L. Cartwright
— (OLC), B. E. White— (W), L. L. Buchanan— (B), A. F. Sat-
terthwait — (AFS), Eugene Ray — (ER), and B. Malkin — (M).

The bibliography includes only such papers as have been pub-
lished since the appearance of Mr. K. C. Cooper’s supplement, or
papers not cited by him at that time.

CICINDELIDiE
(Malkin 1941)

59. Cicindela longilabris Say. Upper Saranac, Jl, Green (G).
CARABIDiE

(Van Dyke 1936, Darlington 1936, 1938, Buchanan 1939,
Malkin 1941c, 1941d)

** . Cardbus sp.f LI: Queens Village, May, under bark,

Funaro , (M).

**166. Carabus mceander Fisch. Dunkirk, Jl., on the shore of
Lake Erie, Funaro, (M).

J651. Bembidion patruele Dej. LI: Rockaway, Aug., (F) ;
Montauk, May, Kremer and Malkin, (M).

J723. Bembidion versicolor (Lee.). LI: Rockaway, Aug.,
(F).

$737. Bembidion quadrimaculatum (L.). LI: Forest Hills,
Apr., Kelley, (M) ; Wantagh, Jl., (F) ; Alley Pond, Nov., Ragot
and Malkin, (F).

f820. Tachyura levipes Csy. SI: Great Kills, Jn., (M).

1 825. Tachyura capax (Lee.). SI : Great Kills, Jl., (M).

f 841. Tacky s cuneatus Csy. SI : Great Kills, May, (M).

June, 1945]

Malkin: Coleoptera

93

**888. Tachyta angulata Csy. Mt. Whiteface, 4000 ft., Jl.,
Green, (G).

**889. Tachyta kirbyi Csy. Mt. Whiteface, 4000 ft., Jl.,
Green, (G).

$1285. Celia patruelis (Dej.) LI: Flushing, Mar., Cooper,
(M).

fl324. Celia schotti Csy. SI : A single specimen from Ch.
Schaeffer’ s collection labelled S.I., 3.20.87, (M).

1511. Platynus stygicus Lee. Saratoga, Aug., Kr enter, (Val-
entine det.).

**1583. Platynus picicornis Lee. Upper Saranac, Jl., Green *

(G).

f 1586. Platynus gemellus Lee. South Fallsburg, Jn., Spector,
(G) ; LI : Alley Pond, Nov., Ragot and Malkin, (F, M).

2039. Selenophorus pedicularius Dej. Van Cortlandt Park,
Jn., (F).

2236. Stenolophus humidus Ham. Bear Mt., Apr., Spector,
(G). This capture verifies old state record of the list.

HALIPLIDiE
(Malkin 1941c, 1941d)

DYTISCHLF

(Fall 1937, Wallis 1939, 1939a, Leech 1940, Malkin 1941c, 1941d)
2346. Hydrocanthus iricolor Say. LI : Cunningham Park,
Queens, Jl., Kelley, (L).

2400. Celina angustata Aube. South Fallsburg, Jn., Spector,
(L) ; LI: Brooklyn, Jl., Spector, (L).

**2425. Coelamhus laccophilinus ( Lee.). LI : May, (G, M).
2637. Dytiscus hybridus Aube. LI : Cunningham Park,
Queens, Jl., Kelley, (L).

** . Hydaticus modestus Shp. White Plains, Apr.,

Squire, (L), misdetermined as H. loevipennis Thom, in collection.

**2670. Cybister flavocinctus Aube. LI : Cunningham Park,
Queens, Jl., Kelley, (L).

GYRINIDiE
(Malkin 1941c, 1941d)

**2703. Gyrinus pect oralis Lee. Upper Saranac, Jl., Green,

(G).

94

Journal New York Entomological Society

[Vol. LIII

19251. Gyrinus bifarius Fall. LI : Wantagh, Apr., Malkin
and Kremer, (M).

HYDROPHILIDiE

(d’Orchymont 1933, Malkin 1941c, 1941d)

*2781. Berosus exiguus Say. White Plains, May, Squire
(M) ; LI : Wantagh, Apr., Kremer and Malkin, (M).

**2874. Cercyon quisquilius (L.). Ithaca, May, Van Nos-
trand, (M) ; South Fallsburg, Aug.-Sept., Sped or, (F) ; Upper
Saranac, Jl., Green, (G) ; LI: Brooklyn, Jl., Spedor, (G).

2879. Cercyon lateralis (Marsh). Upper Saranac, Jl., Green,
(G) ; South Fallsburg, Sept., Spedor, (G).

**2888. Cercyon tristis (Illig). Van Cortlandt Park, Jl.,
(M) ; Port Ontario, Jl., Green, (G) ; Fair Haven, Jl., Green, (G).

19289. Cercyon terminatus Marsh. LI : Brooklyn, Sept.-Dee.,
Spedor, (G).

LEPTODIRUME

**2949. Catops gratiosa Blanch. Upper Saranac, Jl., Green,
(Hatch det.).

J 3022. Leiodes polita Lee. Forest Park, Jn., Kelley, (G).

CLAMBIDiE

3048. Clambus puberulus Lee. ? LI : Brooklyn, Nov., Spec-
tor, (F).

SCYDMAENIDJE

*3073. Euconnus salinator (Lee.) Plattsburg, Jl., Green,

(G).

STAPHYLINIDiE

(Blackwelder 1936, 1939, Voris 1936, Wolcott 1937,

Malkin 1941c, 1941d)

**33 — . Proteinus sp. Upper Saranac, Jl., Green, (G).

**33 — . Proteinus sp. Upper Saranac, Jl., Green, (G).

3332. Megarthrus americanus Sachse. Upper Saranac, Jl.,
Green, (Fall det.).

3425. Acidota subcarinata Er. Ithaca, Oct., Van Nostrand ,

(F).

**3426. Acidota quadrata Zett. Manhattan, Oct., (REB).

June, 1945]

Malkin: Coleoptera

95

**3497. Trogophloeus memnonius Er. Riverside Drive, Jn.,
flying, (REB).

**3577. Oxytelus nimius Csy. Upper Saranac, Jl., Green,

(G).

**3579. Oxytelus invenustus Csy. Upper Saranac, Jl., Green,
(G) ; Darts, Oct., Cooper, (G).

3668. Bledius tau Lee. Phoenicia, Jn., Green, (G) ; Upper
Saranac, Jl., Green, (G).

$3908. Homoeotarsus bicolor (Grav.). LI: Montauk, May,
Kremer and Malkin, (M).

*3910. Homoeotarsus badium (Grav.). Van Cortlandt Park,
May, (M).

f 3926a. Homoeotarsus pallipes var. capito (Csy.). Yonkers,
March-April-Nov., (REB); Piermont, Oct., Kremer, (F) ; SI:
Great Kills, Jn., (REB) ; Princess Bay, Sept., (REB) ; LI: Kew
Gardens, Aug., Kelley , (GOM) ; Forest Park, March, Kelley ,
(G, M).

**3931. Homoeotarsus parviceps (Csy.). SI: Great Kills,
Jl, (REB).

4094. Lithocharis ochracea (Grav.). LI: Brooklyn, Nov,
Sped or, (F).

**4252. Stilicolina tristis (Melsh.). Pelham Bay Park, Jn,
(REB, M).

*4265. Astenus binotatus (Say). Van Cortlandt Park, May,
(M).

4325. Leptacinodes flavipes (Lee.). LI: Brooklyn, Nov,
Spector, (F).

**4386. Philonthus furvus Nordm. Ithaca, May, Van Nos-
trand, (F).

*4429. Philonthus thoracicus (Grav.). Woodlawn Park,
March, (F).

**4461. Philonthus siegwaldi Mann. Upper Saranac, Jl,
Green, (Fall det.).

|4470. Philonthus nigritulus (Grav.). LI: Kew Gardens,
March, Kelley and Malkin, (F).

f4483. Philonthus validus Csy. Upper Saranac, Jl, Green,
(G) ; Copake, Jl, G. Eiten, (F) ; SI: Great Kills, Apr, (F).

f 19349. Philonthus tetragonocephalus Notm. Bronx Park,

96

Journal New York Entomological Society

[Vol. LIII

Apr., (F) ; Van Cortlandt Park, Aug., (F) ; Moshulu, May, (F) ;
Copake Falls, Sept., (F) ; Ithaca, Oct., Van Nostrand, (F) ; SI:
Hugonot, Apr., (F).

** . Ocypus spJ Hunter’s Island, N. Y. C., May, Spec-

tor, (G) ; LI: Amagansett, Sept., Angell, (G). Probably an in-
troduced species.

4545. Staphylinus cinnamopterus Grav. LI : South Hunting-
ton, Apr., Kremer and Malkin, (M).

4553. Ontholestes capitatus (Bland). Slide Mt., Jn., Spector,

(G).

4569a. Heterothops fumigatus fusculus Lee. Lk. Placid, Jl.,
Green, (G).

*4574. Quedius ferox Lee. Yonkers, March, (F).

4586. Quedius capucinus (Grav.). Upper Saranac, Jl.,
Green, (G).

**4592. Quedius virginicus Csy. LI: Jones Beach, May,
Spector, (G).

4660. Tachinus addendus Horn. Upper Saranac, Jl., Green ,
(Fall det.).

**4692. Coproporus Icevis (Lee.). Van Cortlandt Park, Jn.,
very rare, among C. ventriculus, (M).

**4714. Conosoma elongata Blatch. Van Cortlandt Park,
May, (M).

4716. BoletoMus niger (Grav.). LI: Sunken Meadow State
Park, Jn., Kelley, (M, G).

4732. Boletobius qucesitor Horn. Upper Saranac, Jl., Green,

(G).

** . Boletobius spJ Slide Mt., Jn., Spector, (M).

*4738. Mycetoporus humidus Say. Van Cortlandt Park, Jn.,
(REB).

4740. Mycetoporus consors Lee. Bear Mt., Jn., (M).

5508. Datomicra inopia Csy. Lake Placid, Jl., Green, (G) ;
Upper Saranac, Jl., Green, (G).

J5780. Aleochara pubemda Klug. LI : Brooklyn, May, Spec-
tor, (G).

| 5833. Emplenota maritima Csy. LI : Brooklyn, May, Spec-
tor , (G).

PSELAPHIDiE
(Wolcott 1937)

June, 1945]

Malkin: Coleoptera

97

SCAPHIDIIDiE
(Malkin 1941c)

HISTERIDJE

(Reinhardt 1926, Hinton 1935, 1935a, Wenzel 1936, Ross 1940,
Malkin 1941c, 1941d, Bradley 1943)

6531. Hololepta lucida Lee. Hunters Island, Spector, (F) ;
Tuxedo, (M).

6574. Hister memnonius Say. LI : Brooklyn, Sept., Spector,
(G) ; Wading River, Nicolay, (G).

**6575. Hister egregius Csy. South Fallsburg, Aug., Spector.

(G).

**6616. Hister cariniprons Sf. LI : Montauk, Oct., Angell,
(Siepmann det.) ; Amagansett, Sept., Angell, (G. M).

6624. Hister sedecimstriatus Say. LI : Bellport, Jl., Aug..
Oct., Nicolay, (G).

6646. Teretrius latebricola Lew. Upper Saranac, Jl., Green,
(Ballou det.).

**6846. Saprinus minutus Lee. LI: Rockaway Beach, May,
Jl., Spector collection, (G).

6875. Saprinus sphaeroides Lee. LI : Riverhead, Jn., Die-
trich, (G).

** . Saprinus semistriatus Scriba. LI: Jamaica, Jl.,

Spector, (G).

** . Saprinus sp. LI : Sunken Meadows State Park, Jn.,

Kelley, (G).

£6909. Gnathoncus deletus (Lee.). LI: Brooklyn, Sept.,
Spector, (G).

PHENGODIDiE

*7037. Omethes marginatus Lee. Bear Mt., Jn., (G).
CANTHARID^E

(Green 1941, Malkin 1941c, 1941d)

**7054. Podabrus nothoides Lee. East Windham, Jl., Green,

(G).

J7058. Podabrus basillaris Say. LI : Rockaway, Jl., Ragot,
(G).

7078. Podabrus piniphilus (Esch.). Upper Saranac, Jl.,

98

Journal New York Entomological Society

[Vol. LIII

Green, (G) ; Slide Mt., Spector, (G) ; Maplecrest, Jn., Schott,

(G).

7080. Podabrus puberulus Lee. Upper Saranac, Jl., Green,
(G) ; E. Windham, Jl., Green, (G) ; Corinth, May, Marx, (G).

7083. Podabrus laevicoUis (Kby.). Upper Saranac, Jl.,
Green, (G).

**20859. Podabrus appendiculatus Fall. Van Cortlandt
Park, Jn., (G) ; Pelham Bay Park, Jn., (G) ; LI: Brooklyn, Jn.,
Spector, (G).

**20883. Podabrus brevicollis Fall. East Windham, Jl.,
Green, (G).

** . Podabrus sp. nov. Bear Mt., Jn., (KF) ; Schott,

(G) ; Ft. Montgomery, Jn., Schott, (G) ; Upper Saranac, Jl.,
Green, (G).

7094. Cantharis tantillus Lee. Bear Mt., Jn., (G). This spe-
cies is recorded from N. Y., by Green (Green 1941), but without
definite locality.

**7103. Cantharis nanulus Lee. Van Cortlandt Park, Jl.,

(G).

** . Cantharis antennatus Green. New Rochelle, Jn.,

(G) • Bear Mt., Jl., (G) ; SI : Great Kills, Jn., JL, (G).

f7127. Cantharis marginellus Lee. SI : Great Kills, Jn., Jl.,
(G, KF).

7188. Malthodes fuliginosus Lee. Bear Mt., Jn., (KF).
Swept from Rubus.

MALACHIIDJE
(Malkin 1941d)

CLERIDJE
(Malkin 1941d)

LYMEXYLID^K

7740. Melittomma sericeum Harr. Van Cortlandt Park,
Tuxedo, Bear Mt., Sutton Island, SI : Great Kills. Larvae and
pupae reared in great numbers from oak. The adults hatched
out between May 10 and May 26, although a single specimen from
Bear Mt. emerged July 14.

CEPHALOIDiE

7748. Typitium ungulare (Lee.). Upper Saranac, Jl., Green ,
(G) ; Catskill, Oliverea, Shoemaker, (G).

June, 1945]

Malkin: Coleoptera

99

MORDELLIDJE
(Malkin 1941c, 1941d)

**7867. Mordellistena arnica Lee. Mosholu, in American
Mus., coll. (M) ; SI: Great Kills, Jl., (M).

**7911. Mordellistena bihamata (Melsh.). N. Y., in Schaef-
fer’s coll. (M) ; New Rochelle, Jl., Funaro, (M).

f7913. Mordellistena fuscata (Melsh.). New Rochelle, Jn.,
Funaro , (M) ; SI: Great Kills, Jl., (M).

7915. Mordellistena pityptera Lee. Bear Mt., Jl., Schott,
(M).

**7921. Mordellistena fusco-atra Hellm. Whitehall, Jn., Die-
trich, (ER).

**7922. Mordellistena gig as Lilj. Ithaca, Jl., (ER).

MELOIDiE

(Malkin 1941c)

— — . Macrobasis murina Lee. Upper Saranac, Jl., Green,
(G) ; South Fallsburg, Jn., Spector, (G).

8142. Meloe impressus Kby. South Fallsburg, Sept., Spector,
(F).

PYROCHROIDJE

8221. N eopyrochroa femoralis Lee. Bear Mt., Jn., (M) ;
Upper Saranac, Jl., Green, (G).

ANTHICIDiE

8387. Anthicus floralis (L.). LI: Brooklyn, Nov., Spector,
(F).

**8389. Anthicus scenicus Csy. LI : Brooklyn, Nov., Spector,
(M), with floralis L.

*8448. Anthicus melancholicus Laf. Bear Mt., Jn., (F) .

**8455. Anthicus rusticus Csy. Sparkill, Jn., Siepmann, (F).

EUGLENIDJE

8480. Zonantes signatus (Hald.). Bear Mt., Jn., (F).

ELATERIDiE

(Brown 1933, Malkin 1941c, 1941d)

*8596. Monocrepidius lividus (DeG.). Van Cortlandt Park,
Jl., (M) ; Bronx, Jl., (M) ; New Rochelle, Jl., Funaro, (M). A

100

Journal New York Entomological Society

[Vol. LIII

very common species, often swept at night from low grasses,
plants, etc.

8629. Limonius plebejus (Say) . Croton Falls, May, (M).

8654. Pheletes r^mbatus (Say). Bear Mt., Jn., (M).

8791. Ludius rotundicollis (Say). Nyack, March, (F).

** . Dalopius cognatus Brown. Maplecrest, Catskill Mts.,

Jn., Schott, (Lane determined).

8910. Gly phony x inquinatus (Say). Van Cortlandt Park,
Jl., (M) ; New Rochelle, Jl., Funaro, (M).

*8951. Ampedus xanthomus Germ. New Rochelle, Sept.,
Funaro, (M).

*8956. Ampedus pusio Germ. New Rochelle, Jl., Funaro,
(M); Aug., (M).

*8980. Megapenthes limbalis (Hbst.). Montrose, Jl., Ragot,
(M) ; Bear Mt., Jl., (M).

9020. Melanotus canadensis Cand. Bronx, Jl., Under stone,
(M).

9048. Melanotus pertinax (Say). Croton Falls, May, sweep-
ing, (M).

MELASIDJE
(Malkin 1941c)

9133. Deltometopus amoenicornis (Say). LI: Cunningham
Park, Queens, Jl., Kelley, (GOM).

BUPRESTIDyE

(Fisher 1940, 1942, Malkin 1941b, 1941c, 1941d,
Obenberger 1934, Heifer 1941)

9399. Agrilaxia flavimana (Gory). SI : Great Kills, Jl., (M).

9481. Actenodes acornis (Say). Montrose, Jl., Ragot, (M) ;
Napeque, Angell, (M).

9518. Agrilus crinicornis Horn. Van Cortlandt Park, Jn.,
(F) ; New Rochelle, Jn., (F).

9522. Agrilus cephalicus Lee. Copake, Jl., G. Eiten, (F).

HELMIDJE
(Sanderson 1938)

HETEROCERIDiE

*9653. Heterocerus auromicans Kies. Port Ontario, Jl.,
Green, (G).

June, 1945]

Malkin: Coleoptera

101

DASCILLIDiE
(Brown 1944)

HELODID M
(Malkin 1941c)

DERMESTIDiE

(Malkin 1941c, 1941d, Barber 1942, 1942a)

BYRRHID.E
(Malkin 1941c)

9864. Cytilus alternatus (Say). LI: Broad Channel, May,
Spector (F).

OSTOMID.F
(Malkin 1941c)

NITIDULID^F

(Parsons 1938, 1943, Malkin 1941c)

*10060. Carpophilus antiquus (Melsh.). Harmon, Sept., (P).
10061. Carpophilus nit ens Fall. Harmon, Sept., (F).
**10082. Epurcea adumbrata Mann. Upper Saranac, Jl.,
Green, (G).

**10100. Stelidota strigosa (Gyll.). SI : Great Kills, Jl., (P).
*10136. Glischrochilus obtusus (Say). Morris Otsego, Jn,
Angell, (P) ; New Rochelle, May, (M).

**10138a-. Glischrochilus samguinolentus var. rubromaculatus
Reitt. Upper Saranac, Jl., Green, (P).

**21855. Glischrochilus siepmanni Brown. South Fallsburg,
Sept., Spector, (F).

EROTYLIDiE
(Malkin 1941c, 1941d)

**10318. Tritoma erythrocephala Lac. Van Cortlandt Park,
Jl, (M, F).

**10321. Tritoma mimetica Csy. Elbridge, Jn, (G).
CRYPTOPHAGID^H

10361. Loberus impressus Lee. Van Cortlandt Park, Jl, (M).
$10483. Anchicera ovalis Csy. LI : Brooklyn, Nov, Spector,
(F).

102

Journal New York Entomological Society

[Vol. LIII

COLYDIIDiE
(Malkin 1941c, 1941d)

**10531. Synchita granulata Say. LI : Long Island, Ch Bagot
collection. The labelling of this specimen should be verified.

(M) .

**10589. Penthelispa hcematodes (Fab.). Greenwood Lake,
March, Nicolay, (M).

**10602. Cerylon clypeale Csy. SI: Great Kills, Apr., (F).

LATHRIDIHLE
(Malkin 1941c, 1941d)

COCCINELLID^G
(Timberlake, 1943)

(Dobzhansky 1931, 1942, Malkin 1941c, 1941d, McKenzie 1936)
10931. Hyperaspis octavia Csy. Port Henry, JL, Kremer,
Saratoga, Aug., Kremer , (all Chapin and M. det.).

19743. Hyperaspis lugubris (Rand). New Rochelle, Jn.,

(N) ; Bear Mt., Jn., (N).

*11010. Scymnus semiruber Horn. New Rochelle, Aug.,
under bark of willow trees, (M) ; SI: Great Kills, Jn., (S).
**11015. Scymnus rubricauda Csy. Bear Mt., Jn., (M).
*11016. Scymnus chromopyga Csy. Van Cortlandt Park,
May-Jn., (N).

**11076. Scymnus coniferarum Cr. Bear Mt., May, (M).
11188. Coccinetta undecimpunctata L. LI : Sagaponack,

Aug., Kremer , (Chapin and M. det.).

11194&. Adalia frigida var. humeralis (Say) . Van Cortlandt
Park, May, (M).

ALLECULIME

11336. Mycetochara bicolor Coup. Upper Saranac, Jl.,
Green , (G).

TENEBRIONIDJE
(Blaisdell 1934, Malkin 1941c)

**12316. Platydema erythrocerum Cast, and 'Bril. Bear Mt..
Sept., Kremer and Malkin, (F).

12327. Platydema picilabrum Melsh. Van Cortlandt Park,
Nov., (M).

June, 1945]

Malkin: Coleoptera

103

112433. Paratenetus fuscus Lee. Harmon, Sept., (M) ; LI :
Manhasset, May, (M) ; Sea Cliff, May, (M).

MELANDRYID^E
(Malkin 1941c)

12536. Synstrophus repandus (Horn). SI : May, Leng, (G).

PTINIDiE
(Brown 1944)

12601. Mezium americanum Lap. Manhattan, Jl., (M) ; Kew
Gardens, March, Kelley and Malkin, (M).

ANOBIIDJE

12738. Xyletinus lugubris Lee. Upper Saranac, Jl., Green,

(GO-

CISIDJE

**12973. Cis impressa Csy. Yonkers, Sept., from Polyporus ,
Ragot, (M).

SCARABAEIDiE

(Wallis 1928, Kuntzen 1933, Hoffman 1935, Robinson 1936,
Malkin 1941c, 1941d, Cartwright 1943)

13082. Onthophagus orpheus Panz. Montrose, Feb., Ragot,
(OLC) ; Sep., Ragot and Malkin, (OLC) ; Bear Mt., Jn., (OLC).

**13208. Dialytellus dialytoides Fall. Upper Saranac, Jl.,
Green, (G).

fl3216. Ataenius abditus (Hald.). SI : Great Kills, Jn., (M).

13220. Ataenius imbricatus (Melsh.). New Rochelle, Sept.,
Funaro, (M) ; LI : Jamaica, Jl., Spector, (G) ; Brooklyn, Jl.,
Spector, (G).

. Ataenius falli Hinton. Yan Cortlandt Park, May-Jl.,

Sept., (OLC) ; Bronx, Jl., (OLC) ; Bear Mt., Jn., (OLC).

**13237. Dialytes truncatus (Melsh.). Montrose, Sept.,
Ragot, (M).

**13238. Dialytes ulkei Horn. South Fallsburg, Jl., Spector ,

(G).

$13337. Trox capillaris Say. LI: Rockaway, Jl., Ragot,
(OLC).

**19961. S erica opposita Dawson. Cooks Falls, Angell,
(OLC) ; LI: Brooklyn, Jl., Spector, (G).

104

Journal New York Entomological Society

[Vol. LIII

$13517. Phyllophaga drakeiKby. LI : Rockaway, Jl., (OLC).
*13742. Pachystethus oblivia Horn. White Plains, Jl., Squire.
One of the commonest Rutelince in the state.

19986. Anomala orient alis Waterh. LI : Valley Stream, Spec-
tor, (G).

*13803. Ochrosidia villosa (Burm.). Bronx, Jl., L. Faas,
(OLC).

*14022. Trichiotinus assimilis (Kby.). Upper Saranac, Jl.,
Green, (Robinson det.).

PASSALIDtE
(Hincks and Dibb 1935)

CERAMBY CIDH3

(Ran 1935, Van Dyke 1937, Hopping 1937, Dillon and
Dillon 1941)

15137. Ob erea pallida Csy. Upper Saranac, Jl., Green, (G).
CHRYSOMELIDJE

(Heikentinker 1925, Hatch and Beller 1932, Blake 1933, 1935,
1936, 1936a, 1943, Van Dyke 1938, Barber and Bridwell
1940, Barber 1943, Malkin 1941a, 1941c, 1941d)

15256. Lema sexpunctata Oliv. SiPGreat Kills, Oct., (M).
Beaten from Solidago.

** . Exema canadensis Pierce. . Collected by the writer

in several localities in the vicinity of N. Y. C., and in N. J., (W).

**15308. Griburius scutellaris (Fab.). Yonkers, Sept., Bagot,
(W) ; South Fallsburg, Jn., Spector, (G).

15470. Pachybrachys hepaticus (Melsh.). White Plains, Jn.,
Squire, (W).

**15536a. Bassareus lituratus var. castus (Melsh.). Van
Cortlandt Park, Jn., (W).

**15536c. Bassareus lituratus var. niger Blatch. Van Cort-
landt Park, Jl., (M).

*15559. Colaspis favosa Say. New Rochelle, Jl., Funaro,
(M) ; Bear Lake, Aug., Funaro (M).

**15708. Chrysomela interrupta var. quadriguttata Sf. Van
Cortlandt Park, May. A single specimen taken with C. inter-
rupta from Salix. Described from B. C.

June, 1945]

Malkin: Coleoptera

105

** . Galerucella n. sp. Upper Saranac, Jl., Green , (Fall

det. ).

t20196. Galerucella spircece Fall. Flushing, Jn., Van Nos-
trand, (F).

:f20197. Galerucella alni Fall: Flushing, Jn., Van Nostrand,

(F).

**20198. Galerucella perplexa Fall. Upper Saranac, Jl.,
Green, (Fall det.).

*15755. Erynephala maritima (Lee.). Van Cortlandt Park,
Apr., flying, (M).

15865. Oedionychis gibbitarsa (Say). Kensico, Aug., Ragot.
(M),

** . Longitarsus n. sp. Yonkers, Nov., (L. G. Gentner

det.) ; LI : Manhasset, May, (L. G. Gentner det.).

16133. Microrhopala excavata (Oliv.). LI: Manhasset, May,
(M).

MYLABRIDiE

16171. Mylabris chinenis (L). LI: Alley Pond, Aug.,
Kremer, (G, M).

*16218. Mylabris alboscutellatus Horn. Van Cortlandt Park,
Jn-Jl, (M).

BRENTIDJE
(Buchanan 1939)

CURCULIONIDAE

(Buchanan 1934, 1937, 1937a, Hatch 1938, Henderson 1939,
Malkin 1941c, 1941d, Schoof 1942, Tanner 1943)

16325. Rhinomacer pilosus Lee. Saratoga, Aug., Kremer,
(G, M).

16389. Apion molestum Fall. Van Cortlandt Park, Jn., (M).
16396. Apion perminutum Sm. Van Cortlandt Park, Jl., (B).
Listed by Fall without specific locality.

16397. Apion reclusum Fall. SI: Great Kills, Oct., (M).
Listed by Fall without specific locality.

**16407. Apion coxale Fall. Van Cortlandt Park, Jn., (B) ;
SI : Great Kills, Jn., Jl., (B).

16422. Apion patruele Sm. Van Cortlandt Park, Jn., (B) ;
Yonkers, May, Jl., (B) ; Pelham Bay Park, Jn., (B) ; Bear Mt..

106

Journal New York Entomological Society

[Vol. LIII

Jl., (B) ; New Rochelle, Jn., (B) ; SI : Great Kills, Jn., Jl., Sept.,

(B).

16423. Apion walshi Sm. LI: Manhasset, May, (B).

116429. Apion iurlmlentum Sm. Yonkers, May, (B) ; SI:
Great Kills, Jl., (B).

16469. Apion emaciipes Fall. Bear Mt., Jl., (B) ; Van Cort-
land! Park, May, (B). Listed by Fall without specific locality.

16472. Apion carinatum Sm. New Rochelle, Aug., (B).
Confirms Fall’s state record of the list.

16473. Apion attenuatum Sm. Van Cortlandt Park, Jl., (B).

**21444. Apion dilaticolle Fall. Van Cortlandt Park, May,
(B) ; Croton Falls, May, (B) ; Croton on Hudson, Sept., (B) ;
Pelham Bay Park, Jn., (B) ; SI: Great Kills, Jn., Jl., (F, B).

**16723. Aphrastus griseus Blatch. Bronx, Jn., (B) ; Van
Cortlandt Park, Jn., (M). Rare, usually in company of A.
tceniatus Gyll.

**16740. Sitona discoidea Gyll. Tuxedo, May, (M).

** . Sitona cylindricollis Fahr. Van Cortlandt Park, Jl.,

(M) ; Croton Falls, May, (M) ; Croton-on-Hudson, Sept., (F) ;
Highlands, Aug., Kremer, (M) ; Ithaca, May, Van Nostrand,
(M). Abundant on clover, usually accompanied by other spe-
cies.

*16755. Phytonomus eximius Lee. Van Cortlandt Park, May,
(M) ; Montrose, Sept., Ragot, (M) ; Croton Falls, May, (M).

16776. Listronotus callosus Lee. Van Cortlandt Park, Jl.,
(M). Addition to LeConte’s state record.

16786. Listronotus frontalis Lee. Fairhaven, Jl., Green, (G).
Confirms old LeConte. state record.

*16804. Hyperodes cryptops (Dietz). Van Cortlandt Park,
Jl., (M).

**16807. Hyperodes grypidioides (Dietz). Van Cortlandt
Park, Jn., Jl., (M).

16931. Pachyphanes amoenus ( Say). Harmon, Sept., (M).

**16947. Smicronyx tesselatus Dietz. Croton-on-Hudson,
Sept., (F).

**16955. Smicronyx picipes Dietz. SI: Great Kills, Jl., (M).

16973. Smicronyx tychioides Lee. Yonkers, Jl., (M) ; SI :
Great Kills, Jn., (M).

June, 1945]

Malkin: Coleoptera

107

**16975. Smicronyx congestus Csy. New Rochelle, Aug.,
(M).

**16978. Smicronyx apionides Csy. SI : Great Kills, May,
(B).

** . Bagous sp. Bear Mt., Jn., (B) ; LI: Flushing, May,

Bagot, (B).

** . Endalus sp. Pelham Bay Park, Jn., (B) .

** . Endalus sp. Bear Mt., Jn., (B).

**17023. Onychylis longulus Lee. New Rochelle, May,
Funuro, (F).

117284. Anthonomus decipiens Lee. SI : Great Kills, Jn., Jl.,
(F).

1 17290. Anthonomus elongatus Lee. LI : Manhasset, May,
(B).

**21518. Anthonomus nubiloides Fall. SI : Great Kills, Jn.,
(F).

**17298. Anthonomopsis mixtus Lee. Upper Saranac, Jl.
Green, (G).

17351. Acalyptus carpini (Hrbst.). Bear Mt., May, (B).

17360. Miarus hispididus Lee. Van Cortlandt Park, Jn.,
(M) • Bear Mt., Jn., Jl., (M) ; Harmon, Sept., (M).

17415. Lixus musculus Say. Harmon, Sept., (F).

17447. Baris umbilicata (Lee.). Van Cortlandt Park, May,
(B) ; New Rochelle, Jn., (B) ; Pelham Bay Park, Jl., (B) ; SI :
Great Kills, Jl., (B) ; LI : Flushing, May, Bagot, (B).

**17509. Plesiobaris disjuncta Csy. SI : Great Kills, May,
Jn., (B).

^17550. Ampeloglypter ater Lee. Highlands, Aug., Kremer,
(G, M) ; LI: Sea Cliff, Aug., Spector, (G, M).

**17602. Centrinopus alternatus Csy. Van Cortlandt Park,
May, (B).

17646. Limnobaris rectirostris (Lee.). Elbridge, Jn., Spector,
(F).

**17671. Catapastus conspersus (Lee.). Bear Mt., Jl., (B).

17727. Psomus armatus (Dietz). Van Cortlandt Park, May, .
(B).

17735. Acanthoscelis curtus (Say). Phoenicia, Jn., Green,
(B).

108

Journal New York Entomological Society

[Vol. LIII

**17754. Pelenosomus cristatus Dietz. Van Cortlandt Park,
May, (M) ; SI : Great Kills, May, (M) ; LI : Sea Cliff, May, (M).

. Myllocerus castaneus Roelf . LI : Sea Cliff, May, beaten

from oak and maple, (M) ; Alley Pond, Aug., Kremer, (M).

*17758. Acallodes lysimachice Fall. Bear Mt. Jn., Jl., (M).

** . Ceutorhynchus erysimi Fab. Pelham Bay Park, Jn.,

(M) ; Yonkers, JL, (M).

J17781. Ceutorhynchus sulcipennis Lee. LI : Aqueduct, Aug.,
Engelhardt, (G).

J17783. Ceutorhynchus cyanipennis Germ. LI : Manhasset,
May, (M) ; Sea Cliff, May, (M).

** -. Ceutorhynchus sp. Van Cortlandt Park, Jn., (B).

*17824. Perigaster cretura (Hrbst.). Van Cortlandt Park,
Jn., (M).

** . Bhinoncus castor (Fab.). Saratoga, Aug., Kremer,

(B).

17831. Pelenomus griseus Blatch. Van Cortlandt Park, May,
(M) ; LI: Sea Cliff, May, (M).

**17901. Rhyssematus cequalis Horn. SI : Great Kills, Jl., on
milk weed, (M).

17929. Tyloderma punctata Csy. LI: Cypress Hills, May,
Engelhardt, (G).

17973. Cryptorhynchus fuscatus Lee. SI: Great Kills, March,
(B). Hibernating under bark of willow.

**180875. Rhodohcenus 13-punctatus var. 5-punctatus (Say).
Yonkers, Sept., Ragot, (F) ; Port Ontario, Jl., Green, (G) ; Sodus
Pt., Jl., Green, (G) ; Fire Is., N. Y. C., Jl., Spector, (G).

118123. Sphcenophorus minimus Hart. Yonkers, Aug., Ragot,
(AFS) ; SI : Willowbrook, Sept., Ragot, (AFS) ; LI : Jones Beach,
May, Spector, (G).

18141. Sphcenophorus venatus (Say). Pelham Bay Park,
Jn., (AFS).

** . Genus not found. “1 specimen labelled ‘Van Cort.

Park, N. Y.’ Belongs to some exotic species not recorded from
North America. Probably from Japan or neighboring regions.
Looks like an old specimen and locality label may be open to
question.” (Buchanan in litt.)

June, 1945]

Malkin: Coleoptera

109

SCOLYTIDiE

(Blackman 1934, 1938, Pechuman 1937)

. Scolytus multistriatus (Marsh). Ithaca, Aug. 1, 1935,

Van Nostrand, (P) . This is the easternmost record of the species
from the state.

BIBLIOGRAPHY

Barber, H. S. and Bridwell, J. C. 1940. Dejean Catalogue Names. Bull.
Br. Ent. Soc, 35: (1), 1-12. (The following changes proposed:

Hcemonia nigricornis Kby. to Macroplea nigricornis Kby. Gastroidea
should be known as Gastrophysa, Deloyala clavata becomes Plagio-
metriona clavata Fab., while Chirida guttata (Oliv.) becomes Deloyala
clavata (Oliv.).)

Barber, H. S. 1941. Bull. Br. Ent. Soc., 36: (1), 27-28. (Notoxus bicolor
Say should be changed to N. muripennis (Lee.).)

. 1942. Some Synonymy in Dermestes. Bull. Br. Ent. Soc., 37 : (5),

174-176. ( Dermestes vulpinus Fab., a synonym of D. maculatus Deg.)

. 1942a. Raspberry Fruit Worms and Related Species. U. S. Dept.

Agr. Misc. Publ. no. 468, pp. 1-32. ( Byturus sordidus Barber — West

Point. B. rubi Barber-Marion (type, by H. Glasgow), Palmyra, Niagara
Falls. Records of B. unicolor Say of the list unreliable in view of this
new revision. Mr. Barber also believes that the genus and perhaps the
entire group Byturince ought to be removed from Dermestidce.)

. 1943. Notes on Rhabdopterus in the United States. Bull. Br. Ent.

Soc., 38: (4), 111-120. ( Rhabdopterus preetextatus (Say) probably

found in N. Y. area. It. deceptor Barber — New York.)

Blackman, W. M. 1934. A Revisional Study of the Genus Scolytus Geoffroy
(Eccoptogaster Herbst) in N. A. U. S. Dept. Agr. Tech. Bull. No. 431,
30 pp. ( Scolytus sulcatus Lee. — Yonkers (O. L. Wolfberger), Staten
Island (E. A. Firaz). Scolytus multistriatus Marsham. A European
species introduced into the United States. New York, Long Island.
A new record for 11 Continental ’ ’ New York.)

. 1938. Jour. Wash Acad. Sci., 28: (8), 534-545. ( Chramesus

hicorice Lee. — New York.)

. 1942. Revision of the Genus Phlceosinus Chapuis. Proc. U. S.

N. M., 92: (3154), 397-474. ( Phlceosinus rugosus, Swaine probably

erroneously listed this species as its known distribution is restricted to
California and Juniperus occidentalis is the host.)

. 1943. New Genera and Species of Bark Beetles of the Subfamily

Micracinae. Proc. U. S. N. M., 93: (3165), 341-365. ( Micracis rudis

Lee. separated from the original genus and placed in Hylocorus. M.
asperulus Lee. united with M. opacicollis Lee. and placed in Micracisella.)

Blackwelder, R. E. 1936. Revision of the North American Beetles of the
Staphylinid Subfamily Tachyporinae — Part I : Genus Tachyporus Graven-
horst. Proc. U. S. Nat. Mus., 84: 39-54. ( Tachyporus chrysomelinus L.

110

Journal New York Entomological Society

[Vol. LIII

is a European species which does not occur here and therefore should be
omitted. It is probably mixed with jocosus Say and rulomus Blackw. in
collections.)

. 1939. A Generic Revision of the Staphylinid Beetles of the Tribe

Psederini. Proc. IT. S. Nat. Mus., 87 : 93-125. (Several changes in
generic arrangement of Pcederini.)

Blaisdell, F. E. 1934. Studies in the Genus Corticeus. Ent. News. 14,
187-191. ( Corticeus to replace Hypophlceus after Biologia Centrali-

Americana.)

Blake, D. H. 1933. Revision of the Beetles of the Genus Disonycha.
Proc. U. S. N. M., 82: (28), 1-66. ( Disonycha procera Csy. — New York

City, West Point, Whiteface Mt., L. I. T>. arizonce Csy. is a valid spe-
cies— New York. D. alternans Illig. — Long Island. D. admirabilis
Blatch. — West Point, Long Island. T>. gldbrata Fab. — New York. D.
latifrons var. laticollis Sf. — L. I., Wyandach (type). D. uniguttata
Say — Whiteface Mt., Staten Island. T>. fumata Lee. — occurs in the
southwestern portion of the country and series of specimens from New
York in the J. B. Smith collection may be incorrectly labelled. D. penn-
sylvanica parva Blatch, is a synonym of D. pennsylvanica Illig. D. penn-
sylvanica pallipes Cr. is a synonym of D. uniguttata Say. The correct
name for Z>. xanthomelcena Dalm. is xanthomelas.)

. 1935. Notes on Systena. Bull. Br. Ent. Soc., 30: (3), 89-107.

( Systena blanda Melsh. to replace S. tceniata Say and tceniata of the list
probably refers to the former species.)

. 1936. Proc. Ent. Soc. Wash., 38: 13-24. ( Altica ambiens var.

alni (Harris) — New York.)

. 1936a. A Redisposition of the Monoxia puncticollis and Allied

Genera. Jl. Wash. Acad. Sci., 26: 426-430. ( Monoxia maritima Lee.

is a valid species and should replace M. puncticollis of the list under new
generic name Erynephala. The typical E. puncticollis Say is a western
species and does not occur in New York.)

-. 1943. The Generic Position of Eypolampsis pilosa (Illig) and some

Related Species. Ent. Soc. Wash., 45: (9), 207-225. ( Distigmoptera a

new name proposed for Hypolampsis. D. apicalis Blake — L. I., Rock-
away. Also an additional record of D. pilosa (Illig) for Long Island
from the same locality.)

Bradley, J. C. 1943. Notes on Synonymy and Distribution of American
Histerids. Bull. Br. Ent. Soc., 38: (4), 123. ( Hololepta fossularis Say

replaces 3. incequalis Say. The latter a synonym. Similarly, Platysoma
lecontei Mars, replaces P. depressum Lee. Pseudister hospes Lew. most
likely a S. A. species not Neararctic.)

Brown, W. J. 1933. Can. Ent., 65 : 173-182. ( Betarmon geminatus Rand,

should become Agriotella geminata Rand.)

. 1944. Some New and Poorly Known Species of Coleoptera. Can.

Ent., 76: (1), 4-10. ( Eurypogon harrisi (Westw.) — N. Y. C. Mezium
afline Boiel — N. Y.)

June, 1945]

Malkin: Coleoptera

111

Buchanan, L. L. 1934. Proc. Ent. Soc. Wash., 36: 205-207. ( Hylobius

radicis Buch'. — Ballston Spa, Saratoga Co. (type) — O. L. Thompson and
G. H. Harris), Albany (H. L. McIntyre). E. confusus Kby. should be
changed to H. congener Dalla Torre.)

. 1937. Bull. Bk. Ent. Soc., 32: (5), 205-207. ( Ceutorhynchus

americanus Buch. — Ithaca, Illion, Buffalo, Orangeburg, should replace
C. cyanipennis Germ, which is considered to be a synonym of sulcicollis
Payk. and is not found as yet in N. A.)

. 1937a. Notes on Curculionidse. Jl. Wash. Ac. Sci., 27 : 312-16.

( Trachyphlceus bifoveatus Beck. — Barnevald 1917, Oriskany 1931 [In the
U.S.N.M. collection]. This is a European species closely related to T.
davisi Blatch. Gymnetron netum Germ. — New York. The correct name
for G. tetrum Fab. is G. teter. Ceutorhynchus punctiger Gyll. to replace
marginatus Payk. Perigaster lituratus Dietz — New York.)

. 1939. Proc. Ent. Soc. Wash., 41: 79-82. (The following changes

in synonymy are proposed: Agonoderus pallipes Fab. to A. lecontei
Chaud. Eupsalis rinnuta Dru. to Archenodes minutus Dru. Orchestes
pallicornis Say to Bhynchcenus pallicornis. Cryptorhynchus lapathi L.
to Sternochcetus lapathi L.)

Cartwright, O. L. 1943. Bull. Br. Ent. Soc., 38: (3), 108. ( Atcenius

spretulus Hald. a valid species and to be separated from A. strigatus
Say, consors Fall, and falli Hinton.)

Chapin, E. A. 1938. Three Japanese B^eties of the Genus Serica Mac-
Cleay. Jl. Wash. Ac. Sci., 28: 5&4-”£4g£ ( Serica peregrina Chap. — L.

I., Westbury (type), Douglaston. Undoubtedly introduced from Japan.)

Darlington, P. J. 1936. Two Recently Introduced Species of Amara.
Psyche., 43: (1), 20. ( Amara humilis Csy. of the list is a synonym of
European A. familiaris Duft. A. cenea (Deg.) formerly A. devincta
Csy. — Walton.)

. 1938. American Patrobini. Ent. Am., 18: 135-183. ( Patrobus

longipalpus Notman a synonym of P. rugicollis Rand. P. foveicollis
foveicollis (Esch) cited from isolated higher mountains of New York.)

Dillon, L. S. and E. S. 1941. The Tribe Monochamini in the Western
Hemisphere. Reading Public Museum and Art Gal. Sci. Publ. No. 1,
pp. 1-135. ( Goes tessalatus Hald. to replace Hammoderus tessalatus

Hald. — N. Y. Plectrodera scalator Fab. — L. I., Huntington. Mono-
chamus carolinensis Oliv. — N. Y., L. I. — Rockaway Beach, Flatbush.)

Dobzhansky, T. 1931. The North American Beetles of the Genus Coc-
cinella. Proc. U. S. N. M., 80: art. 4, 1-32. ( Coccinella monticola
Muls. a subspecies of the Asiatic nivicola Men. C. perplexa Muls. a
synonym of C. trifasciata L., while the variety C. transversoguttata quin-
quenotata is united with the original species.)

. 1942. Beetles of the Genus Hyperaspis inhabiting the U. S.

Smiths. Misc. Coll., 101: (6), 1-94. ( Hyperaspis lugubris (Rand) ad-

ditional records — West Point, Pinelawn. H. separata Csy. a synonym
of this species. E. lewisi Cr. — West Point, Bear Mt. H. serena Csy. a

112

Journal New York Entomological Society

[Yol. LIII

subspecies of fimbriolata Melsh., additional Long Island records — Baby-
lon, Long Beach, Yaphank. E. disconotata troglodytes Muls. — Cascade,
West Point. The Cascade record, originally referred to H. disconotata
proper, belongs to Mulsant’s subspecies. E. octavia Csy. — West Point,
Ithaca, Buffalo, Mt. Whiteface, Rockaway Beach. E. lateralis Muls., a
record cited from Buffalo, must be erroneous as the species is a western
one. E. signata Oliv. and E . binotata Say both distinct species.

Pall, H. C. 1937. A New Agaporus, Dytiscidae, Coleoptera. Ent. News,
48: 10-12. ( Agaporus latens Fall — Peekskill, 1890 [J. D. Sherman].)

Fisher, W. S. 1940. A New Species of Actenodes. Proc. Ent. Soc. Wash.,
42: (8). ( Actenodes simi Fisher — near N. Y. City, L. I., Yaphank.)

. 1942. A Revision of the North American Species of Buprestid

Beetles Belonging to the Tribe Chrysobothrini. U. S. Dept. Agr. Misc.
Publ. No. 470, pp. 1-274. ( Chrysobothris neopusilla Fisher — Syracuse.

C. adelpha Gemm. and Harold — New York. C. rugosiceps Melsh. — New
York. C. viridiceps Melsh. — New York. C. lecontei Leng a synonym of
C. azurea Lee.)

Green, J. W. 1941. Taxonomic Studies in Cantharis (Coleoptera, Can-
tharidse). Entom. Amer., 20: 159-217. ( Cantharis proximus Green —
L. I., New Lots. C. tantillus Lee. — N. Y. C. nigritulus Lee. a synonym
of C. mandibularis Kby. C. tenuis Green — West Hebron. C. pusillus
(1851) Lee. is a synonym of C. rectus Melsh. C. oriflavus Lee. and C.
imbecillis Lee. are valid species. C. pusillus (1881) Lee. and C. mollis
Fall are synonyms of C. nigriceps Lee. and C. mimus Fall is a subspecies.
C. nigriceps mimus Fall — L. I., Wyandach. C. greeni Fall — N. Y. C.
heterodoxus Green — N. Y.

Hatch, M. H. and Beller, S. 1932. Coleoptera of Washington: Chryso-
melidae. Univ. Wash. Publ. Biol., 1: 93-97. ( Chrysomela to replacc-

Lina and Melasoma.)

Hatch, M. H. 1935. A New Subalpine Genus of Haltieini from North
America. Ent. News, 46: 276-278. ( Crepidodera robusta Lee. sepa-

rated from the genus under name of Orestioides. Here belong also C.
atriventris Kby.)

. 1938. A Bibliographical Catalogue of Injurious Arachnids and

Insects of Washington. Univ. Wash. Publ. Biol., 1: (4), 191. ( Eypera

punctata Fab. becomes Phythonomus zoilus Scop., but this name is not
followed by Winkler [in Cat. Col. Reg. Pal. 13, 1932, p. 1579] who uses
Phytonomus punctatus Fab.)

Heikertinger. 1925. Bestimmungtabelle der Nordamerikanischen Halti-
cinen Gattungen. Kol. Rund., 2: (3-4), 67. ( Crepidodera atriventris

Kby. changed to Ochrosis atriventris Kby.) [See Hatch, 1935.]

Helfer, J. 1941. Revision of the Genus Buprestis of North America North
of Mexico. Entom. Amer., 21: (3), 123-200. ( Buprestis apricornis

Herbst — New York. B. impedita Say a synonym of B. striata Fab.,
while B. consularis Gory is a synonym of B. nutalli Kby.)

Henderson, L. S. 1939. A Revision of the Genus Listronotus. Univ. Kan.
Sci. Bull., 26: (4), 215-320. ( Listronotus callosus Lee. — this is a

June, 1945]

Malkin: Coleoptera

113

southern species and Leconte’s N. Y. citation is a misidentification.
L. incequalipennis Boh. a synonym of L. squamiger Lee. L. tuberosus
Lee. — additional records, Buffalo. L. obliquus Lee., a synonym of L.
sordidus Gyll. L. frontalis Lee. — Additional records, Buffalo, Rochester,
T. B. A. L. latiusculus Boh. a synonym of L. oregonensis Lee. L.
setosus Lee. — New York, N. Y.)

Hincks, W. D. and Dibb, J. R. Junk Cat. Pars 142, Passalidse, p. 16. (Pas-
solus cornutus Fab. changed to Popilvus disjunctus Illig.)

Hinton, H. E. 1935. Can. Ent., 67: (1), 11-18. (Phelister hospes Lee.
separated from the genus under name of

. 1935a. Description of New Neotropical Histeridse with Notes on

Others. Ann. Mag. Nat. Hist., 15: 584-592. (Phelister coquisitus
Lewis is probably a S. A. species and should be removed from the list.)

Hoffman, C. A. 1935. The Biology and Taxonomy of the Genus Trichio-
tinus. Ent. Amer., 15: (4), 133-205. (T. texanus Horn does not occur

in the East and should be omitted. T. assimilis. (Kby.) — Cranberry
Lake, Lake George, Schroon Falls, Buffalo, Rochester, Plattsburg. T.
parvulus Csy. a synonym of T. affinis (G. and P.). T. viridans (Kby.) is
not found in New York.)

Hopping, R. 1937. Can. Dept. Mines Res. Bull. 85, Biol. Serv. 22, Part II,
42 pp. (Stenocorus inquisitor L. to replace Ehagium lineatum Fab.
Parapachyta lacustris Csy. is considered to be a synonym of Centrodera
decolorata Harris. Anthophilax viridipennis Csy. united with A. mala-
chiticus Hald. Evodinus carolinensis Csy. united with E. monticola
Rand.)

Kuntzen, H. 1933. Aus den Verbreitungstatsachen mitgefolgerte neue
Auffassungen uber das System einiger Scarabaeiden-genera vornemlich
Palaarktichen Region. Mitt. Zool. Mus. Berlin, pp. 458-474. (Poly-
phylla occidentalis Eassenkreis variolosa Hentz — New York. According
to Cazier [Ent. News, 51: 134-139, 1940], concept of Rassenkreis cannot
be applied to N. A. species of the genus.)

American species of Chalcoides belong to form Ch. fulvicornis nana
Say, as Ch. helxines L. is questionable. Mr. Gentner believes that speci-
mens placed at present under this name may belong to several groups.

Lacordaire and Boisduval. 1835. Faune Ent. des. env. de Paris, 1835.
p. 613. This publication contains the original description of Agrilus
derasofasciatus Lac., a European form recently introduced in U. S. A.

Leech, H. B. 1940. Description of a New Species of Laccornis, with a
Key to the Nearctic Species (Coleoptera, Dytiscidse). Can. Ent., 72:
122-128. (Generic name of Agaporus changed to Laccornis. Here also
belongs A. latens Fall.)

. 1941. The Species of Matus, a Genus of Carnivorous Water

Beetles. Can. Ent., 73: (4), 77-83. (Matus ovatus Leech — Maiden
Bridge, Peekskill, S. I. — Huguonot.)

Malkin, B. 1941. Bull. Br. Ent. Soc., 36: (1), 28. (Cicindela patruela
Dej. — L. I. — Douglaston [Ch. Ragot]. A new record for Long Island.)

114

Journal New York Entomological Society

[Vol. Lin

. 1941a. Triachus vacuus. Bull. Br. Ent. Soc., 36: (4), 183. (L.

I. — Montauk. )

. 1941b. A European Buprestid in the United States. Bull. Br.

Ent. Soc., 36: (3), 132. ( Agrilus derasofasciatus Lac. — Van Cortlandt

Park.)

. 1941c. Additions to Staten Island List of Coleoptera. Proc. S. I.

Inst. Art. Sci., Oct. 1940-May 1941. Vol. IX, pp. 91-96. (95 species

are listed new to Staten Island. Some of the 'records included in present
list.)

. 1941d. An Addition to the New York State List of Coleoptera

No. 5. Bull. Br. Ent. Soc., 36: 209-212. (81 species are listed, 13 of

which are new additions to the State list. The remainder are new to
Long Island.)

McKenzie, H. 1936. An Anatomical and Systematic Study of the Genus
Anatis of America. Univ. Cal. Publ. Ent., 6: (10), 263-272. (The
variety A. mali Say belongs to the European A. ocellata L. and not to
A. quindecimpunctata.)

Moennich, H. 1941. Supplemental List of Coleoptera Found Living in
and on Various Fungi. Bull. Br. Ent. Soc., 36: (1), 20-24. (Although
this paper contains no new additions to the state, it is of great interest
to N. Y. students, as several rare and rather badly neglected forms are
mentioned, particularly in Stapbylinidce.)

Obenberger, J. 1934. Monografie Rodu Taphrocerus Sole. Sbor. Ent.
Mus. Praze., 12: 5-62. ( Taphrocerus nicolai Obenb.- — New York. The

validity of this species should be verified.)

D’Orchymont, A. 1933. Contribution a 1 ’etude des Palpicornia. Soc.
Ent. Belg., 73: 271-313. ( Hydrophilus occultus d’Orch. — New York.)

Parsons, C. T. 1938. Notes on North American Nitidulidse. II.
Cryptarcha Shuckard. Psyche, 45: 96-100. ( Cryptarcha strigatula

Parsons — Bronx, 1896 (Holotype) .)

* . 1943. A Revision of Nearctic Nitidulidae. Bull. Mus. Comp. Zool.,

92: (3), 121-278. ( Boreades a generic name to replace Cercometes.

Brachypterolus mdrdelloides Notm. a synonym of B. pulicarius L.
Amartus rufipes Lee. records cited are dubious since the species is a
western one. Carpophilus pallipennis Say — N. Y. C. melanopterus
Er. — Rye. C. sayi Parsons — paratypes from N. Y. C. antiquus Melsh.
— Flatbush, an additional record new to L. I.(?). Epurcea umbrosa
Horn — Lake George. Omosita discoidea (Fab.) — New York. Lobiopa
setosa Har. — S. I. Pallodes pallidus Beau, to replace P. silaceus Er.
Glischrochilus obtusus (Say) — Hillburn, also L. I., additional records.
G. sanguinolentus rubromaculatus Reitt. — Buffalo. G. siepmanni Brown
— S. I. Cybocephalus nigritulus Lee. listed from L. I. by Cooper (1935)
apparently not included under Nitidulidce by Parsons. A number of
species may well be expected within N. Y. boundaries, having been
recorded in adjacent states both to the north and south, particularly the
following: E. fulvescens Horn, E. depressa 111., Nitidula flavomacula

June, 1945]

Malkin: Coleoptera

115

Rossi, Meligethes ceneus Fab., Glischrochilus quadrisignatus quadrisig-
natus (Say), and G. q. canadensis Brown.)

Pechuman, L. L. 1937. An Annotated List of Insects Found in the
Bark and Wood of Ulmus Americana L. in New York State. Bull. Br.
Ent. Soc., 32: (1), 8-21. (Coleop. 8-14. Mr. Pechuman overlooked
Cooper’s addition and thus duplicated Scolytus multistriatus Marsh.,
as new to New York.)

Rau, G. J. 1935. A New Variety of Anoplodera Vittata from New York.
Bull. Br. Ent. Soc., 30: (2), 63-64. ( Anoplodera vittata saratogensis

Rau — Saratoga Springs (type).)

Reinhardt, A. 1926. tiber die mit Pachylopus Er. Verwandenten Gat-
tungen. Ent. Blatt., 22: 14. (A genus Bacmceniolus erected to which
belongs Saprinus palmatus Say of the list.)

Robinson, M. 1940. Studies in the Scarabseidse No. 2. Trans. Am. Ent.
Soc., 66: 141-159. ( Trox foveicollis Harold replaces T. insularis Chev.,
the latter probably being not a U. S. species. Trox hamata n. sp. —
Ithaca, Flushing, N. Y. C.)

Ross, E. S. 1940. A Preliminary Review of the North American Species
of Dendrophilus (Coleoptera, Histeridae). Bull. Br. Ent. Soc., 35: (3),
103-108. ( Dendrophilus punctulatus Say united with D. sexstriatus

Hatch and European D. punctatus Herbst., the latter being the proper
one. Among examined material, specimens from Long Island and Rock-
away Beach. This establishes a new record for this locality.)

Sanderson, M. W. 1938. Species of Stenelmis. Univ. Kans. Sci. Bull., 25 :
(22), 637. ( Stenelmis concinna Sand. — North River (type) (Sander-

son). Stenelmis tarsalis Sand. Batavia (Knight) 1915. S. mera Sand.
East Homer (Sanderson), Ithaca, North River (Sanderson). S mus-
gravei Sand. — Batavia (Knight),, this is probably the species recorded
by Knight in the New York State List as new.)

Schoof, H. F. 1942. The Genus Conotrachelus in the North-Central U. S.
111. Biol. Mon., 19: (3), 170 pp. ( Conotrachelus tibialis Schoof prob-
ably occurs in New York. C. erinaceus Lee. — New York. C. carolinensis
Schoof — New York City and vicinity.)

Soraci, F. A. 1940. Distribution in New Jersey of Myllocerus Castaneus.
Jl. N. Y. Ent. Soc., 48: (4), 318. (The author mentions that the species
has been found on Long Island by Mr. R. I. Clement, and causes con-
siderable damage to shrub oaks, feeding on the foliage.)

Tanner, V. M. 1943. A Study of the Sub tribe Hydronomini with a De-
scription of New Species. Study VI. Publ. Dept. Zool. Ent. Brigh.
Young Univ., Provo, Utah, No. 1 and 2, pp. 1-38. ( Bagous longi-

rostris Tanner — Buffalo. B. americanus Lee. — Buffalo, Peekskill,
Esopus, Olcott, vie. of N. Y. C. are additional records. B. blanchardi
Blatch. — Lake Oscawana. B. bicarinatus Blatch. — Penn Yan. B. nebu-
losus Lee. — New York. B. floridanus Tann. — Ithaca. B. magister Lee.
— Penn Yan, an additional record. B. planatus Lee. — Olcott, Penn Yan,
Ithaca as above. Several other species may also be found within the

116

Journal New York Entomological Society

[Vol. LIII

state, most likely B. pusillus Lee., B. cavifrons Lee., and B. transversus
Lee.)

Timberlake, P. H. 1943. The Coccinellidce or Lady Beetles of Koebele
Collection — Part I. Bull. Exp. Sta. Haw. Sug. Plant. Ass. Ent. Ser.
Bull. no. 22. Coleomegilla maculata lengi n. sp. Rochester. This name
replaces Ceratomegilla fuscilabris in part. Also species of American
Neomysia regarded as races of the Palearctic A. oblonguttata.)

Van Dyke, E. C. 1936. A Review of the Subgenus Nomaretus Leconte
of the Genus Scaphinotus Dejean. Bull. Br. Ent. Soc., 31: (1), 37-43.
( Nomaretus degraded to subgeneric rank.)

. 1937. Notes and Descriptions of North American Buprestidse and

Cerambycidse. Bull. Br. Ent. Soc., 32: (3), 105-116. ( Xylotrechus

frosti Van Dyke — Ithaca, 1917 [Van Dyke].)

. 1938. Bull. Br. Ent. Soc., 33: 45-58. ( Chrysolina [not Chryso-

mela ] stibopaca Rogers — Peekskill. This verifies Rogers’ record from
the state.)

V oris, R. 1936. The Rapid Spread of an European Staphylinid in North
America. Ann. Ent. Soc. Amer., 29: (1), 78-80. (Additional records
of Philonthus cruentatus Gmel. — Rochester, Onendango Co.)

Wallis, J. B. 1928. Review of the Genus Odontaeus Dej. Can. Ent., 60:
168-176. ( Odontceus simi — New York. 0. liebecki — New York. 0.
darlingtoni — New York.)

. 1939. Hydaticus modestus Sharp versus Hydaticus stagnalis Fab-

ricius in North America. Can. Ent., 71: 126-127. ( Hydaticus cincti-

pennis Aube and H. modestus Sharp are distinct from E. stagnalis Fab.
which does not occur in this country. Both species should be omitted
from the list and replaced by E. modestus Shp. Likewise, E. Icevipennis
Thom, must be dropped and superseded by Sharp ’s species. This opinion
is also supported by Mr. Leech.)

. 1939a. The Genus Graphoderes Aube in North America. Can.

Ent., 71: 123-131. ( Graphoderes cinereus L. is the same as G. fasciati -

collis (Harr.), while G . elatus L. and G. sonatus Hoppe are synonyms
of G. perplexus Shp.)

Wenzel, R. L. 1936. Can. Ent., 68: (2), 266-272. ( Bister unicus Csy.

a synonym of E. cognatus Lee.)

. 1939. Ohio Jl. Sci., 39: 10-14. ( Bister immunis Er. is the same

as E. interruptus Beauv. Saprinus semistriatus Scriba same as S.
lecontei Csy.)

Wolcott, G. N. 1937. E col. Mon., 7:43-55. (Coleoptera). ( Cryptobium
clavicorne Csy. and Philonthus inquietus Er. new to the list. Tachinus
fumipennis Say new to ‘ 1 Continental ’ ’ New York. Nematolinus longi-
collis Lee., Philonthus thoracicus Grav., and Decarthron longulum Lee.
are additional records.)

June, 1945]

Rapp & Cooper: Psychodid^e

117

CHECK-LIST OF THE PSYCHODID^ OF EUROPE1

By William F. Rapp, Jr., and Janet L. Cooper

The area covered by this check-list includes all Europe, Great
Britain, Russia and Asia Minor.

FLEBOTOMUS Rondani

africanus var. asiaticus Theodor, Bull. Ent. Res., Vol. 24, p. 541.
Palestine.

ariasi Tonnoir, Ann. Soc. ent. Belg., Yol. 61, p. 53.

Barcelona, Spain.

borowskii Khodukin, in Khodukin and Sofiev, Meditz. Muisl’
Uzbekist., Vol. 5, p. 51.

Turkmenistan.

bruchoni Parrot, Arch. Inst. Pasteur Algerie, Vol. 13, p. 252.
Greece.

canaaniticus Adler and Theodor, Proc. Roy. Soc., Series B, Vol.
108, p. 468.

Palestine.

chinensis Adler and Theodor, Bull. Ent. Res., Vol. 21, p. 534.
Persia.

chinensis var. simici Nitzulescu, Ann. Parasit. hum. comp., Vol.
9, pp. 129, 265.

Jugoslavia, Palestine.

chinensis var. longiductus Nitzulescu, Ann. Parasit. hum. comp.,
Vol. 9, p. 128.

Jugoslavia, Palestine.

clydei Sinton, Ind. J. Med. Res., Vol. 15, p. 179.

Afghanistan.

grassii Pierantoni, Bull. Mus. Zool. Torino, Vol. 39 (1924), N. S.
21, p. 5.

Italy.

grekovi Khodukin, Meditz. Muisl’ Uzbekist. Turkmenist. suppl.
1929, p. 101.

Russia.

i Third contribution to a Check-list of Psychodidse of the World.

118

Journal New York Entomological Society

t V ol. LIII

kandelakii Schourenkova, Russ. J. Trop. Med., Vol. 7 (1929), p.
693.

Russia.

larroussei Langeron and Nitzulescu, Ann. Parasit. hum. comp.,
Vol. 9, p. 73.

France.

macedonicus Adler and Theodor, Proc. Roy. Soc., Series B, Vol.
108, p. 468.

Macedonia.

major var. longiductus Parrot, Arch. Inst. Pasteur Algerie, Vol.
6, p. 29.

Turkastand.

major var. syriacus Adler and Theodor, Proc. Roy. Soc., Series
B, Vol. 108, p. 467.

Syria.

mascittii Grassi, Roma Rend. Acc. Lincei, Vol. 17, pp. 681-682.
Italy.

minutus Rondani, Ann. Soc. Ent. France, Vol. 1, ser. 2 (1843),
p. 2 tab.

Mediterranean Region.

minutus var. arpaklensis Perfil’ev, Zool. Anz., Vol. 101, p. 226.
Turkmenistan.

minutus var. meridionalis Pierantoni, Bull. Mus. Zool. Torino,
Vol. 39 (1924), N. S. 21, p. 3.

Italy.

minutus var. sog dianus Parrot, Arch. Inst. Pasteur Algerie, Vol.

6, p. 26.

Turkastand.

neglectus Tonnoir, Ann. Soc. Ent. Belg., Vol. 61, p. 333.

Albania.

newsteadi Sinton, Ind. J. Med. Res., Vol. 15, p. 589.

Afghanistan.

nigerrimus Newstead, Bull. Ent. Res., Vol. 2, p. 68.

Malta.

parroti var . it aliens Adler and Theodor, Bull. Ent. Res., Vol. 22,
p. 107.

Italy.

parroti var. sardous Bogliolo, Ann. Igiene, Vol. 45, p. 42.
Sardinia.

June, 1945]

Rapp & Cooper: Psychodid^e

119

pawlowskyi PerfiPev, Zool. Anz., Vol. 101, p. 222.

Turkmenistan.

perfilievi Parrot, Arch. Inst. Pasteur Algerie, Yol. 8 (1930), p.
383.

Crimea.

perfilievi var. transcaucasicus PerfiPev, Ins. U.S.S.R., Dipt., Yol.
3, No. 2, p. 108.

Transcaucasia.

perniciosus Newstead, Bull. Ent. Res., Yol. 2, p. 70.

Malta.

perniciosus var. nitzulescui Simic, Ann. Parasit. hum. comp., Yol.
10, p. 434.

J ugoslavia.

perniciosus var. tauricus Nasonov., Acad. Sci. U.S.S.R., No. 22
(1927), p. 370.

South Crimea.

perniciosus var. tobbi Adler and Theodor, Bull. Ent. Res., Yol.
21, p. 536.

Persia.

selectus Khodukin, Meditz. MuisP Uzbekist. Turkmenist. suppl.
1929, p. 99.

Russia.

sogdianus Parrot, Arch. Inst. Pasteur Algerie, Yol. 7, p. 303.
Bokhara.

stalinabadi Khodukin, in Khodukin and Sofiev, Meditz. MuisP
Turkmenist., Yol. 5 (9-10), p. 53.

Central Asia.

sumbaricus PerfiPev, Zool. Anz., Yol. 101, p. 225.

Turkmenistan.

tiberiadis Adler, Theodor, and Lowrie, Bull. Ent. Res., Vol. 21,
p. 537.

Palestine.

troglodytes Nitzulescu, Ann. Parasit., Yol. 8, p. 388.

France.

vesuvianus Adler and Theodor, Bull. Ent. Res., Vol. 22, p. 108.
Italy.

wenyoni Adler and Theodor, Bull. Ent. Res., Yol. 21, p. 535.
Persia.

120

Journal New York Entomological Society

[Vol. LIU

LEPIMORIA Enderlein

tatrica Enderlein, Dtsch. ent. Z., 1936 (1937), p. 98.

High Tatras (Polish-Czechoslovakia border).
galicica Enderlein, Dtsch. ent. Z., 1936 (1937), p. 93.

Poland.

latefasciata Enderlein, Dtsch. ent. Z., 1936 (1937), p. 93.

Italy.

MICRODIXA Muller

scutigera Muller, Z. Morph. Oekol. Tiere, Vol. 7, p. 535.
Switzerland.

similis Muller, Z. Morph. Oekol. Tiere, Vol. 7, p. 541.

Germany.

PANIMERUS Tonnoir

maynei Tonnoir, Ann. Soc. ent. Belg., Vol. 60, p. 186.

Belgium.

PERICOMA Walker

albomaculata Wahlgrem, Arkiv. Zool., Vol. 2, p. 16.

Sweden.

ambigua Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX)
(1893), p. 126.

Northern Europe.

andrenipes Strobl, Milt. Ver. Steierm, Vol. 46 (1910), p. 270.
Austria.

advena Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX)
(1893), p. 127.

Northern Europe.

angustipennis Tonnoir, Ann. Soc. ent. Belg., Vol. 60, p. 184.
Belgium.

auriculata Halid, in Curt, : Brit. Ent., 745, Tab., fig. 9 (Psychoda)
( canescens Walk., Ins. Britannica, Dipt., Vol. 3, p. 254, obs.
(1856)) ( fusca Walker (nec. Macq.) Ins. Britannica, Dipt.,
Vol. 3, p. 260).

Northern Europe.

avicularia Tonnoir, Trans. Soc. Brit, Ent., Vol. 7 (1940), p. 37.
England.

blandula Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX)
(1893), p. 123.

Northern Europe, Great Britain.

June, 1945]

Rapp & Cooper: Psychodid^e

121

calcilega Feuerb., Verhand Intern. Ver. f. Linn., 1923, p. 196.
Great Britain.

caliginosa Eaton, Ent. Monthly Mag., ser. 2, Vol. IX (XXIX),

p. 128.

Northern Europe.

canescens Meigen, Klass, Vol. 7 (1804), Vol. 5 (Trichoptera),

p. 45.

Europe, Great Britain.

cognata, Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX)
(1893), p. 121.

Northern Europe, Great Britain.
compta, Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX)
(1893), p. 121.

Northern Europe, Great Britain.
consors Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX)
(1893), p. 127.

Northern Europe.

dalii Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX) (1893),
p. 125.

Northern Europe.

decipiens Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX)
(1893), p. 126.

Northern Europe.

diver sa, Tonnoir, Ann. Soc. ent. Belg., Vol. 60, p. 182.

Belgium.

exquisita Eaton, Ent. Monthly Mag., ser. 2, Vol. 4 (24) (1893),
p. 123.

Great Britain.

extricata Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX)
(1893), p. 121.

Northern Europe, Great Britain.
fallax Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX) (1893),

p. 122.

Northern Europe, Great Britain.
fratercula Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX)
(1893), p. 128.

Northern Europe.

fused Macquaert, Recueil Soc. Sc. Agr. Lille, Vol. 4 (Psychoda)
(1826), p. 167 (calceata Meigen, Syst. Besehr., Vol. 11 (1830)

122

Journal New York Entomological Society

[Vol. Lin

(Psychoda), p. 272) ( tristis Zedderstett {nee. Meigen), Dipt.
Scand., Vol. XII (1855), 6-7 (Psychoda), p. 4887).

Europe, Great Britain.

gracilis Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX)
(1893), p. 123.

Northern Europe, Great Britain.
hibernica Tonnoir, Trans. Soc. Brit. Ent., Vol. 7 (1940), p. 37.
Ireland.

hirticornis Tonnoir, Ann. Soc. ent. Belg., Vol. 62, p. 158.

Austria.

incerta Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX)
(1893), p. 129.

Northern Europe.

labeculosa Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX)
(1893), p. 127.

Northern Europe.

magnicornis v. d. Wulp, Diptera Neerlandica, Vol. I (1877), p.
319.

Central Europe.

manicata Tonnoir, Ann. Soc. ent. Belg., Vol. 60, p. 181.

Belgium.

modesta Tonnoir, Ann. Soc. ent. Belg., Vol. 62, p. 165.

France.

morula Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX)
(1893), p. 127.

Northern Europe.

mutua Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX)
(1893), p. 121.

Northern Europe, Great Britain.
neglecta Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX)
(1893), p. 122.

Northern Europe, Great Britain.
notabilis Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX)
(1893), p. 126 ( canescens Walker, Ins; Britannica, Dipt.,
Vol. 3 (1856), p. 258).

Northern Europe.

nubila Meigen, Syst. Beschr., Vol. 1 (1818) (Psychoda), p. 107
( bombyciformis Schrnk., Fauna Boica, Vol. Ill (Tipula), p.
2347).

Northern Europe, Great Britain.

June, 1945]

Rapp & Cooper: Psychodkue

123

ocellaris Meigen, Klass., Vol. I, tab. II, fig. 19 (Trichoptera)
(1804), p. 44 ( variegata Macquaert, Recueil Soc. Sc. Agr.
Lille, Vol. 2, p. 167 (Psychoda) (1826)).

Europe.

opaca Tonnoir, Ann. Soc. ent. Belg., Yol. 62, p. 154.

Italy.

ornata Tonnoir, Ann. Soc. ent. Belg., Vol. 62, p. 157.

Italy.

palustris Meigen, Klass., Yol. I (Trichoptera) (1804), p. 43.
Europe.

pilularia Tonnoir, Trans. Soc. Brit. Ent., Vol. 7 (1940), p. 38.
England.

plumicornis Tonnoir, Ann. Soc. ent. Belg., Vol. 62, p. 161.
Austria.

pseudequisita Tonnoir, Trans. Soc. Brit. Ent., Vol. 7 (1940),
p. 36.

Great Britain.

pulchra Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX)
(1893), p. 123.

Northern Europe, Great Britain.
revisenda Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX)
(1893), p. 129.

Northern Europe.

soleata Walker, Ins. Britannica, Dipt., Vol. 3 (1856), p. 257.
Northern Europe.

spherica Tonnoir, Ann. Soc. ent. Belg., Vol. 60, p. 180.

Belgium.

subneglecta Tonnoir, Ann. Soc. ent. Belg., Vol. 62, p. 169.
Belgium.

squamigera Tonnoir, Ann. Soc. ent. Belg., Vol. 62, p. 163.
Austria.

trifasciata Meigen, Klass, Vol. I, tab. II, fig. 20 (Trichoptera)
(1804), p. 44.

Northern and Central Europe, Great Britain.
tristis Meigen, Syst. Beschr., Vol. VI, 10 (Psychoda) (1830), p.
272.

Europe.

irwialis* Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX)
(1893), p. 121.

Northern Europe.

124

Journal New York Entomological Society

t Vol. LIII

ustulata Walker, Ins. Britannica Dipt., Vol. 3 (1856), p. 258.
Northern Europe.

PERIPSYCHODA Enderlein

nigritarsis Enderlein, Dtsch. ent. Z., 1936 (1937), p. 99.

Germany.

PSYCHODA Latreille

albipennis Zedderstett, Dipt. Scand., Vol. IX (10) (1850), p.
3708.

Europe.

alternata Say, Long’s Exp. St. Peter’s River App. (1824). (For
synonyms see Journal New York Entomological Society, Vol.
52, p. 204.)

Great Britain.

brevicornis Tonnoir, Trans. Soc. Brit. Ent., Vol. 7 (1940), p. 54.
England.

crassly enis Tonnoir, Trans. Soc. Brit. Ent., Vol. 7 (1940), p. 52.
England, Germany.

erminea Eaton, Ent. Monthly Mag., ser. 2, Vol. IV (XXIX)
(1893), p. 130.

Northern Europe.

flavescens v. Ros., Correspondenzbl. Wurttemberg. landw., Ver.
I, 50 (1840), p. 50.

Central Europe.

gemina Eaton, Ent. Monthly Mag., Vol. 40 (1904), p. 57.
England, Belgium.

grisescens Tonnoir, Ann. Soc. ent. Belg., Vol. 62 (1922), p. 87.
England.

Immeralis Meigen, Syst. Beschr., Vol. I (1818), 7, p. 106 ( bullata
Walker, Ins. Britannica, Dipt., Vol. Ill (3), p. 257 (Peri-
coma) (1856)).

Europe.

lobata Tonnoir, Trans. Soc. Brit. Ent,, Vol. 7 (1940), p. 60.
England, Bulgaria.

lucifuga Walker, Ins. Britannica, Dipt., Vol. Ill, 1 (Pericoma)
(1856), p. 257.

Northern Europe.

obscura Tonnoir, Ann. Soc. ent. Belg., Vol. 50 (1919), p. 140.
England, Europe.

June, 1945]

Rapp & Cooper: Psychodid.®

125

phalcenoides Linnseus, Syst. Nat., Ed. X, 32 (Tipula) (1758),
p. 588 ( muraria Latreille, Hist. Nat. d. Crust, et. Ins., Yol.
XIV (1805), p. 203) ( nervosa Schrnk., Fauna Boica, Vol.
III, 82 (Tipula) (1803), p. 2350).

Europe.

phalcenoides sub. sp. elongata Tonnoir, Trans. Soc. Brit. Ent.
Vol. 7 (1940), p. 51.

England.

primceva Cockerell, U. S. Mus., Vol. 49, p. 493.

England.

setigera Tonnoir, Ann. Soc. ent. Belg., Vol. 62 (1922), p. 85.
England, Belgium, Germany.

severini Tonnoir, Ann. Soc. ent. Belg., Vol. 62 (1922), p. 78.

England, Belgium, Austria, Germany.
spreta Tonnoir, Trans. Soc. Brit. Ent., Vol. 7 (1940), p. 57.
England, Belgium.

silacea Curt., Brit. Ent., Vol. 10 (1839), p. 745.

Northern Europe.

TELMATOSCOPUS Eaton

apicealbus Tonnoir, Ann. Soc. ent. Belg., Vol. 62, p. 179.

Italy.

britteni Tonnoir, Trans. Soc. Brit. Ent., Vol. 7 (1940), p. 47.
England.

eatoni Tonnoir, Trans. Soc. Brit. Ent., Vol. 7 (1940), p. 47.
England.

furvus Tonnoir, Trans. Soc. British Ent., Vol. 7 (1940), p. 49.
Belgium.

rothschildii Eaton, Ent. Mag., Vol. 48, p. 9.

England.

similis Tonnoir, Ann. Soc. ent. Belg., Vol. 62, p. 171.

Holland.

THRETICUS Eaton

compar, Ent. Mag., Vol. XL, p. 57.

Britain, Madeira.

gemina Eaton, Ent, Mag., Vol. XL, p. 57.

England.

126

Journal New York Entomological Society

[Vol. LIII

TRICHOMYIA Haliday

urbica Curt., Brit. Ent., Vol. 9 (1839), p. 745 ( aurea Zett., Dipt.
Scand., Vol. IX (8) (Psychoda) (1850), p. 3707).

Europe.

ULOMYIA Walker

plumata Tonnoir, Ann. Soc. ent. Belg., Vol. 60, p. 187.

Belgium.

(Jour. N. Y. Ent. Soc.) Vol. LIII

(Plate I)

WILLIAM T. DAVIS
(Courtesy of Edwin Way Teale)

litis

June, 1945J

Weiss: Davis

127

WILLIAM THOMPSON DAVIS— 1862-1945

On January 22, 1945, the New York Entomological Society lost
by death its honorary president, William T. Davis, who was one
of its oldest and most distinguished members. Mr. Davis died in
the Staten Island Hospital after a long illness following an opera-
tion in July, 1944. He was in his eighty- third year. Mr. Davis
was born in New Brighton, Staten Island, October 12, 1862. His
parents were George B. and Elizabeth (Thompson) Davis, and his
forebears, at various times, took leading parts in the civic prog-
ress of the Island. After an education in private schools on the
Island he became a clerk in a New York mercantile house.
Shortly afterward he entered the New York Produce Exchange
of lower Manhattan, in the Gratuity Fund Department, where he
stayed for over 26 years, having complete charge of that depart-
ment during the three years previous to his resignation in 1909.
His work in the Exchange was tedious and exacting, involving,
as it did, the keeping in order of more than 3,000 individual
accounts of the members of the Exchange.

Long interested in natural history, Mr. Davis, Mr. Charles W.
Leng and others formed the Natural Sciences Association of
Staten Island at a meeting on November 12, 1881, held in the
home of his maternal grandfather, John C. Thompson, in St.
Mark’s Place at the corner of Hyatt Street. Mr. Davis was only
nineteen years old at the time and Mr. Leng was twenty-one. The
Association was incorporated February 19, 1885, and in May,
1905, the name was changed to the Association of Arts and Sci-
ences. Its present name, Staten Island Institute of Arts and
Sciences was adopted May 18, 1918. Mr. Davis was vice-president
from 1905 to 1929, president from 1930 to 1934, and president
emeritus from 1934 until his death.

After his retirement from business Mr. Davis occupied himself
exclusively with natural history and other interests close to his
heart. Entomology was one of these and probably the most
important. Certainly, it occupied his attention continuously
throughout most of his long life. Mr. Davis was one of the very
few, good, general naturalists in addition to his specialization in

128

Journal New York Entomological Society

[Vol. LIII

entomology. Over the years he thoroughly explored the fauna
and flora of Staten Island and he was capable, at the same time,
of identifying much of his material. Most of his life was spent
on Staten Island, interrupted occasionally by collecting trips to
Long Island, Gardiner’s Island, up along the Hudson River in
New York, to the Pine Barrens and southern counties of New
Jersey, and to California, Arizona, Texas, Florida, Georgia, the
Carolinas and Virginia. In ‘ ‘ A List of the Insects of New York, ’ 9
edited by Dr. M. D. Leonard and published by the Cornell Uni-
versity Agricultural Experiment Station in 1928, will be found
most of his Staten Island and New York records. His New Jersey
records were used by Dr. John B. Smith in “Insects of New
Jersey,” Trenton, 1909, wherein his ability and thoroughness
were recognized. And his observations and descriptions of new
species of dragonflies were utilized in Dr. James G. Needham’s
“Handbook of the Dragonflies of North America,” Baltimore,
1929.

In the prefaces to many works on North American insects will
be found statements of the authors acknowledging their indebted-
ness to Mr. Davis for his large fund of entomological knowledge
and for his help in supplying specimens. Examples of rare spe-
cies were either loaned or given to students as well as notes on
distribution and habits. In the introduction to “Orthoptera of
Northeastern America,” Indianapolis, 1920, Dr. W. S. Blatchley
said, “Mr. Davis, who is not only a student of Orthoptera, but
also a naturalist of high rank and interested in all forms of living
things, has aided me greatly in many ways. ’ ’ The same author,
in his “Heteroptera or True Bugs of Eastern North America,”
Indianapolis, 1926, wrote — “As in my works on Coleoptera and
Orthoptera, so in this instance, my friend and fellow naturalist,
William T. Davis, of Staten Island, N. Y., has been of great
service. Anything I wanted which he could furnish in the way
of specimens or literature, he gladly sent, and he also furnished
many notes on distribution and habits. ’ ’ Such happenings were
a daily part of Mr. Davis’s life.

In addition to helping others Mr. Davis wrote extensively upon
entomological topics. Although most of his contributions ap-
peared in the pages of the Bulletin of the Brooklyn Entomological
Society and the Journal of the New York Entomological

June, 1945]

Weiss: Davis

129

Society, other technical journals carried some of his articles. In
the bibliography of the ‘ ‘ Alphabetical Index to North American
Orthoptera, ’ ’ by Samuel H. Scudder (Bost. Soc. Nat. Hist.,
1901), there are eight references to early papers by Mr. Davis on
the Locustidse. These appeared from 1886 to 1896 in the Ca-
nadian Entomologist, Proceedings of the Entomological Society
of Ontario, American Naturalist, Entomologica Americana and
the Proceedings of the Natural Sciences Association of Staten
Island.

In the 52 volumes of the Journal of the New York Entomo-
logical Society, from 1893 to 1944, there are 132 titles by Mr.
Davis. His first paper in Volume I, 1893, was a “ Catalogue of
the Butterflies of Staten Island,” and his last paper in Volume
LII, 1944, was “The Remarkable Distribution of an American
Cicada: A New Genus and Other Cicada Notes.” Between the
publications of these two papers, 130 notes and lengthy articles
appeared on cicadas, water beetles, dragonflies, butterflies, crick-
ets, local lists of insects, ants, tiger beetles, pink katydids, aphids,
roaches, biographies of entomologists, collecting notes, book re-
views, etc., and in addition the printed Proceedings of the Society
abound with notes and observations by Mr. Davis.

Most of his lengthy papers in the Journal, of which there are
about 35, are concerned with descriptions of new species and
varieties of cicadas from western and southwestern states and
from Mexico, Porto Rico and the West Indies. About 155 new
species and varieties of cicadas were described in the pages of the
Journal together with notes on their songs, distribution and
habits. Other papers, that I know of, on cicadas appeared be-
tween 1891 and 1928 in the Canadian Entomologist, Proceedings
of the Staten Island Institute of Arts and Sciences, American
Naturalist, Entomological News, Bulletin of the Brooklyn Ento-
mological Society and the Proceedings of the Staten Island Natu-
ral Sciences Association. In addition Mr. Davis prepared the
lists of Cicadidas that appear in W. E. Britton’s “Guide to the
Insects of Connecticut, Part IV, Hemiptera (Conn. State Geol.
& Nat. Hist. Survey Bull. 34, 1922) and in M. D. Leonard’s “A
List of the Insects of New York (Cornell Univ. Agric. Exp. Sta.
Mem. 101, 1928), and he also wrote a bulletin on “The Cicadas

130

Journal New York Entomological Society

[Vol. Lin

or Harvest Flies of New Jersey’ ’ (Circ. 97, N. J. Dept. Agric.,
1926).

Mr. Davis was the leading authority on the cicadas of North
America and numerous friends and correspondents all over the
country sent him specimens and notes. Without attempting to
mention all, some of these included S. J. Hunter of Kansas ;
T. D. A. Cockerell of Colorado; Harry H. Knight and Joseph
Bequaert, who were members of the Cornell University Biological
Expedition, organized by J. Chester Bradley in 1917, which ex-
tended from New York to California; Colonel Wirt Robinson of
West Point and Wingina, Va. ; Louise Knobel of Arkansas • E. D.
Ball of Arizona; Douglas K. Duncan and 0. C. Poling who sent
specimens from Arizona; R. W. Harned of Mississippi; W. S.
Wright of California ; R. H. Beamer, who supplied specimens
from New Mexico and Oklahoma ; Alonzo C. Davis of California ;
J. W. Sugden of Utah ; F. F. Bibby of Texas ; Ernest L. Bell, who
sent cicadas from Nevada; and H. Garman of Kentucky. Mr.
Geo. P. Engelhardt during his travels supplied cicadas from
Texas and other places. Howard H. Cleaves also, during his vari-
ous travels, collected many specimens for Mr. Davis, his lifelong
friend. Mr. Davis corresponded with W. E. China of the British
Museum and extensively with E. P. Van Duzee, especially during
the early part of the period in which he was describing new spe-
cies. M. D. Leonard sent him specimens from Spain in 1924.
J. N. Knull supplied specimens from Texas, Arizona and Cali-
fornia during the course of his travels. And in 1922, Joseph
Bequaert took specimens for Mr. Davis to the British Museum for
comparison with Walker’s types. Such activities and exchange
of correspondence took place continually between Mr. Davis and
his friends and such occurrences as I have noted could be ex-
tended over several pages.

In “Insect Singers, a Natural History of the Cicadas,” London,
1929, by J. G. Myers, wherein many of Mr. Davis’s observations
are recorded and which contains a bibliography of his cicada
papers from 1891 to 1906, there is the following statement on
page 82. “That happy walking hand-in-hand of ethology and
taxonomy which has made myrmecology a model for systematic
entomology was never achieved in any degree in tettigology till

June, 1945]

Weiss: Davis

131

Wm. T. Davis brought to bear his very wide field experience.”
In the Staten Island Museum there is Mr. Davis’s extensive col-
lection of Cicadidas and other insects as well as large series of
Coleoptera collected by Charles W. Leng and Mr. Davis.

Edwin Way Teale in his book “Near Horizons, The Story of an
Insect Garden,” New York, 1942, has a chapter entitled “Cicada
Man,” which is an appreciative account of Mr. Davis and his
interest in cicadas interwoven with personal anecdotes about Mr.
Davis, his collecting paraphernalia and habits, his workshop in
the “attic” of the Staten Island Museum with its rattlesnake
skins, Indian skull, boxes of insects, notes, clippings, etc., his
kindliness, his dry humor, and entertaining remarks, his encour-
agement and help to young naturalists all written when Mr. Davis
was close to 80 years of age.

The October, 1942, issue of the Bulletin of the Brooklyn Ento-
mological Society was dedicated to Mr. Davis upon his eightieth
birthday and contains articles by J. R. de la Torre-Bueno,
Edwin Way Teale, Howard Cleaves, and J. Bequaert, all dealing
with different phases of Mr. Davis’s life activities, and person-
ality and illustrated by early and late photographs of Mr. Davis
in action as an entomologist. It is a temptation to quote exten-
sively from this publication and only its comparatively recent
appearance prevents this. From these articles one gets a very
good idea of William T. Davis himself, his qq,iet unassuming
manner, his kindliness and consideration of others, his philoso-
phy, his placidity, his admiring friends, his wide knowledge of
plants and animals, his close association with lifelong friends, his
material generosity to individuals and to natural history institu-
tions, his field trips, his absorbing interest in Staten Island his-
tory and the Staten Island Institute of Arts and Sciences, his
versatility, the endless contents of his pockets and his energy, for
a small man who was, as he said of himself “so poorly put
together. ’ ’

My own friendship with Mr. Davis extended over a period of
25 years. It was the custom of many years for Mr. Davis and
Mr. Leng to arrive at the American Museum of Natural History
late in the afternoon, preceding the evening meeting of the New
York Entomological Society and to gather in Mr. A. J. Mutchler’s

132

Journal New York Entomological Society

[Vol. liii

office for a discussion of entomological matters. Here I used to
enjoy their reminiscences and conversations about old-time ento-
mologists, whom I knew only by reputation. Mr. Davis was fre-
quently burdened by large packages of boxes of cicadas which he
brought along to illustrate his frequently scheduled talks before
the Society and he could always be depended upon for a talk when
the program committee ran short of speakers. My collecting
experiences with Mr. Davis involved only a few trips to Lake-
hurst, N. J., at which time he was interested particularly in
Orthoptera. I recall several pleasant visits to his home on
Stuyvesant Place and to the Staten Island Museum while it was
growing up and where I was entertained by Mr. Davis in the
“attic” where he held forth. After the close of the meetings of
the New York Entomological Society I frequently rode downtown
partway on the subway with Mr. Davis and I used to marvel at
his agility when he was close to 80 and at his continued energy
and varied interests. During my editorship of the Journal Mr.
Davis published many cicada papers therein, all written carefully
in longhand, all requiring little or no editing and all legible to the
printer. His papers were frequently accompanied by checks for
substantial sums to help toward the cost of printing.

At the age of 30, Mr. Davis wrote “Days Afield on Staten
Island” (Published under auspices of the Staten Island Histori-
cal Society, 1892) and a second edition appeared in 1937. This
is a series of records of rambles, over Staten Island made in com-
pany with Charles W. Leng, or Louis P. Gratacap or, that Mr.
Davis made by himself. They are really prose essays dealing with
such subjects as spring in the Clove Valley, South Beach, a de-
serted farm house, native brooks, etc., interwoven with observa-
tions on natural history, trees, insects, plants, picturesque people,
bits of local history, associations, and the author’s own thoughts.
Mostly the book is about the activity of nature that goes on out-
doors, of which so many persons are uilaware. Mr. Davis happily
discovered at an early age the pleasure that may be derived from
making discoveries in natural history and the fact that more joy
may be extracted from such activities than from an exclusive
interest in the acquisition of material possessions. The book is as
pleasing to read now as when it was first written and in it is firmly

4

June, 1945] Weiss: Davis 133

embedded the personality of Mr. Davis which never changed dur-
ing his lifetime. More than any other of his writings this book
reflects the man himself. Mr. Davis occasionally put his thoughts
into verse and several of his poems are printed in this book.

Mr. Davis served the New York Entomological Society long and
faithfully in many capacities. He was treasurer from 1904 to
1928, president during 1929 and 1930 and honorary president
from 1940 until his death. The same can be said of the Brooklyn
Entomological Society of which he was president from 1912 to
1916 and again from 1920 until his death. He was a fellow of
the Entomological Society of America, of the New York Academy
of Sciences, and a fellow and life member of the American Asso-
ciation for the Advancement of Science. The Staten Island Bird
Club, in which he was deeply interested, elected him president
from 1919 to 1930. He was also president of the Staten Island
Historical Society for many years and his offices in the Staten
Island Institute of Arts and Sciences have already been noted.
In addition he was a member of the New York Historical Society,
the Academy of Natural Sciences of Philadelphia, the Boston
Society of Natural History, the American Museum of Natural
History, the New York Zoological Society, the Brooklyn Institute
of Arts and Sciences, and other societies. In 1941 the Park Asso-
ciation of New York City awarded him a citation for his interest
and assistance in the development of parks and nature recreation
centers on Staten Island.

Furthermore, Mr. Davis was actively interested in Staten
Island history. He was the author of “Homestead Graves,”
“Staten Island Names, Ye Olde Names and Nicknames,” 1896,
“Legends, Stories and Folklore of Old Staten Island; the North
Shore” (Charles G. Hine, co-author), and “History of the Staten
Island Historical Society 1936.” In 1925, there was published
under the auspices of the Staten Island Historical Society, by Mr.
Davis, a work by William T. Davis, Charles W. Leng and Royden
Woodward Vosburgh, entitled “The Church of St. Andrew, Rich-
mond, Staten Island, Its History, Vital Records and Gravestone
Inscriptions.” And the following year, 1926, under the same
auspices and publisher there appeared a book on 4 1 The Confer-
ence or Billopp House, Staten Island, New York” by William T.

134

Journal New York Entomological Society

[Vol. LIU

Davis, Chairman Committee on History, Conference House Asso-
ciation. This is the house where representatives of American and
British governments met on September 11, 1776, to discuss the
war of the Revolution.

For several years Mr. Leng and Mr. Davis devoted much time
and effort to the huge task of producing a history of Staten Island
and in 1930 the Lewis Historical Publishing Company, Inc., of
New York City brought out the results of their labors in a monu-
mental work of four volumes entitled, “Staten Island and Its
People, A History 1609-1929,” by Charles W. Leng and William
T. Davis. This work was favorably received and a fifth volume
was published in 1933. Volume three contains biographical ac-
counts of Mr. Davis and Mr. Leng. A bibliography of all Mr.
Davis’s papers would, I am sure, reveal other works outside the
field of entomology and would indicate more fully the breadth of
his activities.

On November 7, 1900, Mr. Davis married Bertha Mary Filling-
ham, daughter of William and Elizabeth (Brook) Fillingham at
Livingston, Staten Island. Her death occurred a little over a
year later on December 17, 1901. Mr. Davis’s father died March
4, 1899, and his mother who lived to be 86 years, and with whom
he made his home at 146 Stuyvesant Place, St. George, S. I., died
on April 9, 1928. No one could know Mr. Davis for any length
of time without being impressed by his innate goodness, his quiet,
cheerful disposition and his abiding interest in natural history.
He was admired by many and by them he will be missed and
remembered. One of his poems “A Day Afield on Staten Island”
is printed below. — Harry B. Weiss.

A DAY AFIELD ON STATEN ISLAND
By William T. Davis

The glorious sun is setting
In the far and distant west,

And the clouds all golden laden
Seem sinking down to rest.

The day was one of glory and
The sun did brightly shine,

No heart in all the wild-wood
Has been so glad as mine.

June, 1945]

Weiss: Davis

135

I went where fancy led me,

Dor fancy is broad and wild;

I stayed where beauty kept me,
For beauty is soft and mild.

But the glorious sun is setting,
And the day is no longer mine ;
Could I but turn the hour-glass
And hold the sands of time.

136

Journal New York Entomological Society

[Vol. Lin

ANTEOS MAERULA, A NEW BUTTERFLY
RECORDED FROM FLORIDA

One bright, windy, sunny day, a female of Anteos maerula
maeurula (Fabricius) flew into one of the gardens at Port Sewall,
Florida. As it settled on one of the flowers, it was caught by
Carolyn Ponsonby, a little girl, who scarcely realized until after-
wards the extraordinary prize of her capture. This particular
specimen bears a closer resemblance to the Cuban and Antillean
members of this species, than to those from Mexico and Central
America.

Anteos maerula is a newcomer to Florida. To my knowledge,
this is the first and only record of this species ever reported from
this state. It is not even mentioned in the latest Check List of
North American Butterflies.: — Leonard J. Sanford

June, 1945]

Schwarz: Wax

137

THE WAX OF STINGLESS BEES (MELIPONIDiE)
AND THE USES TO WHICH IT HAS BEEN PUT

By Herbert F. Schwarz

We are indebted to bees especially for three things: their in-
dispensable service as pollinators of plants, whereby they make
possible life as we know it ; their welcome contribution of a table
delicacy, honey; their production of a substance, beeswax, that
has commercial value. The pollination of plants is the function
of bees of all kinds. The production of honey in edible quantities
and of beeswax is, on the other hand, apt to be associated prima-
rily with the honeybee, Apis. No one would want to under-
estimate the contribution that this insect has made to general well-
being through furnishing in abundance these valued products.
Nevertheless, it is perhaps only proper to point out that before the
discovery of America the indigenes of this Hemisphere were de-
pendent for both honey and wax on the native stingless bees.
There was even an apiculture in Mexico and southward of Mexico
into northern South America.

While honey was the principal object of this culture, beeswax,
too, found employment. In ancient Mexico beeswax was an
essential in the goldsmith’s craft, being used in the preparation
of the molds for the gold ornaments and figurines that are so
characteristic of the art of Mexico and Central America.1 An
earlier and a later technique are indicated and in the later process
workmanship in wax received greater emphasis than it did in the
earlier one.

While beeswax was used in metallurgy even in pre-Columbian
times, there is some doubt whether before the Discovery the native
populations of Mexico and Central America were aware of its
possibilities as a medium of illumination. Indeed, the testimony
of more than one early visitor to the region south of the Rio
Grande is to the effect that this common use for beeswax was not
originally recognized by the aborigines. Gomara, writing about
the middle of the Sixteenth Century, stated that the native people

i Saville, M. H., 1920, The Goldsmith’s Art in Ancient Mexico (Indian
Notes and Monographs, Mus. Amer. Indian, Heye Foundation), pp. 125-142.

138

Journal New York Entomological Society

[Vol. LIII

of Yucatan cultivated large hives and thus had plenty of honey
and wax, but he added that they did not know how to use wax for
illumination until they were taught how to make candles by the
Spaniards.2 Santaren, who is known to have been in the territory
of the Acaxee, a mountain tribe of Durango and Sinaloa in 1600,
wrote with reference to beeswax that the Indians “now know how
to make of it candles for the church,” with the implication that
probably no use was made of wax for lighting purposes by the
Indians prior to their contact with the Whites.3 On the other
hand, Redfield and Villa R4 or at least the senior author expressed
the opinion that probably “The Middle Americans burned wax
as an offering to the gods, as they burned rubber, copal, and other
substances, ’ ’ although conclusive proof could not be offered.
Certain we are of one thing, in spite of these conflicting view-
points, and that is that the use of wax for illumination was
greatly stimulated by ecclesiastical demand. The Indians ap-
parently soon learned the significance of wax for the making of
candles, and as a result wax became an important item of com-
merce not only in Mexico but in other regions, often far distant,
as well.

One of the chief ways in which the wax of stingless bees was
used in South America was for the making of candles.5 Many
of the native tribes apparently made such candles for sale to
Europeans. They were used in rural churches and in the Indian
missions.6 According to Maximilian, Prince of Wied-Neuwied,7
the Tapuya of Brazil prepared such candles by winding about a
thin core of wax a wick of cotton and then rolling the whole

2 Gomara, F. L. de, 1564, Historia delle Nvove Indie Occidental^ Pt. 2,
p. 65; 1940, The Conquest of the Weast India, p. 200.

s Beals, E. L., 1933, Ibero-Americana, Yol. 6, p. 10.

4 Eedfield, E., and Villa E, A., 1934, Chan Kom, A Maya Village (Carnegie
Inst. Washington, No. 448, p. 366).

5 Saint-Hilaire, A. de, 1830, Voyages dans les Provinces de Eio de Janeiro
et de Minas Geraes, Vol. 2, p. 371; 1848, Voyage aux Sources du Eio de S.
Francisco et dans la Province de Goyaz, Vol. 2, p. 164. Also Lepeletier,
A., 1836, Histoire Naturelle des Insectes — Hymenopteres, Vol. 1, p. 414.

e Azara, F. de, 1809, Voyages dans l’Amerique Meridionale, Vol. 1, pp.
161-162.

7 Maximilian, 1820, Eeise nach Brasilien in den Jahren 1815 bis 1817, Vol.
1, p.142.

June, 1945]

Schwarz: Wax

139

firmly into one mass. The same author stated8 that the Camacans
of Brazil arranged the candles intended for sale in long strings,
then placed them in elongated bundles and finally pasted a cover-
ing of large leaves on the outside. The Patacho of the same
country brought great balls of black wax to a settlement for the
purpose of making trades.9 Saint-Hilaire used candles made of
of the purified wax of stingless bees and found such candles satis-
factory although inclined to drip and give forth much smoke.10
Others have indicated that they were of inferior quality. Gard-
ner spoke of these wax tapers as coarse and ‘ ‘ about a yard long. ’ ’
His observations were made in Goyaz.11

To this day apparently beeswax is used in the Mayan village
of Chan Kom for making ceremonial candles. According to
Redfield and Villa R 12 the method of making these wax candles
is identical with that used for those of paraffin. From a wooden
ring placed horizontally are hung about fifty wicks and, as this
ring is revolved, melted wax is poured over the wicks until the
particular diameter desired is attained. Wax candles of this
type are either yellow or black, some hives producing wax of
darker color than do other hives. Candles of black wax are
occasionally lighted at funerals of grown-ups and at that part
of the All Souls’ Day ceremonies when there is commemoration
of the adult dead.

Redfield and Villa R found that beeswax is at Chan Kom used
for black magic. Figures are made of it and “there is a belief
that children who play with it become somnambulists.”

In Java the wax of Trigona ( Teiragona ) iridipennis F. Smith
is used by the natives for the production of ‘ ‘ Battik, ’ ’ a substance
for coloring calicoes and other textiles.13

Peckolt mentioned14 that in southern Brazil the indigenes used the

s Idem, Vol. 2, p. 219.

s Idem, Yol. 1, p. 284.

io Saint-Hilaire, A. de, 1848, Voyage aux Sources du Rio de S. Francisco
et dans la Province de Goyaz, Yol. 2, pp. 164-165.

n Gardner, George, 1846, Travels in the Interior of Brazil, First' Edition,
p. 329 ; 1849, idem, Second Edition, p. 250.

12 Redfield, R., and Villa R, A., 1934, Chan Kom, A Maya Village (Carnegie
Inst. Washington, No. 448, p. 49).

13 Schulz, W. A., 1907, Zeitschr. Wiss. Insektenbiol., Vol. 3, p. 67.

14 Peckolt, T., 1893, Natur, Vol. 42, p. 580.

140

Journal New York Entomological Society

[Vol. LIII

wax for attaching their feather finery and other things. Later15
that author stressed as particularly sought for this purpose, the
very sticky wax of “ Melipona longiceps Smith,” which was
doubtless a slip of the pen for Trigona longipes Smith, but may
be based on a misconception of the true character of longipes , a
synonym of varia Lepeletier.

According to Rayment16 in northern Australia the aborigines
decorated their head by attaching beeswax balls or beads to the
ends of wisps of their hair. Similarly they used beeswax to
form knobs on tassels of various ornaments, heightening the effect
by pressing into the wax the scarlet seeds of the leguminous plant,
Abrussus.

A further use which certain Australian natives have found for
beeswax is as a coping to protect their rock-paintings from rain
that might otherwise run down the sloping surface of the rock and
damage the picture. This is the practice of the Worrora, and in
a picture cave of this tribe Love17 found a semicircle of beeswax
over two representations of Warahninva, the .wedge-tailed eagle.

An Indian tribe (the Pury) living in Minas Geraes, Brazil,
used the wax of stingless bees in the fabrication of their arrows
and bows and also in making candles for sale to the Portuguese.18
By the Xicaque Indians of Honduras, I am informed by Mr. Y.
W. von Hagen, the wax of stingless bees was used for blowgun
sights and for fixing arrows.

In Cuba wax of the stingless bee, Melipona beecheii variety
fulvipes Guerin, was used in lithography. References to this use
of the wax appear both in Felipe Poey ’s account19 and also in
that of his son, Andre Poey.20 A lithographer of Havana by the
name of Marquier was the originator of the plan for using the
dark wax of this bee in the manufacture of lithographic ink, and
the ink made of this native wax proved more suitable for the pur-

15 Peekolt, T., 1893, idem, Yol. 43, p. 91.

16 Rayment, T., 1935, A Cluster of Bees, pp. 512-513.

17 Love, J. A. B., 1929-1930, Jour. Royal Soc. Western Australia, Yol.
16, p. 9.

18 Maximilian, Prince of Wied-Neuwied, 1820, Reise nach Brasilien in den
Jahren 1815 bis 1817, Yol. 1, p. 142.

19 Poey, F., 1851, Memorias sobre la Historia Natural de la Isla de Cuba,
Vol. 1, p. 169.

20 Poey, A., 1855, Bull. Soc. d’Acclim., Yol. 2, pp. 334-336.

June, 1945]

Schwarz: Wax

141

pose than did the imported ink of Europe. Subsequently litho-
graphic pencils were also manufactured from this wax. But
these instances do not cover all the known uses for the native wax
in Cuba. In addition, Felipe Poey noted that it was employed
in fastening artificial flowers, in mending shoes, in removing
corns, in fastening boys’ kites, etc.

Just how corns were removed or “cured” is not made clear but
this was not the only medicinal use for the wax of stingless bees.
The wax of Trigona ( Schwarziana ) quadripunctata variety
bipartite!, (Lepeletier) was, according to Peckolt,21 prized more
highly than any other native wax of southern Brazil for salves
and plasters, and Spix and Martius22 likewise indicated that the
wax of stingless bees was put to these uses. The fragrant wax
of “Yatei” ( Trigona jaty F. Smith) was used, according to
Wappaeus23 in Paraguay “only for perfume or in rural medi-
cine. ’ ’

Writing from Moulmein, Parish24 spoke of the preparation
and use in Burma of propolis (known as Pwai-ngyet) . Boiled
in water and softened thereby, it is then given an admixture of
petroleum and kneaded until it has the consistency of putty.
In this form its principal use is for caulking boats.25 Parish
indicated that Pwai-ngyet in the limited sense is the product
gathered by Trigona (Tetragona) laeviceps , which is a synonym
of Trigona ( Tetragona ) iridipennis F. Smith. A South Ameri-
can species, too, furnishes a resinous material prized for the caulk-
ing of leaking canoes. This species is Trigona fulviventris variety
guiance Cockerell, which, according to Miss L. E. Cheesman, stores
large yellow lumps used by Colombian fishermen for this pur-
pose.26

In East Africa, on the other hand, Morstatt found that wax of
all species of stingless bees was used to make canes and ropes
pliant.27

21 Peckolt, T., 1894, Natur, Yol. 43, p. 90.

22 Spix, J. B. von, and Martius, C. F. P. von, 1928, Reise in Brasilien in den
Jahren 1817 bis 1820, p. 523.

23 Wappaeus, J. E., 1867, Die Republik Paraguay, p. 1157.

24 Parish, C. S. P., 1866, Science Gossip, pp. 198-199.

25 Cook, M. C., 1865, Science Gossip, p. 252.

26 Cheesman, L. E., 1929, Trans. Ent. Soc. London, Vol. 77, p. 149.

27 Morstatt, H. A., 1921, Arbeiten Biol. Reichsanstalt fiir Land- und
Forstwirt., Yol. 10, p. 299.

142

Journal New York Entomological Society

[Yol. LIII

It has long been known that ambergris is a product of the sperm
whale — an origin sufficiently remarkable but far eclipsed by that
assigned to it in a seventeenth century article on stingless bees,28
in which it is stated :

“He (Monsieur Villermont) promises to show me, that Amber-
grise is nothing but wax, mixt with the Honey, which falls into the
Sea, and is beat about in the Waves, between the Tropics.”

One could almost wish that this fantastic explanation had
validity, so 'that yet another use for the wax of stingless bees
might be added to those already cited.

The wax when it issues from the wax-producing glands is ap-
proximately as pale as that produced by Apis, but almost always it
is subsequently mixed by the bees with alien materials that give it
a darker color. It is true that Bertoni29 classifies Paraguayan
bees’ wax as follows: (1) of brown color and much mixed with
woody materials ( amalthea and tat air a)-, (2) of ferruginous
color (various Melipona and Trigona ( Cephalotrigona ) capitata
F. Smith) ; (3) of clear yellow color and soft (various Trigona).
In general, however, the wax used in the nest architecture is far
from pure and tends to be dark in color. It was the conclusion
of Fritz Muller30 that in some species no more than 10% of the
building material is wax, resinous substances and clay accounting
for the other 90%. It is no surprise that, even in cases where the
admixture of foreign materials is much lessj this impure dark
wax is often very resistant to blanching. When Azara was mak-
ing his journey through the South American wilderness in the
late Eighteenth Century no way was known of bleaching it.31 Nor
was any blanching method known in northern South America,
for Fermin in his account of Surinam shortly after the middle
of the Eighteenth Century indicated that the dark wax of the
stingless bees of the region remained permanently dark.32

The Count da Barca, minister of the king, made repeated at-

28 J} Mr., 1685, Philosophical Transactions, Yol. 15, p. 1031.

29 Bertoni, A. de W., 1912, An. Mus. Nac. Hist. Nat. Buenos Aires, Yol.
22 (Ser. 3, Yol. 15), p. 139.

30 Miiller, Fritz, 1874, Nature, Yol. 10, pp. 102—103.

si Azara, F. de, 1809, Yoyages dans l’Amerique Meridionale, Yol. 1, p.
161.

32 Fermin, 1769, Description de la Colonie de^Surinam, Yol. 2, p. 301.

June, 1945]

Schwarz: Wax

143

tempts, according to Saint-Hilaire,33 to give the indigenous wax
of Brazil a light color, yet had not succeeded. On the other hand,
Saint-Hilaire observed at Goyaz a workman who had discovered
a successful method, which consisted of melting it, dividing it
into small bits and exposing these to the sun. He repeated this
process sixteen times, consuming two to three months in doing so,
but in the end the wax was almost as white as that of the domestic
honeybee. Not all wax had to be bleached, however, for according
to Azara,34 a large stingless bee found at Santiago del Estero in
the Chaco produced a pale wax that could be mingled up to a
half with tallow. Inhabitants of the region gathered annually
about 14,000 pounds of this wax. Bertoni35 spoke of the wax of
Trigona ( Plebeia ) mosquito Smith, as “almost white/’ and
Burlamaqui referred to the wax of one Brazilian stingless bee
that was paler than that of the Old World honeybee and superior
in quality. The bee in question, it was said, nested in hollows
of trees in certain regions near the Amazon.36

Peckolt made analyses of the ingredients in the wax of stingless
bees as he did of the ingredients in the honey. The wax of the fol-
lowing species was analyzed : droryana and ruficrus among
Trigona, and fuscata among Melipona. Wax, resin, water, ash,
were present in varying proportions in each case and usually there
was also present a humus-like substance. The wTax in each case
exceeded 50 per cent of the total, ranging from 52 per cent to 59
per cent. Resin was present in the proportion of about 31 per
cent to 42.5 per cent. The analyses are based on too limited a
study of material to be applied more generally. Some species are
known to use other foreign substances than those included by
Peckolt, but it is interesting at least to find the wax content so
high in the conglomerate material studied. However, the per-
centages cited did not apply to the brood-envelope, where the wax
ranged from about 11 per cent to about 20 per cent, and where, in

33 Saint-Hilaire, A. de, 1848, Voyage aux Sources du Rio de San Fran-
cisco et dans la Province de Goyaz, Vol. 2, p. 164.

34 Azara, F. de, 1809, Voyages dans l’Amerique Meridionale, Vol. 1, pp.
161-162.

ss Bertoni, A. de W., 1912, An. Mus. Nac. Hist. Nat. Buenos Aires, Vol.
22 (Ser. 3, Vol. 15), p. 145.

36 Raveret-Wattel, M., 1875, Bull. Soc. d’ Acclim., Ser. 3, Vol. 2, p. 757.

144

Journal New York Entomological Society

[Vol. LIII

addition to resin and humus-like substances, not to mention plant
material, there were in substantial proportions organic sub-
stances soluble in water and organic salts.37 In the analyses of
the brood envelope Peckolt apparently failed to include an analy-
sis of the brood envelope of ruficrus.

The process of preparing the wax in Cuba was described by
Poey.38 The wax was obtained from the provision containers,
which were first thoroughly cleansed of honey and of pollen and,
when dry, placed in a pan over a slow fire. The melted part was
drawn off, and if any dregs remained at the bottom of the pan,
they were thrown away. Another method was to boil the wax in
water, and to skim it from the surface of the brew, or to strain
it through a linen cloth; but this method, which worked well
enough in the case of wax of the European honeybee, is, according
to Poey, unsuited in the case of the dark wax of the stingless bees.
In the Mayan village of Chan Kom the inhabitants clarify wax by
melting it and then placing it in cold water, whereupon the good
wax comes to the surface. “The wax which is no good stays
underneath.”39

From East Africa used to be shipped from time to time wax
that passed under the name of “bumblebee wax,” but as there
are no bumblebees south of the Mediterranean region of Africa,
the designation was obviously a misnomer. The wax in question —
of an inferior quality — was the product of stingless bees.40 Doubt-
less there are many other uses to which the wax of stingless bees
has been put in the Old World and the New, but the instances
here offered indicate at least that this wax has proved its worth
in a great variety of applications.

37 von Ihering, H., 1904, Zool. Jahrb. System. Geogr. und Biol., Vol. 19,
pp. 267-269.

38 Poey, F., 1851, Memorias sobre la Historia Natural de la Isla de Cuba,
Vol. 1, p. 168.

39 Bedfield, R., and Villa R, A., 1934, Chan Kom, A Maya Village. Car-
negie Inst. Washington, No. 448, pp. 49, 50.

40 Morstatt, H. A., 1921, Arbeiten Biol. Reiehsanstalt fur Land- und
Forstw., Vol. 10, pp. 283-284.

June, 1945]

Bromley: House-Fly Enemies

145

INSECT ENEMIES OF THE HOUSE FLY, MUSCA
DOMESTICA L.

By S. W. Bromley, Ph.D.

Bartlett Tree Research Laboratories, Stamford, Conn.

INTRODUCTION

While the house fly, Musca domestica L., is one of the most
abundant of insects and one of the most important from the
standpoint of human health and sanitation, there are very few
accounts in literature of the insect predators of the adult fly.
The following is an annotated record of the writer’s observations
on this subject.

The house fly is much less numerous in New England now than
was the case thirty-five years ago. Better sanitation, screens, fly
sprays have all contributed, but the most important factor, com-
pletely overshadowing all others, has been the decrease of the
horse as it gradually became replaced by the motorized vehicles
of the gas engine era. The principal breeding medium of the
house fly was horse manure. With this staple food of the house
fly gone, house fly populations were bound to decrease.

The passing of old Dobbin brought about other far-reaching-
changes such as the decrease of the English sparrow which de-
pended to a large extent for its winter forage, in the snow-covered
northern states at least, on the undigested grain in horse drop-
pings. As the horse population dwindled, the demand for hay
dropped off with the result that farmers began to abandon their
mowings initiating a succession of vegetation of weeds and brush
to young woods on the old hayfield sites. This affected bird ecol-
ogy to the extent that such grass-nesting birds as meadowlarks
and bobolinks began to fade away from their old haunts with the
passing of the grassland.

The effects on human health were evidenced by a decline of
tetanus or “lockjaw” the causal organism of which throve in soil
or straw in the neighborhood of horses, as well as the decrease of
typhoid fever which was carried largely by the house fly. The
replacement of dirt roads by hard-surface highways for auto

146

Journal New York Entomological Society

[Vol. LIU

transit did away with the clouds of dnst which together with
proximity to horses, in some cases, produced, and in others, aggra-
vated asthma in many humans.

One entomological effect of the decrease of horses was the deple-
tion, in New England, of the horse hot flies, Gastrophilus spp.,
which were completely dependent upon the horse for their exis-
tence. The tabanid horse flies which have a variety of hosts were
affected little if to any extent. Some. of the coprophagons beetles
specialized in equine ejecta and have now become scarce, as for
example, the beautiful Geotrupes semiopacus Jek.

HORNETS VS. THE HOUSE ELY

Back in the days when streets, houses, and farms reeked with
house flies, I spent many hours when, as Mark Twain once re-
marked concerning his own observations on the habits of ants,
“I should have been better employed,” watching the killing of
house flies by their insect enemies.

First and foremost of these insect predators was the common
white-faced hornet, Vespa maculata L. These hornets dogged the
house fly assemblages and persistently hunted down the flies.
Herewith are some of these instances, a very incomplete record
of my observations inasmuch as this was so commonplace an
occurrence I did not take the trouble half the time to make the
record.

Warren, Mass., August, 1908. Around pig sty at barn. Three
captures noted.

Charlton, Mass., August 10, 1909. Around milk bucket. Two
captures noted.

Dudley, Mass., August, 1910. Around barn. Two captures
noted.

North Woodstock, Conn., August, 1910. Around farmhouse
porch. Five captures noted.

Charlton, Mass., August, 1910. On side of barn near door of
farmhouse. Four captures noted.

Southbridge, Mass., July, 1910. Around produce at a meat
market. Hornets noted demonstrating their poor vision by butt-
ing into nailheads which they mistook for flies on the side of the
building. Three captures noted.

June, 1945]

Bromley: House-Fly Enemies

147

Charlton, Mass., August, 1918. Around pig pen at a farm.
Two captures noted.

Certain other social wasps were seen to hunt down house flies
but to a less extent than V. maculata. These were the common
native ground-nesting yellow- jacket, Vespa maculifrons Buy.,
and the English wasp, Vespa vulgaris L.

For V. maculifrons I have the following records of their cap-
turing house flies.

Southbridge, Mass., September 28, 1913. On sunny side of
stable shed. Two captures noted.

Southbridge, Mass., August 12, 1917. Around a rendering
plant. Three captures noted.

On May 4, 1928, on a greenhouse window at Chappaqua, N. Y.,
I saw a brown wasp, Polistes fuscatus Fabr., capture and devour
a house fly, a rather unique instance as this wasp feeds as a rule
on caterpillars.

Among the solitary wasps, Bembex spinolce Lep., Oxybelus
quadrinotatus Say, Stictia Carolina Fabr. (The “Horse Guard”
of the South), and species of Solenius ( Crabro ) have been known
to seize and store up house flies, although such observations have
never chanced to come within my own personal notice.

ROBBER FLIES YS. THE HOUSE FLY

Next in importance to the hornets as house fly enemies were the
robber flies. This was true of New England but throughout the
Southern States the house-fly killing asilids are much more numer-
ous than in the north, and undoubtedly surpass the hornets as
enemies of Musca.

Around restaurant doors and windows or in the open-air
markets for meat, vegetables and fruit where house flies swarmed
in the old days, it was not unusual to see larger, more slender
flies of another breed pouncing on Musca domestica individuals
and sucking their blood. These were robber flies. I have several
records which I have preserved for many years, and now publish
a number of them, some for the first time.

Asilus notatus. Wied. A common asilid at the edge of woods,
fields, and thickets, along fence rows and around buildings. It
used to be attracted to house fly assemblages and was frequently

148

Journal New York Entomological Society

[Vol. LIU

found in towns and cities, feeding on flies. Woodstock, Conn.,
July, 1909, on moving wagon which was swarming with flies.
Southbridge, Mass., July 7, 1910, on side of meat market build-
ing. Southbridge, Mass., July 7, 1910, on box of fruit outside
market. Charlton, Mass., July, 1910, on side of barn. Worcester,
Mass., June 24, 1910, on restaurant window. Worcester, Mass.,
June 24, 1910, on sign on open-air meat counter. Boston, Mass.,
July, 1911, on sunny side of meat market. Boston, Mass., July,
1911, stuck with its prey on fly paper on meat counter. Boston,
Mass., July, 1911, on exposed meat at open-air market.

Asilus sadyates Walker. This is a late summer counterpart of
A. not at us. Southbridge, Mass., August, 1909, on side of building
near kitchen door. Southbridge, Mass., August, 1909, on porch
of house.

Asilus lecythus Walker. Larger than A. notatus and more
local. Rockville, Pa., July 4th. Specimen in State Collection,
Harrisburg, Pa.

Neoitamus ( Asilus ) flavofemoratus Hine. Worcester, Mass.,
June 24, 1910, on restaurant window.

Erax cestuans L. The leading fly-killer of the robber fly group,
east of the Rockies, is the abundant and widespread Erax cestuans
L. This Erax alights in bushes at the edges of woods or fields, on
fences, buildings, domestic animals, and even on man himself
where it seizes house flies, deer flies, mosquitoes and many other
pestiferous Diptera.

A common name throughout the South for Erax cestuans L.,
together with other species of the same genus is the appellation
‘ ‘ fly-hawk, ’ ’ and every farmer has noted its fly-catching proclivi-
ties.

In August, 1927, Dr. C. O. Eddy, then at Clemson College,
S. C., wrote me that this fly-killer had been noted throughout the
summer at the insectary building where it rested on the outside
walls or screens, disposing of a great number of house flies each
day. Here are some records of Erax cestuans feeding on Musca
domestica. Charlton, Mass., July, 1910, on side of farm house.
Charlton, Mass., August, 1911, on barn door. Southbridge, Mass.,
July, 1912, on screen door. Southbridge, Mass., August, 1917, on
side of rendering plant. Columbia, Missouri, July 26, 1923, on

June, 1945]

Bromley: House-Fly Enemies

149

fence near farm building. Eureka, Missouri, August 4 (D. J.
Knull). College Station, Texas, June 1, 1932, on side of small
building; June 11, 1932, on wooden fence along pasture; July 4,
1932, on horse shed; July 8, 1932, on water tank near barn; July
14, 1932, at dairy building. Cedar Creek, Campbell County,
Tennessee, June 10, 1943 (R. M. Goslin).

In sandy farm yards in the Southern States, a number of
ground-resting asilids are common feeders on house flies. Out-
standing are the two “fly-hawks” of the genus Erax: E. barbatus
Fabr. and the so-called “Snorey-Joe,” E. interruptus Macq., as
well as the smaller Proctacanthella. ( Asilus ) cacopilog.a Hine ;
while the much smaller Stichopogon trifasciatus Say occasionally
captures the house fly.

As distinguishing common names for the fly-hawks, I might
suggest bush fly-hawk for Erax cestuans L. ; field fly -hawk for
Erax rufibarbis Macq., and sand fly-hawk for Erax barbatus
Fabr.

Erax barbatus Fabr. This active little asilid feeds on house
flies extensively when it can find them. Cane Springs, Imperial
Valley, California, June 20, 1925, on sand near a filling station.
Weslaco, Texas, June 20, 1933, on sand in citrus grove near pack-
ing shed ; June 21, 1933, on truck in orange grove ; July 2, 1933,
on sand in citrus grove near packing shed ; July 31, 1933, on sand
in citrus grove near packing shed ; August 14, 1933, on sand near
packing shed ; September 14, 1933, on sand in road through citrus
grove.

Erax interruptus Macquart. While this Erax feeds largely on
grasshoppers and Lepidoptera, it also frequently selects house
flies. Yuma, Arizona, July 3, 1925, on sand in horse corral.
Chadbourne, North Carolina, September 12, 1925, on path
through field. Clemson College, South Carolina, August 16, 1926,
on path through field ; August 17, 1926, in barn yard of dairy
farm. Calhoun Falls, South Carolina, August 17, 1926, in road
to farm yard.

Erax rufibarbis Macquart. A common species throughout the
eastern half of the United States, alighting on the ground in open
fields and paths. Southbridge, Mass., September 5, 1914, in path
near dump; September 10, 1915, in path near dump; September
16, 1915, on ground in dry field near stable.

150

Journal New York Entomological Society

[Vol. LIU

Proctacanthella ( Asilus ) cacopiloga Hine. A common ground-
resting species in sandy regions of the South. College Station,
Texas, June 12, 30, 1932, in path near gate to pasture.

Prod acanthus philadelphicus Macquart. Southbridge, Mass.,
August, 1911, in field near stable. A unique record for this large
robber fly whose capture-mechanism is designed for larger prey.

Stichopogon trifasciatus Say. South Union, Ohio, June 29,
1941 (R. M. Goslin). Scioto River, Columbus, Ohio, July 1, 1944
(R. M. Goslin).

DRAGON FLIES YS. THE HOUSE FLY

Certain dragon flies were seen to capture house flies. The late
Dr. Charles Branch Wilson in his comprehensive and interesting
article on ‘ ‘ Dragon Plies in Relation to Pondfish Culture ’ 7
(Bulletin of the Bureau of Fisheries, Vol. XXXVI, Document
No. 882, 1920) on pages 212-214 records several species of dragon
flies; i.e., Gomphus f rat emus, G. vastus, JEschna constrict a, Leu-
corrhinia intact a, Libellula luctuosa, Erythemis simplicicollis and
Plathemis lydia, feeding on house flies. My own records are
rather few. The commonest dragon flies around barns and
stables, Sympetrum rubicundulum Say and vicinum Hagen, I
never saw make any attempt to capture the flies. My only records
are of some of the large aeschnid dragon flies which would occa-
sionally put in an appearance at house fly gatherings, select a
victim and depart.

JEschna constrida Say. North Woodstock, Conn., August,

1909, in farm yard. Southbridge, Mass., September, 1912, in
barn yard ; September 28, 1913, near stable.

JEschna clepsydra Say. Lake Mashapaug, Union, Conn.,
August, 1916, at kitchen door of Summer Hotel. Charlton, Mass.,
August, 1918, in farm yard.

JEschna umbrosa Walker. Southbridge, Mass., August 12,
1917, at rendering plant.

JEschna canadensis Walker. North Woodstock, Conn., July,

1910, near barn.

Boyera vinosa Say. Southbridge, Mass., August, 1910, in late
P.M., near stable.

RECENT RECORDS

With house fly aggregations smaller during late years in the
north at least, their enemies are less in evidence. It usually took

June, 1945]

Bromley: House-Fly Enemies

151

a great abundance of the flies to draw the hornets, robber flies and
dragon flies, where the expenditure of effort in capturing their
prey could be quickly and easily recompensed.

One may still find flies in abundance at piggeries in the dung
of which the house fly will breed. The house fly does not ordi-
narily breed in cow manure. At pig-pens, the white-faced hornet
still finds house flies in sufficient numbers to be tempting, while
garbage dumps attract great numbers of Musca domestica.

Hoping to get the facts on present-day house fly predatism, one
hot Sunday morning in August, 1943, I walked over to the town
dump. Ever since its inception several years ago, this dump had
been, to use a trite expression, the bane of my existence. It would
catch fire every now and then and on cold winter nights, the acrid
and nauseating fumes of smouldering wet paper and decomposing
garbage would cascade down the river valley and collect in my
back yard about a mile and a half away. I organized committees
and tried to make things as miserable as possible for the good
Town Fathers with my angry protests and demands for the abate-
ment of the nuisance but to no avail. The dump went on growing
and waxed mighty both in area and stench, engendering great
numbers of Norway rats, European house crickets, flies and mos-
quitoes.

Finally compensating factors, principally ornithological, began
to develop. The dump attracted flocks of crows. The crow is the
most effective bird enemy of the Japanese beetle, so I didn’t com-
plain about them. The rat populations brought back the horned
owl, which I had heretofore considered extinct, at least as a nest-
ing bird, in the Stamford area. The great horned owl prefers the
brown rat to all else as food: there is no better “ ratter” in the
whole world.

After several years had gone by, particularly as special efforts
were made by the authorities to stop the fires and reduce the
odors by quickly filling with soil, I began to lose my resentment.
So on this particular day, I decided to brave the effluvium and see
what was happening to Musca domestica at the hands of its insect
enemies.

The flies swarming on the garbage were mostly the common
carrion or garbage fly, Phormia regina Meigen, with a scattering

152

Journal New York Entomological Society

[Yol. LIII

of green-bottles, Lucilia sericata Meigen. Musca domestica
swarmed over the old crates and cartons. There were six or eight
white-faced hornets in sight all the time, seizing all three kinds
of flies.

There were ten to fifteen dragon flies, Libellida pulchella
Drury, coursing over the garbage and over the big patches of
scattered broken glass which they evidently mistook for pools of
water. I watched them through the binoculars. They were feed-
ing, not on house flies, but on some very minute insects which they
picked up in the air and then chewed up while resting on some
exposed point. They paid no attention to the house flies, or to the
garbage flies, either.

On September 6, 1943, I revisited the dump. The white-faced
hornets were again the principal fly hunters, although at this time
I counted, in addition, five English wasps, Vespa vulgaris L.,
seizing house flies. At the edge of the dump a dragon fly, JE sclina
umbrosa Walker, swept in and snapped up a house fly which had
just left its perch on a packing box. This was the only time I saw
an 0 donate catch a house fly at this dump.

The foregoing may be taken to indicate that in spite of a
marked reduction of flies in New England during the past few
decades, in spite of an apparent dwindling of breeding places, and
in spite of the pressure of natural enemies, together with man’s
efforts at fly eradication, the lowly house fly is still with us and
will probably be for milleniums to come.

June, 1945]

Richards & Weygandt: Fat Penetration

153

THE SELECTIVE PENETRATION OF FAT SOL-
VENTS INTO THE NERVOUS SYSTEM
OF MOSQUITO LARVAE1

By A. Glenn Richards, Jr., and Jane L. Weygandt
Zoological Laboratory, University of Pennsylvania,
Philadelphia, Pennsylvania

It was pointed out in a previous paper that when certain lipid
solvents are marked with Sudan dyes and introduced into the
tracheal system of mosquito larvae the central nervous system may
become intensely colored with the dye while other tissues remain
uncolored (Richards, 1943). There are reasons for thinking that
the dyes do not leave the tracheae except with the solvent, and
accordingly that the distribution of dye in tissues can, with cer-
tain limitations, be used as an index of the penetration of the
solvent (see discussion). The present paper summarizes data on
over 100 organic compounds of which the majority showed a selec-
tive coloring of the nervous system; the remainder usually were
negative (dye remained in tracheae).

Materials and Methods. These experiments have been per-
formed over a period of a year and a half. In general the entire
series of compounds listed were tested once using fourth instar
larvae of the house mosquito, Culex pipiens ; then with few excep-
tions they were subsequently tested again using fourth instar
larvae of the yellow-fever mosquito, Mdes cegypti. The approach
was to introduce into the tracheae a concentrated solution of a
lipid (Sudan) dye in the solvent being tested, and subsequently
check the distribution and condition of the dye. Notes were also
made on obvious gross pathology.

The standard procedure adopted was to remove the larvae from
the culture singly and place them on filter paper on the stage of
a binocular dissecting microscope. A fine glass capillary tube
from a small hand syringe was introduced into the spiracular
opening at the end of the respiratory siphon when the valves were

1 The work described in this paper was done under a contract, recommended
by the Committee on Medical Research, between the Office of Scientific Re-
search and Development and the University of Pennsylvania.

154

Journal New York Entomological Society

[Vol. LIII

open, and the solution being tested forced out of the capillary and
induced to flow down the longitudinal tracheal trunks until they
were full or nearly so. The larvae were transferred to 10 per cent
formalin in saline for an hour, and then the brains and ventral
nerve cords dissected out and examined in formol-saline under
the compound microscope.

The timing was varied in cases where negative results were first
obtained. Commonly a longer or shorter time was given between
injection and dissection, and the cover glass sealed on and the slide
examined at intervals up to 24 hours. In several cases specimens
were kept in culture water instead of formol-saline to offset the
possibility that the slow formol fixation might be interfering with
penetration (in no case was it found to do so). In some cases the
material was allowed to enter the tracheae from a water-test sub-
stance interface instead of being introduced from a syringe.

As lipid stain, Black Sudan B (National Aniline and Chemical
Company) was usually used because of its intense color and its
chromatic effects. When negative results were obtained with this
complex polyazo dye, Sudan IV or Oil Red 0 was tried because
of its smaller molecular size.

When the dye penetration is considerable the color can be seen
in the nerve cord through the body wall. In these cases it is easy
to see during dissection whether the other tissues are colored to
any similar degree. In all cases other tissues were watched for
the presence of stain. As additional checks on the selective dye
penetration from tracheae, blood injections were made with the
same solutions, and larvae with the respiratory siphons tied off
with a hair-loop were immersed in the solutions to test cuticular
penetration.

Results. In the following table are presented data from over
2,000 tracheal injections. The accompanying blood injections
and cuticle applications were used primarily as controls, and are
not included.

Unfortunately it was not found possible to estimate and inter-
pret the results with sufficient accuracy to permit a precise
quantitative comparison of the penetration and accumulation of
the substances tested. As an attempt to express partially the
obvious gross differences a series of five units was adopted. For

©

©

B

B

B

B

a

g

a

©

©

«H

o

o

o

B

B

B

>

>

©

©

©

o

o

a

g

a

o

o

o

o

o

Eh

Eh

m

OD

QQ

GQ

©ffl®©®®©*

B P

>

>

>

>

>

_i>

P

P

p

P

P

p

P

P

P

B

B

B

B

B

B

B

B

B

be

be

be

be

be

be

be

be

be

©

©

©

©

©

©

©

©

©

£

£

£

£

£

Jz;

£

£

£

©

©

©

©

©

©

s

©

>

>

'>

.3

GQ

>

P

P

p

p

p

p

j

O

P

B

B

B

B

B

B

—<

'o

B

be

be

be

be

be

be

*bjD

GQ

be

©

©

©

©

©

©

©

£

£

Jz;

£

£

P'r

o S
> ©
rH -t-S

© 5Q

B

B "

© N IN (M cq CO Tt<

>

T(H gj

© ©

-M -M 02 CO CO

B B B B 3

be be P P

© <D TrH . Th -rl -r-l

£ £ p ft p p

m ® ®

© ^
.5 ©

<M CO
© ©

SH Sfcl

3

SK ?£

P P

B B

[H &

P tz;

3 B B 3

«K SB

P £ 6 P

w xn m gq

C5

% $P

P

© N N N CO CO

.£h © © ® ® ©

in v m m

IB SB £ SB

P P P P

b o

Jz; P

' H H W

pi p
£13 S§
p^ « h

t^0^s_,rdrdrCJrCrCirCri5 'd rtf 'C r^j rjJ 'B £-1 'B '"B

bebe. 3 OOOOOOO OOOOOOO O O © -i-< -r-i O

©©ajOOOOOOO OOOOOOO O O O B B O

££;PhPPPPPPP OPPPPPP PPPPPP

| 8
£ 'H >

| ° g

COOlOlOOlOOOTtfOi

GO o o o

S«s
Jz; &

C<I ^ CO rtH

IN in IN ^

(NININNNNNINININ

lO il CO CO CO CO IN

©

B

B

© ©

N M

V W ^ O kv Lj

S B s ^ ^ ^ ^ ^ n

B B M

B

* a

© 3

el

/-s P

> S

B g

© P

o

3 ©

© a

• rH Cw

h3 P

?-< rH P

O S P

p

©

*-*3 o 58

gjp P

£ -g »

s ° ®

-s 3*43

P P

?H fH
© ©
Pi PI
© ©
be be
© ©
T3 P?

■ ^
be

p*"»

be be be be
© © © ©

r2 ”

©

pP

-H-3

p

p

o o o o

to! Pp Pp rP

4-3 43 -*-3 -4-3

P P P P
ft P A ft

o ®

pp

p 2
p £

©

>

© © © ©
> > > >

© OQ
>

© © © ©
£ £ £ £

© o

5zi fc

"g O

p

o £

p 2
•n «h

S s

CO s

co

p «

Ph 02

©

P
P
. o
© £
> P

ojcIcStidcjcIrtcjcjsJo

bobobobobobobobcbobobcsS
© © © © ® © © ® © ■© © .P

43 +3 +3

p p p ® p
be be bo $ bo

£££££££££

p ®

&p ”

© gj

h£

3 &>

m ©

>3 fc

© g be
p £
G>pq £

'•§ o g

pt

<D ^
^ ©

©

©

p CO

CO © 02 CO

02 CO
-1-3 -M
© ©

©

© co co co
Ph

P P P
be be §h

© © Ph

p p p p p p

be be be be be be o

®©®©©®?h?-h

©

® © © ® ® ©

P P
‘ O

be

£L| £P Ph P P P
ri o O O O O ©

“ p 5 p n p p

GQ

CO <© 0)

© -ft -A

02 4-3 4-3

p p p
Sh be be

-P © ©

p

g p 8

•J a-g

^ p 00 p

P^ h

$_, P r *. rP rP P P P P i_,rPrP tj P P P P P P

o bc-rt-P o © o bo be bo be -rH o © -3 o o o o o o

o®pp©oo©©®©poopoooooo
Ph^phPPOP^j^^^poopoooooo

o P
o

P-l 43

p

£? be
® ©

P |> Jz;

© 0}

0 P

P5*-1 >

1 ® S

& ^

P O • r-J

| P

CONWWtMNNMOJNNmfM

(M CO N W N

TjHTtlT*HrtlcO'^CO^'cHCOCO

lOCilplOlOlOlOlOO

©

P

P

P

©

P ©
Pi P

£

p

2 ©
a p

? j

a -s

© o

U f-H

£ Jz;

p p
p p
© ©

© 2
£ p
cb ■S

£ *3

2h p
© p

P X
p ©

p -p

2 ^
p r-T
©

<73 t— I

rH (M

4-3 Ph
© P

O O

O p

to os
to ©*

?n. p>

i© oo

m

p

©

©

o

^ ^ 2r
© © p

O O H

-p
©

£ 9

b. b.

© ©

o © o

© © ."S

Ph fts O

O O

©

©

tol _ P

9 p 3

j-H O ©

■S 2 ^

P © -I-H

PQ <1 ft

a

a

o

•rt ^

pi w £

pj

o »

t o £

J> fc.2

•r1 ® ®

&

bf) oa
;3 o
CQ PM

£ > > -g u > > o > .> > > > > > > > > .> >

'43 '-rt S ^ ’43 ‘43 • rH P* "-2 '-rt *43 +3 '43 "43 +3 +5 ’.+j "43 '43

„c3d „.do3'dc3o3dc3c3c3c$c0c3o3c$c3

52 be be pj 52 be bo £ bebebebebebebobebebebebe

CO CO rt*

03 ’ 03 03

GO GO GO

pj d pi

sh sh sh

0) rt<
.fn 03

tl P

03 CO
t> rt

CO

Q P P £ ft

rt-M-M-^coco^co^co
gogogogogogogogogogo
^^„a3®®®a:>a;>‘I)<liOQ;’M

£$ £ S3 ^^^^IPhTM^Tm'Ei'Pi^
bcbeiyooooooooooSH

o

o

PM

Hd trt, fH

o rt o

O 03 O

O PM

& pi p, ^plrdpHrdrd^rdrdrd^rdrdrd^l

biD -rH • rH . rH • rH © *rH O O -H O O O -rH O O O -rH

a> c3 c$ c3c$oo3ooo3oooa3oooa3

£ P pH

tH CO CO
CO (M rH

NtDtOtOHtD^tOlOO

§ ©-,3

£ ’&

i(a3Kl(Mi(COCONi(CONK|(M

COCONtOrtl^tOtOO^tOHHHCO

rtP M3

rP

i— I +2

O 03
03 Pi

^ P

O 03

- - O GO "5

_ r—l rrt rrt H

M~s M> £•■» oj 33

P P ft

p .rt P P P P Hrt

P P PQ PQ P PQ O

03

C$ Pi
03 P
P cO
M2 .33
8 ^

O p

g M2

M2 p

.rt 0

P 0Q

cO [j§

£ '§3

© $>

O H

differences between these two samples are not known.

^5

cd

©

b
o

'■p fd £
oj o pd

fH O

| ^ g
© _ o
bo P ?h
© Id «h

^ -a «

5D B S

® S‘«

§s^.

o Ph

b

_©

P © 58

2

a -s S

C ° «
-m £ "-*3
02 .2 ■*-’

©

©

©

©

©

©

©

©

©

©

©

©

©

©

©

©

©

©

©

B

©

B

_>■

t>

_>

>

>

>

>

>

.!>

>

>

>

>

a

g

a

p

P

P

P

P

P

P

P

P

P

P

P

P

•M

P

•p

P

P

P

P

cd

cd

cd

cd

cd

cd

cd

cd

cd

cd

cd

cd

cd

cd

cd

cd

cd

cd

cd

nd

rd

bo

bo

bo

bjo

bo

bo

bo

bo

bo

bo

bo

bo

bo

bo

bo

bo

bo

bo

bo

cd

cd

cd

©qj©. ©©©©©©

©©©©©©©

W W £ £ £ £ £

© © ©
£ £ £

© © ©

WWW

ri

O 2

’■g | a

5 > ©

-c ^ p

‘ “ GO

rP
m

0>

P‘r

02 02 50 Jd 50 02 02

3 B 1=1 Cl, B 3 B

§B ?B olfcSttSH

SSSpSSSSS

02 N

2 s

& 2

P SB

CQ

H ^ CO
© © ©

B ^ © ©

o ©.fp.fi

F^202O2CO5O,r-'CO-|P-(-J

o.BBBB'to^Bcd

o IB ?B ?B §W ©SB

fH .H -rH -rt tH B >H © ©

qAqqqg>pWW

02 50 02

B

SB SB

P ft P

■ d w se

P .2 g p

d S d

Pp P is

® ^ ^ p ©

© O O .rrl ©

X O O Cd *

P O O P p

© © ©
© © ©
M

P P P

^ 2
W =H >
B © H
B cd

W ^

OlOHCOlfllOHdlO
CO CO r- 1 rH CO t— I i— I

iooiocooohooooo

CQ t— I CO r-l i — I CO ^ I — ICQ

So

Tfl lO CO CQ CO CQ CQ

^ W (M CO

CQCQOOOOOOO

OCirHOOlOOOCOCOrH

© ©
B

© ©

P

P ©

0

© eo ""

?-t r-~o. 1 — C

© ©

4P 6-5 ©

3 H

01 © .g

P d h P

B ©

© Sr-
'S ©

•| s

^ Sd

e ©

F e
© £
W p

CQ

w

p

© ®

© b

M P S

Poo
© ■+■> r~H jb

p © e '3,

o t ^ o

02 £~* SS 02

M p Kp m

^ w

©

© B
B ©

B %

o X

ffl Eh

Number Number bution Distribution Distribution

Material Source of * of in small nervous in °ther Miscellaneous

replicates larvae , ^ , system tissues

©

o

ft

© ^
S

© ,5

© o
be ^

© -ij
x© ©

X X

© ©
P P

g

g

cj

©

>

•m -

>

>

ft

ft

ft

ft

ft

•M

ft

4^>

ft

ft

ft

X©

X©

X©

X©

©

©

©

©

©

©

©

o3

©

©

©

©

©

1©

©

be

be

be

be

be

be

be

be

be

be

<x> cd

w w

© © © ©

02

©

U2

02

C/J

H-i

©

-4—*

©

‘ft

£

s§

5©

be

©

be

©

o

s

s

s

s

£

£

A

© < -

be be be $h

© © © ft!

•'tf <© ^ ®

© .Eh © .Eh

os m 4J

s?s

S5 o

© 03
be be
© ©

£ 5

© © © ©

© © © ©

X X X X

rp © C3 . ^

o © be be .© o

© O O .© .© © O

H H H R fftft

ra H H n ® io
CO ©3 lO I— I r— I

(M N CO CO (M (M (M

CO W W (M N

NWHOmaiOHOlOlO

tH rJH

©

©

O

K> r— '

©

> _
o 2

m r-.

© o
a « ©

© F
© ~ © ^

©

„ ^ 03 'H

Pi S © °
PiO S

^ >-> ft S .5 -h ©

ft X ft pq ft O <1

©

©

©

©

ft

ft

o

-M

©

©

Pi

ft ft

5s S

© ft

^ O .O g

-X ©
J X©

be ^
© ©

Pft q

3 *

ft «w
© ©

, § S'

o ^
o

© 5

ft ft

2 °
S P

© 03 ©

H -fi -M

H CC 02

©0 O rH Cvl CO

X©

©

©

©

be

x©

o

ft

(M CO - ^ l© ft

oo

Glyco Products Company. 14. Shell Development Company.

National Carbon Company. 15. Atlantic Refining Company.

U. S. Industrial Chemicals Company.

160

Journal New York Entomological Society

[Vol. Lin

distribution in the small tracheae and tracheoles the sequence:
excellent, good, fair, poor and negative is used. For the distri-
bution in the nervous system the same scheme is followed but with
the addition that this penetration may be either diffuse or in the
form of droplets; thus “diffuse 1” means an excellent diffuse
staining, “diffuse 2” a good staining commonly restricted to the
fiber tract areas, “diffuse 3” a more erratic or less extensive pene-
tration, and ‘ ‘ diffuse 4 ’ ’ a poor staining that showed in only one
or a few of the specimens. A glance at the table will show that
good distribution in tracheaa is not necessarily correlated with
good penetration into the tissues ( e.g ., compare hexadecane, car-
bon bisulfide, terpin diacetate and xylol).

Materials are arranged according to chemical structure. First
aliphatic hydrocarbons, substituted hydrocarbons, alcohols, ami-
nated alcohols, aldehydes, ketones, » ethers, esters, acids, salts,
cyclo-hexanes, mixed rings; then aromatics: benzene series, aro-
matic petroleums, alcohols, cresols, amines, ketones, ethers and
esters.

Discussion.— Evaluation of Method. The primary difficulty
in any study of this type is to show what relation (if any) the
distribution of the dye bears to the distribution of the solvent
being tested (Bredenkamp, 1942). Theoretically movements of
the solute are independent of those of the solvent. Our best evi-
dence that the dye does reflect movements of the solvent in these
experiments comes from chromatic effects. Black Sudan B is
blue in some solvents, violet or even red purple in others. If a
violet solution in xylol is mixed with paraffin oil the color changes
to blue. In the lipids of mosquito larvae the dye is blue. If a
solvent in which the dye is some violaceous shade {e.g., benzene
series) is introduced into the tracheae and its passage into the
nerve cord watched, one finds that the first color in the tissue is
violaceous and that this changes to blue only slowly and in places
where the dye (and presumably solvent) is not too concentrated.
This is good evidence that in these cases the interpretation is
valid.2

This chromatic effect, of course, cannot be obtained with sol-
vents in which the dye is blue, but in no case did we see any

2 In the case of pyrethrum penetration, can be observed directly by the
yellowish color.

June, 1945]

Richards & Weygandt: Fat Penetration

161

indication that the dye enters the tissue except with the solvent.
In a few cases both the solvent and dissolved dye apparently re-
mained in the tracheae, even for solvents in which Black Sudan is
less soluble than it is in the nerve sheath lipids ( e.g mineral oils) .
However, in some cases the solvent clearly gets into the blood or
tissues leaving most or all of the dye precipitated in the lumen
of the tracheae. Certainly great caution must be observed in
evaluating the distribution of dyes. In general, we feel that the
positive data are valid so far as they go, and that negative results
on accumulation in the blood and non-nervous tissues are valid
for solvents which do show accumulation in nervous tissue.

The term “negative” appearing in the table means only that
no clearly recognizable color was found. For the “distribution
in small tracheae” this almost certainly means that the solvent
never reached these tubes, and so the word ‘ ‘ none ’ ’ might be quite
correct. We find that substances with fair or good water solu-
bility seldom reach the small tracheae. This is true even of those
that are good lipid solvents — acetone, ethyl ether, etc. How much
this is due to poor affinity for the tracheal walls, how much to
volatility and how much to direct penetration from large tracheae
into the blood is uncertain. What is certain is that these vari-
ations decrease the accuracy of the data on penetration. For
negative results with dye penetration into the tissues, however,
we have no way to distinguish between whether the dye is all in
the tracheae or whether so little got into the tissue that it cannot
be recognized. In at least some of the “negative” cases pene-
tration of the solvent takes place because one can see gross patho-
logical changes or precipitated dye in the tracheae. It is possible
that all of the substances tested (molecular weights up to about
375) show some penetration and dye transport. The large size of
the dye molecule would certainly reduce the sensitivity (molecu-
lar weight, Oil Red 0 = 276, Sudan IV = 376, data not available on
Black Sudan B, but it must be considerably greater). We can
only conclude that substances giving completely negative data
cannot be evaluated by this technique.

These data would be much more interesting if we were able to
express the amount of penetration per unit time under constant
conditions. Even if we were to assume that the color is a good

162

Journal New York Entomological Society

[Vol. LIII

index of the solvent concentration, the variations in dye concen-
tration in the solvent, variations in distribution in the tracheae,
and errors in estimation of the amount of color outside the tra-
cheae, would prevent any more quantitative scale than good —
fair — poor.

Evaluation of Results. These data present one portion of
the work showing the structural and toxicological similarity of
vertebrate and insect nerves. It has been generally considered in
entomological literature that insect nerves lack myelinization
(e.g., Lindsay and Craig, 1942). Recently it has been shown that
insect nerves do possess myelin sheaths of extreme thinness but
otherwise structurally comparable to those of vertebrates (Rich-
ards,11943, 1944), and that groups of lipids comparable to those
found in the vertebrate brain and representing approximately
40 per cent of the dry weight ,can be extracted from the insect
nervous system (Patterson, Dumm and Richards, 1945). One
would then expect that lipid-soluble materials would tend to
accumulate in the insect nervous system where the sheath lipids
are readily accessible and in a fluid crystalline state. Such ac-
cumulation has at times been assumed without direct confirmation
from entomological data (Hurst, 1943). The presence of a tra-
cheal system facilitates demonstration of such accumulation since
it permits placing series of lipid-soluble materials in intimate
contact with the various tissues without being limited to the
amount that can pass via the blood in solution.

A wide range of organic solvents have been tested by this dye-
penetration technique. These data show beyond question that
lipid-soluble materials do accumulate in the insect nervous sys-
tem, and that the site of this accumulation is correlated with and
so presumably conditioned by the presence of lipid nerve sheaths.
Of course, smaller amounts (without visible dye penetration)
must gain access to other cells and tissues but the quantitative
difference may be great (Richards, 1943). The fact that some
solvents gave negative results by this technique is not necessarily
counter evidence (see preceding section). The fact that smaller
or even equal amounts get into other tissues in a few cases is con-
firmation rather than otherwise since these seeming exceptions are
with either the best penetrants or with water-soluble materials.

June, 1945]

Richards & Weygandt: Fat Penetration

163

Blood injections were also performed. In most cases the re-
sults were negative since the stained solvent remained as immisci-
ble droplets. Solvents which were significantly water-soluble and
not too cytolytic can give selective staining by this route (e.g.,
diacetone). Blood injections are really superfluous beyond show-
ing that some substances which cannot be satisfactorily intro-
duced via tracheae can be via the blood.

From the tracheal injection data it follows that lipid-soluble
substances will tend to accumulate in the insect nervous system
whatever their mode of entry into the body. Hence general toxins
(e.g., toluene) can give the appearance of being selective nerve
poisons simply because it is there that they first accumulate in
toxic concentrations.

As already stated, the technique employed is primarily quali-
tative but if we accept the quantitative expressions in the table
at face value certain generalizations can be made. So far as one
can reasonably suggest from these data it would seem that the
penetration is not any simple function of chain length, molecular
weight or molecular size. The penetration observed is correlated
with oil and fat solubilities rather than cellulose, nitrocellulose,
vinyl resin, etc., solubilities. However, it seems that more than
the oil-water partition coefficient is involved because substances
with similar water solubility do not necessarily give the same
degree of dye penetration. Somehow the molecular configuration
must be involved but the present data do not warrant saying more
than that in these experiments we obtained the best dye penetra-
tion with the benzene-cymene series and some of the cyclic (ter-
pene) hydrocarbons, and the least dye penetration with the
ketones, amines, chlorinated ethers and nitroparaffins.

Incidentally some remarks can be made concerning the proper-
ties of the tracheal walls of mosquito larvae. It is well known that
these walls must be permeable to water, respiratory gases and the
toxic components of petroleum oils. Obviously they must be
permeable to the substances tested here. Black Sudan B is a
poly-azo dye of fair size. Some of the solvents are also of moder-
ate size. Assuming that the solvents are not radically altering
the permeability, it would seem likely that the tracheal walls are
not greatly different in permeability from the membranes used

164

Journal New York Entomological Society

[Vol. Lin

for dialysis. One would rather expect that the permeability in
life would be controlled, then, more by the surrounding cells ; and
that dialyzable substances would pass through.

Several detergents were also tried. Sudan dyes can be solu-
bilized in water with these agents but all those tested gave nega-
tive results. Tests in beakers showed that such solubilized dyes
pass through a regular dialysis membrane too slowly for the
method to be usable in these studies. Detergents had to be omitted
but one (Ter git ol) is deliberately included in the table to repre-
sent the group.

The dye penetration observed is not necessarily correlated with
toxicity, insecticidal efficiency or pathological effects. None of
these are considered in the present paper.

Summary. Dye penetration tests on over 100 assorted organic
compounds indicate that the penetration from tracheae of mos-
quito larvae results in selective accumulation in the central ner-
vous system. Other tissues seldom show visible accumulation
(mouthbrush muscles are the commonest exception). This ac-
cumulation is correlated with and presumably conditioned by the
presence of extremely thin lipid nerve sheaths. It follows that
lipid-soluble substances with low water solubility will tend to
accumulate in the insect nervous system whatever their mode of
entry into the body. General cellular toxins (e.g., toluene) can
thus give the appearance of being selective nerve poisons.

The data are primarily qualitative but do not seem wholly
interpretable in terms of any one property such as oil-water par-
tition coefficients.

The data presented are not necessarily correlated with the
toxicity or insecticidal efficiency of the various solvents.

It seems likely that the permeability of the tracheal membranes
is of the same general order as that of dialysis membranes.

LITERATURE CITED

Bredenkamp, J. 1942. Zur Kenntnis der Wirkungsweise der Kontaktgifte
mit besondere Beriickichtsigung der Insektencutieula. Zts. angew. Ent.,
28: 519-549.

Hurst, H. 1943. Principles of insecticidal action as a guide to drug re-
activity-phase distribution relationships. Trans. Faraday Soc., 39 :
390-412.

June, 1945]

Bichards & Weygandt: Fat Penetration

165

Lindsay, E., and B. Craig. 1942. The distribution of radiophosporus in
wax moth, mealworm, cockroach and firebrat. Ann. Ent. Soc. Amer.,
35: 50-56.

Patterson, E. K., M. E. Dumm, and A. G. Bichards, Jr. 1945. (The
lipids of bee brains.) In preparation.

Bichards, A. G., Jr. 1943. Lipid nerve sheaths in insects and their prob-
able relation to insecticide penetration. Jour. N. Y. Ent. Soc., 51 :
55-69.

Bichards, A. G., Jr. 1944. The structure of living insect nerves and nerve
sheaths as deduced from the optical properties. Jour. N. Y. Ent. Soc.,
52: 285-310.

June, 1945]

Weiss: Insect Food Habits

167

INSECT FOOD HABIT RATIOS OF THE LLOYD-
CORNELL RESERVATION

By Harry B. Weiss

The Lloyd-Cornell Reservation is a quadrangular, uncultivated
area of approximately 81 acres, fifteen miles northeast of Ithaca,
N. Y. The reservation is more than 1,100 feet above sea-level
and its topographic features consist of a pond, streams, ridges
and bogs. This area and its flora and fauna are described in
Bulletin 27, Entomological Series No. 5, of the Lloyd Library of
Botany, Pharmacy and Materia Medica, entitled ‘ ‘ A Preliminary
Biological Survey of the Lloyd-Cornell Reservation,” by mem-
bers of the scientific staff of Cornell University. This was pub-
lished in 1926.

In addition to other animals, over 2,400 species of insects are
listed in this bulletin as occurring on the reservation. These
have been classified in accordance with their predominating fam-
ily food habits. The weakness of such a classification is appre-
ciated and has been admitted in former papers. The terms phy-
tophagous, saprophagous, etc., are used in their broadest sense
and represent apparent rather than actual food habits.

The distribution ratios of food habit types for the reservation
are as follows :

Lloyd-Cornell

Reservation

2

o

br

c3

p!

SP

o

f-l

SP

ct

m

3

o

be

c$

rP

Pi

o

•+J 4J

2 s

Co <D

Pi S
&

e6 f-i

MO)

Qj

P

2,449

40

26

19

13

Regardless of the numerical abundance of individual species,
these ratios are the percentages of the total number, 2,449. As
various species maintain themselves in certain numerical ratios

168

Journal New York Entomological Society

[Vol. Lin

with respect to factors tending to reduce their numbers and as
such relationships are usually normal, these numerical ratios have
been considered as constant.

It is of interest to compare the Lloyd- Cornell Reservation
ratios with those for the entire state of New York* which are
given below.

'3

©
a

02

o
£

New York State 15,343

There is not as much difference between the two sets of figures
as I had supposed there would be. The entire state of New York
embraces different types of vegetation and I had assumed that
the Lloyd-Cornell reservation, a relatively small area with fewer
vegetation types, would show a greater deviation from the state
ratios. On the other hand the Lloyd-Cornell reservation does
not have a uniform type of vegetation and includes water, bogs,
and ridges. Aside from a reduction in the number of phyto-
phagous species the remaining food habit types do not differ
significantly from those for the state as a whole.

* Jour. N. Y. Ent. Soc., vol, 52 : 283-284, 1944.

June, 1945]

Proceedings of the Society

169

PROCEEDINGS OF THE NEW YORK
ENTOMOLOGICAL SOCIETY

Meeting of January 4, 1944

The Annual Meeting of the New York Entomological Society was held on
January 4, 1944, in the American Museum of Natural History, President
Comstock in the chair; nineteen members and visitors present.

The Treasurer ’s report for 1943 was read. Dr. Schneirla reported that the
Auditing Committee had found the Treasurer’s books in good order. Both
these reports were accepted.

The Nominating Committee’s recommendations for the elective officers of
the Society for the year 1944 were read as follows :

President — Edwin Way Teale
Vice President — George G. Becker
Secretary — Annette L. Bacon
Treasurer — Willis J. Gertsch'

Assistant Secretary — Lina Sordillo
Assistant Treasurer — Margaret L. Guy
Editor — Harry B. Weiss

Executive Committee — Edwin Way Teale, Stanley Bromley, William P.
Comstock, William T. Davis, and Herbert F. Schwarz.

Publication Committee — Harry B. Weiss, T. C. Schneirla, and John D. Sher-
man, Jr.

There were no further nominations from the floor. A motion was made
and passed that the secretary cast one ballot and elect the above proposed
officers of the Society for the year 1944.

Mr. Comstock then turned the meeting over to the new president, Mr. Teale.

The following committees were appointed by Mr. Teale:

Program Committee — William P. Comstock, Henry S. Fleming, and Alan
S. Nicolay.

Field Committee — George C. Furness and Chris E. Olsen.

Dr. Bromley presented the following resolution which was adopted:

“Whereas, in the death of Dr. E. Porter Felt on December 14, 1943, the
New York Entomological Society has lost one of its oldest and most highly
esteemed members, and

“Whereas, the New York Entomological Society wishes to express the
sorrow of its members to the family of the decreased and to bespeak their
tribute to one who had been so faithful and revered a member of this Society,
and one so important to the entire broad field of the science of Entomology.

“Be it resolved, That the Secretary be instructed to prepare a letter of con-
dolence and sympathy to the surviving members of Dr. Felt’s family to be
sent to his youngest daughter, Mrs. Elizabeth F. Deane, 51 Beekman Road,
Summit, New Jersey, Administratrix of his estate.”

A motion was made and carried that Mr. Samuel Harriot, who resigned
at the beginning of last year, be, at his own request, reinstated to member-
ship.

170

Journal New York Entomological Society

[Vol. LIII

A discussion of what to do with the old incomplete sets of signatures of
the Society’s Journal took place. Although several years ago it was sug-
gested that the Society dispose of these signatures after the members had a
chance to pick out any that they wanted, members elected since then did not
know of this plan. Therefore, it was announced that no action would be
taken until the February 1 meeting at which time a vote of the members
would be taken concerning the disposal of the signatures as waste paper.

Mr. Comstock spoke of the Relationships of Antillean Butterflies, the mate-
rial for which was taken from his Puerto Bican Survey paper which is in
press (to be published by the New York Academy of Sciences) and from a
joint paper with Mr. Huntington on Antillean Lycaenidae that has just been
published by the Academy.

Meeting of January 18, 1944

President Teale in the chair; 26 members and visitors present.

The Executive Committee reported as follows : At a special meeting of the
Excutive Committee held on January 18, 1944, with Mr. Comstock, Mr.
Schwarz, and Mr. Teale present, Mr. Wm. T. Davis was appointed the So-
ciety’s delegate to the New York Academy of Sciences for the year 1944.
This report was approved by the members present.

Mr. John Noaks, 500 Washington Avenue, Brooklyn, New York, was pro-
posed for membership.

Father Joseph Asmuth of Fordham University spoke on his Termite Obser-
vations in India.

Meeting of February 1, 1944

President Teale in the chair; 13 members and visitors present.

Mr. John Noaks was elected a member of the Society.

After a short discussion it was decided that the old signatures of the
Journal be turned into the salvage drive for waste paper inasmuch as the
Society had an ample supply of complete issues of the Journal. (The Society
received fifteen dollars for them.)

Mr. Alan S. Nicolay told of the work he had done with Coleoptera during
the last year at museums where he studied other collections and discovered
some synonymies, and in the field, especially in New Jersey but also in the
southern states, where he collected some interesting things.

Meeting of February 15, 1944

President Teale in the chair; 21 members and visitors present.

Mr. Gustav Detgen, 303 W. 42nd St., New York City, was proposed for
membership.

Mr. Comstock discussed the matter of a contribution from the Society for
the International Commission on Zoological Nomenclature to assist it in
carrying on its work.

Dr. Daniel Ludwig of New York University spoke on Relative Humidity
and Insect Development. (A summary of his work appeared in Science
Supplement for February 25, page 12.)

June, 1945]

Proceedings of the Society

171

Meeting of March 7, 1944

President Teale in the chair; 18 members and visitors present.

The following report of the Executive Committee was read and approved:

“At a meeting of the Executive Committee of the New York Entomologi-
cal Society held in the American Museum of Natural History on March 7,
1944, Messrs. Teale, Comstock, Davis and Schwarz were present.

1 1 The principal business under discussion concerned the matter of unpaid
dues for membership and unpaid subscriptions to the J ournal. A motion was
made and passed that dues for membership would be remitted to members
of the armed services on application, the treasurer to 'include such a notice
when bills are sent out. As for delinquent members not in the armed forces,
it was agreed that the treasurer would have to consider each case on its own
merits and is hereby empowered to act according to his best judgment. In
regard to unpaid subscriptions to the Journal, it was moved and passed that
the Journal shall not be sent to anyone a year in arrears. This is according
to postal regulations. (That is, if a subscription for the Journal has not
been paid for 1943, it will not be sent to the subscriber in 1944 and after,
until the subscription has been paid.)

“There was also a discussion of electing more Honorary Members of the
Society. No action was taken at this meeting.”

Mr. Comstock reported that he had collected $78.00 from members of the
Society to be sent to the International Commission on Zoological Nomencla-
ture as a gift from this Society. A motion was then made and passed that
the Society give $100.00 to the Commission, said money to be given by mem-
bers of the Society for this purpose.

Mr. G. Detgen was elected a member of the Society.

Mr. Harold Hagan of the College of the City of New York spoke on the
work he had been doing on the embryonic history of the appendages of the
mantid.

Meeting of March 21, 1944

President Teale in the chair; 75 members and visitors present.

Because of the number of visitors present, all business was postponed to
the next regular meeting.

The speaker of the evening, Mr. Rutherford Platt, showed many unusual
and excellent Kodachrome slides illustrating the relationship of insects and
flowers, or “How Plants and Insects Fit Each Other.”

Meeting of April 4, 1944

President Teale in the chair; 16 members and visitors present.

Mr. Weiss proposed Mr. William M. Boyd, 170 Millerick Ave., Trenton,
N. J., for re-election into the Society. As he was a former member, the
By-Laws were waived and he was elected a member.

Mr. Sanford proposed Mr. Raymond Goelbert, 71-15 65th St., Glendale,
N. Y., for membership in the Society.

Mr. Comstock presented the following report of the Executive Committee
of the Society:

172

Journal New York Entomological Society

[Yol. liii

‘ 1 The Executive Committee of the New York Entomological Society, at
their last meeting, after careful consideration of a long list of candidates,
made a selection of eight eminent entomologists, who, as the result of life-
long devotion to our science, seem to the Executive Committee to be worthy,
and whom they now propose as Honorary Members of the Society.

1 ‘ The selections were controlled by several considerations :

1) Eminence in the science of Entomology.

2) Outstanding qualifications as naturalists and scientists.

3) Services to the Society, past and present.

4) Individual diversity of interest in particular families of insects.

‘ ‘ Our by-laws limit us to 12 honorary members. At present we have but
one honorary member, Dr. Leland O. Howard. The eight eminent men now
to be proposed will raise the number of honorary members to nine and in the
judgment of the Executive Commtitee it seems best not to increase this num-
ber for the present. We now propose as Honorary Members of the New
York Entomological Society:

Prof. Nathan Banks
Prof. T. D. A. Cockerell
Mr. William T. Davis
Dr. J. McDunnough
Prof. A. L. Melander
Prof. James G. Needham
Prof. Herbert Osborn
Dr. Robert E. Snodgrass.”

Mr. Comstock then gave a short biographical sketch of each of the above.

The scheduled speaker of the evening was unable to be present. Mr. Geo.
G. Becker told of the work at the plant inspection house at Hoboken. He
had a number of slides showing the work of the Bureau of Plant Quarantine,
and related some unusual situations that the inspectors are called upon to
meet.

Meeting op April 18, 1944

President Teale in the chair; about a hundred members and visitors
present.

Mr. R. Goelbert was elected a member of the Society.

The eight entomologists proposed at the last meeting as honorary members
of the Society were elected.

The Society heard with regret of the death of Mr. George C. Furness, one
of our active members, on April 10, 1944.

Dr. C. B. Williams from England ^vas one of the guests of the evening.

President Teale was the speaker and told of his “Garden for Insects,”
showing his excellent colored motion pictures.

Meeting of May 2, 1944

A regular meeting was held on May 2, 1944, at the IT. S. Inspection House
in Hoboken, N. J., with 19 members and visitors present.

June, 1945]

Proceedings of the Society

173

Mr. Comstock proposed Mr. Gardiner Mulvaney,/ 3 Patchin Place, New
York City, for membership.

Mr. George Becker, head of the Inspection House, conducted the group
around the Inspection House, showing its facilities for the various activities.
He gave a very interesting and thorough explanation of its operation and
answered many questions during the tour.

Meeting of May 16, 1944

President Teale in the chair; 16 members and visitors present.

Miss Lucy Clausen was appointed a member of the Field Committee to fill
the vacancy caused by the death of Mr. Furness.

Mr. Chris Olsen gave details concerning the Entomological Society ’s picnic
to be held at his home in Nyack on June 4. The Field Committee sent
instructions through the mail as to how to reach there. (Due to Mr. Olsen’s
illness, the picnic was held at the home of the Engelhardt’s — Elizabeth
Sherman — in Hartsdale. )

Mr. Gardiner Mulvaney was elected a member of the Society.

A motion was made and passed that the Society donate its complete set
of the Bevista de Entomologia to the library of the American Museum.

Answers were received from the recently elected Honorary Members of
the Society. They accepted with much pleasure the honor thus bestowed
upon them and conveyed to the members of the Society their appreciation.

There was a general discussion by those present of their winter activities.
Mr. Sherman told of his long trip to the West Coast and his interesting
experiences.

Annette L. Bacon, Secretary.

Lina Sordillo, Assistant Secretary.

Meeting of October 3, 1944

A regular meeting of the New York Entomological Society was held on
October 3, 1944, in the American Museum of Natural History, Mr. Teale in
the chair with twenty-two members and visitors present.

Dr. Clarence Goodnight, New Jersey State Teachers College, Jersey City,
N. J., and Mr. B. C. Casselberry, 55 Edgemont Boad, Scarsdale, were pro-
posed for membership.

A letter from the Treasury Department was read which stated that the
N. Y. Entomological Society is exempt from Federal income tax. The Soci-
ety will be required, however, to file a return with the collector of internal
revenue annually.

A communication from the International Commission on Zoological No-
menclature, London, England, expressed thanks for the $100.00 gift which
the Society has contributed to the Commission’s Special (Publications)
Fund.

The resignation of Annette Bacon, as Secretary of the Society, due to the
pressure of outside work was accepted with regret. The Assistant Secretary,
Lina Sordillo, assumed the duties of Secretary for the remainder of the year.

174

Journal New York Entomological Society

[Vol. LIU

The death of Charles Wunder who died on September 21, 1944, and the
illness of William T. Davis were noted with regrets.

The program of the evening was a general discussion on summer activities
and observations of the members.

Lina Sordillo, Secretary.

Meeting of October 17, 1944

A regular meeting of the New York Entomological Society was held on
October 17, 1944, in the American Museum of Natural History, President
Edwin Teale in the chair with thirty-one members and visitors present.

Dr. C. J. Goodnight, New Jersey State Teachers College, and Mr. R. C.
Casselberry, 55 Edgemont Road, Scarsdale, were elected members of the
Society.

A letter from the Zoological Society of London, asking if it would be
possible for us to send them a donation to the Zoological Record Fund again
this year was read. The Zoological Society of London is being very hard
hit at the present time. Donations to the Society for this worthy purpose
may be mailed to or given personally to Mr. Comstock or Dr. Gertsch.

The speaker of the evening was Dr. S. W. Bromley of the Bartlett Tree
Research Laboratories, Stamford, Conn., who spoke on the ‘ ‘ Insect Enemies
of the House Fly,” the text of which will appear in a later issue of the
Journal of the Society. Dr. Bromley discussed hornets, robber flies and
dragon flies as house fly enemies and presented records going back to 1908
when the house fly was more abundant than at present.

Mr. Teale mentioned reading recently an early account of American
pioneers in which it was stated that it was a custom among the people to
place the nest of a white faced hornet in the cabin during the summer so
that the hornets would destroy the flies, the buzzing wasps living in complete
harmony with the human inhabitants.

Lina Sordillo, Secretary.

Meeting of November 21, 1944

A regular meeting of the New York Entomological Society was held on
November 21, 1944, in the American Museum of Natural History, President
Edwin Teale in the chair with sixteen members and visitors present.

Lt. Edmund C. Berkeley, 36 West 11 St., New York 11, N. Y., was pro-
posed for membership.

Mr. Comstock reported that he had collected $85.00 from members of the
Society, $20.00 promised, to be sent to the Zoological Record Found of Lon-
don, as a gift from this Society.

The speaker of the evening was Dr. J. L. Horsfall, who spoke on “A
method for determining leafhopper populations in grape vineyards, ’ } illus-
trated by lantern slides. A special method for determining grape leafhopper
populations was described. Essentially, it consists of selecting certain
leaves, collecting them, separating the leaf hoppers and determining the total
catch by counting the number in an aliquot of the sample. Studies indicate

‘June, 1945]

Proceedings of the Society

175

that it is rapid, reasonably accurate, sufficiently simple to be used by helpers,
adapted to field studies and requires inexpensive equipment. The errors due
to this special manner of counting appear to be no greater than those occur-
ring in duplicate samples from the same vines, and give very small dis-
crepancies in calculating per cent control especially where the efficiency is
75 per cent or greater.

The amount of variation in populations in different parts of the same
vineyard was found to be practically as great as between different vineyards.

Lina Sordillo, Secretary.

Meeting of December 5, 1944

A regular meeting of the New York Entomological Society was held on
December 5, 1944, in the American Museum of Natural History, President
Edwin Teale in the chair with fifty members and visitors present.

Lt. Edmund C. Berkeley was elected a member of the Society.

Mr. Paul Buxbaum, 360 Central Park West, New York City, was proposed
for membership.

Mr. Comstock reported that he had collected $90.00 from members of the
Society, $15.00 promised, to be sent to the Zoological Record Fund of Lon-
don, as a gift from this Society.

Mr. Sherman told of the illness of Mr. William T. Davis.

Dr. J. L. Horsfall reported that the Joint Meeting of the Entomological
Society of America and the American Association of Economic Entomolo-
gists is to be held on December 13, 14 and 15, 1944, at the Hotel New Yorker.

The President and Trustees of the American Museum of Natural History
invited the Society to attend an informal tea, to open the exhibition of Ani-
mals in Miniature by Louis Paul Jonas, December 15, 1944, from 4: 00 to
6 : 00 in the Portrait Room.

President Edwin Teale appointed a Nominating Committee, consisting of
Mr. Schwarz, Chairman, Mr. Sherman and Mr. Huntington.

The speaker of the evening was Mr. E. Irving Huntington, who spoke on
il Ecology of the West Indies and Panama,” illustrated by lantern slides.

Mr. Huntington had made several trips to the West Indies, Panama and
Colombia, and he made numerous photographs, showing the nature of the
country and the collecting conditions to be met with. He discussed the ecol-
ogy of the Greater and Lesser Antilles, pointing out the Greater Antilles
presumably in the past were a connected land mass, while the Lesser An-
tilles were in the form of two arcs, both of volcanic origin; the northern arc
being more ancient and known as the limestone Caribees, while the southern
arc was of more recent volcanic origin. The fauna of the Greater Antilles
is much richer than that of the Lesser Antilles. Mr. Huntington also spoke
of his collecting experiences in Trinidad and in Panama.

Lina Sordillo, Secretary.

Meeting of December 19, 1944

A regular meeting of the New York Entomological Society was held on

176

Journal New York Entomological Society

[Vol. LHP

December 19, 1944, in the American Museum of Natural History, Mr. George
G. Becker in the chair with twenty-one members and visitors present.

Mr. Paul Buxbaum was elected a member of the Society.

Mr. Comstock reported that he had collected $95.00 from members of the
Society, $15.00 promised, to be sent to the Zoological Record Fund of Lon-
don, as a gift from this Society.

President Teale appointed an Auditing Committee consisting of Wm. P.
Comstock, Chairman; E. I. Huntington, and Herbert F. Schwarz.

The paper of the evening was delivered by Mr. Chris E. Olsen, his subject
being Homoptera which he discussed in a broad way, illustrating his talk
with six boxes of local and exotic specimens. He also exhibited a number
of magnificent water color drawings showing, in an enlarged scale, many of
the astounding variations which occur in these insects. These were the work
of the late Mr. Mataush. Of interest to many of the members were Mr.
Olsen’s reminiscence of the old collectors and members of the Society whom
he knew so well. He mentioned the old meetings in George Franck’s store
and those who used to gather there: Torre Bueno, George P. Engelhardt,
Dr. E. B. Southwick, R. P. Dow, Andrew Mutchler and William T. Davis.
He also spoke of many other prominent entomologists whom he had known
and with whom he had corresponded. He estimated that there were some-
where around 4000 species of Homoptera in North America, excluding plant-
lice and scale insects. His main taxonomic discussion concerned the Homop-
tera, particularly the series Auchenorhyncha.

Lina Sordillo, Secretary.

VoL LIII

No. 3

SEPTEMBER, 1945

Journal

of the

New York Entomological Society

Devoted to Entomology in General

Edited by HARRY B. WEISS

a U U i JL u £%

Publication Committee

HARRY B. WEISS JOHN D. SHERMAN, Jr.

T. C. SCHNEIRLA

Subscription $3.00 per Year

Published Quarterly by the Society
N. QUEEN ST. AND McGOVERN AVE.

LANCASTER, PA.

NEW YORK, N. Y.

1945

CONTENTS

The Ennomid Pupa (Lepidoptera, Geometridae)

By Wm. T. M. Forbes 177

Check-list of Psychodidae of Asia and Australia

By William F. Rapp, Jr., and Janet L. Cooper 211

Book Notice , 218

Insects Feeding or Breeding on Indigo, Baptisia

By S'. W. Frost 219

C. V. Riley’s Pirated Work 226

The Mechanism of Oviposition in Phasnicia (Lucilia)
Sericata Meig. (Diptera)

By Cyril E. Abbott 227

The Longevity of Entomologists

By Harry B. Weiss 231

A Laboratory Aid in the Isolation of Entomogenous Fungi

By E. E. McCoy 238

Phalangida from the United States

By Clarence J. and Marie L. Goodnight 239

Ancient Remedies Involving Insects 246

Check-list of African Psychodidae

By William F. Rapp, Jr., and Janet L. Cooper 247

Early Entomological Manuscripts 258

A Pedicel Gall on Trichostema

By James G. Needham 259

NOTICE: Volume LIII, Number 2, of the Journal of
the New York Entomological Society was published
July 17, 1945.

Entered as second class matter July 7, 1925, at the post office at Lancaster, Pa.,
under the Act of August 24, 1912.

Acceptance for mailing at special rate of postage provided for in Section 1103.
Act of October 3, 1917, authorized March 27, 1924.

JOURNAL

OF THE

New York Entomological Society

Vol. LIII September, 1945 No. 3

THE ENNOMID PUPA (LEPIDOPTERA,
GEOMETRIDiE)

By Wm. T. M. Forbes
Cornell University, Ithaca, New York

The following notes were mostly taken some years ago and have
been set aside in the hope of adding more forms. Since there
seems little prospect of doing so, and the characters developed
throw a definite light on the classification of this now tangled sub-
family, it seems best to put them on record.

Our present knowledge of the geometrid pupa is based on two
papers by Edna Mosher (Bull. 111. State Lab. Nat. Hist., vol. xii
(2), pp. 126 to 133, 1916, and Bull. 259, Maine Agr. Exp. Sta., pp.
35 to 58, 1917). Both of these contain keys, but each has genera
omitted «from the other, and the characters used are expressed
somewhat differently, making them a little difficult to use. Alto-
gether only fifteen genera are discussed.

The present notes are not considered sufficient to base complete
keys upon, and so a key is given to a series of somewhat arbitrary
but convenient groups, and in each group the genera (and a few
species) are separated by summary descriptions. About seventy
genera are now known.

From the point of view of classification, by far the most impor-
tant character developed is the type of cremaster, which in the
boarmid series is a heavy bifurcated process or pair of spines,
with the hooked setae extremely weak or absent; while in the
ennomid series proper there is a simpler cremaster, bearing four
pairs of well-marked hooked setae, one pair (corresponding to the

178

Journal New York Entomological Society

[Vol. LIII

two forks of the Boarmines) being usually though not always
longer and heavier than the rest. This makes almost a sharp
separation, and will be useful in the natural division of this great
subfamily, though a few exceptions have already developed: in
the European Heliothea and Tephrina sepiaria the two forks of
the first group are blunted, and each ends in four hooked setas;
Sericosema has hooked setae on a cremaster of Boarmine type, and
in Heliomata, the cremaster is of the bifid type, although it cer-
tainly belongs to the Ennomine series; while Eumera is so de-
generate as to have lost the characteristic structures.

A second character, which is morphologically significant,
though less consistent, is the character of the suture between the
9th and 10th abdominal segments. In the great mass of En-
nominae, this is deeply marked dorsally and down onto the sides,
and usually has a scalloped or notched posterior edge (the ‘‘dor-
sal groove”), while laterally it is extended back across the seg-
ment in the form of a triangular or oblique depression, which we
will call the ‘ ‘ lateral groove. ’ ’ If one is present the other is usu-
ally also, but in a few forms where the dorsal groove is weak the
lateral is undeveloped ( e.g ., Ectropis crepuscularia and Enno-
mos), while in Patalene the dorsal groove has been lost while the
lateral survives. The primitive condition of the two grooves
appears in Caber a (Deilinea) where the dorsal groove is quite
typical, but the lateral is represented merely by a slight out-
pocketing of the posterior side of the suture. We believe this
structure originated within the Geometridae since it is absent in
Brephos, but it is found in Hydriomeninae and Sterrhinas as well
as Ennominae, so its absence in all but the lowest of the Boarmine
complex is doubtless due to degeneration.

Which is the earlier type of cremaster does not appear, since
while the 8-hooked type would be expected to be most primitive,
and occurs in other subfamilies, Brephos and Alsophila have an
exaggerated form of the bifid type.

Other important characters are shown by the side of the fifth
abdominal segment. We shall have to define the spiracular fur-
row and the flange plate a little more sharply than Miss Mosher
did. We will call spiracular furrow a shallow, usually unsculp-
tured and shining depression crossing or lying immediately in

Sept., 1945]

Forbes: Geometric®

179

front of the spiracle, usually bounded by a sharp ridge behind,
and either by a less definite ridge or the flange plate in front.
In Itarne and a few others, instead of a single spiracular groove
there is a series of sharper ridges in front of the spiracle, and in
this case there are often punctures at the bottom of the inter-
vening grooves. We will restrict the term flange-plate to a sharp
vertical ridge lying well in front of the spiracle, preceded by a
deep groove, which is often heavily punctured or otherwise sculp-
tured at the bottom. In many cases this structure lies far to the
front of the segment, and it has not been possible to check for it
in borrowed material. We have assumed it absent in the whole
series centering around Euchlsena and Cingilia, but frequently
the front of the 5th segment was invisible and it may possibly
have been present in one or another genus. The same structures
are found in the Sphingidae.

The segment numbers given are always of the abdomen. The
dorsal ridge lies between the 9th and 10th. In many Boarmines,
where there is a sharp change of sculpture, we have distin-
guished between the 11th segment (specially sculptured) and the
cremaster proper, but in the majority of the Ennomines in the
restricted sense there is no such distinction, and the whole 11th
segment is called cremaster.

Some time ago (Psyche, xxxii, 106-112) I published a summary
grouping of the Ennominae on certain antennal characters. There
I recognized four main series. Of these the group with pectina-
tions naked and basal on the segments, with sense-cones at their
apices, seems a pretty homogeneous group — Neotropical in origin,
and containing Cingilia with its relatives, Sicya, Metanema, and
many South American genera. The corresponding pupae all fall
in the group with 8-hooked cremaster, femur not exposed and
fully developed dorsal and lateral grooves. But the European
genera with this antennal structure are divergent, and Bupalus
at least shows the Boarmine type of pupa with bifid cremaster,
and dorsal and lateral grooves lost. The only available South
American pupa (Myrmecophantes) falls right into this group.
The group with cones on the shaft and two or more setae of type 2
on naked pectinations also fall pretty consistently in this series,
but so also do a few stray genera with scaled pectinations, and the

180

Journal New York Entomological Society

[Vol. Lin

principal neotropical type with simple antennae, so far as we can
judge by their northern relatives.

In the Boarmine series there is a pretty consistent type, shared
by both the Cleora-like and the Macaria-like genera; with bifid
cremaster, femur exposed and no dorsal or lateral grooves; but
the latter are present in several Old World genera, including
Boarmia itself, and are weakly developed in Ematurga and ves-
tigial in Ectropis crepuscularia. On the other hand they are nor-
mally well developed in the Bistonine type, which are definitely
related to the Boarmines, being absent only in Paleacrita, which
is certainly degenerate. It is interesting that in the Boarmid
series this character is more stable than the fovea or the antenna,
for Paraphia with naked antennae, Vitrinella with scaled antennae
and large fovea, and Anacamptodes, with scaled antenna and no
fovea, are all cpiite like Macaria, which has simple antennae and a
vestigial fovea of peculiar type.

One larval character was also noted in this survey : whether the
suckers on the prolegs do or do not interrupt the series of
crotchets. This was the only larval character found which was
both tangible and of some significance, since the humps and spines
on the body vary even in closely related species (e.p., Therina
quercivoraria and textrinaria, which Capps would not even sepa-
rate specifically). The position of the ventral proleg is probably
of considerable value, but proved very difficult to judge, espe-
cially in inflated specimens ; and the setae are often difficult to see.
The character which seemed useful was not whether the crotchets
were continuous or interrupted as bases, but whether the ones in
the middle, opposite the sucker, had lost their terminal hooks. In
several species the hooks in the middle of the line were somewhat
shortened but still perfectly formed; these were considered to
form a continuous series.

KEY TO GROUPS

1. Cremaster deeply bifid, frequently with a pair of terminal spine-like
setae, but with the other setae vestigial or absent (Boarmine series) 2
Cremaster bifid, but each lobe with a quartet of hooked setae of roughly

equal length (Eur.) Heliothea, Tephrina (1)

Cremaster simple or irregular and knobby, with four pairs of well-de-
veloped hooked setae, equal or unequal 6

Sept., 1945]

Forbes: Geometric

181

2. Dorsal and lateral grooves well developed, the former usually deep,

with scalloped posterior margin, sometimes joining anterior end

of lateral grooves , 3

Dorsal groove weak and shallow; lateral groove shallow and inconspic-
uous or (usually) absent 5

3. Four segments dor sally with deep subdorsal pits, each with a pit in

the center; dorsal and lateral grooves connected as in Cabera.

Heliomata (2)

No conspicuous subdorsal pits; dorsal and lateral grooves normal 4

4. Cremaster more or less conical, tapering from a massive conical base

to a rather short slender cremaster proper Biston group (3)

Cremaster reduced to two spines arising separately from body.

Zamacra (4)

Cremaster with a rounded and heavily rugose base, bearing (? if al-
ways) a slender cylindrical cremaster proper Boarmia group (5)

5. Femur concealed or minutely exposed Exelis, Melanolophia, Pseudo-

boarmia, Paraphia, Paleacrita (6)

Femur well exposed Epimecis, Bupalus, Itame, Ematurga, Macaria,

Ectropis, Anacamptodes, Vitrinella, Glena, Physostegania (7)

6. Dorsal and lateral grooves normal, the latter oblique and meeting the

dorsal at an angle; the dorsal groove rarely obsolescent ( Pero ) when

the crem astral hooks are 2-6 7

A single (dorsal) groove extending two thirds way around the segment,
with a single strong dentation to represent the rudiment of the

lateral groove Cabera ( Deilinea ) (8)

Dorsal and lateral grooves vestigial and ill defined or absent; cremaster
with all hooks short, and the two longer pairs subequal 15

7. Fore femur exposed 8

Fore femur concealed 9

8. Lateral groove small, limited to anterior part of 10th segment, body

with regular puncturation (except in a western species of Pero) ;

exposed part of fore femur small Pero group (9)

Lateral groove large, frequently extending forward onto 9th segment
or backward onto cremaster ; puncturing confused and largely modi-
fied into vermiculate sculpture Sabulodes group (10)

9. Skin smooth or with fine inconspicuous ridging; white or green 10

Skin rough and rugulose or punctured ; brown 11

10. Cremaster with second pair of hooks about as thick as long pair, both

short, but much longer than the more basal hooks; tongue about

2/3 length of wings Xanthotype (11)

Cremaster with two liooks much longer and stouter than the other six;
tongue about as long as wings Cingilia group (12)

11. Cremaster with two pairs of heavy and somewhat longer, and two pairs

of small setae 12

Cremaster with two setae much longer than the other six 13

12. Spiracular furrow absent, but flange-plate present, preceded by a deep

groove Abraxas (13)

182

Journal New York Entomological Society

[Vol. LIII

Spiraeular furrow represented on 5th segment by a series of ridges
separated by series of punctures; small and slender species.

Nematocampa (14)

No trace of spiraeular furrow or flange-plate, the front of 5tli seg-
ment falling off sharply but smooth; very stout, general sculpture

of sparse irregular puncturing Selenia (15)

13. Antenna strongly roughened, in the form of five or six rows of tuber-
cles (one of each row on each segment) Nacophora (16)

Antenna smooth or less extensively sculptured 14

14. Prothoracic callosity prominent Euchlcena, MetarrJianthis (17)

Prothoracic callosity minute and inconspicuous or obsolete.

Therina, Caripeta, etc. (18)

15. Pupa with dense secondary hair Hygrocliroa ( Pericallia ) (19)

Pupa with small primary hairs only 16

16. Antennae with four rows of tubercles, one to a segment in each row;

tongue half as long as legs; cremaster long, pyramidal, with the
basal two pairs of hooks rising far out on it ; preceded by two pair

of raised tubercles Ennomos (20)

Antennae not or less regularly sculptured, normally with only the seg-
ments marked; tongue full length; cremaster normally short and
rough; no raised conical tubercles 17

17. All legs nearly covered ( $ ) ; a series of punctures before and behind

suture 9/10, to indicate the dorsal groove; mesotlioracic callosity

(so-called spiracle) obsolete Patalene (21)

Legs normal; posterior segments without punctures; mesothoracie callo-
sity distinct 18

18. Mesothoracie callosity depressed, indicated mainly by its sharp posterior

edge, giving with the posterior edge of the prothorax almost ex-
actly the impression of a spiracle Opisthograptis ( Bumia ) (22)

Callosity very large, black and chitinized Plagodis (23)

The following supplementary characters, while not always defi-
nitely contrasted, should help in determining some of the pupae,
and may throw a little further light on relationships. They are
listed not in a natural arrangement, but in the order of the key.

1. Heliothea discoidaria (Europe, C.U.). Cremaster in the
form of two rounded but irregular prominences, with a cleft
between them, each bearing four hooks, as in Tephronia, which
is not otherwise related, the most terminal of the four hooks
longer but more slender than the subapical pair. Dorsal groove
well marked, but smooth, suture-like, the lateral groove similar
and slender, branching off from it at an angle and running back
two thirds the width of the segment, the preceding suture also
accented and the segment between raised; sculpture smooth and

Sept., 1945]

Forbes: Geometridje

183

confused, but surface dull, even on terminal segments ; segment 7
with a raised, but rounded, longitudinally rugose keel running
around most of the segment just in front of the spiracle (setae iii
and v), segments 5 and 6 with weaker keels at sides; spiracles
raised. No spiracular furrow or flanges ; callosity well marked,
but pale; appendages mostly normal, but tongue less than half
as long. Femur covered.

The pupa is as queer as the corresponding moth, which belongs
in the waste-basket called ‘ ‘ CEnochrominae. ’ ’ The antenna of
imago is also queer; the pectinations are naked and basal, with
short single type 2 setae, and vestigial abnormal cones on the tips
of a few pectinations.

1. Tephronia sepiaria (N.M. ; Europe). Cremaster bifid, each
half with a little tuft of subequal setae. Dorsal and lateral groove
absent. Sculpture: body punctured, including 9th segment of
abdomen, but not tenth. No spiracular furrow, the front of 5th
segment of abdomen rather more finely but very irregularly punc-
tured ; mesothoracic callosity pretty strong ; legs and tongue nor-
mal, mid-legs short and small.

In this genus, and in all so far as examined with the bifid type
of cremaster (except the following) the series of hooks on the
prolegs is complete.

2. Heliomata cycladata (N.M.). Dorsal groove deep, running
into a short conspicuous lateral groove as in Cabera. Callosity
obscure. Segment 5 with a strong flange-plate in front of spiracle,
the anterior face more finely punctate. Pore femur minutely
exposed or covered. Antenna nearly as in Sabulodes.

Larval hooks interrupted. Imago with simple, prismatic an-
tenna. This genus is as queer in the pupa as in the adult. The
pattern and genitalia suggest a close relationship with the Euro-
pean Abraxas ( Lomaspilis ) marginata, but the pupa of margi-
nata is not available and there is no resemblance to true Abraxas.

3. Amphidasis cognataria. Cremaster minutely bifid at apex
and acutely shouldered near base, longish and rather rugose.
Dorsal groove strong and toothed as usual, lateral strong. Sculp-
ture punctured, especially dorsally, becoming heavier on 6th to
8th segments and absent on 9 and 10. A5 with strong prespiracu-
lar flange with the groove in front. Callosity strong. Tongue a

184

Journal New York Entomological Society

[Vol. LIU

little more than f , .separating the fore legs ; middle legs slender
and normal; antenna very wide basally but not 5-tuberculate ;
femur covered. (See also Mosher, ’16, 131.)

Antenna of imago with pectinations naked, cones on shaft.

3. A. betularia (Dampf coll., C.U.). Similar, the shoulders on
cremaster acute.

3. A. strataria (Dampf coll.). Cremaster with shoulder obso-
lescent. Spiracular furrow very broad and shallow, with the
spiracle in its center; groove in front of flange-plate heavily
barred.

3. Biston hirtaria (Dampf coll.). Similar to Amphidasis; cre-
master more conspicuously but shortly forked, its shoulders sharp
and spine-like. Puncturation present also on anterior side of 9th
segment. Spiracle lying in an emargination of posterior side of
spiracular furrow ; groove in front of flange-plate smooth. Spi-
racular callosity of mesothorax weak. Antenna with two sub-
marginal rows of tubercles, like the two outer rows on Nacophora
or Ennomos. Front strongly rough and bulging below; tongue
almost full length.

Antenna of imago like Amphidasis.

3. Nyssia zonaria (Dampf ). Cremaster obscurely bifid. Punc-
turation as in Biston, the general surface smoother. Flange-plate
and its preceding groove on segment 5 smooth, much shorter.
Spiracular furrow like Biston. Callosity absent. Front and an-
tenna smooth. Otherwise like Biston.

3. Coniodes plumigeraria (N.M.). End of abdomen reduced,
the segments 9-11 together only as long dorsally as 8, heavily
rugose, with unspecialized dorsal groove and lateral groove a
mere pit. Sculpture punctate, heavily chitinized. Spiracular
furrow strong and double, with a more prominent ridge (the
flange-plate) between ; spiracles not produced (unlike Paleacrita).
Callosity obscure; tongue not reduced; mid tibia almost covered
by the very wide (male) antenna; tongue full length (unlike
Amphidasis, etc. ) .

(3) Phigalia not examined. It should presumably be close to
Coniodes, but with more moderate antenna, and probably short-
ened tongue.

3. Erannis tiliaria (Mosher, ’17, 57). Shoulders on cremaster
marked; dorsal groove finely serrate; callosity strong, tongue

Sept., 1945]

Forbes: Geometrid^e

185

nearly full length. Flange-plate of 5th segment characteristic;
a deep narrow elliptical groove at anterior edge of segment, with
sharp black edges; spiracular furrow a smooth glossy band, and
a slight vertical ridge through the spiracle.

3. E. aur antiaria and marginaria (Dampf coll.) essentially the
same, the thick base and thin apical portion of cremaster even
more sharply set off. The dorsal and lateral grooves are very
weak.

(3) Paleacrita belongs to this group, but as it has wholly lost
the dorsal and lateral grooves will be discussed below.

4. Zamacra excavata (Japan, N.M.). Cremaster reduced to
two separate spines arising from body. Body smooth; ( flange-
plate not seen). Dorsal groove rather weak and not toothed;
lateral weak, on front part of 9th segment; callosities indicated
by black color only. Head obscurely 3-tuberculate in front.
Antenna with two rows of tubercles, like Biston; tongue full
length ; femur covered.

5. Phthonosema tendinosaria (Japan, N.M.) . Cremaster stout,
bifid and almost shouldered; dorsal groove slender, but distinct
and toothed, lateral present. Spiracular furrow invisible, appar-
ently absent. Callosity small and obscure. Fore femur exposed.

A curious form, one of the few Old World types with naked
antenna like Paraphia and the Bistonids, but otherwise a Boarmia.
It probably represents the immediate point of origin of the latter.

5. Ematurga atomaria (Dampf) . Cremaster slender and cylin-
drical, arising out of a heavily punctate shortly conical base
(bifid no doubt — all our specimens are broken). Dorsal groove
slender but well marked, continuous far down on sides, a little
scalloped; the lateral grooves forming branches slanting 45° up
and back. Body heavily punctate, the 9th and 10th segments
impunctate but dull ; no spiracular furrow of cleorid type, but a
broad and vague impunctate zone, bounded in front by a sharp
flange-plate preceded by a row of deep crowded punctures (essen-
tially as in Ampliidasis). Callosity in the form of an indefinite
rough spot. Femur covered.

5. E. faxonii (N.M.) is the same so far as my notes extend.
Evidently this genus is not close to Itame, as one might think,
but the naked pectinations of antenna are consistent with a place

186

Journal New York Entomological Society

[VOL LIII

derived from the same early Boarmia type that gave rise to the
Bistonids ( i.e the following).

5. Boarmia roboraria (Dampf). Cremaster (?), apparently
deformed in the only specimen at hand. Dorsal and lateral
grooves strong, the former with many fine dentations. Sculpture
normally punctate, with 9 and 10 smooth and glossy; spiracular
furrow fairly defined, bounded by a ridge through spiracle, and
a flange-plate as in Ematurga. Callosity strong; femur covered.
B. consortaria is essentially the same, the cremaster having a long
slender free portion.

For Boarmia repandaria see Paraphia group.

4 or 5. B. ( Deileptenia ) abietaria (Europe, C.U.). Cremaster
thick to its tip, conical, the two spines arising separately. Spi-
racular grooves distinct, with a ridge just behind the spiracle, and
a front ridge which may represent a flange-plate but is only pre-
ceded by a band of deep but sparse punctures. Sculpture other-
wise of light sparse puncturing, only on anterior parts of seg-
ments.

4, 6. Paleacrita vernata (G.U. ; Mosher, ’17, p. 55). Cremaster
with branches widely divergent and recurved, usually broken off.
Dorsal groove rather rudimentary, even less developed than in
Erannis, the lateral groove in the form of a slight branch of it;
sculpture punctate, strong, covering part of 9th segment and
venter of 8th in male but not female ; flange-plate and furrow in
the form of an elliptical pit, less than twice as high as wide ; cal-
losity fairly well developed ; head with lower part of front bulg-
ing (like Biston), all roughly sculptured, the eye as rough as any
part except for the crescent separating the ‘‘chitinized” and
‘‘glazed” portions. Tongue as long as legs and wings which are
fully developed in both sexes, the female wings showing a normal
venation.

On account of the rudimentary dorsal groove this may be
sought either in the Biston or the Paraphia groups ; in either case
the high raised cones bearing the spiracles are distinctive.

6. Prosopolopha jourdanaria (Europe, C.U.). Cremaster with
basal f rugose in the form of a flattened cone ; terminal portion
smooth, slender and bifid. Dorsal groove rudimentary, a merely
slightly wandering groove, lateral absent. Sculpture densely

Sept., 1945]

Forbes: Geometric

187

punctate, including segment 8, 9 smooth, 10 faintly longitudinally
ribbed above ; spiracular furrow well marked, bounded by a ridge
through the spiracle, and the very strong and sharp flange-plate ;
sculpture in front of flange-plate of dense irregular puncturing.
Callosity well marked, pubescent; appendages normal; antenna
rough with well marked segmentation ; femur minutely exposed,
but covered with glossy varnish, as structures are likely to be
which are normally covered.

Larva with hooks of prolegs continuous. Antenna of imago
distinctive, with three large subterminal setae of type 2 on naked
pectinations, and cones on shaft. By the antenna this genus
would be Ennomine, by pupa Boarmine. The build is stout, and
connections appear to be with the Australasian fauna. A curious
beast. The pupa is essentially Paraphia-like.

6. Nychiodes lividaria (Europe, C.U.). Essentially like Proso-
polopha; cremaster thick clear out to the point of forking, with
.angles half way out like Amphidasis. Segment 10 heavily and
conspicuously longitudinally ridged. Femur covered, callosity
stronger.

6. Exelis pyrolaria (N.M.). Cremaster transversely ribbed at
base, the rest very slender without the thickened bases of the pre-
ceding series. No trace of dorsal or lateral grooves. Sculpture
lieavily punctate, including dorsum of 9th segment (more than in
any of the preceding) ; no spiracidar furrow. Callosities obscure.
Mesothorax rugose, spiracles not raised. Tongue only f length of
appendages.

Imago with antenna scaled, the cones on shaft.

6. Paraphia triplicipunctaria (Rupert) . Cremaster bifid, stout,
less shouldered than usual, smooth, but 11th segment rugose.
Sculpture lightly punctate, running into transverse grooving, 9th
and 10th segments smooth, 8th somewhat pitted. Spiracular
furrow broad, smooth, conspicuous but ill defined, as in Vitrinella
with a faint central ridge, but unlike Melanolophia. Callosity
prominent, pubescent. Flange-plate strong, the groove coarsely
pitted.

Differs from Protoboarmia only in the covered femur, from
Melanolophia mainly by the broad “cleorid” spiracular groove.
Imago with antennal pectinations naked, cones on shaft, as in
both Melanolophia and Protoboarmia, etc.

188

Journal New York Entomological Society

[Vol. Liir

6. Melanolophia (N.M.). Cremaster without shoulders. Sculp-
ture punctured, segment 9 smooth. Spiracular furrow a shallow
fold curving around the spiracle. Callosity small, neat, pubescent.

6. Pseudolooarmia porcelaria (N.M.). No really distinct
shoulders on cremaster. Spiracular furrow strong, its anterior
boundary closely parallel to flange-plate at anterior edge of seg-
ment, its posterior running through spiracle, neither quite so
sharp as in Vitrinella. Callosities strong, setulose. Femur mi-
nutely exposed; antenna extremely narrow in male considering
the width of antenna in the imago.

7. Epimecis hortaria (N.M.). Cremaster a simple spine, defi-
nitely shouldered at the base. No dorsal groove, the suture being
normal, with a suggestion of intersegmental membrane, and 9th
segment lightly punctate. Lateral groove faintly suggested be-
hind. No spiracular furrow of Cleorid type, but 5th segment
with a flange-plate like that of Amphidasis. 6th and 7th seg-
ments with ridges practically tangent to spiracle. No trace of
callosity. Femur narrowly exposed (J tibia).

7. Bupalus piniarius (Dampf). (Determination doubtful.)
Cremaster a short cone, ending in two spikes; the basal half
heavily rugose. No dorsal groove, the lateral faintly visible in
favorable light. Sculpture heavily punctate on abdomen, the 9th
and 10th segments smooth and shining, and thorax nearly smooth.
No spiracular furrow or flange-plate, but the segment falling off
sharply from a ridge close to spiracle. Callosity strong.

The antenna of the imago is as in Melanolophia and Paraphia.
This pupa seems a little closer to Itame, but may be misdeter-
mined.

6 or 7. Boarmia repandaria (Dampf). Cremaster with two
separate spines or a short fork from a thick rugose base (varying
individually). No dorsal or lateral grooves. Sculpture of fine
sparse punctures on a glossy ground. Spiracidar furrow vague,
impunctate ; bounded in front by a strong flange-plate which is
preceded by a band of dense coarse puncturing. Callosity neat
but small. Femur very narrow or obsolescent, but consistently
visible in three specimens.

7. Ectropis crepuscularia (Dampf and N.M.). Cremaster
tapering sharply, bifid, the base rugose as usual. Dorsal groove

Sept., 1945]

Forbes: Geometric®

189

well marked but not strong, very finely dentate on posterior edge,
as in Erannis; lateral groove obsolescent. Sculpture densely
punctured ; 9th and 10th segments clear ; spiracular furrow broad
and vague, bounded in front by a long elliptical fovea sharply
bounded by a chitinized black edge as in Erannis. Callosity small
but well marked.

This is surprisingly like Erannis, especially considering the
rather similar antenna, but can be distinguished by the well ex-
posed fore femur.

7. Vitrinella pampinafia (C.U. andN.M.) (Mosher, ’17, p. 50).
Characters typical for the group, with slender tapering bifid
cremaster, no dorsal or lateral groove ; sculpture of moderate
puncturation ; antenna with segments marked but not otherwise
sculptured, femur exposed, tongue long. Spiracidar furrow broad
and shallow, lying between a ridge behind the spiracle and the
well-marked flange-plate at anterior edge of segment, bisected by
a slight but sharp ridge.

Moth (as in the following) with pectinations scaled and cones
on shaft.

7. Anacamptodes sp. (a female, apparently of fragilaria)
(C.U.). Closely similar to the preceding; cremaster more deeply
forked ; spiracular furrow bounded in front by a ridge passing
through spiracle, undivided. Callosity stronger. A. humaria
appears to be identical.

7. Glena cribrataria (N.M.). Closely similar (fifth segment of
abdomen destroyed). Skin more roughened between the punc-
tures, Spiracle of 8th segment marked by a decided tubercle
(indicated in our specimen of A. fragilaria, but not distinct in
V. pampinaria) .

7. Macaria liturata (Dampf). Most features quite like those
of the preceding three genera, but with no spiracidar furrow nor
flange-plate, there being merely a smooth convex band in front
of the spiracle, preceded by a vague band of close puncturation;
puncturing of posterior segments normal, 8 as heavy as any, 9
and 10 smooth, and base of cremaster rugose.

7. M. eremiata and irrorata (N.M.). Abdomen with segments
9 and 10 slightly rougher ; a deep strong spiracular furrow
flanked by two closely parallel ridges, and with a few punctures
in front of them ventrally.

190

Journal New York Entomological Society

[Vol. LIII

7. M. pervolata (N.M.). Spiracular groove less definite, the
posterior ridge distinct, but anterior ridge partly fused with it,
running into punctures above and below, and preceded by several
irregular ridges.

7. M. punctolineata. Almost like M. liturata, spiracular fur-
row with the posterior ridge only distinct, in front of the spiracle,,
the anterior face with coarse punctures in vertical series as in
Itame.

7. Itame brunnearia (Dampf) and occiduaria (N.M.).- Cre-
master stout and shortly bifid, not really shouldered. No dorsal
or - lateral grooves. Abdomen punctate, including 8 but not 9
and 10. No spiracular furrow , the front of segment falling off,
with coarse punctures in vertical series instead of. the scattered
ones on rest of segment. Callosity smallish, well set off. Femur
well exposed, appendages normal. No flange-plate at all.

7. I. ribearia (C.U.) closely similar, the puncturation on an-
terior slope of segment 5 denser. (See also Mosher, ’17, p. 52.)

7. /. anataria (Mosher, ’17, p. 52) has distinct spiracular fur-
rows.

7. I. wavaria (Europe, C.U.). Similar but the spiracular fur-
rows very well developed, a zone of 4 or 5 close, sharp ridges with-
out punctures between them, the most posterior interrupted
opposite the spiracle, and in front of these, confused and broken
ridges with rows of punctures intervening.

In this group the antennal segments are strongly marked, some-
times with deep puncturing, but without other sculpture ; the
second and third spiracles are partly covered, and fourth fully
exposed. In the specimens of ribearia and brunnearia before me
the second spiracle is less covered than the third.

7. In Physostegania pustularia (Mosher, ’16, p. 130) there is
a deep spiracular furrow with chitinized edges and with punc-
tures in its bottom, unlike the true Itames.

7. Melanchroia chephise (Florida, Hansbery, C.U.). Cremaster
smooth, a nearly regular flattened cone, shortly bifid, with minute
points on side where the dentations of Amphidasis are located.
No dorsal or lateral groove. Sculpture of regular fairly dense
puncturation, as dense on 8th segment as any (except for the
usual smooth posterior stripe) ; 9 and 10 without punctures,

Sept., 1945]

Forbes: Geometrid.®

191

glossy. No spiracular groove, flange-plate short, curving for-
ward at ends to enclose a long-elliptical* area, which is heavily
punctate at bottom, and reaches to front edge of segment. Cal-
losity very heavy, setulose. Appendages normal, smooth and
glossy, tongue full length ; femora exposed ; antenna with segmen-
tation faintly marked.

Larva with sucker interrupting hooks (unlike all with bifid
cremaster except Heliomata). Imago with scaled pectinations
and no visible cones, extremely heavily chitinized. Most of the
pupal characters point to the group of Vitrinella and Anacamp-
todes, and the antenna agrees, but the larval sucker as well as
appearance are wholly wrong. The genitalia are also unique
(Rupert).

8. Cabera erythremaria (N.M.). Cremaster with two longer
hooked setae (unlike the preceding genera). Dorsal groove dis-
tinctive, narrow, as a whole even, but neatly notched on its pos-
terior edge, the lateral groove represented by an oblique lateral
notch hardly if at all deeper than the others. Sculpture of nor-
mal punctures, segment 8 as w^ell as 9 and 10 practically smooth ;
fifth segment rapidly falling off in front, but punctured like the
rest (no spiracular furrow, etc.). Callosity easily seen, but not
raised, femur concealed, other appendages normal.

Larva with sucker interrupting hooks.

Imago with antenna with scaled pectination and cones on shaft.

This is the only genus studied of a rather substantial group,
that may perhaps be ancestral, as the condition of the dorsolateral
groove suggests. There may possibly be some connection (through
South American types) with Heliomata, which also has the larva
with a sucker and a similar dorsal groove. The antenna is very
similar to Apicia (Gaberodes), but this seems to be a case of
parallelism, since other structures diverge. The remaining genera
form the series with 8 hooks on the cremaster, and in them many
genera have the hooks of the prolegs interrupted by the sucker
(except for rudimentary bases) and have the sense cones of the
antenna at the tips of naked pectinations, the combination mark-
ing a distinctive Cingilia group, which is primarily South Ameri-
can.

8. C. pusaria (Dampf). Almost identical. The antenna shows

192

Journal New York Entomological Society

[Vol. LIII

a series of tubercles at the anterior (ventral) edge, marking the
inner series of pectinations.

9. Pero honestarius (N.M.). Cremaster short and rugose, with
two setas much longer than the others. Dorsal groove merely an
accented suture; lateral groove deep, narrow, set at 60°, visible
on 10th segment only, which is long, and ridged behind. Body
regularly punctate, segment 5 not modified, merely falling off
abruptly in front, segments 8 and 9 smooth. Fore femur well
exposed, mid tibia reaching about to front of eye.

Imago with simple antenna.

9. Pero sp. (an undetermined western sp. in Nat. Mus.). Simi-
lar, but sculpture tending to run together into grooves, and dorsal
groove fully developed.

9. Stenaspilates species (in Nat. Mus. from Texas). Dorsal
groove very distinct, sculptured ; lateral groove short, on anterior
part of 10th segment ; fore femur minute.

Pectinations of imaginal antenna naked, cones on shaft.

This tribe may perhaps be distinguished by the regular punc-
turation and small lateral groove. Sericosema agrees on these two
- characters, but has a different 5th segment.

9? Sericosema (N.M.). Cremaster bifid but with recurved
hooks and at least one more pair of hooks besides. Dorsal groove
coarsely notched ; lateral groove well marked, but opening for-
ward into dorsal groove; sculpture coarsely punctate, including
fore portion of 5th segment ; which has a strong flange-plate with
a row of deep pits in front of it; segments 8 and 9 smooth, 10
becoming rough behind, less well set off from cremaster than
usual ; callosities well indicated, but not forming a true tubercle ;
femur exposed; antenna narrow, with a double series of tubercles
toward base ; mid-leg to front of eye, unlike Apicia.

A synthetic type that seems to connect the Boarmine series with
Cabera, but also resembles Pero. By the antenna of the imago it
appears to connect through Ixala to Cabera. The resemblance to
Pero is probably purely superficial.

10. The following group is characteristically neotropical, where
there are many closely related genera. In the imago, one can dis-
tinguish a series with two well-formed accessory cells from one
with more varied venation, but the pupal characters do not cor-

Sept., 1945]

Forbes : Geometrid^e

193

respond. In general the present (Sabulodes) group has simple
antennge and irregular radial system, while the “Therina” group
has the double accessory cell and characteristic antenna with
cones at the tips of naked pectinations. But Abbotana, with the
concealed femur of Therina, has the venation and essentially the
antenna of Sabulodes, while Tetracis, with essentially Sabulodes
pupa and antenna, has the double accessory cell. The genera with
thin glossy nearly unsculptured pupa are not a homogeneous lot,
Xanthotype being near to Euchlsena, Cingilia, etc., to Therina.
Phrygionis is inserted purely because it keys here. It of course
represents the Palyadinge. The pupa which Mosher reported as
Philobia enotata (’16, p. 129) obviously belongs to this group,
but I cannot guess what it really is; it is extremely improbable
that a member of the enotata group (which are normal Macarias)
would have such a different pupa from M. liturata.

10. Sabulodes transversata (N.M.). Cremaster massive, with
weak spines. Dorsal groove normal, lateral very wide in front,
set at 30-45°, encroaching on 9th segment. Scidpture rugulose,
9 smooth, 10 longitudinally ridged, 11 with 4 ridges, 8 with en-
larged tubercle ; middle segments with a long heavy vertical ridge
tangent to spiracle, but no spiracular furrow. Callosity obscure ;
head with a transverse ridge ; femur narrowly exposed (^ tibia),
mid-leg extending beside eye ; antenna with two transverse
grooves to a segment. In other ways much like Abbotana.
(Mosher, ’17, p. 47).

Larva with hooks interrupted; imago with simple, prismatic
antenna.

10. S. forficidaria (N.M.). Sculpture pitted, tending to form
spiracular ridges on 5th segment, ridges on 5 to 7 less prominent
than in transversata; 8 with tubercle i slightly enlarged, the
pitting weak ; 10 irregularly marked, only, not really sculptured ;
11 irregularly rugose.

10. S. caberata. Cremaster longer, with lateral groove con-
tinued onto it; sculpture almost all of vertical ridges, with no
especially high one, smoother than the two preceding ; no enlarged
tubercles ; a keel on vertex.

McDunnough would separate this one species as Sabulodes (of
course with its South American congeners) calling the residue

194

Journal New York Entomological Society

[VOL. LIII

Antepione, but I cannot see so very deep a difference. The longer
cremaster and different sculpture may possibly be generic.

10. Tetrads lorata (N.M.). Cremaster very long, four-ridged
like transversata, with one pair of hooks especially long and
straight. Lateral groove not extending onto cremaster; sculpture
rugulose, practically smooth on 9 and anterior part of 10. Cal-
losity rather stronger; front with a slight suggestion of a knob,
and a slight crest on prothorax.

The pupa show much closer connection to Sabulodes (and
Antepione) than would be supposed from venation and genitalia.
(Studied by Mosher, ’16, but not discriminated from trans-
versata.)

10. Apicia confusaria (N.M.). Similar to Sabulodes, except:
cremaster transversely rugose ; lateral groove less longitudinal,
not invading segment 9. Sculpture punctured and grooved, 9
smooth and 10 more finely longitudinally strigose, especially
behind ; mid-leg falling far short of fore leg at front end (at least
in male). No spiracular groove or distinct keels on segments 5
to 7 ; male antenna twice as broad, with a tendency to pitting
between the pectinations.

Larva with sucker interrupting hooks. Imago with scaled pec-
tinations and basal cones. I have examined four genera of the
Sabulodes group with pectinate antennae, and each differs widely
in structure, so it would appear that pectinations have been secon-
darily developed. In most characters Apicia has antennae like
the Cabera group. Halesa, Patalene and Gynopteryx are each
quite different.

10? Phrygionis argenteostriata (N.M.). Cremaster small.
Dorsal groove strong, lateral nearly longitudinal and not disturb-
ing 9th segment at all; sculpture pitted, fading out behind' 10
rough and heavily chitinized ; spiracular furrow not visible
(probably absent) . Callosity large and brown, but not very high.
Fore femur wide, at least i tibia; antenna narrow, with faint
sculpture ; only fore leg extending past eye ; tongue full length ;
mandible projecting and heavily chitinized, labrum more plainly
than usual 2-segmented (i.e., clypeus and true labrum).

Antennae of imago with scaled pectinations, a few cones on the
shaft, and single strong terminal setae.

Sept., 1945]

Forbes : Geometrid^e

195

11. Xanthotype sospeta (C.U.). Cremaster short, thick, very
rugose and heavily chitinized, with short hooks, the second pair
as thick as the terminal and nearly as long. Dorsal groove with
middle notch much deeper, the side notches numerous and irregu-
lar in depth; lateral groove oblique, conspicuous. Body thinly
chitinized, glossy, wholly without sculpture, the 10th segment
more heavily chitinized but also smooth and glossy ; head darker
but not more heavily chitinized. No spiracular furrow or flanges.
Callosity very large, overhanging behind. Antenna rising to a
slight rounded keel, the segments strongly marked. Tongue only
f appendages. (Mosher, ’16, 129).

Larva without a trace of interruption in the series of hooks;
imago with antenna like Euchlasna, pectinations naked, cones
basal. A close relative of Euchlagna and Angerona in spite of the
different appearance of the pupa.

12. Cingilia catenaria (C.U., Mosher, ’17, p. 43). Cremaster
short, continuing the line of the 11th segment, with the two ter-
minal hooks much longer than the rest; dorsal groove normal,
with middorsal notch a little deeper; lateral groove strong and
oblique. Sculpture of fine vertical strigulation, without any
punctures, the area in front of the spiracle of 5 like the rest, the
ninth and tenth segments smooth and 11th dorsally with deep
longitudinal vermiculation. No distinct callosity. Third spiracle
partly covered.

Larva with hook-series interrupted. Antenna of imago with
terminal cones on naked basal pectinations.

The ventral third of the “glazed eye” in this species shows
striking and regular facetting, and certainly looks as if it should
be functional. This may be correlated with the extremely open
mesh-type of cocoon.

12. Nepytia canosaria and phantasmaria (C.U.). Practically
identical ; the dentations on the dorsal groove perhaps on the
average fewer and coarser. N. canosaria from Ithaca, lacks the
pattern of black dots so conspicuous in Cingilia, but in N. phan-
tasmaria, from Vancouver, they are present, though duller and
more diffuse than in catenaria.

Larval legs not examined, but doubtless as in Cingilia. Pattern
strikingly different. Antennce of imago as in Cingilia.

196

Journal New York Entomological Society

[VOL. LIII

12. Myrmecophantes valens Thierry-Mieg (El Campamiento,
Colony of the Perene, Peru, C.U.). Cremaster a single mass with
11th segment, black, rugose and shapeless, with hooks shorter and
much heavier than in Cingilia; dorsal groove with about 9 regular
dentations, lateral deep, oblique, triangular, not really reaching
posterior edge of segment. Sculpture mostly weak, sparsely
punctured on a lightly chitinized body; anterior halves of seg-
ments 5 and 6 with strong sharp vertical striation, bounded by a
higher vertical keel through spiracle; 7th segment with corre-
sponding portion slightly rugose and lacking the usual punc-
tures; callosity obsolescent. Pattern of black dots much as in
Cingilia with minor differences; e.g., the black on the antenna
takes the form of a solid ventral line instead of a double row of
dots.

I reared this from a cocoon found in the field and do not know
the larva ; the cocoon is of coarse open meshes, like catenaria, but
coarser. Antenna of imago of Cingilia type, more heavily chitin-
ized. This genus represents a large South American group, evi-
dently very close to Cingilia, Nepytia, etc., with which their
antenna and venation also agree essentially.

12. Philtrcea (N.M.). Similar to Cingilia, the sculpture less
markedly transverse ; tongue only half length of wing ; cremaster
not seen.

12. Sicya macidaria (Mosher, ’17, p. 44) (not seen). Similar
to Cingilia; iridescent green or yellow. 11th segment notably
striate (according to figure) with cremaster not distinguished;
body smooth with a few punctures.

12. Cistidia couaggaria (N.M.). Cremaster very short, mas-
sive and rugose, with one pair of longer hooks. Dorsal and
lateral grooves strong. Body practically smooth, punctate only
in front of the keel through front of spiracle, as in Sicya; but
callosity large and rough. Femur not noted. A genus generally
considered related to Abraxas ; the pupa shows a marked likeness
to the Cingilia group, but the characters are not compelling.
Imago with simple antenna.

13. Abraxas grossulariata (C.U.). Cremaster short, flat, semi-
circular, with four short and very heavy hooks radiating from it
at equal distances ; 11th segment and cremaster-base rugose, and
bearing the four small hooks; dorsal groove shallow, with only

Sept., 1945]

Forbes: Geometries

197

three vague notches on its posterior side, cut off below by the
lateral groove, which is oblique, with sharply keeled upper boun-
dary and rounded lower, the upper ending behind in a spur and
lower continued as the suture ventrally; sculpture coarsely and
sparsely punctate on a smooth glossy base, the posterior portions
of segments glossy and impunctate, followed by a shagreened
posterior declivity on segments 4 to 6; segment 5 laterally im-
punctate, without spiracular furrow, but with a strong flange-
plate, preceded by a deep sharply bounded groove, as, e.g., in
Erannis, and unlike most forms with 8-hooked cremaster. Cal-
losity obscure ; appendages normal, femur concealed. The eye is
as smooth and glossy as the rest, with the division into glazed and
sculptured eye barely visible.

Larva with uninterrupted hooks. Imago with simple antenna.
This genus does not fit into the American pattern, having the
cremaster of the second and the flange-plates of the Boarmine
series. But compare Sericosema.

“Abraxas sylvata” (Europe, Dampf, and C.U. from Staud-
inger & Bang-Haas). This pupa is so totally different from the
well known A. grossulariata that I feel sure there must be some
mistake. Perhaps it is a Boarmia in the broad sense.

Cremaster proper glossy, broad at base, and narrowing pretty
abruptly to a short slender tip ; 11th segment rugose. Dorsal
groove rather weak, only the middorsal notch strong; lateral
groove degenerated. Sculpture heavily punctate, with a row of
punctures even on 9th segment, 10th only smooth. No spiracular
furrow, the 5th segment smooth for a limited area in front of
spiracle ; flange-plate very short and sharp, extending up from
level of spiracle ; with a deep, short, smooth groove in front of it
(suggestive of Paleacrita) ; appendages normal, with femur con-
cealed. Callosity small but normal.

14. Nematocampa filamentaria (C.U.). Cremaster short, the
penultimate pair of hooks as stout and almost as long as the
terminal pair, but extending much more laterad; dorsal groove
shallow", with about three posterior notches, only the middle one
well defined ; lateral groove more nearly longitudinal than usual.
Sculpture of sparse puncturing on a dull surface, the 9th and 10th
segment unpunctured, but also dull, posterior part (true cre-
master) only of 11th roughened. Callosity obscure, slightly

198

Journal New York Entomological Society

[ Vol. LIII

raised but not discolored. 8 'piracular furrows of 5th and 6th seg-
ments similar, a series of sharp vertical somewhat anastomosing
ridges, with punctures between them; 7th with a rather sharp
ridge through spiracle, but the rest of the system degenerated.
No flange-plate seen.

We must- assume this genus is degenerate from the Sabulodes
group, but it is quite abnormal ; the well known larva (see Pack-
ard and Mosher) is unique. (Mosher, ’17, p. 42.)

15. Metanema inatomaria (Franglemont). Pupa very strongly
flattened.. Cremaster a shapeless nodule, with very short hooks
on its dorsal surface only; dorsal groove shallow, the posterior
edge hardly dentate, cut off laterally by the unique lateral groove,
which is a shallow area, absolutely continuous with the side of the
9th segment, but extending sharply dorsally and separated only
by a high chitinized ridge from the lower end of the dorsal groove.
Sculpture densely and rather coarsely punctate, the 8th as well
as 9th and 10th segments impunctate, but dull ; no trace of spirac-
ular furrow, the front of 5th segment absolutely continuous with
the rest. Callosity large and high. Femur covered.

Antenna of imago with cones at apices of one series of the
naked pectinations. The pupa and moth remind one strongly of
Selenia, but the antennae are of the Therina type.

15. Selenia alciphearia (N.M.). Cremaster very short, with a
mass of short hooks, wider than long and very rough; dorsal
groove normal, lateral slight and flat, open in front. Sculpture
of sparse irregular puncturing, segments 6 to 10 practically alike
and smooth above ; built very stout. Callosity well developed and
setulose, like the preceding. Face projecting ventrally; femur
concealed; fore tibia projecting and only twice as long as wide.

15. S. tetralunaria (Europe, C.U.). Similar; puncturing very
even all over, but absent from segments 8 to 10; the dorsal and
lateral grooves are shallow, the male antenna keeled, and with a
suggestion of the tubercle-row of Ennomos ; fore tibia not promi-
nent.

Antenna of imago naked, with cones on shaft, unlike Metanema.
Larva with sucker interrupting hooks.

(15) Eumera regina (Japan, C.U.). Entire pupa smooth and
rather glossy, even the eyes and front being smooth and continu-
ous, and only faint rugosities dorsally on thorax, etc. Cremaster

Sept., 1945]

Forbes: Geometric

199

vestigial, flanked by two minute hooks. Dorsal groove strong,
but dorsal only, with about 15 teeth, rather shorter middorsally ;
no lateral groove. No spiracular grooves or flanges. Appendages
smooth, femur covered.

This pupa is far too degenerate to place in a group ; while
smooth it has no resemblance to such genera as Xanthotype, being
fully chitinized. It may possibly be related to Selenia.

16. Nacophora quernaria (Mosher, ’16, 129). Antenna with
5 or 6 rows of tubercles; flange-plates present.

Larva with hooks uninterrupted. Antenna of imago essentially
as in Cingilia group ; of which it is probably a derivative. There
is no connection to the Biston group, where it is commonly placed.
Phceoura mexicanaria is merely a Nacophora, “Biston” arnobia
is a little more distinct, but doubtfully worth a genus.

17. Euchleena tigrinaria (N.M.). Dorsal' groove marked, with
a deep middle notch and a group of two or three subdorsals.
Lateral groove strong, standing at 45°. Sculpture of sparse pit-
ting dorsally, 9th and 10th segments and most of 8th smooth and
shining. Segment 5 with a flange tangent to spiracle (replacing
spiracular furrow). Callosity large but smooth and concolorous.
Antenna moderate, with two rows of pits. Mid tibia extending
forward to front of eye.

17. E. obtusaria decisaria (N.M.). Cremaster with two long-
hooks. Dorsal groove with 7 strong notches ; lateral running into
suture of segments 9-10 below ; sculpture coarser. Antenna with
a strong median keel ; spiracular furrow clearly absent. Callosity
formed of two surfaces at an angle, but both belong to mesothorax
on dehiscense.

The relation of sucker and hooks in the larva varies from spe-
cies to species, but any interruption tends to be weak or absent.
Imago with naked pectinations and cones on shaft. By the an-
tenna this belongs to a well limited holarctic group, including
Lytrosis and Stenotrachelys, which are very close, Angerona and
Xanthotype, which represent each other and are perhaps not dis-
tinct, Metarrhanthis and Cepphis (Priocycla). Many other
genera are similar, but have only two strong setae on each pecti-
nation instead of three.

17. Hemerophila atrilineata (N.M.). Near Euchlaena in both
imago and pupa. Sculpture heavily punctured, with part of seg-

200

Journal New York Entomological Society

[Vol. LIII

ment 8, and 9 and 10 contrasting, smooth ; callosity not very
large; first two spiracles half covered. Antenna keeled, tongue
only f way to apex of appendages, unlike Euchkena.

18. Priocycla decoloraria (C.U., reared by Benjamin). Essen-
tially like Euchkena. Dorsal groove rather short, the lateral
being high on sides, 7-notched; lateral continuous with surface of
9th segment as in Metanema, but normal in position. Sculpture
as in Euchkena, 8 smooth, no traces of spiracular structures on
segment 5. Callosity absent, appendages normal, antenna faintly
pitted. Tongue full length.

The cocoon is of open meshes, but much finer-meshed than
Cingilia.

18. P. ar mat aria (C.U.). Practically identical. In the single
specimen the lateral groove is smaller and branches off from the
dorsal groove, approaching the condition of Cabera.

Larva with series of hooks uninterrupted ; imago with antenna
as in Euchkena.

17. Metarrhanthis hypochraria (Rupert). Closely similar to
Euchlsena. Dorsal groove with several notches, decreasing in size
toward sides; lateral very deep, but merely an extension of the
dorsal. Cremaster dorsally heavily reticulate, the reticulations
drawn out lengthwise ; sculpture coarsely and densely punctate,
including most of the 8th segment. Callosity very strong, in the
form of a half-funnel opening forward. Front sharply raised
above the level of the eyes, which are very coarsely sculptured.
Antenna rather coarsely sculptured, with a double row of pits
which are strongest at middle of its length, and flanked by vague
tubercles. M. warneri and angularia differ in very minor points
of sculpture that may be individual. M. obfirmaria is smoother
and more glossy, with less puncturation on 8th segment, cremas-
ter perhaps a little longer in proportion and shallow very widely
open lateral groove.

17. Gonodontis Mdentata (Dampf). Cremaster short, more
neatly triangular than in Metarrhanthis, the two terminal setee
tremendous and specially socketed, the rest very small and equal.
Dorsal groove with 7 or 8 deep and equal notches, lateral shallow
and very broadly open. Sculpture of rather sparse fine punc-
turing on a smooth and glossy base ; totally absent on 8-10, seg-
ment 9 being only half as wide as 8 and 10; callosity small, but

Sept., 1945]

Forbes: Geometrid^e

201

well set off and of the clearly open half -funnel type of Metar-
rhanthis. No trace of spiracular furrow or flanges. Appendages
normal. So far as the pupa goes this could be a minor differenti-
ation of Metarrhanthis, as they were formerly placed, but the
larvas are strikingly different, Gonodontis having rudiments of
two extra pairs of prolegs. In both the hooks are uninterrupted,
but in Metarrhanthis the middle hooks are shortened to mere
rudiments. The imaginal antenna has lost one of the three termi-
nal setae on the pectinations, a character I believe of some signifi-
cance.

18. Himera pennaria (Dampf). Cremaster rather smooth,
shining, somewhat longitudinally ribbed, the two terminal setae
massive as in Gonodontis, but curved sharply ventrad; dorsal
groove deep with about 8 deep notches like Gonodontis; lateral
vestigial. Sculpture of fine sparse puncturation on a glossy base,
the 8th segment with sparser puncturing behind the middle only ;
callosity vestigial. Appendages normal; no trace of spiracular
furrow or flanges.

The cremaster and dorsal groove suggest Gonodontis, which
the antenna also resembles; the caterpillar has a minute vestige
of the proleg on the 5th segment in the form of a circle of rudi-
mentary hooks, so the connection can be considered solid.

18. Crocallis elinguaria (Dampf). Cremaster with the two
much-enlarged hooked terminal setas; not very rough at base.
Dorsal groove with five deep dentations and a lateral pair of shal-
low ones; lateral shallow and widely open. Sculpture heavily
punctate on a shagreened base ; 8th segment without punctures.
No callosity or spiracular structures. Appendages normal, the
basal portions transversely striate with fine grooves.

Larva with hooks in a complete series, no traces of an extra
proleg. Imago with antenna with cones on apices of both sets of
pectinations, like the following and the Cingilia group, but
unlike Gonodontis and Himera.

19. Abbotana clemataria (N.M. and Mosher, ’17, p. 49). Cre-
master rugose, with setas 2, 6, the terminal extremely large as
given by Mosher. Dorsal groove with a deep central notch,
flanked by a broad riser, and then narrower side notches and
risers; lateral groove very deep, standing at 60° to the longitudi-
nal ; sculpture coarse, of tubercles, continued on 8th segment, 9th

202

Journal New York Entomological Society

[Vol. LIII

smooth, 10th longitudinally ribbed in front ; incisures granulose.
A tendency for the sculpture to form three or four spiracular
ridges. Vertex and thorax with a strong median crest; callosity
obscure. Femur concealed, antenna narrow, with five tubercles
on each segment, arranged 2 and 3.

In the imago in spite of the subpectinate antenna one would
place this next to Sabulodes, but the pupa is wholly different.
Larva with sucker interrupting hooks.

Therina endropiaria (Rupert). Cremaster longer than in the
preceding, four-ridged so as to form an oblong pyramid ; terminal
setae not massive. Dorsal groove with about five strong notches,
lateral deep. Sculpture with punctures running together into
vermiculate grooves, very rough, extending onto anterior part of
8th segment; 9th and 10th smooth, finely shagreened. Sculpture
rather transverse, and forming a hint of spiracular furrows but
not definitely. Callosity minute, but perfectly formed and
faintly setulose.

T. fiscellaria (Rupert) is much smoother, without the rough
vermiculation, but with the punctures tending to lengthen into
shallow .vertical grooves, the 8th segment almost free of sculpture,
and the 5th segment in front of the spiracle with rather more
distinct vertical grooving, but still not really with spiracular
grooves. Dorsal groove with smaller and more numerous notches,
about 9 rather than five to seven. Destutia excelsa shows no sub-
stantial difference, and is also sparsely punctured ; I believe it is
merely a variant Therina.

I consider Therina should be made a conservandum, with type
fervidaria, the one of the two original species which was figured.
While the distinctness of the European “Ellopias” and the
American “Therinas” was never recognized in print- it was well
known to all workers on Geometridse for many years, and usage
was absolutely consistent from Packard’s revision till McDun-
nough’s check-list came out, and practically so for another 20
years after that, except by those who automatically followed the
check-list. I cannot see that the “genera” of Capps are more at
best than very weak subgenera. They are mainly based on small
differences of genitalia, in a group where the genitalic type is
perfectly homogeneous.

Sept., 1945]

Forbes: Geometric

203

18. Caripeta divisata (N.M.). Cremaster of typical 2-6 type,
short and stout. Dorsal groove deep and coarse-toothed, lateral
well-marked. Sculpture of fine punctures, connected by faint
grooves, 8 and 9 completely fused in middle line, dull but not
punctured; 10 smooth. No spiracular furrow , callosity obscure.
Appendages normal, antenna with punctures.

A normal variant of Therina, with which the imaginal venation
agrees. Curiously for this series, the antennal pectination is
scaled ; the cones are on the shaft. Larva with sucker interrupt-
ing hooks (Rupert).

19. Hygrochroa syringaria (Europe). Cremaster with hooks
apparently all very short; no dorsal groove, traces of a lateral
one. Sculpture deeply pitted, and whole pupa densely hairy,
including segment 9 ; segment 5 with a keel through spiracle only.
A cone on face, a ridge on eyes; callosity very large and velvety.

A very curious type, looking like a caricature of Selenia. The
larva also has the dense secondary hair, which is unique in the
Geometridse, and the hooks are interrupted. Imago with pecti-
nations naked, but no cones found.

20. Ennomos magnarius (C.U.). Cremaster a high pyramid,
twice as wide as thick, with four small hooks f way out and the
third pair subterminal and about as heavy as the terminal ; longi-
tudinally ribbed ; dorsal groove obsolete, the suture open ; lateral
represented by a broad triangular slightly depressed smooth area ;
body very thin, white, the sculpture of transverse rugosities ;
segment 9 smooth, 10 longitudinally strigose ; thorax and base of
abdomen dorsally nubbly. Most tubercles well set off, but de-
pressed, the tubercles of segments 8 and 9 conical. No callosity
or spiracular furrows. Antennce with four rows of conspicuous
tubercles, and traces of a fifth; tongue less than half as long as
other appendages, I believe the shortest in the entire Geometridse.

Larva with sucker interrupting the hooks. Antenna of imago
unique in details, with cones on the tips of the scaled pectinations.
An isolated genus, obviously degenerated from a type with dorsal
and lateral grooves^ but far from any other genus studied. While
there are differences of detail, I see no justification for a genus
Deuteronomus. The differences in venation are of a type that
occurs elsewhere, the larva and pupa are almost exactly as in

204

Journal New York Entomological Society

[Vol. LIII

typical Ennomos, and the antenna is substantially the same ; only
the genitalia differ in a group where the structures are highly
plastic anyway, as shown by the striking differences between
species in both the Ennomos and Deuteronomus groups. Even
the coloring is the same except for our single species subsignarius.
The members of both groups are known to hybridize.

20. E. subsignarius (C.U.). Similar to magnarius; cremaster
with hooks more nearly terminal; skin less completely mem-
branous, with rugosities stronger, but almost absent on segment
10 ; quite heavy before spiracle of 5, but not really forming spi-
racular grooves. Appendages as in magnarius except for the
stronger sculpturing. Tubercles ii weaker.

20. E. autumnaria (Europe). In appearance like magnarius,
the skin white but not quite as thin; sculpture longitudinal on
10th segment and subdorsally on 9th ; spiracular furrows rather
distinctly recognizable, multiple, divided by sharp ridges. Cre-
master hooks like subsignarius.

20. E. erosaria (Europe). With the darker color and heavy
skin of subsignarius, heavily rugose, the spiracular furrow near-
est the spiracle rather well marked but extremely harrow. 9th
segment smooth, 10 striate; tubercles i and ii on 9th somewhat
enlarged, black and conspicuous.

21. Patalene puber (N.M.). Cremaster with all hooks very
short, and the longer two pairs subequal; dorsal groove repre-
sented only by series of punctures before and behind the suture;
lateral groove a mere right triangular depression, with a fine
longitudinal ridge above and a rounded oblique one below ; with
separate grooves on the sides of the very large cremaster. Sculp -
ture punctate ; 9 and posterior part of 8 smooth ; mid tibia wholly
and fore tibia mostly covered; femur in the specimen examined
with a linear exposure. Callosity obsolete.

The moth would invariably be placed close to Apicia and the
pupa is quite a surprise, but confirmed by the associated moth.
Larva with hooks uninterrupted. Antenna of imago with pecti-
nations scaled and cones on shaft, and with some scattered extra
type 2 setae on the sides of the pectinations, like Apicia.

22. Opisthograptis luteolata (Dampf). Cremaster short, with
rounded end bearing a row of four practically equal hooks like
Abraxas, the other four a little further basad, as usual ; no traces

Sept., 1945]

Forbes: Geometric

205

of dorsal or lateral grooves. Sculpture punctate on a silky
slightly dull ground, limited to anterior portions of segments,
and absent from 8 to 10. Spiracular ridge well marked, through
the spiracle on segments 6 to 7, slightly in front on 5 ; no flanges.
Spiracles of segments 2 and 3 raised, conical, unlike those of 5
to 7. Appendages normal, only with segmentation marked on the
narrow antenna ; femur exposed.

Larva with prolegs on segments 4 and 5 even better developed
than in Gonodontis; sucker interrupting hooks. Imago with
simple antenna. A peculiar, perhaps a relict genus, apparently
without close relatives, ranging over the Palsearctic. On genitalia
Pierce puts it next to Gonodontis, though the latter has a pair of
structures which may represent the clavus. Both have the type
of gnathos with small dentate tip.

23. Plagodis altruaria (N.M.). Cremaster with all setas minute
and subequal. No dorsal or lateral groove, the suture being an
ordinary ankylosed one. Sculpture fine and punctate, including
anterior portions of segment 5 ; 8 to 10 smooth ; prothorax longi-
tudinally ribbed, but no other special sculpture. Callosity very
large, black and chitinized ; legs and tongue normal, antenna with
one puncture to a segment; fore femur damaged in specimen
examined, apparently with a linear exposure.

Larva with sucker interrupting series of hooks. Imago with
characteristic antenna and venation ; the antenna with scaled
pectinations and cones on shaft. Hyperitis and Anagoga are
closely related by imaginal structures, but the more distant rela-
tionships of the three are obscure.

23. Hyperitis alienaria (Rupert). Similar; cremaster with
terminal pair of setaa much stronger than the rest, though small ;
ridging of prothorax obscure and irregular ; antenna with a vague
ventral terminal depression on each segment rather than a punc-
ture ; femur concealed. The anal notch of fore wing is strongly
shown on the pupa, unlike Gonodontis, where the scalloping is a
secondary development, due to the degeneration of part of the
wing-pad.

SUMMARY

Considering primarily the pupal characters, the Ennomime
divide into two groups :

206

Journal New York Entomological Society

[Yol. LIII

A, a Boarmine series, with cremaster bifid, with hooked setae
absent or vestigial; flange-plate present at least on 5th segment
of abdomen; antenna of imago with cones on shaft, and larva
without the sucker interrupting the series of hooks on the prolegs.

B, an Ennomine series, with cremaster simple, but bearing 4
pairs of hooked setse, the terminal pair or two longer and
stronger; flange-plate absent; femur concealed; the imago nor-
mally with naked pectinations, frequently bearing the cones at
their tips ; the larva most often with the sucker interrupting the
series of hooks.

But there are numerous exceptions and recombinations among
these characters, the principal absent? combinations being a,
flange-plate present, dorsal groove absent; cremaster 8-hooked;
and 1), cremaster bifid and flange-plate absent, except for the re-
duced Macaria group which have spiracular callosity present and
femur exposed.

Aside from this Macaria group, the Boarmine series divide into
Bistonine type with femur concealed but dorsal groove preserved,
and a Paraphia type with dorsal groove lost and femur exposed ;
but there is every possible intergrade between these types.

Symbols on Table

First row: N: pectinations of imaginal antenna naked; S: scaled; A:
absent.

Second row : S : cones on shaft of antenna ; P : cones on pectinations ; O :
no cones on pectinate segments.

Third row : s : larva with sucker on proleg interrupting series of hooks ; o :
series of hooks continuous.

Notes

1 Flange plate not noted (mostly concealed in specimens examined) pre-
sumed as listed.

2 Spiracular callosity of mesothorax not noted.

3 Femur not noted.

4 Femur vestigial, and perhaps sometimes absent.

5 Callosity very weak, should perhaps be counted as absent.

e Lateral groove not set off from dorsal.

7 A faint vestige of dorsal and lateral grooves.

s Dorsal groove vestigial but lateral distinct.

9 A distinct vestige of dorsal and lateral grooves in E. crepuscular ia ;
femur almost covered in E. bistortata.

Sept., 1945]

Forbes: Geometrid^e

207

a

<
& m
g PS
c*3 o

o ft

. ft ft

<D

ft | ;s

Q ft Q, OS 0-(
r— ^r1 GO H

|J^g:

H >“ Er pj ft

3 S|

O O -M

pq ft ft !><! ft ^

O O 02

g ggo2 02 . O
ft ft ft 02 02 < co

® ft c3
^ .— I -rH ‘ft
O o rft £

•3 5'£<ft

V r-H ft, -ft

>y® ft ft

:

QJ

X

: : ^

£ ^ ft pq

; ift

ft

| IS

_ o
O 'CO 02

i i xn.

02

j 1 02

<*H

\ lift

02

; j co

§.«

pq pq

02

oq£

ft OD ft
Prt ® © X

.r- ft

rpH 3 .2 ,S

as

<jHO“

O o

02 co CO o

S M H
C3

n & w

M

o +
W w
' t>

si®
co e

oj o

ft s.

<s

ft :

CD

'ft

<D

•IS 00

CO ,0

© ^

© M
° ft

r~^ S3
e cd

ft

w o

g 5

02 J

S <

3 02
W os

P3 O
d ft

§ be

ft. |s
<d ft
ftCC

si

e

§fV£k<

o © ^ ^ g

o

ft o
*A<

•?=> CO

^ o

•w o ftw

si ^ s
§ § ® .2
rft V. +2

3 13 I

Sg § n3
2

<3 *** ft

<3D

'ft.g

CD fti
ftCQ

i

<v

-l-o

'ft

&
ft O

(D ?H
-ftS <D

CO ft

CO 02

ift

2 2

'3 2

"ft©
<1 EH

DO .
02 <1

208

Journal New York Entomological Society

[Vol. LIII

c$ „

2 .H*

•ssa

2 S ©

&J0 A £*>

WWft

002
xji m
'Amm

o

£

o

5> £
^ ©
§ £

O

s

o

rt i
d

02

to ©

&H-

Ǥ

® 2
fcJOJS

5C' O 02 02

aimmaimm rr,Q-\
'&'A'&'A'A'Am'A

— _ s

■£ -d =0 « H

3 2 ,C) , <M

| e

& 1 3 § 1 1 i I

® -e dH£ =0 ,0 CD

? oj o

c$

H •pH ® . TO

be

© ® ® o

«3g «-g c8

fnH 3j®4J

® cj o ao
g « o -F\o
S P

^ ft ft

Ph ?h ^

cs ^ .a ® ^3 £ 'G ft -2

oq WOPLhO<1

03 PhPhPhPhPhP umww

<5 fr^A^'A'A'A'A'Amm

SSs Cabera6
As Nematocampa
Eumeria

Sept., 1945]

Forbes : Geometrid^e

209

SUPPLEMENT

The following pupas have come in since this paper was in press.
Those of Leucobrephos and. Bapta are of particular interest.

(3) Leucobrephos brephoides (U.S.N.M.). Cremaster proper
short, bearing two massive spines pointing straight back. Dorsal
groove deep, widening backward at middle into a deep semi-
circular or more pointed pit, lateral groove merely a shallow
notch in its side. Sculpture sparsely punctate, including segment
8, but 9 and 10 smooth. No spiracular furrow or flange-plate, the
former represented by a slight smooth ridge,; spiracles, excepting
as usual the first, set well away from edge of wings. Fore femur
and callosity absent ; tongue f length of wings.

Larval and imaginal characters show this is closely related to
Brephos, but the pupa would be considered a normal Ennomine,
and totally unlike Brephos, wdiich is correctly described by Miss
Mosher. The pupa is normal in form, much less cylindrical than
Brephos.

3. Hesperumia sulphuraria (U.S.N.M.). Cremaster bifid, the
base large and highly rugose, ending in a very short smooth
cremaster proper, which bears the usual two spines. Dorsal
groove normal, moderate, lateral strong, a broad deep triangular
depression. Sculpture punctate, 9 more lightly, 10 impunctate;
a slight compound pit at front of segment 1 at middorsal line ;
flange-plate and groove double, the posterior groove smooth, the
anterior pitted, separated and followed by sharp ridges. No
spiracular groove or callosity (Face-plate lost).

By imaginal characters this should belong to the Boarmid
group {5), but the cremaster is on the whole more Amphidasis-
like. Ematurga is somewhat anomalous in the same way, and
may be the closest relative whose pupa is known.

6. Melanolopkia signataria (?) (Rupert). Cremaster regu-
larly tapering to a long bifid tip ; rugose on basal half, the rest
smooth and shining. No dorsal or lateral grooves. Sculpture
punctate, segments 9 and 10 smooth. Spiracular groove shallow,
smooth and shining, the two ends turning back to the level of the
spiracle as if pushed back by the flange-plate. Pit of flange-plate
deep, with sharp anterior as well as posterior edge, smooth at
bottom, tapering to point at ends, the upper end turned sharply

210

Journal New York Entomological Society

[Vol. LIII

back, and anterior edge with a single reentrant point toward
upper end. Fore femur covered ; spiraeular callosity obsolescent,
rugose.

M. imitata (U.S.N.M.) differs, so far as I can tell by brief
notes, mainly in the pubescent, though small, callosity.

(8) Bapta vestaliata (Rupert). Cremaster bearing a pair of
extremely heavy terminal hooks, with their fine apices sharply
bent to the sides, and small hooked setae (only two pair seen), the
base (i.e. segment 11) short, shining and rounded; dorsal groove
deep, with 7 or 8 deep posterior notches, fading out below, but
faintly connected with the shallow rounded pit which represents
the lateral groove. Sculpture punctate, 8 smooth as well as 9
and 10, but with a slight sinuous anterior dorsal ridge. No
spiraeular grooves or flange-plates. Callosity vestigial; mouth-
parts normal, with full length tongue and femur concealed.

Larva with sucker very large, dividing the series of hooks into
two widely separated patches.

This is an important and very distinct form, being the first
pupa seen of its tribe. The general characters suggest the Cabera
(Deilinea) group, the cremaster corresponding to Sericosema
though not to Cabera, but Sericosema has the flange-plates, which
are totally absent in Cabera and Bapta.

22. Plagodis keutzingaria (Rupert). Fore femur concealed.

Sept., 1945]

Rapp & Cooper: Psychodid^e

211

CHECK-LIST OF PSYCHODID^ OF ASIA AND
AUSTRALIA1

By William F. Rapp, Jr., and Janet L. Cooper

The area covered by this check-list includes India, Malay
Peninsula, China, Japan, Philippines, East Indies, Australia, and
New Zealand.

BRUNETTIA Tonnoir

biformis Edwards, Ins. Samoa, Vol. 6, fasc. 2 (1928), p. 68.
Samoa.

sapphiaina Edwards, Treubia, Yol. 6, p. 160.

Sumatra.

travancarica Annandale, Rec. Ind. Mus., Vol. 5, p. 144.

India.

DIPLOMIA Annandale

superstes Annandale, Calcutta Jour. As. Soc. Beng., Yol. 4
(1908), p. 353.

India.

FLEBOTOMUS Rondani

angustipennis Meyere, Tijdschr. Ent., Vol. 52, p. 202.

Java.

annandalei Sinton, Ind. Jour. Med. Res., Vol. 10, p. 744.

India.

arboris Sinton, Ind. Jour. Med. Res., p. 107.

India.

argentipes Annandale and Brunetti, Rec. Ind. Mus., Vol. 2, pp.
101-104.

Calcutta, India.

babu Annandale, Rec. Ind. Mus., Vol. 5, p. 49.

India.

babu var. insularis Theodor, Ind. Jour. Med. Res., Vol. 26, p. 264.
Ceylon.

bailyi Sinton, Ind. Jour. Med. Res., p. 821.

India.

i Fourth Contribution to a Check-List of Psychodidae of the World.

212

Journal New York Entomological Society

[Vol. lixi

bailyi var. campester Sinton, Ind. Jour. Med. Res., p. 821.

India.

barraudi Yao and Wu, Chin. Med. Jour., Vol. 59 (1940).

Kwanysi, China.

barraudi var. kwangsiensis Yao and Wu, Chin. Med. Jour., Vol.
59 (1940).

Kwangsi, China.

barraudi var. siamensis Causey, Amer. Jour. Hyg., Vol. 28, p. 488.
Siam.

ZupfoY Manalang, Philipp. Jour. Sci., Vol. 45, p. 356.

Philippines.

brevifilis Tonnoir, Bull. Ent. Res., Vol. 26,. p. 145.

Australia.

chalami McCombie, Yong and Chalam, Ind. Jour. Med. Res., Vol.
14, p. 849.

Bombay, India.

Christopher si Sinton, Ind. Jour. Med. Res., Vol. 15, p. 33.

Punjab.

colabcensis McCombie, Young and Chalam, Ind. Jour. Med. Res.,
Vol. 14, p. 859.

Bombay, India.

dayapensis Manalang, Philipp. Jour. Sci., Vol. 45, p. 358.
Philippines.

demeijerei Nitzulescu, Ann. Parasit., Vol. 8, p. 543.

Java.

dentatus Sinton, Ind. Jour. Med. Res., Vol. 20,. p. 869.

India.

eadithce Sinton, Ind. Jour. Med. Res., Vol. 20, p. 577.

India.

englishi Tonnoir, Bull. Ent. Res., Vol. 26, p. 144.

Australia.

heiseri Manalang, Philipp. Jour. Sci., Vol. 42, p. 299.

Philippines.

himalayensis Annanda'le,* Rec. Ind. Mus., Vol. 5, p. 50.

India.

hitchensi Manalang, Philipp. Jour. Sci., Vol. 42, p. 291.
Philippines.

hivernus Raynal and Gaschen, Bull. Soc. Path, exot., Vol. 28, p.
582.

French Indo China.

Sept., 1945]

Rapp & Cooper: Psychodid^e

213

hodgsoni Sinton, Ind. Jour. Med. Res., Vol. 20, p. 873.

India.

iyengari Sinton, Ind. Jour. Med. Res., Vol. 21, p. 221.

India, French Indo China.

iyengari var. hainanensis Yao and Wu, 10th Congr. Far East Ass.
Trop. Med., 1938.

Hainan, China.

iyengari var. malayensis Theodor, Ind. Jour. Med. Res., Vol. 26,

p. 266.

Federated Malay States.

kackekensis Yao and Wu, 10th Congr. Far East Ass. Trop. Med.,
1938.

Hainan, China.

keshishiani Shurenkova, Med. Parasit. parasit, Dis., Vol. 5, p. 892.
Pamirs, China, Tibet.

khawi Raynal, Ann. Parasit. hum. comp., Vol. 14, p. 530.

North China.

kiangsuensis Yao and Wu, Chinese Med. Jour. Suppl., Vol. 2.
China.

major Annandale, Rec. Ind. Mus., Vol. 5, p. 46.

India.

major var. chinensis Newstead, Bui. Ent. Res., Vol. 7, p. 191.
China.

malabaricus Annandale, Rec. Ind. Mus., Vol. 5, p. 48.

India.

manganus Manalang, Philipp. Jour. Sci., Vol. 42, p. 283.
Philippines.

marginatus Annandale, Spol. zeyl., Vol. 7, p. 62.

Ceylon.

maynei Sinton, Ind. Jour. Med. Res., Vol. 18, p. 195.

India.

morini Raynal and Gaschen, Bull. Soc. Path, exot., Vol. 28, p. 732.
French Indo China.

nicnic Manalang, Philipp. Jour. Sci., Vol. 41, p. 175.

Philippines.

perturbans Meyera, Tijdschr. Ent., Vol. 52, p. 201.

Java.

philippinensis Manalang, Philipp. Jour. Sci., Vol. 41, p. 175.
Philippines.

214

Journal New York Entomological Society

[Vol. LIU

purii Sinton, Ind. Jour. Med. Res., Vol. 18, p. 1203.

India.

pooi Yao and Wu, Chin. Med. Jour., Vol. 59 (1940).

Kwangsi, China.

punjabensis Sinton, Ind. Jour. Med. Res., Vol. 21, p. 421.

India.

queenslandi Hill, Bull. Ent. Res., Vol. 14, p. 83.

Queensland, Australia.

queenslandi subspecies meridionalis Tonnoir, Bull. Ent. Res., Vol.
26, p. 142.

Australia.

squamirostris Newstead, Ann. Trop. Med. and Parasit., Vol. 17,
p. 531.

Japan.

sylvaticus Raynal and Gaschen, Bull. Soc. Path, exot., Vol. 28,
p. 592.

French Indo China.

taianensis Patton and Hindle, Proc. Roy. Soc. Lond., Vol. 102,
Series B, p. 545.

China.

tonkinensis Raynal and Gaschen, Bull. Soc. Path, exot., Vol. 28,
p. 742.

French Indo China.

torrechantei Manalang, Philipp. Jour. Sci., Vol. 45, p. 361.
Philippines.

zeylanicus Annandale, Spol. zeyl., Vol. 7, p. 60.

Ceylon.

HORAIELLA Tonnoir

consimilis Tonnoir, Rec. Ind. Mus., Vol. 35, p. 60.

India.

prodigiosa Tonnoir, Rec. Ind. Mus., Vol. 35, p. 54.

India.

LEPIDOPSYCHODA Edwards

tie niformis Edwards, Ins. Samoa, Vol. 6, fasc. 2 (1928), p. 72.
Samoa.

NEMOPALPUS Alexander

australiensis Alexander, F. M. S. Mus. Jour., Vol. 14, p. 65.
Federated Malay States.

Sept., 1945]

Rapp & Cooper: Psychtodid^e

215

unicolor Edwards, F. M. S. Mus. Jour., Vol. 17, p. 257.

Borneo.

zelandicus Alexander, Insec. Inscit. Mens., Vol. 9 (1921), p. 158.
New Zealand.

PARABRUNETTIA Brunetti

albohumeralis Brunetti, Rec. Ind. Mus., Vol. 4, p. 311.

India.

flavicollis Brunetti, Rec. Ind. Mus., Vol. 4, p. 314.

India.

longichceta Brunetti, Rec. Ind. Mus., Vol. 4, p. 314.

India.

9-notata Brunetti, Rec. Ind. Mus., Vol. 4, p. 313.

India.

PERICOMA Walker

annandalei Brunetti, Rec. Ind. Mus., Vol. 2, p. 380.

India.

appendiculata Brunetti, Rec. Ind. Mus., Vol. 2, p. 379.

India.

loaner of ti Tonnoir, Ann. Soc. ent. Belgique, Vol. 60, p. 154.

Queensland, Australia.
bella Brunetti, Rec. Ind. Mus., Vol. 2, p. 383.

India.

bengalensis Brunetti, Rec. Ind. Mus., Vol. 2, p. 371.

India.

funebris Hutton, Trans. N. Z. Inst., Vol. 34 (1902), p. 180.

New Zealand.

impunctata Brunetti, Rec. Ind. Mus., Vol. 4, p. 309.

India.

margininotata Brunetti, Rec. Ind. Mus., Vol. 2, p. 381.

India.

margininotata var. gilvipes Brunetti, Rec. Ind. Mus., Vol. 2, p.
382.

India.

margininotata var. lacteitarsis Brunetti, Rec. Ind. Mus., Vol. 2,
p. 382.

India.

metatarsalis Brunetti, Rec. Ind. Mus., Vol. 4, p. 305.

India.

216

Journal New York Entomological Society

[Vol. LIII

metatarsalis var. khasiensis Senior- White, Mem. Dept. Agric.
India, Vol. 7, p. 125.

India.

mixta Brunetti, Rec. Ind. Mus., Vol. 4, p. 306.

India.

multacolorata Hntton, Trans. N. Z. Inst., Vol. 34 (1902), p. 180.
New Zealand.

proxima Brunetti, Rec. Ind. Mus., Vol. 4, p. 308.

India.

spinicornis Brunetti, Rec. Ind. Mus., Vol. 2, p. 378.

India.

squaminervis Brunetti, Rec. Ind. Mus., Vol. 4, p. 303.

India.

townsvillensis Taylor, Bull. Ent. Res., Vol. 6, p. 267.

Queensland, Australia.

unicolor Brunetti, Rec. Ind. Mus., Vol. 4, p. 309.

India.

PSYCHODA Latreille

acutipennis Tonnoir, Ann. Soc. ent. Belgique, Vol. 60, p. 149.
New Zealand.

albonigra Brunetti, Rec. Ind. Mus., Vol. 2, p. 374.

India.

albonotata Brunetti, Rec. Ind. Mus., Vol. 2, p. 373.

India.

albopicta Brunetti, Rec. Ind. Mus., Vol. 4, p. 296.

India.

apicalis Brunetti, Rec. Ind. Mus., Vol. 4, p. 301.

India.

argenteopunctata Brunetti, Rec. liid. Mus., Vol. 2, p. 375.

India.

atrisquamis Brunetti, Rec. Ind. Mus., Vol. 2, p. 376.

India.

acutipennis Tonnoir, Ann. Soc. ent. Belg., Vol. 40 (1920), p. 149.
New Zealand.

conspicillata Hutton, Catal. New Zealand Dipt., etc., Vol. 13
(1881).

New Zealand.

decora Brunetti, Rec. Ind. Mus., Vol. 4, p. 299.

India.

Sept., 1945]

Rapp & Cooper: Psychodid^e

217

distans Brunetti, Rec. Ind. Mus., Vol. 4, p. 296.

India.

distincta Brunetti, Rec. Ind. Mus., Yol. 2, p. 372.

India.

flava Edwards, Treubia, Yol. 9 (1927), p. 364.

Java.

fidvohirta Brunetti, Rec. Ind. Mus., Vol. 4, p. 297.

India.

genicidata Brunetti, Rec. Ind. Mus., Vol. 4, p. 294.

India.

hirtipennis Brunetti, Rec. Ind. Mus., Yol. 4, p. 300.

India.

maculipennis Brunetti, Rec. Ind. Mus., Yol. 4, p. 299.

India.

nigripennis Brunetti, Rec. Ind. Mus., Vol. 2, p. 376.

India.

notatipennis Brunetti, Rec. Ind. Mus., Vol. 8, p. 151.

Burma.

orlicidaris Brunetti, Rec. Ind. Mus., Yol. 4, p. 298.

India.

quadrifilis Edwards, Ins. Samoa, Yol. 6, fasc. 2 (1928), p. 72.
Samoa.

savaiiensis Edwards, Ins. Samoa, Yol. 6, fasc. 2 (1928), p. 74.
Samoa.

squamipennis Brunetti, Rec. Ind. Mus., Yol. 2, p. 375.

India.

transversa Brunetti, Rec. Ind. Mus., Yol. 4, p. 300.

India.

vittata Brunetti, Rec. Ind. Mus., Yol. 2, p. 377.

India.

TELMATOSCOPUS Eaton
horai Tonnoir, Rec. Ind. Mus., Yol. 35, p. 65.

India.

218

Journal New York Entomological Society

[Vol. LIII

BOOK NOTICE

Notas sobre Insectos Daninos Observados en Venezuela 1938-1943.
By Charles H. Ballou. 3a Confereneia Interamericana de
Agricultura. Caracas, 1945. 151 p. 6 figs.

This report is written by a fellow member of the New York
Entomological Society who has been serving since 1938 as Chief
of the Section of Entomology in the Instituto Experimental de
Agricultura in Venezuela. The survey on which the report is
based was begun on March 10, 1938, and terminated on December
31, 1943. The material is divided into two parts. In Part 1 are
listed the hosts — plant and animal — that give sustenance to the
injurious insects considered. Only hosts of economic importance
are included in this compilation, and the arrangement is alpha-
betic. In Part 2 the injurious insects and arachnids are given
the main emphasis and under each insect or arachnid are grouped
the organisms adversely affected by it. Here the arrangement
is by orders and families. Thanks to this division of the subject
matter the reader can readily trace first the different insects or
arachnids that attack a given host and then the different hosts
that furnish sustenance to a given insect or arachnid. In both
Parts the locality where the observation was made, the number
of the week, and the year, as well as the observer’s name, are
noted. The complete destruction of the host is indicated by
placing the number of the week in Italics.

In all 298 kinds of insects are listed but of these only 156 have
been determined as to species and a dozen of these 156 are listed
as doubtful; of the remaining 142, some 75 have been identified
as to genus but the residual 67 still lack generic determination.
In spite of these gaps in the knowledge, the publication will serve
as a very helpful guide to those in Venezuela who want to
know their insect and arachnid foes and to be on guard against
them. — Herbert F. Schwarz.

Sept., 1945]

Frost: Indigo Insects

219

INSECTS FEEDING OR BREEDING ON INDIGO,
BAPTISIA

By S. W. Frost
The Pennsylvania State College

Wild indigo is an interesting group of plants especially from
the standpoint of the insects that feed upon its flowers, leaves,
stems or seeds. The genus belongs to the Leguminosse (the
Fabaceae, according to some authorities) and comprises thirty-
one species that are confined to the eastern portion of North
America and range from Maine to Florida, west to Minnesota
and south to Texas. Some of the species occur in the immediate
portions of Mexico and Canada. Only two species occur in
Pennsylvania namely, Baptism tinctoria (L.), commonly known
as wild indigo, yellow broom, indigo broom or horsefly weed,
and Baptisia australis (L.), known as blue false indigo.1 Bap-
tism tinctoria is the more common of the two species in Pennsyl-
vania and it is from this species that the writer collected many
insects. Baptisia tinctoria ranges from southern New Hamp-
shire to Minnesota and southward to Florida and Kentucky. It
grows rather freely on sandy or gravelly soil and is common in
these localities.

The species of Baptisia are structurally and biologically dif-
ferent from most of the Leguminosae. They possess properties
which apparently are undesirable for many insects and may ac-
count for their relative freedom from insect attack. Three gly-
cosides; baptin, baptism and pseudobaptisin and one alkaloid
baptitoxin have been derived from Baptisia. A phenol, baptinol,
has been extracted from the dried leaves of Baptisia tinctoria by
means of vapors of chloroform and similar substances. The
latter explains the characteristic blackening of injured Baptisia
leaves which Clark (1915) remarks may be produced by any
agent, physical, chemical or mechanical, that disturbs the nor-
mal relationship of the cells containing certain enzymes. First

1 There has been some confusion in entomological literature recording
insects from Baptisia. This is probably due to the fact that Amorpha
fructuosa L. also bears the common name, false indigo.

220

Journal New York Entomological Society

[Vol. LI 1 1

a hydrolytic enzyme decomposes a glucoside and then an oxidase
acts to produce the dark pigment. Wehmer (1911) states that
Baptism tinctoria probably contains a small amount of indican,
a glncoside of indoxyl, from which indigo of commerce may be
produced.

Early in summer, Baptism tinctoria is a graceful, bushy plant
with clean, deep green foliage. Its glabrous, trifoliate leaves
attract some insects which disfigure them in various ways. Sev-
eral insects feed upon the seeds or contents of the pods while a
few bore into the stems. The small, but distinct, yellow flowers
appear about the first of June and are visited by many insects.
During the course of the summer the foliage become riddled by
the attacks of various insects and fungus diseases and by August
the plants in most localities are almost completely defoliated.2

The injects feeding upon wild indigo naturally have a some-
what limited range coinciding with the distribution of the plants.
Many of these insects seem to be confined to species of the genus
Baptism. All but one 'of the insects are native to North Amer-
ica. The coffee bean weevil, Arceocerus fasciculatus (DeG.),
originally introduced from India, has adapted itself to Baptism,
other weeds and- cultivated plants. Authors have occasionally
recorded some of the Baptism insects as ranging to the Pacific ;
however, species of the genus Baptism only occur east of the
Rockies.

1 have had an opportunity to study several rather uncommon
insects that feed upon Baptism tinctoria. These observations
together with previously published records suggested a compila-
tion of the species known to attack Baptism.

A SUMMARY OF THE INSECTS ATTACKING BAPTISIA SPP.

Coleoptera

Curculionidce

Arceocerus fasciculatus (DeG.). On the seeds of wild indigo,
senna, cotton, cacao, coffee, poke berry and dried apple seeds,

2 Eleven parasitic fungi have been recorded from Baptisia. Most of them
are listed by Seymour (1929). They include two mildews, two rusts, three
leaf spots and other disorders. The mildew Brysiphe polygoni D.G., which
occurs on Baptisia tinctoria and Baptisia australis , appears to be the most
prevalent species.

Sept., 1945]

Frost: Indigo Insects

221

Mass., Ohio, D. C., N. J., Ala., Fla. (Blatchley & Leng: 42).
Breeding in St. Ignatius bean ( Strychninos ignatii) in the Philip-
pines (Brown: 116).

Apion rostrum Say. Taken from the pods of false indigo,
Baptisia leucantha in the seeds of which they live, New Hamp. to
Fla., and west to Wis. and Texas (Blatchley & Leng: 81-82).
Infests the seeds of Baptisia leucantha (Beutenmuller, 1890).
Infests the seeds of Baptisia leucantha (LeConte & Horn: 411).
The writer has found these beetles abundant on Baptisia tinctoria
at Arendtsville especially early in June. On June 9 they were
observed feeding on the leaves and flowers. On July 10 there
was evidence of abundant oviposition. The eggs are pushed
through small holes made at the base of the pods, usually one,
sometimes two eggs in a pod. The egg is yellow elongate and
almost as large as the seeds which are small at this time.

Conotrachelus erinaceus Lee. Taken on cotton and Baptisia
in Texas (Pierce: 275). Occurs mainly beneath rubbish along
the banks of streams; also on hickory. Ranges from Ohio and
Northern Indiana to the District of Columbia, Florida and Texas
(Blatchley & Leng: 482).

Tychius sordidus Lee. Breeds in the seed pods of Baptism
bracteata (Sanderson, 1904). Breeds in pods of Baptisia leu-
cantha. The larvae eat pods clean, sometimes two or three larvae
occur in a single pod. They then eat their way out and pupate
in the ground (Pierce: 273). Ranges from Illinois and Iowa to
Louisiana and Texas (Blatchley & Leng: 245).

Lissorhoptrus simplex Say. One specimen on Baptisia, Texas
(Pierce : 265) . Ranges from Canada and New England to Michi-
gan and Iowa, south to Florida and Texas (Blatchley & Leng:
229). Apparently this species dues not breed on Baptisia for
Blatchley states “ Water is the element in which this beetle de-
lights, and it breeds only in plants growing in it.”

Chrysomelidce

Pachybrachys luridus Fabr. Occurs on the foliage of false
indigo, Baptisia leucantha, also on the flowers of Jersey tea.
Scarce in Indiana (Blatchley: 1130).

Pachybrachys trinotatus Mels. New Jersey on Baptisia tine-

Journal New York Entomological Society

[Vol. LIII

222

toria and on Ceanothus americana (Smith: 302). This species
has apparently been confused with P. luridus Fabr.

Lepidoptera

(Ecophoridce

Agonopteryx lecontella (Clem.). Food plant Baptism tinc-
toria (Clarke: 95). Larva green, tubercles shiny black, a dark
spot on each side of the prothoracic shield, a leaf roller on Bap-
tism tinctoria (Frost, 1945).

Tortricidce

Grapholitha tristrigana (Clem.). Larva on “Tinctoria” per-
haps Baptism tinctoria , Massachusetts to Florida, Kansas and
Oregon (Forbes : 394). Food plant Baptism and Lupinus ; larva
lives in seed pods and stems. Illinois, Kansas, Oklahoma, Ala-
bama, Texas, Florida, North Carolina, Pennsylvania, New Jer-
sey, New York, Massachusetts and Ontario (Heinrich: 39).

Pyralidce

Tetralopha baptisiella Fern. Larva on Baptisia, N. J., W. Va.,
and west, “New York.” (Forbes, 1920.) Baptism tinctoria Pa.
(Frost).

Tholeria reversalis Guenee. Larva on Baptisia and Lonicerar
N. Y., 111., to Fla., and Colorado (Forbes, 1920). Baptisia austra-
lis Pa. (Frost).

Hesperidce

Thanaos juvenalis Fab. Food: bean (Apios), wild indigo
(Baptism), Lathyrus and Galactia (Beutenmuller, 1890b: 202).
Ranges from Quebec to Florida and westward as far as Arizona
where it appears to be common (Holland : 335). The writer has
frequently observed the larva of a hesperid feeding on Baptisia
tinctoria. The head and thoracic shield were dark brown or
black, the former roughened and not shiny. The body was pink-
ish or brownish green, often olive in color. At first they ate
small pieces of leaves, later they tied several leaves together to
form a retreat in which they fed.

Thanaos brizo Boisd. Caterpillar feeds on Galactia and pos-
sible Baptism , from the Atlantic to the Pacific, ranging from
New England to Arizona (Holland: 333).

Sept., 1945]

Frost: Indigo Insects

223

Geometridce

Cingilia catenaria Drury. A general feeder, blackberry, cran-
berry, hazel, oak, Myrica, “Wild indigo’ ’ Genista tinctoria, Car ex
Pennsylvania and Rhus toxicodendron (Beutenmuller, 1890b :
221).

Noctuidw

Papaipema baptisice Bird. The caterpillar lives in the stem of
Baptisia tinctoria Providence, R. I. (Beutenmuller, 1902: 434).

Diptera

Agromyza baptisice Frost. A linear-blotch leafminer on Bap-
tisia tinctoria. Arendtsville, Pa. (Frost: 273).

Agromyza species. A petiole miner on Baptisia tinctoria.
Makes a long narrow shallow mine sometimes 31 inches long,
starting at the tip of a branch and working downward. Adults
were not reared (Frost).

Heteroptera

Gelchossa heidmenni 0. & D. This tingid is one of the most
abundant of the insects that attack Baptisia tinctoria in Penn-
sylvania. They make their appearance during May and by the
end of the month their characteristic white stippling is very evi-
dent upon the leaves. During June the plants are still vigorous
although the leaves may become almost white from the punctures
of these insects. By the middle of August the leaves are so
severely affected by these and other insects that they begin to
fall and before the end of the month the plant may be completely
defoliated. On False indigo Baptisia tinctoria , Mass., Pa., N. J.,
Md., D. C., La., Ark. (Weiss & West: 56-60).

Hadronema militaris Uhler. Food plant Baptisia tinctoria,
Long Island and New York (Britton: 501). Colorado, Iowa,
Michigan, Kansas, California, New Mexico (Van Duzee: 385).
On Baptisia leucantha, New York west to the Pacific feeding on
several species of Baptisia, also from Mexico (Blatchley, 1926 :
884).

Homoptera

Macrosiphum solanifolii (Ashm.). Colonies abundant on Bap-
tisia tinctoria during May, June and July at Arendtsville. Ap-

224

Journal New York Entomological Society

[Vol. LIII

parently a summer host. Identified by T. L. Guyton and J. 0.
Pepper. A common aphid with a wide distribution and a wide
range of food plants.

Jassus olitorius Say. Nymphs and adults of this common
species were found on Baptisia (Z. P. Metcalf, in correspon-
dence). A shrub feeding species often taken on Sassafrass dur-
ing July, August and September (Britton: 142). Ont., Me.,
N. Y., N. J., Pa., N. C., Fla., Kans., Ariz., (Bermuda) (Van
Duzee: 60). The writer has found nymphs and adults of a
species of cicadelid on Baptisia tinctoria at Arendtsville, Pa.

Thysanoptera

Sericothrips baptisiae Hood. Apparently confined to Baptisia
tinctoria, Virginia and Maryland (Hood : 113).

LITERATURE CITED

Beutenmuller, Wm. 1890. Food habits of North American Rhynchophora.
Can. Ent., 22: 200-203, 258-261.

. 1890b. Catalogue of the Lepidoptera found within 50 miles of New

York City, with their food plants. Annals N. Y. Acad. Sci., V.

: — . 1902. Descriptive catalogue of the Noctuidae found within 50 miles

of New York City, part II. Bull. Amer. Mus. Nat. Hist., 16 art. 33.
Blatchley, W. S. 1910. Coleoptera or beetles known to occur in Indiana.
Nature Publishing Co.

, and C. W. Leng. 1916. Rhynchophora or weevils of North Eastern.

America. Nature Publishing Co.

. 1926. Heteroptera or true bugs of Eastern North America. Nature

Publishing Co.

Britton, W. E. 1923. Guide to the insects of Connecticut, part IY, He-
miptera or sucking insects of Connecticut. State Geol. and Nat.
Hist., Survey Bull. 34.

Brown, R. E. 1906. Notes on the breeding habits of Arceocerus fasicu-
latus. Jour. N. Y. Ent. Soc., 14: 116.

Clark, E. D. 1915. Notes on the blackening of the leaves of wild indigo
( Baptisia tinctoria ) and the isolation of a new Phenol, baptisol.
Jour. Biol. Chem., 21: 646-660.

Clarke, J. F. G. 1941. Revision of North American moths of the family
CEcophoridae, with descriptions of new species. Proc. U. S. Nat.

. Mus., 90 (3107).

Czapek, F. 1921. Biochemie der Pflanzen. Gustav V. Fischer, Jena.
Forbes, Wm. T. M. 1920. The Lepidoptera of New York and neighboring
states. Cornell University Memoir 68.

Frost, S. W. 1931. North American Agromyzidae (Dipt.). Can. Ent.,
63: 275-277.

Sept., 1945]

Frost : Indigo Insects

225

. 1945. The larva of Agonopteryx lecontella. Jour. Econ. Ent.,

38(1) : 126.

Gorter, K. 1897. Tiber die Bestandteile der Wurzel von Baptisia tinc-
toria. Arch. d. Pharm., 235: 301-302.

Heinrich, Carl. 1926. Revision of the North American moths of the sub-
families Laspeyresiinse, and Olethreutinse. U. S. Nat. Mus. Bull.
132.

Holland, W. J. 1907. The butterfly book. Doubleday, Page & Co.

Hood, J. D. 1916. Descriptions of new Thysanoptera. Proc. Biol. Soc.
Wash., 29: 29-113.

Larisey, M. M. 1940. A monograph of the genus Baptisia. Ann. Mo.
Bot. Gard., 27(2) : 119-224.

Le Conte, J. L., and Horn. 1876. The Rliynchophora of America north of
Mexico. Proc. Amer. Phil. Soc. 15(96).

Sanderson, E. D. 1904. Insects mistaken for the Mexican cotton boll
weevil. Texas Agric. Exp. Sta. Bull., 74: 3-13.

Seymour, A. B. 1929. Host index of fungi of North America. Harvard
Univ. Press.

Smith, J. B. 1900. Insects of New Jersey. Twenty-seventh Ann. Rept.
State Board Agric. for 1899.

Van Duzee, E. P. 1917. Catalogue of the Hemiptera of America north of
Mexico. Univ. Calif. Press.

Wade, J. S. 1935. A contribution to a bibliography of the described imma-
ture stages of North American Coleoptera. Bur. Ent. and Plant
Quarantine, Wash., D. C., E-358.

Wehmer, C. 1911. Die Pflanzenstoffe. Gustav v. Fischer, Jena.

Weiss, H. B., and E. West. 1924. Notes on the false indigo lace bug,
Gelchossa heidemanni O & D. in New Jersey (Hem. Tingidae). Ent.
News 35(2) : 56-60.

226

Journal New York Entomological Society

[Vol. LIII

C. V. RILEY’S PIRATED WORK

In this Journal, vol. 46, p. 194, June, 1938, I published a note
referring to an overlooked title by C. V. Riley on the Colorado
potato beetle, which failed of inclusion in Samuel D. Henshaw’s
“ Bibliography of the More Important Contributions to Amer-
ican Economic Entomology, Part III.” The title in question is
“The Colorado potato beetle with suggestions for its repression
and methods of destruction,” by C. V. Riley. This was published
in London in 1877 by George Rutledge & Sons.

Had I read, at the time, the introduction to Parts I, II, III of
Henshaw’s “Bibliography,” I would have known that the omis-
sion was deliberate. According to the introduction, the book
published by George Rutledge & Sons was a partial reprint of
Riley’s “Potato Pests,” and it was unsanctioned and printed
without the knowledge of C. V. Riley. For this reason Henshaw
omitted this and other pirated works of Riley from his “Bibliog-
raphy.”— Harry B. Weiss.

Sept., 1945]

Abbott: Oviposition

227

THE MECHANISM OF OVIPOSITION IN PH^NICIA
(LUCILIA) SERICATA MEIG. (DIPTERA)

By Cyril E. Abbott

Oviposition is a complex process involving a variety of factors.
Some of the factors involve the physiology of the insect, some are
external to it. All form a complex so involved that the process
is difficult to control experimentally or even to evaluate {rom
observation. In these respects oviposition differs from, let us say,
the feeding response, which, although far from simple, may be
experimentally controlled with some degree of success, and its
characteristics predicted under known conditions with some de-
gree of assurance.

In a living fly, dissected in Ringer’s solution at room tempera-
ture, the ovaries may, and usually do, contract rhythmically.
Contraction is fairly rapid, relaxation slow, and the intervals
between beats are often decidedly unequal. Not only does the
ovary as a whole pulsate, the individual ovarioles exhibit a peri-
odic wave of contraction which passes from the free end of each
toward the body of the ovary. There is no synchrony in this con-
traction, for each ovariole behaves independently in this respect,
and indeed any synchrony between the contractions of the two
ovaries appears to be accidental. These movements are doubtless
responsible for the extrusion of ova which has been observed even
in cases of organs completely excised.

Studies extending over a period of three years had as their
object the determination of factors which influence ovarian pulsa-
tion. The evidence indicates that these are myogenic. Excised
ovaries may pulsate for hours, even in the presence of nicotine,
and there is no evidence that an eserine-acetylcholene solution
has more than a slight effect upon pulsation. Indeed, after
applying every possible type of stimulus, the conclusion was
reached that only three are effective in increasing the rate and
intensity of pulsation. These are : 1, mechanical, 2, osmotic, and
3, thermal.

Poking, pinching, or pushing an ovary which has ceased to
pulsate will often initiate pulsation which, however, is generally

228

Journal New York Entomological Society

[Vol. LIII

of short duration when the mechanical stimulus alone is applied.
The application of solid particles of NaCI, of sucrose, or of con-
centrated solutions of these, initiate pulsations which endure
much longer than those initiated by mechanical stimuli.

But the most effective stimulus is that of temperature. On
several occasions preparations, which had remained overnight,
had cooled to a temperature of 10° C., and ceased to pulsate,
resumed activity when the cold liquid was replaced by Ringer’s
warmed to a temperature of 25°-35° C. Moreover pulsation
then* continued for some time. Further observation demonstrated
that gradual cooling of the fluid was accompanied by a decrease
in the rate of ovarian pulsation. The table is. composed of aver-
ages from several observations.

TABLE I

Decrease in Ovarian Pulsation with Decrease in Temperature

Temperature, ° C.

Number of contractions
. per minute

35

8.1

34

6.0

33

5.4

32

4.5

30

4.3

28

4.2

27

4.0

Of and by themselves these observations indicate nothing con-
cerning oviposition. But consider the following data taken from
my notes.

Still not until I read in a paper by Shannon and Putnam
(1934), that the oviposition rate of Mdes cegypti rises 7.7 per cent
with each rise of 1° C., did the possibility occur to me that ovi-
position by P. sericata may depend more upon certain external
factors than upon nervous mechanisms in the insect.

It has generally been assumed that oviposition by calliphorid
flies is initiated by chemical and mechanical stimuli, the effects of
which are immediate and essential. Now although oviposition
generally occurs in the presence of “meat odors,” and usually,
too, when the insect is in direct mechanical contact with the meat,

Sept., 1945]

Abbott: Oviposition

229

gravid females, confined in small containers ( e.g a test tube),
will deposit eggs, and when confined in a cage will likewise deposit
eggs about their drinking fountain and dishes containing sugar.

Furthermore oviposition is closely associated with feeding.
Detinova (1936) has observed that Anopheles messece, before
ovipositing, probes the water with her proboscis. If the water
contains materials which inhibit feeding, oviposition is likewise
inhibited. Hecht (1930) found that optimum ovipositon in a
given species of mosquito depends in part upon the temperature
of the water where eggs will be laid.

Students are agreed that stimuli which initiate oviposition are
many and seldom operate singly. Kuzina (1940) found this to

TABLE II

Effect of Temperature upon Oviposition of P. sericata

Temperature, 0 C.

Average number of eggs
per female

20

1 ±.0

22

7 ± 1

24

19.5 ± 2

26

38.2 ± 2

28

45.5 ± 3

30

68.5 ± 2

32

75.0 ± 4

34

67.0 ± 5

36

42.5 ± 2

be true of Musca domestica. Mackerras (1933) states that ovi-
position by calliphorid flies depends upon copulation. (It is a
fact that every ovipositing fly which I have subsequently dissected
has contained sperm.) Hobson (1937) mentions as stimuli neces-
sary for oviposition the nutritive condition of the insect, chemical
stimuli, and contact stimuli. Starved, gravid females begin to
feed before they oviposit, and even replete flies, although they will
oviposit in response to distance chemical stimuli, generally require
direct contact with the substrate.

Obviously not all of the factors mentioned are stimuli of a
nervous type, and since even those that are, vary so in their effect;
one is justified in suggesting that the act of oviposition per se

230

Journal New York Entomological Society

[Vol. LIII

involves a release mechanism. This is the more probable because
observation demonstrates that the vagina is generally in a state
of contraction.

There is a definite limit, however, to inhibitory action, whether
at the site of the vagina or elsewhere. High temperatures, con-
finement, and repletion make retention of the ova practically
impossible. It is attraction of the fly to animal matter — not a
stimulus to oviposition — which accounts for the deposition of eggs
upon flesh. This also explains the observations of Salt (1930)
and Vladimirova and Smirnov (1938) that flies deposit more eggs
upon the nutritive substrate than it will support as larvse.

REFERENCES

Detinova, T. S. Med. Parasitol., 5(4) : 525-543. 1936.

Hecht, O. Riv. Malariol., 9 (6) : 706-724. 1930.

Hobson, R. P. Ann. Appl. Biol., 24(4) : 808-814. 1937.

Kuzina, O. S. Med. Parasitol., 9(4) : 340-349. 1940.

Mackerras, M. J. Bui. Entom. Res., 24: 353-362. 1933.

Salt, G. Nature, 125(3145) : 302. 1930.

Shannon, R. C., and Putnam, P. Entom. Soe. Wash., Proc., 06: 185-242.
1934.

Vladimirova, M. S., and Smirnov, E. S. Med. Parasitol., 7(5) : 755-777.
1938.

Sept., 1945]

Weiss: Longevity

231

THE LONGEVITY OF ENTOMOLOGISTS

By Harry B. Weiss

I had often wondered if entomologists as a group had a longer
average life than the average for the population as a whole but
the amount of work involved in assembling the data for a large
number of entomologists always deterred me from trying to find
out. However, upon the appearance of Mathilde M. Carpenter’s
excellent “Bibliography of Biographies of Entomologists” (The
American Midland Naturalist, vol. 32, no. 1, p. 1-116, 1945) I
found that the enormous job of assembly had been done and that
from this source I could draw the type of information that was
needed. Miss Carpenter’s bibliography, among other things,
contains the birth and death dates for 2187 entomologists born
between 372 b.c. and 1920. The term entomologist includes both
professional and amateur workers including some who, achieving
fame in other fields, made some contribution to entomological
science. It also includes a few women, but in numbers these are
so few that the list as it stands consists almost entirely of white
males from all parts of the world. It was impossible to separate
the amateurs from the professionals and this means that other
occupations are included. For the present purpose this is a
defect that is difficult to correct. On the whole, however, it can
be safely assumed that the list represents an occupational group.
The country of birth and cause of death are not known, as these
facts had no place in a bibliography of biographies. In order to
remedy this one would have to read all the individual biographies
and even then the information would not be complete because
many obituaries fail to mention the exact cause of death. In
view of this the mortality of the subjects considered herein must
be put down as due to all causes and the conclusions must be con-
sidered as applying to the entomologists of the world, rather than
of any specific country.

Table I shows the number of deaths and its percentage of the
total for each age from 19 to 98. The largest number of deaths
occurred in the age group 70-74, the highest number occurring

232 Journal New York Entomological Society [Vol. liii

at age 72. The number dying at each age is also shown by the
frequency curve (Figure 1), the peak of which is at 72. The
weighted average age at death for the entire 2187 is 65.48 years.

TABLE I

Mortality Table of 2187 Entomologists Born Between 372 b.c. and 1920
Showing Distribution by Age, Number and Percentage of Total

Age at
death.
Years

No.

dead

Per cent
of total

Age at
death.
Years

No.

dead

Per cent
of total

Age at
death.
Years

No.

dead

Per cent
of total

19

1

0.045

46

21

0.96

73

69

3.15

20

2

0.090

47

21

0.96

74

66

3.01

21

1

0.045

48

27

1.23

75

64

2.92

22

1

0.045

49

24

1.09

76

60

2.74

23

2

0.090

50

25

1.14

77

56

2.56

24

1

0.045

51

23

1.05

78

52

2.37

25

6

0.27

52

21

0.96

79

57

2.60

26

10

0.45

53

29

1.32

80

54

2.46

27

9

0.41

54

19

0.86

81

50

2.29

28

9

0.41

55.

35

1.60

82

42

1.96

29

8

0.36

56

41

1.87

83

35

1.32

30

10

0.45

57

44

2.01

84

36

1.64

31

17

0.77

58

37

1.69

85

29

1.32

32

12

0.54

59

36

1.64

86

27

1.23

33

12

0.54

60

46

2.10

87

24

1.09

34

8

0.36

61

38

1.73

88

24

1.09

35

13

0.59

62

44

2.01

89

18

0.82

36

8

0.36

63

44

2.01

90

11

0.50

37

8

0.36

64

52

2.37

91

10

0.45

38

16 .

0.73

65

59

2.69

92

8

6.36

39

19

0.86

66

49

2.24

93

3

0.13

40

12

0.54

67

62

2.83

94

5

0.22

41

15

0.68

68

59

2.69

95

2

0.090

42

15

0.68

69

51

2.33 •

96

4

0.180

43

13

0.59

70

69

3.15

97

1

0.045

44

14

0.64

71

68

3.11

98

1

0.045

45

22

1.00

72

71

3.24

2187

100.00

According to Table II, which summarizes the distribution of
deaths by age groups, it is apparent that approximately 30 per
cent had died before reaching 60 years. Of the remaining 70

Sept., 1945]

Weiss: Longevity

233

per cent, 23 per cent died between 60 and 69, 29 per cent between
70 and 79 and about 18 per cent between 80 and 98. The same
thing is exhibited graphically by Figure 2 which is a cumulative
curve showing the successive additions of the percentages of
deaths (from Column 3, Table I) at successive ages.

Table III exhibits the average age at death of 2183 entomolo-
gists born between 1500 and 1914. Of the entire original group
of 2187, those born before 1500 and after 1914 were excluded

NO.

DEAD

r

l ■

\

•_

)

i

ifj:

Z'O

V

9

/,!

•

/

i*

»

i

t

1

L

f

t

5

1"

L

4

S

L

r

1

* /

•

4

r

i

J

YV:

V

0 l<

2(

31

40 5

60 7

el

9<

10

IOoTRS.

Fig. 1. Frequency curve showing the distribution of the deaths of 2187
entomologists by ages.

because of the smallness of their number. Entomologists were
scarce during the sixteenth and seventeenth centuries and for
this reason these first two periods cover 100 years each. The
eighteenth century has been split into 10-year periods and the
nineteenth into 5-year periods. The average age at death during
each period within the entire range from 1500 to 1859 shows a
remarkable degree of uniformity. From 1860 to 1914 the aver-
age age at death shows a steady decline. This is because there
are still many entomologists living who were born after 1859.

234 Journal New York Entomological Society [Vol. liii

As practically all who were born between 1500 and 1859 have
now died, the average ages at death for the different periods
between these dates are of special interest. For the 1600 ento-
mologists who were born between 1500 and 1859, the average age
at death for the entire group was 69.09 years. By scanning the
average ages at death for the various periods between 1500 and
1859 in Table III, one may note that the variations from this
average are not extensive. In fact, the standard deviation is
only 3.147.

TABLE II

Distribution of Deaths of 2187 Entomologists by Age Groups

Age group.
Years

Per cent of total

19-24

0.36

25-29

1.94

30-34

2.66

35-39

2.90

40-44

3.13

45-49

5.24

50-54

5.33

55-59

8.81

60-64

10.22

65-69

12.78

70-74

15.66

75-79

13.19

80-84

9.67

85-89

5.55

90-94

1.66

95 and over

0.36 •

100.00

Life tables for early times are mostly incomplete and fragmen-
tary, but Pearl has shown the changing expectation of life at
different periods based on data from various sources. For ex-
ample, in Roman Egypt the expectation of life (or the average
number of years that persons of a given age will probably live)
for a child of about 3 years was a little over thirty years. In
Breslau, for the years 1687 to 1691, the expectation of life at
birth was about 34 years. In Carlisle, England, 1780-1787, the

Sept., 1945]

Weiss: Longevity

235

expectation at birth was close to 40 years, and in continental
United States in 1910, it was about 50 years. In 1940 in conti-
nental United States* the expectation of life at birth for white
males was 62.94 years, and for white females 67.31 years. The
figures in all the life tables show that at all early ages the expec-

X

Fig. 2. Cumulative curve showing the successive additions of the percent-
ages of deaths of 2187 entomologists at various ages. For example at the
age of 60, approximately 30 per cent of the total number had died.

tation of life is longer now than it has been during past times.
And in addition, they indicate that there is no trend toward a
longer life-span.

It is apparent, from Table III, that all during the periods when
the expectation of life at birth was low for populations in general
* Statistical Bull. Met. Life Ins. Co., 22 (12) : 6-8, 1941.

236

Journal New York Entomological Society

[Vol. LIU

TABLE III

Average Age at Death of 2183 Entomologists
Born Between 1500 and 1914

Born from

No.

Range in ages
at death

Average age
at death

1500-1599

12

29-83

60.83

1600-1699

31

34-91

68.03

1700-1709

7 '

54-84

72.85

1710-1719

6

55-72

62.50

1720-1729

18

51-86

67.94

1730-1739

14

31-90

65.64

1740-1749

15

56-88

76.33

1750-1759

23

28-91

70.33

1760-1769

19

45-90

68.42

1770-1779

53

35-89

72.96

1780-1789

53

41-92

71.88

1790-1799

104

38-97

69.66

1800-1804

83

28-96

69.61

1805-1809

78

30-98

70.02

1810-1814

90

33-91

67.53

1815-1819

80

26-91

70.57

1820-1824

103

26-90

68.63

1825-1829

83

26-92

66.53

1830-1834

79

35-94

67.91

1835-1839

124

25-94

68.95

1840-1844

129

29-96

70.17

1845-1849

120

31-94

70.25

1850-1854

128

24-89

67.59

1855-1859

148

28-87

68.20

1860-1864

136

26-82

65.71

1865-1869

118

25-77

62.94

1870-1874

67

37-73

60.01

1875-1879

70 .

27-67

54.18

1880-1884

67

30-63

47.70

1885-1889

45

25-58

44.26

1890-1894

28

27-59

42.25

1895-1899

20

20—47

36.30

1900-1904

14

25-41

36.07

1905-1909

11

23-35

29.90

1910-1914

7

26-29

27.71

and while the average length of life was continually increasing
to its now comparatively high level, entomologists continued to*
live an average of 69 years.

Sept., 1945]

Weiss: Longevity

237

Persons who live long lives generally attribute this fact to
certain habits such as abstinence from alcohol or tobacco, special
diets, exercise or no exercise, fresh air, etc., or to other habits
which personally give them the entire credit for their long life-
spans. As a matter of fact, the length of life of any individual,
barring accidents including infections, etc., is dependent upon
the impetus received during conception and this is largely the
result of hereditary determinants. Karl Pearson reached the
conclusion many years ago that from 50 to 75 per cent of the
general death-rate is determined by the forces of heredity and
is not susceptible of moderation by sanitary measures or preven-
tive medicine. Preventing accidents, and infections, practicing
hygiene, curing diseases, etc., will add to the life-span of many
individuals but the life-span is determined largely at the time
of birth.

In every population from early times to the present a certain
percentage or part of the population has always lived, by reason
of parentage and environment, many years beyond the average
for the balance of the population. Prom the average ages at
death of entomologists who were born between 1500 and 1859,
it is apparent that they have always been recruited, for the most
part, from among that portion of the population that lived the
longest. The gap between the constant high average length of
life of an entomologist in the past and the present average length
of life for the population at large is decreasing because the popu-
lation at large is living longer by reason of the measures taken
to increase national health. Unless the ranks of entomologists
continue to be renewed from that part of the population that
lives the longest, they may not enjoy this advantage in the future.
At any rate, they deserve no special credit for their long lives
however useful they may be. The credit, if any, should go to
their long-lived parents.

238

Journal New York Entomological Society

[Vol. LIII

A LABORATORY AID IN THE ISOLATION OF
ENTOMOGENOUS FUNGI

By E. E. McCoy

The technique here described was developed to facilitate the
preparation of isolation plates of the fungus Beauveria bassiana
occurring on larvas and adults of the Japanese beetle. The dead
insects, even when presenting the typical external growth of
Beauveria spores, are heavily contaminated with a variety of
other microorganisms and the successful preparation of isolation
plates requires that a suspension of discrete cells be made in
water. The spores of most entomogenous fungi are wettable by
water only with difficulty, and the commonly used method for
preparing cell suspensions gives poor results because the mass
is not thoroughly dispersed, so that the contaminating organisms
are carried along with small aggregates of the desired spores.

An excellent dispersal of individual spores may be prepared
by placing a small amount of the fungus growth on a sterile slide
together with a small drop of water, and covering this with a
sterile micro cover. The cover is then moved about in a more or
less rotary fashion by means of a pencil eraser. The disintegra-
tion of the mass can be periodically observed under the com-
pound microscope. When it is seen that the spores are thoroughly
dispersed the cover glass is removed and the suspension on the
slide diluted and pipetted into the usual dilution tubes. Care
must be taken that the grinding action between the slide and
cover glass is not too long continued or the pressure too great,
since the spores are easily crushed or ground and rendered non-
viable.

Sept., 1945]

Goodnight: Phalangida

239

PHALANGIDA FROM THE UNITED STATES

By Clarence J. and Marie L. Goodnight

This present paper is fifth1 in a series in which the authors
are attempting to make the phalangid fauna of the United States
better known. Before the entire group is monographed, it has
been thought better to describe most of the species and to clarify
their relationships and distributions. All holotypes are de-
posited in the collection of The American Museum of Natural
History.

Suborder Laniatores Thorell
Phalangodidae Simon
Phalangodinse Roewer

Wespus arkansasensis Goodnight and Goodnight
(Figs. 7, 8, and 9)

Wespus arkansasensis Goodnight and Goodnight, 1942, Amer.
Mus. Novitates, No. 1188, p. 12, figs. 46, 47, and 48.

Female. — Total length of body, 2.0 mm. Cephalothorax, 0.8
mm. Width of body at widest portion, 1.6 mm.

I II III IV

Trochanter 0.2 mm. 0.2 mm. 0.2 mm. 0.2 mm.

Femur 0.9 0.7 0.8 1.0

Patella 0.4 0.3 0.3 0.4

Tibia 0.7 0.6 0.6 0.9

Metatarsus 0.6 0.7 0.7 1.0

Tarsus 0.8 0.5 0.7 0.5

Total 3.6 mm. 3.0 mm. 3.3 mm. 4.0 mm.

1 See the following papers:

The Genus Protolophus (Phalangida). Amer. Mus. Novitates, No.
1157.

New American Phalangida. Amer. Mus. Novitates, No. 1164.

New Phalangodidae (Phalangida) from the' United States. Amer.
Mus. Novitates, No. 1188.

New and Little Known Phalangids from the United States. Amer.
Midi. Naturalist, Vol. 29, No. 3, pp. 643-656.

240

Journal New York Entomological Society

[Vol. liii

Dorsal scute with five distinct areas, the boundaries of which
are parallel, but difficult to discern. First area without a median
line. Eye tubercle removed from the anterior margin of the
cephalothorax, rounded above, eyes at the base. Eye tubercle
unarmed, but granulate. Entire dorsum roughly granulate, but
without any armature. Venter and coxae roughly granulate.
Third coxa with marginal teeth on the anterior and posterior
margins. Spiracles not visible.

Legs : Clothed throughout with scattered hairs, which are more
numerous on the tarsi. Legs without tuberculations. Metatarsi
of legs divided into a long astragulus and a short calcaneus.
Tarsal segments : 3-6-5-6. Distitarsus of first tarsus with 2
segments, of second 3 segments. The fourth tarsus has 6 seg-
ments rather than 5 as was stated in the original description.
One specimen of the many studied had only 5 segments in the
second tarsus ; the remaining tarsi had the same number as those
of the holotype.

Palpus : Trochanter 0.2 mm. long, femur 0.4, patella 0.4, tibia
0.4, and tarsus 0.3. Total length, 1.7 mm. Palpus armed retro-
laterally as in Figure 9. Prolaterally the femur has two apical
median spines, one of which is larger. The patella has one spine
in the medial portion and one in the apical. The tibia has three
spines, and the tarsus is armed as on the retrolateral side. Dorsal
surface of femur with heavy granulations.

Chelicera normal, clothed throughout with scattered hairs.

Entire body including palpus reddish brown. Free tergites
with darker shadings. Eyes black, contrasting. Appendages
yellowish, lighter than the dorsum.

Male : Total length of body, 2.4 mm. Cephalothorax, 0.8 mm.
Width of body at widest portion, 1.8 mm.

Male similar in appearance to the female ; however each maxil-
lary lobe of the second coxa has a rounded ventral projection.

Records: Washington County, Arkansas, October 6 and 11,
1939, and October 3 and 21, 1941 (Miller and Sanderson). Cen-
treville, Wilkinson x County, Mississippi, July, 1944 (A. F.
Archer) .

Sect., 1945]

Goodnight: Phalangida

241

Suborder Palpatores Thorell
Tribe Dyspnoi Hansen and Soerensen
Nemastomatidse Simon

Nemastoma pallidimaculosa new species
(Figs. 1 and 2)

Male : Total length

of body, 1.4 mm.

Cephalothorax, 0.4 mm.

Width of

body at widest portion

, 0.8 mm.

I

II

Ill

IV

Trochanter

0.2 mm.

0.2 mm.

0.2 mm.

0.2 mm.

Femur

0.9

1.4

0.9

1.3

Patella

0.3

0.3

0.3

0.3

Tibia

1.0

1.8

1.0

1.4

Metatarsus

1.4

3.0

1.7

2.5

Tarsus

1.1

2.2

1.1

1.5

Total

4.9 mm.

8.9 mm.

5.2 mm.

7.2 mm.

Body elongate, eye tubercle low, rounded, near the anterior margin of the
cephalothorax. The male liolotype has colorless eyes at the base. The two
paratypes lack all traces of eyes. Dorsal scute, including the eye tubercle
and cephalothorax, covered with small round granulations. Free tergites
also covered with these small granulations except for the membranes which
separate them. A row of irregularly shaped light splotches along the lateral
margins and the posterior margin of the dorsal scute; these splotches are
without granulations, and extend onto the cephalothorax and up to the eye
tubercle. Free sternites, venter, and coxae thickly covered with the same
type of granulations. First to fourth coxae with teeth on the anterior and
posterior margins. Spiracle very conspicuous.

Legs: Trochanters granulate, remainder of legs smooth, with a few long
scattered hairs on all segments and numerous small, closely adhered hairs on
the tibiae, metatarsi, and tarsi. All femora with a basal false articulation.
Tarsal segments: 8-13-7-7.

Palpus: Trochanter 0.4 mm. long, femur 1.0, patella 0.9, tibia 1.0, and
tarsus 0.4. Total length, 3.7 mm. Palpus very long and slender. All seg-
ments clothed with long hairs which extend at right angles to the segment.
These hairs are more numerous on the tibia and tarsus.

Chelicerge: clothed with scattered hairs. A dorsal median enlargement on
the proximal portion of the second segment.

Entire body brown, with granulations a darker brown. Lateral portion
of the body and membranes between the free tergites lighter. The row of
spots on the abdominal scute lighter and somewhat shiny. Appendages
lighter. Tips of cheliceral claws dark.

Female: Total length of body, 1.6 mm. Cephalothorax, 0.3 mm. Width
of body at widest portion, 0.9 mm.

Similar in appearance to the male, but lacking the enlargement of the
chelicera.

Journal New York Entomological Society

[Vol. LIII

242

Type locality : Male holotype and female paratypes from Rock
House Cave, 1 mile south of Oleander, Marshall County, Ala-
bama, June 23, 1942 (W. B. Jones).

This species differs from Nemastoma inops Packard in the con-
spicuous lateral spots and in the granulation of the body.

Ischropsalidae Simon

Taracus malkini, new species
(Figs. 4, 5, and 6)

Female: Total length of body, 2.0 mm. Cephalothorax, 1.5 mm. Width

of body at widest portion, 3,

.2 mm.

I

II

Ill

IV

Trochanter

0.3 mm.

0.3 mm.

0.4 mm.

0.4 mm.

Femur

1.4

1.2

1.3

1.6

Patella

0.6

0.7

0.6

0.7

Tibia

1.1

1.8

. 1.2

1.5

Metatarsus

1.5

2.3

1.7

2.2

Tarsus

1.3

2.1

1.4

1.8

Total

6.2 mm.

8.4 mm.

6.6 mm.

8.2 mm.

Cephalothorax smooth except for some extremely fine granulations. Eye
tubercle large, not canaliculate, a row of spine-tipped tubercles over each eye.
Just posterior to the eye tubercle is the characteristic spine. It has an en-
larged base, and is situated on a slight elevation. It is directed forward.
Laterad to the spine, on both sides, is a row of several spine-tipped tubercles.
On the last segment of the cephalothorax is a complete transverse row of the
same type of tubercles. Abdomen smooth except for numerous tubercles
which are scattered irregularly over the surface. There is variation in the
size of these tubercles, and little regularity of arrangement. Most, but not
all of the tubercles, are tipped with a small spine-like hair. These hairs are
curved and directed caudad. Venter, coxae, and genital operculum thickly
covered with long black spine-like hairs. These spines are situated on very
small tubercles. Spiracles visible. Maxillary lobe of second coxa club-
shaped and thickly covered with spine-like hairs.

Legs : Trochanters with scattered spine-bearing tubercles which are similar
to those on the coxae. All segments of the legs with numerous small hairs
which are closely applied to the surface. In addition there are the same
spines on all segments but the tarsi. These hair-like spines are somewhat
arranged in rows. All femora with a basal false articulation. Metatarsi
with false articulations. Tarsal segments: 11-16-11-13.

Palpus: Trochanter 0.4 mm. long, femur 1.5, patella 1.1, tifna 0.9, and
tarsus 0.4. Total length, 4.3 mm. Palpus elongate. Femur, patella, and
trochanter with numerous spine-like hairs on small tubercles. Tibia and

V

Sept., 1945]

243

Goodnight: Phalangida

tarsus thickly covered with hairs which extend at right angles to the segment.

Chelicera: Length of proximal segment 1.7 mm. Length of distal segment
2.4 mm. Length of movable claw 0.9 mm. Proximal segment with heavy
spines which are arranged in rows. Distal segment with somewhat less
heavy spines. All these spines are hair-tipped. Chelicera! claw armed as in
Figure 5.

Cephalothorax brown. Eye tubercle brown except for a black ring around
the eye. Spine pale. Abdomen grey with irregular light patches. Spines
on the tubercles black. Legs brown, palpi brown, darker distally. Chelicera
dark brown to black.

Type locality : Female holotype from Manzanita Lake, Lassen
National Park, elevation 5800 feet, California, October 1, 1944
(B. Malkin).

This species is related to T. silvestrii Roewer, differing by hav-
ing the tubercles arranged irregularly on the abdomen and by
having 5 or more tubercles over each eye instead of 3 as in T.
silvestrii.

Tribe Eupnoi Hansen and Soerensen
Phalangiidae Simon
Leiobuninge Banks

Leiobunum gordoni, new species
(Fig. 3)

Female: Total length of body, 5.0 mm. Cephalothorax, 1.8 mm. Width
of body at widest portion, 2.7 mm. Length of femora: I, 9.1 mm.; II, 16.5
mm. ; III, 8.5 mm. ; IV, 12.6 mm.

Dorsum finely granulate. Eye tubercle in the central portion of the
cephalothorax, canaliculate, with a row of spinules over each carina. An-
terior margin of the cephalothorax smooth. Supracheliceral lamella in the
form of an expanded plate. Abdomen subtruncate as seen from above, not
strongly arched. Venter and genital operculum clothed with scattered hairs;
genital operculum and coxae with small tuberculations. Genital operculum
margined with low teeth. Lateral teeth present on the anterior and posterior
margins of coxae I and IV, those on the posterior margin of coxa I very
small; and on the anterior margins only of coxae II and III.

Legs: Trochanters, femora, patellae, and basal portion of tibiae with small
black spines which are more or less arranged in rows. Remaining segments
with a few scattered hairs placed at right angles to the segments and with
many small investing hairs.

Palpus: Trochanter 0.5 mm. long, femur 0.8, patella 0.4, tibia 0.6, and
tarsus 1.2. Total length1, 3.5 mm. Femur elevated only slightly above the
cephalothorax, armed ventrally with small spines. Patella and tibia with
small scattered spines, tibia and tarsus clothed with hairs, palpal claw small
and toothed.

244 Journal New York Entomological Society [Vol. liii

Dorsum yellowish, eye tubercle black (lighter in some specimens) ; base of
tubercle forming the apex of a triangle which broadens and is continuous
with the median dorsal stripe of the abdomen. Lateral portions of cephalo-
thorax bordered with dark brown. Abdominal scute with a dark median
stripe which terminates at the posterior margin of the fifth segment. Second,
third and fourth, abdominal segments indicated by brownish stripes which
begin at the median stripe and terminate at the lateral margins. These have
light spots on them. Other segments indicated by brownish stripes without
the lighter spots. Venter, and coxae slightly lighter than the dorsum.
Palpus light yellowish, distal portion of femur and proximal portion of
patella dark brown. Chelicera light. Legs : trochanters and bases of femora
dark, contrasting with the coxae, remainder of legs lighter brown; patellae
darker. All legs ringed with white at the distal portions of all femora and
tibiae. These white rings are preceded by darker rings.

Male: Total length of body, 3.4 mm. Cephalothorax, 1.3 mm. Width of
body at widest portion, 2.5 mm. Length of femora: I, 10.1 mm.; II, ?; Ill,
9.3 mm.; IV, 12.0 mm.

Similar in appearance to the female. Dorsal stripe less distinct. Penis
alate. The white rings of the tibiae are proceeded by dark brown rings, those
of the femora are not. Remainder of legs yellowish orange, darker on the
distal segments, but much lighter than the legs of the female.

Type locality : Female holotype from Haleyville, Alabama,
July 17, 1943 (D. Sparkman). Paratj^pes from Ranken, Mis-
souri, June 29, 1941 (W. M. Gordon) and from Giant City State
Park, Illinois, July 5, 1944 (Sanderson and Leighton).

This species is closely related to L. relictum Davis differing by
having contrasting trochanters and by the male having an alate
penis.

Roewer in “Die Weberknechte der Erde” has described an
animal under the name of Leidbunum nigropalpi which appears
to have many characters in common with this. The true L. nigro-
palpi, as described by Wood however, has no white annulations
on the legs and has a completely black palpus.

(Jour. N. Y. Ent. Soc.), Vol. LIII

(Plate II)

PHALANGIDA

PLATE II

Figure 1.

Figure 2.
Figure 3.
Figure 4!
Figure 5.
Figure 6.
Figure 7.

Figure 8.
Figure 9.

Nemastoma pallidimaculosa, new species, dorsal view of male
holotype.

Idem, lateral view of eye tubercle of male liolotype.

Leiobunum gordoni, new species, dorsal view of female liolotype.
Taracus malkini, new species, dorsal view of female holotype.
Idem, ventral view of cheliceral claw of female holotype.

Idem, lateral view of eye tubercle of female holotype.

Wespus arJcansasensis Goodnight and Goodnight, lateral view of
eye tubercle of female.

Idem, dorsal view of female.

Idem, retrolateral view of palpus of female.

246

Journal New York Entomological Society

[Vol. LIII

ANCIENT REMEDIES INVOLVING INSECTS

The “Natural History of Pliny” abounds with misinformation
that was once accepted as fact and some of it refers to insects.
For example, a person who carried about with him the beak of
the woodpecker of Mars was never injured by bees, wasps and
hornets. For the bite of a mad dog, one remedy consisted in
attaching to the body of the patient a maggot taken from the
carcass of a dead dog. When the hair was lost, it could be made
to grow again by the application of a mixture of mouse dung and
the heads of flies, applied fresh, the part first being rubbed with
a fig leaf. Other remedies for baldness consisted of applications
of the blood of flies, or of the ashes of burnt flies. In one case
the ashes of burnt flies kneaded with woman’s milk and cabbage
was recommended.

For the bites of all spiders the best remedy was a cock’s brains
taken in oxycrate with a little pepper. Five ants swallowed in
drink was also effective as well as spiders of any kind left to
putrefy in oil.

Nits were destroyed by using dogs’ fat or by eating serpents
cooked like eels. Cantharides were supposed to be produced
from small grubs found particularly in the spongy galls on the
stems of the “dog-rose.” In order to utilize them, they were
put in a small earthen pot, which was covered over with a linen
cloth, on which was placed a layer of full-blown roses. The pot
was then suspended over vinegar boiled with salt, until the steam
penetrated the cloth and stifled them. They they were put aside
for later use. — H. B. W.

Sept., 1945]

Rapp & Cooper: Psychodid^e

247

CHECK-LIST OF AFRICAN PSYCHODID^*

By William F. Rapp, Jr., and Janet L. Cooper

This check-list of African Psychodidae is presented to aid ento-
mologists in working with African specimens.

The genus Flebotomus has been studied by several workers as
some species are carriers of tropical diseases. There are two
notable papers dealing with African species of the genus: one
by Theodor1 giving descriptions and keys and one by Sinton2
dealing with the distribution. Both these papers were written
in the early 1930’s; it should be remembered that much work
has been done since then.

Since none of the other genera is of medical importance, no
particular attention has been paid to them. A great deal of
systematic collecting must be done before we can obtain a com-
plete picture of the species and distribution of African Psy-
chodidae.

The area covered in this check-list includes all Africa, Mada-
gascar, all islands in the Indian Ocean north to the Seychelles
and the Cape Verde Islands in the Atlantic Ocean.

BRUCHOMYIA Alexander

edwardsi Tonnoir, Ruwenzori Exp. 1934-35, Vol. 1, p. 38.
Uganda.

BRUNETTIA Tonnoir

albonotata Brunetti ( Psychoda indica Eaton), Ruwenzori Exp.
1934-35, Vol. 1, p. 76.

Uganda, Seychelles.

gloriosa Tonnoir, Ruwenzori Exp. 1934-35, Vol. 1, p. 73.
Uganda.

grahami Tonnoir, Rev. zool. afric., Vol. 8, p. 143.

Gold Coast.

* Third contribution to a Check-List of the Psychodidae of the World.

1 Theodor, O., “On African Sandflies (Dipt.),” Bull. Ent. Res., Vol. 22
(1931), p. 469-478.

2 Sinton, J. A., “Some New Species and Records of Phlebotomus from
Africa,” Ind. Jour, of Med. Res., Vol. 18 (1930-31), p. 171-193.

248

Journal New York Entomological Society

[Vol. LIII

obscura Tonnoir, Ruwenzori Exp. 1934-35, Vol. 1, p. 77.
Uganda.

pectinata Tonnoir, Bull. Soc. Ent. Egypt, 1921, p. 107.

South Nigeria.

splendens Tonnoir, Rev. zool. afric., Vol. 8, p. 140.

Gold Coast, Uganda.

CLYTOCERUS Eaton

africanus Tonnoir, Rev. zool. afric., Vol. 8, p. 137.

Nigeria.

carbonarius Tonnoir, Ruwenzori Exp. 1934-35, Vol. 1, p. 62.
Uganda.

fasciatus Tonnoir, Ruwenzori Exp. 1934-35, Vol. 1, p. 59.
Uganda.

FLEBOTOMUS Rondani *

adleri Theodor, Bull. Ent. Res., Vol. 24, p. 543.

Gold Coast.

affinis Theodor, Bull. Ent, Res., Vol. 24, p. 545.

Sudan.

africanus Newstead, Bull. Ent. Res., Vol. 3, p. 361.

Algeria, Tunisia, Morocco, Senegal, Gold Coast, Ivory Coast,
Chad Territory, Northern Nigeria, Southern Nigeria, Bel-
gian Congo, Anglo-Egyptian Sudan, Northern Rhodesia,
Nyasaland, East Africa, Portuguese East Africa, Tan-
ganyika Territory, Transvaal, Mauritius.
africanus var. cherifianus Ristorcelli, Ann. Parasit. hum. comp.,
Vol. 17, p. 364.

Morocco.

africanus var. congolensis Bequaert and Walravens, Rev. Zool.
Bot. afr., Vol. 19, p. 38.

Tanganyika (East Africa).

africanus var. longior Parrot, Arch. Inst. Pasteur Algerie, Vol.
14, p. 40.

Ethiopia.

africanus var. magnus Sinton, Ind. Jour. Med. Res., Vol. 20, p.
571.

South Africa.

Sept., 1945]

Rapp & Cooper: Psychodid^

249

africanus var. niger Parrot and Schwetz, Rev. Zool. Bot. afr.,
Vol. 29, p. 226.

Belgian Congo.

africanus var. sudanicus Theodor, Bull. Ent. Res., Vol. 24, p.
541.

Sudan.

antennatus Newstead, Bull. Ent. Res., Vol. 3 (1912), p. 365.

Gold Coast.

bedfordi Newstead, Bull. Ent. Res., Vol. 5, p. 179.

Transvaal.

brodeni Parrot, Rev. Zool. Bot. afr., Vol. 19, p. 185.

Belgian Congo.

buxtoni Theodor, .Bull. Ent. Res., Vol. 24, p. 544.

Gold Coast.

congolensis var. distinctus Theodor, Bull. Ent. Res., Vol. 24,
p. 542.

Gold Coast, Sudan.

decipiens Theodor ( Flebotomus simillimus Adler, Theodor, Par-
rot), Bull. Ent. Res., Vol. 22, p. 473.

East Africa.

duboscqui Neveu-Lemaire, Bull. Soc. Zool. de France, Vol. 31
(1906), p. 65.

French Sudan, Mauretania, Chad Territory, Ashanti, South-
ern Nigeria, Sierra Leone.
dureni Parrot, Rev. Zool. Bot. afr., Vol. 24, p. 266.

Belgian Congo.

fallax Parrot, Arch. Inst. Pasteur Afrique de Nord, Vol. 1, p. 99.
Algeria, Tunisia.

freetownensis Sinton, Ind. Jour. Med. Res., Vol. 18, p. 188.
Sierra Leone.

gig as Parrot and Schwetz-, Rev. Zool. Bot. afr., Vol. 29, p. 224.
Belgian Congo.

ingrami Newstead, Bull. Ent. Res., Vol. 5, p. 179.

Ivory Coast, Northern Ashanti, Uganda.
legeri Mansion, Bull. Soc. Path, exot., Vol. 6. p. 640.

Bastia.

longipes Parrot and Martin, Arch. Inst. Pasteur Algerie, Vol.
17, p. 143.

Abyssinia.

250

Journal New York Entomological Society

[VOL. LIII

katangensis Bequaert and Walravens, Rev. Zool. Bot. afr., VoL
19, p. 35.

East Africa.

langeroni Nitzulescu, Ann. Parasit., Vol. 8, p. 547.

Tunis.

langeroni var. longicuspis Nitzulescu, Ann. Parasit., Vol. 8, p.
551.

Tunis.

langeroni var. orientalis Parrot, Arch. Inst. Pasteur Algerie, Vol.
14, p. 30.

Ethiopia.

mdjor var. perniciosus Newstead, Bull. Ent. Res., Vol. 2 (1911),
p. 204.

Algeria, Tunisia, Morocco, Spanish Guinea.
martini Parrot, Arch. Inst. Pasteur Algerie, Vol. 14, p. 35.
Ethiopia.

matadiensis Theodor, Bull. Ent. Res., Vol. 29, p. 169.

Belgian Congo.

mathisi Parrot, Arch. Inst. Pasteur Algerie, Vol. 13, p. 259.

Senegal, French West Africa.
meilloni Sinton, Ind. Jour. Med. Res., Vol. 20, p. 565.

Africa.

meilloni var. suberectus Sinton, Ind. Jour. Med. Res., Vol. 20, p.
565.

Africa.

minutus Rondani, Annal. Soc. Entomol. France, ser. 2, I (1843), •
p. 265.

Algeria, Tunisia, French Sudan, Sierra Leone, Anglo-
E gy ptian Sudan, Italian Somaliland.
minutus var. africanus, Bull. Ent. Res., Vol. 3 (1912), p. 363.

Gold Coast, Southern Nigeria, Northern Nigeria, Anglo-
Egyptian Sudan, North Eastern Rhodesia, Nyasaland.
minutus var. antennatus Newstead, Bull. Ent. Res., Vol. 3, p.
361.

Northern Ashanti, Gold Coast.

minutus var. fallax Parrot, Bull. soc. hist, nat, Alger., Vol. 12,
p. 37.

Algeria.

Sept., 1945]

Rapp & Cooper: Psychodid.®

251

minutus var. occidentalis Theodor, Bull. Ent. Res., Yol. 24, p.
539.

Gold Coast, Nigeria.

mirabilis Parrot and Wanson, Rev. Zool. Bot. afr., Yol. 32, p. 149.
Belgian Congo.

nairobiensis Theodor, Bull. Ent. Res., Vol. 22, p. 472.

East Africa, Kenya, Uganda.

notatus Parrot, Arch. Inst. Pasteur Algerie, Yol. 16, p. 216.
Ethiopia.

palestinensis Adler and Theodor, Ann. Trop. Med. Parasit., Vol.
21, p. 64.

Algeria.

papatasii (Scopoli) ( Bibo papatasii Scopoli, Deliciae faun., et
flor. Insubriciae, Vol. 1 (1786), p. 55) ( Cynipkes molestus
Costa, Storia dei lavori dell Acald. Aspir. Natural Artie.
Zool. (1840)), ( Hermasson minutus Loew, Stettin. Ent.
Zeil., Yol. 5 (1844), p. 115).

Maltese Islands, Anglo-E gy ptian Sudan, Red Sea Province,
Egypt, Algeria, Tunisia, Morocco, French Sudan, Spanish
Guinea, Italian Somaliland.

papatasii var. bergeroti Parrot, Arch. Inst. Pasteur Algerie, Yol.
12, p. 383.

Central Sahara.

papatasii var. breviventris Ristorcelli, Ann. Parasitol. Humaine
et Comp., Yol. 17 (4) (1939), pp. 364, 365.

Libya, Morocco.

parroti Adler and Theodor, Ann. Trop. Med. Parasit., Vol. 21,
p. 64.

Algeria, Tunisia.

renauxi Parrot and Schwetz, Rev. Zool. Bot. afr., Vol. 29, p. 222.
Belgian Congo.

rodkaini Parrot, Rev. Zool. Bot. afr., Yol. 19, p. 187.

Belgian Congo.

roubaudi Newstead, Bull. Ent. Res., Vol. 3 (1912), p. 361.
Mauretania.

sanneri Galliard and Nitzulescu, Ann. Parasit. hum. comp., Yol.
9, p. 238.

French Equatorial Africa.

252

Journal New York Entomological Society

[VOL. LIII

schwetzi Adler, Theodor and Parrot ( F . symesi Sinton), Rev.
Zool. Bot. afr., Vol. 18 (1929), p. 73.

Kenya.

schwetzi var. cethiopicus Parrot, Arch. Inst. Pasteur Algerie,
Yol. 14, p. 39.

Ethiopia.

sergenti Parrot, Bull. Soc. path, exot., Yol. 10 (1917).

Algeria, Tunisia, Morocco.

sergenti var. scevus Parrot and Martin, Arch. Inst. Pasteur
Algerie, Yol. 17, p. 484.

Ethiopia.

shorttii Adler and Theodor, Ann. Trop. Med. Parasit., Yol. 21,
p. 65.

Egypt,

signatipennis Newstead, Bull. Ent. Res., Vol. 11, p. 310.

Gold Coast.

simillimus Newstead, Bull. Ent. Res., Yol. 5, p. 179.

Gold Coast, Southern Nigeria.

squamipleuris Newstead ( F . ghesquierei Parrot), Bull. Ent.
Res., Yol. 3 (1912), p. 366.

Anglo-Egyptian Sudan, Gold Coast, Mozambique.
squamipleuris var. dreyfussi Parrot, Arch. Inst. Pasteur Algerie,
Vol. 11, p. 605.

North Africa.

squamipleuris var. idicus Theodor, Bull. Ent. Res., Yol. 22
(1931), p. 470.

Africa.

squamipleuris var. inermis Theodor, Bull. Ent. Res., Yol. 29, p.
165.

North Nigeria.

subtilis Parrot and Martin, Arch. Inst. Pasteur Algerie, Yol. 17,
p. 151.

Abyssinia.

symesi Sinton, Ind. Jour. Med. Res., Yol. 18, p. 175.

East Africa.

thomsoni Theodor, Bull. Ent. Res., Vol. 29, p. 167.

Belgian Congo.

transvaalensis Sinton, Ind. Jour, Med. Res., Yol. 20, p. 879.
Transvaal.

Sept., 1945]

Rapp & Cooper: Psychodidje

253

troglodytes Nitzulescu, Ann. Parasit., Yol. 8, p. 388.

Tunisia.

vagus Parrot and Martin, Arch. Inst. Pasteur Algerie, Vol. 17,
p. 147.

Abyssinia.

viator Parrot and Martin, Arch. Inst. Pasteur Algerie, Yol. 17,
p. 153.

Abyssinia.

viduus Parrot, Arch. Inst. Pasteur Algerie, Yol. 14, p. 34.
Ethiopia.

wansoni Parrot, Rev. Zool. Bot. afr., Vol. 30, p. 361.

Belgian Congo.

wurtzi Parrot, Arch. Inst. Pasteur Algerie, Yol. 16. p. 213.
Ethiopia.

yusafi Sinton, Ind. Jour. Med. Res., Vol. 18, p. 181.

East Africa.

yvonnae Parrot and Schwetz, Rev. Zool. Bot. afr., Yol. 29, p. 221.
Belgian Congo.

NEMOPALPUS Alexander

capensis Edwards, Ann. Mag. Nat. Hist., Vol. 3 (10) (1929),
p. 422.

South Africa.

flavus Macquaert, in Webb and Berth : Hist, Nat. d’lles Canaries,
Ent., Yol. 1, Canarienses Dipt. (1838), p. 102.

Canary Islands.

NOTIOCHARIS Eaton

insignis Eaton, Trans. Linn. Soc. Lond., Yol. 15, p. 424.
Seychelles.

PANIMERUS Eaton

scotti Eaton, Trans. Linn. Soc. Lond., Yol. 15, p. 425.

Seychelles.

PERICOMA Eaton

ombigua Eaton, Ent. Mon. Mag., ser. 2, IY (XXIX) (1893),

p. 126.

Northern Africa.

canariensis Tonnoir, Ann. Soc. ent. Belg., Yol. 62, p. 167.
Canary Islands.

254

Journal New York Entomological Society

[Vol. LIII

cxquisita Eaton, Ent. Mon. Mag., ser. 2, IV (XXIX) (1893),

p. 122.

Northern Africa.

meridionalis Eaton, Ent. Mon. Mag., ser. 2, V (XXX) (1894),
p. 194.

Africa.

palustris Meigen, Klass I., 43, 2 (Trichoptera) (1804).

Northern Africa.

pseudocanescens Abren, Mem. Roy. Acad. Barcelona, Yol. (3) 22
(1930), p. 97.

Canary Islands.

unicolor Abren, Mem. Roy. Acad. Barcelona, Yol. (3) 22 (1930),
p. 98.

Canary Islands.

ustidata Walker, Ins. Britannica, Dipt., Vol. 3 (1856), p. 258.
Northern Africa.

vicina Abreu, Mem. Roy. Acad. Barcelona, Yol. (3) 22 (1930),

p. 106.

Canary Islands.

PHILOSEPEDON Eaton

triangulatus Eaton, Trans. Linn. Soc. Lond., Yol. 15, p. 429.
Seychelles.

PSYCHODA Latreille

aberrans Tonnoir, Bnl. Soc. Ent. Egypt., 1921, p. 84.

Egypt,

acuta Tonnoir, Ruwenzori Exp. 1934-35, Yol. 1, p. 54.

Uganda.

albida Tonnoir, Ruwenzori Exp. 1934-35, Yol. 1, p. 52.

Uganda.

albidonigra Tonnoir, Jour. E. Afr. Ug. Nat. Hist. Soc., Yol. 14,

p. 12.

Kenya.

albipennis Zetterstedt, Dipt. Scand., 3708, Yol. 10 (1850).
Northern Africa.

alternata Say, Long’s Exp. St. Peter’s River, App. (1824), p.
358.

(For synonyms see Jour. N. Y. Ent. Soc., Yol. 52, p. 204.)
Northern Africa.

Sept., 1945]

Rapp & Cooper: Psychodid^e

alternaia var. marmorosa Abreu, Mem. Roy. Acad. Barcelona,
Vol. (3) 22 (1930), p. 123.

Canary Islands.

amphorica Tonnoir, Ruwenzori Exp. 1934-^5, Vol. 1, p. 49.
Uganda.

bilobata Tonnoir, Ruwenzori Exp. 1934-35, Vol. 1, p. 50.
Uganda.

dentata Tonnoir, Ruwenzori Exp. 1934r-35, Vol. 1, p. 53.

Uganda.

deviata Tonnoir, Ruwenzori Exp. 1934—35, Vol. 1, p. 46.

Uganda.

dubitata Tonnoir,. Ruwenzori Exp. 1934-35, Vol. 1, p. 57.

Kenya.

efflatouni Tonnoir, Bui. Soc. Ent. Egypt, 1921, p. 81.

Egypt.

erminea Eaton, Ent. Mon. Mag., ser. 2, IV (XXIX) (1893),
p. 130.

Northern Africa.

humeralis var. mauritanica Eaton, Ent. Mon. Mag., ser. 2, IX
(XXXIV) (1898), p. 157.

Northern Africa.

ingrami Tonnoir, Bull. Soc. Ent. Egypt, 1921, p. 88.

Gold Coast.

latipennis Tonnoir, Ruwenzori Exp. 1934-35, Vol. 1, p. 56.
Uganda.

latisternata Tonnoir, Jour. E. Afr. Ug. Nat. Hist. Soc., Vol. 14,

p. 11.

Kenya.

maxima Tonnoir, Ruwenzori Exp., 1934-35, Vol. 1, p. 41.
Uganda.

modesta Tonnoir, Ruwenzori Exp., 1934-35, Vol. 1, p. 43.
Uganda.

nana Tonnoir, Bull. Soc. Ent. Egypt, 1921, p. 93.

Cote de UOr.

nocturna Abreu, Mem. Roy. Acad. Barcelona, Vol. (3) 22 (1930),

p. 115.

Canary Islands.

256

Journal New York Entomological Society

[Vol. liii

nocturna var. nigrithorax Abreu, Mem. Roy. Acad. Barcelona,
Vol. (3) 22 (1930), p. 115.

Canary Islands.

obscura Abreu, Mem. Roy. Acad. Barcelona, Vol. (3) 22 (1930),
p. 109.

Canary Islands.

pallida Tonnoir, Bull. Soc. Ent. Egypt, 1921, p. 96.

Cote de l’Or, Belgian Congo.

pkalcenoides Linnaeus, Syst. Nat., Ed. X, 588, 32 (Tipula) (1758)
( muraria Latreille, Hist. Nat. et Crust, et Ins., XIV (1805),
p. 203) ( nervosa Schink, Fauna Bioca, III, 82, 2350 (Tipula)
(1803).

Northern Africa.

plumosa Tonnoir, Ruwenzori Exp., 1934—35, Vol. 1, p. 56.
Uganda.

pseudoalbipennis Abreu, Mem. Roy. Acad. Barcelona, Vol. (3) 22
(1930), p. 114.

Canary Islands.

pseudomaxima Tonnoir, Ruwenzori Exp., 1934-35, Vol. 1, p. 44.
Uganda.

reducta Tonnoir, Ruwenzori Exp., 1934r-35, Vol. 1, p. 42.

Uganda.

undulata Tonnoir, Ruwenzori Exp., 1934-35, Vol. 1, p. 47-.
Uganda.

solitaria Eaton, Trans. Linn. Soc. Loud., Vol. 15, p. 429.
Seychelles.

SETOMIMA Enderlein

litkocolleta Enderlein, Dtsch. ent. Z., 1936.

French Equatorial Africa.

SYCORAX Curtis

africanus Tonnoir, Rev. zool. afric., Vol. 8, p. 146.

Uganda.

silacea Curtis, Brit. Ent., Vol. 10 (1839), p. 745.

Northern Africa.

SYNSEODA (also as SYNEODAID) Enderlein

flavitarsis Enderlein, Dtsch. ent. Z., 1936, p. 92.

Uganda.

Sept., 1945]

Bapp & Cooper: PsychodiLe

TELMATOSCOPUS Eaton

crassiascoidatus Tonnoir, Ruwenzori Exp., 1934-35, Yol. 1, p. 68.
Kenya.

edwardsi Tonnoir, Ruwenzori Exp., 1934-35, Vol. 1, p. 64.
Uganda.

fryeri Eaton, Trans. Linn. Soc. Lond., Yol. 15, p. 430.

Aldabra, Indian Ocean.

fuscipennis Tonnoir, Rev. zool. afric., Yol. 8, p. 136.

West Africa.

fuscus Tonnoir, Ruwenzori Exp. 1934-35, Yol. 1, p. 65.

Uganda.

meridionalis Tonnoir, Rev. zool. afric., Yol. 8, p. 127.

West Africa.

pallidus Tonnoir from Psychoda, Ruwenzori Exp., 1934—35, Yol.

1, p. 62.

Uganda.

pectinatus Tonnoir, Ruwenzori Exp., 1934-35, Yol. 1, p. 66.
Uganda.

squamifer Tonnoir, Bull. Soc. Ent., Egypt, 1921, p. 102.

Egypt.

THRETICUS Eaton

compar Eaton, Ent. Mag’., Vol. XL, p. 57.

Algeria.

258

Journal New York Entomological Society

[Vol. LIII

EARLY ENTOMOLOGICAL MANUSCRIPTS

After the death of Dr. George H. Horn, the Academy of
Natural Sciences of Philadelphia acquired his library, which
included Dr. Christian Zimmerman’s note book or diary. A. copy
of this diary is also in the library of the Museum of Comparative
Zoology, and at one time, the late Dr. W. H. Wheeler contem-
plated translating it from the German and preparing it for
publication. It is my opinion that this should be done for the
sake of future students of the history of American entomology.

Another set of manuscript notes and entomological drawings
that, should be made available to students are those of Jacob Cist,
an early entomologist who corresponded with Thomas Say and
who lived in Wilkes Barre, Pa. These notes and drawings are
in the possession of Mr. Gilbert S. McClintock, of Wilkes Barre.
Mr. Cist was a versatile person of much skill and inventiveness,
and it is hoped that some day more information will be available
about his entomological activities. — H. B. W.

Sept., 1945]

Needham: Gall

'259

A PEDICEL GALL ON TRICHOSTEMA

By James G. Needham

There is a little mint flower that grows on Longboat Key oppo-
site Sarasota. Florida. It is a bushy species of Trichostema of
boot-top height. It branches from the base and forms a rounded
heap of rather dense evergreen foliage, above which it rears a few
tall leafless open panicles of inflorescence. In the slender termi-
nal pedicels of these panicles I found specimens of the little
woody gall that is the subject of this paper. The moth that
causes the gall is a slender black and white lavernid kindly deter-
mined for me by Dr. W. T. M. Forbes as Mompha sexnotella
Chambers.

The plant is Trichostema suffrutescens. It grows in the outer
edges of rounded tussocks of grass, sharing this situation with the
much commoner horse-mint, Monarda dispersa. Between the tus-
tocks are areas of bare soil; a hard and sterile soil, composed of
marine shells washed up by the waves from the Gulf of Mexico,
and in all stages of disintegration. The narrow strips of this
peculiar meadow lie behind an aggrading storm- wave reef. Here
and there in them are patches of white wind-blown sand, partly
covered by the soft green mats of a lupine, Lupinus diffusus:
Wherever the soil has been disturbed (as around the burrows of
gopher turtles) the low sunflower, Helianthus debilis, spreads its
halberd-shaped leaves and procumbent branches. It blooms there
gloriously in January.

I found the galls there on January 5 at the end of the flowering
season. The massed foliage was fresh and green and scented with
a delicate fragrance, less pungent than that of the horse-mint.
The fruiting panicles at the top were bare and brown, each termi-
nal twig bearing the persistent strongly bilabiate calyx. In some
of the calyces there still nestled four, whitish, pock-marked seeds.

Here and there was a terminal twig that had been made over
into an ocarina-shaped gall of about the size of a peanut, A few
of the galls were still green with a wash of reddish color on the
side -exposed to the sun ; their thicker tissues had dried up more

260

Journal New York Entomological Society

[Vol. LI'II

slowly than the rest of the panicle. Most of them were brown,
some of the older ones were black. Their size made them easy
to find among* the slender twigs.

The gall is short-spindle-shaped, 15 to 20 millimeters long, 5 to
8 wide, smoothly contoured and slightly inequilateral. Its walls
are woody, thick and very hard. The entire gall involves two
internodes of the twig, and the node between the two lies near the
distal end of it at a place that is marked by the vestiges of what
would normally be a pair of opposite branchlets. There is a little
low protuberance on one side near the base — a predetermined
point at which the exit door will be made for the emergence of the
adult insect.

The cavity of the gall lies mainly in the lower internode. It is
oval and wide, loose-fitting for the caterpillar, and it tapers up-
ward to a narrowing tube in the distal internode which serves the
occupant for a. frass receptacle.

Inside the woody layer of the wall there is another hard and
very brittle layer whose components I was unable to determine.
It occupies the position of the layer of nutritive tissue that feeds
the larva during its development ; but in the old galls, containing
only full-fed larvae, this layer is nearly as hard and as thick as
the wood, and much more brittle. At first I found it difficult to
get a larva out uninjured ; for the gall cannot be split and pried
open without crushing this inner hard layer. I found it possible
by sticking a knife point through the softer distal end of the gall,
where the frass receptacle is, and breaking off that portion, then
putting the knife point inside the hole thus laid open and pushing
outward, so chipping the remaining wall away in pieces, the larva
could be uncovered and withdrawn uninjured. In this process
the cylindrical, brittle layer would sometimes remain intact, en-
tirely freed from the wood. Perhaps some frass enters into the
composition of the brittle layer; for the frass in the over-filled
receptacle seems to be continued down the widening sides of the
gall under a thin transparent layer of silk.

In cutting open a score or more of galls I found some of them
empty, the exit hole open, and the empty abandoned pupal skin
of the moth left lying inside. Most of the galls contained fat
larvse, nestled down where the gall narrows to the exit door with

Sept., 1945]

Needham: Gall

261

only a thin pellicle of epidermis closing the door. One gall had
a young and active larva still engaged in clearing the exit passage
way, the silk lining of the gall completed only in its upper half.

Notwithstanding hard walls for its protection, the gallmaker
has enemies that find it. Two of the galls first examined con-
tained adult braconid parasites; adults apparently about ready
for emergence. They had eaten the moth larva and then they
had spun their own slender cocoons obliquely across the gall
cavity. The empty skins of the caterpillars lay beside the cocoons
of the parasites. The braconids had bored their own holes for
exit ; round holes about half as large as the moths would require,
situated near the middle of the gall. They made no use of the
moth’s easier way out at the lower end of the gall. Twelve more
of these braconids emerged later from galls that I had placed in
the rearing cages. The adults were sent to Dr. C. F. W. Muese-
beck for determination. He reports that they represent an unde-
scribed species of the braconid genus Heterospilus. The speci-
mens are now in the United States National Museum.

The moth larvge also have larger enemies. I found about a
dozen of the galls had been gnawed open apparently by the teeth
of rodents; possibly by cotton rats. Mr. William Hegener was
catching these animals in traps in this same strip of shoreland
meadow.

The emergence of the adult moths was slow and irregular.
From galls collected on January 5, the first moth came out on
the last day of the month ; and others came out at long intervals
for two months thereafter.

The living adult is a delight to the ey§ : a slender resplendent
mothlet, ringed and spotted with black and white in perfect
camouflage ; its long wings shedding bronze and coppery reflec-
tions, and on the middorsal line, where fore wings meet and cover
the feathery edgings of the hind wings, tufts of elevated scales
rise in three little rounded heaps that shine like globules of quick-
silver.

262

Journal New York Entomological Society

[Yol. liii

A NEW NAME FOR PERICOMA UNICOLOR
ABREU

By William F. Rapp, Jr,

Recently while working* on a check-list of Psychodidge of the
world I found that Dr. D. Elias Santos Abreu had made a homo-
nym of Pericoma unicolor (Men. Acad. Cie. y Artes, Barcelona,
Vol. 22 (1930), p. 98). Brunetti in the “Record of the Indian
Museum/’ Yol. 4 (1911), p. 309 described Pericoma unicolor from
India. Therefore, I have attempted to contact Doctor Abreu and
bring this matter to his attention, but was unsuccessful. I pro-
pose the name Pericoma abreui nom. nov. for Pericoma unicolor.
The type locality is Canary Island.

Vol. LIII No. 4

DECEMBER, 1945

Journal

of the

New York Entomological Society

Devoted to Entomology in General

Edited by HARRY B. WEISS

Publication Committee

HARRY B. WEISS JOHN D. SHERMAN, Jr.

T. C. SCHNEIRLA

Subscription $3.00 per Year

Published Quarterly by the Society
N. QUEEN ST. AND McGOVERN AVE.
LANCASTER, PA.

NEW YORK, N. Y.

1945

CONTENTS

On the Occurrence of Impaternate Females in the For-
micidse

By Caryl P. Haskins and Ernst V. Enzmann 263

Records and Descriptions of Neotropical Crane-Flies
(Tipulidae, Diptera), XX

By Charles P. Alexander 279

Seasonal Variations in Certain Species of Mosquitoes
(Diptera, Culicidae)

By Charles D. Michener 293

Some Early Entomological Ideas and Practices in America

By Harry B. Weiss 301

New Psychodidae from Barro Colorado Island

By William F. Rapp, Jr 309

Neuropathology in Insects

By A. Glenn Richards, Jr., and Laurence K. Cut-
KOMP 313

NOTICE : Volume LIII, Number 3, of the Journal of The
New York Entomological Society was published
October 8, 1945.

Entered as second class matter July 7, 1925, at the post office at Lancaster, Pa.,
under the Act of August 24, 1912.

Acceptance for mailing at special rate of postage provided for in Section 1103.
Act of October 3, 1917, authorized March 27, 1924.

JOURNAL

OF THE

New York Entomological Society

Vol. LIII December, 1945 No. 4

ON THE OCCURRENCE OF IMPATERNATE
FEMALES IN THE FORMICIDiE

By Caryl P. Haskins and Ernst V. Enzmann
INTRODUCTION

The phenomenon of parthenogenesis in the Hymenoptera has
excited the continuing interest of entomologists for a relatively
long period. The studies of Dzierzon (1845) on the honeybee
belong to a rather early era of entomological thinking, and his
classic “rule” paved the way for much later work. The conclu-
sion of Dzierzon that arrhenotocous parthenogenesis is the in-
variable rule in the honeybee has been confirmed many times,
although it has been questioned by other workers. The fact that
males may be totally absent among certain of the lower Hymen-
optera, so that infertile ova must of necessity develop into
females, was observed by Foerster (1856) as early as 1850 in
the eulophid Astichus arithmeticus. The phenomenon has since
been found to be of common occurrence among the Tenthredinidge
and other sawflies, and to be widespread in the Terebrantia.
Ampherotocous parthenogenesis is also common among the Phy-
tophaga and the Terebrantia.

The fact that, while males are usually of haploid constitution,
females, whether produced uniparentally or biparentally, are
diploid is as well established among the lower as among the
higher Hymenoptera.1 It follows that among species in which

1 The exceptional diploid males of TLabrobracon observed by the Whitings
and their colleagues (1940; 1941) have been found to arise only among
closely inbred stocks, consistently with the hypothesis that heterozygosity

263

©EC. 1 2 '45

264

Journal New York Entomological Society

[Vol. LI1I

ampherotocous or thelytocous parthenogenesis occurs, diploid as
well as haploid ova must be produced. This situation has been
analyzed for the Terebrant genus Habrobracon, where it is known
that diploid ova are produced which develop into females.
Whiting (1928) and Speicher (1934) first accounted for the con-
dition as the result of an abnormal second division of the oocyte,
or its complete suppression, but Speicher and Speicher (1938)
have more recently concluded that such diploid, female-producing
eggs arise from patches of tetraploid ovarian tissue.

The occurrence of uniparental females is such a common phe-
nomenon among the lower Hymenoptera that it is natural to
question whether the conclusion of Dzierzon for the honeybee
is universally applicable among the higher Aculeates. Numerous
apparent exceptions to Dzierzon ’s rule have been reported among
the social Hymenoptera. Many of these have been subjected to
doubt, however, because of the greater difficulty of thoroughly
controlling experiments to prove the point among the social
Aculeates. Thus Onions (1912) and Jack (1917) reported the
production of workers and even queens from worker ova in Apis
mellifica. More recently Mackensen (1943), working with virgin
females of the honeybee; has found indication that uniparental
females may be regularly produced, although only to the extent
of about one per cent of the normal brood. The evidence of
Descy on Osmia (1924), of Stockhert on Halictus (1923), and of
many observers on Bombus and Apis, however, has been entirely
negative.

The case among ants is even less clear and more controversial,
and although the production of impaternate queens or workers
was frequently claimed in the early years of this century, recent
workers have been more skeptical. Thus Tanner (1892) re-
ported the production in the artificial nest of adults of both
sexes and all castes from eggs of workers of Atta cephalotes.
Reichenbach (1902), in a series of observations extending over
three years, described the consistent production of workers as
well as males in a colony of Lasius niger containing originally

of alleles at the sex locus rather than the mere fact of diploidy is the deter-
mining factor of femaleness, and homozygosity rather than haploidy of male-
ness. The production of males from fertilized eggs in the Chalcidid genus
Copidosoma, reported by Leiby (1922) is however less clear.

Dec., 1945]

Haskins & Enzmann: Formiche®

205

only workers. This claim was later substantiated by Crawley
(1912) for the same insect. Comstock (1903) similarly observed
the maturation of workers from worker eggs in L. niger ameri-
canus. Wheeler (1903) has presented a fairly extensive review
of the cases of uniparental females recorded for ants in the late
nineteenth and early twentieth centuries. On the other hand
Forel (1874), Lubbock (1888), Fielde (1901), Janet (1909), and
Tanquary (1913), whose collective observations were both exten-
sive and carefully controlled, reported entirely negative results.

It is to be remembered that this early work was undertaken
without a knowledge of certain pertinent factors in the colonial
economy of the Formicidae which are much better understood
today. It was realized that eggs of worker origin may consti-
tute a considerable proportion of the total number of ova pro-
duced and reared to maturity in normal ant colonies containing
one or more queens. It was not recognized, however, that fecun-
dation of workers — as well as of young infertile queens — may
frequently take place within the parent nest in advance of any
mating flight, and that worker ants are often capable of such
fertilization. Miss Fielde, to be sure, took adequate precautions
in her work to eliminate this possibility. It is much less certain
that this source of error was positively eliminated in the case of
other observers, including many of those reporting the occurrence
of impaternate females.

Recently Goetsch (1937), in a very extensive study involving
six species of the leaf-cutting Myrmicine genus Acromyrmex and
three of Atta, as well as representatives of the genera Pheidole,
Solenopsis, Camponotus, and Lasius , has concluded that unipa-
rental ova among ants produce only males, and Goetsch and
Kathner (1937) report a similar conclusion from a study of
progeny of virgin females of Lasius. These authors have also
reviewed the reported cases of uniparental females among ants
and have decided rather definitely that all are spurious, being
accountable in many cases by intranidal fertilization of 1 1 virgin ’ ’
queens or workers unknown to the experimenter.

In view of the wide divergence in the reports of different
observers and because of the common occurrence of impaternate
females among the lower Hymenoptera, it would seem desirable

266

Journal New York Entomological Society

[Yol. liii

that additional experimental data should be secured before the
final conclusion is reached that only arrhenotocous parthenogene-
sis occurs among the Formicidae. The present program was
undertaken in an effort to secure a certain amount of such data.
The results reported are preliminary and include relatively few
cases. Nevertheless, they seem sufficiently suggestive to publish
at the present time.

EXPERIMENTAL CONSIDERATIONS, MATERIAL, AND
PROCEDURES

Experiments designed to test the occurrence of thelytocous or
ampherotocous parthenogenesis among ants must satisfy a num-
ber of requirements, some of which are inherent in any careful
work with the Formicidae. Optimum conditions of the normal
environment with respect to physical features of the nesting
situation, temperature, humidity, and food supply must be dupli-
cated as closely as possible. These are often complex and spe-
cialized. Colony fragments should be made as small as is con-
sistent with approximately normal social behavior, and the
observation of isolated individuals is a great desideratum when-
ever possible. Both colonies and isolated individuals should be
thoroughly habituated to the experimental situation before work
is begun. Rigid precautions should of course be taken against
the accidental insemination of infertile females or workers within
the nest. Full controls should be kept and the data recorded
should be extensive enough to permit of adequate statistical
treatment. A considerable effort was made to meet all these
requirements except the last one in the work here reported.

Two species of the temperate-zone Myrmicine genus Aphceno-
gaster Mayr were selected for this work, Aphcenog aster fulva aquia
and its variety picea and A. lamellidens. The first species is a
soil-nesting type by preference, is extremely- widespread and
abundant in distribution in the northeastern and Atlantic states,
and occurs in small to medium-sized colonies. Aphcenog aster
lamellidens usually nests in decayed logs, in colonies which at
maturity are somewhat more populous on average than those of
A. fulva. It is somewhat more restricted in distribution, but
is abundant throughout its range. Queens of both species nor-
mally found their colonies in the independent claustral fashion
characteristic of most higher ants.

Dec., 1945]

Haskins & Enzmann: Formicid^:

267

Ants of these species present several advantages as experi-
mental animals. Typically they nest in deeply shaded situations
in woodlands, where the variations of temperature from day to
night and from superficial to deeper layers of soil or wood are
relatively small during the breeding season. This simplifies the
problem of approximating the temperature conditions of the
normal environment very considerably. The species are of both
entomophagous and sweet-feeding habits, and these generalized
alimentary requirements can be fairly accurately met under arti-
ficial conditions. Both ants are unusually tolerant of a wide
range of humidity. The workers of both species are relatively
unspecialized in social function, so that it is more practicable to
work with small colony fragments or even with isolated workers
than is the case with many ants. Workers are usually readily
fertile. The pupae of Aphcenog aster, like those of all Myrmicines,
are naked, and unlike the pupae of higher groups which are
enclosed in cocoons, are able to emerge without assistance. It
is thus possible to rear workers or virgin queens which as adults
have had contact with no other ant. Unlike the condition in
Lasius described by Goetsch and Kathner, fertilization is not
necessary to evoke the normal instincts of colony foundation in
young queens of Aphcenog aster. The entire behavior train in-
volved in colony foundation can be initiated in virgin queens
merely by artificial dealation at maturity. Not infrequently
such females spontaneously dealate themselves if confined in the
artificial nest in the absence of males beyond their normal period
of flight. This situation permits the ready study of single indi-
viduals at a period in the life of the colony when, as Light (1943)
and others have pointed out, it is at its hardiest as well as its
socially simplest stage.

The colonies under investigation were housed in earth-contain-
ing nests of a modified Lubbock type, six by eight inches square
and just sufficiently deep to allow freedom of movement to the
largest members of the colony. The nests were stored in wooden
cabinet drawers seven by eleven inches in inside dimensions and
one and one-quarter inches deep, which were covered with tightly-
fitting glass panes. The upper surfaces of the nests were pro-
tected with sheets of ruby glass except when the colonies were

268

Journal New York Entomological Society

[Vol. LIII

under observation. Each mature colony occupied a separate
draper and the nest-entrances were kept open so that workers
had the opportunity to forage freely at all times within the
confines of the drawers. Small colony fragments, isolated work-
ers, and queens in the process of colony formation were housed
in similar nests but were not permitted to forage. Mature
colonies, colony fragments, and isolated workers were continu-
ously supplied with a diet of strained honey and chopped meal-
worms. Isolated females were fed chopped mealworms from
the time of their isolation but were given no other food. The
inclusion of earth in the nests allowed a much closer approxima-
tion to natural conditions in the physical environment than would
otherwise have been possible. Nearly all colonies of A. picea
maintained an earthen crater of the ragged, scattered form
characteristic of the species outside the nest entrance. It also
greatly facilitated the continuous maintenance of conditions of
normal humidity. Much attention was given to this factor. No
temperature regulation of the nests was attempted. They were
maintained at outdoor ambient temperature during the summer
and at the temperature of a heated room, averaging 21° C.,
during the winter. No effort was made to hibernate the colonies.
This was the experimental condition which deviated most widely
from the normal, but it was not possible to detect significant
abnormalities of behavior or development in colonies maintained
for three or four years under these conditions beyond an acceler-
ated growth of the larvae during the winter and the production
of sexual forms indifferently around the calendar rather than
at sharply circumscribed seasons.

The results presented were obtained with young virgin females
in the isolated condition and with colony fragments composed of
virgin queens and workers of their own colonies. Experiments
with fragments of queenless colonies and with isolated workers
are in progress but are not reported here.

DATA

In the first set of experiments 100 virgin females were dealated
either spontaneously or artificially, were isolated, and were kept
under continuous observation as they attempted to found

Dec., 1945]

Haskins & Enzmann: Formicidjs

269

colonies. They were divided into three groups on the basis of
previous history : those which had been reared from the egg in
old colonies kept in the artificial nest for several years, those
which had been obtained as brood from wild colonies and hatched
under observation, and those obtained from wild colonies as
young winged adults. It was possible to be certain that indi-
viduals of the first two groups had had no contact with males
throughout their development. It was possible that females of
the third class had been fertilized within the nest prior to cap-
ture.

Colony foundation by virgin queens of Aphcenogaster appears
to be difficult and protracted under the best conditions which
can be provided artificially. Only eighteen of the first group of
individuals succeeded. The remainder either rested quiescent
and without brood until the termination of the experiment or
perished at ages of from six months to something over a year.
This is partly to be accounted for by the delayed development
of uniparental larvae and a tendency of the females to destroy
them before maturity. Male pupae were also frequently de-
stroyed as they appeared among the first brood. The period
from dealation of the female to maturity of the first pupa, at
continuous summer temperatures, ranged from three to nine
months with the average well over six months. At the close of
the experiments recorded, which ran between one and two years,
the maximum number of pupae which any infertile female had
reared was five. This was in excess of the number of adults
present in one colony at any time, since young males emerged
continuously if permitted, and perished within the nests if con-
fined to them. This total is to be contrasted with that of the con-
trol, consisting of a young isolated fertile female, which had reared
a single young worker at the time the experiments were begun
and which was given precisely the same treatment as the infer-
tile individuals. At the close of the experiments, the colony of
this control consisted of seventy-six adult workers and a numer-
ous brood.

A detailed account of these eighteen individuals is shown in
Table I.

In the second set of experiments, infertile females were re-
turned to fragments of their own colonies after dealation instead

270

Journal New York Entomological Society

[Vol. LIII

TABLE I

Progeny of Isolated Infertile Females

Parent female

Progeny

Designation

Species

Background

Males

W orkers

Nature and
fate of
pup®

1.

A1

A. picea

*

0

1

Perfect pupa, not seen to hatch

2.

A2

A. picea

*

1

0

Perfect pupa, not seen to hatch

3.

la

A. picea

t

2

0

Both imperfect pupae; destroyed by female

4.

le

A. picea

t

1

0

Perfect pupa, hatched. Wings of one side
imperfectly expanded

5.

Q1

A. picea

t

1

0

Perfect pupa, eclosed perfect adult

6.

Q2

A. picea

t

2

0

Both perfect pupae, eclosed perfect adults

7.

3a

A. picea

t

2

0

One pupa perfect, eclosed perfect adult. Sec-
ond pupa imperfect, destroyed by female

8.

3d

A. picea

t

2

0

One pupa apparently perfect but destroyed.
Second pupa perfect

9.

4b

A. picea

t

1

0

Pupa apparently perfect, but destroyed

10.

4c

A. picea

t

2

0

Both perfect pupae, eclosed perfect adults

11.

4e

A. picea

t

2

0

Both perfect pupae, eclosed perfect adults

12.

4q

A. picea

t

1

0

Perfect pupa, eclosed perfect adult

13.

5d

A. picea

t

5

0,

Four pupae apparently perfect but destroyed.
One pupa perfect, eclosed perfect adult

14.

. 6b

A. picea

t

1

1

Worker pupa perfect, eclosed perfect adult.
Male pupa imperfect, destroyed by female

15.

6d

A. picea

t

3

0

Two pupae apparently perfect but destroyed.
One pupa perfect, eclosed perfect adult

16.

7a

A. picea

t

3

0

One pupa apparently perfect but destroyed.
Two pupae apparently perfect, eclosed
adults with imperfectly expanded wings

17.

7c

A. picea

t

1

0

Perfect pupa, eclosed perfect adult

18.

7i

- A . picea

t

3

0

Two perfect pupae, eclosed perfect adults.
One pupa apparently perfect but de-
stroyed

Totals

32

9

* Female produced in colony long maintained in artificial nest (reared
from egg) .

t Female collected from wild colony as young winged adult.

Dec., 1945]

Haskins & Enzmann: Formicidje

271

of being isolated. Permanent acceptance of such queens under
artificial conditions seems difficult to bring about. Workers ex-
hibit a strong tendency to drive such young females out or to
kill them within a few weeks, and the females on their part fre-
quently strive to leave such colony fragments to establish their
own colonies. From a total of about twenty young females
treated in this fashion, permanent and satisfactory adoption was
secured for only four. The progeny of these females, however,
was much more numerous than in the case of their isolated sisters,
so that the records of numbers are larger. Young were produced
in these colony fragments as indicated in Table II.

TABLE II

Progeny from Colony Fragments Containing Infertile Females

Parent female

Progeny

03

P

©

p

©

Ph

m

bn

rd

P

oS

P

ft

1.

Dl

A. lamellidens *

3

0

Perfect pupae, eclosed perfect adults

2.

D2

A. lamellidens *

22

0

Perfect pupae, eclosed perfect adults

3.

D3

A. lamellidens *

0

8

Perfect pupae, eclosed perfect adults

4.

D4

A. lamellidens *

0

44

Perfect pupae, eclosed perfect adults

Totals

25

52

* Female taken as half -grown larva in wild colony,
ficial nest.

Matured in the arti-

DISCUSSION

Of a total of eighteen isolated infertile young females which
successfully reared young, sixteen produced only males to the
number of thirty-two in all. Two (Al, 6b) each produced a
single worker. One of these subsequently produced a defective
male pupa, which was destroyed prior to maturity and before it
could be thoroughly examined. Of the four colony fragments
containing workers and a sister infertile female which had been
adopted, two (Dl, D2) produced only males, to the number of
25, while two (D3, D4) produced only workers to the number

272

Journal New York Entomological Society

[Vol. LIII

of 52 at the close of the experiments. At that time D4 was still
actively rearing abundant workers, and no males have ever ap-
peared. Thus the four females designated as Al, 6b, D3, and
D4 appear to have exhibited thelytocous parthenogenesis. A
fuller discussion of these cases is warranted, to indicate how
truly their worker progeny can be considered to have been im-
paternate.

Al. Al was a young female of A. picea, matured with a group
of 20 in a colony which had been maintained in the artificial nest
since August, 1940. It accomplished a normal marriage flight
(except for the absence of males and of mating) in January,
1943, and immediately thereafter was isolated. No males were
recorded in the parent colony from the time it was set up in the
artificial nest until January, 1944, after the parent queen had
died and long after Female Al had emerged. There seems vir-
tually no chance, therefore, that Female Al had been acciden-
tally inseminated. This female had been exposed to about 2000
rontgens of x-rays at 85 kvp. as an adult between the time of its
flight and its isolation in connection with another experiment.
There was no external evidence throughout the life of the queen,
however, that this treatment had produced any effect, unless the
premature death of this female can be so considered.

A perfect worker pupa was eclosed in the brood of Female
Al three months after isolation. Several larvae were present at
the time. Unfortunately a prolonged absence from the country
interrupted observations of this colony for three months after
the appearance of this pupa. When observation was renewed
the queen was found intact, but the young worker and all brood
had disappeared. The female attempted to rear a new brood,
but both queen and brood perished from unknown causes late
in 1943.

6b. Female 6b was a young queen of A. picea which was taken
as a winged mature insect from a wild colony, artificially
dealated, and isolated. Three months after isolation a perfect
worker pupa was produced, which hatched successfully as a
first-brood worker of typically minute stature, and remained in
the nest. Younger larvae were present, which developed slowly
but from which a second pupa matured nine and one-half months
after the dealation of the female. This pupa was predominantly
male in character, but small and defective. It was destroyed
by the queen almost immediately, before further examination
could be made of it. The remaining larvae shortly perished or
were destroyed.

Dec., 1945]

Haskins & Enzmann: Formicid^:

273

D3. Female D3, a queen of Aphcenogaster lamellidens, was
produced from a larva taken in a wild colony. It was matured
and hatched under controlled conditions, emerged and was
dealated and returned to a group of its sister- workers, by which
it was accepted permanently. No males were recorded in the
parent nest during its presence there, nor in the colony-fragment
for the duration of the experiment. The first worker pupa ap-
peared in a little over three months after the introduction of the
young female. Seven additional workers were matured during
the next two months. Thereafter the female perished and obser-
vations were discontinued.

D4. Female D4 was a sister of the preceding, and was treated
in exactly the same manner. Worker pupae were similarly ma-
tured within the colony fragment to which this queen was at-
tached, which was composed of older sister-workers. The num-
ber of these worker pupae mounted rapidly in numbers over the
next six months. In October, 1943, six months after the parent
female had been isolated with this colony fragment, which of
course contained no brood at the time of its introduction, worker
production ceased temporarily, with a large residue of larvae
within the nest. In June, 1944, this brood began to mature,
producing only workers. These continued to be produced until
late September, at which time 44 worker pupae had been counted.
It is probable that this count is low, since the colony was not
anaesthetized to make it, and some were therefore probably missed.

The conclusion seems inescapable that the worker progeny
of Female A1 was truly impaternate. Female 6b, since it was
taken as an adult from a wild colony, might have been fecundated
before capture. It would be unusual in such a case, however,
if the second pupa to mature were a male, or a male-worker
mosaic, wrhich latter this pupa may conceivably have been. A
sister of this ant, taken at the same time, produced only males.
Several additional sisters failed altogether to raise broods.

The precautions surrounding the handling of Females D3 and
D4 were such that it is extremely difficult to. believe that either
was accidentally fecundated. However, the young workers
which came to maturity in these colony fragments could conceiv-
ably have been the progeny of workers rather than of these indi-
viduals, and since these workers had been taken as adults one
or more of them might have been fecundated. Female D3 died
during the rearing of the first broods, so that it was impossible

274

Journal New York Entomological Society

[Vol. LIII

to check this point. Female D4, however, survived, and after
rearing the broods recorded was again isolated, this time with
nine of her own worker pupae and semipupae, but without adults.
Under these conditions it was certain that all brood produced
was the progeny either of this female or of young workers known
to be infertile.

Eggs were produced within four days of isolation and a larva
hatched 27 days later. Eighteen days after hatching, this larva
formed a semipupa, from which a perfect, though small, worker
pupa was eclosed eight days later. The pupal period lasted for
15 days, and a normal worker was hatched. A second semipupa
matured nine days after the first, eclosing a perfect small worker
pupa 6 days later, which matured in 20 days. Five additional
adults were brought to maturity after pupal periods of 12, 11,
11, 19, and 13 days respectively. All were workers of the normal
form, though of smaller than normal stature. They closely re-
sembled the usual “first-brood” workers of A. lamellidens in
pilosity, sculpture, coloring, and stature, and all took up norma]
functions in the colony. There seems no doubt, therefore, either
that Female D4 in fact consistently produced workers, or that
they were produced by the young worker progeny of this same
insect. Inasmuch as the first eggs appeared in the colony before
these pupae had completed hatching, and in view of the previous
record of Female D4, it seems highly probable that these ova
were at least in part derived from this insect. It should be
added that, subsequent to the removal of Female D4 from the
original colony fragment, eggs- continued to appear, but none of
these produced workers.

The experiments recorded here involve but few individuals.
They are of preliminary character and are being continued on
a much larger scale. Nevertheless they seem to confirm the sus-
picion that thelytocous parthenogenesis, so common among the
Phytophaga and Terebrantia, may also occasionally occur among
the Formicidge as well as other higher Aculeates. If its occur-
rence is at all frequent among ants, it may have an interesting
bearing upon the methods of colony formation normal to certain
species, and upon other aspects of colonial economy.

These experiments may have a possibly suggestive relation to
one or two questions of the general biology of ants which is worth

Dec., 1945]

Haskins & Enzmann: Formicid^

275

noting in passing. The first question relates to the habit of
dealation. It seems indicated that in Aphcenogaster picea, at
least, the instinct of self-dealation in the young queen may be
fully expressed not only in the absence of a mating flight but
in the absence of fecundation. Thus of the females listed in
Table I five had spontaneously dealated themselves before isola-
tion. None of these had gone through even the semblance of a
mating flight, and all produced males. This may be suggestive
in consideration of the ready self-dealation of the females of
many lower ants in advance of emergence from the parent nest,
and particularly with the observed ground-mating of females of
Stigmatomma pallipes which were already dealate, though still
callow (Haskins, 1928).

The second point concerns the minute stature regularly ob-
served among first-brood workers of most ants which establish
their colonies in the claustral manner. Goetsch (1937) has
shown that eggs of young fertile queens when transferred to older
colonies still give rise to the nanitic workers characteristic of
normal first broods. It is clear that the potentialities of minute
stature are therefore present in first-brood ova, but Goetsch con-
siders this to be a function of the yolk-supply of the egg rather
than a genetic factor. In Aphcenogaster picea and A. lamelli-
dens the males, though of slight stature, are considerably larger
than the ordinary nanitic workers of the first brood. The males
which were brought to maturity as first-brood progeny by the
infertile females reported here were, with a few exceptions, of
fully normal stature. These females, as already noted, had been
well fed with solid proteinaceous material from the time of first
isolation, and their larvae were similarly fed. This might seem
to account for the discrepancy. On the other hand, the worker
daughter of Female 6b, which was treated in exactly the same
fashion, was of nanitic stature, and the worker pupa of Female
Al was also minute. It would be of interest to determine how
far nanism and the sex of the first progeny are really related in
young females which have been abundantly supplied with food
from the time that they wTere eclosed from pupae throughout the
history of their colony foundation. Experiments are in progress
to check this point.

276

Journal New York Entomological Society

[Yol. LIII

SUMMARY

The general question of the occurrence of parthenogenesis in
the Formicidae, and especially the problem of the production of
impaternate females, is considered. The requirements which
must be met in work designed to test the existence of thelytocous
parthenogenesis among ants are outlined and a program of this
sort, at present in its early stages, is described. Preliminary
data are presented which are suggestive of the production of
bona fide impaternate workers in Aphcenog aster fulva picea and
A. lamellidens. The bearing of these results on certain general
questions of the biology of ants is indicated.

LITERATURE CITED
Comstock, A. B. Quoted in Wheeler (1903).

Crawley, W. C. 1912. Parthenogenesis in worker ants, with special refer-
ence to two colonies of Lasius niger Linn. Trans. Ent. Soc. London :
657-663.

Descy, A. 1924. Recherches sur la sexualite et 1 ’instinct chez les Hymen-
opteres. Bull. Biol. Soc. France Belg., 58: 1-37.

Bzierzon, J. 1845-1876. “ Papers on fertilization and parthenogenesis in

the honey bee. Eichstadt Bienen-Zeitung, 1-32, 1845-1876. ’ 3 (Snod-
grass, 1925: 308.)

Fielde, A. M. 1901. A study of an ant. Proc. Nat. Acad. Sci. Phila., 53 :
425-449.

. 1905. Observations on the progeny of virgin ants. Biol. Bull., 9:

355-360.

Foerster, Arnold. 1856. “ Hymenopterologische Studien, 33 Heft 2 — •

‘ 1 Chalcidiae und Proctotrupii. ’ 3 Aachen, Ernst ter Meer. 152 pp.
Forel, A. 1874. Les fourmis de la Suisse. 2nd ed., rev. et cor., La Chaux-
de-Fonds, Imprimerie cooperative (1920).

Goetsch, W. 1937. Ameisen-Staaten. Veroffentlich v. d. Schlesischen f.

vaterland. Cultur. Breslau F. Hirt: 36-41. (109 Jahresbericht

Schels. Gesellsch. f. vaterl. Cultur, 1936. Naturwissensch.-medizin.
Reihe 1.)

. 1939. Die Staaten argentinischer Blattschneider-Ameisen. Zoo-

logica (Stuttgart) Heft 96, Band 35: 1-105. 36 figs.

, and Br. Kathner. 1937. Die Koloniengriindung der Formicinen

und ihre experimentelle Beeinflussung. Zeitschr. Morph, Okol.
Tiere, 33 : 201-260. 6 figs.

Haskins, C. P. 1928. Notes on the behavior and habits of Stigmatomma
pallipes Haldemann. Jour. N. Y. Ent. Soc., 36: 179-184.

Jack, R. W. 1917. Parthenogenesis among the workers of the Cape Honey
Bee. Trans. Ent. Soc. London: 396-403. 2 pis.

Dec., 1945]

Haskins & Enzmann: Formicid^e

277

Janet, C. 1909. Sur la parthenogenese arrhenotoque de la fourmi ouvriere.
Mem. Soc. Acad, de 1’Oise, 8 pp.

Leiby, B. W. 1922. The polyembryonie development of Copidosoma
gelechiae, with notes on its biology. Jour. Morph., 37 : 195-284.
18 pis.

Light, S. F. 1943. The determination of the castes of social insects (Con-
cluded). Quart. Bev. Biol., 18: 46-63.

Lubbock, Sir John. 1888. Ants, bees, and wasps. London, Kegan Paul
& Co., Ltd.

Mackensen, Otto. 1943. The occurrence of parthenogenetic females in
some strains of honeybees. Jour. Econ. Ent., 36: 465—467.

Onions, G. W. 1912. South African ‘ 1 fertile ” worker bees. Agric. Jour.

Union of S. Africa, 3 : 720-728.

. 1914. Same title. Ibid., 7 : 44-46.

Beichenbach, H. 1902. Ueber Parthenogenese bei Ameisen und andere
Beobachtung an Ameisenkolonien in kiinstlichen Nestern. Biol.
Centralbl., 22 : 461-465.

Snodgrass, B. E. 1925. Anatomy and physiology of the honeybee. New
York, McGraw-Hill.

Speicher, Kathryn G. 1934. Impaternate females in Habrobracon. Biol.
Bull., 67 (2) : 277-293.

, and B. B. Speiqher. 1938. Diploids from unfertilized eggs in

Habrobracon. Ibid., 74 (2) : 247-252. 2 figs.

Stockhert, E. 1923. Ueber Entwicklung und Lebenweise der Bienen-
gattung Halictus Latr. und ihrer Schmarotzer. 1, Teil. Die Bi-
ologie der Gattung Halictus Latr. Konowia, 2: 48-64; 146-165;
216-247.

Tanner, J. E. 1892. CEcodoma cephalotes. Second paper. Trinidad Field
Naturalists’ Club, 1: 123-127.

Tanquary, M. 1913. Biological and embryological studies on Formicidae.

Bull. no. 9, Illinois State Lab. Nat. Hist., 417-477, 8 pis.

Wheeler, W. M. 1903. The origin of female and worker ants from the
eggs of parthenogenetic workers. Sci., n.s., 18: 830-833.

Whiting, P. W. 1928. The relation between gynandromorphism and muta-
tion in Habrobracon. Amer. Nat., 62 (678) : 59-62.

. 1935. Sex determination in bees and Avasps. Jour. Hered., 26:

263-272.

. 1940. Multiple alleles in sex determination of Habrobracon. Jour.

Morph., 66: 323-355.

. 1941. Beport on: Investigations on genetics and sex determination

in the parasitic wasp Habrobracon. Yearbook 1940 of the Ameri-
can Philosophical Society: 274-276.

,,

■

Dec.. 1945]

Alexander : Crane-Flies

279

RECORDS AND DESCRIPTIONS OF NEOTROPICAL
CRANE-FLIES (TIPULID^, DIPTERA), XX

By Charles P. Alexander
Amherst, Massachusetts

The preceding part under this general title was published in
March 1945 (Journal of the New York Entomological So-
ciety, 53: 49-61). The majority of the novelties here treated
are from Costa Rica where they were collected by the late Pablo
Schilcl. Such materials were contained in the extensive Melander
Collection which I have been privileged to study through the
kindly interest of Dr. Melander. I am further indebted for the
great privilege of retaining the types of such species as are based
on a single specimen. A few further species are from Panama
where they^ were collected by Dr. C. Howard Curran and are
preserved in the American Museum of Natural History. I am
very indebted to Drs. Curran and Melander for much friendly
co-operation in this survey of the crane-flies of Tropical America.

Genus Teucholabis Osten Sacken

Teucholabis (Teucholabis) circumscripta new species.

General coloration of thorax handsomely patterned with yellow, black and
chestnut, the praescutum with three separate black stripes, the lateral pair
crossing the suture and entirely covering the scutal lobes; postnotum entirely
black; pleura black, with a yellow ventral longitudinal stripe; halteres with
yellow knobs; femora obscure brownish yellow, the tips blackened; wings
whitish subhyaline, characteristically patterned Avith brown, including a nar-
row border that almost encircles the wing ; abdominal segments black, ringed
with yellow; male hypopygium with the inner dististyle terminating in a
single developed spine.

Male. — Length about 6 mm. ; wing 6.2 mm.

Eostrum unusually long, subequal to or exceeding the remainder of head,
black throughout; palpi black. Antennae black; basal flagellar segments
oval, the outer ones more elongate; terminal segment about two-thirds as
long as the penultimate; verticils longer than the segments. Head above
with a large black area that occupies much of the vertex, leaving the front
and part of anterior vertex, together with the posterior portions of head
paler.

Pronotum partly hidden, evidently yellow, the sides more darkened; pre-
tergites pale yellow. Mesonotum with an unusually handsome pattern ; prae-

280

Journal New York Entomological Society

[Vol. LIII

scutum with three black stripes, the median one broadened at cephalic end,
narrowed behind and not reaching the suture; lateral stripes much wider,
crossing the suture and involving all of the scutal lobes; praescutal inter-
spaces light chestnut, the humeral and lateral portions conspicuously light
yellow; central region of scutum and the scutellum conspicuously pale yellow,
the extreme posterior median area of praescutum concolorous; parascutella
dark brown; postnotum entirely black. Pleura conspicuously patterned;
ground color black, including the dorsopleural area and the ventral sterno-
pleurite; a yellow longitudinal stripe extending from behind the fore coxae
to the base of abdomen, more widened behind ; midsternal region again light
yellow. Halteres with stem blackened, knob light yellow. Legs with all
coxae and trochanters black; femora obscure brownish yellow basally, the
tips conspicuously blackened, on the dilated fore femora including more
than the distal third, on the posterior legs involving about the outer fourth
or fifth; posterior femora with basal half infuscated, isolating a broad sub-
terminal yellow ring ; tibiae obscure yellow, the bases and tips narrowly dark-
ened; tarsi brownish black, posterior basitarsi narrowly dilated at proximal
end and with a sensory pocket; claws simple. Wings with the ground color
whitish subhyaline, conspicuously patterned with brown, including a narrow
border that encircles the wing, from h to the axillary angle, including all of
cells C and Sc and remaining of approximately this general width through-
out; further brown spots at origin of Rs, stigma, cord, outer end of cell
1st If a and at end of vein 2nd A, mostly confluent with the marginal dark-
ening ; prearcular field pale ; veins light brown, darker in the patterned fields.
Venation: Sc long' Sc1 ending about opposite three-fifths Rs; R2 slightly
more than one-half longer than _R2+3+4; cell 1st M2 strongly widened out-
wardly, nearly as long as vein Jf1+2 beyond it; m-cu/ about one-third its
length beyond the fork of M.

Abdominal tergites black, the extreme caudal borders of the segments
paler; sternites with the basal segment yellow; succeeding segments con-
spicuously bicolored, black, the incisures broadly yellow, including about the
distal fourth of each segment, as well as the extreme cephalic border of the
succeeding segment; hypopygium chiefly brownish black. Sternal pockets
restricted in area, that on segment five largest, the lateral setae not forming
rows. Male hypopygium with the apical spine of basistyle straight, broad-
based, narrowed rapidly to the acute terminal spinous point, the inner margin
conspicuously fringed with long yellow setae ; mesal lobe conspicuous but not
blackened, provided with elongate setae. Outer dististyle a long simple rod,
the extreme tip apparently broken; surface with long setae. Inner dististyle
relatively narrow, terminating in a single developed spine, the usual basal
armature lacking ; lobe near base of style terminating in a very few setae of
various lengths. JEdeagus relatively narrow, the lower margin at apex
further produced into a long curved spine; ventral margin with a group of
three unusually long setae at near three-fourths the length, with an isolated
additional seta nearer the base; on dorsal edge with a single further strong
bristle.

Dec., 1945]

Alexander: Crane-Flies

281

Habitat. — Costa Rica.

Holotype, J1, Turrialba, November 1922 (Schild).

Teucholabis (T eucholabis) circumscripta is entirely different
from all described regional species in the very characteristic pat-
tern of both the thorax and wings. No approximately similar
species can be indicated.

Teucholabis (Teucholabis) diplaca new species.

Size medium (wing, male, 5 mm.) ; general coloration polished black and
yellow, the black including a discal area on praescutum ; head polished orange
yellow; femora yellow, tips black; posterior tibiae (male) with a blackened
tubercle on outer sixth; wings whitish subhyaline, virtually unpatterned
except for the narrow triangular stigma; base of cell Sc darkened; Sc1 end-
ing about opposite one-third the length of Bs ; abdominal segments black,
ringed caudally with yellow; male hypopygium with the apical lobe of basi-
style appearing as a broad plate, its outer apical angle further produced
into a long slender spine, the inner angle with three smaller spinous points;
outer dististyle a long simple blackened rod.

Male. — Length about 5.5 mm.; wing 5 mm.

Rostrum orange; palpi black. Antennae with scape and pedicel obscure
orange, flagellum black; flagellar segments oval, the outer ones smaller;
verticils of the more proximal segments subequal in length to the segments.
Head polished orange yellow.

Pronotum chiefly orange yellow; pretergites light yellow. Thorax hand-
somely patterned with polished black and light yellow; praescutum chiefly
black, with three areas that form a complete disk behind, split in front to
isolate narrow yellow lines on the cephalic third of sclerite; scutellum,
median area of scutum and an adjoining small median area on praescutum
yellow, the suture entirely yellow ; scutal lobes extensively black ; parascutella
orange; postnotum black, the mediotergite with an extensive yellow lateral
border on the cephalic half or more. Pleura conspicuously patterned with
black and yellow, the former including the polished sternopleurite and anepi-
sternum, with smaller areas on the propleura and metapleura; dorsopleural
membrane and the entire pteropleurite yellow; no pruinose area on pleura,
as in many species in the genus. Halteres brownish black, knobs orange.
Legs with all coxae and trochanters orange yellow; femora yellow, the tips
broadly and conspicuously black; tibiae obscure yellow, the tips darkened;
tarsi brownish black; posterior tibiae near outer sixth with a small blackened
tubercle or knob; proximal fifth of hind basitarsus swollen. Wings whitish
subhyaline, virtually unpatterned except for the unusually narrow triangular
dark brown stigma; base of cell Sc narrowly darkened; veins comprising the
cord and outer end of cell 1st M2 somewhat more intensely darkened but the
membrane not involved; veins dark brown, C, Sc and B more yellowed. Vena-
tion; Sc-l ending about opposite one-third the length of Bs, Sc2 some distance

282

Journal New York Entomological Society

[Vol. LIII

from its tip, Sc± alone nearly as long as R2-, anterior branch of Es gently
sinuous, slightly upcurved at tip so cell E2 at margin is only a little more
extensive than cell Es; outer section of M1+2 arcuated; m-cu at or just before
the fork of M.

Abdomen black, ringed with yellow, the latter color involving the posterior
margins of the segments; hypopygium chiefly darkened. Sternal pocket of
segment five extensive, more or less triangular in outline, the setae converging
toward the midline, the more central ones smallest ; setae of sternite six more
widely separated, including only about six or seven on either side. Male
hypopygium with the apical lobe of basistyle unique, appearing as a broad
flattened plate, the outer apical angle of which is produced into a long
slender spine, the opposite apical angle produced into about three smaller
spinous points; inner margin and apex of plate provided with long yellow
setae; near base of plate with a small lobe bearing about four long marginal
setae; mesal flange of basistyle with smooth margin. Outer dististyle a long
slender simple blackened rod, the tip acute, the surface with conspicuous
appressed spinulae on outer face and with strong tuberculate points. Inner
dististyle with the outer and basal teeth powerful, separated from one an-
other by an oval notch, the margins thickened ; basal lobe of style with about
five very long setae. JEdeagus stout, the apex expanded into obtusely rounded
blades; setae relatively numerous and unusually long.

Habitat. — Panama.

Holotype, Barro Colorado Island, Canal Zone, February
13, 1929 (Curran) ; American Museum of Natural History.

Teucholabis (T eucholabis) diplaca is quite distinct from all
others of the very numerous regional species of the subgenus.
It differs especially in the structure of the male hypopygium,
more particularly the outer lobe of the basistyle. This latter
structure is approximated by species such as T. ( T .) bigladia
Alexander, T. (T.) biramosa Alexander, and T. ( T .) scabrosa
Alexander, all of which are entirely different in other respects
and evidently not closely allied.

Teucholabis (Teucholabis) platyphallus new species.

Allied to furva ; mesonotum black, pronotum yellow; thoracic pleura with
a conspicuous ventral pale stripe; knobs of halteres light yellow; posterior
basitarsi (male) dilated and with an elongated setuliferous pocket; wings
with a brownish tinge, the cephalic border more whitened; stigma oval,
darker brown; Scx ending about opposite midlength of Es ; branches of Es
extending generally parallel to one another ; male hypopygium with the outer
lobe of basistyle a broad flattened blade, its margin fimbriate; outer disti-
style a long simple sinuous rod, microscopically spinulose and with scattered
elongate setae ; inner dististyle trilobed, the basal lobe cylindrical and tipped

Dec., 1945]

Alexander: Crane-Flies

283

with four long setae; aedeagus unusually expanded, subtriangular in outline,
with paired setae on the dorsal and ventral margins.

Male. — Length about 5.5 mm. ; wing 5.5 mm.

Eostrum obscure brownish yellow; palpi black. Antennae with basal seg-
ments pale brown, the outer ones darker ; under face of scape more yellowed ;
basal flagellar segments short-oval, the outer segments more elongate; verti-
cils conspicuous. Head dark liver brown.

Pronotum and pretergites, with the restricted humeral region of prae-
scutum, yellow. Eemainder of mesonotum, including scutellum and pleuro-
tergite, black. Pleura chiefly black, the propleura and a conspicuous ventral
stripe on the mesopleura pale yellow, whitish pruinose, this color including
the whole ventral portion of thorax excepting the restricted ventral sterno-
pleurite. Halteres with stem infuscated, knob light yellow. Legs with the
coxae yellow, the fore pair a trifle darker; trochanters yellow; remainder of
legs long and relatively slender, obscure yellow; tips of femora narrowly and
weakly darkened, on the fore pair a little more extensive and gradually
darkened; tibiae obscure yellow, the tips narrowly dark brown; basal seg-
ments of tarsi obscure yellow, the terminal segments more darkened; claws
simple; posterior basitarsi (male) with proximal end dilated and provided
with an elongated setuliferous pocket. Wings with a brownish tinge, the
prearcular and costal fields, together with areas before and beyond the
stigma, more whitened ; stigma oval, darker brown ;v a weak brown cloud over
anterior cord; veins brown, those in the prearcular field more yellowed.
Venation: Sc ending about opposite midlength of the long Es, the latter very
weakly bent at near midlength; branches of Es extending generally parallel
to one another for their entire lengths; cell 1st M2 subequal to or a trifle
longer than vein M4; m-cu shortly beyond the fork of M.

Abdominal tergites brownish black; sternites paler medially, more dark-
ened on sides; outer segments, including hypopygium, more uniformly black-
ened. Sternal pockets well developed on both segments five and six. Male
hypopygium with the outer lobe of basistyle subapical in position, appearing
as a broad flattened blade, its outer third narrowed into a long terminal
spine; inner margin of blade with a fringe of long yellow setae; darkened
flange of mesal face of style relatively untoothed, the outer portion smooth,
the central part emarginate, the cephalic third with two or three coarse denti-
cles. Outer dististyle a long simple sinuous rod, narrowed very gradually
to an acute spine, the outer surface with microscopic appressed spinulae and
a few long setae. Inner dististyle blackened, trilobed, the outer lobe stout
and obtuse, the inner one a sharp spine; basal lobe cylindrical, tipped with
about four unusually long setae, these subequal to or exceeding in length the
lobe itself. .ZEdeagus unusually expanded, subtriangular in outline, the lower
apical angle a curved black spine; surface smooth except for two powerful
setae on lower edge at near three-fourths the length and two others on dorsal
edge, a trifle more basad in position.

Habitat. — Costa Rica.

Holotype, cf, Turrialba, November 1922 (Schild).

284

. Journal New York Entomological Society

[Yol. liii

The nearest relative of the present fly appears to be TeucJio-
labis (T eucholabis) furva Alexander, which differs in the pattern
of the body and wings, and in the details of structure of the
male hypopygium.

Teucholabis (Teucholabis) serrulifera new species.

Allied to furva; mesonotal praescutum and scutal lobes polished black; a
broad black stripe over the dorsal thoracic pleurites; rostrum and palpi
black; knobs of halteres a little paler than the stem; wings with a weak
blackish tinge, a little more intense on outer fourth; stigma dark brown;
Scx ending opposite midlength of Bs; branches of Bs parallel to one another
for most of their lengths; male hypopygium with the spine of basistyle
slender, glabrous; outer dististyle at base widely expanded into a lobe or
blade bearing spines and conspicuous setae, aedeagus with the apical spine
bent laterad, before the tip with a conspicuous lobe that bears a few setae.

Male. — Length about 5.5 mm.; wing 5.5 mm.

Eostrum and palpi black, the former relatively long, about two-thirds the
remainder of head. Antennae black throughout; flagellar segments oval;
terminal segment shorter than the penultimate; verticils exceeding the seg-
ments in length. Head brownish black.

Pronotal scutum obscure yellow, more infuscated laterally; scutellum and
pretergites light yellow. Mesonotal praescutum and scutal lobes uniformly
polished black, only the reduced humeral region of the former obscure yel-
low; median region of scutum and posterior portions of the lobes narrowly
obscure yellow; scutellum dark brown, parascutella yellow; postnotum black,
the suture between the mediotergite and pleurotergite narrowly reddened.
Pleura conspicuously bicolored, the entire venter light yellow, including the
sternopleurite and meral region; dorsal pleurites occupied by a broad black
longitudinal stripe extending from the sides of the pronotum across the
anepisternum and dorsal pteropleurite onto the postnotum; dorsopleural
region yellow; no well-developed silvery area on pleura. Halteres with stem
blackened, apex of knob a little paler, obscure yellowish brown. Legs with
fore coxae blackened, remaining coxae and all trochanters yellow; femora
yellow basally, the tips blackened, most broadly so on fore legs where about
the distal third is included, narrowest on posterior femora where only the
outer sixth or seventh is blackened; tibiae brownish black; tarsi black; pos-
terior basitarsi strongly dilated, at proximal end with a setuliferous pocket.
Wings with a weak blackish tinge, slightly more intense on outer fourth;
stigma oval, dark brown; a narrow, vague, darkened seam over cord; veins
dark brown, brownish yellow in the prearcular field. Venation: Sc relatively
long, Sc1 ending about opposite midlength of the long Bs; B1 + 2 and B2+s+4
subequal, both a little more than one-half B2; branches of Bs extending
generally parallel to one another for almost their whole lengths, B5 near its
apex diverging, ending at tip of wing; cell 1st M2 narrow, subequal to distal
section of M3- m-cu a short distance beyond fork of M.

Abdominal tergites and hypopygium black; basal sternites a trifle more

Dec., 1945]

Alexander : Crane-Flies

285

piceous. Pocket on fifth sternite oval, with about eight setae on either side,
these directed toward the midline, the area comprised of abundant small
bulbous structures ending in small points; on sixth sternite, setae about ten
on either side, widely separated at midline. Basistyle with apical spine very
slender, subapical in position, without setae ; mesal lobe irregularly erose, the
denticles interspersed with long setae. Outer dististyle a broad flattened
blade that narrows gradually to a slender apical spine, the outer edge micro-
scopically serrulate; near base, style broadly expanded, bearing a long
slender spine, provided with abundant long coarse setae and a few smaller
spinous points. Inner dististyle conspicuously bispinous, the outer spine
larger and more curved; at base of style a broad lobe that bears a transverse
row of about five strong spinous setae. iEdeagus relatively narrow, the api-
cal third bent strongly laterad into a long blackened spine, before the apex
of which, on lower face, bearing a strong cylindrical lobe with two or three
apical setae and one unusually long additional bristle nearer base; on oppo-
site side of aedeagus, at point of bending, with a nearly rectangular lobe,
below which are two strong setae.

Habitat. — Costa Rica.

Holotype, J1, Turrialba, November 1922 (Schild).

This species and various others, center about Teucholabis
(Teucholabis) furva Alexander which they resemble in general
appearance and venation, but differ in all details of the male
hypopyginm, especially the spine of the bastistyle, both dististyles
and the aedeagus.

Teucholabis (Teucholabis) turrialbensis new species.

Male. — Length about 6 mm.; wing 5 mm.

Closely related and generally similar to serrulifera new species, differing
especially in the structure of the male liypopygium. Wings with Sc some-
what shorter, Sc± ending before midlength of Bs.

Sternal pocket of sixth segment of male with the setas fewer in number,
totalling six or seven. Male hypopygium with the outer dististyle much more
slender and subcylindrical, not flattened, the armature much restricted ; serru-
lations of outer surface smaller and inconspicuous ; spinous flange of mesal
face much smaller, closely applied to the style, the outermost spine long and
slender.

Habitat. — Costa Rica.

Holotype, J1, Turrialba, November 1922 (Schild) ; Melander
Collection. Paratopotype, J'; Alexander Collection.

Genus Gnophomyia Osten Sacken

Gnophomyia (Gnophomyia) curraniana new species.

Head and abdomen black; thorax, excepting the postnotal scutum and the
metapleura, orange ; halteres and legs black ; wings broad, dark brown, with

286

Journal New York Entomological Society

[VOL. LIII

a wide whitish band at and beyond midlength; cells C and Sc uniformly
darkened; cell B2 at margin wider than cell B3', cord transverse; cell 1st M2
rectangular.

Female. — Length about 7 mm.; wing 7 mm.

Kostrum and palpi black. Antennae black throughout; flagellar segments
oval, gradually decreasing in size outwardly. Head uniformly black.

Pronotum with scutum black, scutellum orange, the sunken central portion
a little paler. Mesonotum and pleura uniformly orange, only the metapleura
beneath the root of the halteres blackened. Halteres black, the extreme base
of stem orange. Legs, including coxae and trochanters, black. Wings broad,
conspicuously dimidiate; dark brown, with a broad whitish band at and
beyond midlength; cells C and Sc uniformly darkened; basal two-fifths of
wing slightly paler brown than the apical fourth, the latter a trifle more
extensive than the white band; the latter includes most of cell Bx, thence
continued to the posterior border as a nearly parallel-sided area, the outer
edge lying just beyond cord so the bases of cells B± to. M4 are whitened;
conspicuous white streaks along veins M and 1st A, with further similar lines
in basal portion of cell B and near outer end of cell Cu ; veins brown, not
paler in the brightened field, excepting certain elements at cord. Macro-
trichia abundant in all cells beyond cord and in outer ends of cells B1} Cu
and 1st A. Venation: Scx ending just before fork of Bs ; B2 suboblique;
Bs+i present as a short to very short element; veins B2 and B4 more nearly
parallel than in leucoplaca, cell B2 wider than cell B3- in leucoplaca, the
veins divergent so the cells are subequal in area at the margin; basal section
of B5 short and straight, the entire cord transverse; in leucoplaca, anterior
cord oblique, the inner end of cell B4 lying more proximad than the other
cells ; cell 1st M2 more rectangular.

Abdomen black; ovipositor with the cerci unusually slender, dark chestnut
brown, gradually narrowed to the subacute tips, the surface glabrous.

Habitat. — Panama.

Holotype, §, Barro Colorado Island, Canal Zone, February
13, 1929 (Curran) ; American Museum of Natural History.

The most similar described species is Gnophomyia ( Gnopho -
myia) leucoplaca Alexander, from the upper Amazons (Teffe,
Amazonas, Brazil), which differs in the much greater extent of
black coloration, particularly of the praascutum and mesopleura,
and in distinct venational details, as compared above. In leuco-
placa, the cells of the outer radial field are uniformly darkened,
without broad white bases in R± and R5, as in the present fly.
The species is named in honor of the collector, the distinguished
Dipterologist, Dr. C. Howard Curran.

Gnophomyia (Gnophomyia) pulvinaris new species.

Allied to mcestitia ; general coloration of body and appendages black, the
lateral portions of the pronotal scutellum yellow; halteres uniformly black-

Dec., 1945]

Alexander: Crane-Flies

287

ened; wings tinged with brown, with a still darker brown pattern, including
a broad band at cord; male hypopygium with the caudal margin of tergite
gently concave, with a double row of at least one hundred spinous setae ; basi-
style on proximal portion of mesal face with an oval area or cushion includ-
ing about fifty blackened setae; outer dististyle with inner margin with
irregular blackened teeth.

Male. — Length about 4.5 mm.; wing 5.1 mm.

Female. — Length about 4.5 mm.; wing 4.8 mm.

Eostrum and palpi black. Antennae black throughout; flagellar segments
elongate, subcylindrical, with long conspicuous verticils, the longest much
exceeding the segments, unilaterally arranged on the outer face ; in addition
to the coarse verticils, the segments clothed with abundant finer setae. Head
dull black, sparsely pruinose in front; anterior vertex broad, approximately
four times the diameter of scape.

Pronotum dark brown, the lateral portions of the scutellum clear light
yellow. Mesonotum almost uniformly brownish black, the surface more or
less polished. Pleura black, more pruinose on the ventral and posterior por-
tions, including the pleurotergite ; dorsal pleurites, as well as the dorsopleural
membrane, deep velvety black. Halteres black throughout. Legs with the
coxae dark brown, more or less pruinose, especially the middle pair; remain-
der of legs black. Wings with a brownish tinge, rather distinctly patterned
with still darker brown, this appearing especially as a broad dark band at
the cord and darkening in cells M and bases of Cu and 1st A ; stigma long
and narrow, dark brown; veins brownish black. Venation: Es with its basal
section oblique, straight; r-m variable in position, in the type a short dis-
tance before the fork of Es, in the allotype just beyond the fork; E2+3+4 in
direct alignment with E2+s and E3, forming an even arc; E2 + 3+4 and E2+3
subequal; all outer branches of Es extending generally parallel to one an-
other ; cell 1st M2 long-rectangular, with m-cu at near midlength, the cell
about as long as vein M4.

Abdomen, including hypopygium, black. Ovipositor with cerci relatively
long and slender, with setae to the acute tips. Male hypopygium with the
tergite large and conspicuous, transverse, the caudal margin broadly and
gently concave, provided with an unbroken double row totalling at least 100
spinous setae, those at the ends of row not modified into a brush or pencil.
Basistyle short and stout, on mesal face at cephalic end with a dense oval
cushion of long black setae totalling about 50 in number. Outer dististyle
gradually narrowed outwardly, the inner margin with a row of small irregu-
lar blackened teeth, including a larger flange at base of the series. Inner
dististyle about three-fourths as long as last, uniformly dark colored, gradu-
ally narrowed outwardly, the tip a small blackened point; style provided
with long erect setae. Gonapophyses appearing as two separate blackened
plates, one on either side of the slender aedeagus, the tips of the blades
incurved and contiguous.

Habitat. — Costa Rica.

288

Journal New York Entomological Society

[Vol. LIII

Holotype, .J', Turrialba, November 1922 (Schild). Allotopo-
type, J.

Gnophomyia ( Gnophomyia ) pnlvinaris is entirely distinct
from the now numerous species of the genus that center about
G. ( G .) mcestitia Alexander, having in the male an enlarged
ninth tergite that is provided with an armature of strong spinous
setae. The most similar species is G. (G.) nimbifera Alexander,
of Peru, which has all details of the male hypopygium distinct.
The cushion of setae on the basistyle of the present fly provides
a distinctive character.

Genus Neognophomyia Alexander

Neognophomyia schildi new species.

General coloration of mesonotum reddish brown; pleura yellow, with an
almost continuous darkened dorsal stripe; femora yellow, the tips narrowly
infuscated; wings whitish subhyaline, restrictedly patterned with brown,
including a narrow band over the anterior cord; male hypopygium with the
tergal spines strong and powerful, blackened; outer dististyle slender, with
five setae, of which two occupy the apex ; phallosome consisting of two broadly
flattened plates that subtend the shorter aedeagus, the apex of each blade
terminating in a small conical darkened point that is directed laterad.

Male. — Length about 5 mm. ; wing 5 mm.

Rostrum yellow; basal segment of palpus yellow, outer segments brown.
Antennae with scape brownish yellow, pedicel light brown, flagellum brown;
flagellar segments oval, with long conspicuous verticils. Head yellow, darker
behind.

Pronotum very pale brown, the lateral borders of the scutellum, with the
pretergites, pale yellow. Mesonotal praescutum with the disk reddish brown,
the lateral and humeral borders yellow ; scutal lobes a trifle more darkened ;
scutellum obscure yellow. Pleura clear light yellow ventrally, the dorsal
sclerites chiefly infuscated, including the propleura, anepisternum and pleuro-
tergite, somewhat paler brown on the dorsal pteropleurite, the whole forming
a broad dorsal pleural stripe. Halteres with stem light yellow, knob infus-
cated. Legs with the coxae and trochanters light yellow; femora yellow, the
tips narrowly and weakly infuscated ; tibiae and basitarsi yellow ; outer tarsal
segments passing into brownish black. Wings whitish subhyaline, restrict-
edly patterned with brown, including a narrow band over anterior cord and
very narrow seams over the posterior cord and outer end of cell 1st M2; a
very restricted darkening near wing base, especially in cell M ; veins brown-
ish yellow, darker in the patterned areas. Venation: ending about oppo-

site -R2; -K2+3+4 subequal to Es+i- venation of outer radial field normal for
the genus, cell E2 at margin being very reduced, cell E3 greatly widened;
cell 1st M2 short and strongly widened outwardly, m-cu at near midlength.

Dec., 1945]

Alexander: Crane-Flies

289

Abdominal tergites chiefly infuscated, the subterminal ones more yellowed ;
sternites yellow; hypopygium chestnut brown. Male hypopygium with the
tergal spines strong and powerful, blackened, nearly straight, narrowed to
acute points. Dististyle unusually small and simple; outer dististyle slender,
narrowed to the obtuse tip which bears two long setae, with three further
similar setae along outer margin ; near base of style, on lower margin, with
a low lobe or flange; inner dististyle large, generally triangular in outline,
narrowed to the subobtuse apex, outer margin with a row of about six or
seven strong setae ; basal flange low, provided with a similar number of much
longer setae. Phallosome consisting of two broadly flattened plates subtend-
ing the slightly shorter aedeagus, the tips of the blades terminating in small
conical darkened points that are directed laterad (in slide mounts) .

Habitat. — Costa Rica.

Holotype, lCf, Turrialba, November 1922 (Scbild) • Melander
Collection. Paratopotype, J'; Alexander Collection.

Among the described species of the genns that have the male
hypopygium with heavily blackened and powerful tergal spines,
including Neognophomyia colombicola Alexander, N. consociata
Alexander, N. pervicax Alexander and N. scapha Alexander, the
present fly differs conspicuously in all details of structure of the
male hypopygium, particularly of the dististyles and phallosome.
It is perhaps closest to consociata yet amply distinct. I take
pleasure in dedicating this fly to the memory of the collector,
Mr. Pablo Schild.

Genus Gonomyia Meigen

Gonomyia (Lipophleps) lustralis new species.

Belongs to the manca group; size medium (wing, male, 3.8 mm.) ; general
coloration of praescutum cinnamon brown, unpatterned; antennae (male)
long, the flagellar segments with abundant long erect setae, additional to the
long, unilaterally distributed verticils ; thoracic pleura with an ill-defined
whitish longitudinal stripe; knobs of halteres infuscated; legs medium
brown; male hypopygium with the basistyle produced far beyond the origin
of dististyle as a long pale clavate lobe; dististyle conspicuously forked, the
style bearing a slender blackened arm on the outer margin beyond midlength ;
phallosome without blackened elements.

Male. — Length about 3.5 mm.; wing 3.8 mm.

Rostrum yellow; palpi black. Antennae dark brown, relatively elongate;
flagellar segments long-cylindrical, with abundant long erect setae, additional
to the still longer, unilaterally distributed verticils, the latter occurring on
the more proximal segments only. Head above chiefly pale, the central pos-
terior vertex darkened.

290

Journal New York Entomological Society

[Vol. LIII

Pronotum and pretergites very pale yellow. Mesonotal praescutum uni-
formly light cinnamon brown, unpatterned; scutal lobes a trifle darker, the
broad median area yellow ; scutellum yellow, slightly darkened at base ; medio-
tergite brownish gray. Pleura and pleurotergite chiefly yellow, the former
with an ill-defined, more whitened, longitudinal stripe extending to the base
of the abdomen. Halteres with stem pale, knob infuscated. Legs with the
coxae and trochanters testaceous yellow; remainder of legs medium brown.
Wings with a weak brownish tinge, the prearcular and costal fields light yel-
low; stigma scarcely indicated as a very weak darkening; veins pale brown,
including those in the brightened fields. Venation: Sc± ending opposite
origin of Es, the latter about four-fifths to five-sixths the length of its
anterior branch; m-cu close to the fork of M.

Abdominal tergites dark brown, sternites paler ; hypopygium chiefly weakly
infuscated. Male hypopygium with the outer apical angle of basistyle
greatly produced beyond the origin of the dististyle, the lobe being fujly as
long as the basal portion of style, slightly dilated outwardly, the outer end
with unusually long setae. Dististyle conspicuously forked, the main body
pale, bearing the usual two fasciculate setae at and below the apex; on outer
margin beyond midlength, the style bears a slender blackened lobe or arm,
approximately equal in length to but much narrower than the true apex of
the style. Phallosome with all elements pale, undarkened, with two long
flattened blades that are subacute at tips, together with an obtuse median
lobe that juts slightly distad of the paired blades.

Habitat. — Costa Rica.

Holotype, J1, Turrialba, November 1922 (Schild).

Gonomyia (Lipophleps) lustralis is entirely distinct from the
very numerous members of the manca group occurring in Tropi-
cal America. It shows affinity with species such as G. (L.)
batesi Alexander, but differs from all other forms in the struc-
ture of the male hypopygium.

Genus Erioptera Meigen

Erioptera (Mesocyphona) turrialbse new species.

Size small (wing, male, less than 3 mm.) ; general coloration dark brown;
thoracic pleura striped longitudinally with pale; femora very pale brown,
the tips very narrowly and indistinctly whitened; wings uniformly tinged
with pale brown; male hypopygium Avith a single three-branched dististyle,
the axial branch stoutest and longest, the acute apex glabrous; lower branch
pale, slightly more basal in position than the blackened upper arm; gona-
pophyses single, appearing as a very strongly sinuous blackened rod.

Male. — -Length about 2.5 mm.; wing 2.8 mm.

Rostrum and palpi black. Antennae black throughout ; verticils very long.
Head dark brown, the front and orbits narrowly gray.

Dec., 1945]

Alexander : Crane-Flies

291

Pronotum dark brown. Mesonotum almost uniformly dark brown, the
scutellum a trifle more testaceous. Pleura dark brown, with a ventral yellow-
ish longitudinal stripe, the dorsopleural region pale. Halteres with stem
yellow, knob weakly darkened. Legs with the coxae and trochanters obscure
yellow; femora very pale brown, the tips narrowly and indistinctly whitened;
remainder of legs more brownish yellow. Wings with a uniform pale brown-
ish tinge; veins pale brown, trichia dark brown. Venation: Cell M2 open by
atrophy of basal section of ; vein 2nd A with outer third deflected slightly
cephalad.

Abdomen, including hypopygium, dark brown. Male hypopygium with a
single dististyle that is conspicuously three-branched; main axis longest and
stoutest, its apex acutely pointed, glabrous; outer or dorsal branch slender,
blackened, about two-thirds as long as the axial point; lower or ventral
branch arising a trifle nearer base of style than does the outer spine, entirely
pale, the tip obtuse and microscopically setuliferous. Gonapophysis of either
side single, appearing as a very strongly sinuous blackened rod, gradually
narrowed to the acute spinous point, the latter decussate at the midline.

Habitat. — Costa Eica.

Holotype, J', Turrialba, November 1922 (Schild).

This species is allied to various other regional species, includ-
ing Erioptera ( Mesocyphona ) factiosa Alexander, E. ( M .) in-
variegata Alexander, E. (M.) modica Alexander, E. ( M .)
quadrifurcata Alexander, and E. (M.) withy combei Alexander,
differing in the structure of the male hypopygium.

292

Journal New York Entomological Society

[Yol. liii

BOOK NOTICE

The Lost Woods , Adventures of a Naturalist, Illustrated with
200 Photographs by the Author. By Edwin Way Teale.
Dodd, Mead & Company, New York, 1945. 10 x 7 inches,

xv + 326 p.

Mr. Teale ’s fan mail, I am sure, will rise to another peak, after
The Lost Woods gets in circulation. This is the sixth of Mr.
Teale ’s books on natural history to be brought out by Dodd,
Mead & Company, all attractively designed and printed. The
Lost Woods, named after a childish recollection of a mysterious
woods that the author was unable to find after the passage of
many years and which became to him a symbol of outdoor life,
is a collection of fascinating accounts of animals of the land, sea
and air, narrated and illustrated with the skill for which Mr.
Teale is noted.

King crabs, serpents, cloud formations, birds, beavers, insects,
snowflakes, animal tracks, naturalists, and all living things, in-
stead of being ordinary everyday objects of no unusual interest,
suddenly become animated actors in nature’s drama as soon as
Mr. Teale sets foot out-of-doors with his camera. Their behavior,
their amazing and interesting habits, their battles and their
movements are faithfully and accurately recorded by Mr. Teale
for all to enjoy.

Mr. Teale ’s adventures occur in all sorts of places, in Times
Square, in a submarine and in a plane, but most of them happen
on the ground wherever he chances to be. These endless activi-
ties in the world of nature that Mr. Teale exposes to us are avail-
able to all who have the curiosity and interest necessary to be
aware of them. Failing in this, one should read The Lost Woods
in order to be made aware of life other than our own. Mr. Teale
brings the woods, the fields, the sky, the streams, and their in-
habitants right into your living room and his accounts are not
only fascinating, but scientifically accurate. If you can’t afford
an actual vacation in the woods, take one anyhow by reading Mr.
Teale ’s book, and enjoy the company of an interesting, many-
sided naturalist and master photographer, who writes refresh-
ingly about all animals, even jelly fish and slime molds. —
H. B. W.

Dec., 1945]

Michener: Culicidje

293

SEASONAL VARIATIONS IN CERTAIN SPECIES OF
MOSQUITOES (DIPTERA, CULICID^)

By Charles D. Michener
First Lieutenant, Sanitary Corps, A. U. S.

It is the purpose of this paper to point out seasonal variations
in structure, coloration, and size of adults and larvae of certain
mosquitoes of the genus Culex. In some cases such variation may
have led to unnecessary multiplication of specific names. It is of
importance, also, since the apparently less studied winter forms
of some of the species do not run to the correct point in certain
keys.

Seasonal variations are well known in many groups of insects.
In mosquitoes the winter individuals are often larger than the
summer ones and slight color variations have been noted. Thus
the overwintering females of Anopheles maculipennis freehorni
Aitken in California are larger and darker than the summer
forms.1 The same appears to be true of the related Anopheles
quadrimaculatus Say.

The specimens studied in the preparation of this paper have
all come from the southeastern United States. Many of them were
collected at Camp Shelby, near Hattiesburg, Mississippi, by Mr.
Wm. V. Reed, Capt. Basil G. Markos, the author, and others. For
the opportunity to study numerous other specimens from other
parts of the southeast, the author is indebted to Major Stanley J.
Carpenter, Major W. W. Middlekauff, Lieutenant Louis M. Roth,
and other personnel in the entomology department of the Fourth
Service Command Laboratory.

CULEX (NEOCULEX) APICALIS ADAMS

Among the species of Culex occurring in the southeastern states,
the most conspicuous seasonal variation is found in Culex apicalis.
This, holarctic species breeds throughout the year in this area.

1 Freeborn, S. B. 1932. The seasonal life history of Anopheles maculi-
pennis with reference to humidity requirements and ‘ ‘ hibernation. ’ ’ Amer.
Jour. Hyg., 16: 215.

2 94

Journal New York Entomological Society

[Vol. LIII

Larvae, as well as adults, collected in the cooler months of the year
differ markedly from those found during the summer.

The differences which have been noted in adults between a mid-
winter series and midsummer series are indicated in the following
tabulation :

Summer form Winter form

Smaller, wing scarcely over 4 mm. in Usually larger, wing 4 to 5 mm. in
length. length.

Abdominal bands sometimes repre-
sented only by lateral spots, more
often complete but only one or two
rows of scales in width.

Abdominal bands normally broad,
three or sometimes four scales in
width, rarely with a partial fifth
row.

Integument of mesoscutum light
brown, rarely infuscated.

Integument of mesoscutum usually
infuscated or blackish.

Under surfaces of tibiae with scales
mostly pale.

Upper surfaces of femora with dark
scales often not reaching bases.

Uuder surfaces of tibiae with scales
usually almost all black.

Upper surfaces of femora with dark
scaled area frequently reaching
bases.

All the characters listed in the above tabulation vary between
the extremes, and occur in different combinations with one an-
other. It is therefore believed that only one species is involved,
although a large, broad-banded, dark-legged winter specimen with
dark mesoscutal integument looks like a different species from a
summer specimen. No genitalic differences between such indi-
viduals could be found.

Small specimens almost invariably lack or have only narrow
abdominal bands, although they may have an infuscated meso-
scutum. Many large specimens, particularly during spring and
fall, also have narrow bands, and a. few large ones have brown
mesoscuta. The percentage of large individuals and also of those
having infuscated mesoscuta decreases more slowly in spring and
increases more rapidly in fall than does the percentage of indi-
viduals with broad bands. Apparently these characteristics and
band width are affected by different environmental factors or by
the same factor or combination of factors acting at different
thresholds.

Because, as already stated, there are all intergrades between the
extremes for each character, an arbitrary line had to be selected

Dec., 1945]

Michener: Culicid^

295

iii gathering data for the following table, between, for example,
“infuscated” and “not inf uscated. ’ ’ The mesoscntal integu-
ment of the midsummer individuals recorded as infuscated is, as
a rule, paler than that of winter individuals. Table 1, based on
206 specimens collected at localities more than thirty miles from
the Gulf coast in Mississippi, Alabama, and Georgia, shows the
seasonal distribution of certain of the characters discussed above.
The localities from which these specimens were obtained are
Atlanta, Augusta, Hinesville, Macon, Savannah, and Yaldosta,
Georgia; Anniston and Ozark, Alabama; and Centerville, Gre-
nada, and Hattiesburg, Mississippi.

TABLE 1

Per cent with bands three

or more scales wide ......... 88 100 67 33 20 0 0 0 0 0 50 91

Per cent with infuscated

mesoscutum 100 100 67 88 80 43 0 17 10 0 83 100

No. of specimens studied 18 23 30 18 20 7 11 12 10 10 36 11

Similar results were obtained from a study of about 30 addi-
tional specimens from North Carolina (Durham, Fayetteville,
Hoffman, Laurinburg), South Carolina (Greenwood, Spartan-
burg), and Tennessee (Smyrna), except that one specimen from
near Fayetteville, North Carolina, collected in January and one
from Greenwood, South Carolina, collected in November, are
small with narrow bands and not or scarcely infuscated mesoscuta.

The seasonal variation here described does not occur through-
out the range of C. apicalis. In the northern states, and in the
west as far south as Arizona (the type locality of apicalis ), speci-
mens are of the broad-banded type even in summer (according to
a letter from Dr. Alan Stone dated January 28, 1944). In the
southeastern region such broad-banded specimens disappear dur-
ing the summer, being replaced by a narrow-banded form. The
narrow-banded type occurs, as shown by specimens in the United
States National Museum, at least as far north as Maryland. There
is evidence that in Florida, and possibly in a narrow zone along
the Gulf coast, the broad-banded form does not occur even in

296

Journal New York Entomological Society

[Vol. LIII

winter. Eighteen specimens collected at Panama City and Mari-
anna, Florida, in November, December, and February are small
with narrow bands as in summer individuals farther north.
Approximately 40 other specimens from these same localities and
from Jacksonville, Florida, collected from April to June are in-
distinguishable from the winter specimens collected in the same
area. This Floridian population which appears to be narrow-
banded throughout the year does not represent the extreme in
band reduction in this group, however, for in Mexico C. derivator
Dyar and Knab, which is apparently only a subspecies of apicalis,
lacks pale scaling on the abdominal terga altogether or, at most,
has small white areas latero-apically.

Unfortunately, the species appears to be scarce in peninsular
Florida. Through the kindness of Dr. Alan Stone, records have
been received of specimens from Orla Vista, Orlando, Rock
Springs, and Sanford, Florida. Of these, two specimens from
Rock Springs, collected February 25, have “ rather wide” bands,
but the remainder are narrow-banded.

It would be reasonable to recognize the Floridian form as a
distinct subspecies of apicalis. For the present, no name is
applied, since the seasonal variation evident in other southeastern
states suggests that the differences between Floridian and other
populations may be fhe direct effect of the environment rather
than indications of genetic differentiation. It is remotely possi-
ble that two species are involved but if so suitable differentiating
characters have yet to be found. This question will probably
remain unsettled until rearing experiments can be carried out.

The larvse of C. apicalis also exhibit a wide range of variation
in many characters. Certain of these variations appear to be
correlated with the seasons, although not so well so as the vari-
ations of the adults. Those which are best correlated with the
season are indicated in the following tabulation :

S umm&r form W inter form

Smaller, pigmentation light.

Air tube long, slender, about 2|
times as long as antenna, more
expanded at tip than in winter
form, both sides curving outward
apically.

Larger, pigmentation, especially of
head, dark.

Air tube shorter and more robust,
less than 2| times as long as an-
tenna, less expanded at tip, one
side nearly straight.

Dec., 1945]

Michener: Culicid^e

297

All intergradations between these forms occur, sometimes even
in one pool. The size and pigmentation is correlated in part at
least with food supply. In a turbid pool larger, darker specimens
are usually found, while in a clear pond with meager food supply
small pale individuals occur. Thus climatic factors may produce
the variation by their effect on larval food supplies rather than
by a direct effect on the larvae. Typical ‘ 1 summer ’ ’ larvae were
found in certain situations in the middle of February, 1944, at
Camp Shelby, near Hattiesburg, Mississippi, but most collections
throughout March were of the ‘ ‘ winter ’ 9 type.

As with the adults, the typical summer individuals approach
the characters of the Mexican C. derivator, which has an exceed-
ingly long and slender air tube.

Unfortunately larval material, although more easily obtained
in this species than adults, has been preserved in such small
quantities that detailed information on the occurrence of the dif-
ferent types in different regions cannot be given. It is very clear,
however, that in southern Mississippi the larger, darker larvae
wTith shorter, robust air tubes occur during approximately the
months when broad-banded adults are to be found and that paler
larvae with long, slender tubes are collected during the remaining
months. However, narrow-banded “summer” adults have been
reared from dark, short- tubed “winter” larvae.

CULEX (CULEX) NIGRIPALPUS THEOBALD
This primarily Neotropical species has been recorded from
several of the southeastern states, but is common only in Florida.
One hundred thirty-eight female specimens have been studied
from the following localities in Florida : Apopka, Avon Park,
Boca Raton, Leesburg, Mt. Dora, Palm Beach, Panama City, and
Sebring. In this species males show the seasonal differences only
inconspicuously, while no such differences were observed in larvag.

In this species, as in C. apicalis, the specimens found during the
cooler months of the year average larger, with more conspicuous
abdominal bands, than those collected during the summer months.
However, at no time during the year are all specimens of the
large, banded type. Intermediate types are common. In size
they resemble unbanded individuals but have a pale band on the
fourth tergum and sometimes also on the third and fifth terga.
It is such specimens that are listed as intermediate in Table 2.

298

Journal New York Entomological Society

[Vol. Llil

The banded and nnbanded types may be distinguished as
follows :

Unbanded form

Smaller,

Abdomen appearing almost com-
pletely black from above, lateral
patches of pale scales absent on
first two or more segments and
scarcely extending onto dorsum on
following segments.

The seasonal distribution of these forms may be seen in Table 2.

TABLE 2

Mar.

Apr.

May

Jun.

Jul.

Aug.

Sep.

Oct.

Nov.

Per cent unbanded

43

50

50

50

66

100

100

78

35

Per cent intermediate

(slightly banded)

57

17

33

33

34

0

0

22

40

Per cent banded

0

33

17

17

0

0

0

0

25

No. of specimens studied

28

6

6

6

22

7

15

23

40

No specimens are available for December, January, and Feb-
ruary.

It is clear that considerable percentages of banded and slightly
banded specimens persist into the summer months of June and
July. Whether they emerge during these months or are speci-
mens which survive from the spring months is not known.

Specimens studied from more northern localities (Warner
Robins, Georgia; Opelika, Alabama; Hattiesburg, Mississippi)
are all of the unbanded type but all were collected in September
or October.

Culex salinarius Coquillett is closely related to C. nigripalpus.
The banded specimens of the latter resemble salinarius even more
closely than the unbanded ones and do not run clearly to one
species or the other in the key given by King, Bradley, and Mc-
Neel.2 The pale abdominal scales of C. salinarius are yellowish

2 King, W. V., C. H. Bradley, and T. E. McNeel. 1942. The mosquitoes
of the southeastern states, U. S. Dept. Agr. Misc. Publ., 336, 96 p., 26 figs.,
6 pis.

Banded form

Larger.

Lateral patches of pale scales pres-
ent on all exposed abdominal seg-
ments, and extending across terga
as narrow (one scale wide) white
basal bands.

Dec., 1945]

Michener: Culicid^:

299

or golden, while those of nigripalpus are white. Furthermore,
when the pale scales are numerous in C. salinarius they are most
extensive at the apex of the abdomen, often largely covering the
seventh segment, while in banded nigripalpus the pale scales are
most numerous on the middle abdominal segments and do not
cover the seventh segment. The two species are easily distin-
guishable by characters of the male genitalia and the larvae.

CULEX (MELANOCONION) SPP.

Three species of Melanoconion are widespread in the south-
eastern states. Externally they are indistinguishable, or nearly
so, in the females, although the larvae and male genitalia are quite
different. The cibarial armature of the female of one species is
distinguishable from that of the others. These three species are
C. erraticus Dyar and Knab, C. peccator Dyar and Knab, and
C. pilosus (Dyar and Knab).

Winter specimens, chiefly females, of C. erraticus and probably
of one or both of the other species have been noted with white
basal bands two or three scale-rows in width on the abdominal
segments. At other seasons of the year pale scales of the abdom-
inal terga are confined to patches at the sides of the abdomen.
Available material is insufficient to give further significant data
on the occurrence of banded forms of Melanoconion.

The abdominal bands of some winter specimens of Melano-
conion are definitely conspicuous, for which reason they cannot
be properly run through such keys as that of King, Bradley and
McNeel.2

An examination of series of Culex pipiens Linnaeus, C. quinque-
fasciatu's Say, C. salinarius Coquillett, and C. restuans Theobald
showed much individual variation but none clearly correlated
with the seasons.

SUMMARY

Seasonal variation occurs in larvae and adults of Culex ( Neo -
culex ) apicalis and in adults of C. {Culex) nigripalpus and C.
{Melanoconion) spp. in the southeastern United States. In all
three cases the adults have more extensive white areas on the
abdomen in winter than in summer, and in at le'ast the first two
species winter specimens average larger than summer ones.

300

Journal New York Entomological Society

[Vol. LIII

Winter larvae of apicalis are not only larger, but darker and struc-
turally slightly different from summer specimens. Seasonal dif-
ferences were not observed in four other species of Culex.

In Culex apicalis conspicuous seasonal variation in adults is
apparently limited to the southeastern states other than Florida.
In the north and west all specimens are similar to the winter form
of the southeast, while in Florida the summer form of the other
southeastern states appears to occur throughout the year.

Dec., 1945]

Weiss: Entomology

301

SOME EARLY ENTOMOLOGICAL IDEAS AND
PRACTICES IN AMERICA

By Harry B. Weiss

Early ideas and practices in any field should be considered in
connection with the times in which they circulated. When viewed
many years later, in the light of accumulated knowledge and wis-
dom, they frequently appear fanciful and unworkable. On the
other hand some early workers and observers had ideas far in
advance of their time and with the passage of years the soundness
of their judgment has become apparent. The following examples,
drawn from various records, are cited as early expressions of
entomological thought and suggestion.

Occasionally, translations of foreign papers were published in
our early scientific journals. One of these, entitled, “A Memoir
on Animal Cotton, or the Insect Fly Carrier,” by Baudry des
Lozieres, founder of the Society of Sciences and Arts at Cape
Francois, was read before the American Philosophical Society in
1797 (Trans, vol. 5, no. 18, p. 150-159). The author, in this
paper, refers to a caterpillar that destroyed indigo and cassada
plantations at Santo Domingo. He described the caterpillar, its
food, mating of the adults, eggs, natural enemies, etc., but was
particularly interested in the activities of an “ ichneumon ’ ’ para-
site. Parsitized caterpillars changed color, increased in size and
assumed a state of “ factitious pregnancy.” Lozieres described
the cocoons of the parasite, but did not seem to be aware that the
parasitic larvae had been feeding within the body of the host for
he said, as soon as the larvae were hatched, without moving from
the spot where the eggs were laid, they yielded a liquid which
hardened upon contact with air and then spun their cocoons.
These cocoons fascinated him. He said they were made of the
finest cotton and he called it cotton because it was 1 ‘ idio-electric ’ ’
and was pervious to the ‘ ‘ electric fluid. ’ ’ As soon as the parasites
emerged, the cocoons could be carded and spun and in less than
two hours it was possible to collect 100 pints of cocoons as they
were always plentiful. Such 1 1 animal cotton ’ ’ could also be used

302

Journal New York Entomological Society

[Vol. LIII

in hospitals, as it did not inflame wounds like silk and vegetable
cotton. Baudry des Lozieres believed that the use of such cocoons
would introduce a new branch of commerce with the West Indian
colonies and make useful an insect previously known for the
trouble it caused. However, this new branch of commerce never
developed. At the time the idea was suggested, commerce was
uppermost in the minds of many persons and it is likely that
proposals involving any increase in trade always received the
most attention.

A posthumous paper on “Facts, Experiments, and Observa-
tions Relative to Some American Species of Lampyris, or Fire-
flies . . .’’by Dr. Thomas Walmsley appeared in the Medical and
Physical Journal in 1807 (vol. 2, pt. 1, Nov. 27, Suppl. 1, Mar. 7,
Sect. 2, Art. 4, p. 118-123). Dr. Walmsley reported that when
Lampyris was immersed in water, it drowned in fifteen to twenty
minutes, but shone with almost as much brilliancy as when in the
open air. Immersed in spirit of wine, it lived for five or six
minutes, remained “opake” for two or three minutes, then flashed
for three or four times, losing part of its opacity after each flash.
In a perfect Torricellean vacuum it shone for a short time and
then became “opake.” Upon the admission of the least amount
of air, it shone again. Walmsley tried carbonic acid gas, hydro-
gen, oxygenated muriatic acid gas, oxygen and azotic gas and
noted their effects upon the luminosity of the beetle. He then
described the part emitting the light and stated that the air taken
in by the spiracula had no communication with the luminous part.
He believed that the air came into contact with the “ phosphores-
cent ’ ’ substance some other way.

Numerous workers before and after Walmsley have been ex-
cited by luminous organisms and much investigational work has
been done on the biology, physics and chemistry of such forms.
It is now known that the light organs of fireflies consist of a dorsal
mass of reflecting cells and a ventral mass of photogenic cells well
supplied with trachea and nerves. It is in the photogenic layer
that the substance luciferin is oxidized in the presence of an
enzyme, luciferase, under the control of the nervous system. The
brilliance of the flash is determined by the flow of air through the
tracheae and tracheoles.

Dec., 1945]

Weiss: Entomology

303

Dr. Walmsley was born near Philadelphia in 1781. When
nineteen he began the study of medicine at the University of
Pennsylvania, receiving the degree of M.D. in 1803. He prac-
ticed in Chambersburg, Pa., and then in Elizabethtown, Md.
During the years immediately preceding his death at the early
age of 25, he was much interested in the light of fireflies and in
the natural history of these insects.

About 1810 the idea must have prevailed that insect abundance
and human disease went hand in hand because in order to combat
this idea the ‘ ‘ Medical Repository and Review of American Publi-
cations on Medicine, Surgery and the Auxiliary Branches of Phi-
losophy,” published in 1810 (vol. iv, p. 304-305) an unsigned
statement entitled “Insects in Abundance not Necessarily Con-
nected with Sickly Seasons, ’ ’ from which the following is quoted.

“In describing pestilential seasons, it has been very common
to notice the innumerable swarms of insects. Some observers of
the phenomena attendant on times of sickness have even been
led to an opinion that the growth and multiplication of these
tribes of animals is promoted by that condition of the atmosphere
and the waters which is unfriendly to the health of man. We
suspect, however, this is by no means correct. The summer and
autumn of 1800 was very productive of insects in the states of
New York, New Jersey & Pennsylvania and still no considerable
degree of sickness occurred, either in city or country. Locusts
were so thick in West Jersey and the eastern parts of Pennsyl-
vania— as to keep up a continuous noise.” Other cases are cited
of numerous forest tree insects but no remarkeble human illnesses
prevailed. “There is therefore no necessary connection between
swarms of insects & pestilential seasons. But it is nevertheless
true that during some periods of epidemic sickness, there have
been great numbers of insects. These, however, have been merely
accidntal coincidences & by no means indicate a law of nature on
the subject. On the contrary — some insects are killed by pesti-
lential air as has been often observed of common house-flies, vast
numbers of which died on the approach of the sickness in New
York toward the latter part of the summer of 1799. For the
future, physicians & others who describe pestilential seasons,
should be careful not to use the broad & unqualified term ‘in-

304

Journal New York Entomological Society

[Yol. liii

sect’ — but instead particularize the species which is abundant.
This is the safest & will mean more to readers.”

On October 10, 1810, the Rev. William Smith wrote a letter
from Norwalk, Connecticut, to Doctor Mitchill, editor of the
Medical Repository about “Observations on the Decay of Fruit
Trees, and on Lean and Shriveled Fruit, as Caused by Insects,”
which was published in 1811 (vol. 14, no. 4, p. 350-354).

It appears that some peaches on exhibition in New York were
destitute of pulp, juice and flavor. Some resemblance was traced
between their external form and that of hickory nuts, and this led
to a belief on the part of some that they represented a mongrel
production. The Rev. Smith inquired into the facts, found that
they came from the Moses Hanford orchard, of Canaan and con-
cluded that the trouble was caused as follows : 1 ‘ Some fly, to which
the apricot and nectarine trees are more accessible than others,
injects its eggs or semina, which contaminate the juices of the tree
so powerfully as to stop the fructification, at an earlier or later
stage, according to the strength of the trees. ’ ’ The tree in ques-
tion showed exudations of gum and numerous ‘ ‘ insect incisions, ’ ’
and the fruit was dried and shriveled. Probably the trouble was
due to the shot-hole borer, or to peach yellows or to “little-
peach,” or to all.

The Atlantic Monthly in 1832 (vol. 1, no. 1, p. 13-14) pub-
lished an article by Constantine Samuel Rafinesque that does him
no credit although at the time everything probably seemed to be
reasonable and in order. It is entitled “Confirmation of the
Important Discovery of the Property of Sulphur in Trees, to
Destroy all Insects Preying on Them.” The article is quoted
below and in it Rafinesque claims the now doubtful honor of
being the first “tree plugger” in America.

“Farmers and Gardeners ought to hail with rapture a safe,
certain, easy and unfailing mode of driving away or destroying
all the insects, bugs, caterpillars, lice, ants, which prey upon trees
and often kill them.

“Numberless have been the means proposed or devised to get
rid of these troublesome guests, most of which are dirty, costly,
or unavailing. Our farmers appear to have given up in despair
the hope of preventing the deadly attacks of curculios on the roots

Dec., 1945]

Weiss: Entomology

305

of peach trees, and the fruits of the plumb tree. Yet an effi-
cacious mode is said to have been found several years ago in
France, perfectly efficacious and applicable to all cases and all
trees. The name has not even reached us. But we claim the
honor to have been the first to make known the process in
America, in 1823 in Kentucky, and in 1827 in Philadelphia. Yet
the most useful knowledge is so slow to spread, that the fact is
hardly known yet, or doubted by those who know of it.

“We are happy to be able to publish two direct experiments in
support of the fact and discovery.

“First. We bored and plugged with sulphur in the usual way,
a plumb tree which commonly dropped every year all the plumbs
before becoming ripe, the curculios lodging eggs in their germs.
This was done when the tree was in blossom. On that year hardly
any fruit fell, and the tree produced quite well.

“Second. We find in the Genesee Farmer of January 28, 1832,
that a young willow nearly killed by aphis or lice, and pissmires
feeding on their honey, was quite revived in three days, and all
the lice and ants driven off, by boring the tree with an augur five
feet from the ground and three-fourths through the diameter,
filling with brimstone and plugging tight. The tree has thrived
ever since.

“The modus operandi of this singular process is very easy to
explain. The vital energy of the tree and sap, dissolves the sul-
phur, carries it into circulation, and evolves it in sulphuric gas
evaporating through all the pores of branches, leaves and fruits.
This gas is a deadly poison to insects and all animals, it suffocates
them or drives them away as soon as they begin to smell it ; but
no injury whatever results to the tree.

“We have never heard yet of any direct experiment on peach
trees; but we are sure it will answer quite as well. If the sul-
phuric emanation could not reach quick enough the roots of the
trees which are commonly attacked; the plugging must be near
the root or at the time of the descending sap, when it will sooner
reach the roots. Let it be tried and the results made known.
C.S.R,”

The Franklin Society of Providence, R. I., published at Provi-
dence in 1829 an 8-page pamphlet entitled “General Directions

306

Journal New York Entomological Society

[Vol. LIII

for Collecting and Preserving Articles in the Various Depart-
ments of Natural History,” in which it is stated that beetles may
be killed by immersion in spirits of wine, or hot water and also
by touching their heads with spirits of turpentine or by putting
them under an inverted tumbler and filling it with sulphur fumes,
by means of lighted matches held under one edge. However, J. P.
Kirtland, 24 years later, or in 1852, in the American Journal of
Science and Arts (vol. 63, no. 38, March, p. 286-287) said that
compressing the thorax, puncturing the thorax with a needle
dipped into oxalic acid, killing with ether, chloroform, or sulphur
fumes, were all objectionable because they impaired the beauty
of the specimens and mutilated them as well. His method was
to puncture the thorax once or twice with a needle dipped previ-
ously in a strong solution of potassium cyanide. He advised also
that, in order to prevent oily exudations and to preserve colors,
the abdominal contents should be removed and replaced by cot-
ton. Detailed directions were given for the dissection, stuffing
and sewing-up of the body.

At the annual fair of the New Haven Agricultural Society, held
at New Haven, Connecticut, Mr. Noyes Darling, on October 1,
1845, gave a popular talk on the importance of knowing about
insects and much of his information came from Harris ’s ‘ ‘ Insects
Injurious to Vegetation.” Darling recommended the study of
insects as a part of education and said, “If the time ever comes,
as it may come, when the interference of government shall be
required to stay the ravages of insects, a thorough knowledge of
them will be indispensable to enlightened and well directed legis-
lation. ” In addition, he asked, “Why should not government
make war upon cutworms and plum weevils, as well as upon bar-
berry bushes and Canada thistles ? ’ ’ Darling, who was a gradu-
ate of and a tutor in Yale, one time mayor of New Haven, a judge
of the county court, and interested in horticulture and agricul-
ture, would be amazed, were he alive today, at the entomological
activities of government.

A publication called “The Annual of Scientific Discovery” was
started, in Boston, in 1850, to record the most important dis-
coveries and improvements in mechanics, natural philosophy,
chemistry, zoology, botany, geology, etc., etc., etc. As a rule ento-

Dec,, 1945]

Weiss: Entomology

307

mology did not occupy much space in the Annuals, but in the one
for 1851, a statement was copied from the “London Journal of
Arts” about the electroplating of insects, flowers, etc., as a means
of preserving them. The insects were first steeped in a solution
of phosphorus and then in a solution of nitrate of silver. The
phosphorus caused the silver to precipitate upon the insect and
to form a very thin metallic coating over every part of it. Upon
this a thicker deposit of metal was obtained by the “electrotype”
process, after which two or three small holes were made through
the coating and the specimen heated so as to drive off the mois-
ture. This method of preserving insects was ignored by collectors
and museums alike.

Biological control was not highly regarded at first even by some
entomologists. In Brooklyn, N. Y., there appeared, in 1862, a
report on “The measure worm Ennomos subsignaria; a descrip-
tion of the insect in all its metamorphoses, its history and prog-
ress, and a systematic plan for its final extermination, together
with remarks on the state of the shade trees, in the city of Brook-
lyn, N. Y.” This report, prepared by H. A. Graef and Edward
Wiebe, was submitted to a large committee appointed by the
Brooklyn Horticultural Society, which ordered its publication.

The insect in question had been a pest of Brooklyn shade trees
for a number of years. In 1860 the Common Council passed a
resolution “to free the city from the perpetually increasing mea-
sure worm nuisance, even by removing from our streets all trees
infested by this insect.” The control plan of Graef and Wiebe
was extensive. It involved a mapping of the area, the scraping
of all egg masses from all trees during the winter. During April,
tar rings four inches wide, were to be applied to the trunks and
larger branches. Such caterpillars as were found beyond the tar
bands were to be removed with the foliage by means of shears.
Any caterpillars escaping the operations thus far were to be
syringed with a strong tobacco infusion or destroyed by daily
repeated beatings or jerkings of the tree and branches with
proper tools and machinery. If any caterpillars still resisted and
spun cocoons then the cocoons were to be gathered and destroyed.
If, after all the foregoing had been done, adults appeared they
were to be caught in nets or syringed from the trees like the cater-

308

Journal New York Entomological Society

[Vol. LIII

pillars. Such uninterrupted warfare, carried on with energy,
must as a matter of course, so the authors stated, reduce the infes-
tation considerably in a short time. In addition the cooperation
of wrens was to be enlisted by the erection of cheap, useful, simple
and lasting wren houses, in the city streets.

In the preparation of their report, the authors consulted ento-
mologists and other scientific and learned persons. It was re-
ceived flatteringly by almost everyone. There was, however, one
exception. Dr. Trimble, of Newark, N. J., a member of the origi-
nal Committee on Shade Trees, dissented. He said that it was
useless to spend a single dollar, because a little “fly” was destined
to do the controlling. The authors would not have mentioned
this at all if Dr. Trimble had not published his adverse opinion
in the “Newark Weekly Mercury,” of October 14, and sent copies
of this paper to most of the members of the Committee. The
authors claimed that the insect had been getting worse over a
period of 25 years and they did not believe that a parasite, all at
once, would be able to combat an enemy so strong. As practical
men, they preferred to rely on their own exertions, refusing any
help from ‘ ‘ a fanciful agency. ” It is not known if the measures
recommended in the report were attempted or if everything was
left to the parasite. Mr. H. A. Graef, one of the authors of the
report, came from Aix-la-Chapelle, Germany, in 1848, and settled
in what is now known as Bay Ridge, Brooklyn. He established
himself as a florist and he was also interested in natural history
and in collecting local plants. His son, Edward L. Graef, previ-
ous to his death, was a well-known member of the Brooklyn Ento-
mological Society.

Dec., 1945]

Kapp: Psychodid^

309

NEW PSYCHODIDiE FROM BARRO COLORADO

ISLAND

By William F. Rapp, Jr.

Five apparently new Psychodidae together with one new genus
are described in the following pages. The flies were all collected
on Barro Colorado Island by Dr. C. H. Curran. The types are
in the American Museum of Natural History. The writer is in-
debted to Miss Janet L. Cooper for the drawings in this paper.

Psychoda Latreille

Psychoda eburna, new species
( eburneus , ivory-white)

Female. — Antennae (Fig. 1, a, b) thin, beads ovate, brownish- white-
haired; face and occiput yellowish-white, the front with scattered long, white
hairs. Thorax with dense long brownish-white hairs except on the scutum
which is bare and yellowish white. Legs with abundant brownish-white hair,
uniform from coxa to tarsal segments. Abdomen with very thick and long
brownish- white hairs. Wing (Fig. 1, c) thickly brownish -white-haired, espe-
cially along the posterior margin. Length of wing, 2 mm.

Type. — Holotype, female, Barro Colorado Island, Canal Zone,
January 10, 1929 (C. H. Curran).

This species may be easily recognized by the shape of the
antenna beads and the light brownish-white color.

Psychoda maculosa, new species
( maculosus , spotted)

Female. — Antennae (Fig. 2, b, c) thin, light-brown-haired; face and occi-
put with light-brown, the front with light-brown hair. Palpi short, with
brown hair. Thorax with dense brown hair except on the anterior scutum
which is shining light-brown, entire squamae with long brown hair. Legs
with long brown hair, also with brown scales, tarsi with brown white scales,
base of first tarsal segment with a brown ring at base. Abdomen brown-
haired, the dorsum, lighter at the segmental lines and with a black spot at
center of each segment. Genitalia with brown hair. Wings (Fig. 2, a)
thickly brown-haired, with dense clusters of hairs at the ends of the veins,
which form small brown patches. Length of wing, 2.25 mm.

Types. — Holotype, female, Barro Colorado Island, January 7,
1929; four paratypes, Barro Colorado Island, January 8, 1929
(3) and January 10, 1929. All collected by C. H. Curran.

310

Journal New York Entomological Society

[Vol. LIII

This species may be recognized by the shape of the antennas
and the spots at the ends of the wing veins.

Psychoda brevitarsa, new species
( brevis , short; xapabs, tarsus)

Female. — Antennae (Fig. 3, b, c) thick, brown-haired, segments ovate in
shape; densely haired. Face with brown iridescent scales, occiput brown,
the front with brown hair. Palpi long with short brown hair. Thorax with
dense brown hair, except on the sides which are shining brown. Legs with
scattered brown hairs, but completely covered with brown iridescent scales.
Tarsal segment very short. Abdomen brown-haired, last segment with light-
brown scales. Genital segment with two long black hairs. Wings thickly
brown-haired. Length of wing, 1.75 mm. (Fig. 3a).

Types. — Holotype, female, Barro Colorado Island, Canal
Zone, January 8, 1929, three paratypes Barro Colorado Island,
January 8, 1929 (2) and January 10, 1929. All collected by
C. H. Curran.

This species may be recognized by the antennae and the short
tarsi.

Psychoda atraseta, new species
( ater ■ black; seta, hair)

Female. — Antennae thick, the last two segments with thin white hairs, the
rest thick with abundant black hairs; face and occiput black, the front with
black hair. Palpi long, with bushy black hair, the apical segment white.
Thorax with dense black hair except on the area directly below the wing
which is yellowish-white. Legs with abundant black hairs, tarsi with alter-
nate bands of black and white hairs, basal segment, white. Abdomen black-
haired, the dorsum gray. Second and third segments nearly devoid of hair
on dorsum. Genital segment with long brown hair. Wings (Fig. 4, a)
thickly brown-haired, with dense patches of black hair at the ends of the
veins. Length of wing, 2.25 mm.

Type. — Holotype, female, Barro Colorado Island, Canal Zone,
January 10, 1929 (C. H. Curran).

This species may be recognized by the aiitennge, bushy black
with white terminal.

Kupara, new genus

( [Tcupar , cooper; in honor of J. L. Cooper)

This genus may be recognized by the forming of R2 and R3, beyond the
center of the wing. R4 and R5 branch at the center of the wing. Mx and M2
branch past the center and before R2 and R3. M2 does not join with Mj.
(Fig. 5, a).

Genotype. — Kupara albipeda, new species.

Dec., 1945]

Rapp: Psychodid^e

311

Kupara albipeda, new species
(albus, white; yes, foot)

Female. — Antennae (Fig. 5, b, c) slender, dark brown-haired; face dark
brown, occiput with light brown hair, the front with dark brown hair. Palpi
long with dark brown hair. Thorax shining brown, with clumps of long
brown hair. Legs, femur and tibia with thick dark brown hair, tarsi with
white scales. Abdomen brown with white bands at the segmental lines.
First segment with very few hairs, but increasing on the following segments,
genital segment thick with dark brown hairs. Wings (Fig. 5, a) brown-
haired and brown in color, at tips of radial veins a patch of very long dark
brown hair. Length of wing, 2 mm.

Types. — Holotype, female, Barro Colorado Island, Canal Zone,
February 13, 1929; paratype, Barro Colorado Island, Canal
Zone, February 18, 1929 (C. H. Curran).

This species may easily be recognized by the white tarsi and
the antenna.

312

Journal New York Entomological Society

[Vol. LIII

(Jour, N. Y. Ent. Soc.), Yol. LIII

(Plate III)

PSYCHODA

Figure 1.

Figure 2.

Figure 3.

Figure 4.
Figure 5.

Psychoda eburna, n. sp. — (a) tip of antenna, (b) base of antenna,
(c) wing.

Psychoda maculosa, n. sp. — (a) wing, (b) tip of antenna, (c)
base of antenna.

Psychoda brevitarsa, n. sp.— (a) wing, (b) tip of antenna, (c)
base of antenna.

Psychoda atraseta, n. sp.— (a) wing.

Kupara albipeda, n. sp. — (a) wing, (b) tip of antenna, (c) base
of antenna.

Dec., 1945]

Richards & Cutkomp: Neuropathology

313

NEUROPATHOLOGY IN INSECTS1

By A. Glenn Richards, Jr., and Laurence K. Cutkomp

Zoological Laboratory, University of Pennsylvania, Philadelphia,

PENNSYLVANIA2

Neuropathological pictures resulting from insecticide action
have been recorded for the insect nervous system by Kruger
(1931), Klinger (1936), Richards (1941), Wigglesworth (1941)
and especially by Hartzell and his colleagues (1932-1944). The
aim of the present paper is to present the numerous data we
have gathered on various compounds, to compare these data with
other published data, and to discuss the relation between the
various pathological pictures and the cessation of nerve action.
The effects of compounds on other tissues is not considered.
Analysis of the action of pyrethrum is presented in considerable
detail. Other compounds are then treated more briefly, prac-
tically as summaries, without specific reference to the hundreds
of experimental animals involved. Such a space-saving proce-
dure seems warranted in view of the thesis of the present paper;
this thesis being that the visible pathological changes induced in
nerves by insecticides are at least largely postmortem and accord-
ingly too complex for analysis at the present time.

MATERIALS AND METHODS

For most of the original data presented in the present paper
Ave used adults or sometimes large nymphs of the American cock-
roach ( Periplaneta americana). In some cases mosquito larvae
were employed (Cidex pipiens and Aedes cegypti). The sub-
stances tested were introduced in acute dosages (commonly very
large doses) either into the tracheae or hemocoel by means of a
syringe, or fed orally or applied to the cuticle. Some volatile

1 The work described in this paper was done under a contract, recom-
mended by the Committee on Medical Research, between the Office of Scien-
tific Research and Development and the University of Pennsylvania. Valu-
able technical assistance was given by Miss Jane L. Weygandt during the
course of these experiments.

2 Present address of both authors : Division of Entomology, University of
Minnesota, St. Paul 8, Minnesota.

314

Journal New York Entomological Society

[Vol. LIII

substances were also used as vapors. Repeated chronic dosing
was usually not tried. It has been shown in an earlier paper in
this series (Richards & Weygandt, 1945) that organic solvents
of low water solubility definitely tend to accumulate in the cen-
tral nervous system particularly when applied in the tracheae
(Figs. 3, 13, 14). Tracheal injections have several advantages:
the distribution of the substance can be checked with fair accu-
racy on dissection; also the injection may be made at either end
of the animal, one end then serving as the experimental end, the
other end serving as one kind of control (for some purposes this
gives excellent results). The method of tracheal injection was
accordingly commonly employed in studying the effects of lipid
soluble materials on the nervous system.

The majority of the microscopic observations were made by
the polarized light method which is highly sensitive and avoids
questions of fixation artifacts (Schmitt & Bear, 1939; Richards,
1944). Quantitative changes in the birefringence of nerve cords
can be measured with a high degree of accuracy (see preceding
reference for illustrations). It is not possible to repeat the
details of this technique here. Suffice it to say that with this
method it is possible to analyze the effects of various toxins on
both the nerve axoplasm and the surrounding lipo-protein sheath.
In normal nerves there is a balance between positive and negative
components of the birefringence, the positive elements being
slightly stronger (having greater amplitude). Measurements in
saline give the amplitude by which the positive components (axis
cylinder and sheath proteins) exceed the negative components
(sheath lipids). In glycerine (15 per cent in saline) the form
birefringence of the proteins is largely masked, and measure-
ments give the amplitude by which the birefringence of the lipid
sheath exceeds the protein birefringence in this solution. Theo-
retically, elimination of all the positive (protein) birefringence
should show the true negative (lipid) value, and vice versa. It
is not possible at present to do this with nerves. But so long as
conditions of measuring are kept standard the figures from vari-
ous specimens can be compared quantitatively. When the mea-
surements in glycerine remain constant (within the normal
range), the previous measurements in saline indicate the condi-

Dec., 1945]

Bichards & Cutkomp: Neuropathology

315

tion of the positive or protein components ; when both are lowered
by the same or different degrees allowance must be made for the
fact that we are dealing with a balance ; when a nerve is isotropic
in saline but not in glycerine then the two opposed components
are presumably in balance and the measurement in glycerine
gives a measure of the amplitude of each ; and only when a nerve
is isotropic in both saline and glycerine can we say that there
has been an approximately complete decay of the birefringence.

With pyrethrum, valone, aniline, petroleum oils and a number
of other materials, serial sections were prepared following routine
paraffin procedures. Nerve cords or brains were fixed in 95 per
cent ethyl alcohol plus 5 per cent glacial acetic acid, and after
sectioning stained with toluidine blue and acid fuchsin. Polar-
ized light analyses are applicable only to nerve fibers ; investiga-
tion of possible histological effects on the nerve cell bodies must
be done by the more usual methods of histopathology.

In many cases pathological effects can be seen even under the
ordinary light microscope without fixation or other treatment.
This is particularly true of substances which cause opacity,
chromatin clumping or the release of droplets in the highly trans-
parent nervous system of mosquito larvae (Figs. 5, 15).

The condition of the insects was noted after application and
at the time of dissection. A further check was made on the func-
tional condition of the nerve cord by applying a direct electrical
shock to selected parts of the nerve cords and observing any re-
sponses. Platinum electrodes were used either with an ordinary
inductorium or an electronic multivibrator (thyratron controlled,
condenser discharge circuit) adjustable to give shocks with fre-
quency from 0 to 1000 per second, the duration and interval
being separately controlled, and the voltage variable from zero
to several hundred volts.

NOBMAL STBUCTUBE OF THE INSECT NEBYOUS SYSTEM

The central nervous system of insects consists of nerve cells
and their fibers surrounded by thin lipo-protein sheaths (Rich-
ards, 1944) and held together by tracheae, neuroglia and a tough
outer sheath, the neural lamella (Scharrer, 1939). The nerve
cells are of typical structure but range from very small to only

316

Journal New York Entomological Society

[Vol. LIII

moderate in size. The nerve fibers are commonly very small
(to less than 1 micron). The nerve sheaths are always thin, and
in the case of small fibers are not detectable by ordinary micro-
scopic methods and require polarized light methods for their
demonstration.

The central nervous system (Fig. 2) is a solid structure ar-
ranged with the major fiber tracts centrally and the cell bodies
in a peripheral layer in the segmental ganglia (Hanstrom, 1928).
The interganglionic connectives and peripheral nerves consist of
nerve fibers with some tracheae and neuroglia. All ganglia, con-
nectives and peripheral nerves are surrounded by the neural
lamella.

The tracheal network in ganglia is most rich in the boundary
between the cell body layer and the central fiber tract region.
This is readily seen in favorable preparations (Figs. 1, 2). The
distribution of tracheae at this boundary may weaken it, and
also, in cases where toxins are applied via the tracheae, results in
a maximum concentration and effect here. It is not surprising,
therefore, that one of the recorded pathological changes in insect
ganglia is a tendency for cell body and fiber tract regions to
separate (Richards, 1941).

EFFECTS OF PYRETHRUM

For a standard of comparison with a well-known neurotoxic
insecticide, two series of experiments were performed with
pyrethrum concentrates. In the first series a known pyrethrum
concentrate3 was injected into the first thoracic spiracle of 25
adult cockroaches. Individual records were kept on each speci-
men: cessation of general movements, cessation of heartbeat, re-
sponsiveness of the nerve cord to electrical stimulation at the
time of dissection, and the extent of distribution of the pyrethrum

3 Sample received from McLaughlin, Gormley, King Company, Minne-
apolis. Assay: 10.80 per cent pyrethrin I, 9.85 per cent pyrethrin II, total
20.65 per cent pyrethrins. In the absence of chemically pure materials there
is, of course, no way of evaluating how much of the effect may be due to the
other 80 per cent of the material, or whether or not there is any difference
between the effects of pyrethrin I and II. This concentrate, however, is the
kind used in preparing insecticides.

One should also remember that in tracheal injections we are always dealing
with relatively large doses and rapid penetration.

Dec., 1945]

Richards & Cutkomp: Neuropathology

317

concentrate throughout the tracheal system. Most of these speci-
mens were examined with polarized light but representative
specimens were serially sectioned and compared with published
figures and descriptions of pyrethrum lesions. In the second
series 24 last instar nymphs and adults were injected in the first
or second thoracic spiracle with an unassayed pyrethrum concen-
trate, records kept individually hs above, and the nerve cords
later removed and examined with polarized light.

Following the tracheal injection there is an immediate initial
paralysis that may be in part a reflex immobilization.4 A few
minutes later there is a partial recovery followed by a gradual
decline and slower and slower movements of peripheral parts
and eventually complete death. The legs, abdomen and heart
may continue moving for many hours (up to 52 hours in these
experiments). Electrical stimulation to test responsiveness was
routinely performed on specimens still moving appendages at
the time of dissection; specimens treated J to 52 hours prior to
dissection showed in no case any response to direct electrical
stimulation of the affected nerve cord. The fact that muscular
movements may continue for many hours after the nerve cord
is irreversibly paralyzed or even highly degenerate is good evi-
dence for the selective nervous action of pyrethrum.

Analyses with polarized light revealed that nerve cords para-
lyzed with pyrethrum show first an effect on the axoplasmic col-
loid (axis cylinder) of the nerve, then somewhat later an effect
on the lipid component of the nerve sheaths. It seems to be the
latter that gives rise to the most prominent pyrethrum lesions.
Both the axis cylinder of the nerves and the lipo-protein sheath
may lose the ultrastructure responsible for the optical properties
prior to the cessation of movement of appendages but not prior
to paralysis of the nerve cord. These effects on the ultrastruc-
ture responsible for the optical properties are first seen in the
region of maximum penetration, but with the passage of time
extend to all parts of the nerve cord and to at least the larger
peripheral nerves. The experimental technique employed was
not sufficiently refined to permit direct demonstration of the

4 Reflex immobilization from injection of pyrethrum is not a specific action.
It may result from any tracheal injection.

318

Journal New York Entomological Society

[Vol. LIII

status of the optical properties of a nerve at the moment of
paralysis, but we can infer that the visible effects are all post-
mortem since affected nerves are always paralyzed whereas
normal-appearing nerves may or may not be paralyzed. The
paralysis prevents both direct stimulation and the passage of
impulses through to unaffected regions.

No illustrations of pyrethrum effects are given in the present
paper. Adequate illustrations of the more advanced stages of
degeneration as seen in serial sections have already been pub-
lished by Klinger, Hartzell et at., and Wigglesworth. The de-
terioration of optical properties can be measured with consider-
able accuracy but there is no point in publishing photographs of
the various stages. Such a series of pictures would simply range
from those published for normal nerves (Richards, 1944) to a
complete absence of contrast (= no picture).

The two series, totalling 49 specimens, were sufficiently con-
sistent so that the data may be combined and analyzed as follows :

In ten specimens that were dissected from \ to 2J hours after
treatment there was usually a slight but significant decay of the
positive component of birefringence but not any clearly demon-
strable change in the negative component.5 This is interpreted
as meaning that the proteins of the axis cylinder of the nerves
are degenerating but that the nerve sheaths are still normal.
The decay of birefringence of the axis cylinder was most evident
in the injected regions, the abdominal or posterior abdominal
connectives being normal. Another specimen (1-J hrs.) serially
sectioned and stained showed seemingly normal nerve tissue.
The large nerve cell bodies and the fiber tracts appeared normal ;
there was a little homogeneous staining in the vicinity of pyreth-
rum-filled tracheae but this is of questionable significance, and
chromatin clumping was fairly general but not universal in the
medium-sized nerve cells, usually absent in the large nerve cells
and always absent in the neuroglia cells. The “typical” pyreth-
rum lesions were not found in this specimen.

5 In one of the above cases the reading in saline was approximately normal
but an unusually high negative reading in glycerine implies an effect on the
axis cylinder which was masked by slight stretching of the nerve cord during
dissection. This phenomenon can be reproduced experimentally by deliber-
ately stretching pyrethrum-treated nerve cords.

Dec., 1945]

Richards & Cutkomp: Neuropathology

319

In eleven specimens that were dissected in 3i to 7 hours after
treatment the positive component of birefringence was consider-
ably reduced but the negative component was still strong (nerves
isotropic in saline but strongly birefringent in 15 per cent glycer-
ine). This is interpreted as meaning that the optical properties
of the proteins of the axis cylinders have degenerated to half
or less than half their normal value, but that the lipo-protein
sheaths are still normal or nearly so. This ' decay commonly
extended farther posteriorly than the visible distribution of
pyrethrum in the tracheae but was not complete in the posterior
abdominal segments. Another specimen (5J hrs.) serially sec-
tioned and stained showed relatively slight pathological changes,
especially in stainability, throughout the area of penetration.
Histological effects are thus evident but the changes are not of
the extreme type described by Klinger, Hartzell and Wiggles-
worth.

In six specimens that were dissected from 12 to 14 hours after
treatment the positive component of birefringence was low and
the negative component was also somewhat reduced (nerves iso-
tropic in saline but moderately birefringent in 15 per cent glycer-
ine). This is interpreted as meaning an advanced state of de-
generation of the optical properties of the axis cylinders and a
beginning of degeneration of the lipids of the nerve sheaths. The
decay is also more general throughout the nerve cord after this
time interval. No specimen from this set was sectioned.

In eight specimens that were dissected from 24 to 25 hours
after treatment the positive component of birefringence was
further reduced or abolished and the negative component was
also greatly reduced or even abolished in the regions of greatest
penetration (less or not at all affected in regions far removed
from the visible distribution of pyrethrum in the tracheae) . This
is interpreted as meaning a complete or nearly complete degen-
eration of the axoplasmic colloid, plus an advanced degree of
degeneration of the nerve sheaths. Effects are apparent through-
out the nerve cord at 24 hours and are not limited to regions
where pyrethrum is visible in the tracheae. Another specimen
(30 hrs.) serially sectioned and stained showed chromatolysis,
vacuolization, etc., of the types described as “characteristic”

320

Journal New York Entomological Society

[Vol. LIII

for pyrethrum by Klinger, Hartzell and Wigglesworth, but com-
monly the sections illustrated by these authors represent more
advanced stages of degeneration than is to be seen in this speci-
men.

A single specimen that was still moving its metathoracic legs
52 hours after treatment had a nerve cord with seemingly com-
plete decay of all its optical properties (isotropic in both saline
and glycerine). As seen without crossed Nicols, the axis cylin-
der and nerve sheaths were both obviously degenerate.

In nine specimens that appeared to be dead at the time of dis-
section (29J to 56 hrs.) there was generally a complete decay of
both the positively and negatively birefringent components of
the nerve cords. In specimens with shorter exposures (29-30
hrs. ) the negative component had usually degenerated completely
only in the regions where pyrethrum was visible in the tracheae ;
in specimens with the longer exposures (55-56 hrs.) the decay
seemed complete throughout the nerve cord although pyrethrum
was visible in only the thoracic tracheae. As seen without
crossed Nicols, these nerve cords were obviously degenerate. An-
other specimen (55J hrs.) serially sectioned and stained showed
extensive degeneration of the “typical” pyrethrum type.

The above data show that pyrethrum does have a selective ner-
vous action as previously reported by others. As seen with
polarized light, pyrethrum first causes degeneration of the colloid
of the axis cylinder (and likely of the nerve cells). The degen-
eration of the nerve sheaths occurs later. Degeneration proceeds
from the region of application towards and finally to other
regions. The death of the animal does not bear any fixed rela-
tionship to the degree of degeneration of the central nervous
system. All the histological effects seen in our experiments are
subsequent to irreversible paralysis and are accordingly to be
classed as postmortem pictures.6 The appearance of vacuoliza-

6 Hartzell & Scudder (1942) used “moribund” flies four hours after
treatment and obtained both slight lesions and chromatin clumping. Differ-
ences in technique and in the test animals used prevent strict comparison to
these data. However, the use of peripheral movements as an index of life
does not prove that the visibly pathological nerve cells were still living. Our
1 1 living ’ ’ cockroaches had 1 1 dead r ’ nerve cords. It is possible that some
nerve cells may show changes before they die but this would be difficult to
prove and has not yet been proven. Chromatin clumping is discussed in a
later section entitled Suffocation and Acidity.

Dec., 1945] Richards & Cutkomp: Neuropathology 321

tion coincides with the time of breakdown of the lipo-protein
sheaths and may be due to sheath products (Richards, 1943).
The histological effects develop rather slowly following death of
the nerves concerned, and we have found that these effects are
remarkably similar to those seen in the autolytic degeneration of
nerves in saline. It seems questionable, therefore, whether
pyrethrum has any causal relationship to “pyrethrum lesions”
other than killing the nerves. It is quite possible that a lethal
concentration of pyrethrum develops in the central nervous sys-
tem so far in advance of that in other tissues that advanced
autolysis may develop there before other tissues even die.

EFFECTS OF ‘ ‘ THANITE ’ ’

This commercial product and its active ingredient, isoborneol
thiocyanoacetate (96 per cent) were studied mostly for the effect
of the water miscible (soluble ?) fraction in the culture water
of mosquito larvae. Some tracheal injections were also per-
formed with mosquito larvae. Histological examination was
made only by means of stained serial sections. The same results
were obtained with “Thanite” as with isoborneol thiocyanoace-
tate. Selective degeneration of the nervous tissue is not so
marked with this material as with pyrethrum but similar lesions
(vacuolization) are produced in larvae still capable of feeble
movements (Pig. 7).

EFFECTS OF PETROLEUM OILS

The pathological changes resulting to brains and nerve cords
from the application of various petroleum oils in the tracheae
of mosquito larvae have already been described and illustrated in
some detail (Richards, 1941). It was shown that asphyxiation
might or might not occur in the experiments depending on the
technique employed. It was also shown that the lighter oils con-
taining unsaturated compounds produce a degeneration involv-
ing chromatolysis, cell separation and fiber degeneration (stained
sections). At that time it was suggested that the histological
effects might be postmortem insofar as the cells of the nerve cord
were concerned, and that the effect is at least partly intercellular
(i.e., on the nerve sheaths). As seen in stained sections there
are certain similarities between the histological effects of petro-

322 Journal New York Entomological Society [Vol. liii

leum oils and those of pyrethrum and “Thanite.” Cell separa-
tion and fiber separation are more distinct with petroleum oils,
perhaps partly due to a solvent action of the oils on the sheath
lipids.

The saturated, so-called non-toxic, petroleum oils can kill in-
sects. They have not, however, been shown to produce any
noticeable change in the nervous system or other tissues prior to
death of the insect. When asphyxiation is avoided they do not
produce any cytological changes visible in stained sections. Also,
saturated petroleum oils do not alter the optical properties of
the nerve fibers of cockroaches.

EFFECTS OF COBRA YENOM AND LYSOLECITHIN

A number of neurotoxic snake venoms have been studied by
mammalian physiologists. Cobra venom is most commonly used.
This venom is a complex mixture of toxins (see Ghosh, 1940;
De, 1941 ; Macht, 1941 ; etc.) in addition to an enzyme, lecithinase
A, which splits the fatty acids radical from lecithin to form the
potent hemolytic agent called lysolecithin. In some of the older
literature it was assumed that cobra venom acted on mammals
by producing lysolecithin in vivo , the lysolecithin then causing
death (Page, 1937, p. 80). In view of the obvious sheath degen-
eration caused in insects by some insecticides, this hypothesis was
introduced into entomological literature by Richards (1943).
Subsequently, however, experiments designed to test this sug-
gestion have not given confirmatory evidence, and it seems advis-
able to abandon the hypothesis.

Cobra venom7 is quite toxic to insects. It first paralyzes the
nervous system, then later results in the deterioration of the
optical properties of the axis cylinders, and only lastly causes a
breakdown of the lipids of the nerve sheaths. This makes it
seem dubious that lysolecithin formed during sheath breakdown
could be an important factor in the action of cobra venom on
insects. As a further check, lysolecithin was prepared by the
action of crude cobra venom on egg yolks (chicken) following the
techniques of King & Dolan (1933) and Levene et al. (1923, 1924) .
The preparation obtained was potent for hemolysis of erythro-

7 Supplied through the courtesy of Hynson, Westcott & Dunning, Balti-

more.

Dec., 1945]

Richards & Cutkomp: Neuropathology

323

cytes and seemed reasonably pure. Injections of maximal doses'
of saline suspensions into the hemocoel of cockroaches was with-
out effect. One can question whether the lysolecithin penetrated
from the insect’s blood into the cells, but certainly the data do
not support the idea that lysolecithin is toxic to insects. In ver-
tebrates it seems that the effect of lysolecithin is expressed largely
by hemolysis and rupture of capillaries — phenomena that are not
found in insects.8

Bee venom is known to contain lecithinase A (Feldberg, 1940).
Perhaps wasp venom does also. Hartzell (1935) has recorded
nerve pathology with vacuolization in the ganglia of cicadas
paralyzed by wasp stings. Insect venoms, like snake venoms, are
complex mixtures (Beck, 1935). We have not had any venom
of Sphecius speciosus for study, but certainly with cobra venom
the pathological effects on insect nerves are subsequent to the
death of the nerve cells.

EFFECTS OF TRIORTHOCRESYL PHOSPHATE

This is the causative agent of “ginger paralysis” in humans.
It is moderately toxic to mammals but its exact effects are not
clear (Lillie & Smith, 1932). Hartzell (1934) has shown that
it can be used to produce lesions in the nervous system of insects.
We have confirmed this with the polarized light method, but
again the histological effects are found only after paralysis
(death) of the nerve cells.

EFFECTS OF LIPID SOLVENTS

The selective accumulation of various lipid solvents in the cen-
tral nervous system of mosquito larvae has been treated in detail
in a preceding paper (Richards & Weygandt, 1945). It was
reported therein that organic solvents may diffuse from the
tracheae to become uniformly distributed throughout the fiber
tracts (xylol, chloroform, etc., Fig. 14) or they may penetrate
the tracheal walls and collect outside the tracheae as droplets
which disperse gradually (essential oils, Figs. 3, 4; various glycol
derivatives, Fig. 13, etc.). Stained serial sections may reveal
no visible effect in the case of the former, especially those that

8 Several potent toxins of mammals have little or no effect on insects. In
addition to lysolecithin, the list includes histamine and curare.

324

Journal New York Entomological Society

[Vol. LIII

are common ingredients of fixing fluids, perhaps partly because
the same or similar solvents are used in histological techniques.
Stained serial sections of brains into which the organic solvent
has penetrated as droplets may show a considerable degree of
seeming vacuolization, but the so-called vacuoles in these cases
may represent the precipitation of tissue components around the
periphery of invading droplets which are later dissolved away
(Fig. 9).

With the polarized light method we studied the effects of in-
jecting ethyl ether, ethylene dichloride, chloroform, xylol and
toluene as fluids into the tracheae of cockroaches. The effects of
ethyl alcohol on extirpated nerve cords were also studied. When
these solvents are used as fluids (relatively large amounts) they
quickly block metatropic reversal by dissolving the sheath lipids.
They do not destroy the optical properties of the axis cylinder, in
fact they have good fixing properties except for lipids (see Rich-
ards, 1944, Figs. 5-10).

It appears, however, that the detectable solvent effects are not
necessarily concerned with the anesthesia or death produced by
these substances. When cockroaches are anesthetized or killed
with the vapors of chloroform or ethylene dichloride there is no
detectable change in the optical properties of the nerve cords.

EFFECTS OF INSECT EEPELLENTS

Insect repellents are organic compounds, and like, most organic
compounds have lipid solvent properties. Accordingly, it is not
surprising that when introduced into the tracheal system of in-
sects they tend to accumulate in the nervous system, and that
large doses of the fluids so introduced abolish (dissolve) the lipid
component of nerve birefringence. Of more interest is the fact
that they produce no detectable changes other than those which
can be ascribed to their solvent properties. As with the preced-
ing materials, the visible effects are subsequent to paralysis and
presumably subsequent to death of the cells concerned. Mate-
rials tested included dimethyl phthalate, 2-ethyl hexanediol-1,3
(‘ ‘ Rutgers 612 ” ) , aja'-dimethyl-a-carbobutoxydehydro-gamma-
pyrone (“Indalone”), and synthetic Oil of Citronella.

When the above repellents are used as vapors they can kill
cockroaches. It is particularly easy to kill cockroaches with the

Dec., 1945]

Richards & Cutkomp: Neuropathology

325

vapor (saturated atmosphere at room temperature) of synthetic
citronella (12-26 hrs.). Dimethyl phthalate vapors also kill
readily. Cockroaches are more resistant to “Indalone” and
ethyl hexanediol but are eventually killed. The interesting
aspects shown for repellent vapors are: (1) they can kill such a
hardy insect as an American cockroach, and (2) they act like
the vapors of other lipid solvents in not altering appreciably the
optical properties of the nerves they kill.

EFFECTS OF ANILINE

This is an extremely toxic substance for all cells. As such it
is not to be called a selective nerve poison; however, its effects
on the lipids of the nerve sheaths are so striking that we are
treating it separately. When aniline is injected into the tracheae
of mosquito larvae and the nerve cords dissected out in saline im-
mediately, the whole nerve cord (ganglia and connectives) is
diffusely but unusually strongly birefringent (Fig. 17 ; compare
to figures of normal nerve cords in Richards, 1944). If the nerve
cord is now transferred to 10 per cent formalin in saline, bire-
fringent particles appear within a few minutes and rapidly
increase in size and brilliance (Fig. 18). These irregular par-
ticles are found "between the cells and fibers throughout the
ganglia and connectives. They dissolve and disappear within
a few seconds in 95 per cent ethyl alcohol. Birefringent particles
are not found if the nerve cord is left in saline. Untreated nerve
cords in formol-saline retain their normal birefringence. It fol-
lows that aniline does not directly produce these particles but
that it affects the sheath lipids in such a manner that the particles
are produced by the fixation process.9

Similar but less striking effects (smaller particles) are obtained
by the injection of aniline into the tracheae of cockroaches and
subsequently removing the nerve cords into formol-saline.

Serial sections of mosquito larvae treated with aniline, and
fixed first in formol-saline and then in the alcohol-acetic acid
mixture, show many holes (“vacuoles”) in positions comparable
to those occupied by the birefringent particles (Fig. 10). The
presumption is that these holes and also the extensive cell separa-

9 Nerve cell lipids may be involved as well as the sheath lipids. What can
be verified optically is that sheath lipids are involved.

326

Journal ;New York Entomological Society

[Vol. LIII

tion result from the production and solution of such particles.
Aniline is destructive to nerve cells in other ways too. Figure
10 shows that the nerve cell bodies are somewhat shrunken and
stain heavily and nearly homogeneously.

Birefringent particles can be produced in insect nerve cords
by several other means. Immersion of normal nerve cords in
lower concentrations of ethyl alcohol (30-50 per cent) results
in the production of birefringent particles which may appear
similar to the above (Richards, 1943). Autolytic degeneration
of normal nerve cords in saline gives a gradual diminution and
eventually loss of the lipid component of sheath birefringence,
but degeneration of normal nerve cords in 15 per cent glycerine
in saline gives birefringent particles (Richards, 1944, Fig. 16).
Particles that appear somewhat different can be produced by
the action of certain essential oils (see next section). Clearly the
production of birefringent particles can result from the action
of a variety of treatments that seem to have no other obvious
common action. So far as is known, all such particles appear
postmortem.

EFFECTS OF ESSENTIAL OILS

Chemically, essential oils are diverse compounds, commonly
complex mixtures. Included are terpenes, aldehydes, esters,
resins, etc. Some are relatively toxic substances, others less so.
Some are attractants to certain insects ( e.g ., eugenol), others
are repellents (e.g., citronella). All these tend to accumulate
in the insect nervous system (Richards & Weygandt, 1945) where
they usually penetrate as clusters of droplets from the tracheae
(Figs. 3, 4) ; these clusters disperse slowly throughout the nerve
cord and eventually lose their identity in mixing with the degen-
erating nerve substances. Some notes on essential oils have been
included in preceding sections.

In terms of histopathological effects the members of this
diverse group of substances produce various effects. Observa-
tions on gross changes in the nervous system of mosquito larvae
were made during the penetration studies cited above. The list
tested included : oil of thyme, natural oil of citronella, citronella
“tails,” synthetic oil of citronella, eugenol, geraniol, cinnamic

Dec., 1945]

Richards & Cutkomp: Neuropathology

327

alcohol, “Citrola” and several perfume bases called “Petrodars.”
As mentioned above these substances first penetrate as droplets
clustered around the tracheae. Serial sections of mosquito larvae
treated with oil of citronella (Ceylon ST) show clear acidophilic
nuclei in which no chromatin threads can be seen,10 and some
“ vacuoles’ ’ around the tracheae (presumably arising from pre-
cipitation of the tissue elements followed by solution of the
citronella droplets, Fig. 9). Gross pathological changes were
noted only in the cases of eugenol, citronella “ tails’ ’ and oil of
thyme.

Eugenol is striking in that nerve cords fixed in formalin show
many rounded particles or globules (Fig. 15) which are weakly
birefringent (Fig. 16). It seems most likely that the optically
active material in these globules comes from the lipids of the
nerve sheaths. The rounded shape and relatively low amplitude
of birefringence suggest that they are not identical with the par-
ticles resulting from the application of aniline, alcohol or gly-
cerine. These globules in nerve cords treated with eugenol
resemble somewhat the “myelin figures” that can be produced
by the action of water on preparations of phospholipids. Some-
what similar but less striking results were obtained with the
unknown mixture that constitutes the end distillation product
of citronella (“citronella tails”).

Oil of thyme is a destructive material to various types of cells.
The nerve cords of mosquito larvae treated with this material are
opaque and obviously highly pathological. No extensive analysis
of the action of oil of thyme was attempted.

EFFECTS OF “V ALONE” AND ‘ 1 TERTIARY BUTYL
VALONE”

Most of our experiments were performed using “Valone”
(2-isovaleryl-l, 3-indandione), and accordingly this compound
will be discussed first. Individual experiments were performed
on more than 300 adult cockroaches and 200 mosquito larvae in
addition to the accompanying controls. Various types of ap-

10 Similar dissolution of chromatin has been reported for isobutyl undecy-
lene amide by Hartzell & Scudder (1942) and for piperine by Hartzell &
Strong (1944). Several other oils (mustard, croton, cantharidin, colchicine,
etc.) are listed by Haas (1941) as primary nuclear poisons in vertebrates.

328

Journal New York Entomological Society

[Vol. LIII

plications were employed in an attempt to elucidate the erratic
results obtained in the use of this compound in our experiments
on cockroach control. The variations in effectiveness for prac-
tical control programs are not of primary interest to the present
paper where only effects on the nervous system are to be con-
sidered. Intensive study of these practical variations did, how-
ever, lead to the accumulation of an unusually large amount of
pathological data.

The methods of application employed included blood and
tracheal injections of both oil and alkaline-saline solutions, oral
feedings, cuticle applications in sealed wax cells, and in vitro
studies of the effects on extirpated nerve cords.

Cockroaches affected with “Valone” become completely para-
lyzed. The effect from blood and tracheal injections is quite
rapid. With acute doses attainment of complete paralysis may
require only a few minutes or even a few seconds. With smaller
doses the effect is slower. With sublethal doses no effects were
noticed. With tracheal injections the immediate paralytic effect
can be localized to the region of injection and there does not
appear to be any marked stimulation transmitted to the yet un-
affected parts of the nervous system. Electrical stimulation of
exposed nerve cords shows that “Valone” both paralyzes the
affected ganglia and blocks transmission of impulses through to
unaffected areas. With oral and cuticle applications the effect
is slower (may require days) and highly erratic in that many
individuals are never affected. In all cases, irrespective of the
mode of application, once animal are affected they show the same
paralytic symptoms, and in all of our eases cockroaches once
showing paralytic symptoms always died. The variability con-
cerns only the time for the effects to develop and whether or not
the effects ever do develop. In part this probably represents
unexplained variations in penetration or absorption but it also
seems partly due to various uncontrollable degrees of enolization.

Nerve cords of slightly more than 100 cockroaches affected
with “Valone” were examined with polarized light. The results
were entirely consistent irrespective of the mode of application
or the time lapse between application and paralysis. In all cases
the nerve cords or the affected regions thereof showed complete

Dec., 1945]

Richards & Cutkomp: Neuropathology

329

or almost complete loss of the positive component of birefrin-
gence. This is interpreted as indicating destruction of the ultra-
structure responsible for the optical properties of the axis cylin-
ders of the nerve fibers. Correlated with the above is a great
diminution or even loss of the elasticity of the nerve cord, and
also a diminution of the photoelastic properties, but we do not
as yet sufficiently understand the elastic and photoelastic proper-
ties of whole nerve cords to localize or evaluate effects thereon.

It seemed that the effects on the optical properties of the axis
cylinder were always subsequent to irreversible paralysis but
because of the time lag involved in dissection this was checked
by in vitro experiments. Extirpated cockroach nerve cords were
measured with polarized light in saline, and then while observa-
tion was continued a “Valone” solution in alkaline saline was
drawn under the cover glass and the beginning and completion
of the effect timed. Measurable changes, with only one excep-
tion, required 2-4 minutes or longer, and a large effect such as
obtained in dissections after treatment required 10-15 minutes or
longer. This is much slower than the paralytic action under
comparable conditions and so demonstrates that the visible effect
is post-paralysis and presumably postmortem.

The effects of “Valone” differ from those of all other sub-
stances tested not only in the drastic effect on the axis cylinder
but also (and even more strikingly) in the apparent absence of
any demonstrable effect whatsoever on the nerve sheaths.

Serial sections of “Valone’ ’-treated cockroach nerve cords
showed nearly normal histology. Even the axis cylinders of
nerves appeared in reasonably good condition despite the known
extensive degeneration of the optical properties. The only ab-
normal condition found was a moderate amount of chromatin
clumping in a fair percentage of the nerve cells. As will be
pointed out in the section on Suffocation, this indicates an in-
creased cellular acidity presumably brought about by the “Va-
lone ’ ’-injury.

“Tertiary butyl valone” (2-pivalyl-l,3-indandione) was not
studied so intensively as “Valone.” It was used on 75 cock-
roaches and 140 mosquito larvae. The symptoms it produces are
comparable to those found for “Valone,” and so the effects seem
likely to be similar so far as the nervous system is concerned.

330

Journal New York Entomological Society

[Vol. LIII

EFFECTS OF OTHER DESTRUCTIVE AGENTS

In the course of work on penetration a number of very destruc-
tive organic compounds were encountered (Richards & Wey-
gandt, 1945). Octyl alcohol (2-ethyl hexanol) can serve as an
example of the more extreme of these. It accumulates in the
central nervous system when injected into the tracheae of mos-
quito larvae but since it is somewhat soluble in water (0.1 per ,
cent) it can travel readily in the insect’s blood or be used for
in vitro studies. The results are similar whatever method of
application is employed.

The most obvious gross effect of octyl alcohol is to make the
normally transparent brain and nerve cord of mosquito larvae
nearly opaque. Commonly the neural lamella becomes separated
from the nerve cord by a clear space (Fig. 5). Observation of
extirpated nerve cords treated in vitro shows that the neural
lamella is not detectably altered but that the nervous tissue
shrinks away from it. Using high magnifications it is possible
to see that considerable cellular dissolution, as well as shrinkage,
has occurred in these brains. Without any fixation artifacts
being involved, one can see in these whole mounts in saline that
octyl alcohol produces opacity, shrinkage and extensive cellular
dissolution.

In serial sections of fixed brains extensive destruction is appar-
ent. Commonly the peripheral parts of the brain have lost all
appearance of cellular structure and consist of relatively uniform
basophilic granular material (Fig. 11). Nuclei of recognizable
cells are stained a solid dark blue. The fiber tract regions do not
appear to be as badly cytolyzed as with some other agents (Fig.
12). Other tissues are also extensively affected by octyl alcohol,
for instance the cytoplasm (but not nuclei) around muscle fibers
has nearly or quite disappeared and the muscle fibers themselves
show no cross striations.

Obviously the extensive destruction due to octyl alcohol is not
selectively on the nervous system, and the dissolution is so great
that no analysis of the effects is possible.

More or less similar, but usually less destructive, effects were
obtained with a number of other compounds, some of which are
used in insecticides. The list includes methyl diethanolamine,

Dec., 1945]

Richards & Cutkomp: Neuropathology

331

monoisopropanolamine, morpholine, benzyl “ cellosolve, ’ ’ cinna-
mic alcohol, m-cresol acetate, oleic acid, butyl carbitol acetate,
butyraldehyde, oil of thyme, trichlorethane, ethylene dichloride,
and to a lesser extent a number of other organic substances. The
aminated alcohols are especially destructive to tissues.

EFFECTS OF SUFFOCATION AND ACIDITY

In connection with studies on petroleum oils one of us pointed
out that asphyxiation causes a reversible clumping of chromatin
within nuclei (Fig. 6, see also Richards, 1941). This criterion
can, under properly controlled conditions, be used as an index of
suffocation. However, the phenomenon can be produced by other
agencies {e.g., pressure) and so is no specific result of asphyxia
itself. No explanation of the chromatin clumping was offered in
the above paper. More recently it has been called to our atten-
tion that Nassonov analyzed this phenomenon in a paper pub-
lished in 1932 (see also Alexandrov, 1932). He presented strong-
evidence supporting the view that the clumping is due to in-
creased acidity in the nuclei (gut cells of fishes).11

Increased cellular acidity can be obtained in a number of wTays.
Asphyxiation can lower the pH by increasing the C02 concentra-
tion. It is also rather generally accepted that one of the common
effects of injury to cells is increased acidity (Ettisch & Jochims,
1927) . Thus one sees references to “the acid of injury” in physi-
ological literature (see, e.g., Heilbrunn, 1943). Several kinds of
injury can produce clumping of chromatin, e.g., pressure (Buck
& Boche, 1938).

In view of the above, it is not surprising that chromatin clump-
ing has been recently recorded as an effect obtained from the
action of an insecticide (Hartzell & Scudder, 1942) . We too have
found chromatin clumping fairly general but not universal in
pyrethrum-treated nerve cells of cockroaches.11 We have also
obtained chromatin clumping in nerve cells from the action of

11 Chromatin clumping seems to be a general response to asphyxia in cells
with large nuclei. It has been observed and studied in gut cells, gland cells
and nerve cells. It is, however, not found in the small nuclei of the neuroglia
cells of the central nervous system of cockroaches. Since one would expect
asphyxia to lower the pH of all cells it would seem that the chromatin of
some nuclei is unaffected by this degree of acidification (Fig. 6).

332

Journal New York Entomological Society

[VOL. LIII

‘ 1 Thanite ’ ’ and 4 ‘ V alone. ’ ’ Apparently, this effect can be inter-
preted as meaning no more than that the affected cells were in-
jured in such a way (manner unknown) that they become some-
what more acid. This, however, is more than one usually learns
from a histopathological effect, and it is interesting that some
insecticides do this and others do not.12 When this clumping is
produced by asphyxiation it occurs before death and is fully
reversible up to the time of death. It would be very interesting
(and desirable) to find out whether or not it is premortem and.
reversible when produced by insecticide action but this has not
yet been studied.

TOXINS WITHOUT KNOWN HISTOPATHOLOGICAL EFFECTS '

A number of well-known insecticides apparently do not pro-
duce any histologically visible effects in insects. Kruger (1931)
and Hartzell (1934) both failed to find any effects from the appli-
cation of rotenone. Mclndoo (1916) and Hartzell & Wilcoxon
(1933) report no effects from nicotine. The latter workers also
found no histopathological effects in nerve cords of insects killed
with lead arsenate. In our work we were unable to detect any
notable changes in nerve cords treated with formalin, ‘ ‘ non-toxic-’ ’
mineral oils ( e.g Marcol GX), sodium fluoride or “DDT” (2,2-
bis-(p-chlorophenyl)-l,l,l-trichloroethane) . Possible effects from
‘ ‘ DDT ’ ’ have been studied by us in some detail. The nerve cords
of dying cockroaches have normal optical properties and may
even still be capable of transmitting impulses set up by electrical
stimulation. Stained serial sections prepared from cockroaches
dying from the effects of “DDT” showed no clear effects in the
nerve cords or other tissues.13

It has already been noted in a previous section that lipid sol-
vents used as vapors cause no demonstrable histological effects in
insect nerve cords.14 It seems probable that a rather large num-

!2 This criterion, of course, is not applicable to those agents which cause
a dissolution of the chromatin. The work of Haas (1941) is of interest in
suggesting possible differences in terms of nuclear versus cytoplasmic action
of drugs.

13 In addition to nerve cords we examined midgut epithelium, malpighian
tubules, thoracic muscle, “ heart ” and nephrocytes.

14 Shull, Riley & Richardson (1932) concluded, “it is probable that lethal
concentrations of most gaseous compounds do not produce marked visible
changes in the blood ’ ’ of cockroaches.

Dec., 1945]

Richards & Cutkomp: Neuropathology

333

ber of insecticides and other toxins will not give rise to notable
histopathological changes in insects.

DISCUSSION

The term “nerve poison” is encountered rather commonly in
discussions of insecticide action but seldom defined. Actually,
it is a loose term without any very precise meaning. As com-
monly employed in insecticide literature it is used to categorize
the action of compounds that seem to produce symptoms involv-
ing the nervous system. Sometimes there is an implication that
other systems and tissues are less or not at all involved.

Logically at least four degrees or categories of effects on the
nervous system can be recognized : 1 ) A substance may affect all
cells (octyl alcohol, aniline, fixing fluids, etc.), and may cause
either degeneration or fixation of nerve cells non-selectively.
These compounds are not referred to as nerve poisons. 2) A sub-
stance may have a significantly lower threshold for its action on.
nerves. At somewhat higher levels it may affect other tissues.
3 ) A substance may accumulate in the central nervous system and
so reach toxic levels there sooner than in other tissues. This is
rather general for lipid' soluble substances. Since the concentra-
tion of a substance in a bathing solution is not necessarily an
index of the concentration within cells or tissues it is commonly
very difficult to separate truly lower thresholds from accumula-
tion phenomena. In most cases no attempt is made to do so.
Whether or not one is to call substances nerve poisons or selective
nerve poisons when they affect other tissues at slightly higher
concentrations or slightly longer times seems to be a matter of
definition (and accordingly will vary with the purpose for which
the term is used). 4) Lastly, there are some substances such as
atropine which are thought to act specifically on nerve systems.
One could add drugs such as eserine (physostigmine) to this last
category, but eserine not only poisons the specific cholinesterase
in the nervous system but also poisons other esterases from other
tissues. An animal poisoned with eserine apparently dies because
of the anti-cholinesterase action on the nervous system but less
vital esterases of other tissues are also poisoned. With so many
possibilities and gradations the term “nerve poison” cannot have
any specific meaning. By the time the action of a substance is

334

Journal New York Entomological Society

[Vol. LIII

sufficiently understood to state what is meant physiologically by
calling it a nerve poison, a more precise term or statement would
seem possible. Despite all its vagueness, or perhaps because of
its vagueness, the term “nerve poison” is useful at times to
express an action involving the nervous system.

Pyrethrum is not only one of the most standard of insecticides
but also the classical example of an insecticide that is termed a
“nerve poison.” With the above preamble in mind, we would
like to discuss the present state of our knowledge concerning the
neurotoxic action of pyrethrum. The literature is voluminous
and the following citations make no pretense of completeness. In
1924, Juillet, d’Everlange & Ancelin first suggested that pyreth-
rum was a neuromuscular poison because of the paralysis pro-
duced. Saling (1928) added cogent evidence since he could
obtain what he thought were effects on the nervous system but
could detect no effects on the respiratory system or blood. A con-
• siderable series of papers treat or mention the nervous symptoms
of stimulation, narcosis, paralysis and death from pyrethrum,
notably Buchmann (1929), Kruger (1931), Hartzell & Wilcoxon
(1932), O’Kane et al. (1933), Gosswald (1934), Wigglesworth
(1941), Eagleson (1942) and especially Hutzel (1942). Although
the skeletal muscles seem to be stimulated extensively (Kruger,
1931; Klinger, 1936; Hutzel, 1942), the heart is only slowly
affected (Kruger, 1931; Belleuvre, 1938). A number of authors
have recorded and figured extensive histopathological changes in
the central nervous system (Kruger, 1931 ; Hartzell et al., 1932-
1942 ; Klinger, 1936 ; Wigglesworth, 1941 ; and the present paper) .
Kruger and Hartzell also both record histological changes in other
tissues, especially muscles. The recent paper by Sweetman &
Gyrisko (1944) sounds as though they can obtain in firebrats a
progressive localized narcosis involving various tissues but their
data do not yet permit any real analysis of the situation (they
also get the “typical” pyrethrum symptoms). Hurst (1943)
considered the question of cuticle penetration of pyrethrum and
other substances from a physico-chemical viewpoint, and Richards
& Weygandt (1945) showed that pyrethrum, like other lipid sol-
vents or lipid soluble materials, selectively penetrates into and
accumulates in the nervous system. The present paper deals

Dec., 1945]

Richards & Cutkomp: Neuropathology

335

rather extensively with the time relationships between loss of
responsiveness of nerve cords to electrical stimulation, blocking
of nerve transmission, paralysis and the appearance of histopatho-
logical changes. Preliminary oscillographic studies of the effect
of pyrethrum in inducing spontaneous outbursts of impulses and
modifying normal impulses have been presented by Lowenstein
(1942) and Ellis, Thienes & Wiersma (1942). 15 The recent
papers by Roy et al. (1943, 1944) seem to us not convincing and
not significant ; they will not be considered in the following dis-
cussion. The latest review of the action of pyrethrum seems to
be that of Campbell (1942) ; older discussions are given by
Shepard (1939) and Hoskins (1940). A number of papers have
appeared on the effects following injection of pyrethrum into
mammals; the most recent of these is by Leonard (1942) who
agreed with previous authors in reporting nervous stimulation
leading to convulsions and respiratory paralysis but a depression
of muscle contractions in isolated pieces of intestine and an
absence of neuropathology.

These lines of evidence for a neurotoxic action of pyrethrum
may be grouped under five headings: 1) stimulating effect lead-
ing to paralysis and death, 2) histopathological changes, 3)
accumulation and threshold concentrations, 4) early loss of re-
sponsiveness of nerves to electrical stimulation, and 5) effects on
the action potential.

The nervous symptoms and diagnostic effects are thoroughly
discussed by Hutzel (1942) and reviewed by Campbell (1942).
The sequence of activation, convulsions and paralysis, as they
point out, suggest stimulation of peripheral sense organs or
sensory nerves, stimulation of the central nervous system and
then paralysis. These data are quite suggestive and seem cor-
rectly interpreted, but they leave unanswered the question of
possible effects on other tissues and the question of why the heart
is affected so slowly.

The histopathological changes seem to us the poorest line of
evidence. Unlike certain other tissues, a functioning nerve under-
goes no visible changes.16 All one can study is chemical processes

15 This paper deals with the peripheral nerves of crayfish (crustacea) but
the dat’a are nevertheless of entomological interest.

16 With tissues in which one can follow the functioning cytologically, a
stronger ease can be made for histopathology (note for instance how little

336

Journal New York Entomological Society

[Vol. LI II

and electrical phenomena. Using direct electrical stimulation of
nerves and ganglia as a means of verifying the functional status
of the nerve cord, we were unable to produce visible changes with
pyrethrum (or any other insecticide) until after the nerves con-
cerned were dead.17 The degenerative effects and lesions there-
fore seem postmortem and accordingly incapable of being analyzed
at present. Also there is no fixed relation between the degree of
degeneration of the central nervous system and the death of the
experimental insect. More serious is the fact that, except for
possible differences in the time factor, the degeneration of pyreth-
rum-killed nerves follows a course similar to that of nerves de-
generating in saline (present paper) or in the body after suffo-
cation (Richards, 1941). Pathological changes have also been
recorded for muscles (Kruger, etc.). In view of the fact that
the central nervous system degenerates more rapidly than other
insect tissues (Richards, 1941), it seems at least possible that the
degeneration seen in pyrethrum-killed insect nerve cords might
be due to autolysis.18 If this is true, then the recorded histo-
pathology from pyrethrum could be interpreted as indicating no
more than death of the central nervous system prior to that of
other tissues.

The chromatin clumping in nerve nuclei (Hartzell & Scudder,
1942) is a good criterion but seemingly indicates only that the
cells have become somewhat more acid (Fig. 6). It seems likely
that this increased acidity may be due to the pyrethrum but it

reference is made to nefves in Ludford’s review). However, although histo-
pathology is of great use for diagnostic purposes in medicine, it is not viewed
with favor by cellular physiologists or biochemists. It may in some cases
give clues for study but it does not seem likely to explain much of the physi-
ology of toxic action.

17 Data from vertebrates are probably not strictly comparable but we can
note that Schmitt, Bear & Palmer (1941) were unable to affect nerve sheath
structure in vitro with detergents, autolysis, calcium or potassium prior to
the death of the nerves, and Leonard (1942) found no pathology in the brains
of rabbits and mice in convulsions from pyrethrum.

is The difficulty in this connection is to decide what is the il normal’ ’
course of autolysis. Some method must be used to kill the cells. This auto-
matically complicates the analysis. We have found that physical methods
such as heat and cold introduce invalidating errors. Any chemical is suspect.
The two methods recorded here seem to us best although it cannot be claimed
that either is 1 1 normal. 1 ’

Dec., 1945]

Richakds & Cutkomp: Neuropathology

337

would be difficult to prove that an “acid of injury” is really
involved in this case. It is not yet known whether this is pre-
mortem (cell viewpoint) when produced by insecticide action.

Very few data are available on cytopathology of insect nerves
from the action of pyrethrum or any other insecticide. The data
from optical analyses given in the present paper are cytological
(and even based on submicroscopic structure) but cover only the
optical properties of the nerve fibers. The degeneration we found
in these properties was postmortem for the specific cells con-
cerned.17 Except for the phenomenon of chromatin clumping
discussed above, other possible cytological changes in insect nerve
cells (Nissl patterns, mitochondria, Golgi apparatus, etc.) have
been scarcely or not at all studied (see especially review by Lud-
ford, 1942).

Nothing is known about the threshold for the action of pyreth-
rum. MLD determinations can be made accurately but they give
no indication of the quantitative distribution within the insect.
Until actual thresholds for different tissues are determined (or a
specific chemical action to nerves alone demonstrated), it does not
seem possible to state that pyrethrum has a specific effect on
nerves since the material has been shown to penetrate selectively
into and accumulate in the nervous system (Richards & Wey-
gandt, 1945). In any experimental insect one has almost cer-
tainly a greater concentration in the nervous system than in other
tissues. It is conceivable, but not necessarily true, that the appar-
ent selective action is a result of the differences in distribution in
the insect. Some distribution phenomenon such as this may
possibly account also for the slow effect on the insect heart and
its intrinsic nerves.

It was found in our work that pyrethrum paralyzed nerves so
that they would not respond to direct electrical stimulation, and
also blocked the transmission of impulses through affected ganglia.
Yet peripheral movements might proceed for many hours. This
is a direct proof of nerve paralysis. It substantiates the conclu-
sions already drawn by others from the general symptoms and
general paralysis. The paralysis, however, is still open to the
same questions mentioned above ; namely, that other tissues may
be affected and the quicker effect on nerves may merely reflect
the distribution of the pyrethrum.

338 Journal New York Entomological Society [Vol. liii

Preliminary data on action potentials also corroborate the
stimulatory and paralytic effects of pyrethrum. This insecticide
can alter normal action potentials in cockroach nerve cords
(Lowenstein, 1942) and induce spontaneous discharges in cray-
fish peripheral nerves (Ellis, Thienes & Wiersma, 1942).

The one function that a nerve has (transmit an impulse) is
affected by pyrethrum as shown by general body reactions, spon-
taneous discharges, altered discharges, and loss of responsiveness.
Without question, then, pyrethrum has a definite and strong effect
on the insect nervous system, and it seems safe to conclude that
its normal action on an intact insect is to stimulate and then
paralyze. It remains for further work to show how much other
tissues are affected, and in how far the selective nervous action
is due to selective penetration and accumulation in the nervous
system. It seems superfluous to add that as yet we have no idea
as to what specific effect pyrethrum has on nerves or other tissues
(in terms of cellular physiologj^ or biochemistry).

^ # % * *

It may be convenient to some to have a summary of the types
of pathological effects recorded in the present paper.19 The use
of optical analyses of treated nerve fibers has a number of advan-
tages over the usual routine histological procedures (Schmitt &
Bear, 1939; Bichards, 1944). Outstanding among the advan-
tages, the changes in optical properties can be measured accurately
and expressed quantitatively. At least with those substances
studied by us intensively, the method either reveals changes not
detectable by ordinary sectioning methods (since based on sub-
microscopic structures and organization) or is at least more deli-
cate (shows small measurable changes sooner). Fixation and
sectioning, with their attending artifacts, can be avoided. A clear
distinction between effects on the axis cylinder and effects on the
nerve sheaths is usually obtained as a routine result of the several
measurements in different media. The optical method, however,
is applicable only to nerve fibers ; it cannot be used to study the
nerve cell bodies.

19 Of the many treatises on vertebrate, especially human, neuropathology,
we might recommend Spielmeyer (1928), Weil (1933), Speransky (1935),
Page (1937), Young (1942), and Ludford (1942) as particularly useful
references.

Dec., 1945]

Richards & Cutkomp: Neuropathology

339

Nerves may become granular in internal appearance. This is
the first visible change in normal nerves viewed in ordinary light
as they die and begin to degenerate in saline. The granularity
is preceded by a loss of the optical properties of the axis cylinder,
which in turn is preceded by death of the nerve. This granularity
is best seen in intact nerves viewed in saline ; it is commonly not
detectable in stained sections.

Various kinds of larger particles may occur with some mate-
rials. These may be either inside or outside the cells and fibers.
Birefringent particles which are soluble in lipid solvents and
occur outside the fibers most probably originate from the release
of the optically active lipids of the nerve sheaths although one
can not exclude the possibility that some of the lipids might have
been drawn out of the interior of the nerve fibers (Figs. 15-18).
Isotropic particles and globules are of more uncertain origin.

Vacuoles, like particles, may occur either within or between the
cells and fibers, but the largest ones are found outside of the cells
(Figs. 7-10). The holes called “vacuoles” in insect histopathol-
ogy do not or at least do not necessarily represent vacuoles in the
usual cytological sense. They represent the precipitation of tis-
sue constituents around some particle or droplet which is subse-
quently dissolved during preparation of the section. The nature
of those that are found inside cells is unknown, and it does not
seem possible to attribute their presence directly to the action of
the killing agent since they might represent autolytic phenomena
(see Ludford’s review). The holes that are found outside of the
cells and fibers can sometimes be identified with fair certainty.
The possibilities are that they represent breakdown products of
the nerve sheaths produced either by the action of the toxin or by
autolysis, or that they represent actual droplets of the toxin, or
that they represent material withdrawn from or extruded by the
cells. The last possibility is difficult to exclude but examination
of unfixed specimens in saline helps in certain cases. In attempt-
ing to determine from what the holes originate it is desirable to
examine specimens in saline because in several cases {e.g., aniline)
we have been able to demonstrate that the particles and hence
“vacuoles” are fixation phenomena. In stained sections the
“vacuoles” appear simply as holes bearing no label as to their
previous contents.

340

Journal New York Entomological Society

[Yol. liii

There are three other phenomena that seem best studied in
unfixed nerve cords in saline. These are shrinkage, opacity and
chromatin clumping. So much shrinkage is involved in making
sections that this effect should be studied in saline, preferably the
toxin applied after measuring and during observation. Shrink-
age most likely indicates a water loss, but in insecticide studies
the reasons for such a loss do not seem easy to analyze. Opacity
must be examined in unfixed material since fixation produces
opacity, and clearing in mounting media masks it. Opacity may
be used as an index of abnormality, but interpretation of it would
seem difficult. Chromatin clumping can be readily observed
either in transparent tissues in saline (or even in intact trans-
parent animals) or in stained sections. As a reversible physio-
logical phenomenon it should be experimented with either in vivo
or in vitro but as a product of a particular treatment it can be
just as well seen in stained sections. Since it appears to be caused
by changes in the cellular pH, care must be taken to be sure that
the effect is really due to the action of the insecticide. Its possi-
ble reversibility when produced by insecticide action merits
investigation.

Other forms of degeneration are best or sometimes only seen in
sections.19 Chromatolysis or various forms of staining and non-
staining that differ from controls (not necessarily “normals”) is
usually studied in stained sections (Figs. 10 & 12). With cer-
tain dyes this is possible both in vivo and in vitro but such tech-
niques have not yet been applied to insect nerve pathology. An
alteration in staining capacity is the least radical of changes
recorded in previously published literature. Extensive changes
in the staining properties of insect nerve cells and fibers treated
with insecticides are, in our experience, subsequent to irreversible
paralysis of the nerves and so presumably postmortem. It is
quite possible that some changes in stainability and in fine cyto-
logical structure may occur in nerve cells prior to the death of the
cells concerned but this is not easy to study or prove and has not
yet been done.

More extreme forms of degeneration are numerous. One could
apply many terms and describe long series of stages. In general,
two types of extreme degeneration can be distinguished although

Dec., 1945]

Kichards & Cutkomp: Neuropathology

341

they commonly occur together : cell and fiber separation, and cell
and fiber degeneration. Separation is produced by agents which
injure or destroy the nerve sheaths, and may be due directly to
the toxin (Fig. 10) or to autolysis following death caused by the
toxin (Fig. 7). Separation is greatly affected by fixation since
the cells and fibers can shrink independently instead of as a unit.
Separation is always accompanied by more or less degeneration
of the nerves but may be extreme at a time when the nerve cells
and fibers still appear fairly typical. The separation is commonly
most noticeable at the boundary between the central fiber tract
area and the peripheral layer of nerve cell bodies. Probably
several factors are involved : this seems to be the weakest part of
the tissue and also the layer in which most of the tracheae occur
(Fig. 2) and in which “ vacuoles” may be particularly prevalent
(Richards, 1941). The destructive agents which cause radical
degeneration, lytic or otherwise, are recordable but beyond analy-
sis by the methods used in this paper.

As mentioned previously, the data in this and other papers
deal with the effects of single acute doses. No localized action on
particular centers of the insect central nervous system has been
reported. In our work we have not noticed any such local effects
from insecticide action. It seems that the substances studied
affect all nerve cells indiscriminately when applied in the doses
we used. At least one reservation must be made. To date no one
seems to have studied seriously the possible occurrence of localized
effects (more susceptible nerve centers) resulting from prolonged,
repeated chronic doses.

In conclusion on the types of histopathological changes found
in nerves following insecticide action, we can say that a fairly
large number of varieties have already been recorded in the pres-
ent and other papers. There is no reason to think that others
cannot be found. A number of insecticides produce effects which
parallel and so presumably represent autolysis phenomena (py-
rethrum, “Thanite” & petroleum oils) ; certain other insecticides
produce effects more or less distinct from autolysis (“Valone,”
lipid solvents, certain essential oils, etc.). It seems that one could
go on indefinitely performing such experiments and describing in
detail the histological and cytological pictures obtained. The

342

Journal New York Entomological Society

[Vol. Liir

value of such a mass of data seems to us questionable. At least
in the best analyzed case (acute doses of pyrethrum) there is no
fixed relation between the degree of degeneration and death of
the insect. And in no case have we found detectable pathology
from an insecticide prior to the irreversible paralysis (and pre-
sumably death) of the cells concerned. Chromatin clumping is
a possible exception since when it is produced by suffocation it is
premortem and then reversible. Chromatin clumping is a phe-
nomenon that seems to us profitable for further study.

Cytologically the insect nervous system is similar to that of
vertebrates, especially to the non-myelinated fibers such as are
found in the vertebrate autonomic system. The cells are smaller,,
the sheaths thinner, and the connective tissue less, but the basic
structure of the cells and fibers so far as has been determined is
the same (Richards, 1944; Scharrer & Scharrer, 1944-1945). The
chemical or at least lipid components are comparable (Patterson,
Dumm & Richards, 1945). Also insect nerves are cholinergic,
i.e., have an acetylcholine-cholinesterase mechanism, but there are
at least qualitative differences in the cholinesterases of insect and
vertebrate nerves (see Richards & Cutkomp, 1945).

Accordingly, we may conclude that the insect nervous system
is similar in many ways to the vertebrate nervous system, particu-
larly to the vertebrate autonomic system (Jordan, 1928 ; the
Roeders, 1939; the Scharrers, 1944-1945), but that some differ-
ences seem to exist (cholinesterase). A more intensive study of
the insect nervous system and its relation to insecticide action is
needed. It seems to the present authors that histopathology has
little if anything to offer this further analysis. The possible use
of cytopathology of insect nerves has not been sufficiently studied
for evaluation but the slight amount of data available is not
encouraging. Studies of the reactions of insects to drugs (includ-
ing some insecticides), of nerve enzymes and biochemical proc-
esses, and of electrical phenomena are the methods that seem
promising for studying the action of neurotoxic insecticides.

SUMMARY

1. Data are presented on the histopathological effects caused
by acute doses of various materials including pyrethrum, “Tha-

Dec., 1945]

Richards & Cutkomp: Neuropathology

343

nite, ’ ’ petroleum oils, venoms, triorthocresyl phosphate, lipid sol-
vents, insect repellents, aniline, essential oils, “Valone, ” acidity,
and a number of highly destructive compounds of which octyl
alcohol is taken as an example. Cockroaches and mosquito larvae
were used as test animals. No visible elfects were obtained with
“DDT” and certain other compounds. Optical analyses, routine
stained sections and in vitro analyses were used ; electrical stimu-
lation of the nerve cords was employed to determine the physio-
logical state of the nerves being studied.

2. The various types of pathological pictures can be described
by the use of terms such as decrease or loss of one or more of the
components of the optical properties, granularity, chromatin
clumping in the nuclei, and various stages of cell dissolution
ranging from chromatolysis to particle production, “vacuoliza-
tion,” opacity, shrinkage, and extensive cell and fiber separation
and disintegration.

These categories are not sharply defined or mutually exclusive.
A particular toxin may produce a more or less characteristic pic-
ture under a particular set of conditions but at least with pyreth-
rum there is no fixed relationship between the death of the insect
and the degree of degeneration of the nervous system.

3. The physiological and histological effects of pyrethrum are
considered in some detail. It is concluded that previous workers
are correct in calling this insecticide a “nerve poison” but that
the histological (and what is known for cytological) changes are
similar to those produced by autolysis and may not be directly
caused by the pyrethrum. Pyrethrum penetrates selectively into
and accumulates in the nervous system of insects. Its threshold
for nerves, its possible thresholds for and effects on other tissues,
the relation between these thresholds, how it kills and whether
or not it has a specific effect on nerves, are points not yet covered
by the existing literature.

4. Lipid solvents used as fluids in considerable quantity remove
the lipid component of the sheath birefringence. Used as vapors,
however, they kill without producing any visible effect. Accord-
ingly the visible effects that can be produced cannot be the cause
of death from these substances.

5. Insect repellents have a visible effect on nerves comparable

344

Journal New York Entomological Society

[Vol. LIII

to that of lipid solvents. Used as fluids they abolish the lipid
component of birefringence but used as vapors they kill without
visibly altering the structure of the central nervous system.

6. In all cases studied, nerves were paralyzed and presumably
dead prior to the appearance of any abnormalities or lesions with
the possible exception of chromatin clumping. Accordingly all
histopathological pictures recorded for insect nerves, with the
possible exception of chromatin clumping, are to be classed as
“postmortem,” and their further analysis is of questionable
value. The same statement may be made for the little that is
known about insect nerve cytopathology. It seems to us that
histopathology of insect nerves may at times give some slight help
in localizing the action of certain insecticides but that it is at best
a crude and likely to be misleading measure of physiological
effects in insects. The reservation should be repeated that exist-
ing data refer to the effects of acute doses; the possibility of
obtaining more localized effects on particular centers by pro-
longed chronic dosing has not been sufficiently investigated as yet.

7. The term “nerve poison” is a rather vague concept. It is a
convenient term but not a specific one. It can mean either that
the substance has a lower threshold in nerves, or accumulates
there more rapidly or in greater amounts, or it may mean a truly
specific action. However, the analysis of insecticide action on
nerves requires more specific techniques than histopathology and
the demonstration of paralysis.

8. The suggestion (Richards, 1943) that lysolecithin formed
by the breakdown of nerve sheath lipids may be concerned in
insect paralysis, is discredited. Cockroaches are not visibly af-
fected by the injection of maximal quantities of lysolecithin, and
the nerve sheaths are not detectably affected by cobra venom prior
to paralysis. Certain other substances highly toxic to vertebrates
have little or no effect on cockroaches (histamine, curare).

SUPPLEMENTARY NOTE

While this paper was in press Hartzell (1945) published a
paper on the histopathological effects of several compounds used
in insecticides, including “DDT.” He stresses the selective ac-
tion of certain substances on nuclei, nuclear membranes, nerve
cell cytoplasm, nerve fibers or intercellular spaces, and notes that

Dec., 1945]

Richards & Cutkomp: Neuropathology

345

these differences connote something different in the way of action
of the substances concerned. He further suggests that the syner-
gistic effect of certain substances may be due to the activator
attacking one cellular component, the insecticide another. The
paper is subject to the same criticisms pointed out in the above
text, and it is difficult to attempt interpretation on the basis of
such data.

Hartzell records “relatively slight” pathological effects from
“DDT. ” He has no real knowledge of whether or not the nerves
concerned were living or dead but since the time interval was
short (and “DDT” effects are relatively slow) they may well
have been still functional. However, at least in cockroaches,
more variation is seen in long series of controls (or normals)
than Hartzell shows for the differences between normal and DDT-
killed houseflies. Possibly Hartzell did obtain a slight effect
(assuming that histology of the central nervous system is less
variable in houseflies than in cockroaches) but the slight effects
recorded agree with the usual autolysis picture. We can only
repeat that in our “DDT” experiments dying cockroaches which
had responsive nerve cords showed no effects that could not be
ascribed to normal variation and matched by control preparations.

LITERATURE CITED

Alexandrov, W. 1932. Tiber die Bedeutung der oxydo-reduktiven Beding-
urigen fur die vitale F&rbung, mit besonderer Beriicksichtigung der
Kernfarbung in lebendigen Zellen (Chironomus-Larven und Daphnia
pulex). Protoplasma, 17: 161-217.

Beck, B. F. 1935. Bee Venom Therapy. Appleton-Century Co., New York.
Belleuvre, G. 1938. Action physiologique des pyrethrins sur les inverte-
brates. Ann. Physiol. & Physicochim. Biol., 14: 717-733.

Bodian, D. 1942. Cytological aspects of synaptic function. Physiol. Rev.,
22: 146-169.

Buchmann, W. 1929. Untersuchungen iiber die physiologische Wirkung
von Pyrethrum-Insektenpulver auf Fliegenlarven. Zts. Desinf. u.
Gesundh., 21 : 72-76.

Buck, J. B. & Boche, R. D. 1938. Some properties of living chromosomes.
Collecting Net, 13 : 201-203.

Campbell, F. L. 1942. Pyrethrum vs. roaches. Soap, 18(5): 90-93, 103—
104, 18 (6>: 119, 121, 123, 125, 127.

De, S. S. 1941. Crystalline haemolysin from cobra (Naja naja) venom.
Sci. & Culture, 6: 675-676.

Eagleson, C. 1942. Effect of temperature on recovery houseflies. Soap,
18(6) : 115-117, 141.

346

Journal New York Entomological Society

[Vol. L1II

Ellis, C. H., C. H. Thienes & C. A. G. Wiersma. 1942. The influence of
certain drugs on the crustacean nerve-muscle system. Biol. Bull.,
83: 334-352.

Ettisch, G. & J. Jochims. 1927. Dunkelfelduntersuchungen am iiber-
lebenden Nerven. I. Die Wirkung von Elektrolyten. II. Die
Wirkung von Nichtelektrolyten. Pfliiger’s Arch., 215: 519-544,
675-682.

Feldberg, W. 1940. The action of bee venom, cobra venom and lysolecithin
on the adrenal medulla. Jour. Physiol., 99: 104-118.

Frear, D. E. H. 1942. Chemistry of Insecticides and Fungicides. Van
Nostrand. New York.

Ghosh, B. N. 1940. Enzymes in snake venom. Osterr. Chem.-Ztg., 43 :
158-163.

Gosswald, K. 1934. Die Wirkung des Kontaktgiftes Pyrethrum auf
Forstschadlinge unter dem Einfluss der physiologischen Disposition
der Schadlinge und der Einwirkung von okologischen Aussenfak-
toren. Zts. angew. Ent., 20: 489-530.

Haas, H. T. A. 1941. liber die Beeinflussung des Histaminegehalts der
Haut durch Reizstoffe. Naunyn-Schmiedeberg ’s Archiv f. Exp.
Path. & Pharm., 197 : 161-186, 285.

Hanstrom, B. 1928. Vergleichende Anatomie des Nervensystems der wir-
bellosen Tiere, unter Beriicksichtigung seiner Funktion. Springer,
Berlin.

Hartzell, A. 1934. Histopathology of insect nerve lesions caused by in-
secticides. Contrib. Boyce Thomp. Inst., 6: 211—223.

. 1935. Histopathology of nerve lesions of cicada after paralysis by

the killer-wasp. Contrib. Boyce Thomp. Inst., 7: 421-425.

■. 1945. Histological effects of certain sprays and activators on the

nerves and muscles of the housefly. Contrib. Boyce Thompson Inst.,
13: 443-454.

& H. I. Scudder. 1942. Histological effects of pyrethrum and an

activator on the central nervous system of the housefly. Jour.
Econ. Ent., 35: 428-433.

& M. Strong. 1944. Histological effects of piperine on the central

nervous system of the housefly. Contrib. Boyce Thomp. Inst., 13 :
253-257.

& F. Wilcoxon. 1932. Some factors affecting the efficiency of con-
tact insecticides. II. Chemical and toxicological studies of pyreth-
rum. Contrib. Boyce Thomp. Inst., 4: 107-117.

& . 1933. Experiments on the mode of action of pyrethrum

and its effects on insect tissues. Ve Congr. Int. d’Ent., p. 289-294.

Heilbrunn, L. Y. 1943. An Outline of General Physiology. 2nd ed.
Saunders, Philadelphia.

Hoskins, W. M. 1940. Recent contributions of insect physiology to insect
toxicology and control. Hilgardia, 13 : 307-386.

Dec., 1945]

Richards & Cutkomp: Neuropathology

347

Hurst, H. 1943. Principles of insecticidal action as a guide to drug reac-
tivity-phase distribution relationships. Trans. Faraday Soc., 39 :
390-412.

Hutzel, J. M. 1942. The activating effect of pyrethrum upon the German
cockroach. Jour. Econ. Ent., 35: 929-933.

•. 1942. Action of pyrethrum upon the German cockroach. Jour.

Econ. Ent., 35: 933-937.

Jordan, H. J. 1928. Die Theorie der Funktion des zentralen Nervensys-
tems bei niederen Tieren. Tijdschr. nederl. Dierk. Yereenig., 3rd
ser., 1: 51-54.

Juillet, A., M. d’Everlande & M. Ancelin.’ 1924. Le Pyrethre insecti-
cide de Dalmatie. Pyrethrum cinerarifolium Trev. Chrysanthemum
cinerarifolium Yis. iEtude botanique, chimique et agricole. Roume-
gous et Dehan, Montpellier.

King, E. J. & M. Dolan. 1933. The enzymic hydrolysis of phosphatides.
2. Lysolecithin, Biochem. Jour., 27 : 403-409.

Klinger, H. 19.36. Die Insektizidwirkung von Pyrethrum und Derrisgiften
und ihre Abhangigkeit vom insekten Korper. Arb. Physiol, u.
angew. Ent., 3 : 49-69, 115-151.

Kruger, F. 1931. Untersuchungen uber die Giftwirkung von dalmatischen
Insektenpulver auf die Larven von Corethra plumieornis. Zts.
angew. Ent., 18: 344-353.

Leonard, C. S. 1942. Effects of pyrethrins on certain mammals. Jour.
Econ. Ent., 35: 261-264.

Levene, P. A. & I. P. Rolf. 1923. Lysolecithins and lysocephalins. Jour.
Biol. Chem., 55 : 743-749.

, & H. S. Simms. 1924. Lysolecithins and lysocephalins. 2.

Isolation and properties of lysolecithin and lysocephalins. Jour.
Biol. Chem., 58 : 859-871.

Lillie, R. D. & M. I. Smith. 1932. The histo-pathology of some neurotoxic
phenol esters. U. S. Nat. Inst. Health Bull., 160: 54-69.

Lowenstein, O. 1942. A method of physiological assay of pyrethrum
extracts. Nature, 150: 760-762.

Ludford, R. J. 1942. Pathological aspects of cytology. In Bourne, G.
Cytology and Cell Physiology. Clarenden Press, Oxford.

Macht, D. I. 1941. Recent developments in the pharmacology and thera-
peutics of cobra venom. Med. Rec., 153: 369-375.

McIndoo, N. E. 1916. Effects of nicotine as an insecticide. Jour. Agric.
Res., 7: 89-121.

Nassonov, D. 1932. Tiber die Ursachen der reversiblen Gelatinierung des
Zellkerns. Protoplasma, 15 : 239-267.

O’Kane, W. C., G. L. Walker, H. G. Guy & O. J. Smith. 1933. Reactions
of certain insects to controlled applications of various concentrated
chemicals. N. Hamp. Agr. Exp. Sta., Tech. Bull., 54: 1-20.

Page, I. H. 1937. Chemistry of the Brain. Thomas, Baltimore.

348

Journal New York Entomological Society

[Vol. LIII

Patterson, E. K., M. E. Dumm & A. G. Richards, Jr. 1945. Lipids in the
central nervous system of the honey bee. Arch. Biochem., 7 : 201-
210.

Richards, A. G., Jr. 1941. Differentiation between toxic and suffocating
effects of petroleum oils on larvae of the house mosquito ( Culex
pipiens). Trans. Amer. Ent. Soc., 67 : 161—196.

. 1943. Lipid nerve sheaths in insects and their probable relation to

insecticide action. Jour. N. Y. Ent. Soc., 51: 55-69.

. 1944. The structure of living insect nerves and nerve sheaths as

deduced from the optical properties. Jour. N. Y. Ent. Soc., 52:
285-310.

& L. K. Cutkomp. 1945. The cholinesterase of insect nerves.

Jour. Cell. & Comp. Phys. (in press).

& J. L. Weygandt. 1945. The selective penetration of fat solvents

into the nervous system of mosquito larvae. Jour. N. Y. Ent. Soc.,
53: 153-165.

Roeder, K. D. 1939. The action of certain drugs on the insect central
nervous system. Biol. Bull., 76: 183-189.

& S. Roeder. 1939. Electrical activity in the isolated ventral nerve

cord of the cockroach. I. The action of pilocarpine, nicotine,
eserine and acetylcholine. Jour. Cell. Comp. Phys., 14: 1-9.

Roy, D. N. & S. M. Ghosh. 1944. The mechanism of action of a contact
insecticide. Bull. Ent. Res., 35: 161-170.

, & R. N. Chopra. 1943. The mode of action of pyrethrum

on the cockroach, Periplaneta americana. Ann. Appl. Biol., 30 :
42-47.

Saling, T. 1928. tiber das wirksame Prinzip von Pyrethrum-Insekten-
pulvern und eine neue biologische Methodik ihrer Wertbestimmung.
Zts. Desinf . u. Gesundh., 20 : 33-42.

Scharrer, B. 1939. The differentiation between neuroglia and connective
tissue sheath in the cockroach ( Periplaneta americana). Jour.
Comp. Neurol., 70 : 77-88.

& E. Scharrer. 1944. Neurosecretion. 6. A comparison between

the intercerebralis-cardiacum-allatum system of the insects and the
hypothalamo-hypophyseal system of the vertebrates. Biol. Bull.,
87: 242-251.

& . 1945. Neurosecretion. Physiol. Rev., 25: 171-181.

Schmitt, F. O. & R. S. Bear. 1939. The ultrastructure of the nerve axon
sheath. Biol. Rev., 14: 27-50.

, & K. J. Palmer. 1941. X-ray diffraction studies on the

structure of the nerve myelin sheath. Jour. Cell. Comp. Phys., 18:
31-42.

Shepard, H. H. 1939. The Chemistry and Toxicology of Insecticides.
Burgess, Minneapolis.

Shull, W. E., M. K. Riley & C. H. Richardson. 1932. Some effects of
certain toxic gases on the blood of the cockroach, Periplaneta ori-
entalis. Jour. Econ. Ent., 25: 1070-1072.

Dec., 1945]

Richards & Cutkomp: Neuropathology

349

Speransky, A. D. 1935. A Basis for the Theory of Medicine. Interna-
tional Publ., New York.

Spielmeyer, W. 1928. Yasomotorisch tropische Veranderungen bei zere-
braler Arteriosklerosis. Monatschr. f . Psychiat. u. Neurol., 68 :
605-620.

Sweetman, H. L. & G. G. Gyrisko. 1944. Latent injury from pyrethrum
and rotenone dusts. Jour. Econ. Ent., 37 : 746—749.

Weil, A. 1933. A Textbook of Neuropathology. Lea & Febiger, Phila-
delphia.

Wigglesworth, V. B. 1941. The effect of pyrethrum on the spiracular
mechanism of insects. Proc. Roy. Ent. Soc., London, 16A: 11-14.

Young, J. Z. 1942. The functional repair of nervous tissue. Physiol. Rev.,
22: 318-374.

350

Journal New York Entomological Society

[Vol. LIII

Figure

Figure

Figure

Figure

Figure

Figure

PLATE IY

1. Whole mount of supra- and sub-oesophageal ganglia of mosquito

larva showing the distribution of tracheae. The tracheae are
filled with Eugenol saturated with Black Sudan B. Magnifi-
cation 55 x.

2. Obliquely tangential section of prothoracic ganglion of an adult

cockroach showing the distribution of tracheae principally in
the boundary region between the fiber-tracts and cell-bodies.
One half is cut tangentially through this boundary and shows
several tracheae in longitudinal section. Magnification 45 x.

3. Whole mount of supra- and sub-cesophageal ganglia of mosquito

larva showing spotty penetration of stained oil of citronella
from the filled tracheae. Magnification too low to show the
droplet-type of penetration (see figure 4). Magnification 45 x.

4. Portion of whole mount of supracesophageal ganglion of mosquito

larva showing the droplet-type of penetration. Tracheae incom-
pletely filled with stained 1 1 Citrola. ’ ’ Magnification 220 x.

5. Whole mount of supracesophageal ganglion of mosquito larva.

Tracheal injection of octyl alcohol has caused the neural
lamella to separate from the brain (shrinkage of the nervous
tissue). Magnification 55 x.

6. Portion of section of a thoracic ganglion from a suffocated cock-

roach. Shows extreme chromatin clumping in most but not
all of the nerve cells, whereas the nuclei of the neuroglia cells
(vertical row on right side) are normal. Magnification 470 X.

(Jour. N. Y. Ent. Soc.), Vol. LIII

(Plate IV)

352

Journal New York Entomological Society

[Yol. liii

Figure 7.

Figure 8.
Figure 9.

Figure 10.
Figure 11.

Figure 12.

PLATE V

Longitudinal section of fourth abdominal ganglion of mosquito
larva dying from the effects of a tracheal injection of iso-
borneol thiocyanoacetate (active principal of 1 i Thanite? ’■)»
Note “ vacuolated ’ ’ fiber tract region. Compare figure 8.
Magnification 470 x.

Longitudinal section (slightly oblique) of fourth abdominal
ganglion of mosquito larva with tracheal injection of a “ non-
toxic ■ * mineral oil (“Marcol GXM). Control for figure 7.
Magnification 470 x.

Section of supracesophageal ganglion of mosquito larva. Tra-
cheae filled with stained oil of citronella. The arrow points to
a cross-section of a trachea. The holes adjacent to this tra-
chea presumably represent droplets of citronella. Magnifi-
cation 470 x.

Section of subcesophageal ganglion of a mosquito larva killed by
a tracheal injection of aniline. Magnification 435 X.

Portion of a section through the cell-body region of supra-
oesophageal ganglion of a mosquito larva killed by a tracheal
injection of octyl alcohol. Note indistinctness of cells in
central part and disintegration to granular layer in peripheral
part. Magnification 400 x.

Section of suboesophageal ganglion of a mosquito larva killed
by a tracheal injection of octyl alcohol. Magnification 470 x.

(Jour. N. Y. Ent. Soc.), Vol. LIII

(Plate Y)

354

Journal New York Entomological Society

[vol. Lin

Figure 13.

Figure 14.

Figure 15.

Figure 16.

Figure 17.

PLATE YI

Portion of a whole mount of supraoesophageal ganglion of mos-
quito larva showing penetration of stained droplets of propy-
lene glycol monolaurate from injected tracheae. Magnification
470 x.

Whole mount of two thoracic ganglia of a mosquito larva show-
ing the diffuse penetration of stained chloroform. This is an
early stage picked to show gradient from the fluid-filled tra-
cheae into the nervous tissue. Magnification 220 x.

Whole mount of abdominal connective of a mosquito larva killed
by a tracheal injection of Eugenol. Photograph in ordinary
light to show rounded particles. Magnification 300 x.

Same as seen in polarized light between crossed Nicols. The
rounded particles are very faintly birefringent. Magnifica-
tion 300 x.

Whole mount in saline of abdominal nerve cord of a mosquito
larva following a tracheal injection of aniline. Photographed
in polarized light between crossed Nicols. Compare the rela-
tively strong, diffuse birefringence with normal nerve cord
figured by Richards (1944). Magnification 50 x.

Same after fixation for 50 minutes in f ormol-saline. The strongly
birefringent particles are of irregular shapes. Magnification
50 x.

Figure 18.

(Jour. N. Y. Ent. Soc.), Vol. LIII

(Plate VI)

INDEX TO NAMES OF INSECTS AND PLANTS
IN VOLUME LIII

Generic names begin with capital letters. New genera, subgenera, species,
varieties and new names are printed in italics.

.ZEdes

aegypti, 313

Africa, Psychodidae, check-list of, 247
Agonopteryx

lecontella, 222
Agromyza

baptisia, 223
Anteos

maerula, 136
Apion

rostrum, 221
Araeocerus

fasciculatus, 220
Asia, Psychodidae of, 211
Australia, Psychodidae of, 2ll

Baptisia, insects of, 219
Boyd, John, 74
Brachypremna
geijskesi, 49

Butterflies, notes on Mexican, 31
By-laws of Society, 83

Central America, Psychodidae from,
21

Certificate of incorporation of So-
ciety, 79

Check-list, African Psychodidae, 247

Chilopoda, 63

Cingilia

catenaria, 223

Coleoptera, supplement to N. Y.
State list, 91

Collections, Lepidoptera, 62
Conotrachelus

crimaceus, 221
Culex spp., 299
Culex

apicalis, 293

nigripalpus, 297
pipiens, 313

Culicidae, seasonal variations in, 293

Davis, W. T., obituary, 127

Ennomid pupae, 178
key to group, 180

Entomogenous fungi, isolation of, 238
Entomological, early, ideas and prac-
tices in America, 301
Entomologists, longevity of, 231
Entomology, early manuscripts on,
258

Erioptera

turrialbcs, 290

Europe, Psychodidae of, 117

Pat solvents, penetration of into
nervous system of mosquito larvae,
153

Pelt, E. Porter, 169
Fireflies, 302

Formicidae, impaternate females in,
263

Fungi, isolation of entomogenous,
238

Gall, on Trichostema, 259
Gelchossa

heidmenni, 223
Geophilus

duponti, 63
Gnophomyia

apioularis, 59
curraniana, 285
lustralis, 289
pulvinaris, 286
subapicularis, 60

357

358

Journal New York Entomological Society

[Vol. LIII

Gnorimoschema
operculella, 68
Graph olitha

tristrigana, 222

Hadronema

militaris, 223
Helius

phasmatis, 56
scliildi, 54

Housefly, enemies of, 145

Indigo insects, 219
Insect

enemies of house fly, 145
food habit ratios, 167
Insects

ancient remedies involving, 246
biological control of, 307
and disease, 303
electroplating of, 307
of indigo, 219

injection of sulfur in trees to
destroy, 304

killing of collected specimens,
306

neuropathology in, 313

Japanese beetle, distribution and
abundance, 1
Jassus

olitorius, 224

Key, to Ennomid pupal groups, 180
Kupara, 310

albipeda, 311

Leiobunnum

gordoni, 243

Lepidoptera collections, 62
Limonia

cochabambce, 53
ingenua, 52
Lissorhoptrus
simplex, 221

Lloyd-Cornell reservation, insect food
habit ratios of, 167
Longevity of entomologists, 231

Macrosiphum

solanifolii, 223
Mecistocephalus

mecistocephalus, 64
Members, honorary, of Society, 172
Mexican butterflies, notes on, 31
Mompha

sexnotella, 259
Monotarsobius
holstu, 66

Mosquito larvae, penetration of fat
solvents into nervous system of, 153
Mosquitoes, seasonal variation in, 293
Musca

domestica, 145

Nemastoma

pallidimaculosa, 241
Neognophomyia
ineliconicE, 57
schildi, 288

Nervous system of mosquito larvae,
penetration of fat solvents into, 153
Neuropathology in insects, 313

Otocryptops

rubiginosus, 65

Oviposition, mechanism of, Phaenicia
sericata, 227
Oxy discus

destitutus, 56

Pachybrachys
luridus, 221
trinotatus, 221
Papaipema

baptisia, 223

dos Passos, Viola Harriet, 47
Pericoma

abreui, 262
unicolor, 262
Periplaneta

americana, 313
Phaenicia

sericata, 227

Plialangida, from United States, 239
Popillia

japonica, 1

Dec., 1945]

Index

359

Portrait, W. T. Davis, 127
Potato tuber moth, 68
Psychoda

atreseta, 310
brevitarsa, 310
eburna, 309
maculosa, 309
Psychodidse
check-list

of African, 247
for Asia and Australia, 211
of Europe, list of, 117
list of South and Central Amer-
ican, 21

Eiley, C. V., pirated work of, 226

Scolopendra

japonica, 65
subspinipes, 64

South America, Psychodidse of, 21
Stingless bees, wax of, 137

Tanypremna

elegantior, 51
Taracus

malkini, 242

Tetralopha

baptisiella, 222
Teucholabia

circumscripta, 279
diplaca, 281
platy phallus, 282
serrulifera, 284
turrialbensis, 285
Thanaos

brizo, 222
juvenalis, 222
Thereuonema

tuberculata, 67
Tholeria

reversalis, 222
Thysamia

zenobia, 88

Tipulidse, new species, 49, 279
Trichostema, pedicel gall on, 259
Tychius

sordidus, 221

Watson, Frank E., 88
Wax of stingless bees, 137
Wespus

arkansasensis, 239

The

New York Entomological Society

Organized June 29, 1892 — Incorporated February 25, 1893
Reincorporated February 17, 1943

The meetings of the Society are held on the first and third Tuesday of each month
(except June, July, August and September) at 8 p. m., in the American Museum of
Natural History, 79th St., & Central Park W., New York 24, N. Y.

Annual dues for Active Members, $3.00; including subscription to the Journal, $4.50.
Members of the Society will please remit their annual dues, payable in January, to
the treasurer.

A set of the Journal, volumes I to L inclusive, unbound, $90.00.

Officers for the Year 1945

President, GEORGE G. BECKER U. S. Bureau Ent. & Plant Quar., Hoboken, N. J.

Vice-President , DR. STANLEY W. BROMLEY Bartlett Tree Research Lab.,

Stamford, Conn.

Secretary, JOHN C. PALLISTER . American Museum of Natural History

Assistant Secretary, LINA SORDILLO American Museum of Natural History

Treasurer, DR. WILLIS J. GERTSCH American Museum of Natural History

Assistant Treasurer, MARGARET L. GUY American Museum of Natural History

Editor, HARRY B. WEISS Highland Park, N. J.

TRUSTEES

Georqe G. Becker Edwin W. Teale Dr. S. W. Bromley

Wm. P. Comstock Ernest L. Bell

PUBLICATION COMMITTEE

Harry B. Weiss T. C. Schneirla John D. Sherman, Jr.

PROGRAM COMMITTEE

Dr. R. A. Oakley Wm. P. Comstock E. I. Huntington

FIELD COMMITTEE

Lucy W. Clausen Chris. E. Olsen

DELEGATE TO TEE N. Y. ACADEMY OF SCIENCES
Wm. P. Comstock

JOURNAL

of the

NEW YORK ENTOMOLOGICAL SOCIETY

Published quarterly by the Society at N. Queen St., and Mc-
Govern Ave., Lancaster, Pa. All communications relating to
manuscript for the Journal should be sent to the Editor, Harry B.
Weiss, 19 N. 7th Ave., Highland Park, New Jersey; all subscrip-
tions to the Treasurer, Dr. Willis J. Gertsch, American Museum
of Natural History, New York 24, N. Y. Orders, for back issues
should be sent to the Treasurer, Willis J. Gertsch, American
Museum of Natural History, New York 24, N. Y. The society has
a complete file of back issues in stock. The Society will not be
responsible for lost Journals if not notified immediately of change
of address. We do not exchange publications.

Terms for subscription, $3.00 per year, strictly in advance.

Please make all checks , money-orders , or drafts payable to
New York Entomological Society.

Twenty-five reprints without covers are furnished free to
authors. Additional copies may be purchased at the following
rates :

4 pp. 8 pp. 12 pp. 16 pp. 24 pp. 32 pp.

25 copies $2.70 $5.85 $6.15 $6.15 $10.00 $10.65

50 copies 2.90 6.35 6.75 6.75 10.85 11.55

Additional 100’s 75 1.75 2.30 2.30 3.35 3.60

Covers 50 copies, $3.60 ; additional 100 5s, $2.15.

Half-tone prints 1% cents for each half-tone print.

Authors whose papers are illustrated with text figures or
full page plates will be required to supply the electroplates or
pay the cost of making the same by the Journal and also to
pay the cost of printing full page plates on coated paper, when
advisable.

\