■V,

1-

p",

JOURNAL

OF THE

NEW YORK

ENTOMOLOGICAL SOCIETY

BtmUh t0 jEntomology in C^^nrral

VOLUME XLVIII, 1940

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

New York, N. Y.

THE SCIENCE PRESS PRINTING COMPANY
LANCASTER, PENNSYLVANIA

CONTENTS OF VOLUME XLVIII

Page

Alexander, Charles P.

Records and Descriptions of Neotropical Crane-Flies

(Tipulidse: Diptera), XI 105

Barber, G. W.

See Phillips, W. J.

Bell, E. L.

Some Synonymy in Neotropical Hesperiidse (Lepi-

doptera) 116

Bequaert, Joseph C.

An Introductory Study of Polistes in the United States
and Canada with Descriptions of Some New North
and South American Forms (Hymenoptera : Ves-

pidae) 1

Black welder, Richard E.

Some Aspects of Modern Taxonomy 245

Breland, Osmond P.

Some Parasites and Hyperparasites of the Cecropia

Moth 259

Comstock, William P.

Butterflies of New Jersey 47

CuMLEY, Russell W.

Comparison of Serologic and Taxonomic Relationships

of Drosophila Species 265

Evans, W. T.

The Type of the Genus Pyrrhopyge (Lepidoptera, Hes-

peridie) 405

Funkhouser, W. D.

' New Peruvian Membracidae 275

Gaul, Albro Tilton

A Note on Rearing the Brood of Polistes f uscatus Fa-

bricius (Hymenoptera-Vespidae) 391

Hallock, Harold C.

The Sarcophaginae and Their Relatives in New York,

Part I 127

iii

Page

The Sarcophaginae and Their Relatives in New York,

Part II 201

Hatch, Melville H.

Observations on Silphinse with a Note on Intraspecific

Variations and Their Designations 223

Huckett, H. C.

The North American Species of the Genera Lencophora
Robinean-Desvoidy and Proboscimyia (Muscidae-

Diptera) 335

Klots, Alexander B.

A New Brenthis from Alaska (Lepidoptera, Nympha-

lid^) 413

Linsley, E. Gorton

A Reclassification of the Tribe Obriini of LeConte (Co-

leoptera-Cerambyeidae) 367

Phillips, W. J. and G. B. Barber

Seasonal Abundance of Eggs of the Corn Ear Worm

Moth in Virginia 305

Pratt, Harry D.

Studies on the Ichnenmonidae of New England (Hy-

menoptera) 155

Rex, Edgar G.

A Promising Fungous Pathogen of Adult Japanese

Beetles {Popillia japonica) 401

Sanford, Leonard J.

A New Record for Connecticut 244

Atrytone logan Edwards 258

SoRAci, Frank A.

Distribution in New Jersey of Mylloceims castaneus

Roelofs 318

Scotland, Minnie B.

Review and Summary of Insects Associated with Lemna

minor 319

Sherman, John D., Jr.

Henry Clinton Fall 33

Spieth, Herman T.

Studies on the Biology of the Ephemeroptera, II. The
Nuptial Flight 379

IV

Page

SuGDEN, John W.

Characteristics of Certain Cicadas 117

Townsend, Charles H. T.

Review of “Fauna of British India” 395

Weiss, Harry B.

Money Losses Due to Destructive Insects 195

The Death-Feints of Sitophilus granarius Linn., and
Sitophilus oryzce Linn 37

The Death-Feint of Trox unistriatus Beauv 303

White, Ralph T.

The Relation of Ants to the Japanese Beetle and its Es-
tablished Parasites 85

Book Notices 32, 100, 126, 393, 403

Notice to Authors 100

Notice to Members 103

Proceedings of the New York Entomological Society 295

VoL XLVIII

No. 1

MARCH, 1940

Journal

of the ^ ‘

New York Entomological Society

Devoted to Entomology in General

Edited by HARRY B. WEISS

Publication Committee

HARRY B. WEISS J. D. SHERMAN, Jr.

WILLIAM MOORE E. L. BELL

Subscription $3.00 per Year

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

LANCASTER. PA.
liEW YORK, N. Y.

1040

CONTENTS

An Introductory Study of Polistes in the United States and
Canada with Descriptions of Some New North and South
American Forms (Hymenoptera: Vespidae)

By Joseph C. Bequaert ... 1

Book Notice 32

Henry Clinton Fall

By John D. Sherman, Jr . 33

The Death-Feints of Sitophilus Granarius Linn., and Sito-
philus Oryzae Linn.

By Harry B. Weiss 37

Butterflies of New Jersey

By William Phillips Comstock .. .... .. 47

The Relation of Ants to the Japanese Beetle and its Estab-
lished Parasites

By Ralph T. White 85

Notice to Authors 100

Book Notices 100

Notice to Members 103

NOTICE: Volume XLVII, Number 4, of the Journal of the
New York Entomological Society was published on De-
cember 29, 1939.

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. XLVIII March, 1940 No. 1

AN INTRODUCTORY STUDY OF POLISTES IN THE
UNITED STATES AND CANADA WITH DE-
SCRIPTIONS OF SOME NEW NORTH
AND SOUTH AMERICAN FORMS
(HYMENOPTERA; VESPID^)

By Joseph C. Bequaert
Museum of Comparative Zoology, Cambridge, Mass.

Over ten thousand wasps have now been examined for the
forthcoming revision of North American Polistes. Although I
hope to study as many more before completing the work, the
time has come to put on record some of the results, for the benefit
of others. Moreover, the manuscript names distributed to vari-
ous collections should be validated, according to the rules, without
further delay.

In the United States I recognize four ‘‘structural species,”
only one of these entering Canada. Each species is represented
by several color forms, or “varieties.” The males of the four
species are readily told apart; but most of the differentiating
characters of the females or workers are subtle and difficult to
put in words. They become gradually more apparent as one
examines more specimens. This should be kept in mind when
using the subjoined key. In all social species of Polistes with
which I am acquainted, the workers (when present) and fertile
females (or queens) are alike, both in structure and color and
often also in size. It is generally assumed that the smaller fe-
males, sometimes found in the colony, are workers. Moreover,
even unmated females (or workers) may occasionally lay eggs,
which develop by parthenogenesis into male wasps.

APR 2 1940

2

Journal New York Entomological Society [Vol. XLVIII

KEY TO SPECIES

1. Body slender. Abdomen elongate-fusiform, widest about mid-length,

more or less compressed apically; first tergite slightly convex in profile
and gradually sloping onto the base, seen from above slightly longer
than wide at apex. Second sternite little convex, not bulging in pro-
file. Apical sternite of male without medio-basal tubercle 2.

Body thickset. Abdomen ovate-fusiform, widest before middle, more or
less depressed apically ; first tergite strongly convex in profile and
rather abruptly sloping onto the base, seen from above as wide at
apex as long or wider. Second sternite strongly convex, bulging in
profile. Mesopleura with microscopic sculpture only 3.

2. Propodeum distinctly striate transversely throughout. Mesopleura with

many large punctures scattered in the microscopic sculpture. Collar of
pronotum high and sharp, forming raised humeral angles. Clypeus of
female touching the eyes over a distance at most half the length of
the oculo-malar sipace. Clypeus of male subquadrate and flattened,
contiguous to the eyes P. canadensis.

Propodeum very finely striate, except in the median groove. Mesopleura
with the microscopic sculpture only, without larger punctures. Collar
of pronotum low and blunt, weakly projecting at the humeri. Clypeus
of female touching the eyes over a distance about equal to the length
of the oculo-malar space. Clypeus of male irregularly heptagonal and
slightly convex, broadly contiguous to the eyes P. exclamans.

3. Propodeum coarsely striate throughout, with some 20 to 22 strong trans-

verse ridges. Mesopleura often with at least traces of a prepectal
suture running obliquely upward on the anterior convexity. Clypeus of
female touching the eyes over a distance at most half the length of the
oculo-malar space. Clypeus of male subquadrate, with the anterior
margin straight or very slightly convex, the sides always widely sepa-
rated from the eyes. Apical sternite of male without medio-basal
tubercle P. major.

Propodeum finely or moderately striate, usually more strongly in the
median groove, the striae numerous but not ridge-like. Mesopleura
never with a trace of a prepectal suture, the anterior convexity smooth.
Clypeus of female touching the eyes over a distance equal to one-half
or more of the length of the oculo-malar space. Clypeus of male irregu-
larly hexagonal, more or less broadly separated from the eyes (often
scarcely so). Apical sternite of male with a small but distinct medio-
basal tubercle P. fuscatus.

Two names proposed for North American Polistes remain a
pnzzle. In 1872 (Trans. Amer. Ent. Soc., IV, pp. 245-246)
Cresson described P. perplexits and P. generosus, both based on
males only, from Texas. I have studied his types at the Phila-
delphia Academy of Sciences, on several occasions. There can

Mar., 1940]

Bequaert: Polistes

3

be no doubt that P. generosus is only a variant of P. perplexus,
as Cresson surmised, and it should be treated as a synonym of the
latter. These males agree in every important structural charac-
ter with P. fuscatus and at one time I regarded them as aberrant
males of typical fuscatus. They are unusual, however, in being
larger and especially stouter than the regular males of even var.
metricus and var. ruMginosus. 1 have in my collection 12 males
from Texas (some without more definite locality; others from
College Station, Shiloh and Brazos Co.), 1 male from Kentucky
(Woodford) and 1 male from Indiana (Bloomington), which can-
not be separated from perplexus and generosus. I have also 6
females from Texas (some without more definite locality; others
from Austin and College Station) and 1 female from Indiana
(without more definite locality), which appear to be the other sex
of Cresson ’s perplexus and generosus. These females also are
unusually large and stout for forms of P. fuscatus. In both
sexes these supposed perplexus wasps give the impression of
being more powerful insects, with very strong striation of the
propodeum, the head more swollen (particularly the outer orbits)
and the legs and antennae slightly shorter and stouter. I have
been unable, however, to find more tangible structural differences
from P. fuscatus. I should have attached little importance to
the more powerful build of these supposed perplexus, were it not
for the possibility that they may represent a distinct species,
living as a social parasite in the colonies of either var. metricus
or var. ruhiginosus. This parasitic species may have been de-
rived so recently from its social ancestor (presumably some form
of P. fuscatus), that it has as yet developed no cogent structural
differences. Several Palearctic species of Polistes are now known
to be true social parasites (W. Weyrauch, 1937, Zool. Jahrb., Abt.
Syst. Tiere, LXX, pp. 243-290) and I have suggested recently
that some of the Ethiopian species have similar parasitic habits
(J. Bequaert, 1938, Bev. Zool. Bot. Afric., XXXI, p. 130). No
doubt parasitic Polistes will also be discovered in the New World.
Carefully controlled observations in the field alone will enable
one to decide whether or not my surmise is correct in the case of
P. perplexus. If it is a true social parasite, the species will con-
sist only of males and impregnated females. The females will
build no nests nor form colonies of their own. Instead, they will

4

Journal New York Entomological Society [Vol. xlviii

seek out existing colonies of some social Polistes, in which to de-
posit their eggs. The larvae hatching from these alien eggs will
then be raised by the original and rightful builders and inhabi-
tants of the nest.

The wasps recorded as P. perplexus, from Bermuda, by W. F.
Kirby, Verrill and Ogilvie, are P. fuscatus var. nest or (Fab-
ricius).

Vespa Carolina Linnaeus (1767, Syst. Nat. 12th Ed., I, pt. 2,
p. 948), from Carolina, was undoubtedly based upon a North
American Polistes, as I have pointed out before (1931, Entomo-
logica Americana, XII, p. 108). The type is so poorly preserved
and the description so inadequate, that it is impossible to decide
whether it was P. canadensis var. anmdaris or one of the forms
of P. fuscatus (possibly a male of P. fuscatus var. ruhiginosus) .
For this reason, it seems preferable to drop the name altogether.

The following names have been applied erroneously to Polistes
of our territory.

P. crinitus (Felton). — This is a strictly West Indian wasp,
structurally distinct from the four North American species. Its
nearest relative in North America is P. exclamans, which has
sometimes been mistaken for it.

P. minor Palisot de Beauvois, described originally from Santo
Domingo, is also strictly West Indian in my opinion. The wasps
frequently called P. minor in the United States are usually P.
fuscatus var. hunteri, sometimes P. exclamans.

P. carnifex (Fabricius) is occasionally confused with P. major.
True carnifex, as characterized in 1936, Rev. de Entomologia,
VI, pp. 376-383, I have never seen from the United States or the
West Indies.

P. instahilis H. de Saussure was recorded by F. Smith (1857)
from East Florida, Mount Pleasant (Ohio) and New York. True
instahilis, however, is a Central American wasp (See under P.
exclamans).

In the enumeration of species and varieties I have inserted the
published names which I regard as synonyms, without further
discussing the matter. The opportunity has been used to publish
the descriptions of some new color varieties from the West Indies,
Central and South America.

In brief, only three forms of Polistes fuscatus are known with

Mar., 1940]

Bequaert: Polistes

5

certainty from Canada: var. pallipes (Quebec and Ontario), var.
connectens (Alberta) and var. aurifer (British Columbia). — In
the United States the following occur : P. canadensis, typical, P.
canadensis var. annularis, P. canadensis var. comanchus, P.
canadensis var. kaihahensis, P. canadensis var. navajoe, P. ex-
clamans, typical, P. exclamans var. louisianus, P. major, typical,
P. major var. palmarum, P. major var. castaneicolor, P. major
var. kakeri, P. fuscatus, typical, P. fuscatus var. anaheimensis,
P. fuscatus var. apachus, P. fuscatus var. aurifer, P. fuscatus var.
centralis, P. fuscatus var. hellicosus, P. fuscatus var. connectens,
P. fuscatus var. flavus, P. fuscatus var. hunteri, P. fuscatus var.
metricus, P. fuscatus var. montanus, P. fuscatus var. nestor, P.
fuscatus var. pallipes, P. fuscatus yslt. ruhiginosus, P. fuscatus
var. utahensis, and P. fuscatus var. variatus.

The North American Polistes, like those of other parts of the
world, are quite variable in the extent, arrangement and shade of
color markings. The extreme case is perhaps that of P. fuscatus,
in which color seems to run riot and to defy any attempt at
rational analysis. The true meaning of these variations as yet
escapes ns, although a theory is by no means lacking. What we
really should know, before attempting to build up theories, is the
genetic background of the color differences, whether or not and to
what extent they are hereditary in a given population and how
they behave when crossed ; also to what extent they may be in-
fluenced by environmental conditions (nature and amount of
food, climate, resistance to disease, natural selection by predators,
etc. ) . It would seem that only experiments can furnish decisive
answers to these questions. Meanwhile, from observation alone,
the several color variations of one structural species appear to
be of unequal value, hence probably due to different causes.
At least four types of variants may be recognized.

(1) Some variations may be purely individual or fluctuating
and probably due entirely to environmental conditions during
the ontogeny. It is frequently difficult to And two individuals
colored alike in every detail, even within the same population or
among the offspring of a single queen. The differences among
the inmates of one nest may, however, be explained to some extent
by the frequent association of two or more queens in the found-

6

Journal New York Entomological Society [Vol. xiiVllI

ing of a new colony (pleometrosis : See J. Bequaert, 1923, Bull.
Brooklyn Ent. Soc., XVIII, pp. 73-80).

(2) Sometimes two or more variants occur in the same locality,
which, though quite distinct in the extreme or ‘‘typical” indi-
viduals, yet are connected by many intergrades. In most cases
the intergrades are probably hybrids. In the northeastern
United States, for instance, one finds three forms of P. fuscatus:^
(a) typical fuscatus is relatively uncommon in the central part
of this area, from southern Massachusetts to West Virginia; (b)
the var. nestor is common from southern Pennsylvania to South
Carolina and eastern Texas, with an occasional specimen in New
Jersey and on Long Island; (c) the var. pallipes is characteris-
tic of southern Canada, New England, and northern New York,
with a few stragglers farther south in the Alleghany Mountains.
Prom the distribution and relative scarcity^ it would seem that
typical fuscatus (as described by Pabricius) may not be a distinct
“race” at all, but merely a name or label for hybrids between
pallipes and nestor, where the ranges of these two forms overlap.

(3) Two or more variants of a species may occur over much
the same territory, yet be so well defined that intergrades are
hardly ever met with. Thus in the mid-western States, particu-
larly in the drainage of the Mississippi, one finds commonly in
one locality three varieties of P. fuscatus, viz., variatus, metricus
and rubiginosus. Each by itself is more or less variable, but
specimens that might be regarded as connecting two of them are
exceedingly rare or lacking. Their scarcity rather indicates that,
in this case, such intermediate specimens are merely extreme
individual variants, not true intergrades or hybrids. Perhaps
w^e are dealing here with ecological, physiological or even beha-
vioristic races. Phil Ran has shown, for instance, that in the
region of St. Louis, Missouri, three color forms of Polistes fus-
catus nest in different types of environment (1931, Bull. Brooklyn
Ent. Soc., XXVI, pp. 111-116). The “isolation” of such races
may be enhanced by the social behavior, brothers and sisters tend-
ing to mate on or near the nest, while young queens often return
to their mother’s nesting site. In any case, the correlation be-

1 As is the rule in P. fuscatus, these three forms are recognizable in the
female and worker only. The males of this species rarely show the color
differences on which the several varieties are based.

Mar., 1940]

Bequaert: Polistes

7

tween such extra-morphological differences and the visible color
characters can only be fortuitous.^

(4) Finally, in a few cases the variants agree fairly well with
the classic concept of “geographical races” or “subspecies.”
They are sharply defined, apparently without intergrades to other
forms, and inhabit a limited district to the exclusion of other color
variants of the same species. There are few clear-cut cases of
this type among the variants of P. fuscatus, the nearest approach
being perhaps the two forms of the Pacific Coast, var. aurifer and
var. anaJieimensis, although the areas of both overlap to some
extent. Better examples are the variants of P. canadensis: var.
anmdaris (in the southeastern States), var. kaihahensis (in the
Grand Canyon of the Colorado River), var. navajoe (in the Up-
per Sonoran of Arizona), and var. comanchus (in the Upper
Sonoran of southwestern Texas).

A corollary to the differentiation of “races” on a geographical
basis is the convergence in color pattern, or homeochromy, some-
times exhibited by several structural species of Polistes, other
wasps or other insects in general, occurring in the same area.
Illustrations of this are few in the North American Polistes.
There is, however, a decided superficial resemblance between P.
fuscatus var. metricus and P. canadensis var. annularis, in the
southeastern States ; between typical P. exclamans and P. fuscatus
var. hellicosus, in the southeastern States; and between P. cana-
densis var. navajoe and Miscliocyttarus flavitarsis var. navajoe in
Arizona. Whether or not these are cases of “protective colora-
tion, ’ ’ I shall not attempt to discuss.

Polistes canadensis (Linnaeus)

The most widely distributed American species of the genus,
although it does not reach Canada. Linn^us was evidently
misled as to the locality of his specimens, like in so many other
cases. It extends from south of the Great Lakes and Utah to
northern Patagonia. It is, however, unknown in California and
Chile. In this vast territory it has produced several color forms,
thirteen of which have been named thus far. Pour more are

2 A similar fortuitous correlation exists, for instance, between the egg-pat-
tern and the physiological differences of the several races of Anopheles
maculipennis.

8

Journal New York Entomological Society [Vol. xlviii

described below. Of these seventeen varieties, five only occur in
the United States.

KEY TO NEARCTIC COLOR FORMS OF P. CANADENSIS

1. Abdomen either without yellow markings, or the first tergite only with

an apical yellow margin 2.

Abdomen more extensively yellow or orange-yellow, particularly over the
third and succeeding segments, which are often almost entirely of that
color 3.

2. Body fairly uniformly light to dark mahogany-brown, the head and thorax

sometimes lighter than the abdomen. Yellow apical margin of first
tergite more or less distinct, sometimes lacking. Wings purplish-
black, sometimes with a russet tinge, which becomes gradually more
pronounced apically, the veins and stigma either black or russet accord-
ing to the color of the surrounding membrane typical canadensis.

Head and thorax dark mahogany-brown, sometimes blotched with black;
abdomen usually jet-black, blotched with mahogany-brown on the first
segment and occasionally elsewhere. First tergite always with a dis-
tinct yellow apical margin. Wings uniformly deep purplish-black, the
veins and stigma black var. annularis.

3. Head, thorax, legs, and abdomen light reddish-brown, with extensive yellow

markings on head and abdomen. Wings ferruginous, very slightly pur-
plish; costal area, veins and stigma russet var. Icaihadensis.

Ground color of head, thorax, and legs black 4.

4. Head black, extensively marked with orange-yellow; thorax black, only

narrow margins of pronotum and tegulae orange ; abdomen mostly
orange-yellow, elsewhere ferruginous, only the base of first tergite black.
Wings ferruginous, slightly purplish, the veins and stigma russet.

var. comancJms.

Head black, extensively marked with yellow ; thorax black, the pronotum
partly, tegulae and scutellum orange-yellow; abdomen mostly yellow,
elsewhere black, the first and second tergites somewhat reddish at the
limits of black and yellow areas. Wings strongly purplish-black, only
extreme base and costal area somewhat russet, the veins and stigma
black var. navajoe.

1. P. canadensis, typical. — Distributed over most of the Neo-
tropical Region. I have seen it from Mexico, Guatemala, Hon-
duras (Rep.), Colombia, Venezuela, British Guiana, Trinidad,
Brazil, Peru, Bolivia and Paraguay. It barely enters the United
States at the southern border of Arizona. Two females taken at
Nogales by Dr. J. C. Bradley are exactly like a series I have
received from Vera Cruz and Sonora (Bakachaka; Estrella Dis-
trict, Alamos).

Mar., 1940]

Bequaert: Polistes

9

2. P. canadensis var. amazonicus W. A. Schulz. — Known from
Brazil, British Guiana, Panama, Ecuador and British Honduras.

3. P. canadensis var. annularis (Linn^us). — Vespa cincta
Drury (1773), Vespa virginiensis ‘‘Drury” Dalla Torre (1894)
and Polistes annulata Moebius (1856) are synonyms. The most
widely distributed form of the species in the United States. I
have seen it from Connecticut (New Haven), southern New York
(Ft. Montgomery; Hillburn; Orient, Long Island; Ramapo Mts. ;
also taken many years ago on Staten Island by Mr. AYni. T.
Davis), New Jersey (Paterson; Jamesburg; Cape May), Penn-
sylvania (Pittsburgh), Maryland, Virginia, District of Columbia,
West Virginia, southern Ohio (Hocking Co.; Sugar Grove, Ross
Co.; Clifton, Greene Co.), southern Indiana, southern Illinois
(New Columbia; Fort Chartres State Pk.), southern Iowa (Page
Co.), Missouri, Kentucky, Tennessee, North Carolina, South Caro-
lina, Georgia, Florida, Alabama, Mississippi, Oklahoma, eastern
Kansas (Blue Rapids), and eastern Texas (as far west as Bexar
Co.). It will be found also in Delaware, Arkansas, and south-
eastern Nebraska. It is not known from south of 42°, west
of the 100th Meridian, or outside the United States.

4. P. canadensis var. huyssoni BrMlies, 1909, An. Mus. Nac.
Buenos Aires, (3) X, p. 456. — Syn. : Polistes cavapyta var.
thoracica R. du Buysson, 1906, Ent. Medd., Copenhagen, III, p.
19. Not Polistes thoracicus Fox, 1898. This form is common in
certain parts of Argentina.

5. P: canadensis var. cavapyta de Saussure. — Polistes opalinus
de Saussure appears to be a synonjun. A form of northern
Argentina, southern Brazil, Uruguay and Paraguay.

6. P. canadensis var. cinctus Lepeletier. — Vespa (Polistes)
dominica Vallot (1802) may be the same. A form peculiar to
the lesser Antilles, where it is known from Martinique, St.
Vincent, St. Kitts, the Grenadines and Barbadoes. It was also
introduced into Montserrat and St. Lucia, where it appears to
have taken a foothold. This is one of the smaller forms of the
species (fore wing 14 to 17 mm. long).

7. P. canadensis var. comanchus de Saussure. — This form is
characteristic of the Upper Sonoran life zone of southwestern

10

Journal New York Entomological Society [Vol. xlviii

Texas (Brewster Co.; Jeff Davis Co.). I have seen it also from
the Sierra de los Burros, State of Coahuilla, Mexico.

8. P. canadensis var. erythrocephalus Latreille, 1817, in Hum-
boldt and Bonpland, Recueil d ’Observations de Zool. Anat. Comp.,
II, p. 96, PL XXXVIII, fig. 3 (5; South America). — The de-
scription and figure leave no doubt as to the identity of this
form. It is characterized by the jet-black thorax and abdomen,
the ferruginous head and antenna (except for the median black
ring), the black legs with ferruginous or orange knees and tarsi,
and the bluish-black wings. At one time I distributed it as a
new variety with a name derived from Costa Rica. Dalla Torre
overlooked Latreille ’s name. The var. erythrocephalus is found
in Nicaragua (Sioux Plantation, Rio Grande; Eden), Costa Rica
(common), Panama (common), parts of Colombia (Dept.
Santander, Dept. Boyaca and Dept. Tolima), and southern Brazil
(Sete Lagoas, State of Minas Geraes). It is sometimes called P.
rufidens in collections, but de Saussure’s rufidens is structurally
distinct from P. canadensis.

9. P. canadensis var. ferreri H. de Saussure. — ^Uruguay and
northern Argentina.

10. P. canadensis var. kaihahensis Hayward, 1932, Proc. Utah
Ac. Sci., IX, p. 89, PL IX, figs. 4, 12, 13 and 16. This interesting
form is known only from the rim of the Grand Canyon of the
Colorado River, Arizona, where it is common.

11. P. canadensis var. lanio Pabricius. — I have seen this from
Brazil and Peru.

12. P. canadensis var. (or subsp.) mexicanus, new.

Female and WorTcer. — Body and legs bright reddish-brown, only the su-
tures of the thorax, a median line over anterior half of mesonotum and a
narrow stripe in concavity of propodeum (widened below), black; middle
of flagellum infuscated; the following markings are yellow: apical fasciae
on all tergites and second to last sternites (reduced on first and second,
very extensive on the others and more or less divided on each side by a wavy
reddish line or sometimes emarginate) and the knees and hind tarsi more or
less. Wings very strongly infuscate, purplish-black.

Male. — Colored exactly like the female.

Holotype: Cuernavaca, State of Morelos, Mexico, female (G.
Lassman). — Allotype: Cuernavaca, male (A. L. Melander). —
Paratypes: Cuernavaca, four females and two males; Teotihuacan,

Mar., 1940]

Bequaert: Polistes

11

State of Mexico, Mexico, two females (S. B. Jones) ; Valerio,
Trujano, State of Oaxaca, Mexico, two females (M. Embury and
A. Mead). — Holotype, allotype and several paratypes at Museum
of Comparative Zoology, Cambridge, Mass. ; two paratypes in Dr.
R. M. Boliart ’s collection.

This form is n*earest to the Argentinian varieties ferreri,
cavapyta and huyssoni, but the abdomen is more extensively
marked with yellow than in ferreri, while the yellow markings
of head and thorax of cavapyta and huyssoni are lacking. In the
North American fauna it is nearest var. kaihahensis and var.
wheeleri, differing from the former in the color of the wings and
from both in the reddish-brown head.

13. P. canadensis var. navajoe Cresson. — A common form in
Arizona, also found in northern Mexico : south of Nogales, State
of Sonora (P. P. Calvert) ; Escuinapa, State of Sinaloa (J. H.
Batty). Apparently characteristic of the Upper Sonoran life
zone.

14. P. canadensis var. panamensis Holmgren. — Common in
Panama and parts of Colombia (Santa Marta district).

15. P. canadensis var. (or subsp.) satan, new.

Female and Worlcer. — Head, thorax and abdomen jet-black, scarcely with
any ferruginous or yellowish markings; a very narrow inner border of the
eyes, outer orbits, lower portion of clypeus, mandibles, hind margin of pro-
notum and a blotch on tegula3, very slightly suffused with reddish-brown.
Antennse black, base and apex more or less tinged with chestnut-brown.
Legs black; base of tibiae, outer side of fore tibiae, and fore tarsi more or
less ferruginous; mid and hind tarsi dirty yellow to orange. Wings uni-
formly bluish-black, with black veins and costa; stigma dark brown.

Male. — Almost exactly like the female; very narrow inner orbits dirty
yellow.

Holotype: Lassance, State of Minas Geraes, Brazil, female
(Cornell Univ. Exped.). — Allotype: Bello Horizonte, State of
Minas Geraes, Brazil, male (Cornell Univ. Exped.). — Paratypes:
Lassance, one male and six females; Pirapora, State of Minas
Geraes, three females (Cornell Univ. Exped.) ; Fazenda do Dia-
mante, Corinto, State of Minas Geraes, one female (Ynes Mefia) ;
La Esperanza, Dept. Cundinamarca, Colombia, one female (E.
Eoba). — Holotype, allotype and six paratypes in the Department
of Entomology, Cornell University; four paratypes at Museum

12

Journal New York Entomological Society [Vol. XLVIII

of Comparative Zoology, Cambridge, Mass., and one paratype at
the Academy of Natural Sciences of Philadelphia.

The extreme melanistic form of the species, differing from the
var. erythrocephalus in having the head also practically entirely
jet-black.

16. P. canadensis var. (or subsp.) weberi, new.

Fe7nale and WorJcer. — Mostly black, with brownish tinges over some of
the abdominal segments ; most of head, most of pronotum, tegulse, a small
spot in upper corner of mesopleura, and antennse (except for median black
area of flagellum), mahogany-brown. Pale sulphur-yellow markings re-
stricted to the large apical margin of first tergite (emarginate in the
middle and separated from the black base by a reddish-brown area), the
knees, the basal half or more of all tibiae, and all tarsi. Wings strongly
infuscated, purplish-black in basal two-thirds, more russet in apical third.

Male unknown.

Holotype: Macnto near La Gnaira, Venezuela, female (Neal
A. Weber). — Paratypes: Macnto, three females (N. A. Weber) ;
Caracas, Venezuela, three females. — Holotype and paratypes at
the Museum of Comparative Zoology, Cambridge, Mass. ; para-
types also at the American Museum of Natural History.

In general coloration of body and wings the var. weheri re-
sembles the varieties panamensis and amazonicus, but it differs
from these in the large yellow margin of the first tergite. This
margin is much more extensive than in var. annularis, which,
moreover, has uniformly purplish-black wings. It recalls var.
cinctus, of the Lesser Antilles, but that form is fairly uniformly
reddish-brown with entirely yellowish-russet wings. The var.
weheri is also larger (fore wing 20 to 22 mm. long). It was ob-
tained in the arid coastal area of Venezuela, covered with xero-
phytic scrub forest and characterized by the cactus, Cereus griseus.

17. P. canadensis YSLY. (or subsp.) wheeleri, new.

Female. — Body and legs bright reddish-brown; only the ocellar area,
sutures of the thorax, a median line over anterior half of mesonotum and
a narrow stripe in concavity of propodeum (widened below), black; middle
of flagellum infuscated. The following markings are yellow: clypeus, lower
half of face, most of outer orbits and mandibles, under side of scape, apical
fascise on all tergites and on second to last sternites (reduced on first and
second, very extensive on the others and more or less divided on each side
by a wavy reddish line), the knees, the basal third of all tibiae, and most of

Mar., 1940]

Bequaert: Polistes

13

all tarsi. Wings moderately infuscated, somewhat purplish-black and
tinged with russet basally and along anterior margin.

Male unknown.

Holotype: Bochibampo Bay, Guayamas, State of Sonora,
Mexico, female, March 3, 1937 (Wm. M. Wheeler). — Paratypes:
Copete Mine, 30 Kilom. east of Carbo, State of Sonora, 4 females
(F. C. Nicholas). — Holotype at Museum of Comparative Zoology,
Cambridge, Mass. ; paratypes also at the American Museum of
Natural History.

Related to the var. mexicanus, from the Central Plateau of
Mexico, but distinct by the extensively yellow head and legs and
the partly russet wings. It is also near var. comanchus (which
has the thorax almost wholly black) and var. kaibahensis (which
has uniformly russet wings).

Polistes exclamans Viereck

In 1931, G. Salt and I described a Polistes hahamensis, with two
color forms, var. hilineolatus and var. picturatus, all from the
Bahamas. I have since recognized that these Bahaman wasps are
structurally not separable from the continental P. exclamans.
Two forms of exclamans occur in the United States.

Polistes instabilis H. de Saussure (Syn. : P. oculatus F. Smith),
from Mexico, the Republic of Honduras and Costa Rica, resembles
typical P. exclamans in color. The males are readily distin-
guished by the very bulging eyes, the subquadrate clypeus and
the slender antennae (with all segments of flagellum at least twice
as long as thick). I have not yet found reliable structural differ-
ences in the female and worker.

I. P. exclamans, typical. — This is a common wasp in the south-
eastern United States. I have seen it from North Carolina, South
Carolina, Georgia, Florida, Texas, Louisiana (Baton Rouge;
Olivier), Oklahoma, Arkansas, Kansas, Colorado and Arizona.
Although somewhat variable in color, it is always well marked
with yellow, often profusely so; the propodeum bears usually
four yellow stripes ; the wings lack the yellow-russet tinge of the
Bahaman varieties and the mesonotum seldom bears yellow longi-
tudinal lines. Some specimens from Metacumbe Key and else-
where on the east coast of Florida, are somewhat transitional

14

Journal New York Entomological Society [Vol. xlviii

between typical exclamans and the var. l)ilineolatus. Typical
exclamans is readily confused with P. fuscatus var. hunteri or
P. fuscatus var. ~bellicosus, and in collections it is often called also
P. crinitus or P. minor.

2. P. exclamans var. (or subsp.) louisianus, new.

Female and WorTcer. — ^Eeddisli mahogany brown, with only the middle of
the flagellum above, the extreme base of some of the abdominal segments
and the outer side of the mid and hind tibiae infuscate or blackish. Pale
markings ivory-white and scanty: a narrow hind margin to the pronotum,
most of tegulae, anterior angles of scutellum, linear anterior margin of
postscutellum, an elongate spot on mesopleura (beneath tegulae), two nar-
row median stripes on propodeum (sometimes lacking), linear apical mar-
gins on first to third abdominal tergites (more extensive on the first; some-
times lacking on the third), knees, bases of tibiae, and most of tarsi. Wings
as in typical form, uniformly infuscate and purplish.

Holotype: New Orleans, Louisiana, female (Ed. Poster). —
Paratypes: two females from the same locality. — Holotype and
one paratype at U. S. National Museum ; one paratype at Museum
of Comparative Zoology, Cambridge, Mass.

In the absence of the male, the specific identity of this form is
not quite certain. I suspected at first that these three wasps
might be some form of either P. crinitus or P. versicolor, acci-
dentally introduced by man. Structurally, however, the New
Orleans wasps are indistinguishable from P. exclamans. On the
other hand, I have never seen a specimen of either P. crinitus or
P. versicolor, from Central and South America and the Antilles,
colored like them. The reduction of yellow markings is quite
unusual for P. exclamans.

3. P. exclamans var. l)ahamensis Bequaert and Salt (1931, Ann.
Ent. Soc. America, XXIV, p. 793). — Known only from the Ba-
hamas, where it occurs on Andros Island. This is the most dis-
tinct of the several color forms of the species.

4. P. exclamans var. hilineolatus Bequaert and Salt (1931,
op. cit., p. 796). — Known only from the Bahamas, where it is
found on New Providence Island and Eleuthera.

5. P. exclamans var. picturatus Bequaert and Salt (1931, op.
cit., p. 797). — Known only from the Bahamas, where it is found
on Acklin Island, Mariguana, Rum Cay, Crooked Island, Long
Island, Watlings Island and Cat Island.

Mar., 1940]

Bequaert: Polistes

15

Polistes major Palisot de Beauvois

I have discussed this species on two previous occasions (1936,
Ent. News, XL VII, pp. 7-13; 1937, Arch. Inst. Biol. Veget., Kio
de Janeiro, III, pp. 173-174) . It occurs from the southern United
States to Brazil, as well as in the Bahamas, Cuba and Santo Do-
mingo. It has recently been introduced into Puerto Bico.

KEY TO NEARCTIC COLOR FORMS OF P. MAJOR

1. Thorax more or less extensively marked with yellow. Abdomen with

extensive yellow apical margins on all tergites, sometimes covering
them almost entirely 2.

Body almost uniformly dark chestnut-brown; only the tarsi and blotches
on the head yellowish. Wings uniformly brownish, costal area some-
what russet, veins and stigma chestnut-brown var. castaneicolor.

2. Ground color of body light ferruginous-brown. Wings uniformly russet-

fuscous, slightly purplish var. palmarum.

Dark areas of body cinnamon-brown to blackish-brown. Wings dark
cinnamon-brown, with a russet tinge and slightly purplish 3

3. Thorax very extensively marked with yellow, particularly on the pro-

podeum typical major.

Thorax with reduced yellow markings, which are absent on the pro-
podeum var. haTceri.

1. P. major, typical. — Known from Mexico, Guatemala, the
Republic of Honduras, Nicaragua, Brazil, Cuba, Isle of Pines,
Santo Domingo, Navassa Island and the Bahamas ; recently intro-
duced into Puerto Rico. In the United States it has only been
taken in a few localities of southern Arizona ; these specimens are
colored exactly like those from farther south and from the West
Indies.

2. P. major var. (or subsp.) bakeri, new.

Male. — Similar to typical major, but chrome-yellow markings much re-
duced on thorax, being restricted to collar of pronotum (narrowly above,
very broadly on the sides) and part of scutellum and postscutellum (shad-
ing into cinnamon-brown) ; mesonotum, mesopleura, metapleura and pro-
podeum black; major part of pronotum, four elongate spots on mesonotum
and small spot on upper part of mesopleura, reddish-brown.

Female or WorTcer. — Agreeing with the male in the reduction of yellow on
thorax, the propodeum being black, unstriped. The allotype is colored
almost like the holotype. The female paratype has most of pronotum,
scutellum and postscutellum, and a large spot in upper part of mesopleura
chrome-yellow.

16

Journal New York Entomological Society [Vol. XLVIII

Holotype: San Antonio, Bexar Co., Texas, male (G. P. Engel-
liardt). — Allotype: Big Bend Park, Brewster Co., Texas, female
or worker (Rollin H. Baker). — One female paratype with same
data as allotype. — Types at Museum of Comparative Zoology,
Cambridge, Mass. The male was selected as the holotype, being
the only perfect specimen.

3. P. major var. honaccensis J. Beqnaert (1937, op. cit., p. 174).
— This form is known only from Bonacca Island, off the north
coast of the Republic of Honduras. It is homeochromic with
P. carnifex var. ochreata Spinola, which was also taken on Bonacca
Island by Dr. M. Bates. On the neighboring island of Roatan he
found only the typical form of P. major.

4. P. major var. castaneicolor J. Beqnaert. — Known only
from Mexico (State of Jalisco) and a few localities in southern
New Mexico (Alamogordo) and in southern Arizona (Sabino
Canyon, Sa. Cataline Mts. ; etc.).

5. P. major var. palmarum J. Beqnaert. — Known only from
southeastern California and Lower California, Mexico (San Jose
del Cabo).

Polistes fuscatus (Fabricius)

The most common North American species of the genus covers
the entire United States and the southern areas of the Dominion
of Canada, in southern Quebec, southern Ontario, southeastern
Alberta and southern British Columbia (northernmost locality:
Chilcotin, in about 52° N., farthest north for any American
Polistes).^ It extends southward to Mexico, Guatemala and the
Republic of Honduras ; but its distribution is poorly known there.
It has been introduced accidentally by man into the Bermudas,
Jamaica, Barbadoes, the Society Islands and the Hawaiian
Islands, where it is now more or less established. Polistes fortu-
natus Kirby, described from the Cape Verde Islands, appears to
be one of the color forms of P. fuscatus (probably var. nest or
Fabricius), imported by man from the New World. All published
records of P. fuscatus from continental South America appear
to be based upon a confusion with P. pacificus Fabricius, which
is structurally very different.

3 There are as yet no records of Polistes from New Brunswick and Nova
Scotia.

Mar., 1940]

Bequaert: Polistes

17

P. fuscatus is perhaps the most variable wasp in existence, with
regard to color and also to certain structural characters (par-
ticularly the shape of the clypeus of the male). These matters
will be discussed more fully later. In the present preliminary
study, I have been as conservative as possible, retaining most of
the forms named by my predecessors. I recognize eighteen varie-
ties, sixteen of which occur in the United States; five are here
described as new.

KEY TO COLOR FORMS OF P. FUSCATUS
The unusually wide range of variation of most color forms, or varieties,
of P. fuscatus, makes it difficult to construct a workable key for their identi-
fication. Some of the varieties are placed twice in the key; but even this
device will not take care of many transitional specimens, which must be
named more or less arbitrarily. The key, moreover, is based on females
(queens and workers) only. The males are always more extensively yellow
than the corresponding females. They should either be bred or collected
from nests or matched arbitrarily with the forms known to occur in the
same geographical area.

1. Thorax (including mesonotum) predominantly bright yellow or orange-

yellow, with narrow russet or blackish sutures; mesonotum some-
times with small ferruginous or blackish blotches. Abdomen and
legs extensively or mostly yellow. Wings amber-yellow or infuscated
and more or less purplish var. flavus.

Thorax (at least mesonotum) predominantly black or light or dark fer-
ruginous, often more or less marked with yellow 2.

2. Small forms of the Bahamas or of Central America. Very extensively

yellow, particularly on the propodeum and first tergite. Fore wing
10 to 13 mm. long 3.

Forms of the American continent, usually larger (except var. hunteri) ,
much less extensively yellow; propodeum at most Avith narrow

stripes 4.

3. Form of the Bahamas. Mesonotum with four yellow stripes. Most of

second tergite yellow. Wings smoky and rather strongly purplish

var. maritimus.

Form of Central America. Mesonotum Avithout yelloAV stripes. Second

tergite mostly ferruginous. Wings yellowish-russet

var. neotropicus.

4. Dark areas of thorax mainly black, Avithout or Avith yellow or reddish

markings 5.

Thorax mainly light or dark ferruginous, Avith or Avithout yellow or
black markings 15.

5. Small form (fore Aving 10 to 18 mm. long) of the southeastern United

States. Thorax with at least pronotum and scutellum ferruginous;

18

Journal New York Entomological Society [Vol. xlviii

propodeiim almost always with yellow stripes. Most tergites with
yellow margins, the third and fourth usually extensively black.
Wings slightly yellowish russet, more or less purplish in the radial

cell var. Jmnteri.

Larger (fore wdng 15 to 20 mm. long) and of a different color 6.

6. Yellow markings scanty, often reduced to narrow hind margins on pro-

notum and one or more tergites; disk of second tergite without yel-

loAv lateral spots, the sides not or only narrowly yellow 7.

Yellow markings extensive, at least on abdomen; disk of second tergite
either with at least traces of lateral spots or extensively yellow on
the sides ! 10.

7. Mostly black; thorax and second tergite without or with mere traces of

ferruginous blotches; propodeum rarely with yellow .stripes; legs
black or more or less ferruginous, with the tarsi conspicuously yellow

var. palUpes.

Body and legs more or less ferruginous; either thorax or abdomen or
both distinctly blotched with ferruginous (at least on second tergite)

8.

8. Abdomen mostly ferruginous; bases of tergites black, apices of one or

more margined with yellow. Thorax more or less blotched with
ferruginous; mesonotum as a rule uniformly black; propodeum often

with yellow stripes var. nestor.

Abdomen mostly black 9.

9. Large (fore wdng 15 to 19 mm. long). Thorax rather extensively
blotched with ferruginous; mesonotum either mostly ferruginous or
Avith tw^o ferruginous stripes. Abdomen either entirely black or first
and second tergites (or one of them) more or less blotched Avith fer-
ruginous. YelloAV markings often almost lacking; propodeum with-
out yelloAV stripes, usually extensively ferruginous; sometimes first
tergite Avith yellow margin var. metricus.

Medium-sized (fore wdng 13 to 16 mm. long). Thorax not extensively
ferruginous, often only Avith small blotches on pronotum; mesonotum
as a rule uniformly black. Sides of second tergite and sometimes
blotches on first, ferruginous. YelloAV markings more or less devel-
oped; propodeum Avith or AAdthout yelloAV stripes; one or more tergites
Avith yelloAV margins typical P. fuscatus.

10. Propodeum wdthout yellow^ stripes; thorax mostly black. Second ter-

gite black, with yellow margin and two large yellow lateral spots
someAvhat fringed with ferruginous; third to sixth tergites mostly
yellow. Wings broAvnish-yelloAv; stigma and veins rufous

var. utahensis.

Propodeum Avith at least traces of longitudinal yellow stripes 11.

11. YelloAV color extending over most of the tergites; lateral spots (par-

ticularly of second tergite) large, either broadly connected Avith the
yelloAV sides and hind margins or se]3arated by a narrow^ and usually
faint line , 12.

Mar., 1940]

Bequaert: Polistes

19

Yellow color of abdomen restricted to sides and hind margins of ter-
gites and two small, entirely free, lateral spots on some or all ter-
gites (spots of first tergite sometimes lacking or connected with
apical band) 14.

12. Head black and yellow only. Thorax not or scarcely marked with

rufous. Black areas of second tergite as a rule not fringed with
rufous. Wings amber-yellow; stigma and veins rufous

var. aurifer.

At least head and often also pronotum blotched with rufous. • Black
areas of second tergite often more or less fringed with rufous 13.

13. Eufous color of thorax and abdomen extensive. Wings slightly smoky

and more or less purplish; stigma and veins russet var. apaehus.

Eufous color either absent on thorax and abdomen or restricted to
blotches on pronotum and fringes around the yellow areas. Wings
yellowish-gray, very slightly or scarcely purplish; stigma and veins
rufous var. connectens.

14. Black areas of second tergite either more or less replaced by rufous or

broadly fringed with rufous. Pronotum and head usually much
blotched with rufous. Wings smoky, markedly purplish ; veins

blackish, costa and stigma russet var. variatus.

Black areas of second tergite not or barely fringed with rufous. Pro-
notum and head scarcely rufous. Wings yellowish-gray, very slightly
purplish; stigma and veins russet-brown var montanus.

15. Large (fore wing 18 to 22 mm. long). Almost wholly and uniformly

light ferruginous ; only first tergite with narrow yellow margin
(sometimes lacking) ; propodeum and postscutellum sometimes
marked with yellow. Wings purplish-black; stigma and veins black

var. ruhiginosus.

Not uniformly light ferruginous and often smaller, or else several ter-
gites with yellow apical margins 16.

16. Thorax and head mainly light mahogany-brown, more or less blotched

with black, practically without yellow markings. Abdomen mainly
blackish, usually without yellow markings; first and second tergites
often partly ferruginous; first tergite sometimes with narrow yellow

margin. Large (fore wing 14 to 20 mm. long) var. metricus.

Blackish color of abdomen not contrasting with the light reddish-brown
of head and thorax. Yellow markings usually more extensive, par-
ticularly on the abdomen 17.

17. Abdomen very extensively yellow, often almost entirely so ; second ter-

gite either mostly yelloAV, or extensively yellow on the sides, or with

two yellow lateral spots 18.

Abdominal tergites only with apical yellow margins ; second tergite
without lateral yellow spots and not or only narrowly yellow on the
sides. Wings slightly infuscated, somewhat purplish 20.

18. Yellow markings of thorax much reduced or almost lacking; mesonotum

and propodeum without yellow stripes. Wings amber-yellow; stigma
and veins russet. Large (fore wing 16 to 18 mm. long)

var. centralis.

20

Journal New York Entomological Society [Vol. XLVIII

Yellow markings of thorax extensive; propodeum nearly always and
mesonotum often with yellow stripes 19.

19. Small (fore wing 12 to 15 mm. long). Wings amber-yellow. (Cali-

fornia) var. analieimensis.

Medium-sized or large (fore wing 15 to 18 mm. long). Wings slightly
smoky and somewhat purplish. (Southern United States, from Ari-
zona and Utah to Florida) var. apachus.

20. Yellow markings scanty and narrow, often reduced to hind margins of

pronotum and of first tergite. Thorax mostly ferruginous, with
blackish blotches on pleura and mesonotum; propodeum without or
with narrow yellow stripes. Large or medium-sized (fore wing 15
to 20 mm. long) var. nestor.

Yellow markings extensive on thorax and abdomen; as a rule most
tergites broadly margined with yellow; propodeum almost always
with broad yellow stripes 21.

21. Large (fore wing 15 to 20 mm. long). Black areas of abdomen absent

or reduced to extreme bases of tergites var. hellicosus.

Smaller (fore wing 10 to 18 mm. long). Third and fourth tergites
usually extensively black var. hunteri.

1. P. fuscatiis, typical. — Specimens agreeing in every detail
with Fabricius’ description and type, now at the University Mu-
seum in Kiel,^ are by no means common. They are occasionally
found, together with various slight variants, in southern Massa-
chusetts (for instance, on Cape Cod and Martha’s Vineyard),
Rhode Island, Connecticut, southern New York (particularly
Long Island), New Jersey, Pennsylvania, Virginia, West Vir-
ginia, North Carolina (Southern Pines; Raleigh), Ohio (Colum-
bus; Cleveland), Illinois (Havana), Michigan, Indiana (Bloom-
ington), southern Iowa (Decatur Co.), and eastern Kansas (Blue
Rapids). I have not seen typical fuscatiis from Canada, and
what has been recorded from there under that name was probably
all somewhat aberrant var. pallipes. Moreover, it intergrades
with var. nestor and var. pallipeSy as it covers some of the terri-
tory of both. It is particularly hard to draw a line between typi-
cal fuscatus and var. nestor, and it might be more rational to
unite these two, at any rate.

2. P. fuscatus var. analieimensis Provancher. — This color form
is peculiar to California, where it occurs as far north as 38° N.
latitude.

4 Dr. Olaw Schroder kindly compared a series of North American specimens
with Fabricius’ type.

Mar., 1940]

Bequaert: Polistes

21

3. P. fuscaUis var. apachus de Saussure. — P. texanus Cresson
I regard as a synonym. This form is common in Texas, western
Oklahoma (Woodward Co.; Grand), New Mexico, Arizona, south-
western Colorado and southern Utah ; occasionally elsewhere
(Henry Co., southeastern Iowa; Ft. Lauderdale, Florida). I
have also seen it from the States of Sonora and Coahuilla, Mexico.

4. P. fuscatus var. aurifer de Saussure. — The common form of
the species throughout California, Oregon and the State of Wash-
ington. It extends well into southern British Columbia (Ver-
non; Kaslo; Chilcotin), as well as into Idaho (Warren, Idaho
Co. ; Waha ; Coeur d’Alene) and Nevada (Humboldt River) . This
wasp was introduced into the Hawaiian Islands many years ago
(first published record in 1884, by W. F. Kirby) and in 1928 it
was reported from the Society Islands.

5. P. fuscatus var. hellicosus Cresson. — A common form of the
southeastern United States, which, however, does not extend as
far north as metricus. I have seen it commonly from South Caro-
lina (Dewees Id.), Florida, Alabama, Mississippi, Louisiana and
Texas. It intergrades with var. apachus.

6. P. fuscatus var. centralis Hayward, 1933, Proc. Utah Ac.
Sci., X, pp. 141 and 143, PL IX, fig. 13. — This form appears to
be restricted to Utah and western Colorado. In southern Utah
it intergrades with apachus.

7. P. fuscatus var. (or subsp.) connectens, new.

Female and WorTcer. — Head mostly yellow; vertex, occiput, a broad, hour-
glass-shaped spot about the antennae, and outer half of outer orbits, black;
clypeus medially and outer orbits somewhat suffused with russet; antennae
ferruginous, flagellum somewhat infuscate above, scape with black spot near
upper tip. Thorax mostly black, with a few rufous blotches on dorsal areas
of pronotum and on scutellum and postscutellum, variable in extent; narrow
fore and hind margins of pronotum, scutellum and postscutellum anteriorly,
two broad median stripes and lateral spots on propodeum, and a spot on
mesopleura beneath base of wing, yellow; sometimes two yellowish or russet
longitudinal stripes or lines on mesonotum; tegulse russet, spotted with yel-
low. Legs mostly ferruginous; coxae black, streaked with yellow; femora,
tibiae and tarsi more or less yellow on the outer side. Abdomen extensively
yellow; a black spot, irregularly hourglass-shaped, in basal two-thirds of
first tergite, more or less edged with ferruginous ; a larger hourglass-shaped
black spot over basal three-quarters of second tergite more or less edged with
ferruginous which often forms spots at the extreme sides basad of the yellow ;
and median, irregularly quadrate black spots at base of succeeding tergites.

22

Journal New York Entomological Society [Vol. xlviii

often partly ferruginous; the sternites are black with broad yellow apical
margins and lateral spots. Wings subhyaline, suffused with amber-yellow;
veins russet.

Male. — Sometimes similar to the female, though with the face more exten-
sively yellow, without black about the antennse ; mesosternum and entire under
side of coxae yellow. Other males are more extensively ferruginous, being
then hardly separable from those of var. variatus.

Holotype: Badlands west of Grassy Butte, McKenzie Co., west-
ern North Dakota, female (N. A. Weber). — Allotype: same
locality, male (N. A. Weber). — Paratypes: Alberta: Medicine
Hat, several females and males (F. S. Carr; E. H. Strickland) ;
Lethbridge (E. H. Strickland) ; Mayberries (E. H. Strickland).
— North Dakota : Badlands west of Grassy Butte ; Junction of
Cannonball and Missouri Rivers, Sioux Co. (N. A. Weber) ; Me-
dora, one male (0. A. Stevens) ; Fargo (0. A. Stevens) ; Mandan
(0. A. Stevens) ; Medora (C. H. Waldron) ; Sentinel Butte (0.
A. Stevens) ; Mott (J. R. Campbell) ; Marmartli (0. A. Stevens) ;
Beach (R. L. Webster) ; Hague (M. van Soest) ; Trottem (A. C.
Fox). — Montana: Laurel, Yellowstone Co. — Wyoming: Torring-
ton, Goshen Co. (G. B. Fairchild). — Nebraska: Squaw Canyon,
Sioux Co. (H. G. Barber) ; Lodgepole, 4,050 ft., Cheyenne Co.
(H. A. Scnllen). — Colorado: Texas Creek, (J. C. Bradley);
Denver (J. Beqnaert) ; Maniton (J. Beqnaert) ; Clear Creek,
7,000 to 8,000 ft., Jefferson Co. (G. P. Engelhardt) ; vicinity of
Fort Collins (23 mi. up Little Pondre Canyon and Spring Can-
yon) (A. B. Klots) ; Boulder (E. R. Becker) ; Boulder Co. (C.
P. Custer) ; Berkeley. — Holotype, allotype and several paratypes
at Museum of Comparative Zoology, Cambridge, Mass. Paratypes
also in several other collections.

This color form seems to be characteristic of the northwestern
portion of the Upper Sonoran life zone. It probably will be
found also in parts of South Dakota and Kansas. Its occurrence
in southeastern Alberta is of considerable interest.® The region
is apparently an Upper Sonoran island surrounded by Transition
fauna. Professor Strickland informs me that several other south-
ern insects have been taken there. The solitary wasps he sent me
include Odynerus anmdatus var. geminus Cresson, 0. anorniis

5 A female of the var. connectens was also taken on a window pane in a
house at Edmonton, Alberta. This was certainly an accidental importation
by man. No Polistes was ever found in the open in that vicinity.

Mar., 1940]

Bequaert: Polistes

23

Say, 0. aldrichi Fox, 0. dorsalis (Fabricius), Pterochilus quin-
quefasciatus Say, and P. decorus Cresson.

The var. connectens is clearly transitional between the var.
aurifer and the var. variatus, and shows also some gradation to
the varieties apachus, montanus and utahensis. It is most typical
in North Dakota.

At Medicine Hat females were taken in flight during August
and September and males early in September. On November 27
a female was found hibernating “under clod.” On April 17 a
female was caught at the flowers of Buffalo Berry, LepargyrcEa
canadensis (Linnaeus).

8. P. fuscatus var. flavus Cresson. — This is the extreme xanthic
variation of the species, common in Arizona, southern Nevada
(Las Vegas, Clare Co.), southern Utah (St. George, Washington
Co.; Hurricane, Washington Co.), parts of New Mexico (Rio
Grande Canyon, south of Taos; Highrolls; Picacho), and western
Texas (Devils River near Comstock, Valverde Co.; Big Bend
Park, Brewster Co.). It should also occur in northern Mexico,
but I have as yet seen no Mexican specimens. Evidently charac-
teristic of the Lower Sonoran desert country, it intergrades to
some extent with var. apachus.

9. P. fuscatus var. (or subsp.) hunteri, new.

Average size smaller than most other forms of P. fuscatus. Length
(h. + th. + 1. 1 + 2) , of female and worker, 10 to 12 mm.; of male, 10 to 12
mm. Length of fore wing, of female, 10 to 13 mm. ; of male, 10 to 13 mm.

Female and WorJcer. — Variegated black and reddish-brown in fairly equal
proportions and with many bright yellow markings. Head mostly reddish-
brown, with the ocellar area of the vertex and hind face of occiput black, the
upper side of scape and of most of flagellum fuscous, but the flagellum not
ringed with black. Thorax black, with the pronotum mostly, four longitudinal
stripes on mesonotum, and most of scutellum and postscutellum, reddish-brown;
sometimes most of the mesonotum and blotches on mesopleura and sides of
propodeum reddish; or the mesonotum entirely black. Abdomen: first and
second segments mostly reddish-brown, usually with blackish base extending
triangularly behind; third and fourth more brownish-black to black, turning
reddish posteriorly near the yellow hind borders; fifth and sixth reddish-
brown to black. Legs mostly reddish brown, with black coxse and infuscate
bases of femora and apical half of hind tibiae; tibial spurs ferruginous. Yel-
low markings as follows : broad apical margin of clypeus ; lower inner orbits ;
entire outer orbits; narrow collar and hind margin of pronotum; broad
anterior fasciae on scutellum and postscutellum; a large spot on mesopleura

24

Journal New York Entomological Society [Vol. xlviii

beneath base of fore wing ; most of tegulse ; two broad longitudinal stripes and
valvulae of propodeum ; broad apical margins of all tergites and most sternites
(usually narrowed or interrupted medially on the sternites) ; lateral streaks
or spots on first tergite, more or less connected with the apical band; apices
of femora, outer side of tibiie and most of tarsi. Wings moderately yellow-
ish-russet, somewhat darker and slightly purplish over the radial cell; veins
and stigma russet.

Male. — Similar to the female, but the yellow usually more extensive, cover-
ing most of the face, the entire clypeus, the oculo-malar spaces and most of
outer orbits, the mandibles, the under side of the scape, most of the
prosternum and under side of coxae and femora.

Holotype: Paradise Key, Dade Co., Florida, female (Kichard
Dow). — Allotype: Cape Sable Road, 5 mi. W. of Dade Co. Line,
Florida, male (Richard Dow). — Paratypes: Virginia: Falls
Church. — Ohio: Hocking Co. — North Carolina: Southern

Pines ; Statesville ; Raleigh ; Wilmington ; Morgantown ; Kittrell ;
Kingsboro; Laurinbur; Havelock (Lake Ellis) ; Wadesboro;
Conove; Aberdeen; Willard; Liberty; Ft. Bragg; Winston-
Salem; Boardman; Fairmont. — South Carolina: Clemson Col-
lege.— Georgia: Billy’s Id., Okefeenoke Swamp; Roswell.^ — Flor-
ida: Matecumbe Key; Monticello; Paradise Key; Ft. Lauder-
dale ; Gulfport ; Tampa ; Tamiami Trail, Dade Co. ; St. Peters-
burg; Jacksonville; Stuart; Miami; Kelsey City; West Lake,
Dade Co. ; Lutz ; Clearwater ; Titusville ; Biscayne Bay ; Char-
lotte Harbor; Gainesville; Key Largo; Lower Matecumbe Key;
Orlando; Cape Sable; Coconut Grove; Crescent Grove. — Ala-
bama: Thomasville, Clark Co.; Mobile; Leroy, Washington Co.;
Biloxi ; Auburn ; Greenville. — Mississippi : Oxford ; Winona. —
Louisiana : Shriever, Terrebonne Co. ; Darrow, Ascension Co. —
Texas : Williamson Co. ; New Braunfels ; Dallas ; Richmond, Fort
Bend Co.; Carniso Springs; College Station (as prey of a robber
fly). — Oklahoma: Quinton, Pittsburg Co.; Nowata Co. — ^Mex-
ico: Mexico, D. F. ; Guadalajara, State of Jalisco; San Luis (de
Potosi?) ; Cuernavaca, State of Morelos; Valle del Murz. — Holo-
type, allotype and many paratypes at Museum of Comparative
Zoology, Cambridge, Mass. ; paratypes also in several other
collections.

This wasp is one of the most common insects of the southeast-
ern United States. It looks like a diminutive var. hellicosus,
with which it intergrades. In most cases it is readily distin-

Mar., 1940]

Bequaert: Polistes

25

guished from hellicosus by the small size and the predominance
of black on the third and fourth tergites. Although the smallest
hunteri are superficially very different from the other forms of
fuscatus, they have all the structural characters of the species.
In particular, the males always bear a minute tubercle on the
middle of the seventh sternite. In addition it passes gradually
into the var. hellicosus^ so that some specimens must be placed
arbitrarily. Some specimens of hunteri also resemble P. excla-
mans in color, but that species is structurally quite distinct (as
shown in the key). In American collections and publications,
this wasp is frequently called Polistes minor; but Palisot de
Beauvois’ wasp of that name came from Santo Domingo and it is
structurally quite a distinct species from P. fuscatus.^

10. P. fuscatus var. (or subsp.) maritimus, new.

Female or WorJcer. — Head: most of face, vertex and occiput ferruginous,
somewhat black around the ocelli and over a narrow streak on the vertex ;
clypeus, lower inner orbits into the ocular sinuses, oculo-malar space, cheeks
(except in upper hind area) and mandibles, yellow; scape mostly and under
side of flagellum ferruginous, upper side of flagellum black. Thorax :
pronotum, scutellum, postscutellum and propodeum almost entirely yellow;
pronotum and scutellum with small, dark ferruginous blotches; propodeum
with a narrow, longitudinal black streak in the median groove; meso- and
metapleura black with a series of yellow spots ; sternum black ; mesonotum
black with four narrow, longitudinal yellow stripes, two median extending
nearly the whole length, one short on each side above the tegula ; tegula yellow
with a ferruginous spot. Legs yellow, streaked with ferruginous along femora
and tibiae, the hind tibiae more infuscated; tarsi ferruginous orange. Ab-
domen mostly yellow, with the following blackish or dark ferruginous mark-
ings : a small, basal, irregular spot on first tergite ; narrow bases of most
of the segments (more blackish on second and more ferruginous on remainder),
that of second tergite expanding laterally and produced medially into an
hourglass-shaped spot with three narrow apical projections; faint ferruginous
lines set off yellow lateral spots from the apical margins on second to fifth
tergites; sixth segment mostly ferruginous. Wings strongly infuscated, with
pronounced purplish reflection, stigma and costa from base of wing to stigma,
russet; remainder of veins black. Length (h. -|- th. -f- 1. 1 -F 2) : 11 mm.; of
fore wing, 12 mm. *

Male unknown.

Holotype: Mangrove Cay, Andros Island, Bahamas, female,

6 The name hunteri, which I have adopted for this wasp, had been given to
it in manuscript, at the U. S. National Museum, presumably by Ashmead.

26

Journal New York Entomological Society [Vol. xlviii

May-June 1917 (Wm. M. Mann). — Type at Museum of Com-
parative Zoology, Cambridge, Mass.

Apparently a small insular form related to var. hunteri, of the
southeastern United States, but much more extensively marked
with yellow. It resembles superficially the West Indian P. poeyi
Lepeletier and true P. minor Palisot de Beauvois. From the
former it differs in the presence of the mesepisternal suture and
the stronger striae of the propodeum ; from the latter in the shape
of the clypeus and the striation of the propodeum. Since the
male is as yet unknown, the var. maritimus is only tentatively
placed under P. fuscatus.

11. P. fuscatus var. metricus Say. — There is a possibility that
this wasp was described before Say as Vespa geniculata Graven-
horst (1807, Vergleichende Uebersicht Linn. u. einig. neuern
Zool. Syst., p. 276; without locality). Until the type can be ex-
amined critically, I hesitate to adopt the name. — This form is
common in the southeastern and central United States. I have
seen it from New Jersey (Cape May), South Carolina, Georgia,
Florida, Alabama, Mississippi, Louisiana, eastern Texas (Wil-
liamson Co.; Brazos Co.; Dallas; Weslaco), Oklahoma, Arkansas,
Tennessee, Missouri, Kentucky, southern Ohio (Pickaway Co. ;
Perry Co.; Flocking Co.; Buckeye Lake, Licking Co.), southern
Indiana, Illinois (Chicago; New Columbia; Putnam; Urbana),
Iowa (Mt. Pleasant), and Nebraska (Lincoln; Waverly). It
should occur also in eastern Kansas. The distribution is much
the same as that of P. canadensis var. annularis, which is more
or less homeochromic and sometimes confused with it. The var.
metricus extends much farther north in the valleys of the Mis-
sissippi, Missouri and Ohio than along the eastern seaboard. In
certain areas it intergrades with other forms, particularly with
var. nestor.

12. P. fuscatus Ysa\ (orsubsp.) montanus, new.

Female and Worher. — Closely related to var. utahensis Hayward, from
which it differs mainly in the presence of yellow stripes on the propodeum.
Head: mostly black; lower third and broad sides of clypeus, lower inner
orbits up to ocular sinuses, two streaks above bases of antennae, a spot on
upper outer corner of oculo-malar space, and much of mandibles, yellow;
middle and upper part of clypeus (except the upper margin), blotches on
oculo-malar space, a narrow streak on outer orbit, ocular sinuses, margins of

Mar.; 1940]

Bequaert: Polistes

27

mandibles (except the black teeth), and antennae (except a black streak on
upper side of scape), ferruginous. Thorax: black; tegulae ferruginous,
spotted with yellow; narrow anterior and posterior margins of pronotum,
two small spots near anterior margin of scutellum, narrow anterior margin
of postscutellum, a short vertical streak in upper part of mesopleura (be-
neath base of wing), and two longitudinal stripes on propodeum (variable
in extent, sometimes barely indicated), straw yellow; none of the specimens
seen have any ferruginous on the thorax. Legs black; extreme apices of
femora ferruginous on hind legs, ferruginous and yellow on fore and mid
legs; fore and mid tibiae ferruginous with yellow outer streak; hind tibiae
black, blotched with ferruginous and with yellow outer streak ; tarsi mostly
yellow, ferruginous toward apices; tibial spurs and claws ferruginous.
Abdomen: mostly black; moderately large apical margins of tergites 1 to 5
and sternites 2 to 4 (somewhat wavy and more or less interrupted medially)
and free spots of moderate size and irregular shape on the sides of tergites
1 to 5 (very small and sometimes absent on first tergite), placed rather
close to hind margins, straw yellow; sixth segment ferruginous, the tergite
with two small, basal yellow spots; the black of the fifth tergite is usually
tinged with ferruginous and that color may in some specimens surround
more or less the free yellow spots of the other tergites. Wings moderately
infuscated, with a yellowish tinge and marked purplish reflection, especially
in the apical third; veins dark brown; stigma and costa from base of wing
to stigma, russet.

Male. — Differs only slightly from the other sex, with Avhich it is readily
associated. As usual, the yellow color covers the entire clypeus, face and
oculo-malar spaces, and most of the under side of thorax, legs and abdomen ;
the upper side of thorax and abdomen are as in the female.

Holotype: Hamilton, Ravalli Co., Montana, female (W. J. Jel-
lison). — Allotype: same locality, male (W. J. Jellisoii). — Para-
types: Montana: Hamilton, several females (W. J. Jellison) ;
Ravalli Co., one female (AV. J. Jellison) ; AVillow Creek, Ravalli
Co., four females (AV. J. Jellison) ; Blodgett Canyon, Ravalli
Co., one female (AV. J. Jellison) ; Burch Creek, Ravalli Co., one
female (AV. J. Jellison) ; Lake Roiian, Lake Co., one female (R.
D. Eiclimann) ; Baird, Missoula Co., one female (A. L. Me-
lander). — Idaho: AVallace, Shoshone Co., one female; Pine Creek,
Shoshone Co., 2,600 ft., one female. — Oregon: Corvallis (J.
Schuh) ; 23 mi. A\^. of Halfway, 4,700 ft. (R. E. Rider) ; Pine
Creek Canyon, 4,600 to 5,300 ft.. Baker Co. (R. E. Rider) ; Uma-
tilla, Umatilla Co., two females. — Holotype, allotype, and several
paratypes at the Museum of Comparative Zoology, Cambridge,
Mass. ; paratypes also in several other collections.

28

Journal New York Entomological Society [Vol. xlviii

Like the var. utahensis, this is an extreme variant of var.
aurifer, seemingly connecting the two. In Oregon, where it oc-
curs in certain localities together with aurifer, intergrades are
also met with. As it appears to be the dominant color form of
the species in Montana, it would seem to deserve a name.

13. P. fuscatus var. (or subsp.) neotropicus, new.

Closely related to var. hunteri, of which it has about the size, but more
extensively marked with yellow, the propodeum being almost entirely of
that color.

Female and WorJcer. — Head and antennse ferruginous-red, the vertex more
or less blackish about the ocelli ; flagellum somewhat inf uscate above ;
clypeus as a rule entirely, broad lower inner orbits (as far as ocular sinuses),
oculo-malar spaces, broad outer orbits and most of mandibles, pale yellow.
Thorax ferruginous-red on j^ronotum and mesonotum, black on sternum and
pleura (the mesonotum rarely more or less black) ; broad fore and hind
margins of pronotum, tegulse, small spots on sides of mesonotum, most of
scutellum and postscutellum, two spots on the mesopleura (one above the
other), and most of propodeum (except a narrow median black line and
broader black sides), yellow. Abdomen ferruginous-red dorsally, at most
slightly inf uscate at the base of the third and fourth tergites, ventrally
blackish-brown; apex and sides of first tergite very extensively (leaving only
a ferruginous oval patch in the center), apical margins of succeeding ter-
gites (narrow medially, much widened laterally and continued along the
sides of the second tergite) and narrow spots in the hind corners of second
and third sternites (sometimes continued as a complete apical margin on
second sternite), yellow; sixth segment entirely ferruginous. Legs black,
blotched with ferruginous; apical fourth to half of femora above, most of
fore and mid-tibiae, basal two-thirds of hind tibiae, and most of tarsi, yellow;
tibial spurs ferruginous. Wings moderately tinged with yellowish-russet,
somewhat darker and slightly purplish over radial cell; veins and stigma
russet.

Male. — Similar to the female, with which it is readily associated. Yellow
more extensive on the legs, the under side of fore and mid coxas being
mostly of that color. Two yellow spots on sternum.

Length (li. + th. + 1. 1 + 2), of female and worker, 10 to 12 mm. ;
of male, 11 to 12 mm. Length of fore wing, of female and
worker, 10 to 13 mm. ; of male, 11.5 to 13 mm.

Holotype: Puerto Castilla, Eepublic of Honduras, female (J.
Bequaert). — Allotype: Prieta, Republic of Honduras, male (J.
Bequaert). — Paratypes: Puerto Castilla, three females; Prieta,
one female. — Guatemala^ without more definite locality, five
females and two males. — All types at Museum of Comparative
Zoology, Cambridge, Mass.

Mar., 1940]

Bequaert: Polistes

29

14. P. fuscatus var. nest or (Fabricius). — This form is more
extensively ferruginous over the abdomen than typical fuscatus,
with which it intergrades. It is found occasionally on Long
Island (New York) and in New Jersey, more commonly in south-
ern Pennsylvania, Maryland, Virginia, North Carolina, South
Carolina, Georgia, Florida (Tangerine; Monticello), Alabama
(Mobile), Kentucky, eastern Texas, southern Michigan (Midland
Co.), southern Illinois (New Columbia; Herod), Indiana (Bloom-
ington) and Ohio. There are many intergrades not only with
typical fuscatus, but also with the var. variatus. All specimens I
have seen from the Bermudas, Jamaica and Barbadoes were of
the var. nestor, rather than typical fuscatus.

15. P. fuscatus var. pallipes Lepeletier. — P. exilis de Saussure
appears to be a synonym. It is the extreme melanistic form of
the species, characteristic of the northeastern part of the range,
throughout the Transition life zone. I have seen it from south-
ern Quebec (St. Jerome; Montebello; La Trappe; Montreal;
Rigaud; Outremont; St. Remy ; Queen’s Park, Aylmer), south-
eastern Ontario (Frank’ Bay, Lake Nipissing; Gull Lake; Pelee
Id.), Maine, Vermont, New Hampshire, Massachusetts, Rhode
Island, Connecticut (Colebrook), New York (including Staten
Island and Long Island), New Jersey, Pennsylvania (Bartons-
ville ; Pittsburgh ; Braddock ; Green Lane, Montgomery Co. ;
Mauch Chunk), West Virginia, Maryland (Plummers Id.), Ohio
(Mahony Co.; Marietta), Indiana (Winona Lake), Illinois, Mich-
igan (common), and northern North Dakota (Towner). It will
probably also be found in Wisconsin and Minnesota. The north-
ernmost locality is Towner, in about 48° 30' N. In the southern
part of the range is intergrades with nestor and typical fuscatus.
The extent of pale yellow markings varies greatly. Some speci-
mens have only a narrow apical margin on the first tergite, nar-
row margins on the pronotum, the tegulse and two lines on the
propodeum of that color. There are all passages to the other ex-
treme, with broad apical margins on all tergites (often continued
along the sides), yellow margins of pronotum, scutellum and post-
scutellum, two broad stripes on propodeum (in one case even
four stripes), a spot on mesopleura, and markings on the head.
A few specimens may even show small, free, yellow lateral spots
on the second tergite, thus simulating var. variatus and var.

30

Journal New York Entomological Society [Vol. xlviii

montaniis; they lack, however, the rufous blotches of the former
and. the yellowish-gray wings of the latter.

16. P. fuscatus var. ruhiginosus Lepeletier. — Y espa nigripennis
Degeer (1773) may have been this wasp, and if this is the case,
Degeer’s name will take precedence over Lepeletier ’s. This is a
common wasp of the southeastern United States. I have seen it
from southern Ohio (Blue Creek; Adams Co.; Lawrence Co.),
Pennsylvania (Coraopolis), Virginia (Fredericksburg), North
Carolina (Raleigh; Southern Pines), South Carolina, Georgia,
Florida, Alabama, Mississippi, Louisiana, eastern Texas (Comal
Co. ; Fort Bend Co. ; Brazos Co. ; Bastrop Co. ; Kaufman Co. ;
College Station ; Williamson Co. ; San Antonio ; New Braunfels ;
Austin), Arkansas, Oklahoma, Missouri, Kentucky, Tennessee,
and southern Illinois (Olive Branch). It is better defined than
other color forms of the species and shows little intergradation.

17. P. fuscatus var. utahensis Hayward, 1933, Proc. Utah Ac.
Sci., X, pp. 141 and 142, PL IX, figs. 1 and 3-9. — This color
form is known from Utah, Idaho, Nevada, Colorado, and Wyo-
ming (Lava).

18. P. fuscatus var. variatus Cresson. — This is a fairly common
form of the species in the Middle West: North Dakota (Fargo;
Sheldon), Minnesota (Sleepy Eye), Nebraska (Lincoln; Cam-
bridge), Iowa (common), Kansas (Manhattan; Wathena; Hays),
Missouri, Illinois, Indiana, southern Michigan (common), Ohio,
Kentucky (Buckeye; Mammoth Cave), Tennessee, Arkansas
(Spring Dale), Oklahoma (Stillwater; Woodward Co.; Grand)
and eastern Texas. I have also seen a few specimens from south-
ern New Jersey (Lakehurst) and New Mexico (Carlsbad). In
the northern parts of its range it intergrades with var. nestor, in
Iowa with typical fuscatus, and in Texas and Oklahoma with
var. apachus. Hayward’s records of variatus from Canada
(1933, Canad. Entom. LXV, p. 128) were, I believe, all due to a
confusion with other forms of the species. Those from British
Columbia referred to var. aurifer; the one male from Medicine
Hat, Alberta, was my var. connect ens; and the one male from
Nelles Corner, Ontario, probably an aberrant var. pallipes. It is
nevertheless, possible that the var. variatus may yet be found in
southern Ontario.

Mar., 1940]

Bequaert: Polistes

31

Postscript

The following remarks may prevent needless criticism. (1)
For want of space it was not possible to enumerate the many
museums and private collectors to whom I am deeply indebted
for material or other assistance. (2) For the same reason the
distribution of most forms is given by States only, except for the
new and rare forms, or when more definite localities may help to
define the geographical limits. (3) Structural characters could
not now be illustrated nor their variation discussed. (4) More
in particular, the male terminalia are not mentioned, because the
specific differences they show are slight and could not be described
adequately without figures.

32

Journal New York Entomological Society [Vol. xlviii

BOOK NOTICE

Fleas of Eastern United States. By Irving Pox, Ames, Iowa.

The Iowa State College Press, February, 1940. 9x6 inches.

vii + 191 p., 31 pi. $3.00.

This monograph on the fleas of eastern United States is a wel-
come addition to entomological literature. Many working ento-
mologists have little exact knowledge of the various species of
fleas and are apt to assume that a specimen is a certain species
because it was found on a particular host. There is no longer
aii3'" necessity for inconclusive identifications because Mr. Fox’s
work includes 55 species, in five families and 33 genera, found
east of the one-hundredth meridian, exclusive of Texas. A brief
introduction deals with the collection, preservation, morphology,
terminology and life-histories of fleas in general while the balance
of the book is devoted exclusively to keys to the suborders, fam-
ilies, genera, and species, together with descriptions of the males
and females of each species, references to the literature of each
species, detailed records of distribution, and eastern host records.
There is also a synonymic index, a host index and a selected bib-
liography together with 31 plates embracing 166 figures, mostly
of male genitalia and heads of males and females, which are of
systematic importance. It may surprise many to And that vari-
ous mammals are hosts to a half-dozen or more species of fleas.
For example, the dog is host to six species, the cat to five, and
Homo sapiens, that noble and self-important member of the order
Primates, to nine. Fleas are important in view of their implica-
tion in the transmission of diseases and because they parasitize
various mammalian and avian hosts. Entomologists and public
health technicians in the east will And in Mr. Fox’s book the
facilities for identifying the various eastern species, without hunt-
ing through all the literature and in addition they will no longer
have to be concerned with the inadequate and piece-meal bits of
descriptive matter that occur in the general text-books on ento-
mology.— H. B. W.

(Jour. N. Y. Ext. Soc ), Vol. XLVIII

(Plate I)

Mar., 1940]

Sherman: Fall

33

HENRY CLINTON FALL

Henry Clinton Fall was born on Christmas Day, 1862, at
Farmington, New Hampshire, and spent his boyhood at this place.
He graduated (B.S.) from Dartmouth in 1884, and received from
the same institution in 1929 the honorary degree of Doctor of
Sciences. He went to California in 1889, taught at Pomona High
School 1892-1896, and was head of the Science Department
(housed in its own separate building) of the Pasadena High
School from 1896 through 1917. After his retirement he re-
turned East and made his home with his dearly loved sister and
her husband, Mr. and Mrs. Carl Adams Richmond, at Tyngsboro,
Massachusetts, until his death, November 14, 1939. He never
married.

He was a corresponding member of the American Entomolog-
ical Society, and a Fellow of the American Academy of Arts and
Sciences, of the American Association for the Advancement of
Science, and of the Entomological Society of America.

He began to collect beetles at the age of fourteen and soon after
his arrival in California, in 1889, was busily engaged in collecting
and in exchanging specimens with other collectors. At the same
time he was studying his beetles diligently and naming those sent
to him for identification by many other collectors, the writer one
of them. He continued this great service up to the very end. A
few years ago there came to him, as a resulting gift of apprecia-
tion, the immense collection of one of our best known collectors
(still living), Charles Liebeck, of Philadelphia.

Fall offered no papers for publication, however, until he was
thoroughly qualified, and he was almost thirty-five years old when
his first descriptions were published in the October, 1897, issue
of the “Canadian Entomologist” in a paper entitled “A List of’
the Coleoptera of the Southern California Islands with notes and
descriptions of new species.” His final paper, “The North
American Species of Nemadus,” appeared in 1937 in the Journal
OF THE New York Entomological Society. The Leng “Cata-
logue of the Coleoptera of America, North of Mexico,” and its
four Supplements, list the titles of one hundred and thirty papers

34

Journal New York Entomological Society [Vol. xlviii

published by Fall during this period of forty years. Fifteen of
these appeared in the pages of this Journal. Altogether Fall
described 1,453 new species of North American Coleoptera and 37
new genera. This total, though exceeded only by Leconte, Horn,
and Casey, is far less important than the manner in wFich these
descriptions were prepared and presented after the most pains-
taking study of the literature, of the Leconte and Horn types,
and of all available material. As a result very few of his species
have ever been challenged : what little synonymy is recorded, is
of his own discovery.

He did not confine his studies to any particular family but
demonstrated his wide knowledge of the entire order of Coleop-
tera in giving monographic treatment to a great number of diffi-
cult groups, such as Acmasodera (1899), Agabus (1922),
Agathidium (1934), Apion (1898), Bruchus (1910), Coelambus
(1919), Collops (1912), Diplotaxis (1909), Gyrinus (1922),
Hydroporus (1923), Hymenorus (1931), Lathridiidse (1899),
Malthodes (1919), Pachybrachys (1915), Podabrus (1928),
Ptinidae (1905), etc. Among his writings are three faunal lists
of unusual value, “List of Coleoptera of Southern California”
(1901), “Coleoptera of New Mexico” (with Cockerell, 1907), and
a list of the species taken by him in Alaska in 1924, during his
trip to that region with the Richmonds. All his papers relate
exclusively to the fauna of North America ; all are taxonomic.

Early in life he had the good fortune to make the acquaintance
of Frederick Blanchard. Both of them were not only skillful and
tireless collectors, but serious students as well, and they became
the closest of friends. Soon after Blanchard’s death (November
12, 1912) the Richmonds bought the Blanchard place at Tyngs-
boro, and Fall lived with them after he came back East in 1918.
Their home continued to be, more and more, a shrine for students
^of Coleoptera to visit, as it had been during Blanchard’s lifetime.
It was indeed a delightful home to visit, hospitable, harmonious,
inspiring. Seldom does one family, under one roof, have so much
of interest to offer its guests. In this home there were three work
rooms of varied sorts : one for Mr. Richmond, former patent
attorney for the American Telephone and Telegraph Company;
another for Mrs. Richmond, an accomplished genealogist ; and the

Mar., 1940]

Sherman: Tall

35

third Dr. Fall ’s own ‘ ‘ den ’ ’ containing his library, his collection
of both native and exotic beetles, and his other collections of
butterflies, of postage stamps, and of post marks. The deepest
attachment existed always between brother and sister, and he was
blessed indeed to enjoy the loving devotion of this sister at all
times and under all circumstances. The varied interests and
hobbies of the three members of this household never precluded
time and zest for other activities, and friendly discussions en-
joyed by all in common. The Shermans were welcomed many
times to this happy mecca on their way to and from Kandolph,
New Hampshire. Our last stop-over was in May, 1939, on our
return from a week-end in the White Mountains. The hurricane
of September, 1938, wrought great havoc at Tyngsboro, destroy-
ing nearly every tree on the Richmond acres, and the family were
naturally still greatly depressed by this fact, but all w^ere full of
thankfulness that they themselves and their house had escaped
injury. Fall was still as interested as ever in his beetles and
much pleased to receive, while we were there, a letter from Dr.
Reinhard announcing the sending of some of his newly described
species of Texas Phyllophaga.

In 1916 or thereabouts, at my earnest request. Fall assumed
the task of doing something about our much neglected Dytiscid^e.
Chris. H. Roberts and I had been collecting them for a long time
with great enthusiasm and success, but, at least in my own case,
with very little actual study. Fall made various trips to Mount
Vernon making extensive selections from my collection, and as-
sembled countless other specimens from various sources. With
all this material and his own collection before him, he studied at
length our species of the family, in connection with the Leconte
and other types, and many of Dr. Sharp’s specimens loaned by
the 'British Museum. Revisions of the genera Coelambus, Agabus
and Hydroporus and, later on, of Ilybius, resulted and about 100
new species of the family were thus made clearly knowm. At our
various meetings during the course of these studies, besides the
time spent by Fall in very deep application to his entomological
problems, there were various pleasant relaxations and games, in
which we could all participate, and in which Fall was always an
enthusiastic and skillful contestant.

36

Journal New York Entomological Society [Vol. XLVIII

As was expected by bis friends, and following Blanchard’s
example, Fall bequeathed his collection with all his types — the
largest private collection of beetles in the United States — to the
Harvard Museum of Comparative Zoology at Cambridge, where
he had spent so much time studying the Leconte types.

A letter from Charles W. Leng pays this tribute of apprecia-
tion to Dr. Fall ’s life work : —

‘‘Upon the foundation laid by Leconte and Horn, he
continued for forty years with rare conservative judgment
the work of making known our beetle fauna. In the uni-
form excellence of his descriptions and tables he was the
greatest of recent authors.”

John D. Sherman, Jr.

Mar., 1940]

Weiss: Death-Feints

37

THE DEATH-FEINTS OF SITOPHILUS GRANARIUS
LINN., AND SITOPHILUS ORYZiE LINN.

By Harry B. Weiss

The death-feigning instinct, which is well developed in many
insects and which occurs frequently in many of the Coleoptera,
is quite apparent in Sitophilus granarius Linn., and Sitophilus
oryzce Linn., in their reactions to outer stimuli. The following
notes constitute a summary of observations made recently on the
“reflex immobilization” of numerous specimens of these species,
both of which were supplied to me by the kindness of Dr. William
Moore.

Sitophilus granarius Linn.

The granary weevil, while crawling about over grains of corn,
will not feign death if the grains are disturbed so as to touch the
weevils only slightly. However, if the disturbance takes place
persistently and with a little force, the death-feint occurs. And
it takes place instantly when the weevils are touched with the tip
of a pencil, or a pair of forceps. These weevils, which have only
vestiges of wings, appear to be quite sensitive to mechanical
stimuli.

The death-feint posture is unlike that of death. In general,
the following attitude is assumed. The first pair of legs extends
anteriorly close to the body and parallel to the head. The an-
tennae are also parellel with and close to the beak. The femora
of the second pair of legs extend diagonally anteriorly, with the
femora, tibias and tarsi folded upon each other. The femora of
the third pair of legs extend posteriorly diagonally, with the
femora, tibiae and tarsi folded upon each other. The legs are all
held close to the body. Sometimes the femora, tibise and tarsi of
the flrst pair of legs are folded upon each other instead of being
stretched out. Sometimes the antennae stick out at various an-
gles. And sometimes the legs do not all assume the rigid and
flxed positions outlined above, but stick out grotesquely, or assume
somewhat different positions. Apparently the intensity of the
outside stimulus is important in determining the degree of
rigidity.

38

Journal New York Entomological Society [Vol. xlviii

The death-feint response in 8. granarius may be brought about
by various mechanical stimuli, such as turning the specimen over
on its back, by picking it up with fingers or forceps, by dropping
it from a height of two or three inches, by picking it up by one
of its legs, by touching it almost anywhere on its body, although
its dorsal surface is not so sensitive as its ventral one, by breath-
ing upon it, by squeezing the antennse and snout together, by
pressing upon the ventral surface of the thorax, by pressing the
sides of thorax and sometimes by vibrating the medium upon
which it is resting. As a rule, death-feints, when the specimens
were on their backs, lasted longer than when they were on their
ventral surfaces, but there were many exceptions to this gen-
eralization.

The termination of the death-feint may be gradual or sudden.
Usually the antennae move first, then the first pair of legs and
then the remaining pairs. In a gradual termination this sequence
is easily observable. In a sudden termination, the recovery
movements of these appendages appear to take place simul-
taneously.

During the course of the observations, the relative humidity
varied from 36 to 40 per cent, the daylight or brightness from
4 to 150 candles per square foot, as measured by a Weston Pho-
tronic Exposure Meter, and the temperature from 68° P., to 94°
F. As the relative humidity was almost constant, no further
reference will be made to it. The brightness of the field in which
the death-feint reactions were noted may also be neglected. Ex-
cept for an instance of brightness approaching 150 candles per
square foot, with a temperature of 73° F., the observations were
made in fields where the brightness ranged from 4 to 65 candles
per square foot, and this range will be immediately recognized as
rather low. In all cases the degrees of brightness as noted ap-
peared to have no effect upon the duration of the death-feint or
upon the behavior of the insects. Temperatures, however, in-
fiuenced the duration of the death-feint, as will be shown.

In the following Table I, there is shown the average duration
in seconds of successive death-feints at various temperatures, of
twenty-six weevils. The interval between the death-feints was
approximately five seconds in duration. The longest number of
successive death-feints was 104, after which the weevil refused

Mar., 1940]

Weiss: Death-Feints

39

to react. There was considerable variation in the duration of
individual death-feints but the general tendency was for the
death-feints to become shorter as they successively increased in
numbers. It may be noted in Table I that as the temperature
became higher the average duration of the death-feints declined.
At temperatures of 88° F., and above, many weevils refused to
feign death at all. The death-feints that did occur were short
and recovery was rapid. Many refused to perform after two or
three successive death-feints lasting a second or two. When the

TABLE I

AVERAGE DURATION OF SUCCESSIVE DEATH-FEINTS OF SITOPHILUS GRANARIUS
AT VARIOUS TEMPERATURES

Temperature
in degrees F.

Number of successive
death feints

Average (arithmetic)
duration in seconds

68

48*

15

68

72*

27

68

49*

58

70

21

38

70

16

34

70

14

29

70

18

36

70

11

40

72

104*

11

72

52*

23

73

11

35

73

4

37

74

48*

18

80

19

7

80

16

15

80

16

19

80

18

25

88

12*

6

88

29*

11

88

5* "

3

88

2*

2

88

3*

1

94

2*

1

94 •

27*

4

94

6*

2

94

2*

2

* Indicates termination of death-feints.

40

Journal New York Entomological Society [Vol. XLVIII

death-feint was induced at a low temperature, it was ended imme-
diately when the weevil was placed under the influence of the
higher temperatures.

In view of the variation in the duration of successive death-
feints the average time in seconds does not show how long or short
some of the death-feints lasted. This omission is supplied in
Table II. Depending upon the temperatures the death-feints
lasted from one to as many as 207 seconds.

RANGE IN duration OF SUCCESSIVE DEATH-FEINTS OF SITOPHILUS GRANARIUS

TABLE II

AT VARIOUS TEMPERATURES

Temperatures
in degrees F.

Eange in seconds

68

68

68

Ito 60
2 '' 80
2 207

70

70

70

70

70

3 to 118
140
65

5 122

4^‘ 95

72

72

73

73

74

Ito 71
2 79

11 100
25 59
1‘‘ 56

80

80

80

80

Ito 14
4 ‘‘ 29
4 ‘ ‘ 45
7“ 89

88

88

88

Ito 10
2 '' 27
1 7

94

94

94

94

1 to 2
1 6
4

1“ 2

Mar., 1940]

Weiss: Death-Feints

41

The frequency distribution of the variations in the length of
the death-feint at different temperatures is shown in Table III.
At a temperature of 68° F., most of the recorded death-feints
lasted from 1 to 30 seconds. Many longer death-feints occurred
at this temperature but they were comparatively infrequent. At
72° F., most of the death-feints endured from 1 to 20 seconds,
and at temperatures of 88° F., and above, the durations were
from 1 to 10 seconds. Although the figures in Table III present
some inconsistencies and do not indicate a fixed rate of progres-
sion from long to short death-feints as the temperatures increased,
they do show that there is some correlation between the factors
involved.

It is my belief that the variations in the durations of the death-
feints, within certain temperature groups (except 88° and 94°)
are due principally to variations in the intensities of the outside
stimuli. It is impossible, without special apparatus, for an ex-
perimenter to apply mechanical stimuli, time after time, that do
not vary in intensity. During the course of the work, it was

TABLE III

FREQUENCY DISTRIBUTION OF VARIATIONS IN THE DEATH-FEINT DURATIONS OF
SITOPHILUS GRANARIUS AT VARIOUS TEMPERATURES

Length of
death-feint
in seconds

Temperature in degrees F.

68

70

72

73

74

80

88

94

1-10

30

13

72

0

15

24

37

37

11-20

34

11

39

2

16

28

10

21-30

38

18

23

8

7

9

4

31-40

19

11

13

1

8

3

41-50

17

5

3

1

1

4

51-60

13

9

2

2

1

0

61-70

4

4

2

0

0

71-80

7

1

2

0

0

81-90

1

1

0

1

91-100

0

2

1

101-110

1

1

111-120

1

1

121-130

2

2

131-140

1

1

141-150

Over 150

1

42

Journal New York Entomological Society [Vol. XLVIII

noted that long death-feints seldom resulted from slight stimula-
tions. On the other hand, deliberate rough treatment such as
pinching the front legs and beak together very firmly, frequently
resulted in death-feints of comparatively long duration. In such
cases the appearance of the weevil, with its legs drawn tightly
to its body, indicated that the muscles controlling locomotion were
in a tight state of contraction. As this state involved the an-
tennge also, the ‘‘hypnosis’’ probably extended throughout the
nervous system. When the stimulus was slight, the muscular con-
traction was frequently not strong enough to draw the legs close
to the body and they stuck out, sometimes at various angles. In
most of these cases the death-feint was short and recovery was
rapid, although in a few instances such “weak” death-feints
lasted fairly long.

In order to determine how long a death-feint would last under
prolonged stimulation, fifteen weevils were induced to feign
death, and were then continually stimulated by being moved
around and by pressure applied to the ventral surface of the
thorax by the tip of a pencil or a pair of forceps. The durations
of these death-feints in seconds at a temperature of 70° F., were
170, 300, 120, 28, 99, 434, 630, 635, 290, 1,290, 847, 2,160, 916, 434,
420, 384. The shortest was 28 seconds and the longest 2,160
seconds, or 36 minutes. The average length of the death-feint
for the 15 individuals was 591 seconds, or almost 10 minutes.

Sitophihis oryzce

Rice weevils in infested grain crawl over each other and allow
parasites to crawl over them, seemingly without any desire to
feign death. However, if touched by something foreign, death-
feints occur, but they are of short duration. On the whole this
species does not seem as sensitive to outside stimuli as the granary
weevil.

In the death-feigning attitude, the distal ends of the femora of
the first pair of legs extend forward diagonally and are pressed
against the base of the head. The femora of the second pair of
legs extend forward diagonally and are held close to the body.
The femora of the third pair of legs extend posteriorly diagonally,
close to the body. In all cases the femora, tibiae and tarsi are

Mar., 1940]

Weiss : Death-Feints

43

folded upon each other, and held close to the body. The antennae
are parallel and close to the beak. In some cases the femora of
the second pair of legs extend diagonally, posteriorly instead of
anteriorly. Sometimes the antennae participate only slightly in
the muscular tension and do not assume a rigid attitude close to
the beak. Frequently the tibiae and tarsi are not folded close to
each other but extend at wide angles from each other, although
held close to the body. In other cases the legs stick out hori-
zontally from the body. The death-feints of the rice weevil are
comparatively brief. While assuming the death-feint readily if
placed on their backs and with pressure applied to the thorax,
they recover quickly. If handled and stimulated a few times
they refuse to do anything except make efforts to escape. They
are not very sensitive to outside stimuli. However, the death-
feint may be brought about by squeezing the sides of the thorax,
by pressing against the ventral surface of the thorax, by squeez-
ing the antennge and beak together or by dropping the weevil
from a height of one or two inches. If one method fails, fre-
quently another one will succeed. In some cases individuals are
even susceptible to a slight touch against the edge of the body.
All in all, it is a difficult insect to work with, especially if the
temperature is 75° F., or above, and many individuals refuse
to feign death at this and even lower temperatures, except when
squeezed or pressed quite hard.

During the observations with the rice weevil, the same relative
humidity, light and temperature conditions prevailed as were
recorded for S. granarius and the same conclusions apply to both
species.

Table IV shows the average duration in seconds, of successive
death-feints at various temperatures of twenty-four rice w^eevils,
the interval between death-feints having been approximately five
seconds. The longest number of successive death-feints was 29,
after which the stimulus no longer produced a reaction. Indi-
vidual death-feints varied considerably but the tendency was for
the duration to decline as the number of successive death-feints
increased. This table also shows that as the temperature in-
creased, the death-feints became shorter. At a temperature of
74° F., and above, many individuals refused to feign death at

44

Journal New York Entomological Society [Vol. XLVIII

all. When they did, the attitude of the legs was not tense ; they
were not held close to the body, and recovery was rapid.

TABLE IV

AVERAGE DURATION OF SUCCESSIVE DEATH-FEINTS OF SITOPHILUS ORYZ.3E AT
VARIOUS TEMPERATURES

Temperature
in degrees F.

Number of suc-
cessive feints

Average (arithmetic)
duration in seconds

68

16*

6

68

22*

12

68

8*

4

68

9

4

72

8*

4

73

8*

3

73

9*

12

73

3*

2

73

5*

4

74

12

7

74

17*

13

74

6*

6

80

3

1

80

5

2

80

29*

6

80

3

1

88

24*

4

88

2*

1

88

2*

1

88

4*

2

88

3*

1

94

10*

3

94

9*

2

94

2*

1

* Indicates termination of death-feints.

Tables V and VI deal with the range in duration of successive
death-feints and with their frequency distribution with relation
to various temperatures. Death-feints lasted from 1 to 40 sec-
onds depending upon the temperature and the greatest number
of them took place within the range of from 1 to 5 seconds. As
with the granary weevil, it is believed that the variations in
durations of the death-feints in Sitophilus oryzcB, within certain
temperature groups are due principally to variations in the
intensities of outside stimuli.

Mae., 1940]

Weiss: Death-Feints

45

TABLE V

RANGE IN duration OF SUCCESSIVE DEATH-FEINTS OF SITOPHILUS ORYZJS AT
VARIOUS TEMPERATURES

Temperature
in degrees F.

Eange in seconds

68

1 to 12

68

2 ‘‘

7

68

1“

10

68

1

10

72

1

10

73

1

13

73

2

2

73

5“

40

73

1 “

6

74

1“

36

74

1 “

11

74

2

19

80

2

80

1

4

80

1“

20

88

1

9

88

1 “

3

88

1 “

2

94

1“

5

94

1 ‘‘

3

94

1 ‘‘

1

TABLE VI

FREQUENCY DISTRIBUTION OF VARIATIONS IN THE DEATH-FEINT DURATIONS OF
SITOPHILUS ORYZ^ AT VARIOUS TEMPERATURES

Length of Temperature in degrees F.

death-feint

in seconds 68 72 73 74 80 88 94

1-5 27 6 15 15 28 29 21

6-10 13 2 6 9 9 6

11-15 4 2 3 1 ■

16-20 9 14 2

21-25 1 0 1

26-30 1 0 0

31-35 0 2

36-40 1 1

46

Journal New York Entomological Society [Vol. XLVIII

Under continued stimulation at a temperature of 70° F., only
a few could be induced to prolong their death-feints. Seven
weevils, stimulated continuously, remained immobile for the fol-
lowing numbers of seconds before recovering : 33, 19, 40, 11, 26,
3 and 75. The average time was 34 seconds.

Although Sitophilus granarius and Sitophilus oryzce behave
similarly in their reactions to contact stimuli under different
temperatures, there is a great difference in the durations of their
death-feints. In granarius it is brought about more readily and
the locomotion reflexes are inhibited for a greater length of time.
Both species exhibited the same nervous excitability after con-
tinued contact stimuli had brought about a condition in which
the death-feint was no longer produced. And in high tempera-
tures the reflex avoiding reaction took precedence over the
“instinct” to feign death.

Mar., 1940]

Comstock: Butterflies

47

BUTTERFLIES OF NEW JERSEY

A LIST OF THE LEPIDOPTERA SUBORDER RHOPALOCERA OCCURRING IN
THE STATE OP NEW JERSEY ; GIVING TIME OF PLIGHT, FOOD
PLANTS, RECORDS OF CAPTURE WITH LOCALITY AND DATE

By William Phillips Comstock

INTRODUCTION

The butterflies appeal to a great many collectors, though they
probably represent only about one per cent of the listed insects
of the state of New Jersey. Their numbers, as related to other
insects occurring in the state, may be judged from the few pages
devoted to them in the “Annual Report of the New Jersey State
Museum for 1909” containing the Report of the Insects of New
Jersey prepared by the late Professor John B. Smith. Here the
butterflies were quite adequately covered but the list is out of
print and at present copies are difficult to obtain.

The present list adds few to those known to occur thirty years
ago as butterflies are mostly obvious creatures and future addi-
tions to the state list will probably occur more as strays than as
permanent residents. The reason for this list is to record addi-
tional facts about some of the species, as to their w^ay of life, and
occurrence both for dates and new localities.

There are recorded in this New Jersey list 132 species and 39
sub-specifically named. In the list for New York State there are
130 species and 27 sub-specifically named. For Connecticut there
are listed 87 species and 11 sub-specifically named. In a list of the
butterflies of the District of Columbia there are 90 species and 13
sub-specifically named. A recent list for the state of Virginia
gives 133 species and subspecies.^

List of the Insects of New York, Mortimer Demorest Leonard,
editor-in-chief. Cornell University Agricultural Experiment Station, Memoir
101, 1926.

‘ ‘ Check List of the Insects of Connecticut, ’ ’ by W. E. Britton, Ph.D. State
of Connecticut State Geological and Natural History Survey, Bulletin No. 31,
1920.

‘ ‘ The Butterflies of the District of Columbia and Vicinity, ’ ’ by Austin H.

48

Journal New York Entomological Society [Vol. XLVIII

In Smith ’s ‘ ‘ List, ” as it is familiarly known, there are a num-
ber of localities mentioned which no longer exist as good collect-
ing areas. Notable among these are localities in northern New
Jersey, the Orange Mountain area, Paterson and Garrett Rock
and Snake Hill. The old, easily accessible, collecting spots about
the lower end of Greenwood Lake, Hewitt and Lake Hopatcong
are much impaired. The Jamestown area of pine barren terri-
tory and marsh has also been injured. The same is true of the
once wonderfully rich collecting area of Five-Mile Beach. These
localities were formerly extensively collected in and many of the
less common species recorded from them thirty or more years ago
are now scarce or absent. Nevertheless in the absence of other
records it has been necessary to make use of many records from
these places in this list. The dates are reliable and useful as
guides for search in other localities of similar nature, which have
not suffered so much from despoilation.

The arrangement of the species occurring in New Jersey fol-
lows that of several much used text-books upon the butterflies and
diverges but little from the arrangement used by Dr. Smith. The
numbers given are those of McDunnough’s list.^ For the iden-
tification of species there are so many excellently illustrated
books, notably ‘‘The Butterfly Book” by W. J. Holland, revised
edition 1931, that keys or descriptions seem unnecessary.

For convenience and brevity a system of presentation is
adopted in the statements concerning each species. The items to
be covered after the family designation are ordered as follows:
Genus — Species — Important synonym or marked change from
Smith’s “List” — American popular name — General locality —

Clark. Smithsonian Institution, United States National Museum, Bulletin
No. 157.

‘ ^ Preliminary List of the Butterflies of Virginia, ’ ’ by Austin H. Clark and
Leila P. Clark. Proceedings of the Biological Society of Washington. Vol.
50, pp. 87-91, June 22, 1937.

2 ‘ ‘ Check List of the Lepidoptera of Canada and the United States of
America,” by J. McDunnough, Ph.D. 1938.

In the Hesperiidae the arrangement is that of ‘‘The Hesperioidea of North
America,” by A. W. Lindsey, E. L. Bell and R. C. Williams, Jr. Denison
University Bulletin, Journal of the Scientific Laboratories, Vol. XXVI, April
1931.

Specific synonymy is corrected where essential.

Mar., 1940]

Comstock: Butterflies

49

Characteristic environment — Usual time of year when adults are
most numerous — Number of annual broods — Winters as ; egg,
larva, pupa, imago — Peculiarities of habit and remarks — Pood
plants — Recorded dates ; earliest to latest — Selected records ;
locality, date, collector — Subspecifically named forms.

It is not always necessary or possible to give information on
each item as outlined but each listing of a species follows this
general form and if items are omitted the succession of items
remains the same.

In many cases species are very local, confined to very small
areas where their particular food plant grows. The larvae of
some apparently feed upon only one kind of plant. Thus a spe-
cies may be spread over a considerable region but will be found
only in scattered small areas.

Eggs may be laid on the top of the leaf, underneath, singly and
in groups and masses. Some Lycaenidse lay eggs in flower-heads.
The larvae have varied feeding habits. Some are nocturnal feed-
ers, some myrmecophilous, one carnivorous. Some larvae feed
openly ; some hide by day in various ways, as in cover about base
of the plant, in ants’ nests about the lower stem, or in a protec-
tion of leaves spun together. Pupae are generally concealed in
various ways and are difficult to find. Mostly they are of a shape
and color that makes them inconspicuous. Imagoes feeding gen-
erally on the nectar of fiowers and other juices are mostly fond of
the bright sun and go to cover quickly at the slightest clouding
over.

The distribution of species in the state is often closely asso-
ciated with physiography and vegetation. Dr. Smith accom-
panied his report with an excellent map and text defining the
faunal regions. A later and more detailed discussion by Dr.
Fox^ elaborates this.

The state is divided into two portions, — the Northern District
and the Coastal Plain, — by the Fall Line which extends across

3 State of New Jersey Department of Agriculture, Bureau of Statistics and
Inspection, Circular No. 138. ‘‘A Eevised Annotated List of the Dermaptera

and Orthoptera of New Jersey,” by Henry Fox, 1928.

50

Journal New York Entomological Society [Vol. xlviii

the state from Trenton to New Brunswick and thence to Jersey
City. The term Fall Line refers to the line where the hard rocks
of the northern part of the state meet and pass under the loose
deposits of the southern half.

The Northern District has subdivisions of physiographic char-
acter which, extending north from the Fall Line, are the Pied-
mont Legion, the Crystalline Highlands, the Appalachian Valley
and the Kittatinny Range.

The Coastal Plain is subdivided, extending south from the
Fall Line, into the Middle District (inclusive of the Delaware
Valley), the Pine Barrens, the Coastal District (including the
Coastal Strip, Salt Marshes and Barrier Beaches) and the Cape
May Peninsula. These divisions of the Coastal Plain are based
on the nature of the soils and are vegetational. These terms are
used for defining the general distribution of species.

Information about the few butterflies which are of economic
importance is given in the paragraphs on those species. A list of
collectors ’ names is given with corresponding abbreviations which
are used for reference in the text. In recording the plenitude of
individuals of a species and date range the statements are for
average conditions. There are years of scarcity and abundance;
locally favorable conditions may at times produce large broods;
again, normally common species may some years appear to be
entirely absent. For the names of food plants Gray’s ‘‘New
Manual of Botany” (seventh edition) is followed.

I have searched thoroughly the pertinent entomological liter-
ature for records of New Jersey captures. I have also used the
data given in Smith’s “List” of 1909 as required. I am particu-
larly indebted to Mr. Frank E. Watson for many records and
facts and also criticism of the manuscript as a whole. Dr. Wil-
liam T. M. Forbes has given good advice and added information.
Others who have aided by contributing records and suggestions
are Mr. Ernest L. Bell, Mr. Otto Buchholz, Mr. Austin H. Clark,
Mr. Cyril F. dos Passos, Dr. Willis J. Gertsch, Mr. Sidney A.
Hessel, Mr. Charles Rummel and Mr. H. E. Wilford. Other col-
lectors’ records which are cited have been taken from published
statements.

Mar., 1940]

Comstock: Butterflies

51

LIST OF COLLECTORS'

(Aa.) Eugene M. Aaron.
(An.) Elmer Anthony.

(Ang.) John B. Angleman.
(Bar.) Roy Barnett.

(Bl.) E. L. Bell.

(Br.) H. H. Brehme.

(Bt.) Wm. Beutenmuller.
(Bu.) Edward J. Burns.
(By.) Ernest Baylis.

(Bz.) Otto Buchholz.

(C.) Wm. P. Comstock.

(Ca.) J. & J. Cahalan.

(Cy.) John P. R. Carney.
(Dke.) V, A. E. Daeke.

(Dn.) E. L. Dickerson.

(Doll.) Jacob Doll.

(Dow.) Robert P. Dow.

(dP.) Cyril F. dos Passos.
(Ds.) Wm. T. Davis.

(Eng.) George P. Engelhardt.
(Fr.) George Franck.

(Frd.) W. G. Freedley, Jr.
(Fox.) Wm. J. Fox.

(Ge.) Willis J. Gertsch.

(Gr.) John A. Grossbeck.
(Hg.) Herman Hornig.

(HI.) Gaylord C. Hall.

(Hm.) Frank Haimbach.
(Ho.) W.J. Holland.

(Hs.) Sidney A. Hessel.

(Hy.) F. Hoyer.

(Jn.) Charles W. Johnson.
(F.Jn.) Frank Johnson.
(Kl.) Alexander B. Klots.
(Kp.) Stanley T. Kemp.
(Kr.) Wm. D. Kearfoot.
(La.) Roy Latham.

(Lt.) Philip Laurent.

(Lz.) Frank E. Lutz.

(Mt.) Harvey Mitchell.

(N.) Alan S. Nicolay.

(New.) Lloyd Newsom.

(Pm.) Charles Palm.

(Pk.) A. S. Pinkus.

(Ort.) A. E. Ortmann.

(My.) Otto Mayer.

(Ru.) Charles Rummel.

(Sb.) Simon Seib.

(Se.) Otto Seifert.

(Sh.) Ernest Shoemaker.

( Sk. ) Henry Skinner.

(Sm.) John B. Smith.

(St.) Witmer Stone.

(Sto.) Robert W. Storer.
(Su.) C. H. Sunderland.
(W.) Henry W. Wenzel.
(Wat.) Frank E. Watson.
(Wt.) Wright.

(Ws.) Henry Wormsbacher.
(Wi.) Harry B. Weiss.
(Wf.) H. E. Wilford.

DANAID.E

Danaus Kluk

89. plexippus Linnaeus. Monarch, milkweed butterfly. Through-
out state. Open country, meadows. April-November. Three
broods (Wi-Dn). Migrant to south in late fall, does not
overwinter in state, females return in spring to start first
local brood. Records of males returning in spring also.

* Where (Sm) follows another reference it means that the reference occurs
in Smith ’s ^ ‘ List. ’ ’ Occasionally these references are corrected to agree with
the original references.

52

Journal New York Entomological Society [Vol. xlviii

Eggs laid singly, usually on under surface of leaves. Re-
ported as swarming in Florida Everglades in winter, both
males and females. Larvas, Tampa, Fla., March 1-21 (Bl).
Milkweeds ( Asclepias ) .

Lakehurst, April 25, (Ds) — ^Newark, Oct. 7, (C).

Recorded throughout state every month from spring to
fall. Aberration fumosus Hulst, several captures.

NYMPHALIDzE

Dione Hiibner

158. vanillae Linnaeus. Gulf fritillary. Occasional visitor.
Passion flower leaves (Passiflora) .

Cape May; 7-Mile Beach; Camden Co. (Hm) (Sm).

Euptoieta Doubleday

159. Claudia Cramer. Variegated fritillary. Coastal Plain.
Moist open fields. June-October. Three broods. Eggs
laid singly. Occasionally locally common. Passion flower
leaves, violets, pansy (Eng), orpine (Sedum) rarely (Wat).

Engelwood, July 6, (C) — South Amboy, Oct. 4, (C).
Cape May, June-Oct., (Hm) ; Barnegat Pier, Aug. 7-
Sept. 30, (Br) ; Runyon, Sept. 5, (C) ; Lakehurst, July 14,
(C) ; Arlington meadows, July 15, Aug. 20, (Ru) ; Wood-
ridge, Sept. 10, (Mt) ; Hillside, Sept. 15, 20, (Ru).

Argynnis Fabricius

161. idalia Drury. Regal fritillary. Throughout state. Open
country, moist meadows. Late June-September. One
brood. Larva a night feeder, hiding by day. Winters as
first stage larva. Eggs laid singly. The brood is long
extended fresh examples appearing from late June to
middle of August. Violets.

Mashipacong, June 29, (C) — Fairton, Sept. 16, (Br).
Jamesburg, June 30, July 1, Aug. 25, Sept. 3, (C, AVat) ;
Newton, July 5, 23, (C, Wat) ; Englewood, July 13,
(C) ; Ramsey, July 21, (Ge), 28, (C) ; Old Bridge, Aug. 4,
(Wat) ; Waterloo, Aug. 16, (C).

Aberration ashtaroth Fisher. School ey’s Mountain

(Ang) (Sm).

Mar., 1940]

Comstock: Butterflies

53

166. cybele Fabricius. Silver spot. Throughout state. Abun-
dant in southern area. Low open country. June-Septem-
ber. One brood. Larva nocturnal feeder. Winters as
first stage larva. Eggs laid singly. Violets.

Greenwood Lake Glens, June 22, (Wat) — Bowne, Sept.
16, (C).

Newton, July 5, 6, 10, 23, common (C, Wat) ; Jamesburg,
July 1, 4, Sept. 3, (C, Wat) ; Ramsey, July 28, (C) ; Old
Bridge, Aug. 4, (Wat) ; Atlantic Highlands, Aug. 8, (Wat) ;
Waterloo, Aug. 5, 16, (C).

167. aphrodite Pabricius. Northern District. Open country
and meadows. July-August. One brood. Larva noctur-
nal feeder. Winters as first stage larva. Eggs laid singly.
Violets.

Greenwood Lake Glens, June 25, (Wat) — Bowne, Sept.
12, (C).

Mashipacong, June 28, (C) ; Jamesburg, July 4, (C) ;
Newton, July 6, 23, (C, Wat) ; Englewood, July 13, (C) ;
Waterloo, Aug. 5, (C) ; Fort Lee, Aug. 25, (C).

Brenthis Hiibner

200. myrina Cramer. Little silver spot. Throughout state.
Moist fields and meadows. May-September. Two broods.
Winters as partly grown larva. Nocturnal feeder. Eggs
laid singly on stem or leaf. Violets.

Jamesburg, May 24, (C) — Carlstadt, Sept. 23, (C).
Jamesburg, May 30, June 3, 15, 20, Aug. 25, Sept. 3, (C,
Wat) ; Newton, July 23, (C, Wat) ; Ramsey, July 12, (Ge),
July 28, (C) ; Old Bridge, Aug. 28, (Wat) ; Dennisville,
Sept. 5, (C, Wat) ; Point Pleasant, Sept. 17, (C, Wat).

212. bellona Fabricius. Meadow fritillary. Throughout state.
Moist fields and meadows. June-September. Two broods.
The species is long lived and broods drawn out. Winters
as larvae in various stages of growth. Nocturnal feeder.
Violets.

Great Notch, May 2, 10, (Ru) ; Greenwood Lake Glens,
May 30, (C) ; Cranford, July 10, (Ru) ; Ramsey, July 12,
(Ge) ; Bowne, Sept. 12, (C) ; Andover, Sept. 16, (C).

54

Journal New York Entomological Society [Vol. xlviii

Euphydryas Scudder

217. phaeton Drury. Baltimore. Throughout state. Swampy
meadows. June-July. One brood. Sometimes locally
numerous. Gregarious as small larvae, wintering in web,
scattering in spring. Eggs laid in large clusters under-
neath leaf on turtlehead (Chelone glahra). In spring when
larvas scatter they feed more generally. On young ash
shoots (Wat). Figworts {Scrophulariacem) .

Greenwood Lake Glens, May 31, (C) — Ramsey, July 28,

(C).

Greenwood Lake Glens, June 16, 22, 25, (C, Wat) ; Ar-
lington, June 5, (Ru) ; Ramsey, June 9, (Ge) ; Hemlock
Falls, June 28, July 2, (C) ; Cape May Court House, July
16, (HI, Wat) ; Newton, July 23, larvge, (C, Wat).

This species varies greatly from suffusion to absence of
spots leading to a number of named aberrations: superba
Strecker and phaethusa Hulst have been taken in the state.

Melitaea Fabricius

256. harrisii Scudder. Harris’ checker spot. Locally, North-
ern District. Moist meadows. June. One brood. Win-
ters as young larva. Eggs laid in patches underneath leaf.
Larvae gregarious. Doellingeria umhellata.

Ramsey, June 9, (Ge) — Stanhope, June 12, (Ru) ;
Mashipacong, July 5, (HI, Wat) ; Greenwood Lake Glens,
June 22, 25, (C, Wat).

Phyciodes Hiibner

263. nycteis Doubleday & Hewitson. Silver crescent. Locally,
Northern District. Open grassy fields. June-July. One
brood. Winters as partly grown larvae. Eggs laid in
clusters up to 100 underneath leaf. Sunflowers, especially
Helianthus divaricatus, occasionally aster (Wat).

Millburn, June 15, (Ru) — Sloatsburg, N. Y., July 11, (C).

Greenwood Lake Glens, June 16, 22, 25, 27 (C, Wat) ;
Lake Lackawanna, June 20, 30 (Ru).

Aberration milburni Rummel. Type locality Millburn,
June 20, (Ru).

Mar., 1940]

Comstock: Butterflies

55

265. tharos Drury. Pearl crescent. Throughout state. May-
October. Three broods. Winters as partly grown larvae,
gregarious, irregular development due to lethargy makes
great variation in time of emergence of adult. Various
asters.

Staten Island, N. Y., Mar. 29, (Ds) — Mountain View, Oct.

5, (C).

Regularly on the wing from early May to middle October
and our most common butterfly. Form marcia Edwards is
the spring brood. Aberration packardii Saunders. Several
records.

Polygonia Hiibner

285. interrogationis Fabricius. Semicolon. Throughout state.
Borders of woodland and scrub. March-November. Two
broods. Winters as imago, hibernating in hollow trees or
other cover. Abundant, fond of feeding on fallen fruit in
orchards. Eggs laid singly or in chains on under surface of
leaves; larvse partly gregarious. Elm, hop, hackberry.

Hemlock Falls, June 23, (C) — Newark, Sept. 22, (C).

Frequent from the middle of June through September.
Form umbrosa Lintner is the summer brood.

286. comma Harris. Hop-merchant. Throughout state. Bor-
ders of woodland and open woods. March-November. Two
broods. Winters as imago. Sometimes locally abundant.
Eggs laid on under surface of leaf or stem. Larva forms
a nest. Nettle, hop, elm.

Old Bridge, April 26, (C) — Fort Lee, Oct. 21, (C).

Greenwood Lake Glens, June 22, (Wat) ; Newton, July
23, (C, Wat) ; Caldwell, Aug. 20, (C). Form dryas Ed-
wards is the summer brood.

288. faunus Edwards. Green comma. Kittatinny Range. Open
woodland. March-November. One brood. Winters as
imago. Very scarce in state, more common northward.
Eggs laid singly on top of leaf. Birch, willow, currant,
gooseberry.

Mashipacong Pond, July 6, (E. R. Watson in Coll. Wat) ;

56

Journal New York Entomological Society [Vol. xlviii

Lake Lackawanna, Sept. 6, (Rn) ; School ey Mountain (Aa)
(Sm).

294. progne Cramer. Gray comma. Northern District. Open
woodland. March-November. Two broods. Winters as
imago. Not common. Eggs laid singly in notches at edge
of leaf. Currant, gooseberry.

Hemlock Falls, April 20, (Wat) — Hamburg, Oct. 2, (C).

Camden, April 24, (Cy) (Sm) ; Irvington, June 11, 14,
(Bz) ; Greenwood Lake Glens, July 18, (C) ; Stanhope,
July 23, (Ru) ; Lake Lackawanna, July 12, Aug. 13, (Ru),
Aug. 19, (Bz) ; Andover, Aug. 30, (C).

Form 1-argenteum Scudder is the summer brood.
Greenwood Lake Glens, July 1, 2, (Wat).

Nymphalis Kluk

295. j-album Boisduval & LeConte. Large tortise-shell.
Throughout state locally. Open woods and wood roads.
March-November. One brood. Winters as imago. Oc-
casionally locally common. Gregarious larv83. Gray birch
{Betula populifolia) (Wat).

Hemlock Falls, April 6, (C) — Barnegat Pier, Sept. 30,
(Br).

Alpine, June 10, larvae abundant, (Wat) ; Newton, July
6, 10, (C) ; Ramsey, July 7, (Ge) ; Green Village, July 7,
(Ru) ; Barnegat City, Aug. 16, (Br) (Sm) ; Great Notch,
Sept. 5, (Ru) ; Newark, Sept. 12, (C) ; Lakehurst, Sept. 17,
(Ds) (Sm).

297. milberti Godart. Small tortoise-shell. Northern District.
Open fields and meadows. June-October. Three broods.
Winters as pupa and imago. Eggs in masses, sometimes
laid beneath leaf, larvse gregarious. Usually scarce but
some seasons locally very numerous. Nettles {Vrtica
gracillis and dioica).

Hillside, July 10, (Ru) — Camden, Nov. 6, (Cy) (Sm).
Paterson, July 20, Aug. 13, (Gr) (Sm) ; Swartswood Lake,
July 25, (Ds) (Sm) ; Hemlock Palls, Aug. 17, (Wat) ; Ham-
burg, Oct. 2, (C) ; Harrison, Oct. 24, (C).

Mar., 1940]

Comstock: Butterflies

57

298. antiopa Linnaeus. Mourning cloak. Throughout state.
Open woods, wood roads and edges of wet meadows. March-
November. Two broods. Winters as imago. Eggs laid
usually in a single layer encircling a twig ; gregarious larvse,
often making a large colony, defoliate food plant. Control
by arsenical sprays. Elm, willow, poplar, hackberry.

Hemlock Falls,* Mar. 23, (C)— Newark, Oct. 23, (C). On
wing almost everywhere throughout season.

Aberration hygiaea Heydenreich. Synonym lintnerii
Pitch. Occurs in various gradations but is very scarce.

Vanessa Fabricius

299. atalanta Linnaeus. Ked admiral. Throughout state. On
flowers of field and roadside. April-November. Two
broods. Winters as imago. Often abundant in fall. Some-
what migratory. Eggs laid singly on upper surface of leaf.
Larva lives in a nest. Nettle, false nettle {Boehmeria
cylindrica) .

Old Bridge, April 26, (C) — Mountain View, Oct. 5, (C).

On the wing throughout season, but perhaps most com-
mon in September.

300. virginiensis Drury. Synonym huntera Fabricius. Painted
beauty. Throughout state. On flowers of field and garden.
April-November. Two broods. Winters as pupa and
imago. Egg laid singly on top of leaf. Larva forms a
nest. Everlastings (Antennaria) .

Jamesburg, May 17-Oct. 10, (C).

Throughout season, but more common, July through
September.

301. cardui Linnaeus. Painted lady. Throughout state. On
flowers of field and roadside and in waste places. April-
November. Two broods. Winters as imago. In some
years it appears in considerable numbers and again seems
entirely absent. A migratory species with a speed of
flight 20 to 25 miles an hour. Egg laid singly on top of
leaf. Larva makes a nest. Thistle, burdock, hollyhock.

Newton, July 23, (C, Wat) — ^New Brunswick, Sept. 23,
(Wat). There are earlier and later records for New Eng-
land states (Scudder).

58

Journal New York Entomological Society [Vol. XLVIII

Junonia Hiibner

303. coenia Hiibner. Buck-eye. Abundant south of Fall Line,
more sparingly in Northern District. On flowers of field
and roadside. May-November. Two broods. Winters as
imago probably as it does further south. Often common in
Coastal District in late summer. Eggs laid singly on tips
and underneath leaves. Gerardia, snapdragon.

Lakehurst, June 29, (C) — Hillside, Oct. 17, (Ru).

Arlington, Aug. 5, Sept. 5, (Ru) ; Milltown, Aug. 23, (C) ;
Dennisville, Sept. 5, (C, Wat) ; Point Pleasant, Sept. 17,
(C, Wat) ; South Amboy, Oct. 4, (C).

Basilarchia Scudder

322. astyanax Fabricius. Synonym Ursula Godart. Blue vice-
roy. Throughout state. Borders of woodland and road-
side. May-September. Two broods. Winters as small
larva in hibernaculum formed from tip of leaf. Imagoes
often numerous on fallen fruit in orchards. Wild cherry,
apple, thorn, plum, poplar and willow.

Jamesburg, May 30, (C) — Carlstadt, Sept. 18, (C).

More numerous in July and August but on the wing
through five months.

Form albofasciata Newcomb flies with astyanax but is
scarce. Intergrades occur. Mr. Hall records 8 specimens
from Highpoint, 4 in one day.

Mashipacong, June 28, (C) ; Ogdensburg, July 10, (Bl) ;
Elizabeth, July 17, (Bz) ; East Brunswick, July 29, (Dow) ;
Waterloo, Aug. 5, (C) ; Great Notch, Aug. 16, (Ru) ; Still-
water, Aug. 5, (F. Treuting) ; Stanhope, Aug. 15, (Mrs.
F. G. Ruggles).

325. archippus Cramer. Viceroy. Throughout state. Borders
of woods, roadsides and meadows. June-October. Two
broods. AVinters as small larva in hibernaculum. Eggs
laid singly. Willow and poplar.

Jamesburg, June 15, (C) — Mountain View, Oct. 5, (C).

Jamesburg, June 20, July 4, Aug. 25, (C) ; Newton, July
6, 23, larvae, (C, Wat) ; Ramsey, July 28, (C) ; Old Bridge,
Aug. 4, (Wat) ; Dennisville, Sept. 6, (C, AVat) ; Carlstadt,
Sept. 18, (C).

Aberration lanthanis Cook & Watson flies with archippus

Mar., 1940]

Comstock: Butterflies

59

but is scarce. Intergrades occur. Athenia, Aug. 13,
(Wat) ; Elizabeth, Aug. 14, (Bz) ; Arlington, Aug. 20, (Ru).

Asterocampa Rober

327. celtis Boisduval & LeConte. Locally, Northern District.
Open woods. June-September. Two broods. Winters
as larva and perhaps imago (Edwards). Imago rests on
leaves and tree-trunks in woods. Eggs laid singly or in
small clusters on under side of leaf. Hackberry {Celtis).

Newton, July 10, ’36, (C), larvas, July, (dP, Wat) ; Lake
Lackawanna, June 12, 28, July 5, (Ru) ; Great Notch, June
13, ’37, (Bz) ; Burlington (By) ; New Brunswick (Bar) ;
South Orange, July, larvffi (Pk) ; Swartswood Lake, Sept. 4,
’37, (N) ; Lawrence Harbor, Aug. 20, fairly common (W.
C. Frey).

329. clyton Boisduval & LeConte. Emperor. Locally, North-
ern District. Open woods. June-July. One brood. Winters
as small larva. Eggs laid in masses, sometimes 500, on
underside of leaf. Larvne gregarious for three stages, then
after hibernating, scattering. Imago with habits like pre-
ceding species but more frequently seen. Hackberry
{Celtis).

Hemlock Palls, June 28-July 14, (C, Wat) ; Arlington,
July 5, 15, 20, (Ru) ; Newton, July 6, 10, (C, Wat), larvae,
July, (dP, Wat).

Form proserpina Scudder flies with the typical form.

SATYRIDiE

Enodia Hiibner

96. portlandia portlandia Fabricius. Locally, throughout state.
Wood paths and open spots in woods. Julj^-August. One
brood. Winters as larva. Grasses.

Newton, July 11, (C) ; Palisades, July 20, (Eng),
portlandia anthedon Clark. This seems to be the sub-
species most frequently taken in the state.

Mountain Side, June 26, (Bz) ; Hemlock Falls, July 5,
(Wat) ; Lake Hopatcong, July 5, Bear Swamp near Ram-
sey, July 10, Green Village, July 8, (Bz) ; Millburn, July
8, 10, (Ru) ; Ogdensburg, Aug. 10, (Bl).

60

Journal New York Entomological Society [Vol. xlviii

Note. The following records show distribution in the
state without definite record of the form. Ramsey, July
23, (Ge) ; Livingston, July 26, 28, (Hs) ; Newfoundland,
July 27, (Ds) (Sm) ; Paterson, July 16, Aug. 17, (Gr)
(Sm) ; 5-Mile Beach, (Hm) (Sm).

Neonympha Hubner

100. areolatus septentrionalis Davis. As phocion Pabricius
(Sm). Pine Barrens and Coastal District. Swamps with
coarse grass. June-July. One brood. Winters as larva
(Wat). Grasses.

Jones Mills, June 21, (St) (Sm) ; Lakehurst, June 28,
(Wat), July 7, 11, 19, (Hs), type locality, July, (Ds) ;
Richland, July 17, (HI, Wat) ; Brown’s Mills Jc., July 12,
(Dke) (Sm) ; Da Costa, July 17, (Lt) (Sm) ; Toms River,
July 27, (Dke) (Sm) ; Atlantic City, July, (Aa) ; Deal, July
9, (Ca).

Megisto Hubner

103. eurytus Pabricius. Throughout state. Open woodland.
May-July. One brood. Winters as larva. Eggs laid
singly. Grasses.

Bowne, May 28, (C) — Ramsey, July 28, (C).

Common generally through June and July.

Satyrodes Scudder

106. eurydice Johann. Synonym canthus Linnaeus. Through-
out state. Locally common in grassy bogs. June-August.
One brood. Winters as nearly grown larva and possibly
pupa. Eggs laid singly. Coarse swamp grasses and sedges
(Scirpus and Car ex).

Jamesburg, June 30, (C) — Runyon, Sept. 4, (C).

Newton, July 7, 11, (C) ; Ramsey, July 28, (C) ; Livings-
ton, July 26, 28, (Hs) ; 5-Mile Beach, June (Hm).

Minois Hubner

117. alope Pabricius. Blue-eyed satyr. Throughout state.
Open woods and brush. July-September. One brood.

Mar., 1940]

Comstock: Butterflies

61

Winters as larva on emerging from egg. Eggs laid singly.
Grasses.

Newton, July 6, (C) — Carlstadt, Sept. 18, (C).

Green Village, July 15, (Ru) ; Waterloo, Aug. 16, (C) ;
Woodridge, Sept. 10, (Mt).

Form maritima Edwards is more frequent below the Fall
Line. Jamesburg, July 5, (C) ; South Lakewood, July 13,
(Wat) ; 5-Mile Beach, July-Sept., common (Hm).

Form nephele Kirby occurs in the northwestern part of
state. Newton, July 5, 7, (C) ; Mashipacong, July (HI).

LIBYTHEID.E

Libythea Fabricius

336. bachmannii Kirtland. Snout butterfly. Locally, through-
out state. June-September. Three broods. Winters as
imago. Hackberry (Celtis).

Avalon, July 4, (Kp) ; Camden, July 9, (Cy) ; Fort Lee,
July 11, (Ws) ; Greenwood Lake Glens, July 18, (C) ;
Snake Hill, July 20, (Wat) ; Lakehurst, July 21, (Ds)
(Sm) ; Atlantic Highlands, July 26, (C, Wat) ; Arlington,
Aug. 13, (Ru) ; Morgan, larvse (Wat).

RIODINID^

Calephelis Grote & Robinson

346. borealis Grote & Robinson. Northern metal-mark. On
limestone outcrops, Appalachian Valley and Kittatinny
Range. Thickets and fairly thick woods. July. One
brood. Winters as larva. The butterfly rests with wings
open on a sun-lit leaf or a flower in small open spots in
shady woods. Round leaf squaw weed {Senecio ohovatus).

Newton, July 5-18, (Gr, Wt, dP, Wat, C) ; Delaware
Water Gap, (Aa) (Sm).

LYC^NID^

Strymon Hiibner

372. m-album Boisduval & LeConte. Occasional visitor, south-
ern species. Oak.

Orange Mountains, April 28, (Br) (Sm) ; Atlantic City,

62

Journal New York Entomological Society [Vol. xlviii

June 11, (Aa) (Sm) ; Jamesburg, June 21, (Wat) ; Lake
Hopatcong, July 5, (Fr) (Sm).

373. melinus Hiibner. Throughout state. Open woods and
fields on flowers. May-0 ctober. Three broods. Winters
as pupa. Eggs laid singly in flower heads. Flower and
fruit feeder. Bush clover (Lespedeza hirta), other legumes.

Pompton, April 24, (C, dP) — South Amboy, Oct. 4, (C).

Jamesburg, May 8, (C) ; Camden, June 17, Sept. 17,
(Cy) (Sm) ; Kamsey, July 16, (Ge) ; Manasquan, July 24,
(C) ; Paterson, Aug. 3-17, (Gr) (Sm) ; Fort Lee, Aug. 23,
(C) ; Dennisville, Sept. 7, (C, Wat).

380. titus Fabricius. Locally, throughout state. July-August.
One brood. Winters as egg. Egg laid singly on twig
(Scudder). Larva on wild cherry, by day about roots of
small plants, my rmecophilous, night feeder (Wat). Pupae
on wild cherry rootstalk below surface of ground, June 20,
(C). Imago on flowers in bright sun. Sometimes common.

Jamesburg, June 24, (C) — Old Bridge, Aug. 4, common
(Wat).

Kamsey, July 3, (Ge) ; Jamesburg, July 4, (Wat) ;
Englewood, July 6, (C) ; Westwood, July 8-29, (Mt) (Sm) ;
Dover, July 16, (Jn) (Sm) ; Paterson, July 17, (Gr) (Sm).

381. acadica souhegan Whitney. Crystalline Highlands, Ap-
palachian Valley. June-July. One brood. Winters pos-
sibly as egg. Larva by day on leaves and stems of small
plants of willow {Salix discolor), myrmecophilous (AVat).
Imago on flowers, locally common some seasons.

Ledgewood, June 29, (dP) — Kamsey, July 17, (Ge).

Great Meadows, June 30, (dP) ; Hewitt, July 1, (C,
Wat) ; Dover, July 5, (dP) ; Newton, July 6, (C) ; South
Ogdensburg, July 7, (dP) ; Newfoundland May-June, larv«
(Wat).

385. edwardsii Grote & Kobinson. Northern District. June-
July. One brood. Winters as egg. Larvge in ants’ nests
at roots of scrub, scarlet oak, late May (Wat). Imago on
flowers.

Greenwood Lake Glens, June 21, 28, (C) ; Newfoundland,
July 3, (Ds) (Sm) ; Newark, July 4, (Br) (Sm) ; Newton,
July 6, (C) ; Hewitt, May, larvae (Wat).

Mar., 1940]

Comstock: Butterflies

63

387. falacer Godart. calanus Auct. Throughout state. June-
August. One brood. Winters as egg. Larvae feed on
leaves of oak, hickory and butternut.

Paterson, June 24, (Gr) (Sm) — Camden, Aug. 1, (Cy)
(Sm).

5-Mile Beach, July 3, (Hm) (Sm) ; Lakehurst, July 13,
(Hs) ; Newton, July 23, (C, Wat) ; Ramsey, July 28, (C).

389. liparops strigosa Harris. Locally, throughout state. July.
One brood. Winters as egg. Egg laid singly on terminal
twig. Swamp blueberry (Vaccinium corymhosum) (Wat),
shadbush.

Jamesburg, July 4, (Ds) (Sm) — Old Bridge, Aug. 4,
(Wat).

Lakehurst, July 7, 13, (Hs) ; Ramsey, July 16, (Ge) ;
Greenwood Lake Glens, July 18, (C) ; 5-Mile Beach, July
17-26, (Hm) (Sm) ; Hewitt, June, larvae (Wat).

Mitoura Scudder

401. damon Cramer. Green hair-streak. Locally, throughout
state. May and July. Two broods, second partial. Win-
ters as pupa. Egg laid singly at tip of blossoming twig.
Occasionally common. Red cedar {Juniperus virginiana) .

Pompton, April 24, (C, dP) — Westwood, May 20, (Mt)
(Sm).

Almonessen, April 29, (Hg) ; Lakehurst, May 10, (Hs).

Form aestiva smilacis Boisduval & LeConte. Synonym
patersonia Brehme, Paterson, type locality, July 25, (Br).
Greenwood Lake Glens, July 18, (C, Wat) ; Paterson, July
27, (Gr) (Sm).

Incisalia Scudder

403. augustus Kirby. Throughout state. Open scrubby places.
April-May. One brood. Winters as pupa. Larva flower
feeder. Frequently abundant, Pine Barrens. Sheep laurel
{Kalniia angustifolia) , blueberry (Vaccinium).

Paterson, April 8, (Gr) (Sm) — Hemlock Falls, May 30,
(Bz) (Sm).

Jamesburg, April 18, 24, May 8, 17, (C) ; Lakehurst, April
29, abundant (Hs) ; Westville, April 25, (Sk) (Sm) ; Lake-

64

Journal New York Entomological Society [Vol. XLVIII

wood, April 21, May 25, (C) ; Greenwood Lake Glens, May
3, 10, (C, Wat).

405. irus Godart. Locally, thronghout state. April-May. One
brood. Winters as pupa. Egg laid singly at base of flower

stem. Not uncommon. Pine Barrens. Wild indigo {Bap-
tism tinctoria) (Wat), lupine {Lupinus perennis).

Spotswood, April 26, (C) — Greenwood Lake Glens, May
30, (C).

Kanaus Mt., May 4, (dP) ; Jamesburg, May 8, 17, 30,
(C) ; Lakehurst, April 29, (Hs, Wat) ; Anglesea, May 1,
(Lt) (Sm) ; Clementon, May 16, (Jn) (Sm).

407. henrici Grote & Robinson. Locally, Northern District.
Open places with low vegetation. May. One brood.
Rare. Huckleberry {Gaylussacia resinosa).

Greenwood Lake, tops of ridges, April 25, May 10, (C,
Wat).

Note. The several records given in Smith’s ‘‘List” for
this species are omitted as doubtful through misidentifica-

tion.

409. polios Cook & Watson. Pine Barrens. April-May. One
brood. Winters as pupa. Eggs laid among and about
terminal flower buds. Occasionally numerous. Bearberry
{Arctostaphylos uva-ursi).

Lucaston, April 10, (Dke) (Sm) — ^Lakewood, May 25,

(C).

Type locality, Lakewood, April 21, 27, (Wat) ; Milltown,
April 22, (Gr) (Sm) ; Jamesburg, April 18, 24, (C, Wat) ;
Outcault, April 21, (C) ; Lakehurst, April 29, abundant
(Hs).

Aberration davisi Watson & Comstock. Type locality
Lakehurst, April 29, (Wat).

413. niphon Hiibner. Pine Barrens. Open woods. April-May.
One brood. Winters as pupa. Eggs laid singly on pine
sprouts. Not common. Pitch-pine.

Lakewood, April 21, (C) — Camden, June 18, (Sk) (Sm).
Lakehurst, April 29, May 10, (Hs, Wat), May 2 (Bz) ;
Westville, April 29, (Sk) (Sm) ; 5-Mile Beach, May 7,

Mar., 1940]

Comstock; Butterflies

65

(Hm) (Sm) ; Clement on, May 9, (Lt) (Sm) ; Jamesburg,
May 15, 17, 30, (C).

Feniseca Grote

419. tarquinius Fabricius. The wanderer. Locally, throughout
state. May-October. Three broods. Winters as pupa.
Found as occasional individuals resting on a sun-lit leaf in
open woods. Eggs laid singly on alder twigs near lice.
Larva carnivorous, feeding on woolly alder louse (Prociphi-
lus tessellata Fitch), sometimes numerous.

Andover, May 14, bred (C) ; Caldwell, May 26, (C) ;
Hillside, May 30, July 15, (Ru) ; Newton, July 18, (C) ;
Hewitt, July 19, (Hs) ; Waterloo, Aug. 16, (C) ; Ramsey,
Aug., (Ge).

Lycaena Fabricius

424. thoe Guerin. Large copper. Locally, Northern District.
Damp meadows. June- July and August-September. Two
broods. Winters as egg. Eggs laid singly on underside of
leaves and petioles and among seeds. Local and not numer-
ous. Yellow dock {Rumex crispus).

Arlington, June 6, (Ru) — Carlstadt, Sept. 23, (C).

Troy Meadows, June 7, (C) ; Paterson, June 12, Aug. 3,
(Gr) (Sm) ; Snake Hill, July 23, (C, Wat) ; Rutherford,
July 29, (Se) ; Woodridge, Sept. 10, (Mt) ; Waverly, Sept.
19, (Wat).

434. epixanthe Boisduval & LeConte. Small copper. Pine Bar-
rens in cranberry bogs. June-July. One brood. Winters
as egg. Eggs laid singly mostly underneath leaf. Fre-
quently numerous in cranberry bogs. Cranberry.

Brown’s Mills Jc., June 17, (Dke) (Sm) — Toms River,
July 15, (HI, Wat).

Lakewood, June 25, (C) ; Jamesburg, June 21, July 5,
(C) ; Lakehurst, July 2, 14, (C) ; Mashipacong, July, (HI,
Wat).

435. phlaeas hypophlaeas Boisduval. Common copper. Through-
out state. May-October. Three broods. Winters as pupa.
Egg laid singly on stem or leaf. Usually common. Sorrel

66

Journal New York Entomological Society [Vol. xlviii

{Rumex acetosa).

Almonessen, April 29, (Hg) — South Amboy, Oct. 4, (C).

Jamesbiirg, May 8, 17, 24, 30, June 30, July 4, Sept. 3,
(C, Wat) ; Newton, July 6, 10, (C) ; Atlantic Highlands,
Aug. 8, (Wat) ; Bowne, Aug. 21, Sept. 12 (C) ; Carlstadt,
Sept. 18, (C).

This species is subject to considerable variation. Seven
named aberrations have been captured in the state : — fasci-
ata Strecker, obliterata Scudder, fulliolus Hulst, octo-
maculata Dean, banksi Watson & Comstock, fulvus Rum-
mel, neui Rummel. In some restricted areas the tendency
to aberration seems considerable, frequent individuals show-
ing divergence from normal maculation or color but in most
localities the normal form is predominant and the aberra-
tions are very scarce.

Everes Hiibner

447. comyntas Godart. Tailed blue. Throughout state. Open
country. May-September. Three broods. Winters as full
grown larva. Eggs laid singly in flower heads. Usually
common. Flowers of bush clover {Lespecleza) , tick trefoil
(Desmodium) , other legumes.

Almonessen, April 29, (Hg) — South Amboy, Oct. 4, (C).

Continuously on the wing throughout season with over-
lapping of broods.

Lyceanopsis Felder & Felder

475. argiolus pseudargiolus Boisduval & LeConte. Common
blue. Throughout state. Open moist woods. April-Sep-
tember. Three broods. Winters as larva and pupa.
Spring brood often very common in open woods, summer
broods more scattered and less frequent. Eggs laid singly
in flower buds. Larva myrmecophilous (C, Wat). Flower
heads of many plants: — Cornus florida, Vihurnum aceri-
folium, Cimicifuga racemosa and Prunus serotina.

Form vernalis lucia Kirby. Form vernalis marginata
Edwards. These two spring -forms occur with pseudargi-
olus (synonym violacea Edwards). There are intergrades
of every degree. Sometimes the heavily marked forms pre-
dominate.

Mar., 1940]

Comstock: Butterflies

67

Hemlock Falls, March 29, April 20, May 4, (C, Wat) ;
Greenwood Lake Glens, April 25, May 3, (C) ; Lakewood,
April 21, May 18, (Wat) ; Hope, May 5, (C).

Form vernalis neglecta-major Tutt. This form follows
the early spring forms and is intermediate in occurrence
between them and the first summer brood.

Hemlock Falls, May 24, (Wat) ; Jamesbnrg, May 30, (C).

Form estiva neglecta Edwards.

Jamesbnrg, June 15, 20, July 4, Sept. 3, (C, Wat) ; New-
ton, July 5, 10, 18, (C) ; Newark, Aug: 11, 17, (C) ; Point
Pleasant, Sept. 13, (C, Wat).

PAPILIONID^

The Papilionidae in New Jersey are represented by several
species of the genus Pa/pilio which may be divided into three
groups: Aristolochia swallowtails of which P. philenor is the
representative ; Fluted swallowtails including P. ajax asterius,
P. chresphontes, P. glaucus, P. troilus and P. palamedes ; Kite
swallowtails represented by P. marcellus. Three genera may be
used to classify the three groups, respectively : Troicles Hiibner,
Papilio Linnaeus and Iphiclides Hiibner. See ‘‘A Revision of
the American Papilios”, by Hon. Walter Rothschild, Ph.D. and
Karl Jordan, Ph.D. (Novitates Zoologicae, Vol. 13, 1906).

Papilio Linnaeus.

1. philenor Linnaeus. Green swallowtail. Throughout state.
Open fields on flowers, open woods. May-October. Three
broods. Winters as pupa. Sometimes common locally.
Eggs laid in bunches on upper side of leaf or stem. Larvae
gregarious. Dutchman’s pipe (Aristolochia) , wild ginger
(Asarum canadense) .

Greenwood Lake Glens, May 18, (Wat) — Andover, Sept.
16, (C).

Hewitt, May 30, (C) ; Hemlock Falls, June 10, (C) ;
Newton, July 23, (C, Wat) ; Ramsey, July 28, (C) ; Fort
Lee, Aug. 23, (C) ; 5-Mile Beach, July, (Hm) ; Saddle
River, July, (Ge).

4. ajax asterius Cramer. P. ajax polyxenes Fabricius is the
Cuban subspecies. Parsley swallowtail. Throughout

68

Journal New York Entomological Society [Vol. XLVlil

state. Open country. April-October. Two broods.
Winters as pupa. Eggs laid singly. Sometimes so abun-
dant as to be injurious on parsley, carrots and celery.
Hand picking is the safest control. Umhelliferce generally.

Pompton, April 24, (C, dP) — South Amboy, Oct. 4, (C).

On wing throughout season.

Note. Rothschild & Jordan record a transitional speci-
men (female) to calverleyi Grote from Passaic, N. J. (Novi-
tates Zoologicae, Vol. 13, p. 547, 1906).

12. cresphontes Cramer. As thoas Linnaeus (Smith’s ‘‘List”).
Orange-dog, Giant swallowtail. Throughout state. Open
country. June-July and August-September. Two broods.
Winters as pupa. Occasional occurrence, but where prickly
ash is common several may be seen in a day. Eggs laid
singly, usually on upper side of leaves. Prickly ash
{Zanthoxylum americanum), hop-tree (Ptelea trifoliata).

Greenwood Lake Glens, May 18, 30, (Wat) ; Awosting,
May 21, (Hs) ; Newton, July 23, seven specimens (C, Wat) ;
Paterson, Aug. 12, (Gr)(Sm). Records of 1937 — Green
Village, Aug. 12, 16, (Ru) ; Lake Lackawanna, Aug. 10,
(Ru) ; Newton, Aug. 10, Sept. 6, (Ru) ; Newark, Aug. 28,
(C).

15. glaucus Linnaeus. Tiger swallowtail. Throughout state.
Open country. May-September. Two broods. Winters
as pupa. Eggs laid singly on upper surface of leaf. Larva
spins a web on top of leaf where it rests while not feeding.
Wild cherry, tulip-tree, apple, ash and poplar.

Jockey Hollow Park, April 20, (dP) — Bowne, Sept. 16,
( C ) . On the wing for five months.

Form female turnus Linnaeus. This yellow female is
more common than the black one in the state. Intergrades
occur.

20. troilus Linnaeus. Sassafras swallowtail. Throughout state.
Open country. May-September. Two broods. Winters as
pupa. Eggs laid singly on under surface of leaves. Larva
on upper side of leaf which it rolls together and fastens for
concealment. Sassafras, spice bush.

Hemlock Falls, May 4, (Wat) — Carlstadt, Sept. 18, (C).

On the wing for five months.

Mar., 1940]

Comstock: Butterflies

69

22. marcellus Cramer. As ajax (Smith’s “List”). Zebra
swallowtail. Occasionally throughout state. May-August.
Two broods. Winters as pupa. Eggs laid singly on upper
side of leaf. Papaw (Asimina triloha).

Hemlock Palls, June 30, (Br) (Sm) ; Newfoundland,
July, (Ds) (Sm) ; Jersey City, July 3, 4, five specimens
(Ws) ; Lakehurst, July 7, (Hs) ; Woodridge, Sept. 10,
(Mt) ; Anglesea (W).

PIERID^

Pieris Schranck

82. protodice Boisduval & LeConte. Checkered white. Gener-
ally south of and sparingly north of the Fall Line. In
waste places. May-September. Two broods. Winters as
pupa. Large broods frequently occur followed by years of
scarcity. Eggs laid singly. Cruciferse, especially wild
peppergrass (Lepidium virginicum) (Wat).

Riverton, April 16, (Jn) (Sm) — Hillside, Oct. 30, (Ru).

Lakewood, April 26, (Wat) ; 5-Mile Beach, May, (Hm)
(Sm) ; Newark Meadows, June 1, Sept. 10, (Ru) ; James-
burg, July 4, (C) ; Ramsey, July 20, (Ge) ; Paterson, July
30, (Gr) (Sm) ; West Orange, Aug. 25, (C) ; Harrison,
Sept. 25, (An) ; Mountain View, Oct. 5, (C). Spring form
vernalis Edwards is of sparse occurrence.

84. virginiensis Edwards. Records of oleracea (Smith’s
“List”) probably refer to this species. Occasionally
throughout state. May-June. One brood. Winters as
pupa. Toothwort {Dentaria dipliylla).

Garrett Rock, April 30, male (Wat) ; May 6, female
(Kr) ; Paterson, May 6, female (Gr) (Sm) ; Blairstown,
. June 2, male (Kl) ; Camden, (Cy) (Sm).

86. rapas Linnaeus. Cabbage butterfly. Throughout state.
Open country. March-November. Three broods. Winters
as pupa. Eggs laid singly. Introduced from Europe, this
species is often injurious to cabbage, cauliflower, and kale
and occurs on Cruciferse generally.

Newark, April 5, (C) — Stockton, Nov. 19, (C).

On the wing commonly throughout the season. In the
spring specimens without spots are occasionally taken and
called immaculata by various authors.

70

Journal New York Entomological Society [Vol. xlviii

Anthocharis Boisduval

30. midea Hiibner (genutia Fabricius). Orange-tip. Locally,
throughout state. Open woods. April-May. One brood.
Winters as pupa. Sometimes very common in Northern
District. Eggs laid singly, anywhere on plant. Rock cress
{Arahis lyrata and perfoliata) (Wat), 8isym~brinm thali-
ana, Westville (Hg).

Manumnskin, April 24, (Dke) — Awosting, May 27, (Hs).
Greenwood Lake Glens and Awosting, April 25, May 10,
(C, Wat), May 22, 27, (Lis) ; Almonessen, April 29, (Hg) ;
Little Falls, May 14, (Ds) (Sm) ; Old Bridge, April 26,
(C) ; Pompton, May 1, (C, dP, Rn) ; Oakland, May 2, (Ge) ;
7-Mile Beach, May, (Hm).

Phoebis Hiibner

57. sennae eubule Linnaens. Cloudless sulphur. Coastal area
and occasionally inland. Open country. Augnst-October.
Eggs laid singly on young leaves. A strong flier with
migrant habits. Senna (Cassia). Princeton, Stony Brook
Valley, larvae on Cassia marylandica (Ort).

West Creek, Chatsworth, Hammonton, Pleasant Mills,
Camden, Whitings, Aug.-Sept., (St) ; Barnegat Pier, Aug.
7-Sept. 30, common (Br) ; 5-Mile Beach Sept, ‘‘swarms”
(Hm) ; Cai3e May, Sept. 18, ’32, migration observed, 50
counted in one hour (Pk) ; Beach Haven, Sept. 3, (Frd) ;
Sept. 27 (Br) ; Cape May Co. and northward, Spring Lake,
Aug., Sept., (Ho) ; Manasquan, Spring Lake, Sept. 2, Point
Pleasant, Sept. 17, (AVat) ; Newark, Sept. 12, 13, (Ang) ;
Franklin Park, Sept. 14, (Ru) ; Great Piece Meadows, Sept.
2, (C) ; La Vallette, late Aug., common (Joe Scheuerle).

Colias Fabricius

41. eurytheme Boisduval. Alfalfa butterfly. Throughout
state. Open country. May-0 ctober. Two or three broods.
Winters as larva. Eggs laid singly on top of leaf. Of
this species. Dr. Smith remarked, thirty years ago : —
“hardly a regular inhabitant of the state.” Since, in the
last fifteen years, this species has become almost as numer-
ous as philoclice. Alfalfa, clovers.

Mar., 1940]

Comstock: Butterflies

71

Lakeliurst, May 10, (Hs), July 3, (Ru) ; Green Village,
June 24, Aug. 12, Oct. 30, (Ru) ; Bowne, Aug. 31, Sept. 3,
16, Oct. 13, (C) ; Harrison, Sept. 25, Oct. 3, 13, (An) ; Belle-
ville, Oct. 10, (C) ; Hillsdale, Oct. 20, (C).

Form sestiva amphidusa Boisduval is the common form.
The spring form eurytheme is rare. Form female alba
Strecker occurs.

42. philodice Godart. Clouded sulphur. Throughout state.
Open country. May-October. Three broods. Winters as
larva. Egg laid singly on upper side of leaf. Clover and
other legumes.

Lakehurst, April 29, (Hs) — Fort Lee, Oct. 21, (C).

On the wing throughout the season.

Form vernalis anthyale Hiibner is of frequent occurrence
in early spring. Form female alba Strecker occurs through-
out the season.

Eurema Hiibner

67. nicippe Cramer. Locally, Coastal Plain. September-Oc-
tober. Eggs laid singly, usually on underside of leaf.
Senna {Cassia).

Barnegat Pier, Aug. 7-Sept. 30, not common (Br) ; 5-
Mile Beach, Sept.-Nov., (Hm) ; New Brunswick, Aug. 23,
Sept. 22, (C, Wat) ; Point Pleasant, Sept. 13, (Wat) ; Eliza-
beth, Sept. 14, (Gr) (Sm) ; Hillside, Sept. 5, 16, (Ru) ;
Waverley, Sept. 25, (C).

72. lisa Boisduval & LeConte. Little sulphur. Throughout
state. Common in Coastal District, less numerous above
Fall Line. On flowers and moist earth. June and August-
September. Two broods. Eggs laid singly on upper side of
midrib of Cassia marylandica and nictitens.

Jamesburg, May 30, (C) — Oct. 12, (C).

Occurrence is sparse until August and September when
it is often common.

HESPERIID^

Epargyreus Hiibner

484. tityrus Fabricius. Silver-spotted skipper. Throughout
state. May-September. Two broods. Winters as pupa.

72

Journal New York Entomological Society [Vol. xlviii

Eggs laid singly. Larva makes a nest by spinning leaves
together and makes a slight cocoon to pupate. Locust and
other legumes.

Jamesburg, May 17, (C) — Dennisville, Sept. 5, (C, Wat).

On wing throughout season.

Achalarus Scudder

496. lyciades Geyer. Throughout state. Fields, meadows and
wood roads. May-July. One brood. Sometimes locally
abundant. Rests with open wings while feeding on flowers.
Tick trefoil {Desmodium panicidatwm) preferred (Wat)
and bush clover (Lespedeza) .

Arlington, May 18, (Ru) — Greenwood Lake Glens, July
18, (Wat).

Green Village, Jnne 6, (C) ; Lake Hopatcong, June 15,
(Wat) ; Plainfield, June 21, (Ru) ; 5-Mile Beach, June,
July, (Hm).

Thorybes Scudder

503. bathyllus Abbot & Smith. Throughout state. May-Sep-
tember. One brood. Winters as full grown larva. Bush
clover {Lespedeza capitata) preferred (Wat), other
legumes.

Jamesburg, May 24, (C) — Dennisville, Sept. 7, (C, Wat).

Cranford, June 10, (Ru) ; Newton July 6, (C) ; 5-Mile
Beach, May-Aug., (Hm).

505. pylades Scudder. Throughout state. Fields and open
woods. June-July. One brood. Winters as a full grown
larva. Eggs laid singly on under side of leaves. Larva
makes a nest by cutting and folding leaves. Clover and
other legumes.

Hemlock Falls, May 25, (Wat) — Lake Hopatcong, Aug.
10, (C).

Great Notch, May 25, June 6, (Ru) ; Palisade Park, June
7, (Hs) ; Green Village, June 13, 16, (C) ; Greenwood Lake
Glens, June 25, (Wat) ; 5-Mile Beach, May-Aug., (Hm).

Pyrgus Hiibner

515. centaureae Rambur. Crystalline Highlands. April-May.

Mae., 1940]

Comstock: Butterflies

73

One brood. Life history unknown. Formerly numerous at
Paterson and Great Notch.

Normanock, April 25, (C, dP, Wat) ; Paterson, April 25,
May 14, (Gr) (Sm), May 11, (Wat) ; Westwood, April 30,
May 19, (Mt) (Sm) ; Great Notch, May 2, 6, (Ru) ; May 15,
(C) ; Montclair, May 5, (Bent) ; Hewitt, May 18, (Wat) ;
Awosting-, May, (Hs) ; Iona, April 30, (Sk) (Sm) ; Pomp-
ton, May 1-6, (C, dP, Ru).

521. communis Grote. Synonym tessellata Scudder. Piedmont
Region and southward. Dry fields and road sides. Aug-
ust-September. One brood. Winters as larva. Sometimes
locally abundant. Sida spinosa (Wat), Malva rotundifolia,
Abutilon abutilon, Hibiscus trionum. Bowne, Sept. 12, 21.
Eggs and larvae (C).

Lakehurst, May 10, (Hs) — Bowne, Oct. 12, (C).

Stockton, Aug. 8, (C) ; Bowne, Aug. 21, 31, Sept. 12, 16,
(C) ; Fort Lee, Aug. 2, (C) ; Nutley, Aug.-Sept., (New) ;
Dennisville, Sept. 5, (C, Wat) ; Lakehurst, Sept. 10, 11,
(Ru) ; 5-Mile Beach, Aug.-Oct., (Hm) ; Cape May, Aug. 27,
(Pk).

Pholisora Scudder

531. Catullus Fabricius. Throughout state. Roads and waste
places. May-September. Two broods. Winters as full
grown larva, pupating in spring. Eggs laid singly. Larva
forms a nest of leaves. Pigweed (CJienopodium album),
Amaranthus (Wat).

Jamesburg, May 24, (C) — Woodridge, Sept. 10, (Mt).

Throughout season, generally common.

Erynnis Schranck

541. icelus Scudder & Burgess. Northern District. Open
woods. May-June. One brood. Winters as full grown
larva, pupating in spring. Eggs laid singly on upper sur-
face of leaves. Larva makes a nest of leaves and lives
within for winter quarters. Willow, poplar (Wat).

Orange Mountains, May 1, (Bz) (Sm) — Greenwood Lake
Glens, June 26, (C, Wat).

74

Journal New York Entomological Society [Vol. xlviii

Jamesburg:, May 8, 17, 24, 30, (C) ; Hillsdale, May 10,
(C) ; Lake Hopatcong, June 15, (Wat) ; Newton, July 23,
larva (C, Wat) ; Ogdensburg, May 30, (Bl),

542. brizo Boisduval & LeConte. Throughout state. Open
woods. April-June. One brood. Winters as full grown
larva pupating in spring. Oak.

Jamesburg, April 18, (C) — Lake Hopatcong, June 15,
(Wat).

South Orange Mountain, April 19, (Wf) ; Jamesburg,
April 18, 24, May 8, 17, 24, 30, (C) ; Hope, May 5, (C) ;
Hillsdale, May 10, (C) ; Lakehurst, May 22, (Wat) ; Pomp-
ton, May 1, (C, dP, Ru).

547. lucilius Scudder & Burgess. Northern District. May-
August. Two broods. Larva makes a nest of leaves and
when full grown so winters, pupating in spring. Egg laid
singly on under surface of leaf. Wild columbine {Aqui-
legia canadensis).-

Paterson, April 19, (Or) (Sm) — Andover, Sept. 16 (C).

Great Notch, May 3, (Bl) ; Greenwood Lake Glens, May
9, June 25, (Wat) ; Ogdensburg, May 22, June 3, July 11,
Aug. 25, (Bl) ; Newton, July 5, 18, 23, larvge (C, Wat).

546. persius Scudder. Possibly throughout state. Open fields
and roads. April-May. Believed to be single brooded (Dr.
Wm. T. M. Forbes). Winters as full grown larva. Wil-
lows and poplars (Scudder).

Old Bridge, April 26, Jamesburg, May 17, 24, Runyon,
May 19, 26, Lakewood, May 22-24, (C) ; Greenwood Lake
Glens, May 5, (Bl).

. baptisiae Forbes. Possibly throughout state. May- August.

Two broods. Winters as full grown larva. Wild indigo
{Baptisia tinctoria) (Wat).

Greenwood Lake Glens, May 31, Fort Lee, Aug. 2, Water-
loo, Aug. 5, Lake Hopatcong, Aug. 12, Hemlock Falls, Aug.
17, (C)..

Note. For E. lucilius, persius, and haptisue see ^‘The
Persius Group of Thanaos” by William T. M. Forbes.
Psyche, Vol. XLIII, No. 4, pp. lOJ-113 (1936). The

Mar., 1940]

Comstock; Butterflies

75

records given for the above three species were carefully
verified.

550. martialis Scudder. Northern District. Fields and wood
roads. May-June and July-August. Two broods. New
Jersey tea {Ceanothus americanus) .

Great Notch, April 27, (Bl) — Woodbury, Sept. 5, (Sk)
(Sm).

Laurel Springs, April 29, (Hy) (Sm) ; Ogdensburg, May
22, 31, June 3, July 13, Aug. 11, 21, (Bl) ; Hemlock Falls,
May 25, 31, June 8, (C, Wat) ; Greenwood Lake Glens, May
30, June 22, 25, (C, Wat) ; Jamesburg, June 15, (C) ; New-
ton, July 23, (C, Wat) ; Waterloo, Aug. 5, (C).

551. Juvenalis Fabricius. Throughout state. Wood roads, oak
scrub. May-July. Two broods. Winters as full grown
larva. Eggs laid singly. Oaks, less commonly legumes
(Wat).

Jamesburg, April 18, (C) — Newton, July 23, (C, Wat).

Jamesburg, April 24, May 8, 17, 24, 30, June 15, (C),
July 17, (Pk) ; Lakewood, April 21, (C) ; Greenwood Lake
Glens, May 3, 9, 30, (C) ; Bear Lake, Bamapo Mountains,
May 3, (C).

554. horatius Scudder & Burgess. As petronius Lintner
(Smith’s “List”). Throughout state. Open fields. May-
August. Two broods. Oaks, less commonly legumes.

Orange Mountains, April 25, (Ru) — Belleville, Aug. 18,

(C).

Old Bridge, April 26, (C), Aug. 4, (Wat) ; Pompton,
May 1, (C) ; South Lakewood, July 12, (Wat) ; Jamesburg,
July 17, (Pk) ; Ramsey, July 28, (C) ; Waterloo, Aug. 5,
(C).

Ancyloxypha Felder

567. numitor Fabricius. Throughout state. Rich grassy fields.
May-September. Three broods. AVinters as larva. Eggs
laid singly. Larva forms nest of a grass blade. Grasses.

Jamesburg, May 30, (C) — Carlstadt, Sept. 18, (C).

Jamesburg, June 15, 20, 30, (C) ; Newton, July 6, 23, (C,
AVat) ; Ramsey, July 28, (C) ; Old Bridge, Aug. 4, (AA^at) ;

76

Journal New York Entomological Society [Vol. xlviii

Bowne, Aug*. 19, (C) ; Dennisville, Sept. 6, (C, Wat) ;
5-Mile Beach, May-Sept., (Hm).

Hesperia Pabricius

581. metea Scudder. Throughout state. May-June. One
brood. Grasses.

Hemlock Falls, May 6, (C) — ^Lake Hopatcong, June 15,
(Wat).

Jamesburg, May 8, 17, (C) ; Lakehurst, May 10, (Hs),
May 20-27, (Ds) (Sm) ; Clementon, May 10, (Kp), May 15,
(Lt) ; Millburn, May 4, 20, 28, (Ru).

591. leonardus Harris. Throughout state. Dry locations, on
flowers. August-September. One brood. Winters as a
small larva. Eggs laid singly. Grasses, Agrostis.

Port Lee, Aug. 23, (C) — Point Pleasant, Sept. 17, (C,
Wat).

Jamesburg, Aug. 25, (C, Wat) ; Lakehurst, Aug. 30, Sept.
2, (Ru) ; Atco, Sept. 5, (Lt) ; Dennisville, Sept. 5, (C,
Wat) ; Hillsdale, Sept. 9, (C).

595. attains Edwards. Pine Barrens. Swamps. July- August.
One brood.

Lakehurst, June 24, 30, Aug. 14, 29, (Ru), July 22-Aug.
5, (Wf), July 10, 17, (Bz) (Sm) ; South Lakewood, July
12, (C, Wat) ; Da Costa, July 19, (W) (Sm) ; Clementon,
Aug. 3, (Fox) (Sm).

598. sassacus Harris. Northern District. Meadows and fields.
May-June. One brood. Winters probably as pupa (Scud-
der). Eggs laid singly on any low plant in grass. Grasses,
Panicum.

Newfoundland, May 29, (Ds) (Sm) ; Greenwood Lake
Glens, May 31, June 16, 22, 26, (C, Wat) ; Cranford, June
9, (Ru) ; Palisade Park, June 7, (Hs) ; Alpine, June 16,
(C).

Hylephila Billberg

601. phylaeus Drury. Throughout state. Meadows and grassy
places. July-September. One brood.

Englewood, July 21, (Wat) ; Arlington, Aug. 18, (C) ;
5-Mile Beach, Aug. 20, (Hm) (Sm) ; Lakehurst, Aug. 26,

Mar., 1940]

Comstock: Butterflies

77

(Rii) ; Lake Hoptacong, Aug. 29, (Ds) (Sm) ; Dennisville,
Sept. 7, (C, Wat) ; Bowne, Sept. 12, (C).

Atalopedes Scudder

602. campestris Boisduval. Probably from Fall Line south-
ward. August-September. One brood. Bermuda grass
(Cynodon dactylon).

5-Mile Beach, July 26, Sept. 20, (Hm) (Sm) ; Cranford,
Aug. 1, (Bz) ; New Brunswick, Sept. 23, (Wat) ; Cape May,
Sept. (Sk) (Sm) ; Camden, Aug., Sept., (Cy).

Polites Scudder

609. verna Edwards. Throughout state. Damp, open fields.
June-August. One brood.

Elizabeth, June 20, (Bz) (Sm) — Janiesburg, Aug. 25,

(C).

Runyon, July 2, Aug. 4, (C) ; Greenwood Lake Glens,
June 22, 25, (Wat), July 18, (C) ; Lakehurst, July 11,
(Hs) ; 5-Mile Beach, June 23, Aug. 12, (Hm) (Sm).

610. manataaqua Harris. Throughout state. Fields and moist
meadows. June-July. One brood.

Elizabeth, June 20, (Bz) (Sm) — Cape May, Sept. 18,

(Pk).

Greenwood Lake Glens, June 22, 25, (Wat), July 18, (C,
Hs) ; Jamesburg, June 30, (C), July 2, (Pk), July 4, Aug.
25, (Wat) ; Lakehurst, July 7, (Hs) ; Ogdensburg, July 9,
(Bl) ; Ramsey, July 10, (Wat) ; West Cape May, July 16,
(Wat) ; Dennisville, Sept. 5, (C, Wat).

611. themistocles Latreille. Synonym cernes Boisduval & Le-
Conte. Throughout state. Open country. June-Septem-
ber. Two broods. Winters as pupa. Eggs laid singly.
Larva makes a nest and a slight cocoon for pupa. Grasses
{Panicum ) .

Hemlock Palls, June 8, (C) — Bowne, Sept. 12, (C). On
the wing in June and again in August and in September
usually plentiful.

614. peckius Kirby. Throughout state. Open country. May-
June and August-September. Two broods. Winters as
full grown larva and pupa. Eggs laid singly. Grasses.

78

Journal New York Entomological Society [Vol. XLVIII

Jamesburg, May 30, (C) — Point Pleasant, Sept. 17, (C,
Wat). On the wing in May and June and again through
August and September, usually common. There are a few
July records.

618. mystic Scudder. Northern District. In moist locations.
June-July. One brood. Winters as larva. Eggs laid
singly. Larva makes tubular nest of grass blades. Grasses.

Cranford, June 7, 9, 20, (Ru) ; Green Village, June 9,
(Ru) ; Lake Llopatcong, June 15, (Wat) ; Greenwood Lake
Glens, June 16, 22, 25, (C, Wat) ; Newton, July 5, (C) ;
Ramsey, July 21, (Ge).

Catia Godman

621. otho egeremet Scudder. Throughout state. Open fields.
June-August. One brood. Winters as larva. Grasses
(Paniciim).

5-Mile Beach, June 3, Aug. 27, (Hm) (Sm) ; Jamesburg,
June 21, July 4, Aug. 25, 29, (C, Wat) ; Newton, July 6,
11, 23, (C, Wat) ; Greenwood Lake Glens, June 23, July 18,
(C) ; Ramsey, July 28, (C).

Note, otho otho Abbot & Smith occurs in the south.

Poanes Scudder

622. viator Edwards. Locally, throughout state. Marshy land.
July-August. One brood. Occasionally numerous in salt
meadows. Grasses.

Arlington, Julj^ 19, Kearny, July 19, (Ru) ; Snake Hill,
July 11, Aug. 2, (C, Wat) ; Anglesea, July 25, (Hm) ;
Atlantic City (Aa).

623. massasoit Scudder. Locally, throughout state. Fresh
water swamps. July- August. One brood. At times com-
mon. Grasses.

Westville, July 3-10, (Sk) (Lt) (Sm) — Westwood, Aug.
7, (Mt) (Sm).

Jamesburg, July 5, (C) ; Englewood, July 13, 21, (C) ;
Helmetta, July 27, (Hs) ; Ramsey, July 28, (C) ; Green
Village, July 2-22, (Ru).

Form suffusa Laurent. Type locality, Westville, July 4,

Mar., 1940]

Comstock: Butterflies

79

(Lt). Of fairly frequent occurrence there and at Malaga,
July 17, (HI, Wat).

624. hobomok Harris. Northern District. Edges of woods,
fields and meadows. May-July. One brood. Winters as
egg, larva and pupa (Lt). Grasses.

Paterson, May 11, (Gr) (Sm) — Westwood, July 5, (Mt)
(Sm).

Hemlock Falls, May 24, (C) ; Greenwood Lake Glens,
May 31, June 16, 22, 25, (C, Wat) ; Jamesburg, May 31,
June 1, 20, (C) ; Palisade Park, June 7, (Hs).

Form female pocahontas Scudder flies with typical form.

625. zabulon Boisduval & LeConte. Locally, throughout state.
Open woods and fields. May-September. Two broods.
Grasses.

Belleville, May 23, (C)— Hillsdale, Sept. 9, (C).

Jamesburg, May 30, June 15, (C), Aug. 25, Sept. 3,
(Wat) ; Cape May, May 30, (Sk) (Sm) ; Manasquan, July
24, (C) ; New Brunswick, Aug. 19, (Hs) ; 5-Mile Beach,
June 19, Aug. 21, (Hm) (Sm) ; Newark, Aug. 28, (C) ;
Hillside, Sept. 5, (Ru).

628. aaroni Skinner. Coastal District. Salt marshes. June-
September. Two broods. Grasses.

Cape May, type locality (Sk), June 14, (Sk), Sept. (St,
Pk) ; Anglesea, June 12, (Lt) ; 5-Mile Beach, June 15,
Aug. 22-27, common in salt meadows (Hm) ; Dennisville,
Sept. 7, (C, Wat) ; Tuckerton (Ds).

Atrytone Scudder.

634. arogos Boisduval & LeConte. Pine Barrens. Swamps.
July- August. One brood.

Brown’s Mills Jc., July 21, (Dke) (Sm) ; Brookville,
July, (Ds) (Sm) ; Lakehurst, June 29, (Ru), Aug. 30, (Ds)
(Sm). • .

635. logan Edwards. Dr. Smith lists vitellius Fabricius, a West
Indian species. His records refer to this species. Through-
out state. Damp fields. One brood. July. Grasses.

Lakehurst, June 29, (Ru), July 7, (Hs) ; Jamesburg,
July 4, (C, AVat), July 30, (Ds) (Sm) ; Ramsey, July 10,
(Ge) ; Westville, July 10, (Lt) (Sm) ; Lakewood, July 12,

80

Journal New York Entomological Society [Vol. xlviii

(C) ; Stockholm, July 8, (Bl) ; Ogdensburg, July 11, 13,
(Bl) ; Hewitt, July 19, (Hs).

638. dion Edwards. Pine Barrens, Appalachian Valley. July-
September. Grasses.

Great Meadows, June 30, July 4, (dP) ; Anglesea, July 7,
(Hm) ; Stockholm, July 8, (Bl) ; Lakehurst, July 22, Aug.
5, (Bz, Wf), July 19, (Hs), July 29, (Ds) (Sm) ; James-
burg, Aug. 24, 31, Sept. 3, (Wat) ; Brown’s Mills Jc., Sept.
15, (Dke) (Sm).

640. conspicua Edwards. Synonym pontiac Edwards. Through-
out state. Damp meadows. June-August. One brood.
Grasses.

Green Village, June 20, July 14, (Ru) ; Jamesburg, July
4, (Lt) (Sm) ; Westville, July 4, (Sk) (Sm) ; Westwood,
July 4, 28, (Mt) (Sm) ; Newton, July 6, (C) ; Englewood,
July 7, 13, (C) ; Lakewood, July 13, (C, Wat) ; Malaga,
July 17, (HI, Wat) ; Ramsey, July 24, (Ge) ; Old Bridge,
Aug. 4, (Wat).

641. bimacula Grote & Robinson. Pine Barrens, Appalachian
Valley. Swampy spots. June-July. One brood. Grasses.

Greenwood Lake Glens, June 16, 22, (C, Wat) ; Lake-
hurst, June 22, (Ru), June 24, (Wf), June 27, (Ds), July
7, 11, (Hs) ; Oak Ridge, June 26, July 3, (Sh) (Sm) ;
Mashipacong, July 3-5, (HI, Wat) ; Ogdensburg, July 7,
(Bl) ; Stockholm, July 8, (Bl).

642. vestris Boisduval. Synonym metacomet Harris. Through-
out state. July. One brood. Grasses.

Runyon, July 2, (C) ; Jamesburg, July 4, (C) ; Newton,
July 10, 18, 23, (C, Wat) ; Green Village, July 15, (C) ;
Newark, July 25, (C) ; Greenwood Lake Glens, July 18,
(C) ; Ramsey, July 26, (Ge) ; 5-Mile Beach, July 27, (Hm).

Atrytonopsis Godman

644. hianna Scudder. Pine Barrens. May-June. One brood.
Grasses.

South Lakewood, May 20, (Wat) ; Brown’s Mills Jc.,
May 27, (Dke) (Sm) ; Iona (Dke) (Sm).

Mar., 1940]

Comstock: Butterflies

81

Amblyscirtes Scudder

660. vialis Edwards. Throughout state. Moist roadsides. May-
June. One brood in New England but in south two broods
(Scudder). A single brood probably in New Jersey.
Grasses.

Clementon, May 15, (Hg) ; Lakehurst, May 20, June 4,
(Ds) (Sm) ; Lakewood, May 22-25, (C, Wat) ; Jamesburg,
May 24, (C) ; Newfoundland, May 29, (Ds) (Sm) ; Ogdens-
burg. May 30, (Bl).

661. hegon Scudder. Synonym samoset Scudder. May-June.

Ogdensburg, May 31, (Bl) ; Green Village, June 8, (Ru).

Lerodea Scudder

671. fusca Grote & Robinson. Coastal Plain and a little above
the Pall Line. Moist meadows. June- July and August-
September. Two broods. Grasses.

5-Mile Beach, June 9, (Hm) ; Green Village, June 13, 16,
(C), Aug. 19, (Ru) ; Jamesburg, June 15, 20, (C), June 22,
(Ds) (Sm), July 4, (Sk) (Sm), Aug. 25, 29, Sept. 3,
(Wat) ; Camden, June 18, (Sk) (Sm) ; Cape May Court
House, July 17, (HI, Wat) ; Ramsey, Aug., (Ge) ; Mana-
squan, Aug. 24, (C) ; Tuckerton, Sept. 2, (Ds) (Sm).

Prenes Scudder

680. panoquin Scudder. Coastal District. Salt marshes. June-
September. Two broods.

Atlantic City, June 25- July 1, (Aa) ; 5-Mile Beach,
June-Sept., common in salt meadows (Hm) ; Palermo, July
17, (HI, Wat) ; Dennisville, Sept. 7, (C, Wat).

683. ocola Edwards

Bowne, Aug. 21, Sept. 12, ’37, (C) ; Lakehurst, Aug. 24,
’22, (Ru) ; Camden, Sept., common 1899 (Cy) (Sm) ;
Salem (Aa) (Sm) ; Atlantic City (Aa).

Supplemental List

Included here are species of which sixteen have been captured

in the state of New Jersey and three on Staten Island or Long

Island. Of these fifteen species are of regular occurrence south

of New Jersey. The records of these are given and under favor-

82

Journal New York Entomological Society [Vol. XLVlii

able circumstances more records of their occurrence may be ex-
pected but none of these species can now be regarded as a

regular inhabitant of the state.

Phyciodes Hiibner

266. batesii Reakirt. One brood. Wavy-leaf aster (Aster
undulatus). The only record is from Gloucester in the
original description in 1865. The species is of regular oc-
currence in western Pennsylvania and central New York
State.

Basilarchia Scudder

321. arthemis Drury. White admiral. And form proserpina
Edwards. In the last edition of Smith’s ‘‘List” these were
recorded. This is a northern species, frequent in the Cats-
kill region of New York state and northward. It is believed
to be out of range, even in the northwestern part of New
Jersey. The reported captures are believed to be referable
to B. astyanax albofasciata Newcomb. The recorded food
plants of arthemis are black and yellow birch, basswood and
willow. Two broods. Winters as larva.

Megisto Hiibner

101. mitchellii French. Dover, Charles W. Johnson, July 10,
1890 (Sm). Identified by Dr. Henry Skinner. Woodport.

102. sosybius Fabricius. Southern New Jersey. Grasses.
Mount Holly (Aa) (Sm). The species is abundant in the
southern states.

Atlides Hiibner

356. halesus Cramer. Occasional visitor, southern species.
Mistletoe. Cape May, Gloucester, AYestville (Aa) (Sm).

Strymon Hiibner

365. cecrops Fabricius. Occasional visitor, southern species.
Manasqnan, June 29, one male (Br) (Sm) ; East Marion
L. I., N. Y., Aug. 14, two specimens (La).

374. favonius Abbot & Smith. Occasional visitor, southern spe-
cies. Anglesea, Sept. 1, two specimens (Hm) (Sm).

Mar., 1940]

Comstock: Butterflies

83

376. autolycus Ontario Edwards. Newark, bred from larva
found on oak, May 29, emerged June 13, 1892 (Doll collec-
tion). This is a very scarce form of which there are about
a score of records during a half century. These records
extend from Missouri to Massachusetts and Canada.

Erora Scudder

418. laeta Edwards. Atlantic City, July 1, (Aa). Two broods.
This species is apparently rare but extends over a very wide
range occurring abundantly in Arizona, and recorded scat-
ter ingly in Pennsylvania, New York, New England states
and eastern Canada.

Papilio Linnaeus

21. palamedes Drury. Pare visitor from south. Red bay
{Per sea borhonia). West Hoboken, July 8, 1908 (Ws?) ;
Delaware, Aug. 20, 1934, (Sto) ; Fort Wadsworth, S. I.,
N. Y., June 18, (Bu).

Appias Hiibner

78. ilaire Godart. Southern species. Canarsie, L. I., N. Y.,
June 10, 1906, two specimens (My) (det. Wat).

Phoebis Hiibner

58. philea Linnaeus. Southern species. Asbury Park, Sept. 9,
(Eng) ; Riverside Park, New York City, Oct. 13, (Eng).

60. agarithe Boisduval. Southern species. Beach Haven, Sept.
3, male (Frd).

Zerene Hiibner

55. caesonia Stoll. Dog-faced butterfly. Southern species.
Eggs laid on under side of end leaves. False indigo {Amor-
pha fruticosa), also reported on clover. Flushing, L. I.,
N. Y., July 12, (Bl) ; Staten Island, N. Y., June, July, 1896
(Ds) (Sm).

Goniurus Hiibner

486. proteus Linnaeus. Coastal District. August-September.
Very rare as far north as New Jersey. Injurious to beans

84

Journal New York Entomological Society [Vol. XLVIII

in the southern states. Legumes. 5-Mile Beach, (Hg)
(Sm) ; Cape May (Aa) (Sm).

Cecropterus Herrick-Schaeffer

500. cellus Boisduval & LeConte. Southern species. Hog pea-
nut {Amphicarpa pitcheri), Breweria aquatica. Newark
(Sm).

Polites Scudder

619. brettus Boisduval & LeConte. Southern species. August-
September. Camden, 1 female, Aug. 10, 2 males. Sept. 7,
(Cy) (Sm).

Oligoria Scudder

652. maculata Edwards. Southern species. Camden, 1 male, 1
female, July 15, (Cy) (Sm).

Lerema Scudder

653. accius Abbot & Smith. Southern species. Grasses. Salem
(Aa) (Sm) ; Atlantic City, July, (Aa).

Calpodes Hiibner

677. ethlius Cramer. Canna skipper. Southern species. No
record in state but taken at several places on Long Island
in 1911. Occasionally destructive to canna. Yaphank,
L. I., May 27, (Fr) ; Floral Park, L. I., larva. Sept., Oct.,
(Eng).

Mar., 1940]

White: Japanese Beetle

85

THE RELATION OF ANTS TO THE JAPANESE
BEETLE AND ITS ESTABLISHED
PARASITES^

By Ralph T. White

Bureau of Entomology and Plant Quarantine, U. S. Department
OF Agriculture

INTRODUCTION

It has long been known that many ants derive most of their
food from dead and dying insects, and ants have been observed
molesting both adult and larval stages of the Japanese beetle
(Popillia japonica Newm.). Many observations and experi-
ments have therefore been conducted in the vicinity of Moores-
town, N. J., both in the field and in the inseetary, to determine
what relationship, if any, exists between ants (Pormicidse) and
the Japanese beetle or its established parasites in the field.

Observations and diggings of the common ants, chiefly those
frequenting habitats favorable to the various stages of Popillia
japonica in this area, are recorded herein.

OBSERVATIONS OF ANT COLONIES IN 1933

In 1933 diggings were made in and near certain ant colonies in
grub-infested areas in late summer and early fall to determine
the grub population within the colony itself as well as in the
area immediately surrounding such colonies. In general the grub
population within the colonies was about the same as that near
the colonies, although in some cases more grubs were found in the
colony. The following detailed account of three large colonies
examined in 1933 will give the reader a true picture of an average
ant community in pastures of this area.

1 The ants used in this study were identified by M. E. Smith, C. F. W.
Muesebeck, W. M. Mann, and the late W. M. Wheeler. Acknowledgement
is given to E. J. Sim, formerly with the Japanese Beetle Laboratory, for
assistance in collecting specimens from various points in New Jersey, and
to J. L. King for helpful suggestions and counsel in the preparation of this
manuscript.

86

Journal New York Entomological Society [Vol. xlviiI

Colony Site G-5 (Hercher’s Pasture)

Herclier’s pasture, located 3 miles from the Moorestown lab-
oratory, was chosen for these studies because of its nearness to
the laboratory and the willingness of the owner to permit the
making of periodic surveys. Diggings in this pasture on August
8, 1933, showed the grub population to range from 1 to 32 per
square foot, and according to seasonal surveys the average popu-
lation ' for August 1933 was 9 per square foot. In the 18-
square-foot area covered by this colony the average grub popula-
tion was 12.2 per square foot. Grub populations in 10 diggings
at random within a 25-foot radius of this infested area, but con-
taining no ants, were as follows : 2, 8, 12, 16, 1, 8, 5, 11, 9, and 6
per square foot, or an average of 7.8 per square foot.

The species of ants in the main colony chosen was Formica pal-
lide-fulva schaufussi var. incerta Emery, and small colonies of
Lasi'us (Acanthomyops) claviger (Roger), and L. niger var.
americanus Emery were living within the same area. The main
colony of Formica covered 1 square foot. Only one entrance was
visible, and the Lasivs colonies were unnoticed until some soil
was removed. The Formica workers had previously been seen
bringing a dead larva of the eastern tent caterpillar {Malacosoma
americana (F.) ) to their formicary, and also a dead adult ground
beetle {Calosoma sp.). They fed readily on the juices of a freshly
killed Japanese beetle grub placed upon their mound, and later
dragged it beneath the ground.

Colony Site G-3 (Hercher’s Pasture)

This colony, of Formica fiisca var. suhsericea Say, was also in
Hercher ’s pasture but 200 yards to the west and on a lower well-
sodded area. During the summer the refuse heaps were examined
and elytra of various Coleoptera, including one Japanese beetle,
were found. Japanese beetle grubs were found at the edge of
this colony throughout the summer. Diggings within the colony
were limited during the summer, as the colony was under obser-
vation for feeding habits and general activities. On June 6,
however, a small clump of grass within the colony boundary was
removed very carefully, and one grub was found among the roots
just beneath the surface. Ants were comparatively numerous

Mar., 1940]

White: Japanese Beetle

87

here, and when the grub was exposed they attacked it eagerly.
Within one-half hour they had dragged the much weakened grub
into an entrance, and ant activity again quieted down to normal.

Freshly killed grubs were placed directly upon the mound as
well as very near to it. The body walls of some had previously
been punctured to allow the body fluids to exude, and in such
cases the ants would swarm over the remains, first feeding upon
the contents and then dragging the remains into an entrance hole.
The unpunctured cadavers were unmolested until foraging work-
ers later chanced upon them and dragged them within their
formicary.

When a living grub was placed upon the surface of the mound
or at the edge where ants were continually passing, it was at-
tacked by these ants and taken into their formicary. If, on the
other hand, the grub was placed barely beneath the surface of
the soil, it remained unmolested.

On July 14 at the colony site a living Japanese beetle was ob-
served in combat with several ants. The beetle alighted upon the
bare ground, where two ants immediately offered battle, and for
half an hour a struggle was carried on during which other ants
were attracted to the scene. The beetle was Anally overpowered
and dragged into the entrance.

On July 18 a section of the colony was excavated, revealing a
Japanese beetle pupa. This pupa had been unmolested by the
ants, although egg galleries and runaways formed a netwmrk
around it. As soon as it was disturbed, however, ants immediately
attacked and carried it away. In another section of this colony
three Japanese beetle larvse were found unharmed just below the
surface. Three were also found at the edge of the main colony
area. A dead Japanese beetle was also found within the colony
and may have been the one observed 4 days previously.

On September 11 the remainder of this colony was examined.
Systematic diggings extended 4 feet to the north and to the south
of the area of concentrated activity. The colony area was shaded
at times during each day by two trees growing nearby. The area
at the south, being covered by shorter grass than that at the
north, was more attractive to the beetles for oviposition, and
consequently grub population was denser. The total area dug in
this colony was 52 square feet and contained 139 grubs, or an

88

Journal New York Entomological Society [Vol. xlviii

average of 2.6 per square foot. In the central area, where the
ants were consistently most numerous, the grub population aver-
aged 1.5 per square foot, while in the bordering bare-ground area,
where fewer ants existed, only 1 grub was found per square foot,
probably because bare ground is unfavorable for oviposition. If
the ants had molested them, there would have been fewer grubs
within the colony than in the adjacent area with identical ground
cover.

Colony Site S (Haddonfield Pasture)

This colony was in a pasture near the Tavistock Golf Course,
Haddonfield, N. J. It was on a sodded knoll, the surrounding
area sloping toward a swamp. The ants comprising this colony
were the common Lasius (A.) claviger, and were hidden com-
pletely by an old burlap bag, beneath which the ground was en-
tirely devoid of vegetation. When examination of this colony
was begun, on November 1, 1933, thousands of winged males and
females were present. The area not covered by the bag was
densely sodded, and because this species was more or less hy-
pogeal, its presence was not evident. Diggings showed these ants
to be established over an area of approximately 8 square feet.
Grubs in this area averaged 3 to a square foot, whereas the aver-
age in a 17-square-foot area immediately surrounding was only
2.6 per square foot. Additional diggings at random around the
periphery of this colony and not more than 20 feet from the col-
ony center showed an average of 1.3 beetles per square foot. The
observations made in the course of these diggings further exem-
plify the negative effect of this species upon grubs in the soil.

FIELD SURVEYS IN 1934 AND 1935

In 1934 surveys were continued in areas infested with ants
and corresponding areas not thus infested. The data in Table 1
show that in 35 separate diggings of 1 square foot each a total of
210 grubs, or an average of 6 per square foot, was found where
no ants were present, while in the same number of diggings of
the same size and in the same localities with various species of
ants present there was a total of 213 grubs, or 6.1 per square foot.
These surveys indicate rather conclusively that at least these
species of ants had no ill effect upon grubs present in the soil.

Mar., 1940]

White: Japanese Beetle

89

TABLE 1. — Field surveys of populations of Japanese beetle grubs in areas
in New Jersey infested with ants and in surrounding
uninfested areas, 1934

Grubs per
square foot

Date and location

in areas —

of digging

With

ants

With-

out

ants

September 18:

' 8

6

3

7

Vincentown cemetery

Formica f usca var. subsericea Say

4

5

5

9

4

0

Vincentown pasture

Pheidole pilifera (Koger)

4

3

Pheidole vinelandica Forel

3

2

September 19 :

7

4

Vincentown pasture

Lasius niger var. neoniger Emery

3

4

5

7

^ 11

10 .

9

9

Vincentown roadside

Aphcenogaster fulva subsp. aquia

7

7

(Buckley)

7

2

5

2

September 26:

Pemberton pasture

1

f ^

0

Lasius (Acanthomyops) claviger ^

0

1

(Roger)

1 2

0

r 1

■4

Formica pallide-fulva subsp. ]

27

31

schaufussi var. incerta Emery

[ 21

16

September 27 :

27

26

Pemberton pasture

Lasius niger var. neoniger Emery j

1

3

Pemberton orchard

Formica fusca var. subsericea Say •<

1 3
1 4

2

7

1 1

0

October 3 :

r 2

0

Haddonfield pasture

Myrmica group

1 1
1 3

1

0

1 0

2

October 4:

r 2

4

Haddonfield pasture

Lasius {Acanthomyops) claviger
(Roger)

1 0
] 3

0

2

1 5

6

Total

213

210

90

Journal New York Entomological Society [Vol. xlviii

During June and July 1935, 42 colonies of ants were examined
as well as corresponding areas containing no ants. Table 2 gives
data obtained from these surveys. In both the areas containing
ants and in corresponding check areas the average grub popula-
tion was 4.3 per square foot. Although this was less than the
population in 1934, it seems evident from these surveys that the
ants had no ill effect on the grub population.

Established Beetle Parasites in Relation to Ants

EFFECT OF ANTS ON ADULT TIPHIA

Ants are frequently seen foraging in the field on blossoms of
wild carrot {Daucus carota L.) and on the leaves of maple trees
(Acer spp.). Tiphia popilliavora Roh. obtains nectar from the
former and T. vernalis Roh. feeds upon the honeydew found on
the latter. Twice during the 1934 season the writer witnessed
conflicts between T. popilliavora adults and the common black
ant Formica fusca var. suhsericea. In each case a single ant had
obtained a firm hold on a female Tiphia and both fell helplessly
to the ground. In one case the Tiphia gained her freedom* as
they struck the ground, but in the other the ant retained its hold
and, after several minutes of fierce battling, with the aid of two
other worker ants carried the Tiphia to the formicary. Her
wings became ver}^ badly damaged before she was finally lost to
sight.

In May 1934 a battle between individuals of the same species
of ant and a pair of Tiphia vernalis adults was observed at Phil-
mont, Pa. This male and female were mating on a maple leaf
directly over a mound of these ants, when they suddenly fell to
the ground, landing in the midst of ant activity, where they were
instantly seized by workers. During the melee the male escaped,
but the less fortunate female was injured and dragged within
the formicary, where presumably she succumbed.

Since Tiphia adults spend considerable time in the soil and on
low grass above the soil, ants probably cause the death of some.
One such instance was observed at the Overbrook Golf Course in
May 1934, while the author was collecting Tiphia for coloniza-
tion. A female was just emerging from the soil, presumably to
feed on honey solution sprayed on foliage nearby. Several for-
aging ants of Formica fusca var. suhsericea were present, and a

-Field surveys of 'populations of Japanese beetle grubs in areas in Neiv Jersey infested with
ants and in surrounding iininfested areas, 1935

Mar., 1940]

White: Japanese Beetle

91

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TABLE 2 — {Continued)

92

Journal New York Entomological Society [Vol. xlviii

S'S

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Mar., 1940]

White: Japanese Beetle

93

fierce battle ensued in which the wings of the Tiphia became dam-
aged, preventing her flight and enabling the ants to drag her
into their nest without further opposition. Incidents of this type
are exceedingly interesting to watch and may happen rather
frequently, but are probably of little economic importance.

EFFECT OF ANTS ON TIPHIA COCOONS

An instance of damage to Tiphia cocoons in the field by Sole-
nopsis molesta was observed in 1932, when several cocoons held in
vials stoppered with fine wire screens and buried in the soil were
riddled. These tiny yellow ants were able to make entrance into
the vials through the screen. Again during the following season
cocoons buried in a similar manner were destroyed.

Cocoons formed naturally in the field have also been found
completely riddled with holes, and frequently the same species
of ant has been associated with them. Material held in the in-
sectary has sometimes been attacked by this ant, as will be re-
ported under ‘‘Insectary Experiments.” If colonies of Solenop-
sis molesta were more numerous in the Moorestown area, they
might easily prove an important factor working against parasite
establishment and subsequent increase.

EFFECT OF ANTS ON PARASITIZED GRUBS

No evidence has come to the writer’s attention of ants molest-
ing parasitized grubs in the field. It seems entirely plausible,
however, that some such helpless individuals should fall prey to
these industrious workers. This seems especially probable at
times of advanced parasitization by certain dipterous parasites,
and just prior to cocoon formation in case of hymenopterous
parasites. At such times the parasitized grub lies helpless, and
this as well as the parasite larva itself should be appetizing food
for many ants, which often are none too particular in their choice
of viands.

EFFECT OF ANTS ON FLY PUPARIA

In one colony of Formica f usca var. suhsericea the remains of
a Phyllophaga adult containing a fly puparium were found. This
aroused suspicion that certain Japanese beetles containing pu-
paria of Centeter cinerea Aid. might be destroyed by foraging

94

Journal New York Entomological Society [Vol. xlviii

ants, especially since a large proportion of beetles parasitized by
this fly crawl beneath dead leaves and trash to die. These places
are favorite hunting grounds for many ants, and such beetles
may fall victims of rapacious individuals oftener than is sup-
posed. Observations have not materially strengthened this sup-
position although in 1935 one beetle bearing a Centeter egg was
found in a colony of F. fiisca var. suhsericea. In areas of heavy
parasitization by C. cinerea instances of this nature might be of
significance.

Insectary Experiments

SPECIES OF ANTS STUDIED

Twenty- three of the most common species (or varieties) of
ants, found in this area, representing 11 genera, were used in ex-
tensive insectary experiments to determine if any relationship
exists between Pormicidse and Popillia jap^onica. The following
species were included :

Formica pallide-fulva subsp. scliaufussi var. incerta Emery

Formica f usca var. siibsericea Say

Formica fusca var. siibcBnescens Emery

Formica sangumea subsp. ruhicunda Emery

Formica truncicola subsp. integra Nylander

Formica sanguinea subsp. suhintegra Emery

Formica neogagates Emery

Formica pallide-fiUva subsp. nitidiveniris Emery
Solenopsis molest a (Say)

Lasius (AcantJiomyops) claviger (Eoger)

Lasius {Acantliomyops) interjectus Mayr
Lasius niger var. americanus Emery
Lasius niger var. neoniger Emery
Pheidole morrisi Forel
Pheidole pilifera (Eoger)

Pheidole vinelandica Forel

Crematog aster lineolata var. cerasi (Fitch)

Tetramormm ccespitum (L.)

Tapinoma sessile (Say)

Monomorium minimum (Buckley)

Prenolepis imparis (Say)

Aphcenogaster fulva subsp. aquia (Buckley)

Myrmica scahrinodis group

Mar., 1940]

White; Japanese Beetle

95

EXPERIMENTAL PROCEDURE

Two types of cages were used in this work and are hereafter
spoken of as large cages and small cages. The former type was
a wooden box 1 foot square by 2 feet deep. The upper 6 inches
on two sid^s was of glass instead of wood, permitting observa-
tion of ant activity within the cage. The top was hinged and
consisted of wire screen of very fine mesh, assuring ample ven-
tilation and entrance of necessary moisture. All wooden parts
of the cage were soaked in paraffin to prevent excessive decay
when buried. In the bottom several holes 1 inch in diameter were
bored and covered with wire screen similar to that used on the
top, to permit the entrance and escape of moisture. Further to
facilitate drainage, in some of the cages long slits were made in
the sides and likewise covered with screen. However, cages with-
out such slits remained in as good condition as the others, and in
excessively wet periods were in even better condition. During
such wet weather boards were placed over the cages.

These cages were buried in the soil to a depth of li feet, the
glass sides remaining above ground. Grass or wheat was planted
in the cages before the ants and grubs were placed there. Both
crops grow well and sometimes need to be cut back to prevent
undue shading of the ground. Dry leaves and a medium-sized
flat rock were also placed in each cage, as many ants prefer their
formicary beneath shelter, at least at the start, and hypogseic
species remain almost entirely under cover.

The small type of cage consisted of two plates of common win-
dow glass placed on a vertical frame so that there was a solid
bottom and a half-inch space between the plates to hold soil. A
cover consisting of a wedge-shaped piece of wood retained the
ants. By the use of glass sides ant activity was readily observed
without disturbing the enclosed individuals. It was found ad-
vantageous to darken one side of the cage with a black cloth or
heavy paper. Runways were well distributed throughout the
soil, but the ants always reared their young and congregated in
galleries on the darkened side.

The small cages can be kept at room temperature and ant ac-
tivity maintained throughout the year. Moisture must be added
every 5 or 6 weeks during the winter or in cool spring and fall

96

Journal New York Entomological Society [Vol. XLVIII

weather and at intervals of 2 to 3 weeks in the summer. Ants
confined in these cages for 3 months have been removed seem-
ingly in as vigorous condition as when first placed within. The
ordinary time of confinement, however, was approximately 30
days. If ants confined in these small cages for 30 days with
grubs do not molest them, it is doubtful that they would if left
longer. Most species, unless very small, will tunnel throughout
the soil within the first week, but some never penetrate to any
great depth.

Ants and grubs were placed in each of a series of these small
cages, and in some the ants were fed a candy made of white sugar
and honey, whereas in others they were not fed during their con-
finement. The ants that were fed lived longer and were kept
alive indefinitely by feeding. The absence of food did not seem
to force their attentions upon the grubs but merely shortened
their lives.

RESULTS

Accumulated data obtained from the numerous experiments
conducted in the insectary show that 54 cages containing ants
had at the time of examination at least as many grubs as the
corresponding check cages without ants. Of 901 grubs placed in
the cages, only 428 were accounted for after 30 days in the cages
containing ants and grubs, with 36 of this number being dead.
In the corresponding check cages without ants, after the same
number of days and receiving the same treatment, 487 of the 901
grubs were accounted for, of which 75 were dead. In other
words, 39 more dead grubs were found in cages without ants
than in cages containing ants, and likewise 59 more living forms
occurred in the checks than in the cages containing ants. A very
slight indication of the destruction of dead remains by ants
seems evident from these data, which would be of little conse-
quence unless such remains were diseased.

In experiments in which eggs of Poinllia japonica were placed
in the soil and left for development, a total of 270 was used
in experiments with ants and a corresponding number in cages
without ants. A total of 102 first- and second-instar grubs was
found in the cages with ants, whereas 105 grubs in the same stages
were found in the cages without ants at the time of examination.

Mar., 1940]

White: Japanese Beetle

97

From this small difference one cannot attribute serious effects by
the ants on the eggs or on the first and second instars.

In many of the cages some of the third-instar grubs became
adults during the course of the experiments, thus exposing at
some time the helpless prepupal and pupal stages. Ants, however,
paid no more attention to these than to the more active stages.

DAMAGE TO TIPHIA COCOONS

Damage by ants to Tiphia cocoons being held for development
has sometimes been reported. Such an instance was brought to
the attention of the author in 1934, when several cocoons held in
rearing trays were riddled with holes and their contents eaten
by the small yellow ant Solenopsis molest a. These trays are di-
vided into small compartments, and one parasitized larva is
placed in each section before it is filled with soil. The trays are
then stacked and held for development of the parasite larva and
subsequent cocoon formation. Sometimes grubs in these trays
come to the surface of the soil, and on one occasion an individual
confined within a compartment was observed to be overcome by
myriads of these ants. Solenopsis molest a is supposedly fond of
fats and greases, but the writer was successful in attracting them
to a sponge saturated with honey.

Owing to the slight indication noted in field studies that Tiphia
cocoons are occasionally riddled by ants, as well as to similar
observations in the insectary, further tests were made of such
tendencies. Both the small and the large types of cages used in
other experiments were tried. In each of the smaller cages, on
October 3, 1934, six parasitized grubs were placed in the soil with
ants, and corresponding check cages were set up without ants.
Development continued, and cocoons were visible on October 25.
In all cages after 3 months the ants were as active as when first
placed within, and in almost every case had tunneled throughout
the soil. Daily observations of several cocoons that were visible
failed to indicate any damage, although the ants had runways
throughout the soil and one cocoon was in their direct path for 2
months. When the soil was removed on February 15, 1935, only
one cage containing individuals of Lasius (Acanthomyops) clav-
iger gave evidence of possible damage to cocoons. In similar
later experiments with this species no interference resulted.

98

Journal New York Entomological Society [Vol. xlviii

Thirty-seven parasitized grubs were placed in each of 12 large
cages and allowed to form cocoons in the presence of ant colonies.
Four species of ants were used in these tests, namely, Lasius
(Acanthomyops) claviger, an ant of the My r mica scahrinodis
group, Pheidole morrisi, and T etramorium cccspitum. Equal
numbers of parasitized grubs were used in similar cages without
ants. These cages were buried outdoors in October 1934 and
left undisturbed until April 1935, at which time they were ex-
amined for cocoons. Three hundred and twenty-nine cocoons
were found both in the cages containing ants and in the checks
without ants, and none of these was damaged. This was 74
per cent of the total number of parasitized grubs used and in-
dicates a negative effect of at least these species upon parasitized
larvae or the resultant cocoons.

Since Solenopsis molest a had shown a tendency to molest
Tiphia cocoons, individuals were collected for insectary experi-
ments with this species. Several cocoon cells of copper screening
were placed in soil in cages containing these ants. The mesh of
this screen was coarse enough to permit entrance of ants but too
fine to allow soil to sift through readily. These cocoons remained
in good condition and after months only 1 out of 25 showed
signs of fungus. Twenty per cent of these cocoons were riddled
by this tiny ant, showing that the species might easily become a
serious factor working against Tiphia establishment and subse-
quent increase. Fortunately, however, these ants were not
generally abundant in areas containing cocoons.

Damage to Other Popillia Japonica Parasites in
THE Insectary

One instance of damage to insectary material other than Tiphia
cocoons was noted in 1932, when ants of the species Solenopsis
molesta ruined a portion of an experiment by killing flies of
Prosena sihirita (F.) that were being held in a small cloth cage.
The ants were no doubt attracted to the cage by honey which was
sprayed daily upon leaves as food for the flies. After having
been attracted to the scene, they played havoc with the flies as
they larviposited upon the soil.

Mar., 1940]

White: Japanese Beetle

SUMMARY

To obtain information on the part played by ants (Formicidae)
in the destruction of the Japanese beetle {Popillia japonica
Newm.) and its parasites, observations have been made of the
ants occurring in the Moorestown, N. J., area, and 23 species,
representing 11 genera, found to be most common have been
studied in the insectary.

Ants have been observed attacking living larvae as well as adult
beetles in the field.

Comparative data from field surveys in areas infested with ants
show very little difference in grub population.

Kesults of numerous insectary experiments testing any possible
relationship existing between ants and the Japanese beetle seem
to indicate little, if any, harm by the 23 species tried, either to
the egg or to any subsequent stage.

Formica fusca var. suhsericea Say was observed overpowering
adult Tipliia in the field.

Experiments dealing with various formicid species and TipJiia
cocoons in the soil show only one species, Solenopsis molest a
(Say), to damage cocoons consistently. Cocoons riddled by these
tiny workers have been found in the field as well as in the
insectary.

A single record is given of the finding within a colony of
Formica fusca var. suhsericea oi a beetle containing a Centeter
cinerea egg, and of another containing a fiy puparium.

100

Journal New York Entomological Society [Vol. XLVIII

NOTICE TO AUTHORS

Authors whose papers, in past years, were illustrated by line-
cuts or half-tones, may obtain such cuts, insofar as they are still
in existence, by defraying in advance the cost of packing and
shipping them. If claims for such plates are not made within
six months after the appearance of this notice, the Society re-
serves the right to destroy unclaimed plates. Correspondence
should be addressed to the Secretary of the New York Entomo-
logical Society.

BOOK NOTICE

The Principles of Insect Physiology. By V. B. Wigglesworth,
M.A., M.D., F.E.S. Reader in Entomology in the London
School of Hygiene and Tropical Medicine. Large 8vo.,
cloth, 434 pp., 316 illus.. New York. E. P. Dutton & Co.,
1939, $8.00.

This book is of general monographic character, in which par-
ticular emphasis has been given to the functions of organs and
tissues, and it has been so prepared as to amplify and supplement
other recently issued works on entomology, notably, those by
Immes, Weber, Snodgrass and others, in which books primary
emphasis has been given by their authors on diversity of form.
Its subdivisions include discussions of such matters as develop-
ment of the egg; growth; muscular system and locomotion;
nervous system ; sense organs ; vision ; mechanical and chemical
senses; behavior; respiration; circulatory system and associated
tissues ; digestion and nutrition ; excretion ; metabolism ; tempera-
ture relations, and the like. In its preparation special care has
been used to make it particularly suitable to the needs of students
of general entomology, of insect ecologists, and of economic ento-
mologists. Fully illustrated and well documented with a wide
range of bibliographical reference for those who desire to delve
further into various ramifications of the subject, this work,
though primarily on physiology, will be of helpfulness in
enabling its readers to obtain a grasp of the broad basic principles
underlying the ecology and practical control of insect pests. —
J. S. W.

Mar., 1940]

Book Eeviews

101

A German-English Science Dictionary for Students in the Agri-
cultural, Biological and Physical Sciences. By Louis De-
Vries, Professor of Modern Language, Iowa State College,
with collaboration of members of the Graduate Faculty.
12mo., semi-flexible cloth, 473 pp., N. Y., McGraw-Hill Book
Co., 1939, $3.00.

The little book contains 48,000 terms (German-English) per-
taining to the agricultural, biological and physical sciences:
bacteriology, medicine; botany and zoology (including anatomy,
embryology, pathology and physiology), entomology, genetics,
forestry, horticulture, agronomy, chemistry, physics, and mathe-
matics. Many common idioms and other terms likely to be
needed by students are included and past tenses and past parti-
ciples are translated for the benefit of elementary students. A
noteworthy feature is a list of references to other similar works,
over fifty in number, including general dictionaries, German-
English scientific dictionaries and glossaries. Originating at
Iowa State College through the development of complicated inter-
departmental research problems, this book has been in prepara-
tion for some 25 years and is, to a degree, the composite work of
34 members of the Graduate Faculty of that institution. It has
been prepared to meet a definite need for a comprehensive work
effectively adjusted to the steadily increasing overlapping of the
sciences. The small size, the semi-flexible binding, and the ease
with which it lies open, all make it more readily portable and
convenient to use. An exceedingly appropriate quotation for a
work of this type appears opposite the title page and was taken
from Dr. Samuel Johnson: “Every other author may aspire to
praise, the lexicographer can only hope to escape reproach.”- — ■
J. S. W.

The World of Insects. By Carl R. Duncan and Gayle Pickwell.
8vo., cloth, 409 p., 194 Figs. New York, McGraw-Hill Book
Co., 1939. $3.50.

Written by a professor of entomology and botany and a pro-
fessor of zoology respectively in the San Jose (Calif.) State
College, this book is another brief, non-technical resume of the
type of information usually presented in the numerous works of

102

Journal New York Entomological Society [Vol. xlviii

this kind which have appeared in recent years. Consideration
is given to such topics as insect structure and growth; food and
feeding habits; insect voices; social life among the insects; col-
lecting and rearing insects ; book lists for further study, and the
like. Some of the material here presented originally was issued
in publications of the San Jose Teacher’s College. It has, how-
ever, been extensively revised; many new sections and one new
chapter as well as many new illustrations have been added.
Practically all the photographs were taken by the junior author,
and many of these are of considerable excellence. — J. S. W.

Of Ants and Men. By Caryl P. Haskins. Research Associate,
Harvard University. 8vo., cloth, 244 p., 15 illus. New
York, Prentice-Hall Inc., 1939. $2.75.

In this book the author undertakes to make a brief review of
the origin and evolution of ants. Further, the author treats of
their concerted behavior in so far as it illustrates some of the
similarities and some of the differences which exist between their
social life and that of ourselves. The data assembled and pre-
sented form excellent exemplifications of the truism that, almost
inevitably, any group of animals and plants, which superficially
may appear a most compact and homogeneous unit, will upon
minute examination be found to contain the widest diversification
in membership. Some of these similarities and dissimilarities are
here discussed in detail, while others are given scarcely more than
mere mention. Against a well developed background, man is
brought into spotlight. The author holds no halo over his head
either from the standpoint of the present time or from the back-
ground of the past. He does, however, have confidence in human-
ity in dealing with the problems of existence or, as Thoreau would
put it : ‘ ‘ There is more day to dawn. ’ ’ Since this volume is one
of selections of the Scientific Book Club, it is probable that it will
find its way into the library both of the serious scientific worker
and the dilettante. Fortunately, any one can read this little book
with pleasure and profit, and the more the reader already knows
about ants and about men, the keener will be the enjoyment of
it.— J. S. W.

Mar., 19-10]

Notice to Members

103

NOTICE TO members

The Society has on hand a large stock of loose ‘‘signatures” (not
separates) of volumes 14 to 31 inclusive, of the Journal in addition
to the regular stock of ‘ ‘ numbers. ’ ’ These will be distributed free
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parts will be mailed early in May. Address requests to R. E.
Blackwelder, American Museum of Natural History, New York,
N. Y.

,-r^'

The

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JOURNAL

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Vol. XLVIII

No. 2

JUNE, 1940

Journal

of the >

New York Entomological Society

Devoted to Entomology in General ^

Edited by HARRY B. WEISS

Publication Committee

HARRY B. WEISS J. D. SHERMAN, Jr.

WILLIAM MOORE E. L. BELL

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Published Quarterly by the Society
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* LANCASTER, PA.

NEW YORK, N. Y.

1940

I

i

CONTENTS

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

By Charles P. Alexander 105

Some Synonymy in Neotropical Hesperiidae (Lepidoptera)

By E. L. Bell 116

Characteristics of Certain Western Cicadas

By Dr. John W. Sugden 117

New Fleas 126

The Sarcophaginae and Their Relatives in New York,

Part I

By Harold C. Hallock 127

Studies on the Ichneumonidae of New England (Hymen-
optera)

By Harry D. Pratt 155

Money Losses Due to Destructive Insects

By Harry B. Weiss 195

NOTICE: Volume XLVIII, Number 1, op tpie Journal op
THE New York Entomological Society was published on
March 29, 1940.

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. XLVIII June, 1940 No. 2

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

By Charles P. Alexander
Amherst, Massachusetts

The preceding: instalment under this title was published in
September, 1938 (Journal op the New York Entomological
Society, 46 : 327-337 ) . The crane-flies described at this time are
from Potrerillos, “The Valley of the Clouds,” in Chiriqui,
northern Panama, where they were collected by Mr. D. V. Brown,
February to May 1934, and Mr. J. W. MacSwain, May to June
1935. My deep thanks are extended to the collectors of this rich
material. All types are preserved in my private collection of the
Tipulidae of the World. Only a portion of the novelties are
described at this time, the entire collection being rich and varied
in species.

Genus Tipula Linnasus

Subgenus Neotipula new.

Frontal prolongation of head slender; nasus elongate. Antennae 13-seg-
mented; flagellar segments three to nine, inclusive, with a conspicuous lobe
on distal half, this lobe exceeding one-half the length of the segment and
producing a weak pectinated appearance; setae of flagellum reduced, re-
stricted to a series on outer face of segment on opposite side from the pecti-
nation; no basal enlargement on any of the segments; verticils lacking;
terminal segment elongate, subequal in length to the penultimate. Tibial
spur formula 1-1-2 ; claws large, each with an obtuse lobe before midlength.
Sternopleural setae sparse, small and pale. Wings with squama naked; veins
unusually glabrous, beyond cord including a very restricted series of tiny
trichia on vein -R4 + 5. Venation: Bs shorter than m-cu; B^ before fork of
^3 + 4; veins B^ and -R4 + 5 nearly straight, not sinuous at near midlength, as in

juN *1 m

106

Journal New York Entomological Society [Vol. xlviii

Holorusia, cell consequently not constricted at this point; vein M con-
spicuously arched before fork; m-cu long, cell unusually wide, at margin
only a little shorter than cells Jfi to M3, combined.

Type of subgenus. — Tipula (Neotipula) pectinella new species.
(Neotropical Region).

Besides the subgenotype, it would appear that Tipula maya
Alexander likewise belongs here. The strict affinities are very
uncertain. The nature of the antennte, as the lack of a basal
enlargement and accompanying verticils on the flagellar seg-
ments, is more suggestive of Holorusia than of Tipula but the
venation of the radial fleld precludes its reference to the former
genus. I prefer to consider it as being more closely related to
Tipula while emphasizing these points of difference. It seems
entirely possible that the group will eventually be elevated to full
generic rank.

Tipula (Neotipula) pectinella new species.

Size large (wing over 35 mm.) ; general coloration of notum reddish
brown, the scutum behind traversed by a blackened band; pleura yellow,
with a narrow dorso-longitudinal brown stripe; halteres brownish black, the
knobs obscure yellow; legs brownish yellow, the femora with tips broadly
blackened, preceded by a slightly narrower pale yellow ring; wings with a
rich brown suffusion, variegated by sparse darker brown and more abundant
cream-colored areas; E^, shortly before fork of ^^3+4; abdominal tergites dark
reddish brown, with narrow, more blackish stripes.

Male. — Length about 30 mm.; wing 36 mm.; antenna about 4 mm. ■

Frontal prolongation of head and the palpi black. Antennae dark brown,
the first flagellar segment paler; structure as above described. Head dark
brown, the posterior orbits dull yellow; anterior vertex relatively narrow,
about one-half wider than the diameter of the scape.

Pronotum yellow, narrowly darkened on median border. Mesonotal prae-
scutum with the ground-color obscure yellow, almost covered by four reddish
brown stripes, the sublateral portions of the sclerite deepening to black, the
extreme border yellow; scutum with median area obscure yellow, the lobes
reddish brown, the entire distal fourth of scutum traversed by a blackened
band; scutellum dark gray, parascutella brown, the cephalic and caudal por-
tions yellow; mediotergite plumbeous gray, the posterior and lateral por-
tions broadly brownish black. Pleura yellow, with a narrow, dark brown,
longitudinal stripe extending from the cervical margin across the dorsal
pleurites, passing through the anterior spiracle, ending at wing-root. Hal-
teres long, brownish black, the extreme base of stem and apex of knob
obscure yellow. Legs with the coxae and trochanters yellow; femora brown-
ish yellow, darker outwardly, the tips broadly blackened, preceded by a

June, 1940]

Alexander: Crane-Flies

107

slightly narrower pale yellow ring ; tibise light brown, the bases more
whitened, the tips narrowly infuscated; tarsi light brownish yellow. Wings
with a strong rich brown suffusion, sparsely variegated by darker brown and
more numerous cream-colored areas; the chief dark spot lies at near two-
thirds the length of cell M ; the cream-colored areas occupy cell M before
and beyond the dark spot, across the cubital and anal cells, and in the outer
medial field; Cell Sc uniformly dark brown; veins reddish brown, the ob-
literative areas at cord small but conspicuous. Venation: Petiole of cell
a little longer than m ; cell 2nd A wide.

Abdominal tergites dark reddish brown, with three narrow blackish stripes
that are more or less interrupted at the posterior borders of the segments,
especially the median stripe; basal tergite brightened medially; extreme
caudal margins of segments pale; sternites more uniformly reddish brown;
hypopygium a little brightened. Male hypopygium with the eighth sternite
conspicuously produced beyond the level of remainder of hypopygium.

Holotype, J', Potrerillos, altitude 3,000 feet. May 20, 1935
(MacSwain).

The most similar species seems to be Tipula maya Alexander, of
Guatemala, still known only from the unique female type in
which the antennae were broken. The essentials of venation of
the two species are very similar and it seems certain that maya
will likewise be found to belong to this new group.

Tipula browniana new species.

Belongs to the glaphyroptera group ; general coloration yellow, the prae-
scutum with four pale brown stripes, the intermediate pair irregular, broken
before midlength, behind this point closely approximated or fused; antennae
short, bicolored; halteres yellow; femora brownish yellow, with a narrow
pale brown subterminal ring, preceded by a clearer yellow annulus; wings
light yellow, with a tesselated brown pattern; entire; Bs long; abdomen
yellow, the basal tergites narrowly lined sublaterally with dark; male hy-
popygium with a small median lobe in membrane between eighth and ninth
sternites, this bearing long conspicuous setae.

Male. — Length about 13 mm.; wing 14 mm.; antenna about 3 mm.

Frontal prolongation of head yellow; nasus distinct; palpi with basal
three segments yellow, the terminal segment dark brown. Antennae rela-
tively short, if bent backward extending about to wing-root; basal three seg-
ments light yellow, the remainder of flagellum bicolored, yellow, with the
small basal enlargements abruptly black; verticils a little longer than the
segments. Head obscure yellow; posterior orbits and a slight cloud on
anterior vertex pale brown; anterior vertex relatively wide, approximately
five times the diameter of scape; vertical tubercle low, its cephalic end very
feebly notched.

Pronotum yellow, faintly darkened medially above. Mesonotal praescutum

108

Journal New York Entomological Society [Vol. XLVIII

yellow with four pale brown stripes; intermediate stripes irregular in
outline, on cephalic third of sclerite widely separated, slightly broken at
this point and thence forming a single median stripe to the suture; lateral
stripes narrow, the margins laterad of these stripes less evidently darkened;
scutum yellow, the lobes and a narrow but conspicuous median vitta pale
brown; scutellum broadly pale yellow, the parascutella darker; mediotergite
yellow, with a broad, conspicuous, pale brown, median line. Pleura yellow,
the dorsopleural region a little darker ; pleurotergite weakly inf umated. Hal-
teres pale yellow. Legs with the coxae and trochanters yellow; femora
brownish yellow, the bases clearer yellow, with a narrow pale brown sub-
terminal ring, preceded by a much wider, clearer yellow annulus ; tibiae
yellow, the tips weakly infumated; tarsi yellow, the outer segments brown.
Wings light yellow, the costal and prearcular fields more saturated yellow;
a conspicuous tesselated pale brown pattern, as follows: Four subcostal
areas, the first postarcular, including bases of cells and M ; second area at
about one-third the length of cell M, reaching vein M behind; third area at
origin of Bs, not reaching M ; fourth area stigmal, confluent with a major
darkening on anterior cord; remaining cells of wing with zigzag brown
clouds, subequal in extent to the yellow ground; outer half of cell B^ and
most of Ms uniformly pale; veins yellow, infuscated in the clouded areas.
Venation: + o pale but entire; Bs long, about two and one-half times m-cu;

petiole of cell Mi subequal to m.

Abdomen yellow, the basal tergite narrowly lined laterally with darker;
hypopygium yellow. Male hypopygium with the ninth tergite transverse,
the caudal margin crenulate, with a very shallow U-shaped median notch;
dorsal surface of tergite with a median furrow. Outer dististyle relatively
long, narrowed on basal third, sinuous. Inner dististyle relatively broad, the
posterior angle at base rounded. Basistyle produced into a setiferous lobe,
its dorsal angle prolonged into a conical spine immediately behind the inser-
tion of the inner dististyle. Membrane between eighth and ninth sternite
with a small subglobular or short-oval median lobe that bears more than a
score of very long strong setse.

Holotype, J', Potrerillos, altitude 3,500 feet, May 12, 1934
(Brown).

Tipula hrowniana is named in honor of the collector, Mr. D.
V. Brown. The species most resembles forms such as T. absona
Alexander, of Ecuador, and allies, differing especially in the
pattern of the mesonotum and wings, and in the structure of the
male hypopygium. There are several species of the group having
a checkered wing-pattern, these differing among themselves in
the presence or absence of vein the coloration of the femora,
the wing-pattern and venation, and in the structure of the male
hypopygium.

June, 1940]

Alexander: Crane-Flies

109

Tipula scriptella new species.

General coloration of thorax obscure yellow, the prsescutum with dark
stripes; pleura with a complete dark brown girdle on mesepisternum ; hal-
teres obscure yellow; femora yellow, with a relatively narrow and ill-defined
pale brown subterminal ring; wings brown, streaked longitudinally with
whitish and variegated with small darker brown areas; abdominal tergites
yellow, with three brown stripes, the sternites with a similar median vitta.

Female. — Length about 17 mm.; wing 15 mm.

Head behind obscure yellow, the anterior part destroyed.

Mesonotum obscure yellow pollinose, with dark brown stripes on prae-
scutum; central portion of mediotergite darkened, the sides broadly pale.
Pleura obscure yellow, variegated with dark brown, including a complete
transverse girdle from lateral margin of praescutum across the dorsopleural
membrane, involving most of the mesepisternum; pleurotergite slightly
darkened. Halteres obscure yellow. Legs with the coxae and trochanters
yellow; femora yellow, with a relatively narrow and ill-defined pale brown
subterminal ring; tibiae yellow, the tips narrowly darkened; basal two tarsal
segments yellow, the tips very narrowly darkened, the outer tarsal segments
uniformly brown. Wings with the ground color brown, conspicuously varie-
gated with darker brown and whitish subhyaline, the latter chiefly dis-
tributed as a central streak involving the middle third of wing, including
cell 1st M2 and bases of all outer medial cells, reaching the margin in cell
cubital and anal cells likewise extensively whitened; cell C uniform
medium brown; cell Sc with four small dark areas, the second at origin of
Bs, all but the last involving the radial field behind; cells B^ and B^ chiefly
and almost uniformly infumated; darker brown areas at anterior cord, outer
end of cell 1st Mz and as broad seams on outer half of vein Cu, distal section
of Cu-i_ and m-cu; outer medial field infumated, with pale marginal droplets in
outer ends of cells 2nd Mz and M^ ; veins yellow, darker in the clouded areas.
Venation: + a entire; M^ + i only a little shorter than the basal section of M3.

Abdominal tergites obscure yellow, with three brown stripes, the median
one broad on basal and intermediate segments, narrowed on outer tergites,
ending on tergite seven, slightly interrupted at the posterior margins of the
segments; lateral stripes narrow, becoming obsolete behind; sternites yellow,
with a median brown vitta. Ovipositor with cerci nearly straight, very
slender.

Holotype, 5? Potrerillos, altitude 3,500 feet, May 12, 1934
(Brown).

In its general appearance, especially the wing pattern, the
present fly is most similar to the Chilean Tipula pliilippiana
Alexander, differing in the details of coloration and venation.
It is probable that this species will be found to belong to the
glaphyroptera group of species.

110

Journal New York Entomological Society [Vol. xlviii

Tipula ctenopyga new species.

General coloration of mesonotum reddish brown, the usual stripes reduced
to three conspicuous black spots; antennae elongate; femora obscure yellow,
the tips very narrowly but conspicuously black; wings with a brown tinge,
the stigma dark brown, conspicuous; ^1 + 2 entire; abdomen with a subtermi-
nal black ring; male hypopygium with the conspicuous tergite parallel-sided,
the apex convexly rounded and set with abundant black setae; a conspicuous
black lobe on mesal face of basistyle; eighth sternite produced ventrad into
scoop-like flattened lobes that bear marginal spinous setae; aedeagus trifid at
apex.

Male. — Length about 12 mm.; wing 11.5 mm.; antenna about 4.8 mm.

Frontal prolongation of head short and stout, without nasus, obscure
brownish yellow; palpi dark brown. Antennae relatively long; basal three
segments obscure yellow, succeeding segments black, the apices very restrict-
edly brightened; at and beyond midlength of the organ the segments are
uniformly black ; flagellar segments incised ; terminal segment very small.
Head chestnut brown; anterior vertex wide.

Mesonotum reddish brown, the praescutum with three black spots, one
representing the anterior fourth of a narrow median stripe, the others
occupying the anterior ends of the usual outer praescutal stripes, extending
laterad to the margins; scutellum and mediotergite darkened. Pleura ob-
scure reddish yellow. Halteres dusky. Legs with the coxae and trochanters
yellow; femora obscure yellow, the tips very narrowly but conspicuously
black, the amount subequal on all legs; tibiae obscure brownish yellow, the
tips very narrowly blackened; basitarsi brownish yellow, the outer tarsal
segments brownish black. Wings with a brown tinge, prearcular region and
cells C and Sc more yellowish; stigma oval, dark brown, conspicuous; veins
brownish black, paler in the more brightened costal portions. Wings with
conspicuous basal petiole; macrotrichia on veins -E2+3, ^3, -K4 + 5 and very
restricted or lacking elsewhere beyond cord. Venation: i^i + 2 entire; cell
1st M2 long and narrow; petiole of cell Ifi a little exceeding m; cell 2nd A
very narrow, reduced to a linear strip.

Basal abdominal tergites reddish, with a broken black median stripe, on
tergites two and three including the narrow posterior borders; tergites six
and seven uniformly black; sternites reddish yellow, segments seven and
eight black; hypopygium obscure reddish yellow. Male hypopygium with
the ninth tergite unusually long, obliquely elevated, projecting caudad to
opposite or beyond the outer level of the dististyles; parallel-sided, the
caudal margin gently convex, blackened by an abundant concentration of
setae. Lower and inner face of basistyle with a conspicuous, elongate, black
lobe, its margin microscopically • roughened. Outer dististyle small, slender,
black. Inner dististyle a compact compressed blade. Eighth sternite exten-
sive, the apical portion directed ventrad into two flattened scoop-like lobes,
separated from one another by membrane, their margins with several black
spinous setae that pass into longer normal setae on mesal portions. Aedeagus
small and slender, before apex on either side with a slender spinelike arm,
presenting the appearance of a delicate trident.

June, 1940]

Alexander: Crane-Flies

111

Holotype, J', Potrerillos, altitude 3,00d feet, May 20, 1935
(MacSwain).

Tipula ctenopyga is most nearly related to T. detecta Alex-
ander, of Venezuela, differing in the wing-pattern and venation,
and in the coloration of the body and legs. The male sex of
detecta is still unknown to me.

Limonia (Limonia) paucilobata new species.

Belongs to the apicata group; general coloration of mesonotum yellow,
the praescutum with a black median stripe and shorter, confluent lateral
stripes; pleura chiefly yellow; fore femora black; wings with a strong dusky
tinge, the stigma and outer radial field darker; m-cu at fork of M; abdomi-
nal tergites brownish black, the sternites obscure yellow; male hypopygium
with basistyle simple, provided with a single lobe.

Male. — Length about 6 mm.; wings 6.5 mm.

Eostrum and palpi black. Antennae black throughout; basal flagellar
segments oval, the outer segments subcylindrical ; segments with short
glabrous apical necks; terminal segment longer than penultimate, its outer
end pointed. Head gray, clearer gray along posterior orbits and on the
linear anterior vertex; central portion of posterior vertex blackened.

Pronotum obscure yellow, narrowly blackened medially. Mesonotal
praescutum polished yellow laterally, with a conspicuous entire black median
stripe and short confluent lateral stripes; scutal lobes blackened, the median
area obscure yellow; scutellum obscure yellowish brown, paler on posterior
margin; mediotergite infuscated. Pleura and pleurotergite obscure yellow,
the anepisternum and ventral pleurotergite slightly darkened. Halteres
dark brown, the base of stem restrictedly yellow. Legs with the coxae and
trochanters yellow; fore femora black, the bases narrowly yellow; middle
and posterior femora dark brown, pale basally, the tips pa,ssing into black;
tibiae and tarsi brown, the terminal segments of the latter darker. Wings
with a strong dusky tinge, the outer radial field more evidently infumated;
stigma oval, darker brown; veins dark brown. Venation: Sc long, end-
ing shortly before fork of Es, Sc^ close to its tip ; Es exceeding twice the
basal section of -^4 + 5; m-cu at fork of M.

Abdominal tergites brownish black; sternites obscure yellow; hypopygium
darkened. Male hypopygium with the basistyle simple, the mesal face on
proximal half with a single stout lobe. Dististyle simple, gradually nar-
rowed to a slender curved beaklike point. Gonapophyses unusually broad
and flattened, the terminal beak short and curved. Aedeagus narrow.

Holotype, J', Potrerillos, altitude 3,000 feet, May 5, 1935
(MacSwain).

In the structure of the male hypopygium, Limonia {Limonia)
paucilohata is quite distinct from the other members of the

112

Journal New York Entomological Society [Vol. xlyiii

apicata group. The relatively simple nature of the basistyle is
likewise found in L. (L.) acuminata Alexander, L. (L.) hoffmani
Alexander, L. (L.) rapax Alexander, and others, but the details
of structure of all parts of the hypopygium are distinct.

Orimarga (Orimarga) melampodia new species.

General coloration gray, the lateral margin of praescutum, together with
the pleura, striped longitudinally with blue-gray; legs brownish black;
wings whitish subhyaline; Sc short, B1 + 2 relatively long, approximately
three times Bo+s; abdomen brownish black.

Male. — Length about 9 mm. ; wing 6.5 mm.

Eostrum and palpi black. Antenna3 black throughout; flagellar segments
oval, with short verticils; terminal segment larger than the penultimate.
Head gray.

Mesonotum almost uniform dark plumbeous gray, the lateral margins of
praescutum more blue-gray pruinose; a blackened spot on lateral margin of
prsescutum before suture. Pleura above light gray, with a broad blue-gray
stripe extending from behind the fore coxae, passing above remaining coxae
to base of abdomen, narrowly bordered above by blackish; ventral sterno-
pleurite blackened. Halteres with stem pale yellow, knob dark brown.
Legs with coxae and trochanters black; remainder of legs brownish black,
the femoral bases not or scarcely brightened. Wings whitish subhyaline;
veins brown. Venation: Sc short, /Sci ending about opposite one-fifth the
length of Bs and before level of m-cu; B^+z fully three times B2+3 and about
one-third the length of Bs.

Abdomen brownish black, the sternites a little paler. Male hypopygium
with the outer dististyle acicular, blackened, scarcely sinuous on distal half.
Blades of gonapophyses narrow, the tips pale, long-attenuate.

Holotype, J', Potrerillos, altitude 3,500 feet, May 14, 1934
(Brown).

Orimarga {Orimarga) melampodia is closest to 0. (0.) dampfi
Alexander, of Mexico, differing in the blackened legs, details of
venation, as the short B2 + 3 and long B1 + 2, and in slight details
of the male hypopygium.

Helius (Helius) distinervis new species.

General coloration brownish black; legs dark brown, the outer tarsal seg-
ments paling to obscure yellow; wings with a weak brown tinge, stigma
oval, medium brown; m-cu lying far distad, nearly twice its own length
beyond the fork of M ; male hypopygium with numerous small spinulse sur-
rounding the two major apical spines of the outer dististyle.

Male. — Length about 4.5 mm. ; wing 4.6 mm.

Eostrum not apparent in the unique type. AntenngB small, dark brown,
the outer segments broken. Head dark brown.

June, 1940]

Alexander: Crane-Flies

113

Thorax uniformly brownish black, without stripes. Halteres dusky
throughout. Legs dark brown, the outer tarsal segments paling to obscure
yellow. Wings with a weak brown tinge, the prearcular region and cells
C and Sc slightly darker brown; stigma oval, medium brown; veins dark
brown. Costal fringe of moderate length. Venation: Sc long, Scj^ ending
about opposite five -sixths the length of the long Bs, Sc2 at its tip; branches
of Bs rather strongly diverging at outer ends, cell B^ at margin wider than
cell Bo; inner end of cell 1st M2 strongly narrowed; m-cu lying far distad,
about opposite r-m and nearly twice its own length beyond the fork of M.

Abdomen black; hypopygium and sternites a little paler. Male hypo-
pygium with the outer dististyle unequally bifid at apex, with numerous
additional spinulse grouped around the apex. Inner dististyle subequal in
length, gradually narrowed outwardly. Lateral tergal arms directed mesad
as parallel-sided flattened blades, the tips obtusely rounded.

Holotype, J', Potrerillos, altitude 3,000 feet, May 20, 1935
(MacSwain).

Helius {Helms) distinervis is very different from other de-
scribed species, especially in the unusual position of the m-cu
crossvein. The rostrum is invisible in the unique type but the
generic reference seems undoubtedly to be correct unless a new
group might be deemed necessary to receive the species.

Ischnothrix (Ischnothrix) patens ucav species.

General coloration of thorax medium brown; antennse (female) elongate,
exceeding one-half the body in length; legs pale brown; wings with a strong
brownish yellow tinge, the long-oval stigma pale brown ; B^ short, nearly
erect; cell M2 open by atrophy of basal section of M^; m-cu shortly beyond
fork of M; abdominal segments dimidiate, brown, the caudal half light
yellow.

Female. — Length about 5.5 mm.; wing 5 mm.; antenna about 3 mm.

Eostrum pale brown; palpi dark brown. Antennae (female) exceeding
one-half the length of body; scape and pedicel pale, flagellum black; flagel-
lar segments long-cylindrical, with relatively short verticils scattered through-
out the length. Head pale brown.

Mesonotum medium brown, unmarked, the pleura a little brighter.
Halteres yellow, the knobs infuscated. Legs with the coxae testaceous
brown; trochanters obscure yellow; remainder of legs pale brown, the ter-
minal segments dark brown. Wings with a strong brownish yellow tinge,
the long-oval stigma pale brown; cells C and Sc brighter yellow than re-
mainder of ground; veins pale brown. Sc more yellowish. All veins beyond
cord with macrotrichia. Venation: Sc relatively long, Sc^^ ending beyond
midlength of the arcuated Bs, SC2 about mid-distance between origin of Bs
and tip of Sc-^_', E2+3+4 elongate, gently arcuated; -K3 short, nearly erect;
distance on margin between veins B^ + 2 and B^ a little greater than the

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Journal New York Entomological Society [Vol. xlviii

length of latter; a little shorter than -K2+3+4; cell open by atrophy of
basal section of M3; cell 2nd M2 about three times its petiole; m-cu shortly
beyond fork of M.

Abdominal segments dimidiate, brown, the caudal half of each segment
light yellow, the extreme base of the segments more vaguely brightened, cerci
slender.

Holotype, J, Potrerillos, altitude 3,000 feet, May 5, 1935
(MacSwain).

Ischnothrix (Ischnothrix) patens is readily distinguislied from
all previously described Neotropical species by the open cell M2
of the wings. Except for this character, it much resembles I.
(I.) argentinicola (Alexander) of Argentina. It is entirely
distinct from the two species previously known from northern
South America, I. (Z.) hrevisector Alexander, of Venezuela, and
Z. (Z.) Iloydi (Alexander), of Colombia. The genus Ischnothrix
Bigot had not been found in America north of Colombia and is
not included in Curran’s “North American Diptera,” 1934.

Lipsothrix neotropica new species.

General coloration pale yellow; legs whitish; wdngs whitish subhyaline;
m-cu at near two-thirds the length of cell 1st M2; male hypopygium with
the interbase pale, long and sinuous, gradually narrowed to hairlike points.

Male. — Length about 6.5 mm.; wing 7 mm.

Eostrum pale yellow; palpi a little darker. Antennse with scape and
pedicel pale yellow, flagellum brown; flagellar segments oval, the verticils
exceeding the segments in length. Head pale yellow.

Thorax uniformly pale yellow. Halteres pale yellow, the knobs weakly
darkened. Legs whitish, even the last tarsal segment scarcely darkened.
Wings whitish subhyaline, unmarked; veins brown. Macrotrichia of veins
long and conspicuous. Venation: ending just beyond fork of Ms, Sco a

short distance from its tip; Ms in longitudinal alignment with -E2 + 3 + 4; E2 + 3
subequal to M^+2’, veins M^ and M^ diverging at margin so cell M^ is only a
little more than one-half as extensive at margin as cell E3; m-cu at near
two -thirds the length of cell 1st Mo.

Abdominal tergites pale brownish yellow, sternites whitish; hypopygium
pale except for the blackened outer dististyle. Male hypopygium with the
interbases pale, long and sinuous, gradually narrowed to hairlike points.

Holotype, J', Potrerillos, altitude 3,000 feet. May 7, 1935
(MacSwain).

Lipsothrix neotropica is the first Neotropical member of the
genus to be discovered. It is very distinct from the only known
Nearctic species, L. sylvia (Alexander), differing markedly in

June, 1940]

Alexander: Crane-Flies

115

the venation, as the distal position of m-cu, and in the structure
of the male hypopygium, especially of the interbases. The in-
dividual, while pale, is certainly not in teneral condition.

Gonomyia (Gonomyia) remota obtusistyla new subspecies.

Male. — Length about 4.5 mm. ,• wing 4.5 mm.

Close to typical remota Alexander, of southern Mexico and Salvador, dif-
fering in details of structure of the male hypopygium. Inner dististyle
with the outer arm stouter, less spine-like in appearance; inner arm obtuse
at apex, not extended into an acute spine as in the typical form. Aedeagus
with apical blade more expanded.

Holotype, c?, Potrerillos, altitude 3,500 feet. May 12, 1934
(Brown). Paratopotypes, 2 J' J', altitude 3,000 feet. May 14-15,
1935 (MacSwain).

Gonomyia (Lipophleps) macswaini new species.

Belongs to the manca group; general coloration dark brown, the pleura
with a longitudinal whitish stripe; antennse black throughout; legs brownish
black to black; wings with a strong brownish tinge, the stigma a trifle
darker but very poorly delimited; >Sci ending opposite origin of Bs; male
hypopygium with outer apical angle of basistyle produced into a long
fleshy lobe; dististyles subterminal, asymmetrical on the two sides; phal-
losome with a single blackened rod.

Male. — Length about 3.5 mm.; wing 4 mm.

Eostrum and palpi black. Antennae black throughout. Head dark,
variegated on posterior portion by yellow.

Pronotum and lateral pretergites light yellow. Mesonotal praescutum
and scutal lobes dark brown; median area of scutum testaceous; scutellum
yellow, dark at base; mediotergite darkened posteriorly, slightly paler on
cephalic portion. Pleura dark brown, with a broad whitish longitudinal
stripe extending from behind the fore coxae to the abdominal sternites;
dorsopleural membrane pale. Halteres dark, the knobs broken. Legs with
the coxae brownish testaceous; remainder of legs brownish black to black.
Wings with a strong brown tinge, the stigma a trifle darker but very
poorly delimited; prearcular region and costal field slightly paler than the
ground; veins brown. Venation: /Sci ending opposite origin of Bs, Scs re-
moved from its tip; Bs relatively long, exceeding one-half its anterior
branch; cell 1st M2 closed; m-cu at fork of M.

Abdominal tergites dark brown, sternites obscure yellow. Male hypo-
pygium with outer apical lobe of basistyle long and slender, subequal in
length to the body of style. Dististyles of the two sides asymmetrical, one
side terminating in two slender rods, the inner of which is heavily blackened,
the outer straight and paler; at base of these rods an additional tiny black
spine; style of opposite side with a long base, at apex expanded into an

116

Journal New York Entomological Society [Vol. xlviii

asymmetrical head that is conspicuously bispinous. Phallosome complex,
asymmetrical, with a single blackened rod.

Holotype, J', Potrerillos, altitude 3,000 feet, May 5, 1935
(MacSwain).

Gonomyia (Lipophleps) macswaini is named in honor of the
collector of many of the interesting species discussed in this
report. The species is readily told from all other members of
the manca group by the structure of the male hypopygium,
especially the prolonged lobes of the basistyles, in conjunction
with the strongly asymmetrical dististyles.

SOME SYNONYMY IN NEOTROPICAL HESPERIID^
(LEPIDOPTERA)

By E. L. Bell

Kirby in his ‘‘Catalogue of the Collection of Diurnal Lepidop-
tera Formed by the Late William Chapman Hewitson,” p. 243,
1879, lists Nisoniades selma Hewitson’’ from “Cayenne, Ama-
zon, Venezuela, Rio.”

The writer could find no published description of this name by
Hewitson and wrote to Brigadier W. H. Evans of the British Mu-
seum for information about it and has received a reply stating
that it is a manuscript name and that the specimens in the Hewit-
son collection labelled “se^ma” are Chiomara mithrax Moeschler.

Mabille, “Genera Insectorum,” xvii, p. 74, 1903, lists Chio-
mara salma Hewitson,” an evident misspelling of ‘‘selma.’’

Nisoniades selma Kirby (Hewitson ms.) and Chiomara salma
Mabille are thus disposed of as synonyms of Chiomara mithrax
Moeschler.

June, 1940]

SuGDEN: Cicadas

117

CHARACTERISTICS OF CERTAIN WESTERN
CICADAS

By Dr. John W. Sugden
University of Utah, Salt Lake City, Utah

Many differences exist in cicadas other than the morphological
characters nsed in the taxonomic differentiation of genera and
species. The habits, broods, locations, songs, seasons, plant as-
sociations, time and activities may be so constant that they may
be typical of certain groups and may even be of value in the
determination of closely allied species. With a knowledge of
these factors, one may be reasonably certain of what species or
group of species may be expected to be found under certain
conditions.

The season for most of the far western cicadas is chiefly the
early summer and by the latter part of the season, most of their
visible activities are over with. Okanagana and the allied genera
are dominant and Tibicen is represented by only a few species
which are not typical of the genus in the east. By the latter part
of July, the broods have scattered and only rarely can a song be
heard. A record for Okanagana hella Davis is therefore unusual
both for time and altitude. It was observed on September 19,
1932 in Utah County, Utah, east of Mt. Timpanoogas (altitude
11,957 ft. ) in the valley known as Little Provo Hole at an altitude
of approximately 9,000 feet.

All the determinations of the species considered here have been
made by Mr. Win. T. Davis of Staten Island, New York and he
has reviewed the manuscript. Dr. Walter Cottam of the Uni-
versity of Utah determined the plants and John W. Sugden, III
assisted in the collection. The author’s thanks is expressed.

Cacama crepitans Van Duzee

This cicada was found (June 24, 1938) on the hills of the Coast
Range, Orange County, California, east of San Juan Capistrano.
They were on the steep hill densely studded with Opuntia cactus,
but no insects were found on the cacti notwithstanding the

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Journal New York Entomological Society [Vol. xlviii

popular name ‘‘cactus dodger,” but were always located on the
other plants of the region and often were on dead twigs.

The song of the Cacama, markedly different from that of
Okanagana, was given in sequences of ten to twelve seconds.
During the first five or six seconds, the song was rapid and shrill
and then it decreased through the remaining interval to be again
repeated without break in the continuity. The change from the
low, dull song to the high, shrill interval was rapid and that from
the opposites was considerably longer, taking about one second of
time. During the first part, the full song, the abdomen was
elevated and the opercula widely opened. During the remaining
period, the abdomen was depressed and the opercula were then
closed. The depression of the abdomen and the closure of the
opercula damped the sound and with the alternate opening and
closing, the song had an undulating character. The same insect
observed at Point Loma, California, several years ago (June 12,
1931) had the same undulating character but the variations were
slight. Only in cicadas with opercula covering the tympanal
cavities is such a song possible.

When captured there was a high-pitched, shrill, continuous
song. Flying was during the low part of the song, not during
the height with the abdomen raised.

Fig. 1. Song of Cacama crepitans Van Duzee.

Clidophelps pallida Van Duzee

Clidophelps pallida was found in the valleys of Southern Cali-_
fornia (near Elsinore, Calif. June 24, 1938) on Eriogonum.
There was a definite plant association and the insects were found
wherever the plant occurred. When the insects flew they in-
variably lighted on the plant and not on other bushes or plants
in the vicinity.

The characteristic song was a loud rasping rattle, very similar

June, 1940]

Sugden: Cicadas

119

in quality to the rattle of the rattlesnake. Many other cicada
songs have been described as resembling this sound. Putnam
(1881) writes that the note of 0. rimosa is a “rattling noise, ex-
ceedingly like that of a rattlesnake. This resemblance was so
close that one day in 1873, in the Shoshone Mountains, I was at-
tracted by a noise which I took to be one of those insects, and
stopped to pick it up, when I suddenly discovered a large rattle-
snake in its stead” (quotation, Myers, 1929 p. 221 and Davis
1921 p. 50). Englehart states that the song of 0. schoefferi
“closely resembles the whirring noise produced by a rattlesnake
and that on one occasion in June, 1917, in Washington County,
Utah, he came very near being bitten by a snake while searching
in a small bush for the supposed cicada” (quotation, Davis,
1921, p. 50). My notes concerning the same Clidophelps, ob-
served in San Diego County, California (June 12, 1931) were
much the same. The similarity to the rattlesnake rattle was
definite. With the song of any of the Okanaganas, I have never
had the impression of hearing a rattlesnake, however, the dif-
ference in the sounds may be more marked because of familiarity
with the reptile.

During the time of the rasping rattle, the wings were flattened
and slightly opened. This occupied about ten seconds of time
and none that were timed continued longer than twelve seconds.
Many of the insects in a location would give the rattle sound in
unison. When it was started, several could be heard to start
within a fraction of a second and all would stop at almost the
same instant.

In addition, there was a clicking sound, during which the ab-
domen was jerked up and down. This sound was repeated at
a rate as frequent as 140 a minute and could be counted with a
watch, much the same as taking a fetal heart tone. When dis-
turbed by being too close, the clicking was less frequent or dis-
continued. The insects depended upon concealing coloration and
sometimes they could be picked from the stem without attempting
to fly.

Okanagodes gracilis Davis

From a few miles north of Boulder City, Nevada (June 25,
1938) until off the plateau, the insects were found or heard

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Journal New York Entomological Society [Vol. xlviii

wlierever a plant association occurred. Three plants predomi-
nated and were always together. The cicadas were found only
on the first two.

1. The Bottle-plant, Eriogonum inflatum, a pale green,
spindly plant with swollen hollow stems.

2. Burro-weed, Franseria dumosa, a small, grey, sage-like plant,
growing from six to eight inches in height.

3. Creosote-bush, Covillea trident at a, an evergreen shrub with
deep green leaves, growing about two feet in height and giving
the only color to the region. It is the characteristic plant of
the area, the Covillea belt.

At the time, the temperature was estimated to be 110° F.,
which was apparently too conservative as at Boulder City, the
official reading in the shade was 114° F., and at Las Vegas, it
was considered “very warm.” Of course in the desert there was
no shade. Time 11 A. M.

The song, a series of short “zips,” was repeated, when not
disturbed, at intervals of 160 a minute. When approached, the
song was usually stopped or reduced in frequency and when it
was resumed, a few chirps were given rather cautiously before
starting the full song. Invariably the song seemed to be coming
from a plant nearer than the one occupied by the insect. Even
when allowance for error was considered, the site often was
beyond the estimated location. When disturbed and flying, the
male would give a short, rapid, continuous song or as often would
fly without making any noise. The song seemed to resemble a
mechanism that was wound up and released at frequent intervals
for a split second, e.g., a movie camera with the lever repeatedly
pressed. Protective obliterative coloration was especially
marked ; a pale insect on pale plants on a hot desert.

The songs of many of the Okanagana are very similar and
have few distinguishing characteristics. As a rule, the song is
given only during the hottest part of the day, never at evening
and rarely unless the insects are in the bright sunlight, even
being stopped when the shadow of a cloud comes over. Captive
specimens of Okanagana bella and Okanagana striatipes have
been induced to sing when a vacuum cleaner was started.

Oka7iagana ruhrovenosa Davis

At Gold Kun, California, (elevation 3224 Ft., June 13, 1938)

June, 1940]

SuGDEN: Cicadas

121

Okanagana ruhrovenosa were on the manzanita bushes, which in
this locality grew from three to six- feet in height. The cicadas
were usually on the smaller branches and were often out toward
the ends carrying the leaves.

The song, a shrill whistle, was stopped when the insect was ap-
proached and resumed if the observer remained quiet. During
the song, the abdomen was elevated which had the effect of open-
ing wider the tympanal cavities. Sometimes it woidd be con-
tinued while the insect was flying. In addition, a clicking sound
was made by flicking the wings. The females probably can make
the same sound, but those that were observed were quiet and did
not even try to fly when collected. The same insects collected
on manzanita in Washington County, Utah (July 6, 1930 and
June 11, 1932) acted much the same, but did not click their wings.

Okanagana hella Davis

Okanagana hella as frequently observed in the canyons about
the valleys of the Great Salt Lake and at Fish Lake, Sevier
County, Utah (July 2, 1930) and Mt. Hood, Oregon (July 23,
1935) had the same even, high-pitched song of Okanagana
ruhrovenosa. If the two could be heard together, a difference
might be detected. Okanagana hella is found on willows {Salix
sp.) birch, quaking aspen {Populus tremuloicles) and other trees
in the canyons and often at higher elevations than other cicadas.
The wing clicking, however, has not been observed with this
species.

Okanagana vanduzeei Distant

Okanagana vanduzeei were found at several localities. At
Mill Valley and Muir Woods, California (June 17, 1938) they
were on the grass covered hills. The song was more even, but
about the same pitch as that of Okanagana striatipes and their
actions were very similar. They were active, flew readily and
were difficult to approach. After flying, the insects would often
circle back and alight, head upward, at a not very great distance
from the starting point. At Union, Oregon (July 21, 1935) on
a sage brush covered hill, the flight was often at a level above
the sage, when suddenly they would drop to a lower level and
continue until they crashed into another bush. At Soquel and
Elsinore, California (1938) the insects were also on grass.

122 •

Journal New York Entomological Society [Vol. xlviii

Okanagana calif ornica Distant

Okanagana calif ornica at Elsinore, California (June 24, 1938)
was on grass and had a song and actions indistinguishable from
those of Okanagana vanduzeei.

Okanagana luteokasalis Davis

Okanagayia luteohasalis is only associated with the sage brush
and is found over these areas on the plateaus of southern Utah.
The type brood of this species was observed near the town of
Hatch, Garfield County, Utah (June 5, 1931) on the Paunsaugunt
Plateau. (Davis, 1935 p. 302. The type locality is in Garfield
County, not Davis County.)

The brood had emerged just south of the town and to the east
of U. S. Highway 89. In the area, which was about 100 feet in
diameter, the sage was scrubby and only a foot or less in height
and was quite definitely demarcated from the surrounding
country. In this limited area were found thousands of exuvia,
although the adults were beginning to spread out, the skins
exactly marked the limits of the brood. Five to twenty could be
found around every sage brush. It seemed probable that the
large number of nymphs, feeding on the roots, had caused the
plants to suffer over the limited area. With this in mind, the
site was examined from an elevation west of the road and the
area of scrubby sage and exuvia was essentially the same.

Most of the insects were easily captured as is usually the case
with recently emerged broods. Some were copulating and others
had just emerged and were still green, with limp, ghostly, pale
green wings. The pronotum on either side of the mid line was
the first to become dark. The ocelli stood out as orange spots
and the compound eyes were brown. Some of the females were
ovipositing on the sage, but most of them had chosen the rabbit
brush. They would usually take about three minutes to the nest.
One continued to work for ten minutes, but this may have been
the last nest, as it was not filled with eggs and there were
several other completed nests in a line on the stem above. All
the ovipositing females were head down on the stems.

Just north of Nephi, Utah (June 26, 1938) in a sage brush
field, a cicada was heard and when collected, was found to be

June, 1940]

Sugden; Cicadas

123

Okanagana luteohasalis. It was on the stem of the central part
of the sage. No other of this species was found, although they
were searched for. Other broods of the same species have always
been in large numbers so this specimen may have been from a
more distant location. In the same general region, Okanagana
utahensis was also collected, but these insects were always on the
leafy parts of the plant.

Okanagana utahensis Davis and Okanagana
striatipes (Haldeman)

Most of the Okanagana songs are very similar and tend to
be even and continuous. When they are singing in the same
locality, slight differences in the quality become more evident.
Both Okanagana utahensis and Okanagana striatipes are found
on the sage brush and grass-covered foothill of the Wasatch
Mountains, on the terraces of old Lake Bonneville, east of Salt
Lake City. The songs have enough contrast to distinguish the
insects by the sound alone.

At Fort Douglas, Utah (July 4, 1938) Okanagana striatipes
were fairly numerous, alert and well scattered. A few exuvia
were found on the ground or clinging to the plants. A female
flew to a singing male, alighted on the stem, approached the male
and then assumed a parallel position. The male’s song was re-
duced to a few feeble clicks and copulation began. The act was
timed and found to continue for 10 minutes, during which they
were quiet and could be handled without attempting to escape.

In Okanagana striatipes, the song is loud for the size of the
insect and it has a definite rapid undulating character giving a
quivering effect. That of Okanagana utahensis is finer, more
even and somewhat weaker even though the insect is larger.

Okanagana utahensis was the common cicada along the Wasatch
Drive, southeast of Salt Lake City (July 3, 1938) on the sage
brush flats. Most of the insects were located on the leafy parts
of the plants. Only an occasional Okanagana striatipes was
found. A female, Okanagana utahensis, was first observed as it
crawled along side of a male whose song had been heard. Copu-
lation immediately took place and the song was instantly dis-
continued. The act was timed and took 14 minutes during which
time the insects clung to the sage in parallel positions.

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Journal New York Entomological Society [Vol. xlviii

Okanagana synodica (Say)

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 individual. 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 individual 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 resembled the
buff colored soil. Many were mating. The exuvia were on the
ground or attached to the stems.

Okanagana fumipennis Davis

Okanagana fumipennis as observed in San Juan County, Utah
(June 8, 1928 and by H. W. Pickett, June 22, 1932) were on the
wide sage brush flats of the region. They were easily captured,
were poor flyers and if thrown in the air, they would fall to
earth a few feet away. None voluntarily took to wing. The
song was strong and long continued. Many exuvia were on the
ground and clinging to the sage. This species was originally con-
fused with Okanagana schxfferi, (Davis, 1932, 251).

Okanagana schcefferi Davis

The habits and songs of Okanagana schcefferi were entirely dif-
ferent from Okanagana fumipennis when the former was ob-
served along the Mt. Carmel-Zion Park Highway in Utah (June
5, 1931). The insects were found only on the junipers {Juni-
peris utahensis) which were very common in the region. They
were active and flew readily. A few were copulating. The
song had a distinct metallic quality and when several were
singing, the sound resembled the noise made by the safety razor
blade sharpeners in the boot-black stands a few years ago. It
was long-continued and louder than that of the smaller
Okanaganas.

JUNEj 1940]

Sugden: Cicadas

125

Platypedia mohavensis Davis

Platypedia mohavensis were in the same juniper trees with
Okanagana schcefferi. Both sexes made a clicking sound by
flicking the wings and because of the large number, the sound
resembled a shower of hail or shot dropped on wrapping paper.
These insects were not singing.

Platypedia putnami lutea Davis and Neoplatypedia
constricta Davis

Both Platypedia putnami lutea and N eoplatypedia constricta
were collected in South Willow Creek Canyon, Tooele County,
Utah. In May (May 24, 1931) only the former had emerged and
the next month (June 14, 1931) both were present. The songs
were similar, not loud and both sexes of both species were making
the wing-clicking sounds. They were found on rabbit brush
{Chrysothamnus sp.) and sage (Artemisia trident at a) . Platy-
pedia putnami lutea have been collected on juniper in the Kaibab
Forest (June 11, 1924) and on willow (Salix sp.) in the canyons
of Salt Lake County, Utah (May 30, 1930) .

i

Tihicinoides mercedita (Davis) and Tihicinoides cupreo-
sparsus (Uhler)

Tihicinoides mercedita at Dixon, California (June 13, 1938)
had a song very similar to that of Okanagana vanduzeei and
other allied species. The smaller Tihicinoides cupreo-sparsus in
San Diego County, California (June 12, 1931) had an even song,
which was not very loud and was usually stopped before the
insect could be located. The dark color gave a remarkable con-
cealing coloration when the insect was clinging to the dark, dry
twigs and branches of a small shrub.

Eighteen western species of seven genera of cicadas have been
considered. Ten species were observed and collected during 1938
and to these notes have been added other observations made in
recent years.

126

Journal New York Entomological Society [Vol. xlviii

NEW FLEAS

West Coast Crested Fleas Corypsylla and Nearctopsylla (Pacific
Univ. Bull. Vol. 37, No. 1, Mar., 1940, 10 p., 1 pi.) . American
Mole and Shrew Fleas (Pacific Univ. Bull. Vol. 37, No. 2,
April, 1940, 9 p., 2 pL). West Coast Catallagias (Pacific
Univ. Bull. Vol. 37, No. 3, May, 1940, 4 p., 1 pi.) by C. Ander-
son Hubbard.

In these three papers. Doctor Hubbard describes one new
genus and nine new species of fleas, examples of which have been
deposited in colleges, universities and museums in the eastern,
southern, western and central areas of the United States, as well
as in Canada and abroad, thus making the future study of them
easier for students and specialists. Doctor Hubbard questions
the taxonomic value of the “ incrassation ” in Corypsylla and in
his paper on American mole and shrew fleas, presents information
designed to improve the host records, for certain species, as
given by Irving Pox in ‘‘Pleas of Eastern United States,” noted
in this Journal in volume 48, No. 1, p. 32. Other information is
presented, adding to the taxonomic and biologic knowledge of our
American Siphonaptera. — H. B. W.

June, 1940]

Hallock: Sarcophagin^

127

THE SARCOPHAGIN^ AND THEIR RELATIVES
IN NEW YORK, PART I^

By Harold C. Hallock
Pennsylvania State College

INTRODUCTION

An attempt is made in this paper to define the limits of a small
group of muscoid files, the Sarcophagince and their relatives in
so far as the species, which occur in New York, are concerned. A
study of the structures of the male hypopygium of the Sarcophaga
has been included in the hope that a better understanding of the
relationship of the species to each other might be obtained.

The correct understanding of North American SarcophagincB
must be based upon the work of Dr. J. M. Aldrich. Twenty-two
years ago Aldrich (1916) published his Sarcophaga and Allies”
which produced an orderly arrangement of the group for the first
time. Several years later he had an opportunity to visit the
European Museums and study the types of the early authors. As
a result of this study Aldrich (1924, 1930) pointed out the
synonomy of many names that were in common use in American
literature.

Reference must also be made to the important studies by Dr.
R. R. Parker which were published in many widely scattered
papers from 1914 to 1923. Parker described several new species
and made a careful study of the structures of some species of the
Bavinia and Boettcheria groups of the genus Sarcophaga. We
must also depend upon Parker (1916, 1919) for our understanding
of Sarcofahrtia.

A discussion of North American Sarcophagince would not be

1 As this study was completed while the writer was at Cornell University
this article is a contribution from the Department of Entomology, Cornell
University, Ithaca, New York. The writer desires to acknowledge assistance
from Prof. Eobert Matheson, Prof. O. A. Johannsen, Cornell University: Mr.
David G. Hall, U. S. National Museum; Dr. Eichard Dow, Boston Soc. Nat.
History; and Mr. H. G. Eeinhard, Texas Agricultural Experiment Station.
The writer is also grateful for the valuable assistance of his wife, Mabel M.
Hallock, during the proof reading.

128

Journal New York Entomological Society [Vol. xlviii

complete without reference to Mr. David G. Hall who has pub-
lished (1926-1937) about a dozen papers dealing with American
SarcophagidcB. The most important of his papers are ‘[Sarco-
phaga pallinervis and related species in the Americas” (1928)
and ‘‘The Sarcophagince of Panama” (1933).

There are also many taxonomic papers by Dr. C. H. T. Town-
send which must be considered for a complete understanding of
the New York species. The remaining taxonomic, biological and
economic papers, which have been consulted, are widely scattered
in the literature. As a result it seems wise to give a summary of
this information dealing with New York Sarcophagince and their
relatives.

The tribe, Miltogrammini, of the SarcophagidcB will not be con-
sidered in this paper as these flies have been very well treated by
Allen (1926). Very little new information about the New York
Bliltogrammini has been obtained during the past twelve years.
The remaining species of SarcophagidcB, which are known to occur
in New York and a few others which are likely to be found in New
York, are discussed in this paper. Keys are given to separate the
genera of this portion of the family and the species occurring in
the different genera. The discussion of the species will deal
largely with a few distinctive characters, their distribution in
New York and the known hosts or other larval food habits of the
SarcophagincB. The male genitalia will be considered in so far as
they tend to aid in the illustration of the group relationships and
help in the determination of the species.

The known distribution for each species and the total seasonal
occurrence of the adults is given for New York. Credit for the
collection of the different species is not repeated, as this informa-
tion has been given by Leonard (1928) and Hallock (1937).

The writer has found that special methods of preparing the male
genitalia for study are necessary. A combination of several of
the published methods with some variation has proved most satis-
factory. When there is an abundance of material the genitalia
of a few specimens of each species are dissected, soaked in potas-
sium hydroxide to remove the soft muscle parts, etc., washed in
several ch^^nges of water to remove the potash, transferred to cello-
solve and then mounted on slides in Canada balsam in such a

June, 1940]

Hallock: Sarcophagin^

129

manner as not to flatten ont the parts. The minute details of the
specimen, which cannot be seen plainly with a binocular, may now
be examined to advantage with the higher powers of the compound
microscope. This method has been used by Patton (1933) and
Senior- White (1924) but Hardy (1927) objected to the use of
potash. Other specimens should be studied with the genitalia
in situ as described by Parker (1914) and Aldrich (1916) to cor-
rect any errors. The potash may dissolve some of the more deli-
cate parts of the aedeagus or as a result of the dissection and soak-
ing in potash a portion of a sclerite may drop off and be lost.
This is very likely to occur in the case of the portion of the sclerite
immediately surrounding the lower portion of the V of the flfth
sternite.

LIFE HISTORY

The Sarcophagince, as all other Diptera, are insects with com-
plete metamorphoses. They pass through four stages, egg, larva,
pupa and adult during their life cycle.

Egg : Most Sarcophaga do not lay eggs but larviposit flrst instar
larvag, upon the host or larval food. In this case the entire em-
bryological development has taken place within the female fly and
the larvse have hatched before larviposition.

In cases where there is an abundance of meat or other proper
food for the larvae the female fly may oviposit eggs upon the meat.
This oviposition was observed by Smith (1933) during his study
of Sarcophaga securifera. He observed that these eggs either
hatched immediately or within the following eight or nine hours.

The writer has dissected female flies of Sarcophaga bullata,
S. latisterna, and S. misera var. sarracenioides and found larvae
which had already hatched. Several other species of Sarcophaga
have been observed to larviposit when upon the point of death
in a killing jar.

Larva: Each larva passes through three instars during the
development. The first instar is normally very short. Smith
(1933) pointed out that the first instar was completed in thirteen
to sixteen hours at a temperature of 80° F. He also found that
the second molt occurred ten to twenty hours later. The larvae
became completely developed in about six days after deposition
and migrated to the soil to pupate.

130

Journal New York Entomological Society [Vol. xlviii

Under normal temperature conditions Hallock (1929) found
that the larvae of S. latisterna developed in four to five days. At
the same time the larger species, 8. hullata, took an average of
eleven days to develop. It was also observed that hullata devel-
oped slower during cool periods in the summer but its develop-
ment was completed in six to ten days during the hot periods of
the summer. This should be contrasted with the slow develop-
ment of the larvae of 8. fletcheri in pitcher plants. Larvae which
had already reached the third instar stage during July, 1936, were
observed to feed for a period varying from six to ten days before
pupation. This would indicate the possibility that the entire
larval stage may last about a month.

Pupa : The sarcophagid pupal period is subject to considerable
variation at different seasons of the year. The writer found in
early September, 1935, that 8. hullata pupated but the flies did
not emerge when kept at the normal fall temperatures. During
the summer months the pupal period lasted normally ten to four-
teen days. Smith (1933) found that 8. securifera remained in
the pupal stage from ten to thirteen days when kept at a tempera-
ture of 80° P.

Adult : There is very little information available about the
length of life of the adult fly. Smith (1933) reported that the
maximum length of life of 8. securifera was about a month. He
also found that oviposition did not take place until eight to four-
teen days after emergence. It also appears that meat must be
included in the diet of the adult fly in order that larvae will
develop. The writer’s experience in rearing 8. hullata, 8. sar-
racenioides and 8. securifera agree with Smith’s findings that
meat is a necessary part of the fly diet in order that larvae may
develop.

The number of generations in any season depends upon the
species, availability of suitable larval food, and temperature.
8arcophaga cimhicis has been found flying in New York from May
until October. It attacks a wide variety of hosts so it should be
able to find larval food easily. The larval development is rapid
and a cool period, during the summer, does not appear to delay
development as much as in the case of some of the species, such as
hidlata, for example. There must be several overlapping genera-

June, 1940]

Hallock: Sarcophagin^

131

tions although the complete life history of the parasite, cimhids,
has not been worked out. This long period of adult abundance
may be contrasted with Sarcophaga aldrichi which occurs in New
York from late May until early July. This species appears to
be limited to a few Lepidopterous hosts and Malacosoma disstria
is the preferred one. Although the life history has not been com-
pletely studied, available data points to one generation with a
fairly long period of adult emergence. The scavenger groups,
including the dung and dead animal feeders, have several over-
lapping generations.

THE MALE HYPOPYGIAL STRUCTURES

The terminology which has been followed in this study is
adapted from Snodgrass (1935). A few terms which seem more
descriptive of the structure have been adapted from Patton (1933-
1935) after studying his series of papers in the ‘‘Annals of Trop-
ical Medicine and Parasitology. ” A comparison of the terms
which have been used by workers on the Sarcophagidse, is given
in Table 1.

A careful study was made of the hypopygium of about 30 species
from New York and the internal hypopygial structures are illus-
trated in this paper (Figs. 1-97). The structure of Hypopelta
scrofa is described and compared with the variation found in the
other species of Sarcophaginm.

The visceral part of the abdomen of H. scrofa consists of seg-
ments one to five, but the first segment has been obscured by re-
duction and union with the second. As a result the segments (one
to four) mentioned in all systematic papers are segments two to
five of the Dipterous abdomen. In order to avoid confusion the
systematic system has been adapted in the systematic part of this
paper when referring to the first four segments of the abdomen.
The references to the hypopygial structure will be made by a
name assigned to the part or, when mention is made by segment
number; the true abdominal segment number will be used in the
case of all h3rpopygial segments. Segments six to ten and the
fifth sternum are included in the hypopygial structures and many
of these structures have valuable taxonomic characters.

There has been considerable reduction, obliteration, and modi-
fication of the male genitalia structures in H. scrofa and the other

132

Journal New York Entomological Society [Vol. xlviii

Sarcophaginm so it is difficult to correctly name the structures.
The fifth sternum (Fig. 2) is a convex sclerite with a deep U cut
in its caudal margin. Hypopelta scrofa has two raised projections
on the base of the U, which are lacking in the majority of the
Sarcophagince. The U of H. scrofa becomes a V in the Sarcophaga
as illustrated by S. bullata. Segment six appears to be obliterated
as in the case of Pollenia rudis studied by Snodgrass (1935) and
segment seven consists of a smaller tergal plate than in the case
of rudis. The seventh spiracle is present in the membranous walls
on the lateral side of the body just cephalad of the eighth segment.
The eighth segment has a well developed tergum which has often
been mentioned in American taxonomic literature as the first seg-
ment of the hypopygium. The sternum of the eighth segment,
which helps to support the fifth sternum, is reduced to a narrow
sclerotic band, which is incomplete on the right side. The eighth
sternum has no taxonomic value and is not illustrated. The ter-
gum of the ninth segment has been called the second segment of
the hypopygium in taxonomic literature. The sternum of the
ninth segment projects forward and upward from the lower angles
of the ninth tergum and forms the dorsal wall of a large pouch
which is invaginated within the eighth sternum. The phallic
organs normally extend into this pouch. The ninth sternum (Fig.
4) is a broad plate with its posterior angles produced as two arms
which embrace the base of the sedeagus. The plate in the case
of H. scrofa has been reduced and the arms enlarged but Sarco-
phaga hcemorrhoidalis has a normal ninth sternum (Fig. 62).
There are two lateral bars which connect the ninth sternum to the
tenth segment. The tenth segment has been reduced to a mem-
branous area surrounding the anus. There are two lobes con-
nected with this tenth segment which Snodgrass has termed lateral
lobes but in this paper they are called anal forceps (Figs. 1, 3).
The term, forceps, has been used by Aldrich and Parker and is
very descriptive of the sclerite. The lobes which are attached to
the ninth segment and given no definite name by Snodgrass, are
called accessory plates (Fig. 1) which is the name used by Parker
(1914) and Aldrich (1916).

The phallic organs of Sarcophagincc consist principally of a well
developed aedeagus and supporting basal structures. The struc-

June, 1940]

Hallock: Sarcophagin^

133

ture which has been termed sedeagns by Snodgrass has been called
phallosome by Patton, and penis by Aldrich, Parker and several
other taxonomic writers. The sedeagus is a large irregular struc-
ture (Fig. 1). The sclerites called anterior and posterior claspers
(Fig. 1) by taxonomic writers are the lateral processes of the
asdeagus. The shape of the claspers has considerable value as a
taxonomic character but is less important than the aedeagus.
There is a very small sclerite at the base of the claspers which
Parker (1914) termed the fulcral plate. This plate, which varies
very little in the species studied, has no importance in classification
and is not illustrated. The basal apodeme (Fig. 5) is represented
by a low thecal fold. The free end of the basal apodeme extends
cephalad and the attached end surrounds the base of the sedeagus.
The basal apodeme of H. scrofa is broader and shorter than the
average Sarcophagince which is illustrated by Sarcopliaga htUlata
(Fig. 54), a typical Sarcophaga species. The basal apodeme
serves for muscle attachment. The sperm pump sclerite (Fig. 6)
is situated above the basal apodeme and serves for muscle attach-
ment. The sperm pump sclerite is connected with the ejactula-
tory bulb, which connects with the ejaculatory duct. Distal to the
ejaculatory bulb the duct enters the base of the phallus and thus
connects with the asdeagus.

Most of the structures discussed above have been illustrated for
the species of Sarcophaga which occur commonly in New York.
These structures will be discussed further under the genus to show
the relationship of the groups.

CLASSIFICATION

The limits of a small group of New York Muscoid flies, the
Sarcophaginm and their relatives, will be outlined in the follow-
ing pages. These flies belong to the order Diptera and the family
SarcophagidcB.

All the flies in this group have a longitudinal seam along the
upper outer edge of the second antennal segment. The arista is
generally plumose half its length but in some cases it may be only
pubescent or even entirely bare. The mouth parts are well de-
veloped and functional. There is a row of strong bristles on the
hypopleura. The postscutellum is not distinct and the metanotum

Comparison of Hypopygial Terminology Used by Writers on SarcophagincB

134

Journal New York Entomological Society [Vol. XLVIII

b£)

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June, 1940]

Hallock: Sarcophagin^

135

has only a single convexity. There are three or four notopleural
bristles present. The species found in this group are generally
grayish with a more or less tessellated abdomen and three to five
longitudinal stripes on the thorax.

The CalliphoridcB are considered as a distinct family at the
present time. When the world fauna has been completely studied
it may be found necessary to unite the Sarcophagidce with the
CalliphoridcB. As there are always border line forms which can
be placed only with difficulty the Sarcophagidce will be considered
here as a separate family.

In this study the tribe Miltogrammini, of the Sarcophagidce will
not be discussed. This tribe includes species with the accessory
plate as long or longer than the anal forceps. The impression of
the bucca, near the vibrissal angle, is small and not deeply sub-
merged beneath the parafacials and remainder of the bucca. The
sternopleuron never has more than two bristles and the segments
of the abdomen lacks discal bristles.

The group studied in this paper is divided into twelve genera.
In order to make identification as simple as possible all the species
in the several genera are included in the same key (which will be
given in Part II), although keys to the species are given for the
individual genera in some cases.

Key to Genera

1. Arista long plumose 2

Arista short plumose 5

Arista bare or only short pubescent 7

2. First, third and fifth veins with a row of bristles Jolinsonia Coquillett

The fifth vein always bare 3

3. Fourth vein ending at the tip of the wing Opsodexia Townsend

Fourth vein ending far before the tip of the wing 4

4. Front strongly produced, antennal axis strikingly greater than the

vibrissal and frontal profile sloped ; the caudal end of male abdomen

appearing truncate Metoposarcopiiaga Townsend

Front normal ; abdomen not truncate in appearance Sarcophaga Meigen

5. Fourth vein ending at the tip of the wing Opelousia Townsend

Fourth vein ending far before the tip of wing 6

6. Some pale hairs on back of head; epaulets black Hypopelta Aldrich

Only black hairs on back of head ; epaulets light colored.

Sarcofahrtia Parker

7. Median marginal bristles present on first abdominal segment; frontal

bristles not descending below base of antennae; ovipositor of female
drawn out like an awl Macronichia Eondani

136

Journal New York Entomological Society [Vol. xlviii

Median marginal bristles absent on first abdominal segment; frontal
bristles extend below base of antennse 8

8. Abdomen with three rows of shining spots on the second, third, and

fourth segments; these are sometimes confluent; fifth sternite

entire Wohlfahrtia B. & B.

Abdomen pollinose, at most the hind part of segments shining black in
certain lights 9

9. Paraf acials without bristles ; antennae yellow ErytJirandra B. & B.

Parafacials with a row of bristles; antennae black 10

10. Second abdominal segment with median marginal bristles; anterior
acrostichals present; scutellum with a pair of apical bristles.

Oppiopsis Townsend

Second abdominal segment without median marginal bristles; anterior
acrostichals lacking; apical bristles lacking on scutellum.

Laccoprosopa Townsend

DISCUSSION OF GENERA AND SPECIES

Macronichia Roiidani

1859. Eonclani, Dipt. Ital. Prod., vol. 3, p. 229.

1931. Townsend, Ann. and Mag. of Nat. Hist., vol. 8, p. 379.

Townsend (1931) pointed out that the Amohia of authors is
Macronichia Rondani. As a result onr New York species, aurata
and conf undens, fall in the genus, Macronichia.

Genotype. — Macronichia ungulans (Pand.).

Macronichia aurata Coq.

1902. Coqnillett, Proc. U. S. N. M., vol. 25, p. 119.

The two species of Macronichia found in New York are easily
separated as aurata has the fourth abdominal segment and re-
mainder of abdomen posterior to it golden yellow pruinose. The
fourth abdominal segment is crossed longitudinally by brownish
spots.

Length. — 7 to 8 mm.

Records. — Owego ; Ithaca ; Tuxedo. Last of June through
August.

Type.— Cat. No. 6233, U. S. N. M.

Macronichia conf undens Town.

1915. Townsend {Amohio'psis confundens), Proc. Biol. Soc.
Wash., vol. 28, p. 20.

The gray pollinose thorax of this species is marked by three to

June, 1940]

Hallock : Sarcophagin^

137

five black vittse. The abdomen is gray pollinose, marked with
three rows of black spots on the dorsum of the four abdominal
segments.

Length. — 7 to 9 mm.

Records. — Tuxedo. July 24-28.

Type.— Cat. No. 19134, U. S. N. M.

Opelousia Townsend

1919. Townsend, Proc. U. S. N. M., vol. 56, p. 547.

There is very little known about the habits of the Opelousia.
Townsend (1935) recorded that Opelousia have been reared from
snails.

Genotype. — Opelousia ohscura Townsend.

Opelousia ohscura Townsend

1919. Townsend, Proc. U. S. N. M., vol. 56, p. 547.

This species was originally described from three males taken in
Louisiana and one male from North Dakota. The species is mod-
erately common in the south. The writer has seen four specimens
taken near Atlantic City, New Jersey. As the species has a fairly
wide range of distribution it may be expected to occur in New
York.

Length. — 4 to 4.5 mm.

This fly was recorded by Reinhard (1929) as a parasite of the
snail, Succinea luteola. The parasite passed the winter in . the
pupal stage within the shell of its host.

Type.— Male, No. 22249, U. S. N. M.

Opsodexia Townsend

1915. Townsend, Proc. Biol. Soc. Wash., vol. 28, p. 20.

The host relations of this group are unknown. Townsend
(1935) described the female reproductive organs for Opsodexia
and pointed out that the fecundity was very small, ‘‘at most a
dozen at a deposition.” The adult flies have been recorded on
Solidago, Baccharis and other Compositoe.

Genotype. — Opsodexia hicolor (Coq.).

Opsodexia hicolor Coquillett
1899. Coquillett, Jour. N. Y. Ent. Soc., vol. 7, p. 221.

138

Journal New York Entomological Society [Vol. xlviii

1915. Townsend, Proc. Biol. Soc. Wash., vol. 28, p. 20.

1935. Townsend, Manual of Myiology, Pt. 2, p. 256.

The legs are largely yellow and in the light form the abdomen
is yellow except a dark line along the apex of each segment. The
dark form has the femora largely black and the abdomen mostly
dark wdth gray pollinose.

Length. — 5 to 6 mm.

Records. — Ithaca; Hancock; Millwood; Corinth; Lake George;
Kaaterskill; L.I.: Cold Spring Harbor; Wading River; Babylon.
June to September.

Type.— Cat. No. 4121, U. S. N. M.

Opsodexia abdominalis Reinhard

1929. Reinhard, Proc. U. S. N. M., vol. 76, art. 20.

The type locality of Opsodexia abdominalis is Fabyans, N. H.,
so the species can be expected to occur in northern New York.
The fly was taken on flowers of Solidago.

This species may be readily separated from 0. bicolor by the
black densely gray pollinose abdomen. It differs further in that
the fourth vein is broadly bowed and lacks a definite angle, and
the arista has shorter hairs and is practically bare beneath.
Length. — 7 mm.

Type.— Male. Cat. No. 41986, U. S. N. M.

Laccoprosopa Townsend

1891. Townsend, Trans. Am. Ent. Soc., vol. 18, p. 366.

1935. Townsend, Man MyioL, Part II, p. 180.

Curran (1934) placed the species found in this genus in the
Brachycoma although in the New York State List, 1928, he used
the name Laccoprosopa avium Curran for a manuscript species.
It seems best at this time to retain the genus Laccoprosopa.
Genotype. — Laccoprosopa sarcophagina Townsend

Laccoprosopa sarcophagina Townsend

1891. Townsend, Trans. Am. Ent. Soc., vol. 18, p. 366.

The genus Laccoprosopa is represented by a single species in
New York. Leonard (1928) recorded this species in New York as
L. avium Curr. (manuscript name). Mr. David G. Hall, U. S.

June, 1940]

HALLOCK: SARCOPHAGIN.E

139

National Museum, lias kindly studied the specimens and deter-
mined them to be Laccoprosopa sarcophagina Townsend. The
specimens also were compared with a homotype (determined by
Dr. J. M. Aldrich) in the U. S. National Museum.

Length. — 7 to 8 mm.

Records. — Ithaca. Five adults reared from larvae parasitic on
young crows. “Ithaca, N. Y., Cornell University, Exp. No. 1023,
sub. 272 (I. Dobroscky).”

Plath (1922, 1934), Frison (1926) and Townsend (1936)
recorded L. sarcophagina as heavily parasitising five species of
bumblebees (Bombus auricomus, B. feroidus, B. americanorum^
B. bimaculatus, and B. vagans).

Oppiopsis Townsend

1915. Townsend, Proc. Biol. Soc. Wash., vol. 28, p. 20.

1916. Aldrich {Harbeckia) , Sarcophaga and Allies, p. 47.

1918. Townsend, Proc. Ent. Soc. Wash., vol. 20, p. 20.

The genus was erected by Townsend (1915) and in 1918 he
pointed out that Harbeckia Aldrich was synonymous. A single
species Oppiopsis sheldoni has been taken on Long Island.
Genotype— Oppiopsis sheldoni (Coq.).

Oppiopsis sheldoni Coq.

1898, Coquillett {Brachycoma sheldoni), Can. Ent., vol. 30, p.
233.

1915. Townsend {Oppiopsis sheldoni Coq.), Proc. Biol. Soc.

Wash., vol. 28, p. 20.

1916. Aldrich {Harbeckia tesselata), Sarcophaga and Allies, p.

47.

1918. Townsend, Proc. Ent. Soc. Wash., vol. 20, p. 20.

Specimens of 0. sheldoni have been recorded from many locali-
ties throughout the northeastern part of the United States but
the species does not appear to be numerous at any one time.
Length. — 6 to 8-| mm.

Record. — ^Babylon, July 15.

Type.— Cat. No. 4069, U. S. N. M.

Erythrandra B. & B.

1891. Brauer and Bergenstamm, Denkschr. Kais. Akad. Wiss.
AYien, vol. 58, p. 368.

140

Journal New York Entomological Society [Vol. xlviii

1897. Coquillett {Brachycoma) , U. S. D. A. Tech. Ser. No. 7,
p. 131.

1916. Townsend (Eubrachycoma) , Ins. Ins. Men., vol. 4, p. 19.
1924. Aldrich (Erythrandra B. & B.), Ann. Ent. Soc. Am., vol.
17, p. 211.

Aldrich studied the type from the Vienna Natural History
Museum and pointed out in 1924 that onr American species which
Coquillett (1897) described as Brachycoma apicalis belonged in
the genus Erythrandra B. & B. This genus is represented by a
single species in New York.

Genotype. — Erythrandra picipes B. & B.

Erythrandra picipes B. & B.

1891. Brauer & Bergensta, Denkschr. Kais. Akad. Wiss. Wien,
vol. 58, p. 368.

The synonymy of this species is discussed by Aldrich (1924) in
detail. Townsend (1935) states that Eubrachycoma apicalis C. is
distinct from Erythrandra picipes B. & B. on the ground that
Eubrachycoma apicalis has the third longitudinal vein bristled at
least half way to first cross vein while Erythrandra picipes has the
third longitudinal vein bristled only at the base. The specimen
of E. picipes (det. by Aldrich) which the writer has examined
from New York has the third longitudinal vein bristled only at
the base.

Length. — 7 mm.

Records. — Black Mt., Lake George. September.

Type. — Vienna Natural History Museum.

Johnsonia Coquillett

1895. Coquillett, Proc. Acad. Nat. Sci., vol. 47, p. 316.

Hall (1933) pointed out that the diagnostic characters of this
homogeneous genus were the setulose fifth vein and the long
bristles on the cheeks.

Genotype. — Johnsonia elegans Coq.

Johnsonia borealis Reinhard

1937. Reinhard, Bull. Brooklyn Ent. Soc., vol. 32, p. 62.

Reinhard (1937) described Johnsonia borealis from two female
specimens which were taken in Ohio and Michigan. As the spe-

June, 1940]

Hallock; Sarcophagin^

141

cies of this genus are nearly all southern in their distribution this
was the first species of Johnsonia known to occur in northeastern
United States. Hallock (1938) described the male of Johnsonia
borealis and gave the additional distributional records from New
York and Pennsylvania. It should be noted that this species has
been found chiefly in the Upper Austral and Transition Zones.

Length. — 6.5 to 10 mm.

Records. — Poughkeepsie. June to the middle of August.
Figure 7.

Type. — University of Michigan Museum.

Sarcofahrtia Parker
1916. Parker, Psyche, vol. 23, p. 131.

1916. Aldrich (Thelodiscus) , Sarcophaga and Allies, p. 63, 302.

The genus, Sarcofahrtia, was described in detail by Parker
(1916). Later in the same year Aldrich (1916) redescribed the
genus as Thelodiscus and then on page 302 of his ^‘Sarcophaga
and Allies” pointed out the synonym. Parker (1919) added three
new species to the genus.

It is apparent that the genera, Sarcofahrtia and Wohlfahrtia,
are closely related. Both genera have the fifth sternite of the male
undivided, epaulets yellow or brownish, vestiture or back of head
black, the membrane connecting the genital segments to the re-
mainder of the abdomen is unusually short which makes it some-
what more difficult to examine the genitalia of the species in these
two genera than in the case of the Sarcophaga. Parker (1916)
pointed out this relationship.

The habits of the Sarcofahrtia are unknown while the Wohl-
fahrtia are parasites of man and animals and often cause human
myiasis.

Genotype. — Sarcofahrtia ravinia Parker.

Sarcofahrtia ravinia Parker
1916. Parker, Psyche, vol. 23, p. 123.

1916. Aldrich {Thelodiscus indivisus), Sarcophaga and Allies,
p. 64 and p. 302.

1919. Parker, Ent. News, vol. 30, p. 203.

There are numerous records of S. ravinia throughout New
Jersey, New England states, and Quebec, Canada. Although it

142

Journal New York Entomological Society [Vol. xlviii

has never been taken in New York it undoubtedly occurs in the
state.

Length. — 7 to 11 mm.

Type. — Male and female in Mass. Agr. Coll, collection.
Wohlfahrtia B. & B.

1889. Brauer and Bergenstamm, Denkschr. Kais. Akad. Wiss.
Wien, vol. 56, p. 123.

1893. Brauer and Bergenstamm, Verh. Zool.-Bot. Ges. Wien,
vol. 43, p. 501.

When this genus was erected by Brauer and Bergenstamm in-
sufficient distinguishing characters were listed. As a result the
validity of the genus was doubted until Aldrich (1916) stated
the generic characters very clearly.

Townsend (1935) pointed out that the maggots were usually
deposited on the upper lip of man and entered the nostrils, though
at times occurring in the ear and at other times causing dermal
myiasis.

Genotype. — Wohlfahrtia magnifica Walker.

Wohlfahrtia vigil Walker

1848. Walker, List of Dipterous Insects in British Museum, vol.
4, p. 831.

1895. Coquillett {Paraphyto chittendeni) , Jour. N. Y. Ent. Soc.,
vol. 3, p. 105.

1895. Coquillett {Paraphyto chittendeni), U. S. D. A. Ent.

Tech. Bull. No. 7, p. 122.

1916. Aldrich, Sarcophaga and Allies, p. 29.

The three rows of shining black spots on the abdomen separate
this species from all other Sarcophagids found in New York.
Length. — 8 to 14 mm.

Records. — Syracuse; Rochester Junction; Ludlowville; Ithaca;
Lockport; Williamsville ; Utica; Florida. June to August.

This fly has been recorded as a parasite attacking humans by
Walker (1920, 1922, 1931), Felt (1924), Gertson (1933), Mathe-
son (1932) and Ford (1936). Shannon (1923) and Johannsen
(1926) published accounts of W. vigil as a parasite of young
rabbits. Kingscote (1932) gave an account of W. vigil causing

June, 1940]

Hallock: Sarcophagin^

143

myiasis in young fox and mink which resulted in considerable
loss in Canada.

Walker (1937) gave a careful description of the immature
stages of W. vigil.

Type. — Male in British Museum.

Hypopelta Aldrich

1916. Aldrich, Sarcophaga and Allies, p. 49.

When Aldrich (1916) described the genus, Hypopelta, he fully
listed the characters with the exception of the detailed genitalia
differences. The hypopygial studies help to show that the genus
is entirely distinct from other Sarcophagince. The fifth sternite
has a large raised circular projection on each side of the lower
part of the U. The basal apodeme tends to be circular instead
of long and narrow as in the Sarcophaga. The plate portion of
the ninth sternum is much reduced in comparison to its arms
which are larger than the average Sarcophaga. The anterior
clasp er is fused for two- thirds of its length with the ninth sternum.
This fusion has not been observed in the case of any other
Sarcophagince examined.

Genotype. — Hypopelta scrofa Aldrich.

Hypopelta scrofa Aldrich

1916. Aldrich, Sarcophaga and Allies, p. 50.

The males of this species can be easily determined by the row
of very long bristles on the anterior clasper and one long bristle
on the posterior clasper. Both sexes have on each antennae a very
long, thin arista with short plumosity which extends about two
thirds its length.

Length. — 6 to 8 mm.

Records. — Ithaca; Owego. Figures 1 to 6 inclusive.

Type. — Male and female. Cat. No. 20491, U. S. N. M.

Metoposarcophaga Townsend

1917. Townsend, Proc. Biol. Soc. Wash., vol. 30, p. 46.

1919. Parker, Can. Ent., vol. 51, p. 154.

The external characters for separating Metoposarcophaga from
related genera were given only briefly by Townsend when the

144

Journal New York Entomological Society [Vol. xlviii

genus was erected. Aldrich (1916, 1930) placed the genotype,
importuna, in the genus, Sarcophaga. Parker (1919) recognized
Metoposarcophaga, described two new species, and gave a key to
separate the four species placed in the genus.

The study of hypopygial structures helps to show that the genus
is valid. It is unfortunate that M. importuna was the only spe-
cies of Metoposarcophaga available for this study. The V of the
fifth sternum, which is U-shaped in the case of M. importuna, is
much more deeply cut than the average Sarcophagince. The ninth
tergite is very large and gives the abdomen a truncate appearance.
The arms of the ninth sternum are fused at the tip and more
heavily chitinized than the other SarcopJiagince. The internal
portion of the Eedeagus curves around and fuses to the arms of the
ninth sternum as shown in the illustration of the ninth sternum
(Pig. 11). The tip of the sedeagus has a distinct brush-like ap-
pearance on its front side. The pump sclerite is unusually large,
as its diameter is twice the length of the sedeagus, and it has a
definite cap on the small end (Pig. 13). All other Sarcophaginm
studied have a small pump sclerite when compared with the other
hypopygial structures.

Genotype. — Metoposarcophaga importuna (Walker).

Metoposarcophaga importuna (Walker)

1848. Walker {Sarcophaga importuna) , List Dipt. Brit. Museum,
vol. 4, p. 819.

1916. Parker {Sarcophaga pachyprocta) , Jour. N. Y. Ent. Soc.,
vol. 24, p. 171.

1916. Aldrich {Sarcophaga larga, S. pachyprocta), Sarcophaga
and Allies, p. 147, 302.

1919. Parker {M. pachyprocta) , Can. Ent., vol. 51, p. 154.

1930. Aldrich {Sarcophaga importuna), Proc. U. S. N. M., vol.
78, p. 15.

The large abdomen which appears truncate behind, distinguishes
M. importuna from all other Sarcophaginoe which have been found
in New York.

Length. — 6 to 10 mm.

Records. — L.I. : Babylon ; Heckscher State Park ; Dix Hills ;
Long Beach ; Sands Point ; Oak Island. May to August. Pigures
8 to 13 inclusive.

June, 1940]

Hallock: Sarcophagin^

145

LITERATUEE CITED

Aldrich, J. M. 1916. SarcopJiaga and Allies in North America, vol. I.
Thomas Say Foundation, illus.

. 1924. Notes on some types of American Muscoid Diptera in the

collection of the Vienna Natural History Museum. Ann. Ent. Soc.
Am., 17: 209-211.

. 1930. Notes on the types of American two-winged flies of the

genus SarcopJiaga and a few related forms, described by the early
authors. Proc. U.S.N.M., vol. 78, art. 12, no. 2855, pp. 1-43, illus.
Allen, H. W. 1926. North American species of the two-winged flies be-
longing to the tribe Miltogrammini. Proc. U.S.N.M., vol. 68, pp.
1-106, illus.

CoQUiLLETT, D. W. 1897. Revision of the Tachinidse of North America.
U.S.D.A. Tech. Ser. No. 7.

Curran, C. H. 1934. North American Diptera. The Ballou Press, New
York, p. 1-512, illus.

Felt, E. P. 1924. Wohlfahrtia vigil Walker attacking man. Jour. Econ.
Ent., 17: 603.

Ford, N. 1936. Further observations on the behavior of W olilf aJirtia
vigil (Walker) with notes on the collecting and rearing of the flies.
Jour. Parasit., 22: 309-328, illus.

Prison, T. H. 1926. Contribution to the knowledge of the interrelations
of the bumblebees of Illinois with their animate environment. Ann.
Ent. Soc. Am., 19 : 203-234, illus.

Gertson, G. D., Lancaster W. E. B., Larson, G. A. and Wheeler, G. C.

1933. Report of two cases of W oJilf alirtia myiasis in North
Dakota. Jour. Am. Med. Ass., no. 7, pp. 487-488.

Hall, D. G. 1928. SarcopJiaga pollinervis and related species in the
Americas. Ann. Ent. Soc. Am., 21 : 331-352, illus.

. 1933. The Sarcophaginae of Panama. Bull. Am. Mus. Nat. Hist.,

66: 251-285, illus.

Hallock, H. C. 1929. Notes on methods of rearing Sarcophaginae and
the biology of SarcopJiaga latisterna Parker. Ann. Ent. Soc. Am.,
22 : 246-250, illus.

. 1937. A list of the Sarcophagidae of New York (Diptera). Ent.

News, 48 : 258-262.

. 1938. New Sarcophaginae (Diptera). Proc. Ent. Soc. Wash. 40:

95-99, illus.

Hardy, G. H. 1927. Notes on Australian and exotic Sarcophagid flies.

Proc. Linn. Soc. N. S. Wales, 52 : 447-459, illus.

JoHANNSEN, O. A. 1926. W oJilf aJirtia vigil a parasite upon rabbits. Jour.
Parasit., 13: 156.

Kingscote, a. a. 1932. Myiasis in ranch raised foxes. 62nd Ann. Rep.
Ent. Soc. Out., Toronto, pp. 91-93.

Leonard, M. D. 1928. A list of insects of New York. Cornell Univ. Agr.
Exp. Sta. Mem. 101, pp. 1-1121.

146

Journal New York Entomological Society [Vol. xlviii

Matheson, E. 1932. Medical entomology. Charles C Thomas Press, Balti-
more, Md., pp. 1-489, illus.

Parker, E. E. 1914. Sarcophagidae of New England. Males of the genera
Ravinia and Boettcheria. Proc. Boston Soc. Nat. Hist., 35: 1-77,
illus.

. 1916 Sarcopliagidae of New England. III. Sarcofahrtia ravinia,

new genus and new species. Psyche, 23 : 123.

. 1919. North American Sarcophagidae; New species of the genus

Sarcofahrtia. Ent. News, 30: 203.

. 1919. North American Sarcophagidae: flies of the genus Metopo-

sarcophaga Townsend. Can. Ent., 51 : 154-158.

Patton, W. S. 1933. Studies on the higher Diptera of medical and
veterinary importance. Ann. Trop. Med. & Parasit., 27 : 334-337,
illus.

. 1934. Studies on the higher Diptera of medical and veterinary

importance. Ann. Trop. Med. & Parasit., 28: 579-588, illus.

. 1935. Studies on the higher Diptera of medical and veterinary im-
portance. Ann. Trop. Med. & Parasit., 29: 517-532, illus.

Plath, O. E. 1922. Notes on the nesting habits of several North American
bumblebees. Psyche, 29: 189-202.

. 1934. Bumblebees and their ways. The Macmillan Company,

New York City, pp. 1-201.

Eeinhard, H. J. 1929. Notes on the Muscoid flies of the genera Opelousia
and Opsodexia with descriptions of three new species. Proc.
U.S.N.M., vol. 76, art. 20.

. 1937. New North American Muscoid Diptera. Bull. Ent. Soc.

Brooklyn, 32: 62-74.

Senior- White, E. 1924. A revision of the sub-family Sarcophagince in
the Oriental region. Eec. Indian Museum, 26 : 193-284, illus.

Shannon, E. C. 1923. Non-human host records of Wohlfahrtia vigil
(Diptera). Proc. Ent. Soc. Wash., 25: 142.

Smith, C. N. 1933. Notes on the life history and molting processes of
Sarcophaga securifera. Proc. Ent. Soc. Wash., 35: 159-164.

Snodgrass, E. E. 1935. Principles of insect morphology. McGraw-Hill
Book Company, New York City, pp. 1-667, illus.

Townsend, C. H. T. 1915. Proposal of new Muscoid genera for old species.
Proc. Biol. Soc. Wash., 28 : 19-23.

. 1918. Some Muscoid synonymy, with one new genus. Proc. Ent.

Soc. Wash., 20: 19-21.

. 1931. Notes on Old-world Oestromuscoid types. Part I. Ann.

and Mag. of Nat. Hist., 8 (10th ser.) : 369-391.

. 1935. Manual of Myiology, Part II. Charles Townsend & Pilhos

Press, Itaquaquecetuba, Sao Paulo, Brasil, pp. 1-296, illus.

Townsend, L. H. 1936. The mature larva and puparium of Brachycoma
sarcophagina (Town.). Proc. Ent. Soc. Wash., 38: 92-98, illus.

June, 1940]

Hallock: Sarcophagin^

147

Walker, E. M. 1920. WoJilfaJirtia vigil (Walker) as a human parasite.
Jour. Parasit., 7 : 1-7, illus.

. 1922. Some cases of cutaneous myiasis, with notes on the larvae of

WoJilfahrtia vigil (Walker). Jour. Parasit., 9: 1-5, illus.

. 1931. Cutaneous myiasis in Canada. Can. Pub. Health, pp. 504-

508, illus.

. 1937. The larval stages of WoJilfahrtia vigil (Walker). Jour.

Parasit., 23: 163-174, illus.

148

Journal New York Entomological Society [Vol. xlviii

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

Figure 9.
Figure 10.
Figure 11.
Figure 12.
Figure 13.
Figure 14.
Figure 15.
Figure 16.
Figure 17.
Figure 18.
Figure 19.
Figure 20.
Figure 21.
Figure 22.
Figure 23.

Plate II

Lateral view of external genitalia of Hypopelta scrofa.

Fifth sternum of Hypopelta scrofa.

Eear view of anal forceps of Hypopelta scrofa.

Ninth sternum of Hypopelta scrofa.

Basal apodeme of Hypopelta scrofa.

Pump sclerite of Hypopelta scrofa.

Lateral view of external genitalia of Johnsonia loorealis.
Lateral view of external genitalia of Metoposarcophaga im-
portuna.

Fifth sternum of Metoposarcophaga importuna.

Eear view of anal forceps of Metoposarcophaga importuna.
Ninth sternum of Metoposarcophaga importuna.

Basal apodeme of Metoposarcophaga importuna.

Pump sclerite of Metoposarcophaga importuna.

Ninth sternum of Sarcophaga misera var. sarracenioides.

Basal apodeme of Sarcophaga misera var. sarracenioides.
Pump sclerite of Sarcophaga misera var. sarracenioides.

Ninth sternum of Sarcophaga hisetosa.

Basal apodeme of Sarcophaga hisetosa.

Pump sclerite of Sarcophaga hisetosa.

Ninth sternum of Sarcophaga cimhicis.

Basal apodeme of Sarcophaga cimhicis.

Pump sclerite of Sarcophaga cimhicis.

Ninth sternum of Sarcophaga latisterna.

(Plate II)

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

SAECOPHAGIN^

150

Journal New York Entomological Society

Figure

24.

Basal

Figure

25.

Pump

Figure

26.

Ninth

Figure

27.

Basal

Figure

28.

Pump

Figure

29.

Ninth

Figure

30.

Basal

Figure

31.

Pump

Figure

32.

Ninth

Figure

33.

Basal

Figure

34.

Pump

Figure

35.

Ninth

Figure

36.

Basal

Figure

37.

Pump

Figure

38.

Ninth

Figure

39.

Basal

Figure

40.

Pump

Figure

41.

Ninth

Figure

42.

Basal

Figure

43.

Pump

Figure

44.

Ninth

Figure

45.

Basal

Figure

46.

Pump

Figure

47.

Ninth

Figure

48.

Basal

Figure

49.

Pump

Figure

50.

Ninth

Figure

51.

Basal

Figure

52.

Pump

Figure

53.

Ninth

Figure

54.

Basal

Figure

55.

Pump

Figure

56.

Ninth

Figure

57.

Basal

Figure

58.

Pump

Figure

59.

Ninth

Figure

60.

Basal

Plate III

apodeme of Sarcopliaga latisterna.
sclerite of Sarcopliaga latisterna.
sternum of Sarcopliaga latisetosa.
apodeme of Sarcopliaga latisetosa.
sclerite of Sarcopliaga latisetosa.
sternum of Sarcopliaga I’lierminieri.
apodeme of Sarcopliaga I’lierminieri.
sclerite of Sarcopliaga I’lierminieri.
sternum of Sarcopliaga pusiola.
apodeme of Sarcopliaga pusiola.
sclerite of Sarcopliaga pusiola.
sternum of Sarcopliaga stimulans.
apodeme of Sarcopliaga stimulans.
sclerite of Sarcopliaga stimulans.
sternum of Sarcopliaga sueta.
apodeme of Sarcopliaga sueta.
sclerite of Sarcopliaga sueta.
sternum of Sarcopliaga alcedo.
apodeme of Sarcopliaga alcedo.
sclerite of Sarcopliaga alcedo.
sternum of Sarcopliaga aldriclii.
apodeme of Sarcopliaga aldriclii.
sclerite of Sarcopliaga aldrichi.
sternum of Sarcopliaga atlanis.
apodeme of Sarcopliaga atlanis.
sclerite of Sarcopliaga atlanis'.
sternum of Sarcopliaga harhata.
apodeme of Sarcopliaga harhata.
sclerite of Sarcopliaga l)arl)ata.
sternum of Sarcopliaga hullata.
apodeme of Sarcopliaga hullata.
sclerite of Sarcopliaga hullata.
sternum of Sarcopliaga flavipalpis.
apodeme of Sarcopliaga flavipalpis.
sclerite of Sarcopliaga flavipalpis.
sternum of Sarcopliaga fletcheri.
apodeme of Sarcopliaga fletcheri.

[VoL. XLVIII

(Plate III)

I

OUR. N. Y. Ent. Soc.), Vol. XLVIII

SAECOPHAGIN^

152

Journal New York Entomological Society [Vol. xlviii

Figure

61.

Pump

Figure

62.

Ninth

Figure

63.

Basal

Figure

64.

Pump

Figure

65.

Ninth

Figure

66.

Basal

Figure

67.

Pump

Figure

68.

Ninth

Figure

69.

Basal

Figure

70.

Pump

Figure

71.

Ninth

Figure

72.

Basal

Figure

73.

Pump

Figure

74.

Ninth

Figure

75.

Basal

Figure

76.

Pump

Figure

77.

Ninth

Figure

78.

Basal

Figure

79.

Pump

Figure

80.

Ninth

Figure

81.

Basal

Figure

82.

Pump

Figure

83.

Ninth

Figure

84.

Basal

Figure

85.

Pump

Figure

86.

Ninth

Figure

87.

Basal

Figure

88.

Pump

Figure

89.

Ninth

Figure

90.

Basal

Figure

91.

Pump

Figure

92.

Ninth

Figure

93.

Basal

Figure

94.

Pump

Figure

95.

Ninth

Figure

96.

Basal

Figure

97.

Pump

Plate IV

sclerite of Sarcophaga fletcheri.

sternum of Sarcophaga hcemorrhoidalis.

apodeme of Sarcophaga hcBmorrhoidalis.

sclerite of Sarcophaga hcBmorrhoidalis.

sternum of Sarcophaga houghi.

apodeme of Sarcophaga houghi.

sclerite of Sarcophaga houghi.

sternum of Sarcophaga hunteri.

apodeme of Sarcophaga hunteri.

sclerite of Sarcophaga hunteri.

sternum of Sarcophaga johnsoni.

apodeme of Sarcophaga johnsoni.

sclerite of Sarcophaga johnsoni.

sternum of Sarcophaga parallela.

apodeme of Sarcophaga parallela.

sclerite of Sarcophaga parallela.

sternum of Sarcophaga rapax.

apodeme of Sarcophaga rapax.

sclerite of Sarcophaga rapax.

sternum of Sarcophaga reversa.

apodeme of Sarcophaga reversa.

sclerite of Sarcophaga reversa.

sternum of Sarcophaga scoparia var. nearctica.

apodeme of Sarcophaga scoparia var. nearctica.

sclerite of Sarcophaga scoparia var. nearctica.

sternum of Sarcophaga securifera.

apodeme of Sarcophaga securifera.

sclerite oi Sarcophaga securifera.

sternum of Sarcophaga sinuata.

apodeme of Sarcophaga sinuata.

sclerite of Sarcophaga sinuata.

sternum of Sarcophaga uncata.

apodeme of Sarcophaga uncata.

sclerite of Sarcophaga uncata.

sternum of Sarcophaga ventricosa.

apodeme of Sarcophaga ventricosa.

sclerite of Sarcophaga ventricosa.

Four. N. Y. Ent. Soc.), Vol. XLVIII

(Plate IV)

SAECOPHAGIN^

97

June, 1940]

Pratt: Ichneumonid^

155

STUDIES ON THE ICHNEUMONID^ OF NEW
ENGLAND (HYMENOPTERA)

PART V

THE EXTERNAL MORPHOLOGY OF AROTES AMCENUS CRESSON

By Harry D. Pratt

INTRODUCTION

The ichneumon-flies make up unquestionably the largest and
most extensive family in the order Hymenoptera, and it is com-
posed of a vast number of minor groups, representing hundreds
of genera and thousands of species in North America alone. Un-
like other families, the ichneumon-flies, without a single excep-
tion, are all genuine parasites, and destroy or devour the eggs,
larvae, pupae, or imagoes of other insects. Almost every insect,
whose biology is known, has at least one ichneumonoid parasite,
and the more important economic insects (as the gypsy and
browntail moths) have dozens of ichneumon-fly parasites. The
family is, therefore, of the greatest economic importance from the
viewpoint of biological control, and in some cases, as with the
Oriental fruit moth, parasitic control is the only practical type
that is used.

Arotes amoenus Cresson was chosen as the subject of this study
because it is common throughout New England, so that an abun-
dance of material could be collected for study. It is one of the
larger and more primitive of the ichneumon-flies and, for this
reason, is well-suited for such a study.

One of the greatest difficulties in the study of the systematic
groups is the lack of uniformity in the terminology used to de-
scribe the various morphological features of the insects concerned.
This fact is especially noticeable in the descriptions of the
Ichneumonidas when one attempts to use the keys of Asmead,
Schmiedeknecht, or some of the other nineteenth century taxono-
mists, whose descriptions are based either on a purely artificial
system of termiology (as is the case in the naming of the wing

1 Thesis submitted in partial fulfillment for the degree of Master of Science
in 1938 at the Massachusetts State College, Amherst, Massachusetts.

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Journal New York Entomological Society [Vol. xlviii

veins according to the Cressonian system), or on a system con-
taining a great many incorrect terms (as is the case in the mis-
application of the terms “prothorax,” “ postscntellnm, ” “meta-
notnm, ’ ’ and ‘ ‘ metatarsus. ’ ’

The great advances which have been made in comparative
morphology have made it not only wholly unnecessary, but
highly undesirable, to continue such a type of taxonomic work.
The following detailed, purely morphological study of one species
has been made to serve as a basis for future taxonomic work, and
it is hoped that this study will be an aid in determining the
morphological status of many structures which have heretofore
been misnamed, and in clarifying the terminology used in
describing the ichneumon-flies.

The writer wishes to express his indebtedness for the invalu-
able help given by Dr. G. C. Crampton in the morphological
study. To Dr. J. C. Bradley and to Dr. Herbert H. Ross the
writer wishes to express his appreciation for their help in the
study of the wing venation ; and to Dr. Richard Holway for his
help in the study of the pretarsus. To Dr. C. P. Alexander the
writer is indebted for his constant enthusiasm and kindly criti-
cism throughout the course of this research.

THE HEAD

The head of the male is slightly more than two-thirds the size
of that of the female, and its features are less strongly developed
than are those of the female. When viewed from in front, the
head appears somewhat broader than long and is flattened dor-
sally along the parietals. In lateral aspect the compound eyes
appear to be of almost exactly the same width as the gena. The
sutures which demark the areas of the cranium in a typical or-
thopteroid insect have become obsolescent in Arotes, so that the
head is remarkable for its lack of sutures. Two of these sutures,
the occipital and hypostomal, have been replaced, however, by
distinct carinse which are of great importance in bounding the
sclerites of the head.

Head Capsule

The principal areas of the cranium (Pig. 1) are the dorsal
parietals, the median facial, or front o-cly peal area, the lateral

June, 1940]

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157

genm, the occipital arch (composed of the occiput and postgence)
and post-occipnit surrounding the foramen magnum, and the
narrow hypostomal areas upon which the mouth-parts articulate.

The parietals and front o-cly peal areas extend cephalad from
the occipital carina between the compound eyes to a line drawn
between the anterior tentorial pits. The parietals (pa) (Fig. 1)
include the dorsal surface of the epicranium between the occipi-
tal carina and the antennal sclerites, with the exception of the
post-frons which is a more or less pentagonal area extending
caudad from the antennal sclerites to the median ocellus, as will
be explained in greater detail in the discussion of the frons.

The frons (af and pf) is a median unpaired sclerite extending
from the two anterior tentorial pits to, and including, the median
ocellus. According to Crampton (1921), ‘‘When the frontal
suture is absent, if a line be drawn across from the top of one
antennal fovea to the other, and at either end of this line an
angle of forty-five degrees is constructed, the sides of the isosceles
triangle thus formed correspond in a general w^ay to the Y-shaped
epicranial suture.” In Arotes these antennal foveae are rather
deep depressions which serve for the reception of the scape of the
antenna when the antennae are curved back over the body. These
foveae extend caudad almost as far as do the eyes, and then the
frons narrows quite rapidly so as to include the median ocellus.

Because of the noticeable constriction of the frons at the an-
tennae, Crampton (1921) divides the frons by a line drawn be-
tween the bases of the antennae into an antefrons (af) and a ptost-
frons (pf). In Arotes amoenus such a line occurs naturally, ap-
pearing as a darkly-colored hair line extending to the middle of
each antennal sclerite, from the distinct carina on the middle of
the post-frons. Therefore the post frons (pf) is a roughly pen-
tagonal area extending caudad to the median ocellus, while the
ant'efrons (af) is a more or less quadrangular area extending
cephalad to the anterior tentorial pits.

For taxonomic purposes, the morphological antefrons extend-
ing caudad to the base of the antennse might well be called the
“frons.” Similarly, the wdiole dorsal surface of the head could
be called the “vertex,” since the post frons merges completely
with the parietals and the epicranial suture has also disappeared

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Journal New York Entomological Society [Vol. XLVIII

doing away with the necessity of the term ‘‘parietals’’ which
should be used for the paired dorsal sclerites. Therefore, for
taxonomic purposes in Ichneumonidge, it would seem best to use
but two terms, ‘‘vertex” and “frons,” for the frontal aspect of
the head, and to use the antennae as the line of division : the dor-
sal part above the antennae, the ‘ ‘ vertex ’ ’ ; the ventral part below
the antennae, the “frons.”

On either side of the true antefrons (af) lie the areas known
as the parafrontals (paf). These extend caudad to the antennal
sclerites, cephalad to the base of the mandibles, and laterad to
the ocular sclerites (asc) and the suhocular suture (sos). Taxo-
nomically the parafrontals, known as the “inner orbits,” are
important in specific determinations.

The narrow space separating the compound eye from the base
of the mandible, known taxonomically as the “malar space,” is
a very important character in both generic and specific deter-
minations because its length in relation to the basal width of the
mandible is constant. Cushman (1920) sul) ocular suture,

which arises from the ventral margin of the ocular sclerite and
extends ventrad to the dorsal articulation of the mandible, the
‘ ‘ malar suture. ’ ’

The compound eyes are large and convex, with their inner mar-
gins parallel and not emarginate within, or only very slightly so
opposite the antennae. Their surface is reticulate, being com-
posed of the hexagonal facets of the large number of ommatidia.
They are surrounded by narrow, poorly-demarked, elliptical
ocular sclerites (osc) which extend shelf-like into the head cap-
sule. These chitinous ingrowths appear lens-shaped and are
pierced in the middle by a foramen through which the optic
nerves pass connecting the optic lobes with the nerve endings of
the ommatidia.

The endoskeleton of the head, which braces the lower portion
of the head capsule, is called the tentorium. In Arotes two pair
of cuticular invaginations, known respectively as the anterior
and posterior arms of the tentorium, unite within the head to
form a framework arching over the ventral nerve cord, but pass-
ing beneath the stomodceum and supporting the latter. The body
of the tentorium, formed by the fusion of the anterior and poste-

June, 1940]

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159

rior arms, is not clearly demarked and seems to be composed
only of that portion of the tentorium immediately cephalad of
the foramen. The anterior arms arise from the anterior tentorial
pits. Each anterior arm extends candad as a long arched chiti-
nous bar to the body of the tentorium, and serves for the attach-
ment of the muscles moving the mouth-parts. The posterior arms
arise as short invaginations of the hypostomal region, one on each
side of the midgular suture. They are barely distinguishable as
short chitinous bars before they fuse into the body of the ten-
torium which lies immediately cephalad of the foramen. The
dorsal arms are secondary outgrowths of the anterior arms and
are strongly developed in Arotes. They extend cephalad from
the anterior arms to the lateral portions of the antennal sclerites,
and serve as points of attachment for the antennal muscles.

The anterior tentorial pits (at) (frontal pits of Crampton,
1921) are well defined depressions that outwardly mark the two
points from which the anterior arms of the tentorium arise. As
determined by a study of the ental surface, the lateral prolonga-
tions of these arms extend obliquely ventrad to the base of the
mandibles as a pair of lines demarking the clypeus within, but
as in most parasitic Hymenoptera (see Snodgrass, 1935, p. 297),
these ‘‘bars are not connected between the pits.” Posteriorly,
these epistomal bars are continued externally as strong marginal
ridges, differentiated into a pleurostoma (plst) bearing the man-
dibular articulations, and a hypostoma (hp) supporting the max-
illae and labium. In the discussion that follows, the pleurostoma
is considered as the ventral portion of the gena ; while the hypo-
stoma is considered as a separate hypostomal area originally
derived from the postgenae.

The clypeus (cl), which is a flat, trapezoidal area extending
cepahalad from the anterior tentorial pits to the base of the
mandible, is bounded laterally by a line drawn from these depres-
sions to the base of the mandible. The clypeus is slightly in-
flexed at the apex, but since there are no distinct sutures or
Carinas, nor any ental thickenings, it is rather difficult to divide
it satisfactorily into a postclypeus and an anteclypeus. Taxono-
mists usually describe the clypeus as “truncate,” but specimens
boiled in potassium hydroxide and examined under high magnifi-

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Journal New York Entomological Society [Vol. xlviii

cation show that the clypeus is shallowly emarginate with a slight
median projection, as is shown in the figure.

The gence (ge) are long, rounded sclerites beneath and behind
the *eyes and are of about the same width as the compound eyes
when seen from a lateral viewpoint. They are not demarked from
the parietals but may be said arbitrarily to begin on a line drawn
from the most dorsal point of the compound eyes to the occipital
and hypostomal caringe, and to extend anteriorly to the base of
the mandible and the snbocular suture. The ventral part of the
genge, therefore, is the pleurostoma (plst) (Fig. 2) of Snodgrass
(1935). Just laterad of the snbocular suture is a short, blunt
tooth forming the pleurostomal condyle, or the cranial portion of
the dorsal articulation of the mandible. Immediately posterior
to this tooth is a distinct emargination and elevation of the chitin.
Careful examination with high magnification reveals that there
is a corresponding elevation on the mandible, while in between
the two lies the slender extensor muscle which opens the man-
dibles. The reason for this outpouching of the chitin will be ex-
plained in the discussion of the mandible. Many taxonomists
call the genge the “cheeks” or the “outer or posterior orbits.”
The coloration and striation of the gense is much used in the sepa-
ration of subgeneric groups and species of Ichneumonidge.

The occiptal arch. On the postero-caudal surface of the head
(Fig. 2), between the occiptal carina and the postoccipital suture,
is a horseshoe-shaped sclerite called the occipital arch. Although
there is no suture dividing it, the occipital arch is generally said
to be composed of a dorsal part, the occiput (oc), and the ventro-
lateral extensions of this, lying posterior to the genge, called the
postgencB (pg).

The narrow posterior rim surrounding the foramen and de-
marked from the occiput by a post-occipitat suture (pcs) is called
the post-occiput (pc). Snodgrass (1935, p. 112) suggests that,
“The post-occiput probably is a sclerotic remnant of the labial
segment.” If this statement is true, then the postocciput is the
only sclerite entering into the composition of the head to retain
the primitive Annelid alignment.

In Arotes, since there is no subgenal suture, it seems best to
consider the subgenal area as the hypostomal area composed of

June, 1940]

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161

the narrow marginal area on the sides of the cranium upon which
the mouth-parts are articulated and demarked posteriorly by the
hypostomal carina (hpc). The postgenal region of the cranium
has become greatly elongated, thereby creating a long space be-
tween the foramen magnum and the base of the mandibles and,
at the same time, displacing the labium ventrally by mesal out-
growths of the subgense, called the hypostomal bridge, so that the
bases of the labium and the maxillae have become united into a
labio-maxillary complex. According to Snodgrass (1935) the
hypostomal bridge consists of “expansions of the hypostomal
parts of the subgenal margin of the cranium and each is sepa-
rated by a distinct carina from the corresponding postgenal
region. ’ ’

The hypostomal carina (hpc) starts at the base of the mandible
mesal to the mandibular condyle, and extends posteriorly joining
onto the occipital carina and then curving inward towards the
foramen where is finally ends on the midgidar suture determined
by the posterior arms of the tentorium. Due to the approxima-
tion of the posterior arms of the tentorium on the ventral margin
of the foramen, the gula itself has become atrophied and is repre-
sented only by the midgular suture.

Appendages of the Head

The appendages of the head include the antennge, labrum, man-
dibles, maxillge, and labium. Just as the characteristic feature
of the head capsule of Arotes is the development of the hypos-
tomal bridge, so the characteristic feature of the mouth-parts is
the close association or union of the maxillae, the labium, and the
hypopharynx to form the underlip complex, in which the ligula
and hypopharynx are combined in a median lobe on which the
salivary ducts open.

Each antenna (Fig. 4) is composed of about forty segments
and may be divided into three principal parts. The first and
largest segment, by which the antenna is attached to the head, is
termed the scape (scp). Externally it has somewhat the shape
of a funnel with flaring sides, cut diagonally across the top ; in-
ternally, it expands into a basal bulb so that the antenna articu-
lates with the antennal sclerite (as) by a ball and socket joint

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Journal New York Entomological Society [Vol. xlviii

allowing free movement in all directions. The basal bulb has an
opening through which run the muscles moving the scape. These
muscles have their origin on the dorsal arm of the tentorium,
while other muscles arising on the base of the pedicel, move the
pedicel and flagellum together, thus controlling the movements
of the antennae. The pedicel (pdc), or second segment, is quite
short and is sometimes almost hidden from sight, so deeply is it
inserted in the scape. The remainder of the antenna is collec-
tively termed the flagellum (fl) or clavola. The reason for con-
sidering the greater part of the antenna as one of the three prin-
cipal divisions has been explained by Snodgrass (1935, p. 132)
who wrote, ‘‘Since the flagellar divisions in Orthopteriod insects
increase in number from one instar to the next, they appear to be
secondary subdivisions of one primary antennal segment.” The
first flagellar segment is weakly sclerotized a short distance above
the base, giving the effect of a very short segment suggestive of
the ring- joints of Chalcidoidea and high Hymenoptera, but there
is no real articulation or segmentation at this point. The basal
flagellar segment is three and one half times as long as broad, and
each succeeding segment is somewhat shorter than the preceding
one, so that the segment preceding the terminal on.e is but one
and one half times as long as wide. The antennae, therefore, may
be called attenuate. Note that the terminal segment is twice as
long as the preceding one, due, most probably, to a fusion of the
last two segments.

The lahrum (Im) is attached (Fig. 1) to the inner surface of
the clypeus by membrane and projects below it as a semicircular
sclerite. A study of the inner surface of the clypeus reveals that
the labrum is attached to the clypeus at about its middle dorso-
ventrally. The posterior limits of the labrum seem to be deter-
mined by membranous thickenings which have their origin just
mesad of the dorsal articulations of the mandibles. These mem-
branes curve posteriorly a short distance, paralleling the episto-
mal bars, and then converge medially and ventrally on the cly-
peus to form the epipharyyix, a flexible, triangular membranous
lobe which projects below the labrum as a short, sharp, pointed
protuberance about as long as the dorsal and outer tooth of the
mandibles. Along the apical margin of the labrum, on both the

June, 1940]

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163

inner and outer surfaces, are a few long hairs which may be
sensory in nature.

The mandibles (md) (Fig. 7) are placed just behind the epi-
pharynx, one on each side of the mouth opening. They are of
medium size, somewhat curved, and taper apically ending in two
blunt teeth, of which the ventral one is the longer, being half
again as long as the outer and dorsal tooth. Each mandible has
two principal articulations with the head capsule ; an anterior,
or dorsal articulation, and a 'posterior, or ventral articiUation.
Both of these are of the ‘‘ball and socket” type of joints. The
dorsal articulation consists of a pleurostomal condyle situated
just laterad of the subocular suture which fits into a shallow
socket, or ginglymus, on the mandible. The ventral articidation
consists of a deep socket on the pleurostomal region of the gena,
just cephalad of the hypostomal carina, and mandibular condyle
(hypocondyle of Crampton, 1921) which fits into this socket.
Each mandible is moved by two muscles : an extensor or abductor
muscle which opens the mandible ; and a flexor or adductor mus-
cle which closes the mandible. The extensor muscle is relatively
small. It is inserted on a chitinous outpouching of the outer
face of the mandible between the dorsal socket and the ventral
condyle. The outer and anterior face of the mandible is almost
flat between the two articulations and parallels the axis line be-
tween the two so closely that this outpouching is necessary in
order that the base of the extensor muscle be placed sufficiently
far outside the axis line to give effectiveness as a muscle. The
insertion of the extensor muscle is contiguous to the anterior
articulation of the mandible, but it is twice its width from the
posterior articulation. The flexor muscle is huge and is com-
posed of several bundles of fibers inserted on a large apodeme
attached at the inner angle of the mandible almost posterior to
the anterior articulation. When not in use the tip of one man-
dible extends over and covers the tip of the other. There seems
to be no special arrangements as to which is outer, for, in speci-
mens examined, the right seemed to cover the left as often as the
left covered the right.

The labio-maxillary complex of Arotes is attached to the poste-
rior wall of the head between the postgenal margins of the era-

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Journal New York Entomological Society [Vol. xlviii

Ilium by ample membranes, which allow it free movement on the
snspensoria formed by the maxillary cardines. As explained by
Snodgrass (1935) a line of flexure crosses the posterior part of
this complex through the stipito-cardinal sutures of the maxilla.
The basal portion composed of the cardines is bent abruptly
toward the head, where it is attached ; and the distal portion com-
posed of the labium and the body of the maxillae lies parallel
with the ventral surface of the head. The entire organ can thus
be extended by swinging distally on the maxillary cardines. The
maxillEe lie one on each side of the labium and are connected
to it by the membranous submental area and by maxillary mem-
branes.

Maxillce (Fig. 6). In each maxilla the car do (cd) is a heavily
sclerotized sclerite, somewhat triangular in shape, which tapers
to the base and is bent entally. Its extreme base is modifled to
form two processes which articulate deep in the head capsule
with two arm-like projections of the hypostomal bridge, one on
each side of the midgular suture. There is apparently no cardo-
condyle by which the cardo articulates with the post-genal area,
but the whole maxilla is capable of some movement due to the
concavity of the hypostomal area and the convexity of the cardo
and stipes. The cardo (cd) articulates with the stipes (st) by
means of a distinct membranous hinge, and their planes form a
distinct angle at the union. The stipes (st) is the broad and
elongate sclerite forming the body of the maxilla. On its ventral
surface is a distinct carina, or ridge, extending diagonally across
the stipes from the cardo to the palpus (mxplp). There is no
trace of the parastipital area. The 5-segmented palpus (mxplp)
is inserted latero-ventrally on the distal end of the stipes. Just
distal to the base of the palpus is attached a large, fleshy lobe
called the galea (ga), which is distinctly hollowed out for the
reception of the lacinia. The lacinia (la), which is hinged to the
distal end of the stipes, is a large, flat, weakly sclerotized lobe
with two slight tooth -like expansions (as is shown in Fig. 6).

Lahium (Fig. 5). The labium (lb) is composed of a mem-
branous sul)niental region (smt) and a distal prementum (prmt)
bearing the palpi, glossa, and paragloss^e. The submentum is
not sclerotized and probably is situated in the membranous re-

June, 1940]

Pratt: Ichneumonid^

165

gion attached posteriorly to the stipes at the cardostipital suture
and extending anteriorly about two-thirds of the length of the
stipes. The jjrementum (prmt) (mentum of many authors) is
a large, strongly convex, heavily sclerotized plate, the basal and
lateral portions of which are strongly produced and bent over so
as to extend dorsally to meet the hypopharynx which is situated
on the dorsal surface of the labium. The prementum (prmt) is
expanded centrally and is very convex, sloping abruptly to the
glossas beyond. At about the middle of the prementum are at-
tached the 4-segmented labial palpi (laplp) which are only about
as long as the first three segments of the maxillary palpi and are
much shorter and somewhat more slender than the latter. The
glossa (gl) is a large, bilobed, membranous pad attached to the
distal portion of the prementum. Its apical margin is covered
with short, plush-like hair, while the rest of the glossa lacks this
covering. The ventral surface of the glossa lacks this covering.
The ventral surface of the glossa seems to be longitudinally striate
throughout, but the dorsal surface is transversely striate on the
distal two-thirds and is smooth and rather heavily sclerotized on
the basal third. On either side of this narrow sclerotized base of
the glossa occurs a small membranous pad, very similar in struc-
ture to the glossa, and these pads apparently represent the
paraglosscB (pgl). These paragloss^e are closely appressed to the
prementum and are best seen from the dorsal view. Viewed
from below, they appear as membranous lobes dorsal and cephalad
to the palpi (Fig. 6). A membranous papilla-like protuberance,
which is the hypopharynx, occurs on the median line of the dor-
sal surface of the prementum just caudal to the base of the
glossa. The identity of the hypopharynx is determined by the
presence of the opening of the salivary duct at the base of this
distinct papilliform protuberance.

NECK OR CERVIX

The neck or cervix connects the head with the thorax and covers
the antero-ventral region of the thorax. It is composed of a
single pair of plates known as the lateral cervicals (Ic) (Fig. 9)
which articulate with the head by means of the cephaligers and
with the cox^ by a coxal process.

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Journal New York Entomological Society [Vol. xlviii

The lateral cervicals (Ic) are broad, truncate sclerites, taper-
ing from base to apex, and are joined with membrane along the
median line. As shown by Crampton (1926), each lateral cer-
vical results from the fusion of the enlarged lateral cervical, the
proepisternum, and the proepimeron; while the prosternum, for
which the lateral cervicals have so frequently been mistaken, is
represented by a tiny plate partially concealed between the coxae.

At the anterior end of the lateral cervicals, the cephaligers
arise on the inner surface and enlarge into knob-like projections
which are the actual fulcral points of the head with the cervix.

A blunt, tooth-like projection, which is called the coxal process,
is found on the ental surface of each lateral cervical near the lat-
eral posterior angle of the sclerite and this projection forms a
part of the apparatus for the attachment of the coxa of the fore
leg.

THORAX

Prothorax

In Arotes the prothorax (Pig. 9) consists largely of the pro-
notum (Ni) for the propleura have fused with the lateral cervi-
cals, as explained before in the discussion of the cervix, and the
prosternum is reduced to a tiny sclerite lying between the cox®.

The pronotum (Ni) is a narrow transverse plate produced lat-
erally and posteriorly into distinct, triangular lobes which extend
posteriorly to the tegulse, and ventrally to the bases of the fore
cox£e. The anterior margin of the pronotum is broadly emargi-
nate, while the posterior margin is arcuately emarginate and
overlaps the front margin of the prescutum of the mesothorax
(pSCs).

The pro sternum is a small sclerite composed of an anterior
hasisternum and a posterior furcasternum. The triangular
basisternum seems to send chitinous processes to the basal end of
the lateral cervicals, while the furcasternum forms a broad
rounded lobe between the fore coxae and is the base to which the
sternal apophyses of the endoskeleton of the prothorax are
attached.

Mesothorax

The tergum of the mesothorax (9) is divided into a prescutum,
scutum, scutellum, and parascutellum.

June, 1940]

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167

The prescutum (psca) is the large, arched, anterior plate of
the tergum which extends cephalad under the pronotum and
there gives rise to the prephragma of the mesothorax. It is sepa-
rated from the scutum by the notaulices (usually called the
notauli by systematists and variously called the V-shaped suture,
convergent sutures, or the parapsidal sutures by morphologists).

The scutum (SC2) is the large, rather convex plate rather defi-
nitely divided by the notaulices into two lateral portions some-
times called the parapsides, to which the tegidce and the wings
are attached. Behind the scutum lies the scutellar fovea which
is twice as long and contains several indistinct longitudinal carinae
along its middle.

The scutellum (scB) is twice as long as this scutellar fovea
and is connected with the scutum by a chitinous bar on each end
of the fovea. Its apex is truncate. The postscutellum is hidden
beneath the scutellum, where it is fused with the endoskeleton of
the mesothorax.

The parascutellum consists of two triangular plates extending
from the scutum and scutellum to the wing bases, where they
form the posterior margin of the fore wing base and the anterior
margin of the hind wing base.

The mesepisternal and mesosternal plates are fused into a
single sterno-pleural plate making the identification of the lateral
and sternal plates rather difficult. Snodgrass (1910) states that
the “mesopectus consists of three principal plates, the combined
sternum and episterna, and the two epimera. ’ ’

The mesepisternum and mesepimeron were originally sepa-
rated by a pleural suture corresponding to the strong, internal
pleural ridge, forming the wing process above and the coxal proc-
ess below. In Arotes this suture is represented by a row of small,
pit-like depressions similar to those in Trogus which are well
shown in the fine figures by Snodgrass (1910). Although this
row of punctures is scarcely recognizable as a suture, the pres-
ence of the pleural ridge directly beneath it proves that it is the
pleural suture, the important landmark on the mesopleura in the
division into mesepisternum and mesepimeron.

The mesepisternum (eps2) is the large, more or less triangular
plate forming the greater part of the mesopleura. Its anterior

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Journal New York Entomological Society [Vol. xlviii

margin lies concealed beneath the lateral portions of the prono-
tnm (Ni) while its posterior, or dorso-caudal, margin, is de-
marked by the pleural suture extending from the wing base to
the coxal process. While the mesepisternum is not definitely de-
marked from the mesosternum, it is divided into an anterior and
posterior part by the prepectal carina (pete). This is a strong,
transverse carina originating on the median line of the sternum,
which curves posteriorly away from the coxa, and then curves
dorso-cephalad on the mesepisternum. Viereck (1916) used the
presence or absence of this carina as a primary character for the
separation of the various Ichneumonoid families in his Hymen-
optera of Connecticut key, but this view is not accepted today by
most taxonomists.

The mesepimeron (epm2) is the narrow band lying dorso-
caudad of the mesepisternum and separated from it by the pleural
suture.

The mesosternum, as already mentioned, is fused with the
mesepisternum to form a single sterno-pleural plate and there is
no line of demarkation between the two areas.

Along the median line of the sternum, starting at the prepectal
carina, there is a row of pit-like depressions which outwardly
mark the line of attachment of the apophyses of the mesothoracic
endoskeleton. The pleurosteriial plate is modified posteriorly
to form a coxal process projecting into the mesothoracic coxal
cavity from its antero-median margin, while the pleural coxal
process, formed at the end of the pleural ridge, projects into the
coxal cavity from the postero-lateral margin thus forming the
two points of articulation for the mesathoracic leg.

Mesad of the coxal cavities lies a triangular sclerite known as
the furcasternum which is poorly demarked by carin^e from the
rest of the mesosternum (eusternum of the mesothorax). This is
deeply invaginated along the median line and bears the internal
structure called the furca.

Metathorax

The tergum of the metathorax (Fig. 9) is composed of four
sclerites which are called the prescutum, the scutum, the scutel-
lum, and the postscutellum (all designated in figure 9 as Ns).

June, 1940]

Pratt: Ichneumonid^

169

The prescutum is the depressed area directly behind the meso-
scutellum and attached beneath it to the mesopostscntellum. It
merges into the large, convex plate composed of the fused meta-
scutum and metascutellum. This plate has usually been called
the “ postscutellum ” or “metanotum.” Crampton (1931), how-
ever, has shown that in the closely allied genus Megarhyssa the
large dorsal plate lying directly posterior to the prescutum is the
metascutellum while the narrow saddle-like hand extending over
the tergum between the metascutellum and the propodeum is the
metapostscutellum.

The metapostscutellum curves cephalad as a narrow band
forming the posterior margin of the hind wing base and at the
lower margin of the wing base fuses with another narrow sclerite.

This last plate, which lies between the wing base and the meta-
thoracic spiracle (sps), Crampton (1931) has shown to be com-
posed of the fused anepisternum and anepimeron of the meta-
thorax (aess and aemg). Below the spiracle the band widens out
into a broad, slightly convex plate, usually called the ''meta-
pleura,” but actually composed of the katepisternum and katepi-
meron of the meta thorax (kess and kerns).

The lateral longitudinal carina extending cephalad from the
metathoracic coxal cavity to a point just posterior to the meso-
thoracic coxal cavity seems to mark the line of division between
the metapleural and metasternal regions. On the metasternum
the line of attachment of the sternal apophyses is clearly
demarked.

THE WINGS

The system of wing venation proposed by Boss (1936) has been
used in this paper in preference to the artificial Cresson system
usually used by taxonomists, or the Comstock-Needham system
which is often figured in the older text-books. The system pro-
posed by Bradley (1931) is somewhat similar to that of Boss
(1936), but the latter seems to have worked out the origin of the
main veins, particularly in the posterior portion of the wing,
more carefully and on sounder grounds — i,e., axillary sclerites
instead of axillary furrows — and, for this reason, the Boss (1936)
interpretation is used in this paper. The discussion of the wings
follows the interpretation which Dr. Boss outlined to me in a

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Journal New York Entomological Society [Vol. xlviii

letter dated December 22, 1936, while the drawings of the wings
are exact copies of a figure which he labelled at the same time.

Fore-Wings

Costa (C) (Fig. 12) forms the thickened anterior, or front,
margin of the wing. It arises just distad of the basicostal plate
and extends along the anterior margin of the wing to the ptero-
stigma, from which it is separated by a distinct break, the costal
hinge.

Subcosta (Sc), present in the wings of primitive sawflies, has
frequently been considered as fused into a ‘ ‘ principal vein ’ ’ with
radius and media, but Bradley (1931) writes, “It is both sim-
pler and more probably truthful to consider that it has atrophied
entirely . . . therefore the marginal vein should be labelled C.”
Many writers have shown that Sci becomes atrophied at an early
stage in the phylogeny of the Hymenoptera, either in the Cim-
bicidas or in the Siricidae, while in the primitive Macroxyela
type Sc2 becomes separated from Ri and extends along the ante-
rior margin of the wing forming with Ri the cell Sc2 and Ri.
This cell has been retained in the intermediate forms becoming
gradually thickened and growing by fusion, so that in Arotes the
cell labeled Sc2 and Ri is actually the pterostigma, or “stigma.”
The break in the chitinization just before the pterostigma is
called the costal hinge. It marks not only the termination of
costa and the point at which Sc2 attains the costal margin of the
wing, but also indicates the point at which the radial sector turns
inward.

Badius (R) is the second longitudinal vein. It forms the poste-
rior margin of the very narrow costal cell, a cell so narrow that
in most family keys to the Hymenoptera it is regarded as absent
or “lost through coalescence,” in contradistinction to the much
wider costal cell of Aulacid^ and Gasteruptionidse. Basally
radius (R) (which is united with costa before the latter articu-
lates with the basicostal plate) articulates with the second axil-
lary sclerite. R forks but once, giving off an anterior branch Ri
(which marks the posterior margin of the stigma and continues
beyond the stigma albng the anterior margin of the wing, and a
posterior branch, the radial sector (Rs), which fuses with the

June, 1940]

Pratt: IcHNEUMONiDiE

171

first abscissa of Media just basad of the costal hinge. In all the
Chalastogastra the first abscissa of Rs + M (indicated in the figure
by the dotted line ) cuts across the cell R5 + 1st M and touches the
first medio-cubital cross-vein (1 m-cu) ; but this first abscissa of
Rs - M has been lost in all Ichneumonidse and only a spur, called
the ramellus, remains to prove the former existence of such a
vein, as in Cryptus and Exetastes. In Arotes this spur is very
tiny. Rs 4- M now extends antero-distad for a short distance and
then divides into Rs and M, with Rs forking at almost a right
angle just beyond 2 m-cu and then continuing in a gentle arc to
the apex of the wing where it joins Ri to form cell Ri.

The cross-vein arising from the middle of the stigma, and con-
necting it with the radial sector, is the second radial cross-vein
2r.

The areolet, present in the wings of most Ichneumonidse, is
formed with the first abscissa of radius by the second radio-
medial cross-vein.

Media (M), fused basally with cubitus, forms the third longi-
tudinal vein. At its origin, media forms part of the compound
vein complex just distad of the basicostal plate (bp), and is more
definitely associated with the median plate (m). Just distad of
the pteralia, media and cubitus separate from radius and proceed
postero-distad along the basal third of the wing. Here media
and cubitus fork, cubitus continuing to the anal angle, while
media turns abruptly toward the stigma and joins the radial
sector (Rs) a short distance from the costal hinge. (Note that
according to the Ross interpretation, media never actually attains
the base of the stigma, or the costal margin, although it does come
very close.) As previously explained, Rs + M cuts across the
cell Rg and 1st M in the primitive Hymenoptera, but this ab-
scissa of Rs + M has atrophied leaving only the tiny stump to
mark its former location. From here Rs + M proceeds antero-
apicad dividing abruptly into the radial sector (Rs), which
eventually attains the apex of the wing, and M which extends in
a gentle arc to the outer margin of the wing.

The first and second medio-cubital cross-veins connect the
medial and cubital fields, 1 m-cu marking the terminus of the
first abscissa of Rs + M and 2 m-cu marking the terminus of the
second abscissa of Rs + M.

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Journal New York Entomological Society [Vol. xlviii

Cuhitus (Cu), as explained previously, is associated with the
base of radius and media, so that actually none of the four prin-
cipal veins, namely costa, radius, media, and cubitus, arise inde-
pendently. Cubitus apparently articulates with the anterior-
distal corner of the median plate (m), and after proceeding for
the basal third of the wing as a fused vein with M, divide into
M and cu. Eoss, following Tilljmrd’s hypothesis of the loss of
Cu2 in the fossil Permoraphidia and Martynovia, and the re-
sultant carry-over in modern Holometabola, believes that Cu2
has been lost in Hymenoptera and that the apical division of Cu
should be called merely a division of Cui, with an anterior branch
Cuia and a posterior branch Cuib.

The first cubito-anal cross- vein (1 cu-a) lies between cubitus
and the first anal vein and connects the cubital and anal fields.
The position of 1 cu-a with reference to the first abscissa of M is
of considerable taxonomic importance, i.e., whether 1 cu-a
(called the nervulus by taxonomists) is opposite M (interstitial),
proximal to it (antef ureal), or distal to it (postfur cal).

Anal Veins. The homologies of the anal veins are indeed a dif-
ficult problem. According to Eoss (1936), there were originally
four anal veins in the Macroxyela type, all articulating with the
third axillary sclerite. In the lower Hymenoptera — Macroxyela
as figured by Eoss (1936) and Itycorsia (a pamphilid) as figured
by Snodgrass (1910) — the' bases of these anal veins are all dis-
tinct and separate. In the higher sawflies, however, a fusion of
the bases of these anal veins begins to develop, associated with
the dropping out of the second anal vein. In the Ichneumonidae,
as shown by Megarhyssa and Arotes, this fused base of the anal
veins becomes very large and triangular in shape as a result of
the fusion of the anals and only the first anal remains distinct.
Both Snodgrass (1910) and Eoss (in a personal letter) have
shown that in Ichneumonidse this is the first anal vein (lA).
The second anal vein has dropped out in the sawflies (Eoss, 1936)
while the third anal vein has been identified as the tiny stub in
the basal membrane arising from the middle of the fused base
of the anals. The fourth anal vein is present only in the mem-
bers of the Xyelidse, and even in them is represented by only a
tiny stump in the basal membrane.

June, 1940]

Pratt: Ichneumonid^

173

The first anal vein (lA) is the fourth longitudinal vein and
follows almost a straight course parallel to the inner margin of
the wing, ending some distance before the anal angle of the wing.

The anal fold, or furrow, lying anterior to the first anal vein
is indicated in the figure by a dotted line. It is clearly demarked
by a line of dark setffi, which contracts strongly with the hyaline
membrane at the base of the wings. As it extends to the anal
angle of the wing, this anal fold breaks the chitinization of the
veins 1 cu-a and Cuib, and the resulting membranous areas in
these veins are called hullce. There are three other bullag in the
fore-wing, one in the middle of the second abscissa of Rs + M, and
the two others about equidistant from each other on the second
medio-cubital vein (2 m-cu).

Hind Wings

Although at first glance the venation of the hind wing (Fig.
13) appears to be radically different from that of the fore-wing,
yet a closer examination shows that the scheme of modification
has been essentially similar in both wings.

Costa (C) is the slightly thickened, anterior margin of the
wing, extending at most only along the basal third of the wing.

Suhcosta (Sc) is completely fused with radius.

Radius (R) divides into Ri (which bends toward the anterior
margin of the wing, and attains it, at the apical third) and Rs,
which attains the outer margin of the wing just below the apex
of the wing.

The radio-medial cross-vein (r-m) connects the radial sector
(Rs) with the medial field (M).

The hamidi, a row of a dozen tiny hooks which fit into a pocket
in the fore wings and hold the two wings together in flight, are
borne on the basal part of Ri.

Media (M) separates from radius near its base, fuses with Cui
for a considerable distance, then separates from it and proceeds
to the margin of the wing. It is connected to Rs by r-m, but the
medio-cubital cross-veins are lacking.

Cubitus (Cu) is represented only by Cui, which does not di-
vide into Cuia and Cuib, as in the fore wing, but continues to the
margin of the wing simply as Cui. It is fused to Media to about

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Journal New York Entomological Society [Vol. xlviii

the middle of the wing and then separates sharply at a right
angle, until it touches the cnbito-anal cross-vein (cu-a), at which
point it again becomes a longitudinal vein which parallels M to
the outer margin of the wing.

Cui and cu-a make up the very important structure known to
taxonomists as the nervellus. In using this important structure
for the separation of genera and species, taxonomists have three
alternatives: it may be ‘‘broken above the middle,” i.e., first ab-
scissa of Cui shorter than cu-a ; it may be ‘ ‘ broken at about the
middle,” i.e., first abscissa of Cui about equal to cu-a; or it may
be “broken below the middle,” i.e., first abscissa of Cui longer
than cu-a.

In Arotes taxonomists describe the nervellus as “broken at
about the middle, and reclivous. ” This last term was first de-
fined by Cushman and Rohwer (1920) with two other alterna-
tives, as follows:

“A 'perpendicular nervellus is one in which the anterior end
is opposite the posterior end, that is, one in which a line drawn
touching both the anterior and posterior ends is at right angles
to the longitudinal axis of the submediallan cell (Cell Kg).
Other authors have called this a continuous or interstitial ner-
vellus.

“An inclivous nervellus is one in which the anterior end is
nearer the base of the wing than is the posterior end. This has
been spoken of as a postfurcal nervellus.

“A reclivous nervellus is one in which the posterior end is
nearer the base of the wing than is the anterior end. This has
heretofore been spoken of as a postfurcal nervellus.”

The anal furrow is indicated in the figure by a dotted line.
Behind it lies a distinct longitudinal vein arising from the third
axillary (3 Ax). This is the first anal vein (1 A).

Behind this first anal vein, extending into the first anal cell is
a tiny spur of the third anal vein (3A).

Pteralia of the Fore- Wing

Each wing is attached to the thorax by a membranous basal
area containing several tiny sclerites which Snodgrass (1935) has
called the pteralia. These include in Hymenoptera an anterior
hasicostal plate (which Snodgrass calls the humeral plate), a

June, 1940]

Pratt: Ichneumonid^

175

group of four axillary sclerites, and a single median plate (which
would seem to be the outer median plate, labeled m' by Snod-
grass (1935, p. 219)).

In the fore wing (Fig. 11) the hasicostal plate (bp) lies be-
tween the anterior notal process and the base of costa. It is a
large plate resembling a second tegula somewhat and has a deep
suture crossing it diagonally so that there appear to be two
basicostal plates.

The first axillary sclerite (1 ax) is a peculiar twisted sclerite
which articulates on its inner margin with the thorax and on its
outer margin with the axillary sclerites and the base of the prin-
cipal veins. Its inner margin articulates with the anterior notal
process and with the tergal margin. On its outer margin, its
anterior process forms a fulcral point for the basicostal plate
and the second axillary, and on its under side it apparently also
forms a fulcral point for a portion of the enlarged base of costa,
radius, media, and cubitus.

The second axillary sclerite (2 ax) is hinged obliquely to the
outer margin of the first axillary. Anteriorly it articulates with
the base of radius and media (and through these with costa and
cubitus) and with the median plate. Posteriorly, the second
axillary articulates with the median projection of the third
axillary.

The third axillary sclerite (3 ax) lies in the posterior part of
the articular membrane just before the axillary cord. Ante-
riorly it articulates with the enlarged, triangular base of the
first and third anal veins, mesally with the posterior end of the
second axillary, and posteriorly with the fourth axillary.

The fourth axillary sclerite (4 ax) is a small sclerite lying be-
tween the third axillary and the posterior notal process.

The median plate (m) is probably the distal median plate
labelled m' by Snodgrass (1935, p. 219). It is a triangular plate
lying in the median area of the articular membrane. By its basal
corner it articulates with the second axillary; but its anterior-
distal corner, with the base* of Cu; and by its posterior-distal
corner, with the large, triangular base of the first and third anal
veins.

Pteralia of the Hind Wing

The pteralia of the hind wings (Fig. 10) are similar to those

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Journal New York Entomological Society [Vol. XLViii

of the fore-wings. There is apparently no tegula in the hind
wing and the base of the wing continues simply the articular
membrane.

The hasicostal plate (bp) articulates by its apex with the com-
bined bases of costa, radius, media, and cubitus.

The first axillary sclerite (1 ax) is a V-shaped sclerite which
articulates with the basicostal plate by its anterior process and
with the second axillary by its posterior arm.

The second axillary sclerite (2 ax) in addition to its articula-
tion with the first axillary, articulates by its anterior corner with
the fused base of radius and media.

The third axillary sclerite (3 ax) articulates anteriorly with
the base of the anal vein, mesally with the second axillary and
posteriorly with the posterior notal process. There is no fourth
axillary in the hind wing, as shown by Salman (1928) and
Snodgrass (1910). Posterior to the third axillary the membrane
is thickened to form the axillary cord.

The median plate (m) articulates posteriorly with the fused
base of media and cubitus, and more particularly with the
latter, and anteriorly with the base of the first anal vein.

Cells of the Wings

The names of the various cells have been figured by Cresson
(1887), Comstock (1930), Bradley (1931), and Boss (1936). A
comparison of the various systems as they apply to Arotes is

given below.

Cells of the Fore Wing

Cresson

Comstock-

Needham

Bradley

Eoss

Pterostigma

Pt

SCa "t" El

SCa

Marginal, or Eadial

2nd El - Eg

2nd El + Eg

2nd El

Cubito-discoidal

M^

1st Ml + 1st El

1st El + 1st M

Median

M

M

E

Third Submarginal

Rs

1st + 2nd + 3rd Eg

R5

First Apical

2nd Mo

2nd Ml

CUia

Second Apical

M,

.2nd Ma

3rd M

Second Discoidal

Cui

Cui

Cuib

Third Discoidal

1st Ma

1st Ma

2nd M

Submedian

Cu

Cu + 1st A

Cui

Anal

A

2nd A

1 A

June, 1940]

Pratt: Ichneumonid^

177

Cells of the Hind Wing

Costal

C

C

C

Median

M

M

R

Radial

Ri 4" R2

El

Ri

Cubital or submarginal

R5

R5

R5

Submedian

M4 + 1st M2 + Ml

M + Cu + Cui

1st Cui

First Discoidal

Ml

M2

M

Second Discoidal

M2

Ml

2nd Cui

Anal

1st A

2nd A

1st A

THE LEGS

The legs are rather long and slender in proportion to the size
of the body. The front pair is the smallest while the hind pair
is the longest and largest, the hind femora of the male reaching
to, or almost to, the tip of the abdomen.

Each leg (Fig. 8) is made up of a rather large coxa (cx), a
distinct two-segmented trochanter (tr), a rather short and almost
cylindrical femur (fe), an elongate tihia (ti), a five-segmented
tarsus (ta), and a pretarsus (ptar) bearing a pair of claws.

In the hind legs the coxae are almost pear-shaped. On the
large basal end, each coxa is deeply incised on its postero-laterad
surface by a hasicoxal suture. The narrow band of chitin thus
demarked expands above this suture into a semi-globular hasi-
coxite which fits into the metathoracic coxal cavity, so that the
articulation strongly resembles a ball and socket joint. The coxo-
trochanteral articulation is dicondylic, for the distal end of the
coxa is emarginate on either side and into these emarginations
fit peg-like projections of the trochanters, sometimes known as
trochanter-condyles acting as pivotal points. Posteriorly a leva-
tor muscle is attached to the trochanter ; anteriorly the depressor
muscle is attached.

The trochanter (tr) is almost as long as the coxa but is more
slender and almost cylindrical. As in all Ichneumonidse, the
trochanters are two-segmented. Cushman (1929) calls the
apical segment the trochantella, which is a very appropriate term
since in this genus and in most other Ichneumonidse the second
segment is the smaller of the two segments. Snodgrass (1935),
by a study of the muscle attachments, has shown this two-seg-
mented trochanter is formed not by a division of the trochanters
(as in Odonata) but by a basal subdivision of the femur; and
Crampton (1925) thinks it is preferable to call this segment the

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Journal New York Entomological Society [Vol. xlviii

hasifemur “rather than by the terms which imply that it is a
part of the trochanter.” The two segments are joined to each
other by an articular membrane.

The femur (fe) is a strong, robust segment attached by articu-
lar membrane to the trochanter along a trochantero-femoral
hinge. It is perhaps the most important leg segment from a
taxonomic viewpoint because the coloration of this segment offers
primary characters for the separation of the eight species of
Arotes occurring in North America. The femora-tibial articu-
lation is similar to that between the coxa and trochanter. The
pivotal points of the femur, sometimes known as the tihiafers,
are found on the anterior-lateral portions of the apical margin.
Basad of this articulation, the proximal end of the tibia is pro-
duced into a distinct head, called the tibiacaput (Crampton,
1923), which is received in a distal pocket of the femur or the
“femur cup.” This device allows the tibia to be indexed close
to the under side of the femur.

The tibia (ti) is the longest and slenderest segment of the leg.
It is narrower at the base and gradually expands toward the
tip where it is of about the same diameter as the trochanter. On
the apical margin of the tibia are two immovable spines about as
long as the third tarsal segment. The mid tibia is similar to the
hind one, both bearing two immovable, apical spines usually
known as tibial spurs. The fore tibia, however, is relatively
broader at the apex and bears on its apex one movable spine
which is modified to form a part of the antennal cleaner. This
spine is drawn between it and the modified inner basal surface of
the basitarsus. The apex of the tibia is hollowed out to form
a tibiatJieca, or cup of the tibia, which receives the head of the
basitarsus; so that the tibio-tarsal joint is in the nature of a
ball and socket joint.

The tarsus (ta) is composed of five segments, of which the
basal one, known as the basitarsus (formerly metatarsus) , is as
long as the next three segments together. The tarsal segments
each expand slightly from base to apex. They are clothed with
a fine short pubescence, but I have seen no trace of tarsal pulvilli
or euplantulce. The fourth tarsal segment (ta4), which is the
shortest segment of the tarsus, is half as long as the apical tarsal
segment. The subsegments of the tarsus are freely movable on

June, 1940]

Pratt: Ichneumonid^

179

one another by inflected connecting membranes, but apparently
as Snodgrass (1935) states, ‘‘the tarsus never has intrinsic
muscles. ’ ’

The fifth tarsal segment, or distitarsus, bears the pretarsus
which is composed of two lateral claws and a complex series of
structures between them.

The dorsal surface of the distitarsus (Fig. 14) bears on its
apical margin a triangular-shaped sclerite, the unguifer (uf) to
which the claws, or ungues (un), are attached.

On the ventral surface of the distitarsus, and partially con-
cealed by it, lies a median plate, the unguitractor (ut). On its
base it bears an apodeme (ap) to which is attached the depressor
muscles of the pretarsus, often called the retractor of the claws.

The claws, or ungues (um), which are attached by flexor mem-
brane to the unguifer and unguitractor, are large and curved.
Each claw is noticeably cleft at the apex and typically bears two
or three long spines on its base just laterad of the flexor mem-
brane. The plant a (pi) is a rather broad sclerite attached
distally to the unguitractor plate. It is rather easily seen because
of the three or four setae it bears.

The orhicida (or), which is apparently attached to the
unguifer, lies above the planta on the dorsal surface, and the
girdle-like camera lies between the two.

The camera (cm) is a heavily sclerotized band which by its
contractions and expansions changes the arolium (ar) from a
small, membranous pad into a large, bilobed structure.

THE ABDOMEN

The propodeum (Pig. 3), long, erroneously, called the “meta-
notum” by taxonomists, is morphologically the tergite of the
first abdominal segment. It is derived entirely from the tergum,
the pleural and sternal plates having become atrophied as ex-
plained by Crampton (1931). Its hind margin broadens out
into a semicircular band forming the dorsal half of the aperture
through which the petiole of the abdomen projects and articulates
with the thoracic region.

The propodeal spiracles are situated on the anterior lateral
margin of the propodeum. These are long, oval, or slit-like
spiracles, as contrasted with the short oval, or circular spiracles

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Journal New York Entomological Society [Vol. xlviii

of Coleocentrus, the other genus in the Acoenitini. The spiracles
are about twice their own length from the base of the propodeum,
and are connected with the base by carinae.

Various systems of naming the areas of the propodeum have
been proposed from time to time. The systems introduced by
Davis (1897) and by Morley (1903) seem to be most used by
modern taxonomists, and for this reason are used in this paper.

1. Basal, or first median area (basareola)

2. Superior, or second median area (areola)

3. Apical, petiolar, or third median area (petiolarea)

4. External, or first lateral area

5. Area dentipara, external median, or second lateral area

6. Internal, middle apical, or third lateral area

7. Spiracular, or first pleural area

8. Lateral, middle pleural, or second pleural area

Snodgrass (1935) divides the abdomen proper into three

groups : The pregenital , or visceral segments which, in Arotes,
comprise the second to seventh segments; the genital segments,
composed of the eighth and ninth segments in the female, and the
ninth in the male ; and the postgenital segments.

The abdomen in both sexes is slightly petiolate at the base.
In the male (Fig. 17), the abdomen is more or less cylindrical
throughout, expanding but slightly from base to apex, and is only
slightly compressed apically. The abdomen of the female (Fig.
15) is cylindrical at the base, but expands medially, and becomes
compressed from side to side on its apical third, in a manner
suggestive of the Ophioninse. The most remarkable feature of
the female abdomen (and the structure by which the female
Acoenitini may be at once separated from all other Ichneu-
monidse) is the very large vomeriform hypopygium which is
heavily sclerotized and acute at the apex.

The abdomen consists of nine distinct segments, exclusive of
the propodeum, which is the true first tergite, and it is possible
that the membranous area around the anus may well represent
the eleventh tergite, in which case the abdomen proper is com-
posed of ten segments.

The connection between the abdomen proper and the pro-
podeum is made by the insertion of the basal end of the petiole,
or true second abdominal segment, into the aperture formed by

June, 1940]

Pratt: IcHNEUMONiDiE

181

the propodeum and the metasternum. The petiole apparently
articulates with the propodeal part of this aperture by lateral
projections of the base of the petiole. On the base of the petiole
lies a median, dorsal ligament, called the funicle, which is at-
tached anteriorly to the endoskeleton of the metathorax by
muscles, and acts as a levator of the abdomen.

The petiole, or second abdominal segment (2t), according to
Crampton (1931) is composed entirely of the second tergite which
has extended ventrally and fused in the anterior region, crowd-
ing the second sternite (2s) backward in the process. The
petiole is about three times as long as it is wide at the apex, and
expands from base to apex. The oval spiracles lie in the middle
of the petiole.

Female Abdomen

In the female (Fig. 15), the tergite is bent at its basal third,
and is produced into a small protuberance somewhat similar to
the large basal protuberance in the petiole of such Mutillidae as
Dasymutilla. The second pleurite is a narrow, longitudinal band
of membrane originating beneath the spiracle, and extending
caudad to the third pleurite. It is demarked by a fold from the
membranous second sternite.

The third to fifth abdominal segments are similar in general
form and structure, each consisting of a tergite, sternite, and a
pair of sclerites of uncertain affinities, which in this paper are
called the pleurites. These three segments increase noticeably in
size and sclerotization as they extend caudad. The tergites grow
successively shorter, wider, and more convex; the pleurites be-
come more strongly sclerotized; and the sternites show larger
sclerotized areas in the sternal membrane.

In the sixth segment, the tergite (6t) fuses with the pleurite.
The spiracle of the sixth segment, however, is just as far from
the^ latero-pleural line as are the spiracles of the three preceding
segments (which fact may indicate either that the pleurites are
of recent origin, or more probably that the fusion of the sixth
tergite with the pleurite has taken place quite recently), whereas
the spiracles of the seventh, and eighth segments are quite near
the latero-pleural line. The sixth sternite is almost completely
sclerotized.

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Journal New York Entomological Society [Vol. xlviii

In the seventh abdominal segment, the tergite is similar to that
of the sixth. The seventh segment is the largest segment of the
abdomen, and combines the two features by which the females
of this tribe are easily told from all others : the abdomen begins
being compressed from side to side with the seventh tergite, and
the seventh sternite (7s) is enormously elongated, very heavily
sclerotized, and acute at the apex, producing what taxonomists
call ‘Hhe large, vomeriform hypopyginm characteristic of the
tribe Acoenitini. ” It is about one-third as long as the whole
abdomen, or about as long as the three preceding segments to-
gether. This plate is concave, so that the ventral portions of the
genitalia fit into it, and are concealed and protected by it. For
this reason, it is often called the suhgenital plate or hypopyginm.

Female Genitalia

The genital segments of the female (Fig. 16) are composed of
the eighth and ninth tergites and their appendages, for in Arotes,
as in all Hymenoptera, there are no pleural or sternal plates on
these segments in the females. The eighth tergite (8t) is slightly
smaller than the seventh and its spiracle lies on the anterior
edge of a distinct fovea. The ninth tergite (9t) is an elongate
plate forming the fulcral point for both the ovipositor and the
valves of the ovipositor. It has no spiracle.

The basal part of the ovipositor consists of the first and second
valvifers (1 vf and 2 vf), which lie beneath the ventral margin
of the ninth tergite. The first valvifer articulates by its dorsal
angle with the ninth tergite, and by its posterior, ventral angle
with the second valvifer. Anteriorly, the first valvifer has the
form of a very thin, transparent plate, which is continuous with
the ramus of the first valvulge.

The second valvifer (2 vf) is a narrow, elongate plate, which
lies half hidden beneath the ninth tergite. Anteriorly, it widens
into a large, rounded process articulating with the first valvifer
so that there is actually no articulation with the ninth tergite
as in most typical insects; posteriorly, it broadens very slightly
and becomes modified to form the long, slender third valvulce
(3v). It is continuous with the ramus of the second valvulse
along its extreme anterior margin, and its antero-dorsal margin

June, 1940]

Pratt: Ichneumonid^

183

is produced into a long, pointed apodeme for the attachment of
muscles.

The shaft of the ovipositor is elongate and compressed with
an acute apex. The ovipositor itself is composed of the first and
second valvuloe (1 v and 2 v) which form respectively the ventral
and dorsal parts of the ovipositor. The apical portion of the
ovipositor is usually enclosed between the concave third valvulce
(3v) which are often called the valves or sheaths of the ovipositor.

Postgenital Region

This is a membranous, apical area surrounding the anus. At
the base there are weakly sclerotized dorsal and ventral sclerites
which may represent the tenth tergite and sternite (lOt and 10s)
(Pigs. 15, 16, 17). The cerci (ce) are borne in the membranous
area between them and the anus at the extreme apex.

Male Genitalia

In Arotes the copulatory ossicles are located on the ventral
side of the genitalia, and studies of the more primitive Macro-
xyela and certain Cephid genera would indicate that this is the
more primitive, orthandrous condition, and that the torsion of
the parts that occurs in some sawflies in the family Tenthredinidag
is a secondary development characteristic of the Tenthredinid
complex called the Strophandria by Crampton (1919) and by
Ross (1937).

The basal ring (hr) (Pigs. 18, 19) is a narrow, basal band en-
closing the rest of the genitalia. According to Snodgrass (1935,
p. 605), “it is said to differentiate as a circular fold of the wall
of the genital chamber.”

The genital claspers, or parameres, were first homologized with
the parameres of Coleoptera by Wheeler in 1910, and his views
on the subject have been accepted by Pruthi (1924), Balfour-
Brown (1932), Snodgrass (1935), and Crampton (1936). The
parameres (pr) arise on the ventral side of the basal band or ring
and expand laterally into wide basal plates, which narrow again
on the dorsal side meeting along a dorso-median caudal groove.
Apically the parameres are lengthened into blunt, tooth-like pro-
jections which serve somewhat similar functions as the distal

184

Journal New York Entomological Society [Vol. xlviii

segments of the parameres in lower saw-flies. These tooth-like
projections, however, are probably not homologous with the distal
segments of the parameres (which Crampton, 1919, called the
cochlearia in the lower saw-flies) since these distal segments seem
to drop out in the phylogeny of the Hymenoptera at about the
Cephid level.

The copulatory ossicles are composed of a large basal portion
known as the volsella (vol) and a smaller, slightly cylindrical
apical segment called the sagitta (sag), which is connected to
the volsella by membrane and is the true copulatory ossicle.
They arise mesally on the parameres and project posteriorly, one
lying on either side of the sedeagus. It should be emphasized
especially, however, that in Arotes and also in Coleocentrus,
which two genera comprise in America the tribe Acoenitini of
Cushman and Rohwer, there is no distivolsella. Therefore, the
inner forceps structure is lacking, although it is present in all
the other Ichneumonidas known to the present writer. The
absence of the distivolsella in these two genera would seem,
therefore, to be a character of tribal value in separating the
Acoenitini from the other tribes of Ichneumonidae, and it is pos-
sible that this character may be correlated with the huge
development of the plow-shaped, or vomeriform, hypopygium in
the female.

The central structure of the male genitalia is the cedeagus,
which is composed of a pair of penis valves (ae). At the base of
the asdeagus, on the dorsal side are a pair of parapenes (pap)
which help to strengthen the pallic structure.

Postgenital Segments

The postgenital segments (Fig. 17) are membranous but do
include small sclerotized areas which would seem to be the tenth
and eleventh tergites (lOt and lit) and the corresponding
sternite. Snodgrass (1935) and other writers would call the
small cercus-like appendages of the postgenital segments socii and
would homologize them with similar structures in Trichoptera
and Lepidoptera, but it is more reasonable to compare these ap-
pendages with those of male Mecoptera (since the genitalia of
Hymenoptera are closest to those of Mecoptera) and to call them

June, 1940]

Pratt: Ichneumonid^

185

true cerci as Crampton (1936) has done. The anus is situated
at the very apex of the abdomen in both sexes.

LITERATUKE CITED

Balfour-Browne, F.

1932. A Textbook of practical entomology. E. Arnold and Co., VIII +
191 pp., 161 figs.

Bradley, J. C.

1931. Laboratory Guide to the study of the wings of insects. Daw,
Illston and Co., Ithaca, N. Y.

Comstock, J. H.

1918. The Wings of Insects. Comstock Publishing Co., Ithaca, N. Y.
1930. An introduction to entomology. Comstock Publishing Co., Ithaca,

N. Y.

Crampton, G. C.

1918. The genitalia and terminal abdominal structures and genitalia

of male Neuroptera, and Mecoptera, with notes on the
Psocidae, Diptera, and Trichoptera. Psyche S5: 47-59, No.
3 ; Pis. II, III.

1919. The genitalia and terminal abdominal structures of males, and the

terminal abdominal structures of the larvae of Chalasto-
gastrous Hymenoptera. Proc. Ent. Soc. Wash., SI: 129-151;
Pis. 9-12.

1920. A comparison of the genitalia of male Hymenoptera, Mecoptera,

Diptera, Trichoptera, Lepidoptera, Homoptera, and Strepsip-
tera, with those of lower insects. Psyche S8 : 34-44 ; PI. IV.

1921. The sclerites of the head and the mouthparts of certain immature

and adult insects. Ann. Ent. Soc. Amer., 14 : 65-103 ; Pis.
2-8.

1923a. Preliminary notes on the terminology applied to the parts of an
insect’s leg. Can. Ent., 65: 126-132; PI. 3.

1923b. A phylogenetic comparison of the maxillae throughout the orders
of insects. Journ. N. Y. Ent. Soc., 31: 77-106, Pis. 12-17.
1923c. A comparison of the labium in certain Holometabolous insects
from the standpoint of phylogeny. Proc. Ent. Soc. Wash.,
S5: 171-180; PI. 15.

1925. A phylogenetic study of the labium of Holometabolous insects,

with particular reference to the Diptera. Proc. Ent. Soc.
Wash., S7: 63-91; Pis. 6-8.

1926. A comparison of the neck and prothoracic sclerites throughout the

orders of insects from the standpoint of phylogeny. Trans.
Amer. Ent. Soc., 5S: 199-243; Pis. 10-17.

1928. The eulabiuiji, nientum, submentum, and gular region in insects.

Pomona Journ. Ent. and Zoo., SO: 1-18; Pis. 3.

1929. The terminal abdominal structures of female insects compared

throughout the orders from the standpoint of phylogeny.
Journ. N. Y. Ent. Soc., 37: 453-496; Pis. 9-16.

186

Journal New York Entomological Society [Vol. XLVIII

1931. A phylogenetic study of the posterior metathoracic and basal

abdominal structures of insects, with particular reference to
the Holometabola. Journ. N. Y. Ent. Soc., 39: 323-357;
Pis. 20-24.

1932. A phylogenetic study of the head capsule in certain Orthopteroid,

Psocoid, Hemipteroid, and Holometabolous insects. Bull.
Brook. Ent. Soc., S7 : 19-55 ; Pis. 4-8.

1936. Suggestions for a new interpretation of the postabdomen in male
cyclorraphous Diptera. Bull. Brook. Ent. Soc., 31 : 141-148 ;
PI. VII.

Cresson, E. T.

1887. Synopsis of the families and genera of Hymenoptera of America
north of Mexico. Trans. Amer. Ent. Soc., Supplementary
volume for 1887.

Cushman, E. A.

1929. A revision of the North American Ichneumon-flies of the genus
Mesostenus and related genera. Proc. U. S. Nat. Mus., 74:
1-58 ; 8 flgs.

Cushman, E. A. and Eohwer, S. A.

1920a. The North American Ichneumon-flies of the tribe Acoenitini.
Proc. U. S. Nat. Mus., 57 : 503-523.

1920b. Holarctic tribes of the Ichneumon-flies of the subfamily lehneu-
moninae (PimplinaB). Proc. U. S. Nat. Mus., 57: 379-396.

Davis, G. C.

1897. A review of the Tryphoninae and descriptions of new Ichneu-
monidae. Trans. Amer. Ent. Soc., 24: 193-372.

Franklin, H. J.

1912. The Bombidae of the New World. Trans. Amer. Ent. Soc., 38:
178-486.

Holway, E. T.

1935. Preliminary note on the structure of the pretarsus and its possible
phylogenetic significance. Psyche 42: 1-20; Pis. 1-3.

Hooker, C. W.

1912. The Ichneumon-flies of America belonging to the tribe Ophionini.

Trans. Amer. Ent. Soc., 38: 1-177 ; 3 pis.

MacGillivray, a. D.

1906. A study of the wings of Tenthredinoidea. Proc. U. S. Nat. Mus.,
29 : 569-653 ; figs. 49 pis.

Merrill, J. H.

1913. On some genera of the Pimpline Ichneumonidae. Trans. Amer.

Ent. Soc., 41 : 109-154.

Morley, Claude

1913. The fauna of British India including Ceylon and Burma.

Hymenoptera III, Ichneumonidae I. Taylor and Francis,
London.

June, 1940]

Pratt: Ichneumonid^

187

Peck, Oswald

1937. The male genitalia in Hymenoptera especially the family lehneu-
monidae. Can. Journ. Eesearch 15: 221-274; Pis. 5-10.

Pruthi, H. S.

1924. On the post-embryonic development and homologies of the male
genitalia of Tenebrio moUtor L. Proc. Zool. Soc. London:
857-868, 3 pis., 3 figs.

Eoss, H. H.

1936. The ancestry and wing venation of the Hymenoptera. Ann. Ent.
Soc. Amer., £9 : 99-109 ; Pis. 1-2.

Salman, K. A. .r-

1929. The external morphology of Pepsis elegans Lepeletier. Trans.
Amer. Ent. Soc., 55: 119-153; Pis. 6-9.

Snodgrass, E. E.

1910. The thorax of the Hymenoptera. Proc. U. S. Nat. Mus., 39: 37-
91; Pis. 1-16.

1935. Principles of insect morphology, first edition. McGraw-Hill Book
Co., New York City.

ViERECK, H. L.

1916. The Hymenoptera of Connecticut. Bull. 22 State Geo. and Nat.
Hist. Survey, pp. 1-825; figs., 10 pis.

Wheeler, W. M.

1910. Ants, their structure, development, and behavior. Columbia Uni-
versity Press, New York. XXV -t- 663 pp.

LIST OF ABBEEYIATIONS

ae

— aedeagus

0

— ocellus

aem

— anepimeron

oc

— occiput

aes

— anepisternum

or

— orbicula

af

— ^antefrons

osc

— ocular sclerite

ap

— apodeme

ar

— arolium

P

— pleurite

as

— antennal sclerite

pa

— parietals

at

— anterior tentorial pits

paf

— parafrontals

ax

— axillary sclerite

pap

— parapenes

pc

— postocciput

bp

— basalar plate

pcs

— postoccipital suture

br

— basal ring

pctc

— prepectal carina

C

— Costa

pdc

— pedicel

cd

— cardo

Pf

— postfrons

ce

— cercus

Pg

— postgena

cl

— clypeus

Pgl

— paraglossa

cm

— camera

pi

— planta

Cu

— Cubitus

plst

— pleurostoma

Cx

— coxa

ppct

— prepectus

pr

— paramere

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Journal New York Entomological Society [Vol. xlviii

epm

— epimeron

print

— prementum

eps

— opisternum

psc

— prescutum

pscl

— postscutellum

fe

— femur

ptar

— pretarsus

fl

— flagellum

E

— Eadius

ga

— galea

Es

— radial sector

ge

— gena

gl

— glossa

s

— sternite

sag

— sagitta

hp

— hypostoma

Sc

— Subcosta

hpc

— hypostomal carina

sc

— scutum

scl

— scutellum

kem

— katepimeron

scp

— scape

kes

— katepisternum

smt

— submentum

SOS

— subocular suture

la

— lacinia

sp

— spiracle

laplp

— labial palpus

st

— stipes

lb

— labium

Ic

— lacinia

t

— tergite

Im

— labrum

ta

— tarsus

ti

— tibia

M

— Media

tr

— trochanter

m

— median plate

m-cu

— medio-cubital cross-vein

uf

— unguifer

md

— mandible

un

— ungues

mxplp — maxillary palpus

ut

— unguitractor

n

— notum

V

— valvula

vf

— valvifer

vol

— volsella

Plate V

Figure

1.

Dorsal view of the head.

Figure

2.

Ventral view of the head.

Figure

3.

Dorsal view of the propodeum.

Figure

4.

Antenna.

Figure

5.

Labium.

Figure

6.

Maxilla.

Figure

7.

Mandible.

Figure

8.

Fore Leg.

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

(Plate V)

7

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Journal New York Entomological Society

[VoL. XLVIII

Figure 9.
Figure 10.
Figure 11.
Figure 12.
Figure 13.
Figure 14.

Plate VI

Lateral view of the pro-, meso-, and metathoraces.
Pteralia of the hind wing.

Pteralia of the fore wing.

Fore wing.

Hind wing.

Pretarsus of the hind leg.

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

(Plate VI)

192

Journal New York Entomological Society [Vol. xlviii

Plate VII

Figure 15. Lateral view of the female abdomen (ovipositor and sheaths
cuts off at tip of the abdomen).

Figure 16. Lateral view of the female genitalia.

Figure 17. Lateral view of the male abdomen.

Figure 18. Dorsal view of the male genitalia.

Figure 19. Ventral view of the male genitalia.

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

(Plate VII)

June, 1940]

Weiss: Insect Losses

195

MONEY LOSSES DUE TO DESTRUCTIVE INSECTS

By Harry B. Weiss

It is customary, from time to time, for entomologists to call
attention, in text books, special articles, newspapers, etc., to the
enormous money losses due to destructive insects. These figures
sometimes reach dizzy heights and carry the conviction that cold
type seems to have for many readers. Without meaning to lessen
the importance of destructive insects in their ability to create real
crop losses of considerable magnitude, we would like, in this little
article, to cast a few doubts upon the accuracy of the method in
use at present whereby these losses are converted into dollars.

In arriving at the percentage of loss by insects, it is customary
to obtain estimates indicating to what extent a particular insect
has reduced the normal production of a crop to the production
actually harvested. Having arrived at a figure indicating a loss
in bushels, the value of this lost portion is obtained by basing it
on the prevailing average farm price of the crop actually har-
vested, disregarding the reduction in value which usually follows
the marketing of a larger crop. A few authors of papers on the
subject of crop losses due to insects admit this error in their
figures, but hold to them because of the absence of a better method
and because they believe that the enhanced value given to the
destroyed portion is offset by other losses chargeable to insects,
such as the cost of control, which they have not included.

A quite recent publication cites the potato crop of the United
States for 1936 as having been damaged to the extent of 15 per
cent by insects. The total actual production of potatoes for that
year was estimated at approximately 330,000,000 bushels. Be-
cause the supply was below the average that year the farm price
went up and was about $1.13 per bushel. Now, if the total pro-
duction for that year had been 388,000,000 bushels, i.e., the actual
production plus 58,000,000 bushels, the estimated loss due to
insects, the farm price would, in all probability, have gone down
to around 60 cents per bushel, as it was in 1935 or 1937 or 1938,
when production stood in proximity to 388,000,000 bushels. The

196

Journal New York Entomological Society [Vol. XLVIII

farm value in 1936 reached a figure of about $370,000,000. If
there had been no insect injury and if the production had been
388,000,000 bushels, the farm value at the lower price per bushel,
provided the production could have been absorbed by the market,
would have been about $233,000,000, which is $137,000,000 less
than the farm value of the crop that was reduced by insects and
actually produced.

To mention another example, apple production in the United
States was supposed to have suffered a loss of 20 per cent in 1936
due to insects. Production for 1936 was estimated as 117,506,000
bushels, or 80 per cent of what the production should have been
had there been no insect injury. The farm price for 1936 is
quoted as $1.05 per bushel. If the insect damage had been elimi-
nated, the total production would have reached 146,883,000 bush-
els. Taking into account the purchasing power of the population
since 1933, if 146,883,000 bushels had been thrown on the market,
the price, in all likelihood, would not have gone higher than 75
cents per bushel, and even this is a generous allowance. The
estimated farm value of the apple crop in 1936 was about
$123,381,000. With insect injury eliminated, resulting in a crop
of 146,883,000 bushels, the total value at 75 cents per bushel would
have been $110,162,000, or more than $13,000,000 in favor of the
status quo. Of course having a crop of 146,883,000 bushels, more
labor would be required to harvest, grade, pack and deliver it,
provided the bottom did not drop out of the apple market com-
pletely. With the complete elimination of insect damage, a state
of affairs not likely to happen, there would be some violently
painful and long adjustments in the economics of the apple indus-
try. A smaller number of trees would suffice, less labor would
be employed and there would be changes all along the line. We
have no desire to forecast what would happen to the farm and
retail prices, or to the growers, etc., but if insect damage to crops
were eliminated or greatly curtailed beyond the present amount,
during the adjustment period surpluses would arise to plague us.
But of course these could be handed over to some ‘‘Agricultural
Surplus Commission” to worry about and would be of no concern
to entomologists. If the large surpluses could be sold to countries
where there were shortages, this would solve the problem, but in

June, 1940]

Weiss: Insect Losses

197

the case of perishable crops, the surpluses might easily be calami-
tous. In the case of commodities that could be stored, the situa-
tion- would be better, but even stored commodities in large
amounts have a depressing effect upon market prices.

The percentages of damage, to various crops in the United
States as a whole, by insects, as given in our text books, seem, for
the most part, to run in multiples of 5, such as 5, 10, 15 and 20.
Estimates seldom are less than 5 per cent or more than 20 per
cent, and these are said to be conservative. To us, these estimates,
for the most part, appear rather high for the country at large.
They probably represent the opinions of a comparatively small
number of technical men rather than the observations of numer-
ous producers. The difficulty of obtaining reliable estimates
from either source is fully appreciated. Numerous factors con-
trol production and the hazards of farming include not only
insects, but plant diseases, defective seed, deficient moisture, ex-
cessive moisture, frost, hail, hot winds, storms, etc. Of the fac-
tors reducing normal production to the production actually
harvested, adverse climate is the most important, and for the most
part the effects cannot be avoided. Factors such as insects, plant
diseases, poor seed, etc., may be overcome to a certain extent and
the losses, due to them, reduced.

If the yield of a certain crop is 10 per cent less than the so-called
normal yield, how is one going to apportion this loss to insects,
plant diseases, deficient moisture, etc., etc., etc., on the basis of
our present knowledge, with any degree of accuracy? With
stored products, of course, where only one factor is at work reduc-
ing the volume, the operation is not difficult. And there are
other instances where insect damage is apparent and separable
from injury by other causes and where it can be estimated or
arrived at fairly accurately. In the case of a growing crop being
injured by one species of insect, perhaps the loss in yield due to
the insect can be estimated with some degree of accuracy, but
even in this instance, there may be other factors tending to reduce
the yield, including adverse climate, plant diseases, etc., and the
assignment of loss due to each would be difficult. In the case of a
growing crop being injured by several species of insects, the case
becomes more difficult, in fact, the difficulties increase as the num-

198

Journal New York Entomological Society [Vol. xlviii

ber of factors increases. Experimental work designed to isolate,
and measure the effect of, single adverse factors in reducing yields
would be costly, difficult and perhaps inconclusive. Even if we
obtained, by counts in sample areas of a field, some idea of the
numerical abundance of different species of insects, there would
still remain the difficulty of converting this information into
terms of bushels lost due to specific, adverse agents.

The possibility of arriving at a solution seems almost out of the
question. On the other hand it would appear to be possible to
increase the accuracy of the estimates of the percentages of insect
damage to crops, by increasing the number of estimators and by
giving them some common, fundamental basis for their estimates
and by educating them in the importance of weighing the dif-
ferent factors. We have no confidence in the flat estimates by
single individuals covering widespread areas, even whole states,
unless it can be shown that such estimates are based upon ex-
tensive field observations and counts, where it has been possible
to make them, and a full appreciation of the various factors in-
volved. In the case of some estimates we have no doubt that such
care is exercised, but entomologists as a rule are not conservative
in estimating insect damage, and are inclined to extend to a very
large area the estimates that have been based on a very small and
unrepresentative sample, and that may represent a special case.

In Mr. J. A. Hyslop’s useful paper on ‘‘Losses Occasioned by
Insects, Mites, and Ticks in the United States” (U.S.D.A. Bur.
Ent. and PL Quar., Wash., D. C., July, 1938, mimeographed)
there is a wide variation in the percentage losses, due to various
insects, as gathered by him from different sources and this one
would expect. They are, no doubt, as correct as existing facili-
ties and interest in the subject permitted them to be. Many of
them seem conservative, especially if one has no definite informa-
tion with which to check them. Some were no doubt arrived at
on the basis of surveys and counts. And they forcibly call at-
tention to the seriousness of insect injury to crops. However, we
do not believe that they are all as accurate as it is possible to
make them.

When these losses are converted into dollars they total to a
staggering sum which includes the enhanced value given to the

June, 1940]

Weiss: Insect Losses

199

destroyed portions of the crops. It is the enhanced values that
we believe should be written off in the interest of accuracy. As a
matter of fact, the destroyed portions have no market or other
value. They donT exist. If insect damage could be eliminated
completely, the larger production during the readjustment period
would result in a lower price. In trying to give a unit value to
something that does not exist, and did not actually come into
existence, we either have to give it the same unit value of the
crop that does exist, which is not correct, or assume that the in-
creased production due to the elimination of insects could ac-
tually be marketed at the lower price. As this is only an assump-
tion and as increased production through elimination of insects,
in many cases, would result in a farm value much less than if the
reverse happened, i.e., lower production due to insect damage,
why is it necessary to attempt to convert bushel losses into dol-
lars? Why not allow these losses in production, when they are
arrived at as accurately as possible, to remain in bushels ? It is
too bad that our standards of value require so many things such
as insect damage, college educations, etc., to be valued in terms
of dollars.

From an economic viewpoint large farm surpluses at present
would not be regarded as blessings. If our present surpluses,
due to better methods of production, etc., the dislocation of for-
eign trade, industrial unemployment, etc., are still further aug-
mented by the elimination of all insect damage, which is highly
theoretical, our social and economic life would have to undergo
severe readjustments. Before large surpluses can be sold to low
income groups, a change in our methods of distribution will have
to take place. Economic entomologists should not dream of the
complete elimination of all losses due to insects, nor talk as if it
would really be desirable for this to happen. They should be
content to see their recommendations employed sufficiently to
prevent insect damage from reaching the proportions of disaster,
or in seeing that such damage is kept doAvn to a reasonable level.
Even though they are more ambitious than this, there are always
enough lax producers, and insects, to keep production at a level
more or less consistent with the economics of the times.

The

New York Entomological Society

Organized June 29, 1892 — Incorporated June 7, 1893
Certificate of Incorporation expires June 7, 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
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Members of the Society will please remit their annual dues, payable in January, to
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Honorary President, WILLIAM T. DAVIS

Officers for the Year 1940

President, DE, ALEXANDER B. KLOTS College of the City of New York, N. Y.

Vice-President, MAX KISLIUK, JR Federal Bldg., New York, N. Y.

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EXECUTIVE COMMITTEE

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PUBLICATION COMMITTEE
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DELEGATE TO THE N. ¥. ACADEMY OF SCIENCES
William T. Davis

JOURNAL

of the

NEW YORK ENTOMOLOGICAL SOCIETY

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No. 3

1^

/ol. XL VIII

SEPTEMBER, 1940

Journal

of the

New York Entomological Society

Devoted to Entomology in General

Edited by HARRY B. WEISS

Publication Committee

HARRY B. WEISS J. D. SHERMAN, Jr.

WILLIAM MOORE E. L. BELL

Subscription $3.00 per Year
Published Quarterly by the Society

N. QUEEN ST. AND McGOVERN AVE.
LANCASTER, PA.

NEW YORK, N. Y.

1940

CONTENTS

The Sarcophaginae and Their Relatives in New York, II

By Harold C. II alloc k 201

Observations on Silphin^ with a Note on Intraspecific
Variations and Their Designation

By Melville H. Hatch 233

A New Record for Connecticut

Leonard J. Sanford 244

Some Aspects of Modern Taxonomy

By Richard E. Blackwelder 245

Atrytone logan Edwards

Leonard J. Sanford 258

Some Parasites and Hyperparasites of the Cecropia Moth

By Osmond P. Breland 259

Comparison of Serologic and Taxonomic Relationships of
Drosophila Species

By Russell W. Cumley 265

New Peruvian Membracidae (Homoptera)

By W. D. Funkhouser 275

Proceedings of The New York Entomological Society 295

The Death-Feint of Trox unistriatus Beauv.

By Harry B. Weiss 303

NOTICE: Volume XLVIII, Number 2, of the Journal of
The New York Entomological Society was published
ON June 19, 1940.

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. XLVIII September, 1940 No. 3

THE SARCOPHAGIN^ AND THEIR RELATIVES
IN NEW YORK. II^

By Harold C. Hallock

Lancaster, Pa.

Sarcophaga Meigen

1826. Meigen, Syst. Beschr. Zwei. Ins., vol. 5, p. 14.

1916. Aldrich, Sarcophaga and Allies, p. 5.

Aldrich (1916) gave a fnll account of the history of the genus,
Sarcophaga, and the distinguishing characters which will not be
repeated here. The hypopygial studies have given additional evi-
dence that there can be some grouping of the species within the
genus. These structures are more important as the final char-
acters to be used in distinguishing some of the closely related
species.

The New York species {hisetosa, cimbicis, and latisterna) , which
Parker (1914) placed in the genus Boettcheria, have several com-
mon genitalian characters. There is a small rounded projection
on each side of the V of the fifth sternum a short distance before
its point of union. The sperm pump sclerite of each of these
three species is thickened along the margin of its larger end and
they are similar in size and shape. The posterior claspers of the
three species are more or less enlarged on the front side near the
base.

1 This article is a continuation of ‘ ^ The Sarcophaginse and their relatives
in New York. I.’’ Jour. N. Y. Ent. Soc. vol. 48, pp. 127-153, 1940.

Contribution from the Entomological Laboratory, Cornell University,
Ithaca, N. Y.

SEP i t mo

202

Journal New York Entomological Society [Vol. xlviii

The New York species {laakei, latisetosa, Vherminieri, pusiola,
stimulans, and suet a) of the Ravinia group have many similar
li3^popygial structures. When the anal forceps are viewed from
the back it is noticeable that each is united near its base, becomes
widely separated and then generally tend to be closer together at
the apex. There is a remarkable resemblance in the shape of the
aedeagus of these five species as will be seen by examining the
illustrations. Three species, Vherminieri, sueta, and pusiola have
a similar arrangement of short, stout bristles along each inner
margin of the V of the fifth sternite. The species, laakei, lati-
setosa, and stimulans, do not have this striking bristle arrange-
ment but are similar to each other as they have an enlargement
on the inner margin of the V of the fifth sternite.

It is evident that harhata and securifera are closely related.
Although the anal forceps can be distinguished they have a strik-
ing resemblance in their shape. Each species has a broad acces-
sory plate, a row of bristles along the posterior margin of the
anterior claspers, two bristles on the front margin near the apex
of the posterior claspers, considerable similarity in the shape of
the fifth sternites and the arrangement of the bristles along the
inner margins of the V of their fifth sternites.

There is some similarity in the aedeagus aiid anal forceps of
alcedo and uncata but the shape of the fifth sternites would not
lead one to expect very close relationship between these two spe-
cies. Aldrich (1916) compared both of these species to excisa.
There are several species in this group that have many characters
in common but they are not as closely related to each other as the
species within the Ravinia or Boettcheria groups.

It is true that there are distinct groups within the genus Sarco-
phaga. Yet it does not seem wise to attempt to express this rela-
tionship by creating separate genera for these related species. In
almost all the cases it would be nearly impossible to find generic
characters which would place the females in the proper genus.
The writer desires in the near future to study the hypopygial
structures of the female Sarcophaga which may help to show rela-
tionships within the group. It would also be wise to supplement
these characters by comprehensive larval studies of the type con-
ducted by Knipling (1936) and studies of the habits of the related
species.

Sept., 1940]

HALLOCK: SARCOPHAGINiE

203

Although there is a very striking similarity in the appearance
of the adult Sarcophaga, the habits of the immature stages show
considerable variation. As Aldrich (1916) brought to our atten-
tion these flies are now in the interesting transition from the habit
of devouring dead animal tissues to parasitism upon living tissue.
The larvae of the Bavinia group develop in excrement. There are
many Sarcophaga which have the habit of developing in dead
flesh as illustrated by Sarcophaga hullata. The aquatic Sarco-
phaga, which develop in pitcher plants, feed largely upon dead
insects but they will attack and kill living insects which occur in
the water found in the pitcher plant. There are many true para-
sitic forms in the Sarcophaga and these may be illustrated by
S. atlanis, which attacks grasshoppers, S. aldrichi, which attacks
chiefly the forest tent caterpillar, and S. ciml)icis, which has a
wide variety of hosts. Although S. cimbicis may develop as a
true parasite the larvge have also been reared on meat. The habits
of the New York species will be discussed more fully under each
individual species.

The keys given in the present paper include all SarcophaginaB
known to occur in New York state in order that the identification
of the flies may be made simpler and confusion avoided. Char-
acters other than those found on the genitalia have been used
whenever possible in the first key to the males. A second key to
the males, which is based entirely upon genitalia characters, has
been given in order to aid the student in the study of this group.
The third species key deals with the female Sarcophaginge of New
York where known material has been available for study.

Genotype. — Sarcophaga carnaria (Linn.)

Key to Separate Male Sarcophagin^

1. Three distinct postsutural dorsocentrals 2

Four postsutural dorsocentrals, or else the anterior ones of the post-
sutural series indistinct, scarcely differentiated from hairs 31

2. Hind tibiae with villosity, the hairs of which are at least longer than the

diameter of the tibiae 3

Hind tibiae without erect villosity (the appressed hairs at most not longer
than the diameter of the tibiae) 9

3. Hypopygium entirely black 4

At least the second segment of the hypopygium red or yellow 5

204

Journal New York Entomological Society [Vol. XLVIII

4. Abdomen pollinose, at most the hinder parts of the segments shining
black ; middle femur with a patch of yellow hairs on the apical third
of front sides SarcopJiaga sinuata Meig.

Abdomen with three rows of shining black spots on the second, third, and
fourth segments ; middle femur without yellow hairs

Wohlfahrtia vigil Walker

5. Anterior acrostichals present 6

Anterior acrostichals absent 8

6. Prescutellar bristles present, the third abdominal segment with a com-

plete row of marginal bristles SarcopJiaga sarracenice Eiley

Prescutellar bristles absent, the third abdominal segment with only
median marginal bristles 7

7. Accessory plate long, slender, finger-like, only the distal end of the row

of median bristles on the hind femur present

SarcopJiaga latisterna Parker
Accessory plate not finger-like, a complete row of median bristles on
hind femur SarcopJiaga cimloicis Townsend

8. Third abdominal segment with a complete row of marginal bristles,

fourth abdominal segment yellow apically SarcopJiaga sima Aldrich

Third abdominal segment with only a median pair of marginal bristles,
fourth abdominal segment entirely black

SarcopJiaga bisetosa Parker

9. Hypopygium entirely black 10

At least second segment of the hypopygium red or yellow 17

10. The frontal bristles extending below the base of the antennae 13

The frontal bristles not extending below the base of the antennae 11

11. Arista short plumose 12

Arista long plumose; legs largely yellow; abdomen largely yellow in the
light form and abdomen dark in the dark form

Opsodexia bicolor Coq.

12. Legs entirely black ; palpi brownish and nearly as long as the proboscis

Oppiopsis sJieldoni Coq.

Tibiae yellow; palpi yellow and only about one-third the length of
proboscis Opelousia obscuria Town.

13. Anterior acrostichals wanting 16

Anterior acrostichals present and well developed 14

14. Ground color of ventral portion of abdomen largely red, second and

third abdominal segments with only median marginal bristles

SarcopJiaga fletcJieri Aid.
Ground color of ventral portion of abdomen black 15

15. Accessory plates nearly as long as anal forceps; the V lacking on the

fifth ventral sternum Laccoprosopa sarcopJiagina Town.

Accessory plates short; fifth sternite has a well developed V

SarcopJiaga nox Hall

16. Palpi yellow, forceps with strong protuberances on dorsal half, on which

are striking tufts of long black hair SarcopJiaga Jmnteri Hough

Sept., 1940]

Hallock: Sarcophagin^

205

Palpi black, forceps slender, protuberances and tufts of hair lacking

Sarcophaga atlanis Aid.

17. First, third and fifth longitudinal veins hairy, fourth abdominal segment
and hypopygium yellowish red Johnsoni horealis Reinhard

Only first and third longitudinal veins hairy 18

Third longitudinal vein only hairy 19

18. Anterior acrostichals well developed, second abdominal segment with

median marginal bristles MetoposarcopJiaga import una Walker

Anterior acrostichals not differentiated from the surrounding hairs, sec-
ond abdominal segment without median marginal bristles

Sarcophaga rapax Walker

19. Palpi yellow 20

Palpi black 22

20. First segment of hypopygium with a row of stout bristles near its base,

fifth sternite entire 21

First segment of hypopygium without row of bristles near base, the row
of bristles at apex often indistinct, fifth sternite deeply cleft form-
ing a distinct V Sarcophaga hunteri Hough

21. Arista plumose; parafacials and paraf rentals light golden yellow pol-

linose Sarcofahrtia ravinia Parker

Arista bare or slightly pubescent; parafacials and paraf rentals gray
pollinose Erythrandra picipes B. & B.

22. Arista long and thin, short-plumose; the second segment of the large

hypopygium red and with long bristles Hypopelta scrofa Aid.

Species not as described 23

23. Anterior acrostichals absent or scarcely differentiated from surrounding

hairs Sarcophaga parallela Aid.

Anterior acrostichals well developed 24

24. Epaulets yellow, frontal rows of bristles not divergent below

Sarcophaga pusiola V. d. W.
Epaulets black, the frontal rows of bristles suddenly divergent in the
last two or three bristles 25

25. The third abdominal segment with a pair of median marginal bristles,

median marginal bristles absent on the second abdominal seg-
ment 26

Species lacking the above combination of characters 29

26. Wings slightly smoky, hind calypters light brown with a white margin,

parafacials and parafrontals yellow pollinose

Sarcophaga alcedo Aid.

Species lacking the above combination of characters 27

27. Fourth abdominal segment yellow apically; accessory plate broad and

deeply excised at apex Sarcophaga excisa Aid.

Fourth abdominal segment black; accessory plate round at apex 28

28. Apex of sedeagus with a long tube-like portion which is incurved at end

to form a loop that can be seen through Sarcophaga reversa Aid.

Tip of sedeagus without loop, but with thin expanded margins which are
whitish or transparent Sarcophaga uncat a V. d. W.

206

Journal New York Entomological Society [Vol. xlviii

29. Euf0e-like expansions lacking on aedeagus, the minute incurved copula-

tory tube concealed by the stout divergent prongs at the tip of the

aedeagus Sarcophaga aculeata Aid.

On the discal portion of the aedeagus appear lateral, thin expanded
margins which are white or transparent and shaped similar to a
ruffle 30

30. A pair of erect median marginal bristles on the second abdominal seg-

ment, the distal portion of the aedeagus distinctly notched to form

two sharp points at the apex Sarcophaga setigera Aid.

Median marginal bristles rarely present on second abdominal segment
(when present small and widely separated) ; distal segment of
aedeagus swollen and whitish at end with no sign of notch

Sarcophaga coloradensis Aid.

31. Hind tibiae with villosity, the hairs of which are at least longer than

the diameter of the tibiae 32

Hind tibiae without erect villosity (the appressed hairs are at most not

longer than the width of the tibiae 45

32. Hypopygium entirely black 33

At least the second segment of the hypopygium red 37

33. Middle tibiae with long dense villosity; front approximately as wide as

one eye Sarcophaga johnsoni Aid.

Middle tibiae without villosity; front much narrower than the width of
the eye 34

34. Anal forceps broad, approximately the same width for three-fourths

their length; anterior claspers deeply grooved on back

Sarcophaga aldrichi Parker

Species lacking the above characters 35

35. HSdeagus globose with several long slender processes at apex

Sarcophaga misera var. harpax Pand.
The long slender processes at apex of the aedeagus lacking 36

36. The aedeagus with a large hump on the back ; second abdominal segment

with median marginal bristles Sarcophaga houghi Aid.

The aedeagus without conspicuous hump on the back; the second ab-
dominal segment without median marginal bristles

• Sarcophaga montanensis Hallock

37. First segment of hypopygium black on apical half 38

Both segments of hypopygium yellow or reddish in color 41

38. The prescutellar bristles absent; the forceps with a notch behind near

base Sarcophaga hcemorrhoidalis Fall.

Species lacking the above characters 39

39. A single row of black hairs back of eyes, abundant white beard which

extends over the back of the head up to the row of black hairs

behind eyes Sarcophaga securifera Vill.

With more than one row of black hairs back of eyes and white beard
less extensive 40

Sept., 1940]

Hallock: Sarcophagin^

207

40. Forceps without emarginatioii and hump behind; sedeagus with slender

processes at tip; two rows of black cilia behind eyes; white hair on

posterior portion of cheeks Sarcophaga misera var. exuberans Pand.

Forceps with an emargination behind followed by a hump, terminal
portion of aedeagus much enlarged and no slender processes

Sarcophaga harhata Thompson

41. The middle tibiae with long villosity, which begins about the middle 42

Middle tibiae without long villosity 43

42. The length of villous hair on hind tibiae approximately twice the diameter

of the tibiae; length of villous hair on middle tibiae approximately
equal to the diameter of tibiae

Sarcophaga scoparia var. nearctica Parker
The length of villous hairs on hind tibiae approximately three times the
diameter of the tibiae; length of villous hairs on middle tibiae twice
the diameter of tibiae Sarcophaga fulvipes var. triplasia V. d. W.

43. Third abdominal segment without median marginal bristles

Sarcophaga cooleyi Aldrich
Third abdominal segment with median marginal bristles 44

44. Forceps not tapering to a point but with an apical tooth formed by an

excision on the front edge; distal segment of aedeagus not unusually
broad, with long slender processes at tip ; three rows of black cilia
behind eyes; hairs on cheeks black

Sarcophaga misera var. sarracenioides Aid.
Forceps tapering to a sharp point ; distal segment of aedeagus broad and
blunt Sarcophaga bullata Parker

45. First vein hairy 54

First vein bare 46

46. Epaulets yellow 47

Epaulets black 48

47. Fourth segment of abdomen and first segment of hypopygium widely

yellowish red; parafrontals and parafacials yellow pollinose

Sarcophaga sueta V. d. W.
Fourth segment of abdomen and first segment of hypopygium black;
parafrontals and parafacials gray pollinose

Sarcophaga I ’herminieri Eob.-Des.

48. Palpi yellow, pale hairs of beard conspicuous in front of metacephalic

suture Sarcophaga flavipalpis Aid.

Palpi entirely black 49

49. Scutellum with a pair of apical bristles between the large marginal

bristles 50

The apical bristles on the scutellum lacking 52

50. Frontal bristles extending below the base of the antennge and the last

two or three bristles in the frontal rows divergent; parafrontals

and parafacials gray pollinose 56

Frontal bristles not extending below the base of antennae and not diver-
gent; parafrontals and parafacials golden yellow pollinose 51

208

Journal New York Entomological Society [Vol. xlviii

51. Fourth abdominal segment and the hypopygial segments covered with

golden yellow pollinose hairs Macronidhia aurata Coq.

Fourth abdominal segments and the hypopygial segments covered with
only gray pollinose Macronicliia confundens Town.

52. Front approximately as wide as one eye Sarcopliaga cistudinis Aid.

Front much narrowed below the ocelli so that it is about half as wide

as one eye 53

53. Anterior acrostichals present; fourth abdominal segment black apically;

legs yellow Sarcopliaga ventricosa V. d. W.

Anterior acrostichals absent; fourth abdominal segment yellow apically;
legs black Sarcopliaga cingarus Aid.

54. First segment of the hypopygium red and fourth abdominal segment

partially red Sarcopliaga laalcei Hall

First segment of the hypopygium black and fourth abdominal segment
black 55

55. Outer vertical bristles present Sarcopliaga latisetosa Parker

Outer vertical bristles absent Sarcopliaga stimulans Walker

56. Forceps with dense erect short hairs almost at tip Sarcopliaga utilis Aid.

Short erect bristly hairs lacking near tip of forceps 57

57. Forceps parallel to the tip, cut bluntly at tip, and the two forcexis curved

beneath so as to give a spoon appearance when viewed beneath

Sarcopliaga niagarana Parker
Forceps parallel more than three-fourths the length but tips widely
separated so as to form a semi-circle when viewed from behind

Sarcopliaga yorMi Parker

Key to Separate Male Sarcophagin^

Based on Hypopygial Characters

1. Fifth sternite cleft, so that two lobes extend caudad, generally forming

a V 3

Fifth sternite entire with no sign of a V on its caudal margin 2

2. Both segments of hypopygium black W olilf alirtia vigil Walker

Both segments of hypopygium red Sarcof alirtia ravinia Parker

3. First segment of hypopygium large, flat behind so that the abdomen

apears truncate; pump sclerite unusually large as its diameter is
twice the length of the sedeagus

Metoposarcopliaga importuna Walker
First segment of hypopygium normal; pump sclerite small 4

4. Fifth sternum with a pair of median pad-like projections at the point

where the gradual sloping V suddenly becomes nearly parallel sided

on the caudal margin {Bcettclieria group) 5

The V of the fifth sternum without pad-like projections 7

5. The tip of the gedeagus long and extending cephalad with a number of

slender processes beneath; posterior claspers normally slender

Sarcopliaga latisterna Parker
The tip of the sedeagus strongly curved back caudad; posterior claspers
with a decided enlargement near their middle 6

Sept., 1940]

Hallock: Sarcophagin^

209

6. Anal forceps short and wide, basal half or more with tufts of very

striking long, black hair; anterior clasper with tooth of apex

SarcopJiaga cimhicis Town.
Anal forceps longer, curved at tip and without striking tufts of hair;
anterior claspers curved at apex Sarcophaga bisetosa Parker

7. Apex of sedeagus with two black transverse arms which curve around

forward and make an almost perfect ring and each arm is divided
into two points at the apex

Sarcophaga scoparia var, nearctica Parker
.^deagus without the chacteristic arms 8

8. Anal forceps cleft approximately two-thirds or more of the distance

from the tip to the base 9

Anal forceps cleft only a short distance or not more than half way from
the tip to the base 22

9. Anal forceps nearly straight in profile with a notch behind near base

of cleft; arising from the front side of the sedeagus a little before
the tip are a pair of curved arms reaching forward which together
with the flat and enlarged tip suggest the figure of a small dog
sitting up in the attiude of begging

Sarcophaga hcemorrhoidalis Fall.
Species lacking the above characters 10

10. Distal segment of the aedeagus in the form of a long tube which is

incurved at the end and forming a loop that can be seen through

in profile beyond the lateral processes Sarcophaga reversa Aid.

Distal portion of asdeagus without loop that can be seen through 11

11. Forceps when viewed from back show a deep depression on the caudal

side about the middle 12

Forceps without the depression 16

12. The yellow forceps rather square at the tip, and the base of the forceps

enlarged into a prominent lobe which bears a striking tuft of
straight black hairs twice as long as the forceps

Sarcophaga hunteri Hough
Anal forceps without lobe and tuft of black hairs 13

13. Fifth sternite large, the V is cut out somewhat in the form of a semi-

circle, the lower portion of the V margined with strong bristles;
distal segment of aedeagus large with a keel-like hind margin
which extends in a curve over the tip into a pronounced free

hook turned forward Sarcophaga flavipalpis Aid.

Species lacking the above characters 14

14. Inner margins of anal forceps parallel for half of distance to the

points, then suddenly turned outward so that the points of forceps
are widely separated; the margins of the V of the fifth sternite
parallel over half the distance to the point and then suddenly
divergent so that points are widely separated, a group of stout
bristles on inner margins about one-third distance from points

Sarcophaga alcedo Aid.

Species lacking the above characters 15

210

Journal New York Entomological Society [Vol, xlviii

15. Fifth sternite with a bristly projection on each of the inner margins of

the V at about its middle; sedeagus without ruffle

SarcopJiaga ailanis Aid.
Fifth sternite without projection on the inner margins of the V ;
sedeagus with a lateral white fringe or ruffle on its apical half

Sarcophaga uncata V. d. W.

16. Anal forceps widely separated from each other at tip and at base

attached to each other only for a short distance; bristles confined
largely to outer margins of forceps; the usual V-shaped cleft of the
fifth sternite is a linear fissure in this species and on each side of
the fissure behind is a small black hood-like point

Sarcophaga fletcheri Aid.

Species without these characters (Eavinia group) 17

17. Anterior claspers with a row of bristles extending the entire length of

caudal margin 18

Anterior claspers without bristles on caudal margin 19

18. Aedeagus when viewed from behind decidedly narrowed before the

apex; anal forceps less widely separated at tip than at one-fourth

of distance toward base Sarcophaga I’herminieri E.-D.

.^deagus when viewed from behind not narrowed; anal forceps
gradually divergent toward apex Sarcophaga sueta V. d. W.

19. ^deagus ending in two rounded enlargements separated by a median

notch, on front side well before the apex are two pairs of processes

projecting forward Sarcophaga pusiola V. d. W.

The sedeagus without rounded enlargements 20

20. Anterior clasper terminating in point and nearly as long as aedeagus;

posterior clasper slender and smooth on front side; fifth sternite

broadly excised Sarcophaga stimulans Walker

Anterior clasper much shorter than aedeagus; posterior clasper with two
small humps on front margin near tip ; the fifth sternite less broadly
excised 21

21. Anterior clasper hallowed out or dished on front margin at tip so that it

has the appearance of being toothed Sarcophaga latisetosa Parker

Anterior clasper ending in a point without any dished appearance on
front margin Sarcophaga laakei Hall

22. The V of fifth sternite narrowed almost to a linear fissure ; forceps when

viewed from side are turned back near the apex to appear somewhat

like the barb of a hook Sarcophaga ventricosa V. d. W.

The V of fifth sternite normal; anal forceps without barbed-like ap-
pearance 23

23. Anal forceps black, broad and with shining area on lower half when

viewed on the caudal side; anterior claspers large and deeply
grooved on front side almost to the apex

Sarcophaga aldrichi Parker
Anal forceps without broad shining area; anterior claspers not deeply
grooved 24

Sept., 1940]

Hallock: Sarcophagin^

211

24. Hypopygium very small; a single very long bristle on anterior margin

of posterior claspers; a few long curved bristles at tip of V of

fifth sternite Sarcophaga rapax Walker

H}T)opygiuni normal size; posterior claspers without long bristles; fifth
sternite without long curved bristles at apex of V 25

25. .liEdeagus with a large hump on the back side near the apex; the

diverging lobes of the V of the fifth sternite covered with short,

stout, black bristles Sarcophaga houghi Aldrich

.i3Edeagus without hump 26

26. Arms of the V of the fifth sternite very widely divergent, the portion

of sternite around the union of V covered with strong bristles;

distal segment of sedeagus very broad and blunt 27

Species with different characters 28

27. Accessory plate covered with short bristles only; tip of sedeagus with a

process curving forward which is cut off bluntly on front apex and
.aedeagus only slightly narrowed at junction of its two segments

Sarcophaga cooleyi Parker
Accessory plate with long bristles at its apex; the process at tip of
aedeagus ends in a sharp point and the aedeagus is very much nar-
rowed at junction of the two segments Sarcophaga hullata Parker

28. Arms of V of fifth sternite long, slender and sparsely covered with

slender hairs, at the union of the V arms are two circular pads
which are densely covered with bristles; anal forceps have abundant
short, stiff erect hairs on the front and back sides just beyond
their lower angle but the extreme tips of forceps are bare

Sarcophaga johnsoni Aldrich
Circular pads lacking on fifth sternite and some true strong bristles on
fifth sternite; characteristic groups of short, stiff erect hairs lack-

ing on anal forceps 29

29. Tips of aedeagus with five characteristic slender processes {miser a

group) 33

Tips of aedeagus without the -long slender processes 30

30. The entire hypopygium black 31

At least the second segment of hypopygium red 32

31. Posterior clasper with a long bristle near base; apex of aedeagus with

a raised light colored piece extending around its tip

Sarcophaga nox Hall

Posterior clasper without long bristle near base; raised light colored
piece lacking on apex of aedeagus Sarcophaga sinuata Meig.

32. Anal forceps with a hump on the caudal side just before the apex; and

the front side concave to form a hooked tip ; the apex of the
aedeagus with two long slender processes which are distinctly

widened at the apices Sarcophaga securifera Vill.

Anal forceps without hump ; processes when present on the aedeagus
lack the enlarged apices 35

212

Journal New York Entomological Society [Vol. xlviii

33. Both liypopygial segments black; processes arising from apex of

sedeagus slender and the ones curved around to the front are

toothed at the apex Sarcophaga misera var. harpax Band.

At least the second segment of hypopygium wholly red; processes at
apex of sedeagus not toothed at tip 34

34. First segment of hypopygium generally brownish black; distal processes

of aedeagus long and very slender

Sarcophaga misera var. exuherans Band.
Both segments of hypopygium red ; distal processes of the aedeagus
shorter and thicker Sarcophaga misera var. sarracenioides Aid.

35. Anal forceps in back view contiguous to the tip 41

Anal forceps not contiguous to the tip 36

36. Bosterior claspers much enlarged and attached to ninth sternite for

at least half their length, a row of long bristles on caudal margin
of posterior claspers; anterior claspers greatly curved, toothed, and

with a long bristle near the base Hypopelta scrofa Aldrich

Anterior and posterior claspers normal 37

37. Anterior clasper slender, widened apically, the tip excised to form a

distinct claAV ; arms of fifth sternite black

Sarcophaga sarracenice Biley

38. Apex of aedeagus ending caudad in a keel which turns sharply forward

in the form of a hook; anal forceps excised in front to form an

apical tooth 40

Apical segment of aedeagus very wide and without a distinct keel at
apex; anal forceps not excised in front to form an apical tooth 39

39. Apex of anal forceps with a row of short bristles on front and back

sides just back of tip; tip of anal forceps from side view resembles

a head of a bird Sarcophaga utilis Aldrich

Apex of anal forceps without bristles and not resembling a head of
a bird 42

40. Anterior clasper dished on the anterior surface; the anterior process

of the apex of the aedeagus, when viewed from the back, appears
like a broad scoop with its center curved back and the caudal keel

comes up around it Sarcophaga sima Aldrich

Anterior clasper not dished but with a notch at the apex, the front
margin of apex of aedeagus somewhat rectangular in shape

Sarcophaga parallela Aldrich

41. Forceps with a lateral process near tip Sarcophaga cm gar us Aldrich

Forceps without lateral process near tip

Sarcophaga niagarana Barker

42. Forceps cleft nearly half their length; aedeagus large and apical circle

lacking Sarcophaga harhata Thompson

Tips of anal forceps separated only a very short distance; aedeagus
moderate size and its apical end shaped to form a semicircle when
viewed toward the tip Sarcophaga yorlcii Barker

Sept., 1940]

Hallock: Sarcophagin^

213

Key to Separate Female Sarcophagin^

1. Three distinct postsutural dorsocentrals 2

Four postsutural dorsocentrals, or else the anterior ones of the post-
sutural series indistinct, hardly differentiated from hairs 19

2. Genital segments mostly black 3

Genital segments red or reddish yellow 5

3. Arista bare; epaulets yellowish brown

Laccoprosopa sarcopliagina Town.
Arista long plumose; epaulets black 4

4. Anterior acrostichals absent, mid-femur with patch of yellow hairs on

apical third of front side Sarcophaga sinuata Meig.

Anterior acrostichals present; mid-femur without yellow hairs; palpi
yellow Sarcophaga hunteri Hough

5. Anterior acrostichals absent, not differentiated from hairs 6

Anterior acrostichals present 10

6. First vein hairy, scutellum with apical bristles, small species

Sarcophaga rapax Walker
First vein bare, larger species 7

7. Second abdominal segment with median marginal bristles, ground color

of ventral portion of abdomen largely red

Sarcophaga fletcheri Aldrich
Second abdominal segment without median marginal bristles, ground
color of abdomen largely black 8

8. Prescutellar bristles absent; arista long and thin, short plumose; palpi

black Hypopelta scrofa Aldrich

Species lacking the above grouping of characters 9

9. Palpi black ; arista long plumose Sarcophaga bisetosa Parker

Palpi yellow; plumosity of arista only median length

Sarcofahrtia ravinia Parker

10. Second abdominal segment with erect median marginal bristles 11

Second abdominal segment without erect median marginal bristles ... 13

11. Epaulets yellow, abundant whitish beard Sarcophaga atlanis Aldrich

Epaulets black, beard black 12

12. Median marginal bristles on second abdominal segment erect and close

together, fourth abdominal segment entirely black, larvipositor

sharp and bent forward Sarcophaga setigera Aldrich

Median marginal bristles when present on second abdominal segment
not erect and considerable distance apart, fourth abdominal seg-
ment yellow apically, larvipositor rounded and not bent forward

Sarcophaga excisa Aldrich

13. Three maginal pairs of bristles on scutellum 14

Two long marginal pairs of bristles on scutellum 16

14. Sides of fourth abdominal segment nearly covering fourth sternite,
apical edges of fourth segment rapidly separating, forming a V,
and exposing the red fifth sternite

Metoposarcophaga importuna Walker

214

Journal New York Entomological Society [Vol. xlviii

15.

16.

17.

18.

19.

20.

21.

22.

23.

24.

25.

26.

27.

Fourth sternite widely exposed as the fourth tergites are widely sepa-
rated ventrally 15

Anterior cross-vein infuscated SarcopJiaga cimbicis Town.

Anterior cross-vein not infuscated SarcopJiaga latisterna Parker

Epaulets yellow, mid tibiae with two bristles on outer front side

SarcopJiaga pusiola V. d. W.

Epaulets black 17

Wings slightly smoky, hind calypters light brown with a white margin

SarcopJiaga alcedo Aldrich

Wings hyaline, calypters white 18

Fourth abdominal segment yellow apically, genital orifice rounded and

surface of dorsal half without bristles

SarcopJiaga excisa Aldrich
Fourth abdominal segment entirely black, yellow larvipositor pointed
and the surface of dorsal portion covered with bristles

SarcopJiaga reversa Aldrich
Impression of the bucca, near the vibrissa! angle, large, conspicuously
submerged below the surface level of surrounding sclerite; vibris-
sae placed at least the length of second antennal joint above the

oral margin; ovipositor drawn out into a long slender point 20

Species lacking the above combination of characters 21 ,

Fourth abdominal segment and the hypopygial segments covered with

golden yellow pollinose hairs MacronicJiia aurata Coq.

Fourth abdominal segment and the hypopygial segments covered with

gray pollinose hairs MacronicJiia confundens Town.

Genital segments mostly black 22

Genital segments ■ entirely red or reddish yellow 24

Palpi black SarcopJiaga aldricJii Parker

Palpi yellow 23

Third abdominal segment with only a pair of median marginal bristles

SarcopJiaga joJinsoni Aldrich
Third abdominal segment with complete row of marginal bristles; ab-
domen with three rows of shining black spots on second, third,
and fourth segments. Eastern species ...WoJilfaJirtia vigil Walker

Anterior acrostichals present 25

Anterior acrostichals absent 29

Epaulets black, dark yellow palpi which are rather long and broad

SarcopJiaga flavipalpis Aldrich

Epaulets largely yellow; palpi black 26

First vein hairy 27

First vein bare ' 28

Middle femur with two bristles on outer front side

SarcopJiaga stimulans Walker

Middle femur with more than two bristles on outer front side

SarcopJiaga latisetosa Parker

Sept., 1940]

Hallock: Sarcophagin^

215

28. Parafrontals, parafacials and front half of bucca with strong ochraceous

pollinose; thorax and abdomen with yellowish pollen

Sarcophaga sueta V. d. W.
Yellowish pollen replaced by gray pollinose, with reflections that are
rather brownish than yellow Sarcophaga I’herminieri Eob.-Des.

29. Prescutellar bristles present 30

Prescutellar bristles absent Sarcophaga hcemorrhoidalis Fall.

30. Fifth vein with strong setae along vein on the upper side of the wing

Johnsoni horealis Eeinhard
Fifth vein without setae 31

31. At least one-fourth of beard white 32

Beard containing no white hairs 37

32. Whitish beard of metacephalon extends well forward of the suture,

blending with the black hairs 33

Whitish beard not extending beyond the suture and blending with the
black hairs 35

33. Single row of black bristles behind eyes, bucca with abundant white

beard which also extends over back of head up to row of black
hairs, dorsal marginal bristles lacking on third abdominal segment

Sarcophaga securifera Vill.
Not with above combination of characters 34

34. A single row of bristles in cheek groove extending nearly the entire

height of parafacials Sarcophaga harbata Thompson

Bristles on lateral sides of parafacials irregularly arranged and not
in a single row , Sarcophaga bullata Parker

35. Middle tibiae with three bristles on the outer front, a group of para-

facial macrochaetae present, three distinct rows of black hairs behind
eyes, genital orifice triangular in shape and fringed with bristles

above Sarcophaga misera var. sarracenioides Aid.

Characters not in the above combination 36

36. Fourth abdominal segment yellow apically, parafacial bristles not ar-

ranged in a distinct row Sarcophaga cistudinis Aldrich

Fourth abdominal segment entirely black, a row of strong bristles ex-
tending nearly the entire length of parafacials

Sarcophaga scoparia var. nearctica Parker

37. The hind and mid-femur yellowish red Sarcophaga ventricosa V. d. W.

The hind and mid-femur dark colored (black)

Sarcophaga cingarus Aldrich

MISERA GROUP

The species placed in this group have been recognized only as
varieties of misera, the first species described in the group.
Parker (1919) and Hardy (1927) gave the history of this group
and so it will not be repeated here. Although there is an unusu-
ally close resemblance of the male hypopygium in all the species

216

Journal New York Entomological Society [Vol. xlviii

in the misera group the males may be separated from each other.
On the other hand it is very difficult to separate the females from
other Sarcophaga which do not belong in the misera group.
Additional studies may show that these species should be raised
from the rank of varieties to species and place them in a sub-
genus. The writer would not go further as it does not seem wise
to erect a genus based upon the characters of only one sex. A
clear understanding of the group can be secured only by exten-
sive rearings and a careful study of all stages.

Sarcophaga misera var. exuherans Pand.

1896. Pandelle, Rev. Ent., vol. 15, p. 186.

1912. Bottcher {S. tuberosa var. exuherans P.), Deutsch. Ent.
Zeitschr., p. 735.

1916. Aldrich, Sarcophaga and Allies, p. 232.

1919. Parker {S. dux var. exuherans P.), Bull. Brooklyn Ent.
Soc., vol. 14, p. 41.

1927. Hardy {S. misera var. exuherans P.), Proc. Linn. Soc.
N. k Wales, vol. 52, p. 452.

There are several very closely related forms that have been
placed in the misera group by the different students of Sarco-
phaga which can be separated only by close study. Their separa-
tion depends largely upon genitalia characters which have been
given in the keys. It should be kept in mind that exuherans has
two rows of black cilia behind the eyes and white vestiture on the
posterior portion of the cheeks.

Records. — Ithaca. June to August. Figure 98.

Jack (1935) reports this species to have been reared from dead
insects in South Rhodesia, Africa. Callot (1935) reared the
species from adults of Stauroderus hicolor Ch.

Type. — In the Pandelle collection in Paris.

Sarcophaga misera var. harpax Pand.

1896. Pandelle, Rev. Ent., vol. 15, p. 189.

1912. Bottcher {S. tuherosa var. harpax P.), Deutsch. Ent.
Zeitschr., p. 735.

1916. Aldrich {S. tuherosa var. harpax P.), Sarcophaga and
Allies, p. 171.

Sept., 1940]

Hallock: Sarcophagin^

217

1919. Parker {S. dux A^ar. harpax P.), Bull, Brooklyn Ent. Soc.,
vol. 14, p. 41.

1927. Hardy {8. misera ysly. harpax P.), Proc. Limi. Soc. N. S.
Wales, vol. 52, p. 452.

1930. Aldricli, Proc. U. S. N. M., vol. 78, p. 27.

Only a single specimen of Sarcophaga misera var. harpax P.
has been taken in New York State. This form is easily distin-
gnished from all other varieties of misera that occur in America.
The genitalia segments are entirely black. The processes at the
apex of the ^deagns, which curve anteriorly, are distinctly
toothed at their tips. The cheeks are clothed with black hairs.
Length. — 8 to 12 mm.

Record. — Pelham Park, New York City. Figure 99.

Aldrich (1916) states that the fly has been reared from Gypsy
moth caterpillars. Senior- White (1927) records the rearing of
harpax from decaying snails.

Type. — In the Pandelle collection in Paris.

Sarcophaga misera \mr. sarracenioicles Aid.

1916. Aldrich {8. tuherosa var. sarrace7iioides) , Sarcophaga and
Allies, p. 227.

1919. Parker {S. dux var. sarracenioides A.), Bull. Brooklyn
Ent. Soc., vol. 14, p. 41.

1927. Hardy {S. misera var. sarrace7iioides A.), Proc. Linn. Soc.
N. S. Wales, vol. 52, p. 452.

This species, sarrace7iioides, is one of the most common Sarco-
phaga occurring in New York. There is a single row of strong
black cilia and two other rows of weaker black cilia behind the
eyes. The vestitnre of the cheeks is black.

Length. — 8 to 13 mm.

Records. — Axtoii; Ithaca; New York City; L. I.: Cold Spring
Harbor ; Babylon ; Half-way Hollow Hills. May to September.
Figures 100, 101 and 102.

A very Avide range of rearings have been reported for this spe-
cies. It is certain in many cases that living insects have been
parasitized. Aldrich (1916) reported 45 adults from grasshop-
pers at widely distributed localities; 1 adult from the western
cricket in British Columbia; 5 adults from Heliopila imipuncta
in Oklahoma ; 23 adults from Lachnosterna sp. in Kansas ; 2

218

Journal New York Entomological Society [Vol. xlviii

adults from Eleodes tricostata in Oklahoma ; 2 adults from Allo-
rhina nitida in Virginia and 66 adults from carrion in Texas.
Davis (1919) found the fly parasitizing Phyllophaga sp. in Illi-
nois. Treherne and Buckell (1924) reported parasitism by this
fly on grasshoppers in British Columbia. Knipling (1936) reared
this species from meat. The writer in 1935 found that sarraceni-
oides would larviposit on the inner surface of a funnel of a beetle
trap above dying Asiatic garden beetles and that the first instar
maggots would drop about six inches to reach the bodies of the
beetles.

Holotype and allotype. — Male and female, No. 20551, U. S.

N. M.

BOETTCHERIA GROUP

Parker (1914) described Boettcheria as a new genus and placed
three species, hisetosa, cimhicis T. (fernaldi P.) and latisterna,
in the genus. The females were excluded from consideration in
Parker’s (1914) paper. Aldrich (1916) stated that the species
are closely allied but expressed the doubt, ‘‘if the fact is best
expressed by raising them to generic rank.” The writer, under
the discussion of the genus Sarcophaga, has pointed out addi-
tional hypopygial characters which are common to the flies in this
group. It is hoped that opportunity may be secured to conduct
additional studies later with the immature forms and the adult
female flies which may show that it is advisable to place these
species in a separate genus.

Sarcophaga hisetosa Parker

1914. Parker, Proc. Boston Soc. Nat. Hist., vol. 35, p. 69.

1916. Aldrich, Sarcophaga and Allies, p. 81.

Parker (1914) separated this species from the closely related
cimhicis mainly by the absence of anterior acrostichals and only
two sternopleurals while cimhicis has three sternopleurals. The
enlarged tip of the aedeagus, which turns back caudad, separates
this species from all the closely related species.

Length. — 8 to 12 mm.

Records. — Ithaca ; Albany ; White Lake ; Trenton Falls ; Yon-
kers ; L. I. : Half-way Hollow Hills. May to September. Figures
103, 104 and 105.

Sept., 1940J

Hallock : Sarcophagin^

219

Information is not available about the immature stages.

Type. — Mass. Agri. College.

Sarcophaga cimhicis Townsend
1892. Townsend, Can. Ent., vol. 24, p. 126.

1896. Van der Wulp {8. sehUosa), Biol. Cent. Am., vol. 2, p. 276.
1914. Parker {8. fernaldi), Proc. Boston Soc. Nat. Hist., vol. 35,
p. 72.

1916. Aldrich, 8arcophaga and Allies, p. 79.

1930. Aldrich, Proc. U. S. Nat. Mus., vol. 78, p. 32.

The characters given in the two male and female keys easily
separate this species from the two very closely related species
{hisetosa and latisterna). In most localities cimhicis is more
plentiful than either hisetosa or latisterna.

Length. — 5 to 10 mm.

Records. — Brie Co. ; Ithaca ; Albany ; Carmel ; W. Nyack ; New
York City; Tuxedo ; Brewster; Poughkeepsie; Oneonta; Rome;
Troy; 8.1. : Watchogue; Ft. Wadsworth; L.I.: Babylon; Jericho;
Riverhead; Heckscher State Park. May to October. Figures
106, 107 and 108.

Kelly (1914) recorded the species as a parasite of adult grass-
hoppers in Kansas. Aldrich (1916) recorded that the type spe-
cies was reared at Brookings, S. Dak., from pupa of Cimhex amer-
icana. Hayes (1917) found the fly breeding in dead insects.
Davis (1919) reared this species in Illinois from adult Pliyllo-
phagd sp. Sherman (1920) reared cimhicis from the green clover
worm larva {Plathypena scahra). Breakey (1929, 1931) re-
corded the larva of the iris borer as a host of cimhicis. The
writer (1929) found 40% of the adult beetles (Xyloryctes
satyr us) had been parasitized by this fly at Jericho on Long
Island. Experiments at that time tended to indicate that cimhicis
would not larviposit upon dead insects. Knipling (1936) pressed
larvae from the female flies and reared them on meat. Decker
(1931, 1932) in Iowa recorded the parasite from Papaipema
nehris and Epihlema otiosana.

Type. — In University of Kansas.

8arcophaga latisterna Parker

1914. Parker {Boettcheria) , Proc. Boston Soc. Nat. Hist., vol. 35,
p. 67.

220

Journal New York Entomological Society [Vol. XLVlir

1916. Aldrich, Sarcophaga and Allies, p. 77.

Two striking characters distinguish this species from the closely
related forms found in New York. It has a long slender acces-
sory plate and on the hind femur a group of striking long bristles
which are present only on the distal end of the median row.

Length. — 8 to 13 mm.

Records. — Grand Island; Ithaca; Karner; King Perry; Rens-
selaer; S.I.: Arrochar; Wadsworth; New Dorp ; New Brighton;
Watchogue; Richmond; L.I.: Babylon; Half-way Hollow Hills.
May to August. Figures 109, 110 and 111.

Aldrich (1916) and Chittenden (1926) listed the species as a
parasite of the imported cabbage worm {Pieris rapce). Phillips
and King (1923) reared latisterna from the corn ear worm.
Porter and Alden (1924) recorded a rearing from the larva of
the canker worm. Breakey (1929) gave a record of rearing
latisterna from the iris borer larva in Wisconsin. Knull (1932)
reared the fly from the pupae of Ennomos suhsignarius in Penn-
sylvania. The writer (1929) recorded on October 1, 1926, adult
latisterna flying thickly around the larvae of Anisota sp. at
Taunton Lakes, New Jersey. Although gravid female flies were
abundant no parasitism was secured. Maggots were removed
from gravid female flies and placed on dead insects. These mag-
gots developed normally. Knipling (1936) also reared this spe-
cies on meat after removing larvse from the fly.

Type. — Male, in Mass. Agri. College.

RAVINIA GROUP

The genus Bavinia was erected by Desvoidy (1863) with Sarco-
phaga haematodes Meig. as genotype. Bottcher (1912, 1913)
recognized the group only as a sub-genus. Parker (1914) re-
turned Bavinia to generic rank and pointed out that, in addition
to the frontal bristles diverging anteriorly, there was a striking
resemblance in the male copulatory organs. Aldrich (1916) and
Hall (1928) reduced the genus to sub-generic standing because as
Aldrich stated, “the characters of the frontal row is less devel-
oped in the males,’’ and, “there are many species outside the
group which show it.” Hall (1928) added the statement, “the
course of the frontal bristles range from parallel in some species

Sept., 1940]

Hallock: Sarcophagin^

221

to strongly divergent in others; all species possessing striking
similar copnlatory organs.” No additional characters have been
observed so the group is considered only as of sub-generic rank.

Sarcophaga laakei Hall

1931. Hall, Ann. Ent. Soc. Am., vol. 24, p. 181-182.

It is difficult to separate this species from the closely related
form, latisetosa. The characters given in the key will separate
the two species. This species occurs fairly common in the south
but it has been taken only a few times on Long Island, New York,
by Mr. F. S. Blanton.

Length. — 6 to 7 mm.

Records. — L.I.: Babylon. June, August. Figure 112.
Knipling (1936) reared larvae of laakei on meat.

Holotype.— No. 43264, U. S. N. M.

Sarcophaga latisetosa Parker

1914. Parker (Ravinia), Proc. Boston Soc. N. H., vol. 35, p. 63.
1916. Aldrich, Sarcophaga and Allies, p. 299.

1928. Hall, Ann. Ent. Soc. Am., vol. 21, p. 343.

The outer vertical bristles are present in this species but they
are lacking in the closely related stimulans and the posterior
clasper is much longer in latisetosa than in stimulans.

This species is very common in the northern part of the United
States and Canada but it is taken in fewer numbers south.
Length. — 6 to 7 mm.

Records. — Ithaca ; McLean ; Duck Lake ; Herkimer ; Tuxedo ;
Troy; Poughkeepsie; L.I.: Cold Spring Harbor; Babylon. June
to September. Figures 113, 114 and 115.

Aldrich (1916) states that the larvae of this species are excre-
ment feeders. Professor G. W. Herrick reared twelve adults of
this species at Ithaca from dog excrement in July 1912.
Holotype. — Male, in collection of Mass. Agri. College.

Sarcophaga Vherminieri R.D.

1830. Robineau-Desvoidy, Essai Sur les Myodaires, p. 339.

1837. Walker (anxia, querula, avida, rediviva, aspera, rahida,
acerka, comes), Trans. Linn. Soc., vol. 17, p. 818-825.

222

Journal New York Entomological Society [Vol. xlviii

1868. Thompson {8. pallinervis) , Eugenies Resa, p. 535.

1914. Parker {Ravinia communis), Proc. Post. Soc. N. H., vol.
35, p. 55.

1916. Aldrich {8. communis), 8arcophaga and Allies, p. 253.
1930. Aldrich, Proc. U. S. N. M., vol. 78, p. 13, 27, 28.

.This scavenger species is the most common Scarophagid in
New York State. It occurs in all parts of the state.

Length. — 6 to 12 mm.

Records. — Tuxedo ; Middleport ; Rome ; Trumbull Corners ;
Tear; Potsdam; IJnionport; Buffalo; Ithaca; Mosholu; 8.1. :
New Brighton; Port Wadsworth; L.I.: Flatbush; Sea Clift; Cold
Spring Harbor ; Gardiners Island ; Wyandanch ; Brooklyn. May
to September. Figures 116, 117 and 118.

Parker (1914), Aldrich (1916) and Wilson (1932) have all
recorded this species abundantly feeding and larvipositing on all
kinds of mammal excrement. Knipling (1936) found that larvae
of this species would develop when placed upon either excrement
or meat.

Type. — Museum of Natural History in Paris.

8arcop}iaga pusiola V. cl. W.

1896. Van der Wulp, Biol. Cent. Am. Dip., vol. 2, p. 278.

1914. Parker {Ravinia peniculata) , Proc. Bost. Soc. N. H., vol.
35, p. 58.

1916. Aldrich {8. peniciilata) , 8arcophaga and Allies, p. 121.
1930. Aldrich, Proc. U. S. N. M.,vol. 78, p. 33.

The superficial characters of this species are similar to
riierminieri. This species belongs to the three postsutural dorso-
central group while I’herminieri has four postsutural dorsocen-
trals. The average size of pusiola is smaller than I’herminieri.
Length. — 6 to 9 mm.

Records. — Redhook; Tuxedo; Tear; Upper Ausable; Olcott;
Ithaca ; L.I.: Cold Spring Harbor. June to September. Figures
II9, 120 and 121.

Parker (1914) recorded rearing the larvas of this species from
dung. Knipling (1936) found that larvae normally developed
upon excrement but that they could develop upon meat.

8arcophaga stimulans Walker

1848. Walker, List of Dip. Brit. Mus., p. 817.

Srpt., 1940]

Hallock; Sarcophagin^

223

1895. Van der Wulp (S. vagabunda) , Biol. Cent. Am. Dip., vol.

2, p. 270.

1900. Coqnillett {S. quadrisetosa) , Ent. News, vol. 12, p. 17.

1914. Parker {Bavinia quadrisetosa)^ Proc. Bost. Soc. N. H.,
vol. 35, p. 60.

1916. Aldrich {S. quadrisetosa) , Sarcophaga and Allies, pp. 296.

1917. Townsend (Cheetoravinia quadrisetosa), Proc. Biol. Soc.

Wash., vol. 30, p. 195.

1928. Hall (S. quadrisetosa) , Ann. Ent. Soc. Am., vol. 21, p. 346.
1930. Aldrich, Proc. U. S. Nat. Mus., vol. 78, p. 15, 29.

Although this species occurs in New York it has never been
taken as commonly as the closely related latisetosa. Hall (1928)
reported stimulans more numerous south of the Ohio River than
in the northern part of its range.

Length. — 6 to 8 mm.

Records. — Yonkers; 8.1. : Clove Valley; L.I.: Babylon. August.
Figures 122, 123 and 124.

Aldrich (1916) and Greene (1926) reported that this fly is a
very common excrement feeding species in its larvae stage. Knip-
ling (1936) reared this species from excrement but failed to
secure development of larvae upon meat.

Type. — In British Museum.

Sarcojjhaga sueta V. d. W.

1896. Van der Wulp, Biol. Cent. Am. Dip., vol. 2, p. 281.

1916. Aldrich {8. communis var. ochracea), 8arco])Jiaga and
Allies, p. 255.

1928. Hall {8. ochracea), Ann. Ent. Soc. Am., vol. 21, p. 340.

1930. Aldrich, Proc. U. S. N. M., vol. 78, p. 34.

For many years sueta was considered in America as a variety
of Vherminieri but Greene (1926) pointed out distinct differences
in the immature stages. Hall (1928) raised this form to the rank
of species. This species occurs commonly from southern Ohio to
the southern part of the United States. It has been taken on rare
occasions on Long Island, New York.

Length. — 6 to 12 mm.

Records. — L.I.: Babylon. August. Figures 125, 126 and 127.
Aldrich (1916), Lindquist (1936) and Knipling (1936) have
pointed out that the larvse of this species develop in excrement.

224

JOURNAL New York Entomological Society [Vol. XLVIII

typical sarcophaga group

The remaining species of the genus Sarcophaga are placed in
this group. Additional study may show that it is desirable to
divide this residuary group into smaller groups.

Sarcophaga aculeata Aldrich

1916. Aldrich, Sarcophaga and Allies, p. 143.

Although this species has not been recorded in New York State
it is so widely distributed in the United States and Canada that
it can be expected to occur here. According to Aldrich (1916)
this species has been reared from the adults of several species of
grasshoppers.

Holotype and allotype. — Male and female. No. 20526, U. S. N. M.

Sarcophaga alceclo Aldrich

1916. Aldrich, Sarcophaga and Allies, p. 132.

This species is close to S. excisa but it can be separated by the
distinct yellow cast on the parafrontals and parafacials, wings
slightly smoky and hind calypters light brown with a white mar-
gin. The genitalia differences are illustrated in the figures.
Length. — 8 to 9 mm.

Records. — Although this species is widely distributed in the
United States it has only been recorded in New York at Babylon,
L.I., where Mr. F. B. Blanton has taken 12 specimens. August.
Figures 128, 129 and 130.

The hosts of alcedo are not known.

Holotype and allotype. — Male and female, No. 20517, U. S.
N. M.

Sarcophaga aldrichi Parker

1916. Parker, Jr. Econ. Ent., vol. 9, p. 438.

1916. Aldrich, Sarcophaga and Allies, p. 167.

This species belongs to a small group of flies with four post-
sutural dorsocentral bristles, hind tibiae villous in the males and
the hypopygium entirely black. The males of this species may
be separated from the closely related species in this group by the
broad anal forceps with a large bare, shining area on their lower
central portion and the deep groove on the large thick anterior

Sept., 1940]

Hallock; Sarcophagin^

225

clasper. The female has black genital segments with an oval
aperture which is surrounded by dense bristles.

Length. — 8 to 10 mm.

Records. — Ithaca ; Oneonta ; Axton ; Mt. 'Whiteface ; L.I. :
Babylon; .Herscher State Park. May to the middle of July.
Figures 131, 132 and 133.

Adults of this species occur commonly around infestations of
the forest tent caterpillar {Malacosoma disstria). C^sar (1916)
reported in Ontario that 90% of the larvae of the forest tent cater-
pillar were parasitized by this fly. Parker (1916) recorded a
rearing from Porthetria dispar. It has also been reared accord-
ing to Glendenning (1914) from Stilpnotia salicis in British
Columbia.

Holotype. — Male, in collection of Mass. Agri. College. Para-
types are in the U. S. N. M.

Sarcophaga atlanis Aldrich

1916. Aldrich, Sarcophaga and Allies, p. 100.

Although this species may be confused with hunteri it may be
distinguished by the black palpi and the lack of a tuft of long
black hair which is found on each anal forcep of hunteri.

Length. — 5 to 8 mm.

Records. — McLean; Herkimer; L.I.: Islip. June to August.
Figures 134, 135 and 136.

Aldrich (1916) pointed out that this species was a parasite of
grasshoppers. Mi^ J. W. H. Rehn very kindly permitted the
writer to publish data taken during the summer of 1936 when he
secured information by roarings to show that approximately 25%
of the adults of Dendrotettix quercus P. at Mt. Misery, New Jer-
sey were parasitized by Sarcophaga atlanis.

Holotype and allotype. — Male and female. No. 20506, U. S.
N. M.

{To he continued)

226

Journal New York Entomological Society

[VOL. XLVIll

Figure 98.

Figure 99.

Figure 100.

Figure 101.
Figure 102.

Figure 103.

Figure 104.
Figure 105.
Figure 106.
Figure 107.
Figure 108.
Figure 109.

Plate VIII

Lateral view of external genitalia of Sarcophaga miser a var,
exuberans Pand.

Lateral view of external genitalia of Sarcophaga misera var.
harpax Pand.

Lateral view of external genitalia of Sarcophaga misera var.
sarracenioides Aid.

Fifth sternum of Sarcophaga misera var. sarracenioides A.

Eear view of anal forceps of Sarcophaga misera var. sar-
racenioides Aid.

Lateral view of external genitalia of Sarcophaga bisetosa
Parker.

Fifth sternum of Sarcophaga bisetosa Parker.

Eear view of anal forceps of Sarcophaga bisetosa Parker.

Lateral view of external genitalia of Sarcophaga cimbicis T.

Fifth sternum of Sarcophaga cimbicis Townsend.

Eear view of anal forceps of Sarcophaga cimbicis Town.

Lateral view of external genitalia of Sarcophaga latisterna
Parker.

SAECOPHAGIN^

228

Journal New York Entomological Society

[VoL. XLVIII

Figure 110.
Figure 111.
Figure 112.
Figure 113.

Figure 114.
Figure 115.
Figure 116.

Figure 117.
Figure 118.
Figure 119.

Figure 120.
Figure 121.

Plate IX

Fifth sternum of Sarcopliaga latisterna Parker.

Eear view of anal forceps of Sarcopliaga latisterna Park.
Lateral view of external genitalia of Sarcopliaga laalcei Hall.
Lateral view of external genitalia of Sarcopliaga latisetosa
Parker.

Fifth sternum of Sarcopliaga latisetosa Parker.

Eear view of anal forceps of Sarcopliaga latisetosa Park.
Lateral view of external genitalia of Sarcopliaga I’lierminieri
E.-D.

Fifth sternum of Sarcopliaga I’lierminieri E.-D.

Eear view of anal forceps of Sarcopliaga I ’lierminieri Eob.-Des.
Lateral view of external genitalia of Sarcopliaga pusiola

V. d. W.

Fifth sternum of Sarcopliaga pusiola V. d. W.

Eear view of anal forceps of Sarcopliaga pusiola V. d. W.

(Jour. N. Y. Ent. Soc.), Vol. XL VIII

(Plate IX)

IIT

SAECOPHAGIN.^

230

Journal New York Entomological Society [Vol. xlviii

Figure 122.

Figure 123.
Figure 124.
Figure 125.
Figure 126.
Figure 127.
Figure 128.

Figure 129.
Figure 130.
Figure 131.

Figure 132.
Figure 133.
Figure 134.

Figure 135.
Figure 136.

Plate X

Lateral view of external genitalia of Sarcophaga stimulans
Walker.

Fifth sternum of Sarcophaga stimulans Walker.

Eear view of anal forceps of Sarcophaga stimulans Walk.
Lateral view of external genitalia of Sarcophaga sueta V. d. W.
Fifth sternum of Sarcophaga sueta V. d. W.

Eear view of anal forceps of Sarcophaga sueta V. d. W.
Lateral view of external genitalia of Sarcophaga alcedo
Aldrich.

Fifth sternum of Sarcophaga alcedo Aldrich.

Eear view of anal forceps of Sarcophaga alcedo Aldrich.
Lateral view of external genitalia of Sarcophaga aldrichi
Parker.

Fifth sternum of Sarcophaga aldrichi Parker.

Eear view of anal forceps of Sarcophaga aldrichi Parker.
Lateral view of external genitalia of Sarcophaga atlanis
Aldrich.

Fifth sternum of Sarcophaga atlanis Aldrich.

Eear view of anal forceps of Sarcophaga atlanis Aldrich.

(Jour. N. Y. Ent. Soc.), Vol. XL VIII

(Plate X)

SARCOPHAGIN^

Sept., 1940]

Hatch : Silphin^

233

OBSERVATIONS ON SILPHIN^ WITH A NOTE ON
INTRASPECIFIC VARIATIONS AND THEIR
DESIGNATION

By Melville H. Hatch

The following observations on Silphinae are supplemental to
two previous papers on the same group. ^ I am indebted to Mr.
Roy D. Shenefelt for permission to study the collection of Wash-
ington State College.

Apteroloma tenuicorne LeC. — While I have taken this species
from as low an elevation as 1900 feet at Cle Elum, Washington,
and Mr. M. C. Lane has taken it at about 1800 feet at Ritzville,
Washington, it appears to be especially prevalent at somewhat
greater elevations in extreme eastern Washington and northern
Idaho. Thus I have a series of thirteen specimens taken by Mr.
Lane at an elevation of about 4000 feet on Cedar Mt. near Moscow,
Idaho, on May 25, 1935 ; and Mr. Clifford J. Burner and I se-
cured fifty or more specimens on May 30, 1937, at an elevation of
from five thousand to fifty-two hundred feet on Mt. Spokane,
northeast of Spokane, Washington. In this latter situation the
beetles occurred on the damp or wet ground under the edges of
the melting snow fields.

Silpha (Thanatophilus) tritiiherculata Kb^c — A second Wash-
ington specimen was taken by Mr. Joseph Bruzas near the Dry
Falls of the Grand Coulee, May 7, 1938, and given to me.

Silpha {Thanatophilus) lapponica Hbst. — There are, in the
Washington State College collection, two series of this species
taken in Chicago by A. L. Melander and in '‘N. 111.” by F. M.
Webster. This extends the range of this species in this direction
and makes its absence from Indiana and southwestern Michigan
more noteworthy than ever.

Silpha {Blitophaga) opaca Linn, and hituherosa LeC. — Essig^

1 Hatch, Melville H. Studies on the Silphinae. JouR. N. Y. Ent. Soc.,
XXXV, 1927, pp. 331-371.

Hatch, Melville H., and William Eueter, Jr. Coleoptera of Washington:
Silphidae. Univ. Wash. Publ. Biol. I, 1934, pp. 147-162.

2 Ins. w. N. Ainer., 1926, pp. 381-383.

234

Journal New York Entomological Society [Vol. xlviii

lists the first of these species from Alaska to California and the
second from .Washington. I have no knowledge of either oc-
curring in Washington.

Nicrophorus (Necropter) vespilloides Hbst. {defodiens Mann,
of Portevin and Hatch). — I follow Leech^ in his inability to dis-
tinguish specifically the Nearctic and Palaearctic forms of this
species and in his use of defodiens Mann., as a Pacific Coast sub-
species of the same. Leech’s notes are incorporated in the
following revised tabulation of the Nearctic forms of this species,
which should be used in connection with my original table
vespilloides Hbst. {defodiens Mann, of Portevin and Hatch)
typical form {hehes Kby., pygmceus Kby., humeralis Hatch)
ab. ruher Hatch

ab. near cticns HOY. {defodiens typical form Port, and Hatch,
nec. Mann.)
ab. nicolayi Hatch
ab. oregonensis Hatch
subsp. defodiens Mann.

typical form {nunemacheri Hatch, nunenmaclieri Leech)
ab. Mnotoides nov. {hinotatus Hatch nec. Port.)
ab. conversator Walk, {defodiens var. b of Mann., lateralis
Port.)

ab. pacificce Hatch
ab. gaigei Hatch

ab. walkeri nov. {conversator Port, and Hatch, nec. Walk.)
ab. kadjakensis Port,
ab. mannerheimi Port.^
ab. hmotatus Port.^

It should be noted that, with the exception of the typical form,
none of the Nearctic aberrations of vespilloides occur hi the
Palaearctic Region and that, moreover, the types of variation in
the two regions are very different. None of the Palgearctic aber-
rations, for instance, appear to lack an orange spot from the base

3 Bull. Brook. Ent. Soc., XXXI, 1936, p. 156.

4 Hatch, Jour. N. Y. Ent. Soc., XXXV, 1927, p. 356.

5 The anterior elytral fascia in this aberration is divided into two. I was
in error in describing it as constricted.

6 The posterior elytral spot is absent, the anterior fascia reduced to a
single spot in this aberration.

Sept., 1940]

Hatch: Silphin^

235

of the epipleuron/ which is lacking in all the American aberra-
tions except the typical form and the ab. ruher. Leech® suggests
that the subsp. defodiens extends to Japan, and the ab. sylvivagus
Reitter from that locality may be referable to that subspecies, ap-
parently resembling the ab. lateralis Port. On the other hand,
sylvivagus is, according to Portevin,® related to the eastern
Asiatic varieties, borealis Port, and sylvaticus Reitt., in which the
abdominal pubescence is yellow rather than black.

Leech’s observations^® on the habits of defodiens are of great
interest. They confirm to a suprising degree those previously
published by Pukowski^^ for a series of European species, in-
cluding vespilloides. These observations of Pukowski’s are ab-
stracted at length in English by Balduf in the Bionomics of
Entomophagous Coleoptera (John S. Swift and Co.), 1935, pp.
69-75.

Nicrophorus (Necropter) investigator Zett. subsp. investigator
Zett. and nigritus Mann. — have now seen examples of both these
forms from Pullman and the latter in addition from Spokane and
Wawawai, all in eastern Washington.

Nicrophorus {Necropter) guttulus Mots. — Of the subsp. guttu-
lus, I have a single example of the ab. vandykei Angell from
Pullman and ab. woodgatei from Port Townsend, both in
Washington.

INTRASPECIFIC VARIATIONS AND THEIR DESIGNATION^^

Intraspecific variation should be studied for the following
reasons :

(1) The fundamental obligation of taxonomy is to describe
accurately and precisely the variation presented.

7 This term is, according to Torre Bueno’s Glossary of Entomology, 1937,
pp. 92, 132, definitely superior to “hypomeron,” which I used formerly.

8 Lx., p. 156.

9 Bull. Mus. Paris, XXX, 1924, p. 375; Encycl. Ent., VI, 1926, p. 235, 259.

10 Proc. Ent. Soc. B .0., XXXI, 1935, pp. 36-40.

11 Zeit. Morp. Okol. Tiere, XXYII, 1933, pp. 518-586.

12 These remarks are supplemental to my original statement in Jour. N. Y.
Ent. Soc., XXXV, 1927, p. 341, and are largely prompted by Leech’s com-
ments on my classification of the aberrations of Nicrophorus vespilloides
subsp. defodiens Mann, in Proc. Ent. Soc. B. C., XXXI, 1935, pp. 36-40, and
Bull. Brook. Ent. Soc., XXXII, 1937, pp. 156-159.

236

Journal New York Entomological Society [Vol. xlviii

(2) What appear at first to be intraspecific variations are fre-
quently the elements out of which subspecies and species are
later recognized. The indication of such forms by one author,
though he holds them to be entirely intraspecific in nature, may
enable a subsequent author to reach conclusions of a very different
sort.

(3) Many intraspecific variations are the materials out of
which new species evolve in the course of geologic time. Their
detection and study is the first step in their consideration as fac-
tors in evolution. To argue that this is more suited to genetics
than to normal systematic entomology^^ is to adopt a wholly
untenable view of the discreteness of taxonomy from other
branches of biological knowledge. It is in important measure the
taxonomist passing in review large series of different life forms
who is in a position to discover this sort of data.

There are several ways in which intraspecific variation may
be treated.

(1) It can be entirely ignored! Every working taxonomist
can probably bring to mind descriptions in highly variable groups
Avhich mention only the supposed specific characters with never
a word as to the variation.

(2) Then there is the traditional method, which is a prevalent
procedure in America, of devoting a special paragraph to the
intraspecific variation. This method is applicable to strictly
continuous variation, where it can be supplemented by curves of
variation and other graphs. It is also useful where the variation
is vague or extremely complicated. When applied to more or
less discontinuous variation, it shows that the study of the vari-
ants is in an early pre-Linn^an stage, since subsequent references
to the variations mentioned must repeat the entire definition of
the variety.

(3) The early coleopterists lettered intraspecific variations.
Thus Illiger, in his Verzeickniss der Kdfer Preussens (1798),
and Schonherr, in his Synonymia Insectorum (1806-1817), and
LeConte as late as 1880 gave Greek letters to their varieties,
“alpha” being reserved for the typical form. The current pro-

13 Leech, l.c., p. 39.

Sept., 1940]

Hatch : Silphin^

237

cedure would undoubtedly be to substitute Latin for Greek letters,
and I have employed letters myself on several occasions.

(4) The transition from letters to names was gradual. One
author would describe a species which another author would
regard as a variety. The extra name would be retained as a
varietal name and the tendency would be to invent new names for
other varieties of similar rank.

For about half a century continental coleopterists have em-
ployed the term aherratio (English, aberration) to apply for the
most part to color varieties. Most of their valid species were
described, so they turned to the problem of intraspecific vari-
ation. Aberrations were used by Ganglbauer in Die Klifer von
Mitteleuropa in 1892 and are to be found employed in practically
every continental work on coleopterology that has appeared since
1900, but only during the past decade has this usage made much
headway in English speaking countries. Leech^® finds the term
so unusual that he puts it in quotation marks and there is an
unfortunate tendency^® in some quarters to substitute the English
for the Latin meaning of the word and thus limit it to freakish,
monstrous specimens. In reality the word is to be derived from
the figurative use of the verb aberro, ‘‘to wander, deviate, depart
from.”

There has been a general tendency to regard all intraspecific
names as being nomenclatorially on a par with specific and sub-
specific names,^^ and this is, perhaps, one of the chief sources of
the prejudice against them. Many systematists have disliked
seeing names based on trivial or supposedly trivial features on
a par with names based on supposedly fundamental characters.
There is, accordingly, some cogency for :

(5) The procedure suggested in the Entomological Code of
Banks and Caudell (1912, p. 9, sec. 37) and that of the British
National Committee on Entomological Nomenclature^® releasing

14 Tech. Publ. N. Y. St. Coll. For., 17, 1924, p. 307 ; Univ. Wash. Publ.
Biol., I, 1932, p. 100.

15 Bull. Brook. Ent. Soc., XXXII, 1937, p. 158.

16 Guilder, Ent. News, XXVIII, 1927, p. 265; Carter, Ann. Mag. Nat.
Hist., 104, 1934, p. 552.

11 This is my own preference.

238

Journal New York Entomological Society [Vol. XLVIII

aberrational names from the application of priority. The British
code would release them entirely, the Banks and Candell code
would require priority to operate within the limits of the single
species.

(6) The logical consequence of releasing aberrational names
from the operation of priority are the somewhat similar proposals
of BlaisdelP® and Heikertinger^® who suggest the use of de-
scriptive or semi-descriptive words in place of aberrational names,
these words being used over and over again in different species
even of the same genus as often as required, being purely de-
scriptive, and absolute synonyms of ordinary specific and varietal
names entirely free from the operation of priority or other nomen-
clatorial rules. Blaisdell called the category to which he applied
these names a ‘‘forma,” employed words like typica, glahra,
interstitialis, punctata, annectans, catalince, communis, emar-
ginata, horealis, etc., and accompanied them with descriptions.
He used them in connection with his monograph of the Eleodini
referred to above, and where it has become subsequently desirable
to elevate some of them to the rank of variety or species, he
considers^^ the names so used to date from the , time of their
elevation rather than from that of their original proposal.

Heikertinger calls his terms “ Kennworten, ” “recognition
words, ’ ’ and has attempted to use words of self-evident meaning,
not requiring attendant descriptions. He works out this sug-
gestion in the Halticinae section of Winkler’s Catalogus Cole-
opterotnim regionis palcearcticce (1930, pp. 1317-1352), using
terms like viridicollis, ruficollis, vitipennis, maculipennis , nigro-
suturata, latilimbata, etc. Heikertinger ’s system would appear

18 Proc. Ent. Soc. London, III (1), 1928, p. HE.

19 U. S. Nat. Mus. Bull., 63, 1909, pp. v-vi.

20 Kol. Eund, XV, 1930, pp. 213-230. In this connection it is interesting
to note the suggestion of Croneis in Science (LXXXIX, 1939, pp. 314-315)
of a series of categories paralleling the LimiEean ones for use in classifying
fossil remains whose true biological affinities are not ascertained. This is
significant as an insistence that taxonomy not neglect its basic function of
describing, classifying, and designating for the important but sometimes
impossible task of interpreting. Hubbs’ arguments (Science, LXXI, 1930,
pp. 317-319) in favor of a uninomial as opposed to a binomial system of
species designation is another suggestion looking in the same direction.

21 Pan-Pac. Ent., II, 1925, p. 77.

Sept., 1940]

Hatch : Silphin^

239

to be applicable only to relatively simple types of variation rather
than to those involving complicated color pattern changes.

(7) For complicated color pattern variations formnlse can be
used. The spots or other elements of the pattern are numbered
or lettered, various combinations of these symbols indicating the
different forms. This system was used by Johnson in Coccinel-
lidas^^ and I have myself employed it on several occasions.

(8) The ultimate stage in the nomenclature of intraspecific
variation is attained when it can be defined in terms of the
constituent genes or other hereditary or environmental factors
involved. This is possible at present for only a very few species,
like some of the Drosophila and a few of the Chrysomelidse and
Coccinellidse.

The principal requirement of a nomenclature for intraspecific
variations is that it be clear cut, unequivocal, and of such a na-
ture as to make it possible for subsequent authors to refer to the
forms described precisely without having to repeat the entire
description in pre-Linnaean fashion. Any one of the last six
methods cited is available with the second one in reserve for
vague or imperfectly understood variation. Whether one uses
numbers, letters, formulae, or names with or without priority, or
all in combination, is of secondary importance so long as is met
the initial requirement of precision.

Objections to studies of intraspecific variation are voiced on
various grounds. It is said that many variation studies, espe-
cially those involving scarcely more than the pointing out of the
existence of the several variations, are of little value because of
their superficial preliminary nature, because they are confined
to a single prominent variable character, or because no attempt
is made to correlate them with the environment or heredity of
the form under consideration. One might equally criticize the
describer of a new species because he fails to work out its life
history or genetics before going on to the next species. These
problems are important and some day will be studied by some-
body. But the task, or one of the tasks, of taxonomy is a descrip-

22 Johnson, Eoswell H. Determinate evolution in the color-pattern of the
lady-beetle, Carnegie Inst, of Wash. Publ. No. 122, 1910, 104 pp.

23 Jour. N. Y. Ent. Soc., XXXV, 1937, pp. 347-348 ; Univ. Wash. Publ.
Biol., I, 1932, pp. 98-99.

240

Journal New York Entomological Society [Vol. xlviii

tive survey of the animal kingdom — a survey that would be indefi-
nitely delayed if each investigator attempted to see each of his
problems through to an ultimate ecological or genetical conclu-
sion. It is stated, again, that such studies are ‘ ‘ genetics ’ ’ rather
than “normal systematic entomology”! They are criticized
because they are of no use to the economic entomologist.^^ Ap-
parently “normal systematic entomology” may “make up” to
the economic entomologist, but must “shy off” from the
geneticist !

One of the main results derived from designating a form an
aberration or color variety or forma is that it is thereby almost
certainly removed from the attention of the economic entomologist
and the general ecologist,^® except as they find it convenient not to
be led astray by extreme phases that the species may assume,
melanistic, immaculate, or depauperized forms that might at first
be mistaken for distinct species. The classification of aberrations
is usually of no more significance for such persons than, for in-
stance, the designation of sex; in fact, it may be of far less im-
portance, since the recognition of the sexes is of practical concern
in many experimental procedures. Thus, in citing an insect like
the asparagus beetle, Crioceris asparagi L., for the Insect Pest
Survey Bulletin, it would be absurd to give more than the species.
The numerous aberrations of the asparagus beetle^® and other
species are available for those who need or are interested in such
matters ; they may be overlooked by others.

A more cogent objection to the validity of studies in intraspe-
cific variation is that such variations may intergrade; they may
intergrade in series, the right and left side of the body may exhibit
different variations, different variations may occur in the progeny
of the same pair of parents. A good deal of the force of this
objection is due to a misapprehension. The principal difference
between species and intraspecific forms of all sorts is that the

24 Hopping, Proc. Ent. Soc. B. C., XXXI, 1935, p. 34.

25 The person making a detailed ecological study of a single species may
be very much interested in aberrations, especially if any of them prove to
have an origin that is immediately environmental.

26 See Hatch, Bull. Brook. Ent. Soc., XXII, 1927, p. 211; Univ. Wash.
Publ. Biol., I, 1932, pp. 72-74, for some account of these aberrations and
their literature.

2" Leech, lx., p. 158.

Sept., 1940]

Hatch: Silphin^

241

former are characterized by an absence of intermediates, at least
in theory. If the intraspecific variations were marked off as
distinctly as it is intimated they should be, they would not be
intraspecific variations but species.

The matter, then, comes to this : Is it worth while to attempt to
distinguish characteristic phases of a more or less continuously
varying series? And, if we do, what is to prevent us from run-
ning off into non-significant ^‘unnamables” and individual vari-
ants? The question of ‘‘unnamables” in Lepidoptera has been
discussed by Guilder^® and Forbes^^ and should offer little diffi-
culty in the long run. Unless the causes of the variation have
acted acted from within the living organism,®® the forms produced
are clearly of little significance®^ for the student of intraspecific
variation. Moreover the variation must be of such a type that it
occurs or probably occurs in series. Otherwise it is ‘ ‘ individual ’ ’
or ‘Mnnamable.” Even as regards size, it is sometimes con-
venient to designate minor or depauperized specimens, especially
when this is accompanied by the specific characters in an en-
feebled form.

Leech (?.c.) specifically questions the utility of distinguishing
four stages in the disintegration of the anterior elytral fascia in
one of the Pacific Coast forms of Nicrophorus. In 1927 I de-
scribed these and other aberrations of this species on the basis of
series too small to show their nature. In 1934,®® in a series of
178 specimens' from western Washington, I showed that these
four stages form an approximately normal curve of variation with
the apex of the curve close to one of the intermediate types. Leech
could have continued the analysis in other portions of the range.
He preferred, however, sim^^ly to express the view that such a
studjq based on color pattern variation alone, was without signifi-

28 Ent. News, XX’XIX, 1928, pp. 201-204, pi. VII-X.

29 Bull. Brook. Ent. Soc., XXIX, 1934, pp. 65-67.

30 By which I do not mean to limit the causes to hereditary ones. En-
vironment may, for instance, produce melanism, which I would tend to
regard as ^ ‘ significant. ’ ’

31 1 prefer ‘^significance” to “ nameability ” because, throughout the
present discussion, I maintain that whether a variant is ‘ ‘ named ’ ’ or merely
designated by a number, letter, or formula is of no importance.

32 Univ. Wash. Publ. Biol., I, 1934, p. 158.

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Journal New York Entomological Society [Vol. xlviii

cance, and a large number, perhaps the majority, of American
coleopterists would agree with him.

The principal objection to studies of intraspecific variation is
not directed against the studies as such but against the naming
of the variations. As has been noted, the author’s contention is
that whether or not the variants are named is a matter of sec-
ondary importance, so long as they are precisely designated.
This, however, is not the view of opponents of names of this class.
They consider the so-called ‘ ‘ cluttering up ” of the literature with
such names an olfense than which scarcely any other is greater !
as though naming a variant could give it any importance and
dignity it did not have before ! Names are merely combinations
of letters — symbols for designating biological concepts. They are
symbols, however, that, because of their great associational pow-
ers, are so much more convenient than letters or numbers or
formulas that the tendency to use them for this class of variation
is very great. They, varieties, color varieties, aberrations, forms
— the precise term applied is of little moment — named or merely
designated — should be taken at their face value for exactly what
they are — categories intermediate between the subspecies and the
individual.

Where then are we going to stop ? Lacking data derived from
genetical experimentation, we must depend on our judgment.
The benefit of the criterion of the presence or absence of inter-
mediate specimens is denied to us, since all intraspecific variations
are subject to such a connection. Forbes {l.c.) suggests that if
we can recognize 90 per cent of the specimens from a given area
as belonging to a given geographical race the race is valid. Per-
haps that or some other percentage will serve for the recognition
of aberrations. The matter is really akin to that of genera and
subgenera. As many are recognized as are convenient. And the
number, as with the higher taxonomic categories, is almost certain
to increase as more and more extensive series of specimens are
passed in review and more and more precise methods of study are
employed.

There seems to be concern in some quarters that a continued
accumulation especially of named varieties, aberrations, etc., will
involve the literature in hopeless confusion. Such a fear is

Sept., 1940]

Hatch: SiLPHiNiE

243

groundless. The matter is self-correcting, just as is the over
multiplication of categories at any of the other systematic levels.
Only such designations persist as subsequent authors find recog-
nizable and useful. Authors who do not care to pay attention to
non-geographical intraspecific variations ignore them, treating
such names as may have been proposed as absolute synonyms,
which, indeed, they truly are, from their point of view. Other
authors, who are interested in such categories, recognize as many
of their predecessor’s “designants” as the nature of the charac-
ters cited in the original descriptions or inherent in the types,
where types are available, and the nature of the organisms them-
selves permit — exactly as with categories of other types. It is
probably too much to expect that there will be any cessation of
vituperativeness between persons holding different views !

The whole matter may come down to personal preference. But
the groiying continental usage would seem to indicate that, as the
beetle fauna becomes better and better known, more attention is
directed to intraspecific variation.

Studies of these intraspecific categories may be largely over-
looked by the economic entomologist and the ecologist, and must
be overlooked by the naturalist whose aim is simply to get into
his cabinet a ‘‘set” of two, six, or any other fixed number of
specimens of each species. They are, however, almost inevitable
for the student who desires to study his material with some regard
for its complexity — an inadequate forecast of the time when
taxonomists will analyze their species genetically.

SUMMARY

It is held that intraspecific variants, such as varieties, color
varieties, aberrations, forms, etc., should be as precisely described
and designated as the material will permit. Whether this desig-
nation is accomplished by means of numbers, letters, formulae,
or names with or without priority is secondary. The author pre-
fers to allow the principal of priority to operate as generally as
is conveniently possible, but this likewise is a secondary con-
sideration.

I must not close this discussion without acknowledging the
invaluable critical assistance I have received in its preparation

244

Journal New York Entomological Society [Vol. xlviii

from two of my former students, Dr. Donald L. Frizzell and Dr.
Harriet Exline Frizzell, especially the former, neither of whom,
however, are responsible for any of the opinions expressed herein.

A NEW RECORD FOR CONNECTICUT

Along a wooded road near Sharon, Conn., on June 16, 1940, a
small skipper, Carterocephalus palmmon, race mesapano Scndder
was discovered. This is a rather nnnsnal record for this butter-
fly which hardly ever ocnrs south of the Adirondacks or the
White Mountains. So far as is known, this is the one and only
instance that it has ever been found in Connecticut. — Leonard J.
Sanford.

Sept., 1940]

Blackwelder: Taxonomy

245

SOME ASPECTS OF MODERN TAXONOMY*

By Richard E. Blackwelder

It should not be nepessary to define for the readers of this jour-
nal the word taxonomy. Perhaps we have not each taken the
trouble to think out what it means to us, but we nevertheless un-
derstand it well enough. I was therefore somewhat surprised
recently to find out that I was attaching to the word in my own
thoughts certain meanings which it did not hold for some other
taxonomists to whom I talked. So I desire to make it clear that I
am now using the terms taxonomy, classification, and systematics
as absolutely synonymous, since they all inherently refer to the
science of arrangement or classification.

The study of taxonomy in its broadest sense is probably the
oldest branch of biology or natural history as well as the basis for
all the other branches, since the first step in obtaining any knowl-
edge of things about us is to discriminate between them and to
learn to recognize them. It is therefore natural that for the first
one hundred years or more of the existence of this branch as a
science, it was concerned primarily with the segregation and
recognition of species. The direct results of this trend are some-
times underestimated, for they include the following among
others : First, taxonomists made known some idea at least of the
tremendous number of organisms that exist or have existed upon
the earth. This fact has had a very great influence on some of
man’s long-established and much-cherished conceptions. In par-
ticular, it gave a very great impetus to the warfare of science with
theology. Second, the knowledge accumulated by the taxonomist
was the principal basis for Darwin’s ‘‘Theory of Evolution.” It
is not necessary for me to point out the tremendous affect on
human thought which was produced by that series of studies.
Third, taxonomists have built up an enormous mass of knowledge
which is in some degree classified and available. The system is
far from perfect, we must admit, and many of its parts are of
very inferior quality, but, nevertheless, the general pattern has
stood the test of time and has proved its usefulness. Fourth,

* Delivered before the New York Entomological Society, December 12, 1939.

246

Journal New York Entomological Society [Vol. xlviii

taxonomy lias furnished to other biologists many of the funda-
mental questions to which they are seeking answers. The geneti-
cists and experimental biologists spend much of their time on
problems which are fundamental to taxonomy today and yet were
recognized because of the implications of studies on classification
and evolution.

We might multiply these examples at some length, but surely
this is sufficient to show that taxonomy during its early years was
a worthy subject for research, and that it did actually contribute
greatly to the advance of knowledge in many fields.

As I have said, there was a time when taxonomists made large
contributions to scientific thought through the mere naming and
cataloging of species. As the other branches of biology devel-
oped, the relative importance of this study of faunas and floras
diminished, not because fewer people worked at it, not because
there was less work to be done, not because there were fewer con-
tributions to scientific thought to be made, but simply because the
growth of these other branches was more rapid, and they sur-
passed it in popularity and possibly in the quality of the work.
We must then consider whether taxonomy is no longer capable of
contributing to biologic science, whether it has lost its position of
fundamental importance among all the branches of biology.

In a practical way, then, let ns outline the place of taxonomy at
the present time. Consider the plight of experimental biologists,
geneticists, students of geographical distribution, stratigraphers,
and workers in the applied natural sciences if they had no means
of recognizing and recording the various species with which they
deal. It would be absolutely necessary for them to develop for
themselves a system that would serve them and they would
therein become systematists. How could the tremendous mass of
experience be recorded and consulted without the taxonomists to
name, identify, and classify the organisms with which all the
others deal. A few examples should not be out of place here.
Many problems of the geologist have been solved at least in part
by the taxonomist. Properly used, the distribution of animals
can aid in the problem of the distribution of ancient land masses
and the seas of past geologic eras. Stratigraphy, or the correla-
tion of rock formations, depends in large part on the recognition

Sept., 1940]

Blackwelder: Taxonomy

247

and classification of fossil organisms. The physical anthropolo-
gist, with his problems of human races and human origins, is
merely a specialized taxonomist working in a very limited field.
Many problems of the geneticist, such as the recognition and
separation of mutants and the experimental unfolding of the
processes of evolution, are really specialized problems in taxon-
omy. I have not mentioned the sciences of comparative anatomy,
embryology, and evolution which are so closely interwoven into
taxonomy that frequently they must be considered to be part
of it.

There remains a branch of biology which is of unusual interest
to many entomologists. This is applied biology. Just as the dis-
crimination of species is the basis of systematic work, it is also the
starting point of many of the problems of the economic entomol-
ogist. If an insect pest be discovered, taxonomy tells us whether
it is native or introduced, what its natural enemies are, where
they will be found, what its normal distribution is, and many
other things. Without taxonomy how could w^e have biological
control, which depends upon identification of both the pest and
the parasite or predator.

We have then several aims in systematic biology which are also
our basis for a claim to recognition as one of the important sci-
ences. It is our aim first to name and describe species so that they
can be recognized and referred to, so that we may study them and
catalog them, and assemble data about them. It is our second aim
to discover through any means at our disposal the facts of descent
and blood-relationship between species. And our third aim is to
arrange these facts and these species into a classification or
scheme of arrangement which will express as nearly as can be the
relationships and the lines of descent of these species.

It is obvious that if we merely asemble a multitude of data con-
cerning a multitude of species, we ourselves, not to mention scien-
tists in other fields, will be unable to comprehend the larger im-
plications of our discoveries, the interrelation of them, and their
vast significance in related studies unless we arrange them in
some sort of order, one based on some fundamental concept to
which all are related. The concept which we have used for this is
the theory of evolution, the assumption of the community of

248

Journal New York Entomological Society LVol. XLVIII

descent of organisms. We will return to this subject of classifica-
tions after considering the means to be employed in fulfilling the
aims we have outlined.

In the study of the relationships of insects, which we call tax-
onomy, data from various sources have been used. Among tlies'e
we find characters of morphology, of geographical distribution, of
geological distribution, of genetics, of ontogeny or development,
of ecology, of physiology (which includes chemical and serolog-
ical studies), of host-parasite relationships, of teratology (or de-
formities), and of experiment. All of these have been used in de-
fining species or in building classifications, but by far the greater
number of species and classifications are based entirely upon the
first one — morphology. This is at it should be, since it has been
found that none of the others yields as readily as complete a pic-
ture of relationships as does structure. The other fields are em-
ployed in special cases in which morphological data are not suffi-
cient. There is a real danger, however, that taxonomists will for-
get that morphological data do not invariably furnish us with
the complete picture. We should remember that situations can
easily arise in which data from the other fields can be properly
and usefully utilized to supplement the characters of morphology.

Still more important, however, is an understanding of the in-
escapable fact that the taxonomist is absolutely bound to consider
all the data that may be of value in whatever problem confronts
him. If we had, for example, spread out before us, all the species
that exist on the earth of a certain group of animals, it would suf-
fice for some purposes to select one or two characters that distin-
guish each from its fellows, and consider that these were enough
to make the species known. In actual practice, however, we can
never say with certainty that we have all the species that exist
before us and the best of us will often not be able to anticipate
what the missing ones will be like. It therefore becomes neces-
sary for us to record all the characters of each species that can be
of use in separating it from the others (whether known or not).
In most cases and particularly in the case of categories higher
than species, a careful study of comparative morphology of the
group will show definitely which characters are of sufficient con-
stancy to be used for separating the known groups, and these are
the ones which we must record.

Sept., 1940]

Blackwelder: Taxonomy

249

The ideal, then, is to record in each instance all the characters
on the insect which the stndy of comparative morphology shows
to be significant for the systematic work at hand. It should not
be necessary for me to point out how far from this goal nearly all
of onr taxonomic work today really is. But inasmuch as the com-
monly accepted standards do not come even close to fulfilling the
above requirement, let us pry a little into the reason for this
failure.

During the early years of the study of taxonomy the investiga-
tion of insects was strictly limited by the equipment available.
Microscopes such as we use today were unknown, and technique
of various sorts had not been discovered. Many of the early tax-
onomists strived toward the goal of recording all the characters
that they could see, and, because of the limiting factors mentioned
above, a certain standard of description was gradually set up.
Taxonomists became used to these standards and when better ap-
paratus and techniques were devised they were not commonly util-
ized. The result is that the large majority of our systematic
work today is at exactly the level of much of the better work of
100 years ago. For example, it is difficult to find among modern
studies on Coleoptera one which can stand comparison with that
of Erichson in 1840. This is not an exceptional case, for the ma-
jority of large groups of insects are being studied today in exactly
the same way and in only slightly greater detail than they were
50 or 100 years ago.

Two things seem to be responsible for this situation. The first
is our reticence to change our methods of study. It is not pos-
sible to examine the intersegmental membranes of the abdomen
of a beetle, or the waxy capsule of a scale-insect, or the muscula-
ture of any insect, or the minute dermal organs which are so
commonly present but so seldom seen, or any of hundreds of other
structures, without subjecting the insect to some preparation or
technique which is not commonly employed. Yet these structures
are of great significance in nearly every group in which they have
been studied and often are clearly worthy of consideration in our
scheme of classification. We have simply been content with the
established way of doing things and have forgotten our ideal of
constructing a classification that will reflect relationships and
which, therefore, must utilize all possible characters.

250

Journal New York Entomological Society [Vol. XLVIII

The second factor which has held our descriptions down to the
old standards is the lack of knowledge of comparative morphol-
ogy. Is there a single large order of insects in which the funda-
mental nature of all the parts has been worked out and in which
the variation of each character throughout the group has been
determined ? It is certainly not so in the beetles, one of our best
known orders in some respects. If it is true of any order the
fact has escaped me. Several smaller groups, such as the Coc-
coidea, do come near to this ideal.

Ten years ago it was not known that the order Coleoptera con-
tains two radically different types of thoracic structure. Even
at the present time there are numerous questions of homology
that have not been settled in this order or any of the others.
If we should look at a complete series of drawings of the morpho-
logical details of a beetle, I think you would be surprised at the
number of structures which have not been used in classification,
and yet this condition is typical of nearly all the orders.

Let me recall to you the ideal which I mentioned before for
finding the characters we are to use in reconstructing the rela-
tionships of species. We must use all the characters which the
study of comparative morphology shows to be significant. We
are exceedingly far from that ideal.

In addition to the segregation and description of species the
taxonomist should be engaged in something much more far-reach-
ing in science, something which will be more universally accepted
as a truly scientific endeavor. This is classification. It is the
arrangement of species into groups to show their relationships.
The groups are then brought together into larger groups and so
on, until the degree of relationship is expressed by the category
in which union takes place. In spite of the fact that the organ-
izing of our data into systems or classifications, into a form which
will make it useful in other branches of science, is much the most
important part of taxonomy from the point of view of the ad-
vance of knowledge, entomologists have been very slow to make
contributions of this sort. If a certain group of animals, for
example an order, has been so thoroughly studied that a com-
plete classification is available, with definitely known and de-
scribed categories throughout, all we would need to do upon the

Sept., 1940]

Blackwelder; Taxonomy

251

discovery of a new species would be to describe it and place it in
its proper genus. This would automatically place it in the rest
of the system. In reality this is the way in which many taxon-
omists appear to work. The assignment of a new species to a
genus is taken to be sufficient indication of its relationships. I
venture to say that there is not a single large order of insects in
which there is more than the flimsiest classiflcation in the sense
that I have tried to give to that word.

Let me illustrate with an example from my own field, the
StaphylinidaB. In the very large subfamily Aleocharinse there
has been a large amount of work done by several profilic taxon-
omists. They have established more than 1000 genera, which
contain over 7000 species. New species are being added con-
tinually and each is being placed by its describer in what he
deems to be its proper genus. There is nothing unusual in this ;
it is being duplicated, perhaps on a smaller scale, in many other
groups of insects. However, if we take the trouble to probe
more deeply, we may be surprised to find that not a single one
of these 1000 genera has ever been adequately described, and
many of them not described at all, being based merely upon
known species. And further, when we examine other groups we
find that this is not an exception drawn from a badly neglected
group but is in fact the ‘‘ normal’ ’ condition, or at least the
“usual” one. In a family which is as well known as the Coc-
cid^, the scale-insects, a study of the genera is even now being
published that will for the first time enable us to make generic
assignments with certainty. The large amount of taxonomic
work which has been done in this group and its great economic
importance would have led us to imagine that its classification
must by now be on a firm basis, yet a recent article on the sub-
ject states: “. . . the student of the Coccoidea ... is forced to
wander in a maze of generic names the application of most of
which can not be determined from the existing literature.” If
this is true in the Coccidse, where can we find a group which can
claim to be better known in these respects. Certainly not any-
where in the Coleoptera.

Hoav can we have any confidence in the validity of the horde
of new species that are described each year if we know in ad-

252

Journal New York Entomological Society [Vol. xlviii

vance that their generic assignments are based entirely upon the
author’s conception of the genus to which he assigns it. How
often is the same species described in different genera by dif-
ferent workers, merely because the genera have never been firmly
established? As one would expect, it happens with great fre-
quency in groups of wide distribution and accounts for a large
part of our overburdened synonymy.

You may think that I have painted a very dark picture. But
it is a picture of a condition which exists and which will con-
tinue to exist until taxonomists take the necessary steps to cor-
rect it. I am glad to be able to say that there is evidence of in-
creasing realization of the seriousness of this condition, and there
are an increasing number of attempts to help correct it. I have
mentioned the study of the genera of the Coccidae, and we may
note also an increasing number of studies of genotypes and of
groups of genera. There have even been a few studies on the
principles and bases of higher classification. This is the track
that we must follow if taxonomy is not to continue to merit the
bad reputation it has acquired among biologists. We must be
more than mere describers of new species and lawyers on arbi-
trary points of procedure.

There is one more point that should be mentioned in this
regard. A division of labor is not the solution of this problem.
It will not suffice for us to describe new species ad infinitum and
leave the classification to someone else. We cannot possibly claim
to know that a species is new unless we know definitely where it
belongs in our classification. And we will have a hard time to
justify our labors to science in general if we do not complete our
work so that the results are available for others to use.

If I have succeeded in convincing you that our taxonomy has
fallen far short of its ideal and that we come close to deserving
the scorn of our fellow biologists, if I have established in your
minds the idea that taxonomy must be more than the mere de-
scription of new species, then you will ask what is to be done and
how can we do it. My answer is, of course, that we must make
classification a major part of our work; we must arrange our
knowledge, as our species, in a system which will express what
we have been able to discover, by all means in our power, of their

Sept., 1940]

Blackwelder : Taxonomy

253

interrelationships, of their origins and potentialities, and conse-
quently of their evolution. This is the logical goal of systematic
work and one which is in all respects fully worthy of onr en-
deavors. If we can attain this goal we will find that most of our
other problems in taxonomy, such as the identification of species,
will be solved as by-products of the major problems. Let ns see,
then, what a classification is and how it is made.

A classification is a grouping or arrangement of things with
regard to some group of attributes. We can classify insects ac-
cording to their food, their place or means of living, their distri-
bution in space or in time ; or by their structure. In taxonomy
we are interested primarily in a classification based upon amount
of similarity in structure because we believe that this will give
us the truest and most complete picture of lines of descent and
degree of relationship.

There are two principal methods of recording a scheme of
classification. They have different uses and different advantages
and disadvantages. The first is a purely linear arrangement.
We place the most primitive at the first of our list, next to these
the ones which resemble them most, then the next, and so on to
the most highly specialized. Our arrangement is rather arbi-
trary because one group must follow after another. Relationship
can be shown with only two other groups, the one preceding and
the one following, and it is not possible to give any indication
of the degree of relationship, the amount of similarity. This
system is most commonly employed because it is readily adapted
to printing. Examples are to be found in all our catalogs and
check-lists and all our textbooks of taxonomy.

The other system of recording a classification is by means of a
branching arrangement, usually called a ‘ ‘ tree. ’ ’ There are two
kinds of trees in use. The most familiar is that used in paleon-
tology to indicate the relationships of animals in time. As one
passes down the time-scale the various groups merge to form a
tree which indicates in some measure the degree of relationship,
the time at which the separation of the two groups occurred, the
number and proximity of related groups, the lines of descent,
and perhaps even the ancestors of each group. This tree is gen-
erally based upon very meager information, but is useful and illu-
minating in proportion to its accuracy.

254

Journal New York Entomological Society [Vol. XLVIII

The type of tree which can help us most is one which we see
very seldom in entomological literature. It is not of exceedingly
great importance in itself but serves several very useful pur-
poses— or might serve them. Its preparation would crystallize
and demonstrate many of the broader aspects of classification
in the mind of the creator of the tree. It would enable other
entomologists as well as other scientists in general to see the
results of the detailed work of taxonomy. This type of tree is
in a strict sense ‘ ‘ a classification. ” I do not want to start a dis-
cussion of what a species is, so let us take the word species as
each of you would define it for yourself. Among these species
those which have certain characters in common we group to-
gether into a genus ; genera which are more like each other than
they are like still others we group together into a family ; and so
on through orders, classes, phyla, and kingdoms. This is a classi-
fication. If we examine simply the species, they appear in a
linear arrangement but the other categories can show us the
degree of interrelationship and can bring this meaningless series
of species into a system in which each is related by one degree or
another with every other.

Such classification as this gives us a broad picture which in-
cludes not only our own species but all the others as well, relating
them to each other and to the entire scheme of life and of evo-
lution. Such a classification can help to demonstrate to us in-
consistencies in our use of such categories as genus and species,
it can demonstrate the need for a real understanding of the
higher categories and the limits of each.

It is obviously not practicable to draw diagrams of this sort
for all the groups of insects. But it is not the diagram that is of
value so much as it is the idea of relating our groups to each
other by means of successively higher categories. For example :
In one subfamily of the Staphylinidse we have four readily
recognized groups which have all been named as tribes, thus :
Xantholinini, Staphylinini, Xanthopygini, and Quediini. There
is no question that these groups exist and that they are more like
each other than any of them is like any of the other subfamilies
of the Staphylinidse. At first glance then it would seem to be
adequate to rank them all as tribes as has been done heretofore.

Sept., 1940J

Blackwelder : Taxonomy

255

However, careful examination of many characters shows that the
Xantholinini differ more from any of the others than they do
from themselves. Some writers, recognizing this, have made it
a separate subfamily, but I have already pointed out that it is
more like the rest of the subfamily Staphylininae. If we separate
it, we obscure the fact of its similarity in subfamily characters,
and if we make it a tribe we obscure the fact of its greater
divergence.

What is really needed in this case is another category between
subfamily and tribe — we may use supertribe. This is what we
get then : The subfamily Staphylininae composed of tAVO super-
tribes, the Xantholinina and the Staphylinina. The Xantholinina
contains only a single tribe, the Xantholinini, but the Staphy-
linina contains three tribes, the Staphylinini, the Xanthopygini,
and the Quediini. Now we are able to see how much each of these
groups resembles the other and how great the degree of difference
is in each case.

The point I wish to show here is that there is much more to
taxonomy than the segregation and naming of species. These
should be used merely as tools to enable us to handle the groups
Avhile Ave combine and arrange them into a scheme Avhich Avill
show their interrelationships. How are we to knoAV Avhether a
particular group of specimens is a species unless Ave knoAV in just
what degree they differ from or resemble the numerous other
groups we call species? We must knoAv more than just the tail-
end of the scheme of CA^olution if we are to be able to say that a
group of individuals is a species, a subspecies, a genus, or some
other category.

The study of classifications Avill not, of course, be a cure-all for
our problems of speciation, of descent, or of relationship, but it
will go a long way toward giving us an understanding of hoAV
groups relate to each other, of what rank should be assigned to
each group, and of AAdiat the course of evolution of these groups
has probably been.

It is more the attitude of mind which is important, and \Adien
new facts are brought to light which affect our established system,
we must not only be prepared to accept them for what they are
worth, but Ave must have a background of thought Avhich will en-

256

Journal New York Entomological Society [Vol. XLVIII

able ns to see how they affect our ideas. To many a taxonomist
the idea that he has consistently misused a category such as a
genus or a species, giving it a higher or a lower rank than is con-
sistent with the facts, is something that he is utterly unable to
understand. The standard that he uses is so firmly fixed in his
mind as the true standard of that category (let us say of a spe-
cies) that he cannot believe that any facts could upset the
standard.

On the other hand we have a taxonomist who is used to think-
ing in terms of a more or less complete classification, where the
categories indicate the degree of known relationship. If evidence
is discovered that shows that his species are in reality only sub-
species (or races), he is not so likely to object because the groups
are still intact and usable but are simply changed slightly in the
system and a further opportunity is opened for showing the
development of the system.

This subject of classification is a very broad one. It has rami-
fications in many directions. I have touched on certain phases
only, without intending to give these an exclusive claim to im-
portance. But let me summarize some of the principles that
should be basic in the study of taxonomy. First of all, we must
consider all the available data. This means that we must use
characters from whatever field of science we can; in the case of
morphology, which is most important, we must aim to use all the
structures on the insect, whether they are obscure or hard to ex-
amine or little known, and our only guide here shall be that com-
parative studies must show them to be of value for what we are
trying to do. This principle, then, demands a complete knowl-
edge of morphology and homology as basic to any study of sys-
tematics.

The second principle is this. Kegardless of other considera-
tions, we must use methods of study or procedures in working
that will give the most complete and accurate results. We must
not let our methods depend on the habits of the past. The im-
portant thing is not to follow any set procedure, but to treat
every case on its merits and requirements and to employ every
means possible to arrive at the complete truth. For example, if
we find an important character on an insect that requires dissec-

Sept., 1940]

Blackwelder : Taxonomy

257

tion of our specimens, we cannot neglect to dissect them if we are
to be worthy of the name of scientist. Or if we should find that
facts from genetics, ecology, development, or paleontology are sig-
nificant, we must not fail to consider them adequately.

Our third basic step is to carry our work on to its logical con-
clusion, at least as far as taxonomy is concerned. If we merely
describe species and genera and are not able to show exactly how
they relate to other known species and genera, where they fit into
the whole structure of evolution, how they add to our knowledge
of the whole scheme of life, we will have missed our principal op-
portunity to make real contributions to science. We speak of the
species that are “known to science,” yet how many of these are
really known in any sense beyond the publication of a name
and description ? Many are, of course, but not a very large per-
centage of the enormous numbers named. The placing of a spe-
cies in a genus, and the genus in a family, without very careful
consideration of the foundations of these groups, adds little to the
general picture of relationships which we are attempting to set
up.

This brings us to the fourth, and in some ways the most impor-
tant, principle. We must make our results available for other
scientists to see and understand. Endless series of new species
and new genera not only do not aid the workers in other branches
of science but actually serve to make taxonomy appear to them as
meaningless and purposeless. I’m sure you will agree that this
is the condition in which we find ourselves today, for many other
biologists have no conception of the part which taxonomy can
play in science and are inclined to look upon taxonomists merely
as egotists trying to attach their own names as authors of ueAv
species. Taxonomy was once much more than that and can be
again, but we will have to enlarge our horizons, raise many of our
standards of thinking, and make our results available and useful
to the rest of science.

258

Journal New York Entomological Society [Vol. XLVIII

ATRYTONE LOGAN EDWARDS

Another record for Sharon, Connecticnt, is a pair of Atrytone
log an Edwards caught July 21 and 22. These two specimens as
well as the Carterocephalus palaemon race mandan were caught
by the author and are now in the collection of The American
Museum of Natural History, New York City. — Leonard J. San-
ford.

Sept., 1940]

Breland: Parasites

259

SOME PARASITES AND HYPERPARASITES OF THE
CECROPIA MOTH^

By Osmond P. Breland
The University of Texas

When parasitic insects are bred from a lepidopterons cocoon
or chrysalis, it is too often concluded that these insects are para-
sites of the lepidopteron, without much thought of hyperpara-
sitism. Thus any worker interested in the specific parasitic reac-
tions of a particular group of insects is prone to regard with sus-
picion reports which simply state that a certain insect is a para-
site of a certain moth or butterfly.

This study was undertaken in the hope of indicating some of
the possibilities that may be obtained from one of the larger moth
cocoons. The writer of course realizes that a study of this kind
from a limited locality, in which only the end results are ex-
amined, is far from being complete. However, it is hoped that
some of these results may be suggestive. Some of this work was
done at the North Dakota State College.

So far as the writer could determine, the most thorough recent
work on the parasitism of the cecropia moth was done by Marsh
(1934). Parts of this investigation have been published (1936)
and (1937).

The cecropia moth cocoons which were used in this study were
all collected at Brooklyn, New York, by Mr. J. H. Cohen of 1532
Sterling Place. During the summer of 1937, some parasites bred
by Mr. Cohen were examined, so he was really responsible for
the writer’s initial interest in the problem. Three lots of co-
coons were received from Mr. Cohen : one lot collected November
11, 1937, one collected March 19, 1938, and the third lot Decem-
ber 15, 1938. The first group contained 113 cocoons, most of
which were supposedly parasitized. Mr. Cohen stated that in

1 The writer Avishes to express sincere appreciation to the following men
for assistance in determining some of the insects mentioned in this article :
Mr. C. F. W. Muesebeck, Mr. A. B. Gahan, who determined the chalcidoids;
Mr. E. A. Cushman, who determined the ichneumonid, and Mr. D. G. Hall,
who determined the Diptera.

260

Journal New York Entomological Society [Vol. XLVIII

procuring this material he handled 228 cocoons and determined
the parasitized insects by shaking the cocoons. The second lot,
which were selected for parasitic Ichneumonidae, contained only
20 selected cocoons. In obtaining these, 250 cocoons were ex-
amined. The third group consisted of 40 cocoons, most of which
were parasitized.

In most instances, a parasitized cecropia cocoon is somewhat
lighter in weight than a healthy one. Parasitized cecropia larvae,
after they spin a cocoon, will in many cases become hard and dry,
and break into several parts. Thus, the cocoon will rattle if it is
shaken. Actual dissection of cocoons indicated that one can de-
pend upon the “shaking test” to a very high degree in distin-
guishing between parasitized and healthy cocoons.

In obtaining the following results, all the cecropia cocoons were
opened, and the parasites examined. The parasitic larva© and
pupae were then placed in watch glasses in the laboratory until
they emerged as adults. Since these insects were kept in the lab-
oratory at a fairly constant temperature, they presumably
emerged earlier than they would have in nature.

It has been thought best to center this discussion about several
of the insects which were primary parasites of the cecropia, and
which also served as hosts for some other parasite.

Pseudogaurax anchora (Lw.) (Chloropidae)

This insect was by far the most prevalent parasite of the
cecropia cocoons examined. Out of 100 cocoons upon which ac-
curate records were kept, this dipteron occurred in 54. The
larvae occurred within all parts of the dried tissues of the host.
The number of parasites present varied from only a few in some
cases to 81 in one cocoon. Many of the dipterous larvae were
dead when discovered, and many were themselves parasitized.
Possibly because of laboratory conditions, and because of the
heavy parasitism by a small chalcidoid, only one adult specimen
was obtained from the first shipment of cecropia. Several adults,
however, emerged from the second and third groups. It is inter-
esting to note that Kaston and Jenks (1937) report this same
species as a parasite of spider egg cases.

The chalcidoid parasitizing the larvge of Pseudogaurax anchora

Sept., 1940]

Breland: Parasites

261

has been determined by Mr. Gahan as a new species of Pleuro-
tropis (Enlophidae). This insect is an internal parasite, and is
difficult to discover until the host forms a puparium, or just before
the puparium is formed. The pnparia, although small, may easily
be dissected without injuring the parasites to any great extent,
by partially burying the pnparia in household cement, and allow-
ing this to dry. Dissecting needles may then be used to pull
away the puparial skin. The number of parasites found within,
or emerging from, a single puparium, varied from 2 to 10.

Achcetoneura samice Web. (Tachinidas)

This insect occurred occasionally, as a primary parasite, within
all three lots of cocoons, and several adult specimens were reared.
In addition, a single specimen of Sarcophaga misera var. sarra-
cenioides Aid. (Sarcophagidae) emerged. A large number of
dipterous pnparia were found which varied somewhat in size and
coloration. Since many of these did not emerge as adults, the
writer could not determine to his entire satisfaction that these
were all of the same species. It is thought probable, however,
that most of these puparia were those of Achcetoneura samice,
since only a single specimen of any other large dipteron was
obtained. But the possibility that other species might have been
represented should not be overlooked. From the first lot of
cocoons, only 19 out of 100 cocoons were parasitized, or showed
signs of having been parasitized, by a large dipteron. The para-
sitized cecropia in some instances had formed a pupa within the
cocoon before dying. In a few cases, the parasites had emerged
from the host and formed puparia loose in the cocoon; in other
cases, puparia were formed within the body of the host.

Several of the dipterous puparia were parasitized. Within one
puparium, 22 chalcidoid pupae were found, and several other dis-
sected puparia yielded additional parasites. In addition, many
of these insects emerged in the breeding bags. This parasite
proved to be Dimmockia incongrmis Ashm. (Enlophidae). All
these insects were in the pupal stage, and some of these pupae were
likewise parasitized. Within the pupal skin of some of these
insects were the larvae or pupae of Pleurotropis tar satis (Ashm.).
Normal pupae of D. incongrmis were of a uniform glossy black

262

Journal New York Entomological Society [Vol. xlviii

color, but the parasitized pupae were brownish, and the pupal
skin was broken in many cases. Pleurotropis iarsalis was thus
in this instance a tertiary parasite.

Spilocryptus extrematus Cress. (Ichneumonidae)

From the first lot of cecropia cocoons examined, only a single
cocoon was parasitized by this ichneumonid. Within this one
cocoon, however, there were 54 cocoons of this parasite, all of
which were still in the larval stage when they were examined
January 6. None of these ichneumonids was parasitized.

The second lot of cecropia, however, had been collected spe-
cifically for this parasite in a somewhat different region, and
some of the cocoons had been opened in order to determine defi-
nitely the nature of parasitism. Thus, in the second lot, there
were 12 cocoons, out of the 20 received, which were parasitized
by Spilocryptus extrematus Cress. Two species of chalcidoid
parasites were parasitic upon ichneumonids from 6 of these 12
parasitized cecropia cocoons. These insects were Dihrachys cavus
(Walk.) (Pteromatidce) , and Monodontomerus sp., (F) (Calli-
momidae) .

The pteromalids were present within two of the cecropia co-
coons, and at the time of dissection, March 31, these insects were
in the pupal stage. Possibly because of the warm laboratory
temperature, these insects emerged a few days later as adults.
Females oviposited readily into living ichneumonid larvae.

Monodontomerus sp., was obtained from 4 of the parasitized
cecropia cocoons. Presumably these insects were external para-
sites of Spilocryptus extrematus, and at the time of dissection,
March 30, occurred in both the larval and pupal stages. The
parasites were loose in the ichneumonid cocoon, and in most in-
stances the shriveled-up larva of the host was still present.

In the laboratory, the females did not show particular interest
in cocoons or naked larvae of Spilocryptus extrematus, and no
attempt was made to oviposit. When, however, a cecropia cocoon
containing ichneumonid cocoons was placed with the females,
several attempts were made to oviposit through the cecropia co-
coon. One female also crawled into the cocoon and stayed sev-
eral minutes. Whether or not oviposition occurred is not known.

Sept., 1940]

Breland: Parasites

263

It would seem, however, that in this case, the combination of the
cecropia cocoon plus the ichneumonid cocoons, was a stronger
stimulus to oviposition than the ichneumonid larvse or cocoons
alone.

It is thought probable that the females of Monodontomerus
sp., since these insects possess a comparatively short ovipositor,
crawl through the valve of the cecropia cocoon, and oviposit
directly into the cocoon of their host. In all cases of parasitism
examined, the callimomids were inside the host cocoon. Only a
few of the total ichneumonid cocoons were parasitized.

MULTIPLE PARASITISM

In a few cases, cecropia cocoons contained several types of
parasites. One cocoon was of particular interest. Within this
cocoon were several cocoons of Spilocryptus extrematus, all in-
sects of which were in the pupal stage. In addition there were
several larvae of Pseudogaurax anchora, and some large dipterous
puparia. Within two of these dipterous puparia were pupae of
Dimmockia incongruus, and some of these in turn were para-
sitized by Pleurotropis tarsalis. This cecropia had, therefore,
directly or indirectly, supported insects of 5 species. It is prob-
able that under normal conditions most of these insects would
have emerged as adults. But in this particular case, there was
a hole in the cocoon, indicating that a bird had enjoyed a meal
at the expense of the enclosed occupants. Consequently several
of these insects were dead.

I believe this study emphasizes the fact that parasitic popula-
tions of a given host may var}^ considerably from locality to local-
ity and even may vary in regions of the same locality. Marsh
(1934), in a study of cecropia cocoons collected in the vicinity
of Chicago, reared 9 species of parasitic insects. Of the 9 species
reared by the writer, only three, Dimmockia incongruus, Spilo-
cryptus extrematus, and Pleurotropis tarsalis, were identical with
species obtained by Marsh. In the Chicago area. Marsh dis-
covered that Spilocryptus extrematus was the principal primary
parasite of the cecropia. In the present study, the principal
primary parasite was Pseudogaurax anchora, and parasitism by
Spilocryptus extrematus was extremely limited.

264

Journal New York Entomological Society [Vol. xlviii

LITEEATURE CITED

Kaston, B. J. and Jenks, G. E. 1937. Dipterous parasites of spider egg
cases. Bull. Brook. Eiit. Soc., 32: 160-165.

Marsh, F. L. 1934. A regional study of Sarnia cecropia L. and nine asso-
ciated parasites and hyperparasites. A master’s thesis. North-
western University library, Evanston, Illinois.

. 1936. Egg placing by DihracJiys Itoucheanus Eatzburg. Can. Ent.,

68: 215-216.

■ . 1937. The biology of the ichneumonid Spilocryptus extrematus

Cresson (Hymenoptera) . Ann. Ent. Soc. Am., 30: 40-42.

Montgomery, B. Elwood. 1933. Preliminary studies of insect parasites
of Indiana. Can. Ent., 65: 185-190.

Sept., 1940]

CUMLEY: Deosophila

265

COMPARISON OF SEROLOGIC AND TAXONOMIC
RELATIONSHIPS OF DROSOPHILA
SPECIES*

By Bussell W. Cumley
The University of Texas

INTRODUCTION

During the past two years studies have been made of extracts
prepared from various species of Drosophila with the view of dis-
tinguishing the species through the use of serological procedures.
Several different technics have been tried, and the reagents which
were tested have been prepared in different ways. The results of
these experiments have been published in several reports, to which
reference will be made later. The purpose of this work is to com-
pare the species relationships, as revealed by the serologic investi-
gations of their antigens, with the relationships which are recog-
nized on the basis of the more commonly accepted taxonomic
criteria.

Although the different serological tests have yielded similar
results, seldom have they offered exactly the same relations among
the several species. It is impossible, at the present time, to state
with certainty which technic reveals most nearly the real bio-
chemical relationships among the species. Boyden (1936) has
stated that no two technics are of equal worth; and Chester
(1937), in his extensive review of plant serology, has been unable
to conclude which of the many technics and modifications most
nearly reveal the truth. This lack of agreement of the various
tests and the consequent indecision regarding the relative value
of serologic technics present a problem not greatly different from
that of the taxonomist. The taxonomist is confronted with the
difficulty of determining w^hich characteristic or group of charac-
teristics relates most truly the various species.

* The methods herein discussed were presented in demonstration at the
Sixth Annual Meeting of the Genetics Society of America, Indianapolis,
1937. An abstract may be found in Eecords of the Genetics Society of
America, No. 6, p. 146, 1937.

This article was taken from the author’s Ph.D. Dissertation, The Univer-
sity of Texas, June, 1938.

266

Journal New York Entomological Society [Vol. XLVIII

Aside from the difficnlty in appraising the systematic worth
of any morphological characteristic, another problem of species
inter-relations presents itself. A given characteristic which is
thought to be of specific value may serve well to distinguish one
species from a second, but may in turn be shared by a third spe-
cies. This fact, no doubt, has contributed considerably to the
confusion regarding onr knowledge of evolution and inter-rela-
tions of species. In serological investigations of animals relation-
ships the same feature has been noted by Irwin (1938), who
found that each dove or pigeon species possessed cellular sub-
stances, determined by the agglutinin-absorption technic, which
were not found in any of the other species. Also, he showed that
some of the serological characteristics which distinguished one
species from another could in turn be shared with still other spe-
cies. Hence, difficulties of evaluating specific qualities are com-
mon to both the morphologic and immunologic procedures. In
serologic tests, however, the reactive substances are biochemical
elements of the body tissues of the animal. Through the applica-
tion of technics which assay these fundamental chemical proper-
ties of the organism, we should be able ultimately to determine
with a reasonable degree of exactitude the extent to which the
protoplasm of one species is similar to that of others.

MATERIALS AND METHODS

In the serological tests that have been performed the relation-
ships of about a dozen Drosophila species have been studied.
Extensive data, however, have been accumulated only for the
four species: D. carMea, D. melanogaster, D. mulleri, and D.
virilis. At least a dozen tests, representing several different pro-
cedures, have been performed on the antisera to each Of these
species. The taxonomic data used in this report were taken
largely from Professor Sturtevant’s work (1921). These data
were supplemented by those of other authors (Metz, Moses, and
Mason, 1923) and by some of my own observations. Recognizing
our incapacity to appraise the relative specific worth of any mor-
phological unit or serological reaction, I have assumed that all
of the various immunological tests are of equal value, and that all
of the fourteen taxonomic characters are of equal value.

Sept., 1940]

CuMLEY: Drosophila

267

The serological reactions which were employed in this com-
parison were as follows :

1. Complement-fixation reactions, using as antigens the saline
extracts of dried flies, without any further extraction (Cnmley
and Haberman) ;

2. Precipitation reactions — ring tests, using the same antigens
as in the preceding (Haberman and Cnmley) ;

3. Complement-fixation reactions, using as antigens the saline
extracts of the ether-insoluble fractions of dried flies (Cnmley,
1939) ;

4. Optimal antigen-antibody ratio reactions, using the same
antigens as in the tests immediately above (Cumley, 1938) ;

5. Precipitin absorptions, using the same antigens as in the
preceding (Cumley, 1939, a).

The systematic characteristics employed in this comparison
were as follows :

1. Costal index : Length of 2nd section of costal vein/Length
of 3rd section of costal vein ;

2. Fourth-vein index: Length of 4th (distal) section of 4th
vein/Length of 3rd section of 4th vein;

3. 4c index: Length of 3rd section of costal vein/Length of
3rd section of 4th vein ;

4. 5x index: Length of 3rd (distal) section of 5th vein/
Length of posterior vein ;

5. Number of branches of arista ;

6. Approximate width of the “front ’’/width of the head;

7. Size of 2nd orbital bristle/Size of the other two;

8. Size of the 1st oral bristle/Size of the 2nd ;

9. Greatest width of the cheeks/Greatest diameter of the eyes ;

10. Number of rows of acrostichal hairs ;

11. Number of filaments on eggs ;

12. Body length ;

13. Wing length ;

14. Number and type of chromosomes.

In order to rank the species serologically, each serologic test
was considered separately, and the species were arranged in the
order of the extent of reactivity of their antigens to the particu-
lar antiserum in question. This procedure was repeated for each

268

Journal New York Entomological Society [Vol. XLVIII

of the several tests. Since all the tests of a given antiserum did
not rank the species in the same order, the result was that any
one species would sometimes assume second, and sometimes third
or fourth place. From data taken in this way from the various
tests, it was possible to observe the number of times a particular
species assumed each of the four serological ranks made possible
by a consideration of only four species. The percentage ranks
were calculated from these observations and are shown in Table 1.

TABLE 1

SEROLOGIC RANKING

Percentage ranks

Species foi comparison

1

2 3

4

Species in question; Drosophila melanog aster

D. melanogaster

100.0

0

0

0

D. carihhea

0

96.5

3.5

0

Z>. mulleri

0

3.5

64.3

32.2

D. virilis

0

0

32.2

67.8

Species in question:

Drosophila carihhea

D. carihhea

100.0

0

0

0

D. melanogaster

0

71.1

13.5

15.4

D. mulleri

0

11.5

57.8

30.7

D. virilis

0

17.4

28.7

53.9

Species in question :

: Drosophila mulleri

D. mulleri

100.0

0

0

0

D. virilis

0

88.5

11.5

0

D. carihhea

0

7.7

69.3

23.0

D. melanogaster

0

3.8

19.2

77.0

Species in question

: Drosophila virilis

D. virilis

100.0

0

0

0

D. mulleri

0

82.2

3.6

14.2

D. carihhea

0

17.8

50.0

32.2

D. melanogaster

0

0

46.4

53.6

In this table one may observe that, with reference to D. melano-
gaster, the four Drosophila species are ranked as follows : melano-
gaster, 1st place in 100 per c«nt of the tests; carihhea, 2nd place
in 96.5 per cent, and 3rd place in 3.5 per cent of the tests ; mulleri,

Sept., 1940]

Cumley: Drosophila

269

2nd place in 3.5 per cent, 3rd place in 64.3 per cent, and 4th place
in 32.2 per cent of the tests; virilis, 3rd place in 32.2 per cent and
4th place in 67.8 per cent of the tests. The same type of informa-
tion is to be seen in the remainder of the table. That is to say,
all four of the species are ranked in their respective relations to
a given species.

The taxonomic ranking has been accomplished in much the
same way, except that instead of considering serological tests, the
individual morphological units have been applied. The species
are then ranked in the order of their relationship in terms of a
given systematic criterion. As in the preceding rankings, the
percentage ranks of the various species with reference to a given
species have then been calculated. These percentage ranks have
been presented in Table 2. In this table one may see that the

TABLE 2

TAXONOMIC RANKING

Percentage ranks

SpociGS f 01 C’Omp9.(Tison

1

2 3

4

Species in question: Drosophila melanogaster

D. melanogaster

100.0

0

0

0

D. carihhea

0

77.0

23.0

0

D. mulleri

0

7.6

42.4

50.0

D. virilis

0

15.4

34.6

50.0

Species in question :

Brosophila carihhea

D. carihhea

100.0

0

0

0

B. melanogaster

0

84.8

11.5

3.7

D. mulleri

0

7.6

46.2

46.2

B. virilis

0

7.6

42.3

50.1

Species in question:

Brosophila mulleri

D. mulleri

100.0

0

0

0

B. virilis

0

69.3

7.6

23.1

B. melanogaster

0

11.5

53.8

37.7

B. carihhea

0

19.2

38.6

42.2

Species in question

: Brosophila virilis

B. virilis

100.0

0

0

0

B. mulleri

0

77.3

4.5

18.2

B. melanogaster

0

9.1

59.1

31.8

B'. carihhea

0

13.6

36.4

50.0

270

Journal New York Entomological Society [Vol. xlviii

various Drosophila species are related to D. melanogaster as fol-
lov^s: melanogaster, 1st place in 100 per cent of the characters;
carihhae, 2nd place in 77.0 per cent and 3rd place in 23.0 per cent
of the characters; mulleri, 2nd place in 7.6 per cent, 3rd place in
42.4 per cent, and 4th place in 50.0 per cent of the characters ; and
virilis, 2nd place in 15.4 per cent, 3rd place in 34.6 per cent, and
4th place in 50.0 per cent of the taxonomic characters. Similarly,
all four of the species are ranked taxonomically in their relations
to a given species.

RESULTS

Figure 1 has been prepared in order that a more graphic viev7
may be obtained of the comparison of the serologic and taxonomic
rankings. From a perusal of this figure and the data of Tables
1 and 2 several facts become apparent :

1. The serological and taxonomic technics have always pre-
sented the same species for second rank.

2. With reference to ranks three and four, the taxonomic and
serologic methods have given essentially the same results in two
of the comparisons (Z>. melanogaster and D. carihhea), and have
failed to correspond, apparently, in the other two comparisons.
In the two cases where the results failed to coincide, however,
there is some doubt as to the real ranking on the basis of taxo-
nomic data. That is to say, when the species were ranked taxo-
nomically in relation to D. virilis, the species caribhea assumes
fourth rank in 50.0 per cent of the characteristics, whereas
melanogaster ranks fourth in only 31.8 per cent of the tests. By
virtue of these figures alone, caribbea obviously would receive
fourth rank. However, caribbea also ranks second in 13.6 per
cent of the systematic features, whereas melanogaster ranks
second in only 9.1 per cent. Therefore, it is impossible to deter-
mine offhand which of the two species should be considered as
ranking third place. The same situation exists when the species
are given morphological ranks, with reference to D. mulleri.

In general, then, we may conclude that the taxonomic and sero-
logic methods agree definitely to the extent of the second rank,
and when they disagree as to the third and fourth ranks, the
taxonomic method has not clearly indicated which order of rela-
tionship is correct.

FIGURE 1

SEROLOGIC BANKING OF DROSOPHILA SPECIES

Sept., 1940]

CuMLEY: Drosophila

271

272

Journal New York Entomological Society [Vol. xlviii

3. The results of the serologic ranking apparently are more
specific than are those of the taxonomic ranking. This fact is
indicated by several features of Figure 1 and Tables 1 and 2.
First, a species which ranks third serologically always is repre-
sented by a sharply-peaked curve. Not only is that not the case
with the taxonomic ranking, but there is always some doubt as
to which species really ranks third. This fact is readily apparent
since all of the curves representing the third and fourth places,
taxonomically, cross each other or coincide in at least three
points; whereas in the curves representing the serologic ranks,
only twice do the 3rd and 4th rank curves coincide more than
once. Furthermore, in none of the taxonomic rankings, as ob-
served in Table 2, is it possible to determine which species ranks
third and which fourth ; whereas in the serologic data of Table 1,
in only one ranking (carihhea) is this the case. Second, in two
of the serological rankings those species which rank fourth never
ranked second in any of the tests. In the case of the taxonomic
ranking, all the species that ranked fourth on the basis of some
morphological unit, ranked second on the basis of some other.
Third, those species which are ranked fourth serologically are so
ranked by virtue of from 53.6 per cent to 77.0 per cent of the
tests indicating this rank. Those species ranked fourth taxonomi-
cally are so ranked by virtue of 42.2 per cent to 50.1 per cent of
their characteristics. Similar figures may be observed for the
second and third ranks.

4. The two subgroups designated by Sturtevant (1921) as sub-
group 1 and subgroup 2 of Group F have been definitely indi-
cated by the serologic methods, as well as by the taxonomic
studies. Sturtevant ’s subgroup 1 includes the yellowish or red-
dish species of which D. melanogaster and D. carihhea are mem-
bers. Subgroup 2 includes the blackish or grayish species of
which D. virilis and D. mulleri are members. In the serologic
ranking as well as in the morphologic ranking, D. virilis and D.
mulleri always rank close together, and D. carihhea and D. mel-
anogaster always rank close together. If one assumes that these
two subgroups arose from a common stock and that the species of
each subgroup diverged much later, then this is more or less the
relation one would expect to find.

Sept., 1940]

CuMLEY: Drosophila

273

CONCLUSIONS

Serologic technics recently have been employed by several
investigators in relating or ranking species of molluscs (Makino,
1934), helminths (Eisenbrandt, 1936), amphibia (Boyden and
Noble, 1933), moths (Martin and Cotner, 1934), and other animal
species. Problems of hybrids, likewise, have been attacked by
these biochemical methods (Irwin, 1938; Irwin and Cole, 1936
a & b; Irwin, Cole and Gordon, 1936). In general the results
have been sufficiently encouraging to warrant the application of
serologic technics to an analysis of Drosophila species. The pres-
ent paper and those which have preceded serve to indicate the
results which may be expected from such procedure. Obviously,
the work has only begun, and several refinements are necessary.
It is believed, however, that these methods eventually will provide
valuable data relative to problems of speciation and phylogeny.
Furthermore, the recent work of Levit, Ginsburg, Kalinin, &
Feinberg (1936) suggests the possibility of applying immuno-
logic technics to the study of the expression of individual chro-
mosomes or even genes. To what extent the method may be uti-
lized remains at present a matter of conjecture.

These comparisons have demonstrated that on the basis of
morphology, species cannot always be ranked to the third and
fourth places, but with the serologic methods this can be accom-
plished. In cases where morphological differences are insignifi-
cant or absent, the serologic technics may provide adequate means
of determining species relationships.

LITERATUEE CITED

Boyden, A. A. 1936. Serology and biological problems. Sigma Xi Quar.,
24: 152-160.

Boyden, A. A., and Noble, G. K. 1933. The relationships of some com-
mon Amphibia as determined by serological study. American
Museum Novitates. No. 606, pp. 1-24.

Chester, K. S. 1937. A critique of plant serology: Part I, The nature
and utilization of phytoserological procedures, Quar. Rev. of Biol.,
12 : 19-46 ; Part II, Application of serology to the classification of
plants and the identification of plant products. Ihid., 165-190;
Part III, Phytoserology in medicine and general biology, — bibli-
ography. Ihid., 294—321.

CuMLEY, R. W. 1937. Serology of Drosophila. Rec. of the Genetics Soc.
of America., No. 6, p. 146 (Abstract).

274

Journal New York Entomological Society [Vol. xlviii

. 1938. Serological Investigation of Drosophila Antigens. Dis-
sertation, The University of Texas.

. 1939. The relations among Drosophila species, as determined by

the complement fixation reaction using ether-insoluble fractions.
Jour. Exp. ZooL, 80: 299-314.

. 1939, a. Precipitin absorption with Drosophila antigens. The

American Naturalist, Vol. LXXIII, pp. 375-380.

CuMLEY, E. W., AND Haberman, S. 1938. Serological investigation of
Drosophila antigens Avith the complement fixation reaction. Jour.
N. Y. Ent. Soc., Vol. XLVI, pp. 401-415.

Eisenbrandt, L, L. 1936. Precipitin reactions of Helminth extracts.

Proc. Soc. Exp. Biol, and Med., 35: 322-325.

Haberman, S., and Cumley, E. W. 1939. Serological investigation of
Drosophila antigens using the precipitation reaction. Jour. N. Y.
Ent. Soc., Vol. XLVII, pp. 219-226.

Irwin, M. E. 1938. Immuno-genetic studies of species relationships in
Columbidse. Jour, of Gen., 35: 351-373.

Irwin, M. E., and Cole, L. J. 1936a. Immunogenetic studies of species
and of species hybrids in doves, and the separation of species-spe-
cific substances in the backcross. Jour. Exp. ZooL, 73: 85-108.

. 1936b. Immunogenetic studies of species and of species hybrids

from the cross of Columha livia and Streptopelia risoria. Jour.
Exp. ZooL, 73: 309-318.

Irwin, M. E., Cole, L. J., and Gordon, D. D. 1936. Immunogenetic studies
of species and of species hybrids in pigeons, and the separation of
species-specific characters in back-cross generations. Jour. Exp.
ZooL, 73: 285-308.

Levit, S. G., Ginsburg, S. G., Kalinin, V. S., and Feinberg, E. G. 1936.

Immunological detection of the Y-chromosome in Drosophila
melanogaster. Nature, 138; 3480; 78-79.

Making, K. 1934. Beobachtungen uber die Immunitatsreaktionen bei Mol-
luskenarten. Zeit. fur Immunitats. Bd. 81, Heft 3/4, 316-335.
Martin, S,, and Cotner, F. B. 1934. Serological studies of moth proteins
with special reference to their phylogenetic significance. Ann.
Ent. Soc. Amer., 27 : 372-383.

Metz, E. W., Moses, M. S., and Mason, E. D. 1923. Genetic studies on
Drosophila virilis with considerations on the genetics of other spe-
cies of Drosophila. Carnegie Institution of Washington. Publ.
No. 328, 94 pp.

Sturtevant, a. H. 1921. The North American Species of Drosophila.
Carnegie Institution of Washington. Publ. No. 301, iv 1-150 pp.

Sept., 1940]

Funkhouser : Membracid^

275

NEW PERUVIAN MEMBRACID^ (HOMOPTERA)

By W. D. Funkhouser
University of Kentucky

A large collection of Membracidse recently received from Peru
contains several new species which are here described and figured
as follows :

1. Lycoderes luteus sp. nov. (Fig. 1)

Large, yellow, coarsely punctate, not pubescent; head quad-
rate, subf oliaceous ; pronotum extended into flattened porrect
horn; posterior process flattened, tectiform, separated very little
from anterior portion of pronotum and reaching to the internal
angles of the tegmina ; tegmina hyaline with basal and anal areas
coriaceous; legs yellow and f oliaceous; undersurface of body
yellow.

Technical description :

Head quadrate, broader than high, roughly sculptured, finely punctate;
base feebly arcuate and sinuate; eyes flattened, brown; ocelli small, incon-
spicuous, much farther from each other than from the eyes and situated close
to the upper margin of the head, far above a line drawn through centers of
eyes ; inferior margins of gense lobate, f oliaceous and sinuate ; clypeus broad,
subovate, not extending below inferior margins of gense, tip truncate.

Pronotum flattened laterally, yellow tinged with brown, coarsely punctate,
not pubescent, bearing a porrect frontal horn and a strong posterior process
with very little constriction between the two ; pronotal horn porrect, sub-
triangular, laterally flattened, tip blunt with a weak lateral extension on each
side; posterior process heavy, laterally flattened, coarsely punctate but not
laterally carinate, sharply tectiform, median carina strongly percurrent, tip
blunt and extending just to the internal angles of the tegmina; humeral
angles weak and rounded ; scutellum well exposed on each side, below a weak
inferior constriction between the frontal horn and the posterior process.

Tegmina broad, entirely exposed, hyaline; base broadly brown, coriaceous
and punctate ; anal margin narrowly brown, translucent and punctate ; five
irregular apical cells ; one discoidal cell ; apex rounded ; anal margin truncate ;
no apical limbus.

Legs yellow and f oliaceous; all tarsi about equal in length. Sides of
mesonotum and metanotum extended into pointed teeth. Under-surface of
body yellow; abdomen greenish.

276

Journal New York Entomological Society [Vol. xlviii

Length from front of head to tips of tegmina 6.2 mm,; width between
humeral angles 2 mm.

Type: Female.

Type locality: Guaybamba, Amazones, Peru.

Described from a single specimen collected in August 1936, Type in
author’s collection.

2. Stylocentrus rubrinigris sp. nov. (Fig. 2)

Shining black with the tegmina and abdomen strongly marked
with red ; base of head strongly bitubercnlate ; pronotnm gibbous
with a pair of slender, curved frontal horns and a slender, arched
posterior horn, all three arising from a very short stalk; no pos-
terior process ; scutellum entirely exposed ; tegmina hyaline with
heavy black veins and with membrane strongly tinged with red ;
legs and undersurface of body rufous ; abdomen red.

Technical description :

Head triangular, black with white pubescence, roughly sculptured, finely
punctate; base arcuate and sinuate and bearing two prominent tubercles;
eyes large, globular, glassy; ocelli small, amber-colored, situated on rounded
elevations more than twice as far from each other as from the eyes and
located near the base of the head far above a line drawn through centers of
eyes ; inferior margins of genge straight and sloping ; clypeus sharply pointed,
extending for one-third its length below inferior margins of gense.

Pronotum convex and gibbous, black, finely punctate, and bearing a
linear white tomentose patch below the horns ; a very short porrect stalk gives
rise to a pair of long, slender, recurved frontal horns and a slender, tri-
carijiate posterior horn which arches high above the body and then curves
downward, well above the abdomen and reaching almost to the internal angles
of the tegmina; scutellum entirely exposed; triangular, tip sharp; humeral
angles weak and rounded; inferior margin of pronotum projected downward
in a tooth.

Tegmina entirely exposed, long, narrow, hyaline; base and entire clavus
coriaceous and punctate; membranes of basal half bright red; veins heavy
and black ; five irregular apical and two irregular discoidal cells ; anal margin
truncate; no apical limbus.

Sides of thorax reddish-black with white pubescence; legs simple, reddish-
black, hind tarsi longest ; abdomen red.

Length from front of head to tips of tegmina 7.2 mm.; width between
tips of frontal spines 5 mm.

Type: Male.

Type locality: Callanga, Peru.

Described from a single specimen in the author’s collection. This species
is close to S. championi Fowler but is much larger and differs particularly in
the structure of the pronotal spines.

Sept., 1940]

Funkhouser : Membracid^

277

3. Alchisme laticornis sp. nov. (Fig. 3)

Heavy bodied, reddish-brown mottled with black, coarsely
punctate, sparingly pubescent; humeral angles produced into
long, sharp horns ; median pronotal horn broad, flat and ridged ;
posterior process heavy, tectiform and nearly straight; tegmina
subhyaline, bronze, with very broad veins ; legs and undersurface
of body reddish-brown ; hind tarsi very short.

Technical description:

Head subquadrate, twice as broad as high, roughly sculptured; base
arcuate and sinuate; eyes large, gray, globular; ocelli large, prominent,
glassy, much nearer to each other than to the eyes and situated about on a
line drawn through centers of eyes; inferior margins of gense straight and
swollen ; clypeus reflexed, extending for half its length below inferior margins
of genae, tip blunt.

Pronotum convex, rounded, reddish-brown with a transverse black band
extending across the top of the metopidium and covering the dorsal surfaces
of the humeral spines, coarsely punctate, sparingly pubescent, produced at
the shoulders into long spines, above into a broad median horn and behind
into a heavy posterior process; metopidium sloping, broader than high;
median carina strongly percurrent ; humeral angles extended into heavy,
flattened, sharp spines extending outward and forward; median pronotal
horn erect, laterally flattened, irregularly ridged on both sides, higher than
its breadth at base, anterior margin nearly straight, posterior margin rounded,
tip blunt; posterior process heavy, nearly straight, tectiform, tip sharp and
extending a little beyond the internal angles of the tegmina.

Tegmina subhyaline, smoky-brown, entirely exposed ; basal and costal areas
coriaceous, brown and punctate ; veins very broad ; five apical and three
discoidal cells ; apical limbus narrow. Hind wings with four apical cells.

Legs reddish-brown, stout ; femora cylindrical ; hind tibiae flattened and
spatulate at distal end ; hind tarsi very short.

Length from front of head to tips of tegmina 8.2 mm. ; width between tips
of humeral spines 4 mm.

Type : Male.

Type locality: Pumamarca, Peru.

Described from a single specimen in the author ’s collection.

4. Alchisme pinguicornis sp. nov. (Fig. 4)

Small, uniformly dark brown, punctate, pubescent; humeral
angles cylindrical, blunt, extending outward and slightly for-
ward; dorsal crest erect, thick, heavy, rounded, tricarinate on
each side with the posterior ridge extending to the inferior
lateral margin of the pronotum ; posterior process long, slender.

278

Journal New York Entomological Society [Vol. XLVIII

acute, reaching beyond the internal angles of the tegmina ;
tegmina dark brown with strongly pilose veins; legs and under-
surface of body brown.

This species is near the preceding but is much smaller and
differs particularly in the structure and sculpturing of the head
and pronotal process. It is also apparently near A. turrita Ger-
mar but is not identical.

Technical description :

Head subquadrate, three times as broad as high, black, finely punctate,
sparingly pilose; base arcuate and feebly sinuate; eyes large, black, ovate;
ocelli small, inconspicuous, amber-colored, equidistant from each other and
from the eyes and situated about on a line drawn through centers of eyes;
inferior margins of genae regularly curved; clypeus short, rounded, extend-
ing only a little below inferior margins of genae and continuing the rounded
inferior outline of the face made by these margins.

Pronotum dark brown, coarsely punctate, sparingly pubescent ; metopidium
sloping, broader than high; median carina strongly percurrent; humeral
angles extended into heavy, subconical, blunt horns projecting outward and
slightly forward; median dorsal crest thick, heavy, arising from just behind
the humeral angles, only slightly flattened laterally, about as high as its
width at base, tip broadly rounded, three ridges on each side, the posterior
ridge very heavy and reaching to the inferior lateral margin of the pro-
notum; posterior process long, slender, tectiform, nearly straight, tip acute
and reaching to a point about half-way between internal angles and tips of
tegmina.

Tegmina entirely exposed, long, narrow, brown; basal and costal areas
coriaceous and punctate ; veins strongly pilose ; five apical and three discoidal
cells ; apical limbus broad.

Undersurface of body dark brown; legs simple, slender, very dark brown;
hind tibiae spatulate; hind tarsi much shorter than the others.

Length from front of head to tips of tegmina 6.8 mm.; width beUveen
tips of humeral spines 3.5 mm.

Type: Male.

Type locality : Santo Domingo, Peru.

Described from a single specimen collected in November 1937 by Mr. Felix
Woytkowski. Type in author’s collection.

5. Alchisme spinosa sp. nov. (Fig. 5)

Large, pale greenish-yellow, coarsely punctate, not pubescent;
humeral angles long, slender, projecting outward, upward and
forward; dorsal spine long, sharp, slightly curved; posterior
process tectiform, straight, sharp,' just reaching tips of tegmina ;
tegmina hyaline, free ; undersurface yellowish-green.

Sept., 1940]

Funkhouser: Membracid^

279

Technical description :

Head subovate, twice as broad as high, yellow, smooth; base high and
strongly bisinuate ; eyes large, ovate, brown ; ocelli large, conspicuous, yellow,
equidistant from each other and from the eyes and situated on a line drawn
through centers of eyes; inferior margins of genae sinuate and sloping;
clypeus extending for half its length below inferior margins of genae.

Pronotum greenish-yellow, coarsely punctate, not pubescent, margins
smooth and strongly carinate; metopidium triangular, higher than broad;
median carina strongly percurrent; humeral angles extended into long spine-
like processes, longer than the width of the metopidium, extending strongly
outward and slightly upward and forward, base impinging on the eyes, some-
what flattened dorso-ventrally, tips rounded ; dorsal spine long, acute, arising
from behind humeral angles, laterally flattened, slightly recurved, margins
ridged; posterior process long, slender, tectiform, straight, tip sharp and
just reaching tips of tegmina.

Tegmina hyaline, entirely free ; basal and costal areas narrowly coriaceous
and punctate ; veins strong ; five apical cells ; discoidal cells small and vari-
able in number ; apical limbus broad.

Sides of thorax and undersurface of body yellow; legs yellow, simple;
hind tarsi much shorter than the others.

Length from front of head to tips of tegmina 12 mm.; width between
tips of humeral spines 9 mm.

Type: Female.

Type locality: Santo Domingo, Peru.

Described from two females collected at the type locality in November
1937 by Mr. Felix Woytkowski. Type and paratype in author’s collection.

6. Paragargara nigra sp. nov. (Fig. 6)

Small, black, finely punctate, sparingly pubescent; anterior
pronotum strongly convex ; median carina very sharp ; no anterior
processes ; posterior process sharply set off by a deep indentation ;
tegmina hyaline with inconspicuous veins ; legs simple.

Technical description:

Head subf oliaceous, smooth, black, finely punctate, not pubescent, vertical ;
base arcuate and weakly sinuate; eyes large, globular, white; ocelli small,
inconspicuous, glassy, twice as far from each other as from the eyes and
situated high up near the base of the head far above a line drawn through
centers of eyes ; inferior margins of gense short, sinuate and sloping ; clypeus
very broad, flat, extending for two-thirds its length below inferior margins
of genae, tip truncate.

Pronotum strongly convex, gibbous, dull black, very finely punctate, spar-
ingly pubescent, slightly carinate on each side ; metopidium vertical, broader
than high; median carina sharply percurrent; humeral angles heavy, tri-
angular, blunt; no anterior horns or processes; posterior process arcuate,

280

Journal New York Entomological Society [Vol. xlviii

carinate, short, separated from the anterior pronotum by a deep depression
which gives it an up-turned appearance, tip sharp, depressed and extending
just to the internal angles of the tegmina.

Tegmina broad, hyaline, entirely exposed; base broadly black, coriaceous
and punctate; veins indistinct; five apical and two discoidal cells; apical
limbus very broad.

Sides of thorax and undersurface of body black ; legs simple, ferruginous ;
all tarsi about equal in length.

Length from front of head to tips of tegmina 3.8 mm.; width between
tips of humeral angles 1.7 mm.

Type: Female.

Type locality : Leonpampa, Peru.

Described from two specimens, a male and a female, the female from the
type locality and the male from Marcapata, Peru. The male agrees with the
female in all respects except that it is slightly smaller. Type and allotype
in author’s collection.

7. Aconophora erecta sp. nov. (Fig. 7)

Large, ferruginous, punctate, pubescent; anterior horn long,
porrect, laterally flattened, tip recurved ; posterior process heavy,
decurved, reaching almost to tips of tegmina; tegmina fuscous-
hyaline, entirely exposed; sides of thorax ferruginous; legs fer-
runginous, heavy, femora cylindrical, tibige triquerate ; tarsi
equal.

Technical description :

Head triangular, punctate, puebscent; base arcuate and sinuate; eyes
globular, black ; ocelli large, conspicuous, brown, equidistant from each other
and from the eyes and situated somewhat above a line drawn through centers
of eyes; inferior margins of genee sloping and strongly sinuate; clypeus
broad, extending for half its length below inferior margins of genae, tip
pointed and hirsute.

Pronotum uniformly ferruginous, finely punctate, pubescent; metopidium
projecting forward, triangular, broader than high; median carina percurrent;
humeral angles heavy, rounded, blunt; anterior pronotal horn slender, later-
ally flattened, edges subfoliaceous, extending strongly upward and slightly
forward, as long as the distance from its base to the humeral angles, tip
broadly rounded, slightly recurved; posterior process heavy, convex, not
carinate, tip sharp, depressed, and reaching almost to the tips of the
tegmina.

Tegmina entirely free, fuscous-hyaline; base coriaceous, brown and punc-
tate; basal area slightly pilose; tip pointed; veins strong; five apical and
two discoidal cells; apical limbus broad.

Sides of thorax, undersurface of body and abdomen uniformly ferruginous ;
legs heavy, ferruginous; femora cylindrical; tibiae triquerate and spined; all
tarsi equal in length.

Sept., 1940J

Funkhouser : Membracid^

281

Length from front of head to tips of tegmina 6.3 mm. ; width between tips
of humeral angles 3.2 mm.

Type: Male.

Type locality: Napo Eiver, Peru.

Described from a single specimen collected in June 1920. Type in author’s
collection.

8. Aconophora brunnea sp. nov. (Fig. 8)

Small, brown, punctate, pubescent ; anterior horn thick, heavy,
curving forward; posterior process stout, reaching to a point
half-way between internal angles and tips of tegmina; tegmina
entirely exposed, hyaline with brown markings ; legs and under-
surface of body brown.

Technical description:

Head triangular, dark brown, punctate, pubescent ; base arcuate and feebly
sinuate; eyes ovate, glassy; ocelli small, inconspicuous, amber-colored, equi-
distant from each other and from the eyes and situated a little above a line
drawn through centers of eyes; inferior margins of gense sloping and sinuate;
clypeus broad, extending for half its length below inferior margins of genae.

Pronotum light brown, finely punctate, densely pubescent; metopidium
extending forward, subtriangular, broader than high; median carina per-
Qurrent ; humeral angles weak, rounded, blunt ; anterior pronotal process
heavy, stout, as long as the distance from its base to the humeral angles,
flattened laterally, extending upward and forward and curving distinctly
forward over the head and metopidium, tip broadly rounded ; posterior
process heavy, weakly tectiform, tip acute and reaching half-way between
internal angles and tips of tegmina.

Tegmina narrow, entirely free, hyaline with a brown transverse band across
the center, a brown spot half-way between this band and the base, and the
apex tinged with brown; base narrowly coriaceous and punctate; veins strong
and brown ; five apical and two discoidal cells ; apical limbus broad.

Sides of thorax, undersurface of body, abdomen and legs all uniformly
brown. Legs stout; femora cylindrical; tibiae triquerate and pilose; all
tarsi about equal in length.

Length from front of head to tips of tegmina 5 mm. ; width between hum-
eral angles 2.7 mm.

Type: Female.

Type locality: Celendin, Peru.

Described from seven females and four males, three females collected at the
type locality in May 1936, all of the others collected at Limon, Peru in June
of the same year. Type, allotype and all paratypes in author ’s collection.

9. Sundarion nigromacula sp. nov. (Fig. 9)

Large, dull brown, coarsely punctate, not pubescent, with a
large black spot on each side of the inferior margin of the pro-

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Journal New York Entomological Society *[Vol. xlviii

notum just behind the middle; suprahumerals short, sharp,
extending directly outward ; posterior process slender, tectiform,
reaching half-way between internal angles and tips of tegmina;
tegmina free, smoky hyaline; sides of thorax and undersurface
of body brown; legs simple, brown, hind tarsi longest. Near
S. hrunniventris Fairmaire but smaller and with very differently
shaped suprahumerals.

Technical description ;

Head subtriangular, rough, brown with coarse black punctures and a black
spot on each side of upper margin; not pubescent; base regularly arcuate;
eyes large, ovate, glassy ; ocelli conspicuous, a little nearer to each other than
to the eyes and situated about on a line drawn through centers of eyes; in-
ferior margins of gense strongly sloping and weakly sinuate; clypeus feebly
trilobed, extending for half its length below inferior margins of genae, tip
broadly rounded and weakly pilose.

Pronotum dark brown, coarsely punctate, not pubescent, convex in front;
metopidium vertical, broader than high; median carina percurrent; humeral
angles weak, triangular, acute; suprahumeral horns short, triquerate, not
half as long as the distance between their bases, tips very sharp and black;
dorsum roundly sloping; sides of pronotum weakly impressed near margin
behind humeral angles; posterior process slender, tectiform decurved, tip
acute and reaching to a point half-way between internal angles and tips of
tegmina.

Tegmina entirely exposed, smoky hyaline; base narrowly brown, coriaceous
and punctate; veins strong; five apical cells, median apical cell truncate at
base ; one elongate discoidal cell ; apical limbus broad.

Sides of thorax light brown; undersurface of body brown; legs simple,
brown; hind tarsi longest.

Length from front of head to tips of tegmina 8 mm.; width between tips
of suprahumerals 5.3 mm.

Type: Female.

Type locality : Tingo-Maria, Peru.

Described from a single specimen taken in September 1937 by Mr. Woyt-
kowski. Type in author’s collection.

10. Ceresa luteimaculata sp. nov. (Fig. 10)

Small, brown with yellow markings, punctate, pubescent ;
pronotum convex, strongly impressed above lateral margins;
suprahumerals short, sharp extending directly outward ; posterior
process long, acute, decurved, reaching beyond internal angles of
tegmina; sides of pronotum and tip of posterior process marked
with bright golden yellow ; abdomen black ; legs ferruginous.

Sept., 1940]

Funkhouser : Membracid^

283

Technical description:

Head subtriangular, wider than high, roughly sculptured, shining, brown
mottled with black; base feebly arcuate; eyes large, globular, black; ocelli
large, prominent, brown, much nearer to each other than to the eyes and
situated about on a line drawn through centers of eyes; inferior margins
of genae sinuate and sloping; clypeus extending for half its length below
inferior margins of genae, tip rounded and pilose.

Pronotum brown, gibbous, coarsely punctate, sparingly pubescent;
metopidium vertical, subquadrate, a little wider than high; median carina
strongly percurrent; humeral angles heavy, triangular, blunt, extending out-
ward slightly farther than the eyes ; suprahumeral horns short, conical,
blunt, no longer than half the distance between their bases, extending di-
rectly outward; a broad yellow band extending from base of suprahumerals
to the lateral margins of the pronotum; sides of pronotum impressed and
black; posterior process slender, subconical, not ridged, tip sharp, bright
yellow, somewhat depressed and reaching to a point about half-way between
internal angles and tips of tegmina.

Tegmina entirely free, hyaline; base narrowly yellow, coriaceous and
punctate; veins heavy and curved; five apical and two elongate discoidal
cells; apical limbus broad.

Sides of thorax brown and pubescent; legs uniformly ferruginous, simple,
pilose; abdomen black.

Length from front of head to tips of tegmina 6 mm.; width between tips
of suprahumerals 2.7 mm.

Type : Female.

Type locality: Shishmay, Peru.

Described from a single specimen collected in September 1937 by Mr.
Felix Woytkowski. Type in author’s collection.

11. Ceresa grisescens sp. nov. (Fig. 11)

Greenish with dark grizzly markings, coarsely punctate, spar-
ingly pubescent; suprahumerals robust, extending directly out-
ward, distinctly depressed at tips; sides of pronotum deeply
impressed above lateral margins ; posterior process heavy,
strongly depressed, reaching to a point half-way between internal
angles and tips of tegmina; tegmina hyaline with tips clouded;
sides of thorax greenish; undersurface griseus; hind femora
strongly marked with black.

Technical description:

Head subtriangular, lightly punctate, green with strong black longitudinal
striae; base regularly arcuate; eyes globular, greenish; ocelli large, amber-
colored, conspicuous, equidistant from each other and from the eyes and
situated a little below a line drawn through centers of eyes; inferior mar-

284

Journal New York Entomological Society [Vol. XLVIII

gins of genae strongly sloping and weakly sinuate; clypeus long, greenish
with black stripe on each side, extending for more than half its length below
inferior margins of genae, tip blunt and densely pilose.

Pronotum convex, greenish with strong black scattered punctures which
give a grizzly appearance; metopidium sloping, broader than high; median
Carina percurrent; humeral angles strong, triangular, blunt; suprahumeral
horns robust, subconical, as long as half the distance between their bases,
extending directly outward, tips blunt and strongly depressed; sides of
pronotum strongly semicircularly impressed above lateral margins; dorsum
regularly arcuate; posterior process heavy, subconical, distinctly curving
downward, tip sharp and reaching a point half-way between internal angles
and tips of tegmina.

Tegmina entirely free, hyaline; base narrowly coriaceous and punctate;
apical and anal areas marked with brown; veins strong and curved; typi-
cally five apical and two discoidal cells but with the costal apical cells
inclined to be subdivided; apical limbus broad.

Sides of thorax greenish, abdominal segments black edged with white ; legs
simple, hind femora strongly marked with black; hind tarsi longest.

Length from front of head to tips of tegmina 8 mm,; width between tips
of suprahumerals 4.2 mm.

Type: Male.

Type locality: Limon, Peru.

Described from two males and one female all taken at the type locality
by Mr. Woytkowski in June 1936. Type, allotype and paratype in author’s
collection. The female is a little larger and less griseus than the male.

12. Stictolobus nitidus sp. nov. (Fig. 12)

Brilliant shining reddish-brown with a golden spot in the
middle of the dorsum ; pronotum convex anteriorly and tectiform
posteriorly ; no suprahumerals ; posterior process straight, reach-
ing just beyond internal angles of tegmina; tegmina smoky-
hyaline clouded with shining brown ; undersurface and legs bright
shining brown.

Technical description :

Head subquadrate, twice as broad as high, smooth, shining, not punctate
nor pubescent; base lightly arcuate and sinuate; eyes large, globular,
glassy; ocelli conspicuous, amber-colored, about equidistant from each other
and from the eyes and situated on a line drawn through centers of eyes;
inferior margins of genae sloping and sinuate; clypeus broad, not extending
below lateral margins of genae and continuing the inferior outline of the
face made by those margins, tip truncate and pilose.

Pronotum bright shining reddish-brown with a golden mark in the center
of the dorsum, weakly punctate, not pubescent; metapidium sloping, much
broader than high ; median carina almost obsolete over metapidium but

Sept., 1940]

Funkhouser : Membracid^

285

sharp on posterior process; humeral angles weak and rounded; sides of
pronotum strongly impressed behind humeral angles; posterior process
straight, tectiform, tip acute and reaching just beyond internal angles of
the tegmina.

Tegmina entirely free, smoky-hyaline ; base, central area and tip marked
with shining brown ; veins heavy and much curved ; five apical and two
discoidal cells; median apical cell petiolate; apical limbus broad.

Sides of thorax, undersurface of body and abdomen bright reddish-brown;
legs simple, shining brown; hind femora strongly marked with black;
hind tarsi longest.

Length from front of head to tips of tegmina 7.2 mm.; width between
humeral angles 3 mm.

Type : Female.

Type locality: Limon, Peru.

Described from a single specimen collected in June 1936. Type in
author’s collection.

13. Stictolobus marginatus sp. nov. (Fig. 13)

Shining jet-black with a bright yellow stripe on the inferior
lateral margins of the pronotum, coarsely punctate, sparingly
pubescent ; pronotum convex anteriorly and rounded posteriorly ;
no suprahumerals ; sides of pronotum deeply impressed above
lateral margins ; posterior process subcylindrical, depressed,
reaching to a point about half-way between internal angles and
tips of tegmina ; tegmina entirely free, subhyaline ; undersur-
face of body black; legs slender, ferruginous; hind tarsi very
much longer than the others.

Technical description;

Head subquadrate, black, broader than high, roughly sculptured, shining,
not punctate nor pubescent; base weakly arcuate and sinuate; eyes ovate
and gray; ocelli large, amber-colored, nearer to each other than to the eyes
and situated on a line drawn through centers of eyes; inferior margins of
genae sloping; clypeus extending for half its length below inferior margins
of genae, tip pointed and pilose.

Pronotum shining black with a bright yellow stripe on lateral margins,
coarsely punctate, sparingly pilose with long bristly hairs ; metopidium
sloping, wider than high; median carina very faintly percurrent; humeral
angles heavy and rounded, edged with yellow; no suprahumeral horns; sides
of pronotum deeply impressed in a semicircle above lateral margins;
posterior process swollen at base, then suddenly narrowed and extending to
a point about half-way between internal angles and tips of tegmina, a
yellow band just before the tip.

286

Journal New York Entomological Society [Vol. XLVIII

Tegmina entirely exposed, cloudy hyaline ; base narrowly coriaceous,
brown and punctate ; veins weak • five apical and three discoidal cells ; apical
limbus broad and wrinkled.

Sides of thorax dark brown; abdomen black; legs simple, slender, fer-
ruginous, tibiae margined with yellow; hind tarsi very long.

Length from front of head to tips of tegmina 6 mm.; width between tips
of humeral angles 2.4 mm.

Type: Female.

Type locality: San Martin, Peru.

Description from a single female collected by Mr. Woytkowski in August
1936. Type in author’s collection.

14. Amastris peruviana sp. nov. (Fig. 14)

Large, dull green, punctate, pubescent; pronotum roundly
arcuate, laterally compressed, highest behind humeral angles;
metopidium nearly vertical ; posterior process triangular as seen
from the side, almost reaching the tips of the tegmina; tegmina
about half covered by sides of pronotum, smoky-hyaline ; under-
surface and legs ferruginous. Near A. project a Funkhouser but
the pronotum does not overhang the head, and the tegmina are
distinctly clouded.

Technical description:

Head subtriangular, finely punctate and pubescent; base weakly arcuate
and strongly sinuate; eyes large, ovate and black; ocelli conspicuous, equi-
distant from each other and from the eyes and situated slightly below a
line drawn through centers of eyes; inferior margins of gense sloping and
weakly sinuate; clypeus subtriangular, projecting very little below inferior
margins of gen^ and continuing the apical outline of the face made by those
margins, tip rounded and pilose.

Pronotum dull green, finely punctate, sparsely pubescent; metopidium
triangular, about as broad as high, nearly vertical above the head with only
a slight backward slope; median carina strongly percurrent; humeral angles
strong, triangular and sharp ; pronotum highest behind humeral angles,
roundly arcuate and then gradually sloping to the tip of the posterior
process; sides of pronotum without ridges or impressions; posterior process
heavy, tectiform, apex acute and extending almost to tips of tegmina.

Tegmina about half covered by sides of pronotum; dark smoky hyaline
with brown spot at tip; base narrowly coriaceous and punctate; veins not
prominent; five apical and three discoidal cells; apical limbus broad.

Sides of thorax and undersurface of body ferruginous; legs stout, simple^
ferruginous, hind tarsi longest.

Length from front of head to tips of tegmina 6.5 mm.; width between
tips of humeral angles 3.4 mm.

Sept., 1940]

Funkhouser: Membracid^

287

Type: Female.

Type locality: San Martin, Peru.

Described from a single specimen collected by Mr. Woytkowski in August
1936. Type in author’s collection.

15. Tynelia nigra sp. nov. (Fig. 15)

Large, entirely black, punctate, pubescent; head projecting
strongly forward with lateral margins distinctly flanged ; dorsum
gradually arcuate; posterior process extending just to tips of
tegmina ; tegmina about half exposed with very heavy black veins ;
legs and undersurface of body black.

Technical description:

Head very long and projecting strongly forward, smooth, shining, dark
brown, deeply sulcate on each side of lateral margin; base curved downward
in middle; eyes large, subtriangular, gray-black; ocelli large, prominent,
glassy, about equidistant from each other and from the eyes and situated
above a line drawn through centers of eyes; genae foliaceous, thin, plate-
like, strongly flanged; inferior margins of genae strongly sloping and sinu-
ate; clypeus flat, not extending below inferior margins of genae, tip rounded
and hirsute.

Pronotum black; flnely punctate, sparingly pubescent; metopidium slop-
ing, wider than high; median carina strongly percurrent; humeral angles
heavy, triangular, blunt; dorsum low, gradually arcuate, somewhat flat in
middle; sides of pronotum without ridges or impressions; posterior process
heavy, subconical, apex sharp and just reaching tips of tegmina.

Tegmina about half exposed; hyaline with very heavy, black veins; base
narrowly coriaceous and punctate; flve apical and three irregularly shaped
discoidal cells; third apical cell petiolate and transverse; apical limbus
narrow; extreme tip black.

Sides of thorax and undersurface of body black ; legs heavy, simple,
black; hind tarsi longest.

Length from tip of clypeus to tips of tegmina 7.8 mm.; width between
humeral angles 3 mm.

Type : Female.

Type locality: Leonpampa, Peru.

Described from a single specimen taken in December 1937 by Mr.
Woytkowski. Type in author’s collection.

16. Vanduzea decorata sp. nov. (Fig. 16)

Jet black with a bright orange band across the middle and with
the humeral angles edged with yellow ; eyes white ; dorsnm slightly
depressed at middle; posterior process heavy, blunt and not
reaching the tips of the tegmina; tegmina about two-thirds ex-

288

Journal New York Entomological Society [Vol. XLVIII

posed, hyaline with heavy black veins; undersurface of body
black ; legs yellow.

Technical description:

Head vertical, subtriangular, shining black edged with yellow, smooth, not
punctate, not pubescent; base extended slightly downward at middle, feebly
sinuate; eyes large, ovate and white; ocelli large, prominent, conspicuous,
pearly, farther from each other than from the eyes and situated about on a
line drawn through centers of eyes; inferior margins of gense rounded, slop-
ing, edged with bright yellow; clypeus broad, not extending below inferior
margins of genae, tip rounded and pilose.

Pronotum shining black with a large orange band across the middle, very
finely punctate, not pubescent; metopidium sloping, twice as broad as high;
median carina obsolete; humeral angles large, triangular, acute, edged with
yellow; dorsum nearly straight but with a slight indentation at middle;
posterior process heavy, broad, tip rounded and reaching a point about half-
way between internal angles and tips of tegmina.

Tegmina about two-thirds exposed; hyaline with heavy black veins; base
narrowly black, coriaceous and punctate ; five apical and three discoidal cells ;
base of median apical cell petiolate and transverse; apical limbus narrow.

Length from front of head to tips of tegmina 4.5 mm. ; width between
humeral angles 2.2 mm.

Type: Female.

Type locality: Leonpampa, Peru.

Described from a single specimen collected in December 1937 by Mr. Felix
Woytkowski. Type in author’s collection.

17. Thrasymedes virescens sp. nov. (Fig. 17)

Large, green, coarsely punctate, not pubescent; femora and
abdomen marked with black; dorsum flat and nearly straight;
posterior process acute and reaching beyond internal angles of
tegmina; tegmina entirely free and entirely hyaline with one
discoidal cell ; under surface strongly marked with black ; legs
simple.

Technical description :

Head subquadrate, three times as broad as long, yellow, smooth, shining;
base irregularly sinuate, lowest just mesad of the eyes on each side; eyes
large, ovate, black; ocelli small, glassy, equidistant from each other and
from the eyes and situated below a line drawn through centers of eyes;
inferior margins of genas sloping and sinuate ; clypeus ovate, extending very
little below inferior margins of genae and continuing the ventral outline of
the face made by these margins, tip rounded and pilose.

Pronotum low, flat, green, coarsely punctate, not pubescent; metopidium
sloping, twice as wide as high; median carina obsolete; humeral angles

Sept., 1940]

Funkhouser: Membracid^

289

weak, rounded, broad; posterior process long, heavy, cylindrical at base and
tricarinate at apex, tip tectiform, sharp, reaching to a point about one-
fourth the distance from the internal angles to the tips of the tegmina;
sides of pronotum concolorous, not ridged nor impressed.

Tegmina broad, entirely free, hyaline; base narrowly coriaceous and
punctate; veins weak and brown; five apical cells; one discoidal cell; tip
rounded; apical limbus broad.

Sides of thorax and undersurface of body black; margins of abdominal
segments black; legs simple and fuscous; femora cylindrical and strongly
marked with black; tibiae triquerate; hind tarsi much longer than the
others.

Length from front of head to tips of tegmina 7 mm.; width between
humeral angles 2.7 mm.

Type: Female.

Type locality: Santo Domingo, Peru.

Described from two females both collected at the type locality in Novem-
ber 1937. Type and paratype in author’s collection.

18. Micrutalis nigromarginata sp. nov. (Fig. 18)

Fuscous with margins of head and pronotum bordered with
black, finely punctate, not pubescent; metopidium broad and
sloping; dorsum straight; posterior process acute and reaching
just beyond internal angles of tegmina; tegmina hyaline, veins
weak and obscure ; undersurface of body yellow ; legs simple and
yellow.

Technical description :

Head subquadrate, three times as broad as high, yellow, smooth, not punc-
tate, not pubescent, shining; base weakly sinuate and strongly margined
with black; eyes large, ovate and greenish; ocelli large, prominent, yellow,
a little farther from each other than from the eyes and situated above a line
drawn through centers of eyes; inferior margins of gense sloping and sinu-
ate; clypeus subovate, smooth, extending for about one-third its length below
inferior margins of genaB, tip rounded.

Pronotum fuscous, shining, finely punctate, darker on metopidium; meto-
pidium sloping, twice as broad as high; median carina obsolete; humeral
angles broad, heavy and blunt; sides of pronotum strongly margined with
black; dorsum straight and only weakly convex; posterior process straight,
very sharp, tip black and reaching just beyond the internal angles of the
tegmina.

Tegmina entirely free, hyaline; base narrowly coriaceous and punctate;
veins weak and very obscure in basal area; four apical and no discoidal
cells; tip roundly pointed; apical limbus broad.

Sides of thorax and undersurface of body yellow; area of abdomen marked
with black; legs simple and uniformly yellow; hind tarsi longest.

290

Journal New York Entomological Society [Vol. XLVIII

Length from front of head to tips of tegmina 4.4 mm.; width between
humeral angles 2 mm.

Type: Female.

Type locality: Tingo-Maria, Peru.

Described from a single specimen collected in September 1937 by Mr.
Woytkowski. Type in author’s collection.

Plate XI

1.

2.

3.

4.

5.

6.

7.

8.

Lycoderes luteus sp. nov.
Stylocentrus rubrinigris sp. nov.
Alchisme laticornis sp. nov.
AlcMsme pinguicornis sp. nov.
Alchisme spinosa sp. nov.
Paragargara nigra sp. nov.
Aconophora erecta sp. nov.
Aconophora hrunnea sp. nov.

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

(Plate XI)

7

292

Journal New York Entomological Society

[VoL. XLVIII

Plate XII

9. Sundarion nigromacula sp. nov.

10. Ceresa luteimaculata sp. nov.

11. Ceresa grisescens sp. nov.

12. Stictolohiis nitidus sp. nov.

13. Stictolohus marginatus sp. nov.

14. Amastris 'peruviana sp. nov.

15. Tynelia nigra sp. nov.

16. Vandusea decorata sp. nov.

17. Thrasymedes virescens sp. nov.

18. Micrutalis nigro'marginata sp. nov.

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

(Plate XII)

Sept., 1940]

Proceedings of the Society

295

PROCEEDINGS OF THE NEW YORK
ENTOMOLOGICAL SOCIETY

Meeting op January 3, 1939

The annual meeting of the New York Entomological Society was held on
January 3, 1939, in the American Museum of Natural History; President
Moore in the chair with thirty members and visitors present.

The following were elected to active membership: Mr. A. Davidoff, Mr.
M. Gadol, Dr. J. Johnston, Dr. R. E. Blackwelder, Mr. George E. McKenna.
The report of the auditing committee was accepted as read.

The following officers were elected for the year 1939 :

President

Vice-President

Secretary

Treasurer

Librarian

Curator

Dr. H. T. Spieth
William T. Davis
Lucy W. Clausen
Paul T. Richard
Frank E. Watson
A. Glenn Richards

Executive Committee:

H. E. Schwarz
Dr. William Procter
Dr. F. E. Lutz
Dr. A. B. Klots
Henry Bird

Publication Committee :
Harry B. Weiss
John D. Sherman, Jr,
E. L. Bell
Dr. William Moore

Auditing Committee:
Dr. J. L. Horsfall
Max Kisliuk
G. B. Engelhardt

Delegate to the New York Acad-
emy of Sciences:
William T. Davis

The resignation of Mr. G. C. Hall was accepted with regret.

A report of the Richmond meetings was given by Dr. Horsfall, Dr. Argo,
Dr. Spieth and Mr. Engelhardt.

Lucy W. Clausen, Secretary.

Meeting of January 17, 1939

A regular meeting of the New York Entomological Society was held on
January 17, 1939; President Spieth in the chair with forty-three members
and visitors present.

Mr. Mutchler was appointed chairman of the meeting. Dr. Spieth spoke
on ‘‘Some Methods of Biological Control.’’ He gave the factual evidence
upon which the theory of control is built, the methods of control of pests,
relative state of present development and relative importance of each pest,

296

Journal New York Entomological Society [Vol. XLVIII

history of biological control, mechanical problems, and concluded his talk on
the general value of biological control.

Lucy W. Clausen, Secretary.

Meeting of February 7, 1939

A regular meeting of the New York Entomological Society was held on
February 7, 1939 ; President Spieth in the chair with fifty-five members and
visitors present.

Mr. Baumann spoke on color photography, and showed about 220 colored
pictures, explaining the lens, timing, aperture and equipment used.

Mr. F. S. Blanton brought a breath of spring to the meeting by giving
narcissus blooms to those present.

A. Bacon, Secretary pro tern.

Meeting of February 21, 1939

A regular meeting of the New York Entomological Society was held on
February 21, 1939; President Spieth in the chair with forty-five members
and visitors present.

The Secretary was instructed to write a letter of sympathy to Dr. William
Moore and to extend wishes for his speedy recovery.

Dr. Payne, the speaker of the evening addressed the members on ‘ ‘ Differ-
ential Growth in the Locust, Locusia migratoria.”

Lucy W. Clausen, Secretary.

Meeting of March 7, 1939

A regular meeting of the New York Entomological Society was held on
March 7, 1939, in the American Museum of Natural History; President
Spieth in the chair with sixty members and visitors present.

Dr. Euckes announced with regret the death of the eminent zoologist. Dr.
Wilson, of Columbia.

The speaker of the evening, Mr. Edwin W. Teale, before showing his
motion pictures, related some of the trials and tribulations he went through
in his attempts to record the ways of insects on colored motion film. He
felt, however, that the studies he made last year were only a beginning and
this year he hopes to accomplish a great deal more because of his experience.

Dr. Louis Pyenson of the School of Applied Agriculture, Farmingdale,
Long Island was nominated for membership.

Lucy W. Clausen, Secretary.

Meeting of March 21, 1939

A regular meeting of the New York Entomological Society was held on
March 21, 1939, in the American Museum of Natural History; President
Spieth in the chair with eighty members and visitors present.

Mr. Kisliuk reported the death of Dr. W. E. Britton.

Dr. L. Pyenson of the Institute of Applied Agriculture, Farmingdale,
Long Island, was elected to active membership.

Sept., 1940]

Proceedings of the Society

297

Dr. Blackwelder presented a suggestion whereby all the old accumulated
books, pamphlets and separates could be offered for sale to interested parties
and in this way the Society would be able to realize some money on an other-
wise unusable collection of material. Dr. Blackwelder and Dr. A. Glenn
Kichards were appointed by Dr. Spieth as a committee to consider this sug-
gestion, and act upon it.

Mr. Vladimir Tuma then spoke upon the ‘ ‘ Breeding of Cockroaches ’ ’ the
title of which, he explained, hardly covered the subject. Mr. Tuma’s chief
concern was to standardize tests for roach insecticides. In order to do this
he used two types of roaches, German and American, but so far has concen-
trated on the German. Over a period of three years work he found that
German roaches 17 weeks of age (at which time the females are about to lay
their eggs) are the least susceptible to insecticides. This is also true of the
Periplaneta. Although his work is not yet as complete on the American as
on the German, Mr. Tuma found that the resistance of the American roach
is greater to liquid insecticides as well as to pastes than that of the German.

Mr. L. E. Chadwick of Harvard University gave a very detailed account
of his work on the flight of insects. By the use of lantern slides he pointed
out the course of movement of the wings. This was then followed by high
speed motion pictures.

As an added lesson in flight he showed Dr. Emerton ’s motion pictures of
homing pigeons' and humming birds.

Lucy W. Clausen, Secretary.

Meeting of April 4, 1939

A regular meeting of the New York Entomological Society was held on
April 4, 1939, in the American Museum of Natural History; Vice-President
W. T. Davis in the chair with thirty-five visitors and members present.

The committee in charge of the sale of the Society’s books and pamphlets
reported that bidding was brisk and that there was every prospect of a good
income from this sale. Bids may be received until the close of the meeting,
Tuesday, April 18.

Dr. E. E. Blackwelder then spoke on ^^The Wiles and Trials of a West
Indian Staphylinid Collector.” His talk was illustrated by lantern slides.
A motorcycle with a fully equipped side-car was his chief mode of traveling
while on the islands. Dr. Blackwelder utilized all the well known tricks of
collecting and improvised a few of his own. One of the innovations was the
use of a large net on the top of a car driven along the countryside slowly
at dusk. In that way he was able to get numerous rare and desirable
species.

It was his impression that Island entomologists were conspicuous by their
scarcity.

A general discussion of Dr. Blackwelder ’s paper brought the meeting to
a close.

Lucy W. Clausen, Secretary.

298

Journal New York Entomological Society [Vol. xlviii

Meeting of April 18, 1939

A regular meeting of the New York Entomological Society was held on
April 18, 1939, in Eoosevelt Memorial; President Spieth in the chair with
forty-five members and visitors present.

It was reported that the sale of books and pamphlets sponsored by the
Society was proceeding at a rapid pace and bids would be received only
until the close of the evening’s meeting.

Mr. H. L. Fellton, Box 207, Liberty Corners, N. J., was proposed for
membership.

Dr. Bromley spoke upon the ^ ^ Hurricanes and Their Effect on Insects. ’ ’
The effect of hurricanes are felt by trees as well as insect life. After the
Texas hurricane of 1933 the insects most affected were those of large and-
more fragile structure such as Lepidoptera, certain Orthoptera, Neuroptera
and Odonata. Some insects which were favored were the soil-inhabiting
species which were aided by rainfall accompanying the storm. There was
also an increase of mosquitos.

The hurricane of 1938 was too late in the season, however, to affect insects
adversely. We will probably experience an increase in bark borers and
beetles. In a survey of the territory hit by the hurricane of last fall Dr.
Bromley f)ointed out that the youngest and oldest trees survived. There
had been four days of rain preceding the hurricane and thus tree roots
pulled easily. He predicted an increase in woodland insects, followed by a
similar increase in predators and parasites.

After a general discussion of Dr. Bromley ’s paper the meeting was
adjourned.

Lucy W. Clausen, Secretary.

Meeting of May 2, 1939

A regular meeting of the New York Entomological Society was held on
May 2, 1939, in the American Museum of Natural History; President Spieth
in the chair with forty-eight members and visitors present.

Mr. H. L. Fellton, Box 207, Liberty Corners, N. J., was elected to active
membership.

The program committee announced that the next meeting of the Society
would be in the nature of a social gathering. Dr. Needham had kindly
consented to address the members at this time.

Dr. Eichards reported that the income from the sale of books and pamph-
lets of the Society was $42.06. The question of what to do with this money
was referred to the Executive Committee.

Mr. Dietrich as chairman of the Field Committee announced that plans
were being made for a field trip of the Society to take place on June 10 at
the home of Mr. Olsen of West Nyack.

The speaker of the evening, Mr. Girth, began his discussion with a history
of the entrance of the Japanese beetle into this country about 1916. The
adult beetles begin to emerge about June 25 and reach a peak about July 25.

Sept., 1940]

Proceedings of the Society

299

Since the eggs hatch in the fall the larvae do the most damage in the fall
and early spring.

After experimenting with various media it was found that the nematodes,
Neoplectana glaseri, grow best in a veal infusion agar. Feeding upon this
substance nematodes will increase 20 fold within a week. After being aer-
ated and agitated for a week the nematodes are then ready to be placed in
the soil. It takes one week to culture them and one week to aerate them and
then they may be stored for three months.

Upon being spread on soil infested with Japanese beetles in the grub form
the nematodes enter the grubs. They then begin to multiply in the host.
In a week it dies and the nematodes enter the soil and are ready to attack
other grubs.

The nematodes are applied to infested areas with ordinary sprinkling cans.
This gives immediate quick coverage of areas. A period of two weeks is
allowed to elapse before a recount is made.

Mr. Girth’s talk was illustrated with slides showing the laboratory phase
of the work.

Lucy W. Clausen, Secretary.

Meeting of May 16, 1939

A meeting of the New York Entomological Society was held on May 16,
1939, in the American Museum of Natural History; President Spieth in the
chair with seventy members and visitors present.

Because of the social nature of the meeting it was decided that the read-
ing of the minutes of the last meeting be omitted.

Mr. Dietrich announced that there would be a field trip on Sunday, June
11, to West Nyack. Notices of routes and train schedules Avere sent out
giving further details.

Dr. Spieth brought to the attention of the Society the fact that there were
twenty-five members and subscribers who have been delinquent for at least
the past three years. A motion was made and carried that these members
receive a letter asking them to pay their debts and either pay for the Jour-
nals they have received or return them.

Mr. M. H. Sartor was elected to membership.

Dr. Needham addressed the members on his recent trip to the West Indies.

Lucy W. Clausen, Secretary.

Meeting of October 3, 1939

A regular meeting of the New York Entomological Society Avas held on
October 3, 1939, in the American Museum of Natural History; President
Spieth in the chair with twenty-six members and visitors present.

Dr. Blackwelder reported that the Society Avould sponsor another auction.
Bids may be made in three ways: (1) on the Avhole volume (2) on individual
issues and (3) on individual articles.

Mr. Edwin Hunger of 4341 Eichardson Ave., N. Y. C., was proposed for
active membership.

300

Journal New York Entomological Society [Vol. XLVIII

The Secretary read a letter from Mr. Paul Sampson, Manager of the
DuPont Co., extending to the members of the Society an invitation to visit
the DuPont building at the New York World’s Fair. Of particular interest
is the Pest Control Section under the direction of Dr. M. D. Leonard.

Mr. F. E. Watson was elected to honorary membership and the Journal
was included in this membership.

The meeting was then given over to a discussion of notes on summer col-
lecting by members.

Lucy W. Clausen, Secretary.

Meeting of October 17, 1939

A regular meeting of the New York Entomological Society was held on
October 17, 1939, in the American Museum of Natural History; President
Spieth in the chair with forty members and visitors present.

Dr. Blackwelder reported that more issues of the ‘‘Canadian Entomolo-
gist” had been found and were now in the auction. The auction was ex-
tended to the meeting a month from now.

Mr. Edwin Hunger was elected to active membership.

Dr. Spieth spoke to the members on his trip to Europe this past summer,
especially his work on the Walker and Eaton types of mayflies in the British
Museum of Natural History.

The meeting was adjourned at 9 : 00 p.m., and then turned over to the
auction and informal discussion.

Annette Bacon, Secretary pro tern.

Meeting of November 21, 1939

A regular meeting of the New York Entomological Society was held on
November 21, 1939, in the American Museum of Natural History; President
Spieth in the chair with fifty-five members and visitors present.

Dr. Spieth appointed a nominating committee consisting of Mr. E. L. Bell,
Dr. C. H. Curran and Dr. Herbert Euckes.

Miss Majorie Schwarz spoke on her trip through East Africa. Her talk
was illustrated with lantern slides.

Lucy W. Clausen, Secretary.

Meeting of December 5, 1939

A regular meeting of the New York Entomological Society was held on
December 5, 1939, in the American Museum of Natural History; President
Spieth in the chair with twenty-eight members and visitors present.

Mr. William Bjerke, 655 E. 233rd Street, New York City, was proposed
for membership.

Upon informing the Society of the death of Dr. Fall, Mr. Angell made a
motion, which was carried, that the Secretary be instructed to write a letter
of condolence to Mrs. Richmond, sister of Dr. Fall.

Sept., 1940]

Proceedings of the Society

301

A motion made by Mr. Bell, and carried, instructed the Secretary to write
a letter to Br. Moore telling him that the Society is happy to learn of his
recovery from his recent illness.

Mr. Davis exhibited a cave cricket, Tachycines asynamorons Adelung, new
to the state of New York.

Lucy W. Clausen, Secretary.

Meeting of December 19, 1939

A regular meeting of the New York Entomological Society was held on
December 19, 1939, in the American Museum of Natural History; President
Spieth in the chair with thirty-five members and visitors present.

A motion was made by Mr. Sherman and carried that the recommendations
of the Executive Committee be accepted as follows:

1. That the sums realized from the auctions of publications recently held
by the Society be placed in the general fund of the Society.

2. That the residue of odd numbers of the Casey Memoirs still in posses-
sion of the Society be placed with the individual in charge of Coleoptera
at the American Museum with power to give these odd numbers to any
individual or individuals interested in beetles who in his estimation can use
them to advantage.

3. That it be recommended to the Society that an announcement be in-
serted in the Journal to the effect that the line-cuts and half-tones of illus-
trations published in the Journal in years past shall, in so far as they are
still in existence, be available to the authors of the articles in connection
with which they originally appeared provided that the cost of packing and
shipping such line-cuts and half-tones shall be defrayed in advance by those
claiming them. If claims for such plates are not made within six months
of the time when the issue of the Journal containing the announcement
makes its appearance, the Society reserves the right to destroy the plates
that are unclaimed.

4. That it be recommended to the Society that the insect parts of volumes
73 and 74 of the Zoological Eecord for the years 1936 and 1937 be purchased
(payment to be made preferably on delivery) in order to insure the continu-
ity of this series, from which the volumes in question have unfortunately
disappeared apparently beyond recovery.

A motion was made by Mr. Bell, and carried, that the offer of $30.00 for
600 signatures on hand belonging to the Society be accepted.

A motion was made by Mr. Kisliuk, and carried, that the residue of these
signatures be distributed to anyone interested.

Mr. William Bjerke was elected to active membership.

Dr. Blackwelder then addressed the members on ‘ ‘ The Modern Basis of
Taxonomy^’ which will appear in the Journal.

Lucy W. Clausen, Secretary.

,\) ■ ; . • , j v; ■ ’ i> r ; '’'t> r r? vi.f

.(lAt ■ ■

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Sept., 1940]

Weiss: Trox Unistriatus Beauv.

303

THE DEATH-FEINT OF TROX UNISTRIATUS
BEAUV.

By Harry B. Weiss

These brief notes represent the results of some observations
made upon the death-feints of six specimens of Trox unistriatus.

These beetles feign death readily upon the slightest touch,
unless they are handled constantly. In the death-feint attitude

Duration of Successive Death-Feints of
Trox unistriatus Beauv.

Beetle A

Beetle B

Beetle C

Beetle D

Beetle E

Beetle F

(Seconds)

(Seconds)

(Seconds)

(Seconds)

(Seconds)

(Seconds)

25

70

112

20

930

6

45

55

61

4

70

4

10

30

17

5

125

1

14

29

21

4

30

1

90

10

29

3

21

25

22

9

4

8

22

14

12

' 2

7

15

13

23

1

8

9

11

14

1

7

4

6

35

1

5

5

4

10

4

2

2

7

2

2

1

6

3

7

6

4

4

1

2

3
1
1

4

5
2
1

268 267 394

20 20 15

45 1,220 12

4 94 3

Total

Average ...

304

Journal New York Entomological Society [Vol. xlviii

the head is bent down and the femora of the first pair of legs
advance toward the head. As the head bends down, the antennae
take a position under it, and before the femora of the first pair
of legs move up, the tibiae and tarsi assume a position parallel to
and close against the anterior edges of the femora. The second
and third pairs of legs are held close to the ventral surface of
the body, with the femora, tibiae and tarsi in the positions com-
monly assumed by other beetles in death feigning attitudes. The
termination of the death-feint is usually gradual, the antennae
emerging first, or the tarsi of the first pair of legs. This relaxa-
tion then extends to the other legs more or less simultaneously.

The foregoing table exhibits the duration in seconds of the suc-
cessive death-feints of six beetles, with an interval of four seconds
between each death-feint, at a temperature of 80° F., and a rela-
tive humidity of 60 per cent.

Apparently there is a wide variation in the length of the death-
feint of different specimens. In the above cases, it varied from
one second to as many as 930 seconds, when the intensities of the
outside stimulations were approximately constant. With the
exception of beetles C and F, the number of successive death-
feints ranged from 10 to 13. In F it was 4, and in C, it was 26.
At the end of these numbers the death-feints were no longer
produced and the beetles made continuous efforts to escape. It
may also be noted that the tendency is for the death-feints to
become shorter as they successively increase in numbers.

The

New York Entomological Society

Organized June 29, 1892 — Incorporated June 7, 1893
Certificate of Incorporation expires June 7, 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, 77th Street and Columbus Avenue.

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

Officers for the Year 1940

President, DR. ALEXANDER B. KLOTS College of the City of New York, N. Y.

Vice-President, MAX KISLIUK, JR Federal Bldg., New York, N. Y.

Secretary, LUCY W. CLAUSEN American Museum of Natural History

Treasurer, DR. HERMAN T. SPIETH College of the City of New York, N. Y.

Librarian, DR. R. E. BLACKWELDER American Museum of Natural History

Curator, C. E. OLSEN American Museum of Natural History

EXECUTIVE COMMITTEE

H. F. Schwarz Dr. F. E. Lutz Dr. Wm. Procter

Dr. Herbert Ruckes Henry Bird

Harry B. Weiss

Dr. W. J. Gertsch

Dr. J. L. Horsfall

J. W. Angell

PUBLICATION COMMITTEE
Dr. Wm. Moore

E. L. Bell

PROGBAM COMMITTEE

F. A. SORACI

AUDITING COMMITTEE
E. L. Bell

FIELD COMMITTEE
A. S. Nicolay

John D. Sherman, Jr.

E. Teale

G. B. Engelhardt

G. Rau

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. Herman T. Spieth, American Museum
of Natural History, New York, N. Y. Orders for back issues
should be sent to the Librarian, Dr. R. E. Blackwelder, American
Museum of Natural History, 77th St. and Columbus Ave., 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 $o.58 $9.00 $9.60

Additionals 100 ’s 60 1.44 1.92 1.92 3.00 3.00

Covers 50 copies, $2.00 ; additional 100 ’s, $1.50.

Half-tone prints It^ 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.

VoL XLVIII

No. 4

DECEMBER, 1940

Journal

of the

New York Entomological Society

Devoted to Entomology in General

Edited by HARRY B. WEISS

Publication Committee

HARRY B. WEISS J. D. SHERMAN, Jr.

WILLIAM MOORE E. L. BELL

Subscription $3.00 per Year

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

NEW YORK, N. Y.

1940

CONTENTS

Seasonal Abundance of Eggs of the Corn Ear Worm Moth
in Virginia

By W. J. Phillips and G. W. Barber 305

Distribution in New Jersey of Myllocerus castaneus

By Frank A. Soraci 318

Review and Summary of Studies of Insects Associated
with Lemna minor

By Minnie B. Scotland 319

The North American Species of the Genera Leucophora
Robineau-Desvoidy and Proboscimyia Bigot (Muscidae,
Diptera)

By H. C. Huckett 335

A Reclassification of the Tribe Obriini of LeConte (Cole-
optera, Cerambycidas)

By E. Gorton Linsley 367

Studies on the Biology of the Ephemeroptera. II. The
Nuptial Flight

By Herman T. Spieth 379

A Note on Rearing the Brood of Poilstes fuscatus Fabricius
(Hymenoptera, Vespidae)

By Albro Tilton Gaul 391

Book Notices 393

Book Review — “Fauna of British India”

By Charles H. T. Townsend 395

A Promising Fungous Pathogen of Adult Japanese Beetles
(Popillia japonica)

By Edgar G. Rex 401

Book Notice 403

The Type of the Genus Pyrrhopyge (Lepidoptera, Hes-
periidae)

By W. T. Evans 405

A New Brenthis from Alaska (Lepidoptera, Nymphalidae).

By Alexander B. Klots 413

NOTICE: Volume XLVIII, Number 3, of the Journal of
The New York Entomological Society was published
ON September 18, 1940.

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. XLVIII December, 1940 No. 4

SEASONAL ABUNDANCE OF EGGS OF THE CORN
EAR WORM MOTH IN VIRGINIA

By W. J. Phillips and G. W. Barber
Cereal and Forage Insect Investigations
Bureau of Entoaiology and Plant Quarantine

INTRODUCTION

The determination of the seasonal abundance of the different
stages of the corn ear worm {Heliothis ohsoleta (F.)) is attended
by so many difficulties that, in studying the seasonal history of
the insect, investigators have usually restricted themselves to
insectary rearings of isolated individuals under artificial condi-
tions. A more complete knowledge of the seasonal history and
abundance of the insect as it is found in nature was needed.

The fact that this insect passes part of its life in the soil makes
field counts of pupation and emergence impracticable. The
moths are rapid fliers and move about so freely that it is difficult
to make accurate observations of their habits. Observations on
the infestation of corn ears in the field may give information rela-
tive to seasonal abundance and life history, but it is by no means
certain with what degree of accuracy the numbers of larvas occur-
ring at any time may be determined. On the other hand, obser-
vations to determine the abundance of eggs can be made much
more easily and with much greater accuracy. The seasonal vari-
ations in egg abundance should indicate the prevailing moth
population with a fair degree of exactness and should also indi-
cate what the subsequent larval population will be. The occur-

306

Journal New York Entomological Society [Vol. xlviii

rence of eggs, therefore, should give the most accurate informa-
tion obtainable of seasonal history and abundance of the insect.

One of the difficulties encountered in the determination of the
seasonal history and abundance of the insect by observation of
oviposition lies in the fact that corn is more attractive to the
moths for egg laying at certain stages of its growth than at others.
It is most attractive while the plants are in the fresh-silk stage,
and least attractive while the plants are small or after they have
ripened. In the latitude of Virginia corn may be planted during
a period of more than 2 months. An early-maturing variety
planted the first of April may begin to silk by mid- June, whereas
corn planted in June may not begin to silk until September.
Since plants with fresh silks are more attractive than others for
oviposition, the concentration of eggs on plants during June, for
instance, would be far greater on the early-planted than on later-
planted corn. Therefore, the examination of a single planting
of corn would not show the actual abundance of moths through-
out the season, for, though eggs might be laid late in the season in
great numbers, they would be deposited on plants in fields that
were in more attractive stages of growth.

To determine the seasonal occurrence and abundance of eggs,
numerous plantings of corn were made throughout the growing
season at Charlottesville, Ya., each year from 1921 to 1927, ex-
cepting 1923. Similar plantings were made at Richmond, Ya.,
from 1924 to 1927. Such plantings, except during the early part
of a season, provided plants continuously in the stages of growth
favorable for oviposition. Fresh silks often appeared by the
second or third week of July and were continuously present there-
after until the first week in October.

A number of representative plants were chosen from each plot,
and, except when weather prevented, daily records were made of
the number of eggs deposited on each plant. All eggs were re-
moved each day except those on certain plants set apart for deter-
mination of the fate of the eggs. These eggs were marked and
their positions noted, so that new depositions could be easily
recognized. Observations of a given plant were discontinued 3
days after the silks had completely dried, or somewhat before
roasting-ear stage. From these records it was possible to deter-

Dec., 1940]

Phillips and Barber: Corn Ear Worm

307

mine the average deposition of eggs per plant per day and to
chart the seasonal occurrence and abundance of eggs. The ad-
vantage of this method of recording the eggs deposited on a num-
ber of plants in various stages of growth was that field conditions
as found in the section in which these studies were made were
approximately simulated except, of course, for seasonal variation
in acreage. No definite planting date for corn is usually observed
by farmers, and for this reason plants in various stages of growth
are present in the field over a period of several months.

In the manner described, 16 studies were made of the local
seasonal occurrence of the eggs of the corn ear worm in the locali-
ties and years mentioned. In all, complete oviposition records
on 891 corn plants, representing 267 plots, were obtained. A
total of 43,828 eggs were recorded, or an average of 49 eggs per
plant. A total of 52,818 plant observations were made. The
average rate of oviposition per plant per day was 0.83 for all
years and localities.

PERIODS OP OCCURRENCE OF THE CORN EAR WORM IN VIRGINIA

In the latitude of Virginia several generations of the insect
occur annually. Moths appear first between the last of May and
the middle of June, and they continue to emerge from hibernation
throughout June and July. Emergence is somewhat irregular
because of the operation of several natural factors, such as pre-
cipitation and soil temperature. Since a generation may be com-
pleted in about 5 weeks, an overlapping of broods occurs from
July onward through the season. For convenience in this study,
and because generations cannot be recognized, the season was
divided into two periods. The first period comprised roughly
May, June, and July, and the second period consisted of August,
September, and October.

CORN PLOTS USED IN THESE STUDIES

At Charlottesville, Va., three series of plots were used. These
were designated as upland, bottom, and garden plots. The up-
land plots were located in a field of clay loam which sloped toward
the west. Each plot consisted of three rows of field corn across
the field. The bottom plots, also of field corn, were located in a
river-bottom field about one-half mile south of the upland plots.

308

Journal New York Entomological Society [Vol. XLVIII

The garden plots each consisted of two rows of sweet corn in a
vegetable garden in rich loam about 600 yards south of the upland
plots. The plots at Kichmond, Va., on level upland in sandy
loam soil, consisted of field corn, and were similar in size to the
upland plots at Charlottesville.

The upland and bottom plots at Charlottesville were planted to
the same varieties of field corn and on similar dates. Common
varieties of sweet corn were used in the garden plots, plantings
being started earlier and being made at greater intervals than in
the other plantings. The plantings in four localities, in different
soil types, and at dates spread throughout the season, covered as
wide a range of environment as it was possible to observe.

SEASONAL OCCURRENCE OF EGGS

The studies were begun each year when the earliest corn plants
were about 8 inches tall, and continued until all corn had lost its
attractiveness to the moths and egg laying on it had ceased.

The date when eggs were first found on corn in any year de-
pended on two factors, (1) when moths emerged from hibernating
pup80, which was dependent on spring weather, and (2) the time
when corn was planted and the rate at which it grew, also depen-
dent on spring weather, making for earliness or lateness of the
season. The date on which the first eggs were laid on corn varied
about a month in different years. The earliest occurring eggs
were found on May 21, 1925, at Richmond. The lateness of oc-
currence of corn in attractive stages of growth in the fall, and
consequent occurrence of ear worm eggs on it, varied for similar
reasons. The latest egg recovered on corn was on October 10 in
Charlottesville upland plots in 1927. A record of egg recovery
is given in table 1. A general summary of the results of this
study is given in table 2.

Eggs were deposited on many days in each period. For each
period, however, the proportion of days on which eggs were de-
posited, based on the total number of days of observation applica-
ble to that period, was variable. Figure 1 shows the percentage
of days in each period on which eggs were deposited. Within the
first period the range was from 16.67 (bottom plots at Charlottes-
ville in 1925) to 100 per cent (upland plots at Charlottesville in

TABLE 1. — Average occurrence of corn ear worm eggs on corn plants during each 5-day period during the season {nuniher of eggs per plant per day)

TABLE 2. — Summary of seasonal occiirrence of eggs of the corn ear worm in Virginia

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June 19
July 13
July 11
July 23
July 29
Aug. 1
July 28
June 24
July 22
June 18
July 21
June 23

July 28
June 12
July 11
June 24

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312

Journal New York Entomological Society [Vol. XLVIII

1921 and at Richmond in 1927). In the second period the range
was from 53.85 per cent to three records of 100 per cent.

These studies showed that, because of the widespread occur-
rence of the eggs throughout each season, corn plants would
probably receive eggs regardless of planting date.

SEASONAL ABUNDANCE OF EGGS

The abundance of eggs in the first period was frequently some-
what irregular, but in 10 of the studies the distribution was so
uniform that no definite peaks of occurrence were found. In four
of the seasonal histories, where eggs were less evenly distributed,
more than one peak of occurrence was found, and a single definite
peak of abundance was found only in two instances.

DESCRIPTION OF LOCALITY
VIRGINIA

1921 CHARLOTTESVILLE UPLAND

1922 CHARLOTTESVILLE UPLAND

1922 CHARLOTTESVILLE BOTTOM

1924 CHARLOTTESVILLE UPLAND

1924 CHARLOTTESVILLE BOTTOM

1925 CHARLOTTESVILLE UPLAND

1925 CHARLOTTESVILLE BOTTOM

1925 CHARLOTTESVILLE GARDEN

1926 CHARLOTTESVILLE UPLAND

1926 CHARLOTTESVILLE GARDEN

1927 CHARLOTTESVILLE UPLAND

1927 CHARLOTTESVILLE GARDEN

1924 RICHMOND UPLAND

1925 RICHMOND UPLAND

1926 RICHMOND UPLAND

1927 RICHMOND UPLAND

PERCENTAGE OF DAYS ON WHICH PLANTS RECEIVED EGGS

FIRST PERIOD
0 20 40 60 80 100

SECOND PERIOD

0 20 40 60 80 100

1 i ■ 1

— 1 — 1 — 1 — 1 — 1 — 1 — \ — 1 — 1 — 1 — r-

HHB

Figure 1. Percentage of days during the several studies of the seasonal
history of the corn ear worm on which moths deposited eggs
on corn plants. Based on the total number of days on which
observations were made for the respective periods.

Numbers of eggs in the second period began to increase from
the middle to the last of August, and reached clearly defined
peaks of abundance from the middle to the last of September.
During a period of about a month eggs usually were laid very
plentifully. The egg laying during the second period was, there-
fore, usually concentrated within a relatively few days, in con-

Dec., 1940]

Phillips and Barber: Corn Ear Worm

313

trast to that of the first period, which was usually distributed over
a larger number of days. The time of maximum abundance of
eggs of the second period in 12 of the seasonal histories occurred
either on a single date or extended over several days. However,
in four instances the eggs were deposited more or less evenly dur-
ing several weeks, and no well-marked peaks of abundance were
found.

In all the seasonal occurrences studied, several days were found
when the plants were relatively, sometimes entirely, free of eggs.
The duration of the egg-free days, which usually occurred during
the last week of July or the first week of August, ranged from a
few days to 2 weeks or more.

In figure 2 the relative abundance of eggs of the first and second
periods for each of the 16 seasonal histories studied is given.
Eggs of the first period ranged in abundance from 0.01 egg per
plant per day (bottom plots at Charlottesville in 1925) to 2.04
eggs per plant per day (garden plots at Charlottesville in 1927).
In the second case eggs were 204 times as abundant as in the first.
Eggs of the second period ranged in abundance from 0.06 egg per
plant per day (upland plots at Richmond in 1924), to 7.89 eggs
per plant per day (upland plots at Richmond in 1925). Second-
period eggs at Richmond in 1925 were, therefore, 131.5 times as
plentiful as in the same location in 1924.

NUMBERS OF EGGS DURING THE TWO PERIODS

A comparison of the numbers of eggs which occurred per plant
per day in the two periods showed remarkable variation. This is
illustrated graphically in figure 2. In each period, in the differ-
ent seasonal histories, eggs ranged from scarce to abundant.
Eggs of the second period were more abundant than those of the
first period in 5 out of 6 years at Charlottesville, and in 2 out of
4 years at Richmond, and in 13 out of the 16 seasonal histories
studied. In one seasonal history eggs of each period were equally
abundant (Charlottesville upland plots of 1927). The numbers
of eggs of the second period ranged from 0.13 to 92 times the
number of eggs of the first period. These data are given in
table 2.

The years 1924 and 1927 were apparently unfavorable for
increase of this insect, while 1925 was the most favorable by far.

314

Journal New York Entomological Society [Vol. XLVIII

COMPARISON OF ABUNDANCE OF CORN EARWORM E0GS

DESCRIPTION OF LOCALITY
VIRGINIA

FIRST PERIOD

SECOND PERIOD

1921

CHARLOTTESVILLE

UPLAND

■

■

1922

CHARLOTTESVILLE

UPLAND

■

■

1922

CHARLOTTESVILLE

BOTTOM

■

■

1924

CHARLOTTESVILLE

UPLAND

■

■

1924

CHARLOTTESVILLE

BOTTOM

■

■

1925

CHARLOTTESVILLE

UPLAND

•

■

1925

CHARLOTTESVILLE

BOTTOM

•

B

1925

CHARLOTTESVILLE

GARDEN

■

■

1926,

CHARLOTTESVILLE

UPLAND

■

■

1926

CHARLOTTESVILLE

GARDEN

■

■

1927

)

CHARLOTTESVILLE

UPLAND

■

■

1927

CHARLOTTESVILLE

GARDEN

■

■

1924

RICHMOND

UPLAND

■

■

1925

RICHMOND

UPLAND

■

■

1926

RICHMOND

UPLAND

■

■

1927

RICHMOND

UPLAND

■

■

Figure 2. Comparison of abundance of eggs of the corn ear worm during
two periods of the year for 16 seasonal-occurrence studies.
The areas of the squares indicate the number of eggs deposited
per plant per day.

RELATIONSHIP OP PRECIPITATION TO ABUNDANCE OP EGGS

While many factors influence the population of this insect, as
determined by the number of eggs found on corn plants, none
seems to be more important than precipitation during the oviposi-

Dec., 1940]

Phillips and Barber: Corn Ear Worm

315

tion period. Dry weather is definitely favorable in enabling the
moths to lay their full complement of eggs, and in permitting a
high rate of hatching and survival of young larvae. Since
drought is usually accompanied by high temperatures, such con-
ditions cause more rapid development of the various stages of the
insect. During the course of these studies one year of severe
drought occurred, 1925, especially during September. The insect,
in response to favorable conditions, built up enormous popula-
tions in September. Eggs of the second period at Charlottesville
were 15.17 times, and at Kichmond 14.35 times, as plentiful as
eggs of the first period.

The greatest rainfall occurring during any year of the study
was at Richmond in 1927, and as an effect of this, eggs of the
second period were 0.13 times as plentiful as eggs of the first
period. The records of 4 years of study at Richmond showed
that the proportion of eggs of the second to those of the first
period varied directly with the precipitation during the months
from June to September. For Charlottesville this relationship
was less direct, probably because of the influence of other factors.
These data are given in table 3.

summary

Because of the habits of the corn ear worm, it is not easy to
determine the seasonal occurrence or abundance of the insect by
means of counts of the pupae, moths, or larvae. It was thought
that this information could be obtained by counts of eggs de-
posited on corn plants, as the numbers of eggs might reflect moth
abundance and later larval populations indirectly.

Daily examination of selected corn plants of successive plant-
ings in two localities and four environments, during 6 years, gave
data on 16 seasonal records of egg occurrence.

Much difference was found in the seasonal occurrence of eggs.
This depended in part on earliness or lateness of the spring or
fall.

In each seasonal occurrence studied there were a number of
days in the last week of July or the first week of August when
eggs were extremely scarce or wholly wanting. This time of egg
scarcity was used to divide each season into two periods, the first

316

Journal New York Entomological Society [Vol. xlviii

CO

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observation).

Dec., 1940]

Phillips and Barber: Corn Ear Worm

317

comprising roughly May, June, July and the second consisting of
August, September and October.

Abundance of eggs varied greatly in different years. Eggs of
the first period ranged from 0.01 to 2.04 per plant per day. Eggs
of the second period ranged from 0.06 to 7.89 per plant per day.
When most plentiful, eggs of the first period were 204 times as
numerous as in the year of least abundance, and eggs of the
second period were 131.5 times as numerous as when least
plentiful.

While eggs of the second period were usually more plentiful
than those of the first period, in 3 of 16 instances they were less
abundant, and in one case the numbers were equal. Numbers of
eggs of the second period ranged from 0.13 to 92 times those of
the first period.

Precipitation seemed to be a principal factor in determining
the abundance of the eggs. During seasons of much rain the ear
worm population increased little. During seasons of little rain
populations increased greatly.

DISTRIBUTION IN NEW JERSEY OF (MYLLOCERUS)
CORIGETUS? CASTANEUS RCELOFS

This weevil, catalogued in the “Fourth Supplement 1933 to
1938 (inclusive) to the Leng Catalogue of Coleoptera of Amer-
ica, North of Mexico,” by Richard E. Blackwelder, Mount Ver-
non, N. Y., Dec. 1939, as “ Curculionidse : Myllocerus castaneus
Roelofs. Jap. Sib.? N. A.,” was taken (probably from oaks) at
Montclair, N. J., in July, 1933, by A. S. Nicolay. A note by
A. C. Davis, Takoma Park, Maryland, appearing in Bui. Brooklyn
Ent. Soc., Vol. 30, 1935, p. 19, recorded this find, and the fact
that L. L. Buchanan, of the Bureau of Entomology and Plant
Quarantine of the United States Department of Agriculture, to
whom a specimen was submitted for identification, “states that
it evidently does not belong in the genus Myllocerus, and refers
it tentatively to Corigetus. ”

In November 1939, a New York State Nursery Inspector, Mr.
R. L. Clement, reported to the New Jersey Department of Agri-
culture that this insect had been found on Long Island and that
it appeared to be causing considerable damage by feeding on oak
foliage.

In the late summer of 1940, a rough survey was made in New
Jersey to determine the distribution of the insect and the amount
of damage caused by it. The present infestation can be described
as extending from the northern tip of Bergen County to Denville
in Morris County to Plainfield in Union County to Port Lee in
Bergen County. The insect was also found, close to the New
Jersey shore, from Atlantic Highlands to Allenhurst. The few
observations on feeding habits of this insect would indicate that
although it does feed extensively on deciduous foliage (the oaks,
primarily), heavy damage is not apparent until late in the season
(late August or early September) and therefore injury to the
host would be slight. — Frank A. Soraci.

REVIEW AND SUMMARY OF STUDIES OF INSECTS
ASSOCIATED WITH LEMNA MINOR*

By Minnie B. Scotland
N. Y. State College for Teachers, Albany, N, Y.

From 1931 to 1939 I have made observations of the animal
population of Lemna minor found in the vicinity of Ithaca, New
York. My interest has been centered upon insects that were
more or less dependent upon Lemna, especially upon the Lemna
fly, Lemnaphila scotlandce Cress. This concentration led to the
further study of parasites that emerged from the Lemna fly’s
puparia.

In making these surveys I have had little difficulty in obtaining
duckweed, Lemna minor, for it abounds generally where there
is quiet water, as in ditches, marshes, ponds, and small nooks
along the banks of slowly flowing streams. Associated with
Lemna minor are often large amounts of Wolffia and Spirodela,
and infrequently small quantities of Lemna trisidca. The rain-
fall of a summer determines the available regions of quiet water
on which Lemnacese can propagate. I have found tanks at the
Cornell University fish hatchery (Plate XIV, Figure 4) and small
ponds entirely covered by dense growths. The summer of 1939
was an unusually dry one for the vicinity of Ithaca and conse-
quently many previous collecting spots were entirely bare.
However, there was enough Lemna minor gathered to continue
my records of animal life of this interesting association.

When I began these studies relatively few species of insects
had been recorded as associated with duckweed. Among these
there were tw(5 beetles entirely dependent upon this plant. A
duckweed beetle, S dries tibialis Guer., was described in 1918 by
W. C. Kraatz, who found it feeding entirely on Lemna minor in
waters near Madison, Wisconsin. The other, a duckweed weevil,
Tanysphyrus lemnce Fab., was reported by C. von Urban in the
Entomologische Blatter (1922) as dependent upon duckweed for
food. Lemna had been listed as the food plant of two aphids.

* Contribution of the Entomology Laboratory, Cornell University, Ithaca,

320

Journal New York Entomological Society [Vol. XLVIII

In 1910 T. A. Williams gave duckweed as food of Bhopalosiphum
nympJmm Linn, and in 1918 H. F. Wilson and R. A. Vickery re-
corded Lemna minor as a food plant of Aphis samhuci Linn.

Records of other uses of duckweed by insects include a paper
in 1911 by Wm. T. M. Forbes in which be mentioned Lemna thalli
as the material used for making the case of the aquatic caterpillar
Elophila. In the same report he said of this case, “It is distin-
guished from those of Nymphula and Hydrocampa by the lack of
a sharp lateral edge,” implying that the cases of these aquatic
caterpillars were also constructed of Lemna. He observed that
the “leaves of the case do not seem to be eaten.” Another paper
which appeared in 1919 was H. B. Hungerford’s, “The biology
and ecology of aquatic and semi-aquatic Hemiptera,” in which
he reported the use of the under surface of Lemna thalli for the
attachment of the eggs of the aquatic bug, Trepohates pictus H.
S. These appear to have been the only definite references to the
relations of insects with Lemna minor before my observations be-
gan in 1931.

Since that time I have published one general paper (1934) on
“The Animals of the Lemna Association,” in which I listed
fifteen species of insects, and a second more specialized paper
(1939) on “The Lemna Fly and Some of its Parasites.” In 1938
F. X. Williams described and figured a similar Ephydrid,
Hydrellia williamsi Cresson, as being found “on Lemna-covered
water” in Hawaii and having “much the same habits as
Lemnaphila scotlandce Cress.”

My own observations have been made on the following more
permanent resident insects in the Lemna association : one
Collembolan {Sminthurus aquaticus Bourlet), one Hemipteran
{Mesovelia hisignata Uhl.), one Homopteran {Bhopalosiphum
nympJmce Linn.), one Coleopteran {Tanysphyrus lemnce Fab.),
one Lepidopteran {Nymphula ohliteralis Wlk.), one Dipteran
{Lemnaphila scotlandce Cress.), and three Hymenopteran
parasites {Opius lemnaphilce Muesebeck,, Trichopria angusti-
pennis Muesebeck, Trichopria paludis Muesebeck).

The habits of these Lemna insects may be summarized as
follows :

The diminutive spring-tail, Sminthurus aquaticus Bourlet,

Dec., 1940]

Scotland: Lemna Insects

321

jumps about on the floating mats of duckweed. Occasionally
pairs will be seen to jump together, the yellow colored male being
carried by the mauve colored female. The male has a clasping
device on the second and third antennal segments composed of
hooks and tubercles. These lock around the straighter antennae
of the female. The pair meet face to face, lock antennae, and
then, the female raises the male at an angle varying from a few
degrees to as many as ninety. Sometimes a pair will remain to-
gether in this fashion for more than an hour. Duckweed serves
this species as a source of food and also as a place for depositing
the eggs. Many small holes are made in the thalli in the process
of the prolific egg laying. Some idea of the damage to the plant
can be gained from Plate XIII, Figure 3.

The bright green nymphs of the water strider, Mesovelia
bisignata Uhl., are common on duckweed. The nymphs run
quickly over the thalli and can easily dart out over the open
water without breaking through the surface film. Lemna minor
functions as a convenient plant in which the adults can lay their
eggs. The elongate-oval eggs with curved necks are inserted
singly between the epidermal layers of the Lemna thallus, their
tips being exposed on the upper side, where they appear as shiny,
membranous, ringed spots. Kelatively little destruction of
Lemna is caused by these carnivorous bugs.

The “pond-lily Aphid,” Ehopalosiphum nymphcece Linn., is
always very abundant in my collections. These aphids move
about on the thalli and on the water, apparently unhindered by
any moisture on their tarsi. They are very prolific. Viviparous
females produce young almost daily during August. They feed
upon the sap in Lemna thalli. One aphid may remain on a single
plant for hours. No appreciable injury to the duckweed seems
to result from the microscopic feeding punctures of this aphid
and evidently the loss of sap is easily overcome by the natural
metabolic processes in the plant.

The duckweed weevil, Tanysphyrus lemnce Fab., is commonly
found in the egg, larva, and adult stages on floating Lemna.
The pupa is less frequently seen. In the laboratory pupge
were obtained by placing grown larvae in masses of sphagnum
and leaving them there to pupate. This is the method I offered

322

Journal New York Entomological Society [Vol. XLVIII

in 1937 (Culture Methods, p. 480). Scattered clumps of Lemna
thalli on moist earth also served as satisfactory cover under which
wandering larvae pupated. The mining larvae devour the green
interior of the thalli, leaving whitened plants, while the adults
use their snouts to eat out spherical holes from the upper side
of the Lemna. The destruction is very great, as may be seen
in Plate XIII, Figure 2.

An aquatic case-making moth, Nymphula {Ilydrocampa)
ohliteralis Wlk., is usually present in collections of duckweed. A
dirty-whitish larva binds together overlapping thalli for a case.
When deprived of one case, it proceeds to make a new one imme-
diately. It nibbles off bits of duckweed for food. In the labora-
tory a single larva was placed with a quantity of Lemna in a
Syracuse watch glass. This insect ate so many of the plants
that only a few were left for its pupal case. Certainly Lemna
minor is used for the growth of this moth.

The Lemna fly, Lemnaphila scotlandm Cress., was discovered
by me in 1933 and sent to Mr. E. T. Cresson for determination.
It was found by him to be a new genus and species of the
Ephydridae. It is very small (length 1.2 mm.) ; so small that it
has the unique ability among all the Lemna insects of being able
to pass through its whole life cycle on and within a single thallus.
However, it is not strictly limited to a single plant. It may
be found moving from one thallus to another, even crossing
intervening stretches of open water. The larva is a miner, and
on hatching it bores into the center of the thallus for food and
shelter (Plate XIV, Figure 9). Here, also in the mine, the
amber-colored puparium is found (Plate XIV, Figure 6). Nor-
mally there is only one pupa in a plant. However, recently, I
found two pupae crowded in a single thallus. The adult fly
(Plate XIV, Figures 7, 8, 10-12) crawls out of the puparium be-
tween the epidermal layers of the plant and walks about on the
duckweed. Very soon it begins to eat by gouging out portions of
Lemna, leaving characteristic parallel streaks (Plate XIII, Figure
1 ) . These peculiar markings on thalli prove an aid in detecting
the presence of the Lemna fly in any quantity of duckweed. The
feeding activities of this fly result in the almost complete de-
struction of the thalli. The female attaches her dainty melon-

Dec., 1940]

Scotland: Lemna Insects

323

shaped eggs to the margin of the thallus, as shown in Plate
XIII, Figure 1 and Plate XIV, Figure 5.

Three specific parasites on the Lemna fly (Plate XVI) were
likewise unknown. One has been determined as a new species
of Opius (0. lemnaphilce Muesebeck) and two as new species of
Trichopria (T. angustipennis Muesebeck and T. paludis Muese-
beck). All of these parasites emerge from fly puparia, leaving
tell-tale holes in the empty shells, as shown in Plate XVI, Figures
20 and 21. Comparison of the numbers of these parasites in
1938 and 1939 revealed a marked reduction in the total number
found in 1939 but a relatively constant ratio of the different
species. In 1939 as in 1938 there were more individuals of Tri-
chopria than of Opius. Of the Trichopria there were three times
as many T. angustipennis as T, paludis. Males predominated.

In addition to these resident Lemna insects, all of which are
of some importance in the economy of the plant, there are several
transient visitants and chance associates commonly present.
These are included below in a comprehensive list^ of insects
of the Lemna association.

COLLEMBOLA

Sminthurus aquaticus Bourlet

Podura aquatica — Larger than 8. aquaticus, found rarely.

ORTHOPTERA

Undetermined nymphs of crickets — Observed running over a
mat of duckweed. Captured individuals nibbled edges of the
thalli.

EPHEMERIDA and ODONATA

Undetermined nymphs — Foragers among the roots of duck-
weed.

thysanoptera

Limothrips cerealium Hal. — Probably from nearby grasses.

* The taxonomic sequence is that followed by Leonard in A List of the
Insects of New Yorh. Cornell Univ. Agr. Ex. Sta. Mem. 101, Aug. 1926.

324

Journal New York Entomological Society [Vol. XLVIII

HEMIPTERA

Mesovelia hisignata Uhl.

Trepohates pictus H. S. — Hungerford (1919) described and
showed figures of eggs on the under surface of floating duck-
weed.

Gerris sp. Found on surface of Lemna mat.

Notonecta sp.

Plea striola I Common aquatic bugs that move among the
Banatra sp. ( roots of Lemna.

Corixa sp.

HOMOPTERA

Aphis saml)uci Linn. Not observed by me.

Bhopalosiphum nymphcece Linn. Very common.

COLEOPTERA

Tanysphyrus lemnce Fab.

Scirtes tibialis Guer. Not observed by me.

Haliplus sp. Larvae and adults scurry among roots.
Laccophilus sp. Larvae.

Hydroporus sp. Lays its eggs singly on the pendant Lemna
roots.

Tropistermis lateralis Fab. Larvae observed foraging among
roots.

TRICHOPTERA

Limnophilus combinatus Walk. Used bits of Lemna in con-
structing its case.

LEPIDOPTERA

Nymphula obliteralis Wlk.

Elophila sp. Not observed by me.

DIPTERA

Lemnaphila scotlandce Cress.

Hydrellia williamsi Cress. Not observed by me.

Corynoneura scutellata Winn. Larvae, pupae, adults.
Dasyhelea traverce Thom. Larvae, pupa.

Tetanocera sp.

Culex sp. Eggs, larvae, pupae.

Dec., 1940]

Scotland; Lemna Insects

325

Anopheles sp. ‘‘ “ “

Chironomus sp. Cases.

Odontomyia sp. Larvae and pupae.

Eristalis tenax — Larvae.

HYMENOPTERA — All parasltes.

Opius lemnaphiloe Muesebeck 1

Trichopria angustipennis Muesebeck ^Parasites of Lemna fly

Trichopria paludis Muesebeck J

Undetermined genus near Aphanta Foerster

Polynema sp. Parasite of Mesovelia eggs embedded in Lemna

thalli.

Tiphodytes gerriphagus
Anaphoidea pullicrura

In a summary like this I cannot refrain from mentioning the
unexpected manner in which new fields of investigation often
present themselves. It was while I was searching for Hydro-
porus eggs in 1931 that these studies of the Lemna association
began. I found the beetle eggs glued to the roots of duckweed.
Upon looking through library sources I could discover no ecolog-
ical study of the very common pond plant, Lemna minor.
Thereupon, I undertook what has been a labor of love and what
promises to be a life long inquiry into the wonders of this diminu-
tive society.

BIBLIOGEAPHY

Cresson, E. T. 1933. A new genus and species of the dipterous family
Epliydridse reared from duckweed. Ent. News 44: 229-231.
Forbes, Wm. T. M. 1911. Another aquatic caterpillar (Elophila). Psyche
18: 120-121.

Hungerford, H. B. 1919. The biology and ecology of aquatic and semi-
aquatic Hemiptera. Kansas Univ. Sci. Bull. 11: 99-106.

Kraatz, W. C. 1918. Scirtes tibialis Guer., (Coleoptera-Dascyllidae), with
observations on its life history. Ann. Ent. Soc. Amer. 11 : 393.
Muesebeck, C. F. W. 1939. Three new hymenopterous parasites of the
Lemna fly. Proc. Ent. Soc. Wash. 41: 58-62.

Scotland, M. B. 1934. The animals of the Lemna association. Ecology 15 :
290-294.

. 1937. The duck -weed weevil, Tanysphyrus lemncB, in Culture meth-
ods for invertebrate animals by J. G. Needham and others, Ithaca,
N. Y.: pp. 480-481.

. 1939. The Lemna fly and some of its parasites. Ann. Ent. Soc.

Amer. 32: 713-718.

326

Journal New York Entomological Society [Vol. XLVIII

Urban, C. von. 1922. Zur lebensweise der Tanysphyrus lemnce Payk.
Ent. Blatt. 18 : 73-75.

Williams, F. X. 1938. Biological studies in Hawaiian water-loving insects.

Part III. Biptera or flies. A, Epliydridse and Antliomyiidse.
Proc. Haw. Ent. Soc. 10 : 86-90.

Williams, T. A. 1910. The Aphididae of Nebraska. Univ. Studies 10: 71.
Wilson, H. F. and E. A. Vickery. 1918. A species list of the Aphididae
of the world and their recorded food plants. Wis. Acad. Sci., Arts
and Letters, Trans. XIX : 153.

Plate XIII

Figure 1.

Figure 2.

Figure 3.

Infested Lemna, showing injuries

The work of the adult Lenina fly, Lemnaphila scotlandce Cress.,
which appears as parallel gougings. Nine of the thalli show
eggs of this fly on their margins.

The work of the duckweed weevil, Tanysphyrus lemnce Payk.,
showing the round feeding punctures made by the adult weevil
and the irregular linear mines of the larva. One larva is
visible through the transparent epidermis in a mine in the
upper right.

The work of the collembolan, Sminthurus aquaticus Bourlet. The
round holes are the excavations in which the eggs were de-
posited singly.

(Jour. N. Y. Bnt. Soc.), Vol. XLVIII

(Plate XIII)

328

Journal New York Entomological Society [Vol. XLVIII

Figure 4

Figure 5

Figure 6

Figure 7
Figure 8

Plate XIV

. A tank at the Fish Hatchery of Cornell University, Ithaca, N.
Y., showing a thick floating mat of lemna, the joint habitat of
the lemna fly, the case-bearing moth, the Lemna weevil, the
pond-lily aphid, the water strider, and the Collembolan.

. The ribbed and sculptured egg of Lemnaphila scotlandce Cress.
The micropyle is seen on the right.

. Empty puparium showing the rupture through which the fly has
emerged.

. Wing of the Lemna fly.

. Antenna of same.

(Jour. N. Y. Ent. Soc.), Vol. XL VIII

(Plate XIV)

330

Journal New York Entomological Society

[VOL. XLVIII

Plate XV

Figure 9. Tip of the mouth-hooks of the larva of Lemnaiihila scotlandce
Cress., showing the serrated margin of the inner surface of
one hook. The other hook of the pair, diverging in position,
shows dimly, being out of focus. A palpus shows faintly in
the lower left.

Figure 10. Adult Lemna fly.

Figure 11. Tip of proboscis of same with labella retracted.

Figure 12. The same, more enlarged and with labella expanded, showing the
hooks that gouge the parallel rows of holes in the Lemna
^ thallus. A labial palpus appears at the left below.

(Jour. N. Y. Bnt. Soc.), Vol. XLVIII

(Plate XV)

332

Journal New York Entomological Society [Vol. XLVIII

Figure 13.
Figure 14.
Figure 15.
Figure 16.
Figure 17.
Figure 18.
Figure 19.
Figure 20.

Figure 21.

Plate XVI

Parasites of Lemnaphila scotlandce Cress.

Opius lemnaphilce Muesebeck, adult female.

Antenna of same.

Terminal segments of same, showing sensillae.

Antenna of Trichopria angustipennis Muesebeck.

Terminal segments of same, showing pit-like sensillse.

Wing of Opius lemnaphilce Muesebeck.

Wing of Trichopria angustipennis Muesebeck.

Empty puparium of Lemnaphila scotlandce Cress., showing the
large hole from which Opius lemnaphilce Muesebeck has
emerged.

Empty puparium of Lemnaphila scotlandce Cress., from which
Trichopria angustipennis Muesebeck has emerged.

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

(Plate XVI)

20

21

' ''v. ’

T-'^ '' ' '^ . ". '"'r • # m V

THE NORTH AMERICAN SPECIES OF THE GENERA
LEUCOPHORA ROBINEAU-DESVOIDY AND
PROBOSCIMYIA BIGOT (MUSCID^,
DIPTERA)

By H. C. Huckett

Eiverhead, N. Y.

The genus Leucophora Robineau-Desvoidy^ belonging to the
subfamily Anthomyiinge is of particular interest to students of
insect life on account of the fact that the larvae, so far as is known,
live as inquilines^ or parasites® in the nests of solitary bees and
wasps. It is recorded that the flies of many of the species be-
longing to this genus have the habit of shadowing bees and wasps
as the latter approach their burrows, which are situated in sandy
or gravelly soils.^ In this way the nests of hymenopterous hosts
are probably detected by the female for purposes of oviposition.
Despite these peculiar habits the adults of the group are not
to be readily distinguished on the basis of generic characters from
many species occurring in allied groups associated with Hylemyia
sens.-lat. The flies of the genus Leucophora differ principally
but not invariably by having the genal, facial and occipital
regions of the head more buccate, interfrontalia in female
narrower, frons in this sex approaching the proportions of that
in male. The thorax is robust and abdomen in male conical and
rarely longer than thorax, ovipositor armed with recurrent spines
on anal palpi except in maculata, legs proportionately stout, wings
with costal thorns vestigial, m-cu cross veins in many species

1 In the sense of Hylephila and Hammomyia (Eondani) Coqnillett. Proc.
U. S. Nat. Mus., 1910 XXXVII p. 560.

2 Huie, L. H. The habits and life history of Hylemyia grisea Fall., an
anthomyid fly new to the Scottish fauna. The Scottish Naturalist, 1916 No.
49 Jan. p. 13-20.

3 Seguy, E. Contribution a Petude des dipteres anthomyides du genre
Hylephila. Comptes rendus du Congres des Societes savantes en 1925, 1926
p. 474.

4 Collin, J. E. A contribution towards the knowledge of the anthomyid
genera Hammomyia and Hylephila of Eondani (Diptera). Trans. Ent. Soc.
London, 1921 Pt. Ill p. 305-326.

336

Journal New York Entomological Society [Vol. XLVIII

oblique and sinuate and veins B. 4 + 5 and M. 1 + 2 convergent
distad. Hypopygiuni and copulatory appendages in male of
normal structure, not shining; in both sexes the apical scutellar
setulag are more robust than those of Prosalpia. The arbitrary
nature of such a combination of characters may, without further
knowledge, serve only to confuse rather than to clarify the
definition of the genus. Undoubtedly one of the most important
sources of information relating to the scope of the genus is to be
found in the facts concerning the peculiar habits of the species.

Most authors have treated the group as composed of two
segregates, and have aiDplied to them Rondani’s names Hylephila
and Hammomyia respectively. In dealing with North American
species this classification seems scarcely practical owing to the
intergrading of many of the characters proposed for the separa-
tion of the different segregates. I have been unable to find addi-
tional characters that might help to support Rondani’s action,
and hence have assembled the species into one genus under the
name Leuocophora Robineau-Desvoidy, as proposed earlier by
Coquillett.®

The following species are recorded as occurring in North
America

Leucophora alhiseia (von Roser)
johnsoni (Stein)
maculata (Stein)
marylandica (Malloch)
ohtusa (Zetterstedt)
sociata (Meigen)
unilineata (Zetterstedt)
unistriata (Zetterstedt)
fusca n. sp.
annexa, n. sp.

Genus Leucophora Robineau-Desvoidy

Leucophora Robineau-Desvoidy, Essai Myod., 1830 p. 562 . . .

Coquillett, Jour. N. Y. Ent. Soc., 1901 IX p. 138 .. .

Coquillett, Proc. U. S. Nat. Mus., 1910 XXXVII p. 560.

5 Loc. cit., p. 560.

Q Hammomyia setigera Johannsen (Trans. Amer. Ent. Soc., 1916 XLII p.
387) belongs to Hylemyia, being closely related to H. inornata Stein.

Dec., 1940]

Huckett: Leucophora

337

Chortophila Macquart, Hist. Nat. d. Ins. 1835 II p. 328 . . .

Meade, Descr. List Brit. Anth., 1897 II p. 43.

Anthomyia Schiner, Faun. Austr., 1862 I p. 638.

Hylephila Rondani not Billberg, Dipt. Ital., Prodr., 1877 VI p.
13, 233 . . . Strobl, Verb, zool.-bot. Ges. Wien, 1893 XLIII
p. 263 . . . Schnabl and Dziedzicki, Abb. K. Leop. -Carol.
Deutscb. Akad. Naturforsch., 1911 XCV Nr. 2 p. 91 . . .
Stein, Arch. f. Naturgesch., 1916 (1915) LXXXI A belt 10
p. 158, 222 . . . Johannsen, Trans. Amer. Ent. Soc., 1916
XLII p. 388 .. . Stein, Arch. f. Naturgescb., 1920 (1918)
LXXXIV A heft 9 p. 84 . . . Collin, Trans. Ent. Soc. Lon-
don, 1921 Pt. Ill p. 311, 306 . . . Seguy, Faune de France,

1923 VI p. 72 . . . Seguy, Compt. Congr. Soc. Savantes,
1926 p. 473 . . . Karl, Tierwelt Deutschlands, 1928 XIII Pt.
3 p. 192 . . . Malloch, Diptera of Patagonia, 1934 Pt. VII
fasc. 2 p. 187 . . . Seguy, Gen. Insect., 1937 Fasc. 205 p. 135.

Hammomyia Rondani, Dipt. Ital., Prodr., 1877 VI p. 13, 236 . . .
Strobl, Verb, zool-bot. Ges. Wien, 1893 XLIII p. 265 . . .
Aldrich, Misc. Coll. Smithsn. Inst., 1905 XLVI No. 1444 p.
554 . . . Stein, Kat. Paliiark. Dipt., 1907 III p. 698 . . .
Williston, Man. N. A. Dipt., 3rd ed., 1908 p. 336 . . . Schnabl
and Dziedzicki, Abh. K. Leop. -Carol. Deutscb. Akad. Natur-
forsch., 1911 XCV Nr. 2 p. 91 . . . Stein, Arch. f. Natur-
gesch., 1914 (1913) LXXIX A heft 8 p. 32 . . . Stein, Arch,
f. Naturgesch., 1916 (1915) LXXXI A heft 10 p. 163, 222 .. .
Johannsen, Trans. Amer. Ent. Soc., 1916 XLII p. 387 . . .
Stein, Arch. f. Naturgesch., 1919 (1917) LXXXIII A heft 1
p. 152 . . . Stein, Arch. f. Naturgesch., 1920 (1918)

LXXXIV A heft 9 p. 84 . . . Malloch, Canad. Ent., 1921
LIII p. 78 . . . Collin, Trans. Ent. Soc. London, 1921 Pt.
Ill p. 306, 307 . . . Seguy, Faune de France, 1923 VI p.
70 . . . Huckett, Mem. 77 N. Y. (Cornell) Agr. Exp. Sta.,

1924 (1923) p. 13 . . . Karl, Tierwelt Deutschlands, 1928
XIII Pt. 3 p. 191 . . . Malloch, Diptera of Patagonia, 1934
Pt. VII Fasc. 2 p. 189 . . . Curran, Fam. Gen. N. A. Dipt.,
1937 p. 391.

Anthomyia (Hammomyia) Pandelle, Rev. ent. France, 1901 XX
p. 203, 299.

338

Journal New York Entomological Society [Vol. XLVIII

Genotype Leucophora cinerea Eobineau-Desvoidy ( = Aricia
albescens Zetterstedt).

The genus Leucophora was described by Eobineau-Desvoidy
(1830)^ for the reception of five nominal species, none of which
have since been recognized with any degree of confidence by sub-
sequent authors. A careful study of the original description of
genus and species has lead me to the conclusion that the same
general group is here depicted that was later to be described by
Eondani as two segregates.

Macquart (1835) apparently did not recognize Eobineau-
Desvoidy ^s genus, and forthwith cited the first four species of
Leucophora as a part of his heterogeneous group Chortophila.

Eondani (1877) in dealing with the classification of Italian
diptera erected the allied genera Hylephila and Hammomyia, and
designated Musca huccata Fallen as the type of the former genus
and Aricia albescens Zetterstedt as the type of the latter. In the
case of albescens the author included in synonymy the species
Leucophora cinerea Eobineau-Desvoidy.

Coquillett (1901) in a preliminary list of the types of
anthomyid genera designated L. cinerea, the third of five original
species, as the genotype of Leucophora. Stein (1907) in his
catalogue of palagarctic diptera cited Leucophora and Hylephila
as synonyms of Hammomyia Eondani, and recorded cinerea and
albescens among the synonyms of Hammomyia albiseta (von
Eoser). Coquillett (1910) in his later compilation of the type
species of North American genera of Diptera included both of
Eondani ’s genera as synonyms of Leucophora, accepting the
name Leucophora cinerea in the sense of Aricia albescens Zetter-
stedt.

Subsequent authors (Stein, 1916, 1920) (Collin, 1921) (Seguy,
1923, 1925) (Malloch, 1921, 1934) (Karl, 1928) have main-
tained in a large measure the classification proposed by Eondani,
although some doubt has been expressed concerning the validity
of Hylephila as a name for one of the genera, owing to its possible
preoccupation in Lepidoptera.

7 Citations of literature are given in abbreviated form in the synonymy,
the year of publication being inserted within parentheses.

Dec., 1940]

Huckett: Leucophora

339

Seguy (1937) in his recent monograph on the Muscidse of the
world combined both of Rondani’s segregates into one genns
under the name Hylephila Rondani.

In my opinion the name Leucophora was rendered valid by
Coquillett’s action in designating Leucophora cinerea Robineau-
Desvoidy as the type of the genus. I have regarded the identity
of cinerea as equivalent to that of Aricia albescens Zetterstedt,
according to the concensus of opinions expressed in the literature.
Zetterstedt ’s species is the genotype of Hammomyia Rondani,
and on that account I have considered Hammomyia as a synonym
of Leucophora. I have come to the conclusion that the generic
name Hylephila as used by Rondani cannot stand owing to its
preoccupation by Billberg.® If Rondani’s group is to be main-
tained it seems that a new name should be proposed to displace
Hylephila. This genus possesses priority over Hammomyia,
hence in any attempt to merge the two groups it would appear
that the name Hammomyia would be sunk in synonymy. How-
ever in my opinion the need for a new name does not arise, if
for no other reason than the fact that there already exists the
prior claims of Leucophora Robineau-Desvoidy.

I have not thought is advisable, at least for the purposes of this
study, to make any further reference concerning the possible
status of the remaining species described in Leucophora because
of lack of evidence or authority regarding their probable identity
for specific purposes.

8 The name Hylephila was employed firstly in 1820 by Billberg (Enumer-
atio Inseetorum in Museo Billberg, 1820 p. 81) to denote a genus in Lepi-
doptera. Although the validity of Billberg ’s genera has been indirectly
questioned by Walsingham and Durrant (Eevision of the nomenclature of
micro-lepidoptera. Ent. Month. Mag., 1902 XXXVIII p. 163-170) and later
by others on account of the unfortunate circumstances surrounding Billberg ’s
work these authors evidently did not take into consideration the attempt
made by Scudder (Historical sketch of the generic names proposed for
butterflies: A contribution to systematic nomenclature. Proc. Amer. Acad.
Arts Sciences, 1875 X p. 193) to revive Billberg ’s genus Hylephila. In
Scudder ’s treatment of the genus the names of three original species are
retained, of which Papilio pJiylaeus Drury is selected as the genotype of
Hylephila. The authorship of the genus is credited to Billberg. In my
opinion this action served to validate the name Hylephila Billberg for the
group.

340

Journal New York Entomological Society [Vol. XLVIII

Key to Males

1. Mid tibia with a bristle on anteroventral or mid ventral surface 2

- Mid tibia with no bristle on either ventral surface 7

2. Notopleural callosity with several setulse 3

- Notopleural callosity devoid of setulae 4

3. Scutellum fuscous on lateral and ventral surfaces; lower caudal sterno-

pleural bristle weakly developed fusca n. sp.

- Scutellum uniformly grayish on lateral and ventral surfaces; lower

caudal sternopleural bristle stoutly developed annexa n. sp.

4. Tibiae, parafacials and cheeks normally largely reddish; aristal hairs

longer than basal diameter of arista; prealar bristle absent; cross
veins cloudy; cruciate setulse present johnsoni (Stein)

- Tibiae normally largely blackish; arista minutely pubescent or bare;

cross veins clear 5

5. Mid tibia with a bristle on anteroventral surface; fore tibia with a

robust apical posterodorsal bristle; cruciate setulae present.

unilineata (Zett.)

- Mid tibia with a bristle on mid ventral surface ; fore tibia lacks a robust

apical posterodorsal bristle; cruciates absent 6

6. Prealar bristle as long as posterior notopleural bristle; planes of dorso-

central and acrostical bristles usually noticeably setulose; abdominal
vitta blackish ohtusa (Zett.)

- Prealar bristle shorter than posterior notopleural bristle, or absent;

planes of dorsocentral and acrostical bristles not noticeably setulose;
abdominal vitta brownish unistriata (Zett.)

7. Prealar bristle present 8

- Prealar bristle usually absent 11

8. Notopleural callosity with several setulae; parafacials broader ventrad

than at base of antennae; proboscis slender annexa n. sp.

- Notopleural callosity invariably devoid of setulae; parafacials broader

at base of antennae and narrower ventrad; proboscis not slender 9

9. Processes with a fringelike series of long bristles from base to apex that

is ventrally directed; prealar bristle short maculata (Stein)

- Processes with no pronounced series of longish bristles directed ventrad,

if long directed caudad; prealar bristle as long as posterior noto-
pleural bristle 10

10. Bristles on inner border of processes short and stiffish, stouter developed

than bristles on outer border (fig. 13) ; bristles on antero- and postero-
ventral surfaces of hind femur not longer than greatest breadth of
femur alhiseta (von Eoser)

- Bristles on inner border of processes fine and slender, not stouter devel-

oped than those on outer border (fig. 15) ; bristles on antero- and
posteroventral surfaces of hind femur longer than greatest breadth of
femur sociata (Meigen)

11. Aristal hairs longer than basal diameter of arista; tibise reddish tinged;

processes with no noticeable series of bristles directed ventrad 12

Dec., 1940]

Huckett: Leucophora

341

- Aristal hairs not longer than basal diameter of arista; tibiae black;

processes with a noticeable series of downwardly directed bristles.

maculata (Stein)

12. Processes reddish yellow ; mid tibia with apical anterodorsal bristle more
robust than apical posterodorsal; m-cu cross vein usually cloudy and
semierect joJmsoni (Stein)

- Processes blackish; mid tibia with apical posterodorsal bristle more

robust than apical anterodorsal; m-cu cross vein clear and usually
obliquely directed marylandica (Malloch)

Key to Females

1. Mid tibia with a bristle on anteroventral or mid ventral surface 2

- Mid tibia with no bristle on either ventral surface 7

2. Tibiae reddish yellow; cross veins cloudy; aristal hairs longer than basal

diameter of arista johnsoni (Stein)

- Tibiae blackish; cross veins clear; aristal hairs not longer than basal

diameter of arista 3

3. Notopleural callosity with several setulae 4

- Notopleural callosity devoid of setulae 5

4. Scutellum fuscous on lateral and ventral surfaces ; fore tibia with at least

three well developed apical bristles; setulae on mesopleura encroaching
cephalad onto declivity dorsad of mesothoracic spiracle fusca n. sp.

- Scutellum grayish on lateral and ventral surfaces, concolorous with

dorsum; fore tibia with two well developed apical bristles; meso-
pleural setulae not encroaching onto declivity dorsad of mesothoracic
spiracle annexa n. sp.

5. Fore tibia with a robust apical posterodorsal bristle; mid tibia with

bristle on anteroventral surface; cruciate setulae present; outer pair
of verticals bristlelike unilineata (Zett.)

- Fore tibia lacks a robust apical posterodorsal bristle; mid tibia with

bristle on mid ventral surface; cruciates absent; outer pair of verti-
cals setulose 6

6. Prealar bristle as long as posterior notopleural bristle; abdominal vitta

blackish; first abdominal sternum with several setulae; middle four
bristles of marginal series on tergum 5 situated further from caudal
margin than those laterad obtusa (Zett.)

- Prealar bristle shorter than j)Osterior notopleural bristle, or absent;

abdominal vitta brownish; first abdominal sternum bare; middle four
bristles of marginal series on tergum 5 situated about the same dis-
tance from margin as those laterad unistriata (Zett.)

7. Prealar bristle present 8

- Prealar bristle usually absent 9

8. Palpi flattish, narrowly spatulate; hind femur with a stoutish bristle on

posteroventral surface sociata (Meigen)

- Palpi slender, filiform; hind femur with no stoutish bristle on postero-

ventral surface albiseta (von Eoser)

342

Journal New York Entomological Society [Vol. XLVIII

9. Erons at narrowest about equal to one-third width of head viewed from
above; aristal hairs longer than basal diameter of arista; ovipositor

armed with recurrent spines on anal palpi marylandica (Malloch)

- Erons less than one-third width of head ; arista minutely pubescent ; ovi-
positor lacks recurrent spines on anal palpi (figs. 19, 20).

maculata (Stein)

Leucophora alMseta (von Eoser)

Leucophora cinerea Kobineau-Desvoidy, Essai Myod., 1830 p.
563 . . . Coquillett, Jour. N. Y. Eiit. Soc., 1901 IX p. 138
. . . Coquillett, Proc. U. S. Nat. Mus., 1910 XXXVII p. 560.
Chortophila cinerea Macquart, Hist., Nat. d. Ins., 1835 II p. 328.
Anthomyia alhiseta von Poser, Wiirttemb. Corrbl., 1840 I p. 59.
Aricia albescens Zetterstedt, Dipt. Scand., 1845 IV p. 1520.
Anthomyia albescens Scbiner, Faun. Austr., 1862 I p. 638 . . .

Neuhaus, Diptera marchica, 1886 p. 228.

Hammomyia albescens Rondani, Dipt. Ital., Prodr., 1877 VI p.
236 . . . Strobl, Verb, zool.bot. Ges. Wien, 1893 XLIII p.
265 . . . Collin, Trans. Ent. Soc. London, 1921 Pt. Ill p. 308
. . . Seguy, Faune de France, 1923 VI p. 71 . . . Karl,
Tierwelt Deutschlands, 1928 XIII Pt. 3 p. 191.

Chortophila albescens Meade, Ent. Month. Mag., 1882 XIX p.

146 . . . Meade, Descr. List Brit. Anth., 1897 II p. 44.
Anthomyia {Hammomyia) albescens Pandelle, Eev. Ent. France,
1901 XX p. 300.

Hammomyia albiseta Stein, Kat. Palaark, Dipt., 1907 III p. 698
. . . Schnabl and Dziedzicki, Abh. K. Leop.-Carol. Deutsch.
Akad. Naturforsch., 1911 XCV Nr. 2 p. 91 . . . Stein, Arch,
f. Naturgesch., 1914 (1913) LXXIX A heft 8 p. 32 . . . Stein,
Arch. f. Naturgesch., 1916 (1915) LXXXI A heft 10 p. 163
. . . Tiensuu, Acta Soc. Faun. Flor. Fenn., 1935 LVIII No.
4 p. 14.

Hylephila albiseta Seguy, Gen. Insect., 1937 Fasc. 205 p. 136.
Records

Alaska, 2J', Fairbanks, July 1 1921 (J. M. Aldrich).

British Columbia, 3J', Nicola, June 27 1923 (C. B. D. Gar-
rett) ; Seton Lake, Lillooet, June 2 1926, 2^?, June
4 1926 (J. McDunnough).

California, IJ, Felton, St. Cruz Mts., 300-500 ft.. May 15-19
1907, (J. C. Bradley).

Dec., 1940]

Huckett: Leucophora

343

Colorado, IJ', IJ, Campus of University of Colorado, Boulder,
July 1910, (T. D. A. Cockerell).

Idaho, 1$, Clementsville, July 22 1926 (R. W. Haegele).

Nova Scotia, IJ', Kentville, Aug. 14 1917.

Oregon, IJ, Niger’s Island, June 11 1925 (J. Wilcox).
Washington, SJ', Clarkston, June 12 1930 (J. M. Aldrich)
[U. S. N. M.]

The male of all)iseta has been commonly characterized as hav-
ing a plain dove gray color on thorax and whitish dust on frontal
regions of head. The above specimen from Nova Scotia is typical
in these respects, but those from Alaska and British Columbia
possess a much darker aspect and resemble in many ways the
allied species sociata (Meigen). If it were not for the structure
of the copulatory appendages (figs. I, 12, 13) and for the fact
that many of the species represented in this study exhibit a
similarly wide range in tonal forms it would be a difficult matter
to reconcile the fact that these western specimens represented the
same species as the dove gray examples of albiseta from Europe.
The female of albiseta has the palpi filiform, whereas in sociata
they are narrowly spatulate in this sex.

Leucophora johnsoni (Stein)

Hylemyia johnsoni Stein, Berl. Ent. Zeitschr., 1898 (1897) XLII
p. 215 . . . Aldrich, Misc. Coll. Smithsn. Inst., 1905 XLVI
p. 552 . . . Britton, Bull. 31 Conn. Geol. Nat. Hist. Surv.,
1920 p. 198 . . . Johnson, Occ. Pap. Boston Soc. Nat. Hist.,
No. VII 1925 p. 234.

Hammomyia johnsoni Stein, Arch. f. Naturgesch., 1919 (1917)
LXXXIII A heft 1 p. 152 . . . Stein, Arch. f. Naturgesch.,
1920 (1918) LXXXIV A heft 9 p. 84 . . . Malloch, Canad.
Ent, 1921 LIII p. 78 . . . Huckett, Mem. 77 N. Y. (Cornell)
Agr. Exp. Sta., 1924 (1923) p. 13 . . . Hallock and Parker,
Circ. 103 N. J. Dept Agr., 1926 p. 16.

Hylephila johnsoni Seguy, Gen. Insect., 1937 Ease. 205 p. 138.
Records

Idaho, 2J^, Mt. Moscow, June 1924 (J. M. Aldrich) [U. S.
N. M].

New York, IJ', Glen Head, Long Island, April 14 1921; 13J',
1?, Valley Stream, Long Island, April 27 1921; 3^;^, Lake-

344

Journal New York Entomological Society [Vol. XLVIII

ville, Long Island, May 22 1921 ; IJ, Babylon, Long Island,
May 22 1933, 1?, June 15 1935 (F. S. Blanton) ; 3c?, Heck-
scher State Park, Long Island, May 30 1935, IJ, Babylon,
Long Island, May 24 1934, Ic?, May 22 1935, IJ', May 26
1935, 1$, June 9 1934, 1$, June 15 1934, Half Way
Hollow Hills, Long Island, May 18 1935 (Blanton &
Borders) ; 4c?, Middle Island, Long Island, May 30 1931;
15, Kiverhead, Long Island, May 27 1934, IJ', June 26
1927 ; IJ', 15, Ithaca, May 28 1922.

Ontario, Ic?, Niagara Glen, June 23 1926, 1$, June 24 1926 (G.
S. Walley).

Quebec, IJ, Covey Hill, June 15 1927 (G. S. Walley).

Texas, 2J', College Station, April 13 1935, 2c?, April 18 1935,
15, April 10 1932, !<?, April 21 1936 (R. J. Reinhard).

The typical form of johnsoni may be described as possessing
yellowish tibiae, cloudy cross veins, longish aristal pubescence,
but lacking the prealar bristle. There are however specimens
which differ in one or more respects to the above combination of
characters. In occasional examples the cross veins may be clear
and the presence or absence of prealar bristle difficult to deter-
mine on account of the development of setae in this region. In
the male type, which is preserved in the collections of the Field
Museum of Natural History, the mid tibia possesses a mid ventral
bristle. This character is lacking in most of the above male
specimens, but is present in the female sex. I have tentatively
regarded the two male specimens from Idaho as aberrant forms
of this species. In these specimens the fore and mid tibiae are
largely blackened, arista apparently subnude, abdomen grayish
drab, and aedeagus not conforming in structural details to that
of eastern examples of johnsoni. The proper status of such
specimens may be more confidently determined on the examina-
tion of a larger series of specimens from western States.

Leucophora marylandica (Malloch)

Hammomyia marylandica Malloch, Trans. Amer. Ent. Soc.,

1920 XLVI p. 185 .. . Malloch, Canad. Ent., 1921 LIII p.

78 . . . Huckett, Mem. 77 N. Y. (Cornell) Agr. Exp. Sta.,

1924 (1923) p. 14 . . . Hallock and Parker, Circ. 103 N. J.

Dept. Agr., 1926 p. 16.

Dec., 1940]

Huckett: Leucophora

345

Hylephila marylandica Johnson, Occ. Pap. Boston Soc. Nat.

Hist., No. VII 1925 p. 236 . . . Johnson, Insect. Fann. Biol.

Surv. Mt. Desert Region, 1927 p. 211 . . . Seguy, Gen.

Insect., 1937 Fasc. 205 p. 138.

Records : -

Alberta, IJ, Waterton, July 8 1923 (H. L. Seamans).

British Columbia, IJ, Kaslo, June 18 (R. P. Currie)

[U. S. N. M.]

Colorado, IJ', Paonia, July (G. E. Quinter) [U. S. N. M.]

Georgia, IJ^, no data [U. S. N. M.]

Massachusetts, IJ, Brewster, June 25 1937 (Campau).

Michigan, IJ', Stevensville, Berrien Co., May 29 1938 (C.
W. Sabrosky).

New Jersey, IJ^, Rancocas Park, April 23 1925 (H. C. Hal-

lock) ; 12, Delaware Water Gap, June 21 . (A. T.

Slosson).

New York, 2J', Babylon, Long Island, May 11 1935 (Blanton
& Borders) ; 2(^, Selden, Long Island, May 12 1924; 7J',
Riverhead, Long Island, May 6 1923, May 18 1923, Ic?,
May 20 1923, 2^2, 12, May 27 1923, Icf, May 29 1923, 3J', 12,
May 3 1927, 2,^2, May 8 1927, IJ^, May 15 1927, 3^, 42, May
22 1927, 2c?, May 4 1938.

Ontario, Ic?, Low Bush, Lake Abitibi, June 20 1925 (N. K.
Bigelow).

Quebec, 12, Meach Lake, June 21 1916 (W. T. M. Forbes).

Texas, 2c?, College Station, April 13 1935 (H. J. Reinhard).

The species marylandica resembles the european form Musca
grisea Fallen, from which it may be distinguished by the shorter
aristal hairs, and by the absence of a robust apical posterodorsal
bristle on fore tibia. Among North American species mary-
landica may be associated with Hylemyia johnsoni Stein on ac-
count of the longish aristal hairs and absence of prealar bristle,
but the female of marylandica may be readily separated from
that of johnsoni owing to the lack of a mid ventral bristle on mid
tibia and by the broader frons, the latter being at least as wide
as distance between the second pair of dorsocentral bristles. In
the male the distinguishing characters can not be so readily de-
fined owing to the marked extent of variation within johnsoni.

346

Journal New York Entomological Society [Vol. XLVIII

The male of marylandica, in common with the female, invariably
possesses a more obliquely directed m-cu cross vein that not
unusually divides the distal half of vein M.1+2 into two nearly
equal sections; whereas in johnsoni the cross vein is usually
placed in a more upright position and the penultimate section of
ilf.1+2 is thereby noticeably shorter than the ultimate.

Leucophora sociata (Meigen)

Anthomyia sociata Meigen, Syst. Beschr., 1826 V p. 98.
Hylemyia sociata Neuhaus, Diptera marchica, 1886 p. 224.
Hydrophoria sociata Stein, Kat. Palaark. Dipt., 1907 III p. 687.
Hammomyia gallica Schnabl and Dziedzicki, Abh. K. Leop. -Carol.

Deutsch. Akad. Naturforsch., 1911 XCV Nr. 2 p. 236, 355.
Hammomyia sociata Stein, Arch. f. Naturgesch., 1916 (1915)
LXXXI A heft 10 p. 163 .. . Collin, Trans. Ent. Soc. Lon-
don, 1921 Pt. Ill p. 309 . . . Seguy, Panne de Prance, 1923
No. VI p. 72 . . . Karl, Tierwelt Deutschlands, 1928 XIII
Pt. 3 p. 192.

HylepJiila sociata Seguy, Gen. Insect., 1937 Pasc. 205 p. 138.
Records : -

Alberta, IJ', Calgary, July 14 1923 (G. Salt).

British Columbia, IJ, Oliver, May 30 1923, 1$, June 3 1923,
IJ, Keremeos, June 27 1923 (C. B. D. Garrett) ; 1^, Seton
Lake, Lillooet, June 1 1926, 1.^, June 4 1926 (J. McDun-
nough).

California, IJ, San Gabriel Canyon, Oct. 21 1929.

Colorado, 1$, Tennessee Pass, 10240 ft. alt., July 11 (J.

M. Aldrich) [U. S. N. M.]

Idaho, IJ', Mt. Moscow, July 12 1924, 2^, Lake Waha, June
14 1930 (J. M. Aldrich) ; 2J', Mts., Moscow, June 25 1920
(R. C. Shannon) ; 1,J^, Emigration Canyon, Aug. 24 1934
(T.) 0. Thatcher).

Manitoba, 1$, Winnipeg, June 22 1927 (H. J. Brodie) ; I5,
Aweme, Oct. 7 1924 (N. Criddle).

Maryland, I5, Plummers Island, June 22 1916 (A. K. Pisher).
New Hampshire, IJ', Pranconia, (A. T. Slosson).

Utah, IJ', Uintah Mts., Tryol Lake, (J. C. Pechser).
Washington, IJ', Ewan, June 13 1920 (R. C. Shannon) ; IJ^,

Dec., 1940]

Huckett: Leucophora

347

Asotin, April 15 1923, IJ, May 20 1923 (Virgil Argo) ; IJ',
Mt. Rainier, Shadow Lake, 6200 ft. alt., July 29 1932, IJ',
July 31 1932; 1^, Potlatch, May 28 1930 (J. M. Aldrich).
[U. S. N. M.].

Wisconsin, IJ, Fountain City, Buffalo Co., Aug. 12-17 1910.
The species sociata may be regarded as allied to aVbiseta, the
darker forms of the latter superficially resembling this species.
In sociata the bristles on antero- and posteroventral surfaces of
hind femur are more robust than in alhiseta^ being abnormally
stoutly developed in some specimens. There are also specific
differences observable in the structure of male copulatory ap-
pendages (figs. 4, 9). In the female of sociata the palpi are
narrowly spatulate, being pointed apicad.

Leucophora maculata (Stein)

Hammomyia maculata Stein, Berl. Ent. Zeitschr., 1898 (1897)
XLII p. 229 . . . Aldrich, Misc. Coll. Smithsn. Inst., 1905
XL VI No. 1444 p. 554 . . . Malloch, Trans. Amer. Ent. Soc.,
1918 XLIV p. 303 .. . Malloch, Canad. Ent., 1921 LIII p.
78 . . . Stein, Arch. f. Naturgesch., 1919 (1917) LXXXIII
A heft 1 p. 152 . . . Stein, Arch. f. Naturgesch., 1920 (1918)
LXXXIV A heft 9 p. 84.

Hylephila maculata Seguy, Gen. Insect., 1937 Fasc. 205 p. 138.
Records : —

Alberta, IJ', Wainwright, June 24 1938 (E. H. Strickland) ;
2J', 15, Waterton, July 9 1923, 1^, Lethbridge, July 6 1921
(H. L. Seamans).

British Columbia, IJ', Lillooet, June 15 1917, IJ', June 14 1917
(J. D. Tothill) ; Ic?, Chilcotin, July 26 1920 (E. R. Buckell).
California, IJ', Corona, March 25 1932, IJ', Riverside, March
14 1933, 1^, March 30 1933, 2^?, April 10 1933, 1^, April 1
1933, IJ', April 3 1933 (A. J. Basinger).

Colorado l^f, Florissant, June 19 (Cockerell) [U. S.

N. M.].

Idaho, 3c?, 1$, Moscow, June 24 1912, 2c?, 1?, June 16 1910
(U. S. N. M.) ; Ic?, McCall, June 22 1926, 10525 ft. alt.; 1?,
Fairfield, Aug. 27 1926 (R. W. Haegele).

Manitoba, IJ, Balmoral, Aug. 24 1924 (A. J. Hunter).

348

Journal New York Entomological Society [Vol. xlviii

Montana, IJ, Missoula, July 13 (U. S. N. M.).

North Dakota, IJ', Mandan, June 16 1918 (U. S. N. M.).

Oregon, 2^, Hood Kiver, no date (Childs).

South Dakota, IJ', Hot Springs, July 13 1924.

Utah, IJ, Payson, no date (M. H. Peterson).

Washington, IJ', Friday Harbor, June 19-26 1909 ; 2^^, IJ,
Lind, June 11 1919, 1^?, June 16 1919; 1^, Edwall, July 8

(J. M. Aldrich) [U. S. N. M.] ; 1^, Ritzville, June 12

1920, 15, Lake Paha, June 20 1920, IJ, Sprague, June 20
1920 (R. C. Shannon) ; IJ', 1$, Toppenish, June 19 1923,
15, Mt. Rainier, White River, July 20 1924 (A. L. Melan-
der).

Wyoming, IJ', Yellowstone Park, Madison Jnct., July 27 1923
(A. L. Melander).

The species maculata is apparently western in its distribution,
and does not occur in New York as previously recorded (Huckett
1924, Leonard 1928). The latter citations are based on a female
specimen collected at Ithaca which agrees closely with the female
cotype of Stein. Stein’s female specimen is, in my opinion, not
conspecific with the male cotype, and differs from the females of
the above records in having recurrent spines on anal palpi, and in
the spatulate form of oral palpi. In the above female specimens
the terminal sclerites of the ovipositor have a characteristic
padded or cushionlike appearance owing to the enlarged form
and dense vestiture of the subanal plate ; the recurrent spines are
not developed (figs. 19, 20). The male of maculata may be dis-
tinguished by the notable series of long bristles arising from
abdominal processes. The bristles are conspicuous on account of
their ventral position. The species covers a wide range in gen-
eral coloration and development of markings. Occasional speci-
mens may possess a weak prealar bristle and three posterodorsal
bristles on hind tibia.

Leucophora unilineata (Zetterstedt)

Anthomyza unilineata Zetterstedt, Ins. Lapp., 1838 p. 675.

Aricia unilineata Zetterstedt, Dipt. Scand., 1845 IV p. 1518.
Anthomyia unilineata Schiner, Faun. Austr., 1862 I p. 638.
Chortophila unilineata Meade, Entom. Month. Mag., 1882 XIX

p. 146 . . . Meade, Descr. List Brit. Anth., 1897 II p. 44.

Dec., 1940]

Huckett: Leucophora

349

Hylephila unilineata Strobl, Verb, zool.-bot. Ges. Wien, 1893
XLIII p. 265 . . . Stein, Arch. f. Naturgesch., 1920 (1918)
LXXXIV A heft 9 p. 84 . . . Collin, Trans. Ent. Soc. Lon-
don, 1921 (1920) Pt. Ill p. 311 . . . Segny, Faune de
France, 1923 No. VI p. 77 . . . Segny, Compt. rend. Congres
Societes savantes 1925, 1926 p. 475 . . . Johnson, Occ. Pap.
Boston Soc. Nat. Hist., 1925 No. VII p. 236 . . . Ringdahl,
Ark. f. ZooL, 1930, Bd. XXIA No. 20 p. 5 . . . Ringdahl,
K. Svensk. Vetenskaps. Skrift. Naturskydds., 1931 Nr. 18 p.
18 . . . Segny, Mem. Acad. Cienc. Exact. Fisico-Qnim.
Natnr. Zarag., 1934 III p. 51 . . . Segny, Gen. Insect., 1937
Fasc. 205 p. 139.

Hammomyia unilmeata Stein, Berl. Ent. Zeitschr., 1898 (1897)
XLII p. 230 . . . Aldrich, Misc. Coll. Smithsn. Inst., 1905
XLVI No. 1444 p. 554 .. . Stein, Kat. Palaark. Dipt., 1907
III p. 699 . . . Smith, Ann. Kept. N. J. State Mnsenm 1909,
1910 p. 791 . . . Stein, Arch. f. Natnrgesch., 1914 (1913)
LXXIX A heft 8 p. 32 . . . Cole and Lovett, Proc. Cal.
Acad. Sci., 1921 XI No. 15 p. 313 . . . Leonard, Mem. 101
N. Y. (Cornell) Agr. Exp. Station, 1928 (1926) p. 837 . . .
Strickland, Canad. Jonrn, Research, 1938 Sec. D XVI p. 209.
Anthomyia (Hammomyia) huccata Pandelle not Fallen, Rev. ent.

France, 1901 XX p. 301.

Records : —

Alberta, IJ, Wabamnn, April 21 1934, 2^?, IJ, Edmonton,
April 25 1937, April 28 1937, 1?, May 13 1937 (E. H.
Strickland) ; IJ, same locality, April 30 1937 (F. 0. Morri-
son).

Manitoba, IJ', Aweme, April 7 1935 (N. Criddle).

Michigan, IJ, Grand Rapids, May 15 1937.

New York, IJ, Sea Cliff, Long Island, April (N. Banks) [U. S.
N. M.] ; 25, Valley Stream, Long Island, April 27 1921; IJ',
Deer Park, Long Island, April 20 1935 ; 1^, Dix Hills, Long
Island, April 18 1935 (Blanton & Borders) ; Riverhead,
Long Island, April 28 1924, 1 J', 1?, May 1 1927 ; IJ, May 13
1923, 42, May 4 1926, 22, May 8 1926, 12, Sept. 12 1926 ;
IJ', Sonthold, Long Island, April 11 1925 ; 12, Ithaca, May
2 1900.

350

Journal New York Entomological Society [Vol. xlviii

Wisconsin, 4J, Madison, April 14 1936, 12J, April 29 1936,
45, May 3 1936, 2$, May 11 1936, 4J^, 2% Dane County, April
18 1937, 4c^, 4?, May 1 1937, 3^?, 2?, May 6 1937, 1^, April
22 1937, 1$, Devils Lake, May 7 1937 (F. M. Snyder).

The species unilineata may be distinguished from related forms
in that the mid tibia possesses a weak bristle near middle of
anteroventral surface. In both sexes the fore tibia has a robust
posterodorsal apical bristle and interfrontalia a pair of weak
cruciate setulse. Collin (1921) has recorded that Stein in his
monograph on the european Anthomyiidee appeared to have mis-
taken ohtusa for unilineata, and that Schnabl and Dziedzicki in
Die Anthomyiden have erroneously illustrated the copulatory
appendages of unistriata as belonging to this species. I have also
in a study of the Anthomyiinaae of New York mistakenly repre-
sented the figures of the male genitalia of ohtusa as belonging to
unilineata.

Leucophora ohtusa (Zetterstedt)

Anthomyza ohtusa Zetterstedt, Ins. Lapp., 1838 p. 682.

Aricia ohtusa Zetterstedt, Dipt. Scand., 1845 IV p. 1571.
Hylephila ohtusa Stein, Arch. f. Naturgesch., 1920 (1918)
LXXXIV A heft 9 p. 84 . . . Collin, Trans. Ent. Soc. Lon-
don, 1921 Pt. Ill p. 311 . . . Seguy, Faune de France, 1923
No. VI p. 76 . . . Seguy, Compt. rend. Congres Societes
savantes 1925, 1926 p. 475 . . . Johnson, Occ. Pap. Boston
Soc. Nat. Hist., 1925 No. VII p. 236 . . . Karl, Tierwelt
Deutschlands, 1928 XIII Pt. 3 p. 193 . . . Tiensuu, Acta
Soc. Faun. Flor. Fenn., 1935 LVIII No. 4 p. 13 . . . Seguy,
Gen. Insect., 1937 Fasc. 205 p. 138.

Hammomyia ohtusa Leonard, Mem. 101 N. Y. (Cornell) Agr.
Exp. Sta., 1926 (1924) p. 837 . . . Hallock and Parker, Circ.
103 N. J. Dept. Agr., 1926 p. 16.

Kecords : —

California, I5, Los Angeles Co., March (U. S. N. M.) ; IJ,
Artesia, Febr. 10 1935 (M. W. Stone) ; IJ', Santa Cruz,
April 1924 (Latta) ; 1$, Cazadero, April 12 1918 (J. C.
Bradley).

District of Columbia, 2$, Washington, March 26 (J. M.

Aldrich) [U. S. N. M.].

Dec., 1940]

Huckett: Leucophora

351

Idaho, 2?, Moscow, May 6 1912 (U. S. N. M.) ; IJ', Kendrick,
April 1912 (U. S. N. M.).

Illinois, 12, Algonquin (Coq.) [U. S. N. M.] ; 12, Dubois,
April 24 1914.

Indiana, 2J', Lafayette, no date, IJ', May 10 1917, 5J', 12,
April 22 1917, 1^, April 17 1916, 12, April 28 1916, 22, May
1 1915 (J. M. Aldrich), IJ', same locality, April 21 1915
[U. S. N. M.].

Kansas, 12, Ellsworth Co., May (R. H. Painter).

Michigan, IJ', Detroit, no date (Hubbard & Schwartz) [U. S.

N. M.].

Montana, 12, Gallatin Co., 4800 ft. alt., April 26 1902 (R.
Benton) .

New York, IJ', 52, Valley Stream, Long Island, April 27 1921 ;
12, Dix Hills, Long Island, May 19 1935, 12, Half Way
Hollow Hills, Long Island, May 18 1935 (Blanton &
Borders) ; IJ', Yaphank, Long Island, May 3 1924; 2J', 12,
Riverhead, Long Island, May 13 1923, 12, May 8 1926 ; 12,
Ithaca, no date, 12, April 16 1917 (R. C. Shannon) ; IJ',
Taughanic Falls, near Ithaca, April 21 1917 (R. C. Shan-
non).

Ohio, IJ', Marion, May 1 1915 (R. C. Smith).

Oregon, IJ', Kiger’s Island, April 8 1930 (J. Wilcox) ; 12,
Corvallis, March 27 1919 (A. L. Lovett), 12, April 8 1930
(J. Wilcox).

Pennsylvania, IJ', Harrisburg, April 1917, 12, April 1929
(W. R. Walton) ; 12, Allegheny, no date [U. S. N. M.].

Quebec, 1(2, Mount Royal, April 12 (J. Ouellet).

Texas, 1(2, College Station, March 1 1937 (H. J. Reinhard).

Utah, 1(2, Spanish Pork, no date (D. E. Hardy) ; 22, Logan
Canyon, April 24 1938 (Wm. P. Nye).

Virginia, 12, Great Falls, May 11 1917 (C. T. Greene).

Washington, 18(2, Pullman, May 3 1924, 2(2, 12, May 13
1923, 4(2, April 22 1923, 2(2, April 25 1924, 12, Jnne 4 1925
(A. L. Melander) ; 12, Rainier National Forest, Sawmill
Flat, May 26 1935 (J. Wilcox).

Wisconsin, 2(2, Madison, May 5 1936, 12, May 11 1936, 2(2,
April 29, 1936, 4(2, Dane County, May 6 1937, 1(2, May 10
1937, 1(2, Devils Lake, May .7 1937 (P. M. Snyder).

352

Journal New York Entomological Society [Vol. XLVIII

The species ohtusa has undoubtedly been confused with uni-
lineata and unistriata (Zetterstedt) in many local collections.
Structurally the species resembles unistriata {-paludis Joh.),
but ohtusa is usually larger and typically is clothed with more
abundant setulse. In the above series of specimens there are few
if any safe diagnostic characters that may serve usefully to sepa-
rate the species irom unistriata. The more satisfactory distinc-
tions have been presented in the keys to species.

Collin (1921) has concluded that Stein’s european records of
ohtusa do not refer to that species but to another for which he
has proposed the name personata. I have also concluded from an
examination of the specimen on which Stein® based his record of
ohtusa in North America that the specimen does not probably
belong to that species but is more likely conspecific with the form
named herein as fusca.

Leucophora unistriata (Zetterstedt)

Anthomyza unistriata Zetterstedt, Ins. Lapp., 1838 p. 677.

Aricia unistriata Zetterstedt, Dipt. Scand., 1845 IV p. 1512.
Hammomyia unistriata Stein, Entom. Nadir., XVIII 1892 p. 325
. . . Stein, Kat. Paliiark. Dipt. 1907 III p. 700 . . . Knowl-
ton, Trans. Utah Acad. Sci. Arts Letters, 1936 XIII p. 239
. . . Strickland, Canad. Jour. Res., 1938 D XVI p. 209.
Hammomyia paludis Johannsen, Ent. News, 1917 XXVIII p.
323 . . . Malloch, Canad. Ent., 1921 LIU p. 78 . . . Huckett,
Mem. 77 N. Y. (Cornell) Agr. Exp. Sta., 1924 (1923) p. 14.
Hylephila unistriata Collin, Trans. Ent. Soc. London, 1921
(1920) Pt. Ill p. 319 . . . Seguy, Faune de France, 1923
No. VI p. 77 . . . Seguy, Compt. rend. Congres Societes
savantes 1925, 1926 p. 475 . . . Karl, Tierwelt Deutschlands,
1928 XIII (3) p. 193 . . . Seguy, Gen. Insect., 1937 Fasc.
205 p. 139.

Records : —

Alberta, 2^?, Edmonton, May 10 1924, IJ', May 9 1937, 3J', 3$,
May 6 1924, 1$, May 23 1937 (E. H. Strickland) ; Ed-
monton, May 11 1938 (A. W. E. Eriksson) ; IJ, same local-
ity April 30 1937 (F. 0. Morrison) ; IJ', Wabamun, May 28
1936 (E. H. Strickland).

9 Stein, P. Nordamerikanische Anthomyiden, 2 Beitrag, Arch. f. Natur-
geseh., 1920 (1918) LXXXIV A heft 9 p. 84.

Dec., 1940]

Huckett: Leucophora

353

Indiana, IJ', Lafayette, April 27 1918 (J. M. Aldrich) [U. S.

N. M.].

Michigan, IJ, Albion, May 16 1936, IJ', E. Lansing, May 9
1937, 12, May 14 1937 (C. W. Sabrosky).

New York, 2^?, IJ, Valley Stream, Long Island, April 27 1921 ;
22, Babylon, Long Island, May 10 1935 (Blanton & Bor-
ders) ; 12, Ithaca, April 30 1915, IJ, April 25 1920, IJ',
April 29 1922; IJ', 6-mile Creek, Ithaca, April 29 1922
(L. S. West) ; 1(^, Cayuta Lake, May 8 1935; 2J, Danby,
May 14 1916 (E. G. Anderson).

Nova Scotia, IJ', Truro, June 24 1917, IJ, June 18 1917 ; IJ',
Kentville, June 27 1917.

Washington, 1(^, Pullman, April 21 1924 (A. L. Melander).

The species comes closest to oMusa^ but is usually smaller and
of paler hue. The m-cu cross vein in unistriata is quite fre-
quently more upright than in ohtusa. In some male specimens of
unistriata the prosternum may possess bristles. The species may
usually be distinguished from ol)tusa and other related species by
the characters given in the keys.

Collin (1921) is of the opinion that unistriata was included by
Stein in his later European records under inflata Rondani, and
that Schnabl and Dziedzicki have mistaken the species for uni-
lineata Zett. Johnson^® and Leonard^^ have listed paludis Jo-
hannsen as a synonym of ohtusa Zetterstedt. I have seen a female
paratype of paludis in the collections of the United States Na-
tional Museum that I regard as conspecific with ohtusa, but the
type specimens of paludis are, in my opinion, representative of
unistriata.

Leucophora fusca new species

Male, parafrontals and parafacials silvery pruinescent, cheeks reddish
tinged; antennae and palpi blackish, proboscis lightly pruinescent; thorax
and abdomen with dark reflections, the former subshining and with trace of
three brownish vittae, scutellum infuscated laterad; abdomen more densely
pruinescent, abdominal vitta brownish, paler on basal terga. Legs blackish,
narrowly reddish at knees. Wings faintly tinged, lower calyptral scale
whitish; halteres deep yellow with trace of reddish tinge.

10 Johnson, C. W. List of the diptera or two-winged flies. 15. Fauna of
New England. Occ. Pap. Boston Soc. Nat. Hist,, 1925 VII p. 236.

11 Leonard, M. D. A list of the insects of New York. Mem. 101 N. Y.
(Cornell) Agr. Exp. Sta., 1928 (1926) p. 837.

354

Journal New York Entomological Society [Vol. XLVIII

Erons at narrowest scarcely wider than breadth of third antennal segment,
frontal bristles continued to ocellar callosity, cruciate setulae present; buccal
region strongly developed, parafacials wider ventrad, lower half of occipital
region swollen, marginal bristles and vibrissse not robust; arista subnude,
proboscis slender; thorax with one or more pairs of presutural acrostical
bristles, prealar bristle longish, notopleural callosity with setulse, mesopleural
setulas invading the declivity dorsad of mesothoracic spiracle, scutellum
setulose laterad, ventral bristle of caudal pair of sternopleurals weakly de-
veloped; abdomen stout, truncated, scarcely as long as thorax, processes short
and blunt, sterna clothed with longish bristles, tergal marks slightly tapering
toward caudal margin.

Fore tibia with two or more posteroventral bristles, apical anterodorsal
robust, apical posterodorsal weak; mid tibia with a mid ventral bristle, one
anterodorsal, one posterodorsal and two or more bristles on posterior surface,
apical anterodorsal robust, apical posterodorsal weak; hind femur with a
complete series of anteroventral bristles with a series of finer bristles on
proximal half of posteroventral surface ; hind tibia with three to five antero-
ventral bristles, four or five anterodorsal, three posterodorsal, four or five
weaker posteroventral bristles. Wings with m,-cu cross vein sinuate and
markedly oblique; cell slightly narrower distad. Length 9 mm.

Female similar to male, paler, width of frons as in male, marginal bristles
of fifth abdominal tergum stoutly developed, ovipositor with a pair of short
recurrent spines and numerous spinules on anal palpi. Fore tibia with a mid
posterior and a weak anterodorsal bristle; mid tibia with a posteroventral
bristle adjacent mid ventral bristle; hind femur with bristles on proximal
half of antero- and posteroventral surfaces weaker, hind tibia with a series
of three or more weak bristles on posteroventral surface.

Type, c? Mt. Moscoav, Idaho, June 5 1930 ( J. M. Aldrich) [U. S.

N. M.].

Allotype, 5 Lafayette, Indiana, May 14 1918 [U. S. N. M.].

Paratypes, J' Edmonton, Alberta, May 9 1937, 2^ Clymont,
Alberta, May 24 1937 (E. H. Strickland) [C. N. C.].

Records :

Alberta, Edmonton, May 13 1937 (F. 0. Morrison).

Idaho, 7J', Mt. Moscow, June 5 1930 (J. M. Aldrich) [U. S.

N. M.].

Indiana, IJ', Lafayette, April 26 (U. S. N. M.), 22, same

locality. May 14 1918 (U. S. N. M.).

Oregon, IJ', Corvallis, June 2 1929 (John Wieting).

Washington, 4J', Almota, May 20 1923 (A. L. Melander) ; 12,
Sprague, June 13 1920, 12, Ewan, June 13 1920 (R. C.
Shannon).

Dec., 1940]

Huckett: Leucophora

355

The species fusca resembles the European form personata Col-
lin, from which it may be distinguished by the scutellar infusca-
tion, weakly developed ventral bristle of caudal pair of sterno-
pleurals, and by the presence of weak cruciate setulse. This is
the species recorded by me in error as personata in Strickland’s
list^^ of the dipterous flies occurring in Alberta. Another closely
allied species is described as follows.

Leucophora annexa new species

Male and female resembling fusca, differing in that the mesonotiim has
only a dorsocentral vitta, scutellum entirely pale grayish. In the male the
lower sternopleural bristle of caudal pair is well developed and the jjrocesses
are longer. Fore tibia with one or two posteroventral bristles, apical antero-
dorsal weak, mid tibia with a weak mid ventral bristle, hind femur with
bristles on proximal half of posteroventral surface weaker and shorter, not
as long as breadth of hind femur where situated, hind tibia with a weaker
series of posteroventral bristles. In the female the lower sternopleural bristle
of caudal pair is weakly developed, fore tibia lacks a mid posterior bristle
as in fusca and has a weakly developed apical anterodorsal, apical postero-
dorsal absent, mid tibia with a stout mid ventral bristle, one anterodorsal, one
posterodorsal and three posterior bristles, apical anterodorsal robust, hind
femur with no series of bristles on posteroventral surface, hind tibia with
one or two inconspicuous longer setulse on posteroventral surface.

Type, c?, Lind, Washington, May 30 1919 (F. W. Carlson) ;
Allotype, Lewiston Hill, Idaho, May 31 1934 (A. L. Melander).
In the collection of Dr. A. L. Melander.

Superficially annexa resembles a paler form of fusca, and in
this respect conforms closely to the appearance of personata
Collin. I have regarded the species as distinct from personata
owing to the presence of cruciate setulas, weaker series or lack of
bristles on posteroventral surface of hind femur, and further
owing to the greater length of abdominal processes in male.
Since the mid ventral bristle on mid tibia of male is notably weak
in the specimens examined I have included the species under both
categories in the first couplet of the male key, lest additional
specimens may show that this character is not dependable for
comparative purposes.

12 Strickland, E. H. An annotated list of the diptera (flies) of Alberta.
Canad. Journ. Eesearch, Sec. D XVI 1938 p. 209.

356 Journal New York Entomological Society [Vol. XLVIII

Genus Proboscimyia Bigot

Proboscimyia Bigot, Bull. Soc. Ent. France, 1883 No. 4 p. 30 . . .
Huckett, Mem. 77 N. Y. (Cornell) Agr. Exp. Station, 1924
(1923) p. 50 . . . Curran, Fam. Gen. N. A. Dipt., 1937 p.
391.

Proboscidomyia Bigot, Ann. Soc. Ent. France, 1884 ser. 6 IV p.
266 .. . Aldrich, Misc. Coll. Smitlisn. Inst., 1905 XLVI No.
1444 p. 548 . . . Stein, Arch. f. Naturgesch., 1919 (1917)
LXXXIII A heft 1 p. 152 . . . Seguy, Gen. Insect., 1937
Fasc. 205 p. 128.

Dolichoglossa Stein, Berl. Ent. Zeitschr., 1898 (1897) XLII p.
230 .. . Aldrich, Misc. Coll. Smitlisn. Inst., 1905 XLVI No.
1444 p. 554 . . . Williston, Man. N. A. Dipt., 1908 3rd ed.,
p. 335.

Genotype Prohoscimyia siphonina Bigot.

The genus is not to be readily distinguished from Leucophora
as denoted by the adult flies. Nothing, as far as I am aware, is
known concerning the habits of the species. The character that
has served to distinguish the group has been the abnormally
slender proboscis in the genotype. It is to me doubtful whether
this character is of generic value in view of the fact that another
species belonging to the genus occurs in North America whose
proboscis is not abnormally slender. The lower calyptra in both
species is protruded distinctly beyond margin of upper scale.

Key to Species

Distal section of proboscis longer than fore tarsus, oral vibrissse shorter than

palpi siphonina Bigot

Distal section of proboscis not as long as fore tarsus, oral vibrissae as long as
or longer than palpi b7'evis n. sp.

Prohoscimyia siphonina Bigot

Prohoscimyia siphonina Bigot, Bull. Soc. Ent. France, 1883 No.
4 p. 30.

Proboscidomyia siphonina Bigot, Ann. Soc. Ent. France, 1885
ser. 6 V p. 267 . . . Aldrich, Misc. Coll. Smithsn. Inst., 1905
XLVI No. 1444 p. 548 .. . Stein, Arch. f. Naturgesch., 1919
(1917) LXXXIII A heft 1 p. 152 . . . Seguy, Gen. Insect.,
1937 Fasc. 205 p. 129.

Dec., 1940]

Huckett: Leucophora

357

Dolichoglossa americana Stein, Berl. Ent. Zeitschr., 1898 (1897)

XLII p. 230, 286 . . . Aldrich, Misc. Coll. Smithsn. Inst.,

1905 XLVI No. 1444 p. 554.

Records : —

Illinois, IJ', 15, Carlinville (Robertson) [IT. S. N. M.].

New Mexico, IJ', Koehler, Ang. 12 1914, IJ', no date (W. R.
Walton), 25, Socorro, 1916 (Williston) [U. S. N. M.].

South Dakota, IJ, Elmira (J. M. Aldrich), cotype of Dolicho-
glossa americana Stein, Lectotype [U. S. N. M] ; IJ, Onida,
July 12 1932 (G. B. Spawn).

Texas, IJ', Comanche Co., June 8 1928 (V. A. Lityle) ; 1$,
Taylor, May 27 1929, IJ', IJ, Hidalgo Co., June 6 1930
(J. C. Gaines) ; IJ, Bexar Co., June 8 1928, IJ', IJ, same
locality. May 29 1932 (H. B. Parks) ; IJ', College Station,
May 25 1930, IJ, same locality. May 29 1920 (H. J. Rein-
hard) [Texas A. & M.] ; 2J, no data, IJ', Belfrage [U. S.
N. M.].

All previous records have been credited to this species. I am
however doubtful whether those of eastern regions may be unre-
servedly accepted as correct in view of the evidence provided by
the material before me. The species siphonina is represented
exclusively by specimens collected in western parts of North
America, whilst in New York, Nova Scotia and the District of
Columbia another species has made its appearance.

In siphonina the proboscis is much longer, oral vibrissas weaker,
palpi slightly longer and notopleural callosity typically more
setulose than in the following species. In both forms the male
lacks and the female possesses a mid ventral bristle on mid tibia.
In the male of siphonina the processes have a chitinous callosity
at middle of inner margin (fig. 25).

Proboscimyia brevis new species

Similar to siphonina Bigot, but having the parafrontal bristles and oral
vibrissae more robust; proboscis shorter, distal section scarcely as long as
fore tarsus; palpi proportionately shorter; mesonotum with fewer accessory’
setulae on notopleural callosity; notably so at basal region of posterior noto-
pleural bristle ; abdominal processes devoid of a .chitinous callosity at middle
of inner margin; bristling of legs in male as follows: — Fore tibia with one
or two posteroventral bristles and a weak mid anterodorsal bristle ; mid tibia
with one anterodorsal, one posterodorsal and two posterior bristles; hind

358

Journal New York Entomological Society [Vol. XLVIII

femur with a complete series of anteroventral bristles and a weaker series of
bristles on proximal half of posteroventral surface; hind tibia with two
anteroventral, three anterodorsal and three posterodorsal bristles.

Female similar to male, with a series of fine recurrent spines on anal palpi
of ovipositor ; sternopleural bristles arranged 2 : 2, the lower bristles weakly
developed ; mid tibia with a strong anterodorsal and mid ventral bristle, hind
femur with bristles on anteroventral surface much weaker proximad and those
on posteroventral surface absent except for one at base, hind tibia with one
or more weak bristles on posteroventral surface. Length, 6-8.5 mm.

Type and allotype, J' Babylon, Long Island, New York, Sept.
18 1937 (F. S. Blanton) [U. S. N. M.].

Records : —

District of Columbia, IJ, Oct. (Coqnillett) [U. S. N. M.].

New York, IJ', IJ, Babylon, Long Island, Sept. 14 1933, 1$,
Aug. 15 1933, Ic?, 1?, Sept. 12 1933, IJ', Sept. 15 1937, 1?,
Sept. 23 1935, 1?, Sept. 19 1935, 5J, 1?, Sept. 23 1933, 6^?,
4$, Sept. 18 1937 (F. S. Blanton) ; IJ', IJ, Babylon, Long
Island, Sept. 14 1935, 2^, 1$, Sept. 19 1934, 4c?, Sept. 12
1935, Ic?, Sept. 17 1935, 2c?, 1$, Sept. 10 1935, 1?, Sept. 7
1935 (Blanton & Borders) ; IJ, Riverhead, Long Island,
Sept. 21 1926.

Nova Scotia, IJ, Truro, Sept. 15 1913 (R. Matheson).

Plate XVII

Dorsal or caudal aspect of male copulatory appendages

Figure

1.

Figure

2.

Figure

3.

Figure

4.

Figure

5.

Figure

6.

LeucopUora alMseta (von Eoser).
Leucopliora maculata (Stein).
Leucophora marylandica (Malloch).
Leucopliora sociata (Meigen).
Leucophora fusca new species.
Leucophora unilineata (Zetterstedt).

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

(Plate XVII)

LEUCOPHOEA

360

Journal New York Entomological Society [Vol. XLVIII

Figure

Figure

Figure

Figure

Figure

Figure

Plate XVIII

Lateral aspect of male copulatory appendages

7. Leucophora marylandica (Malloch).

8. Leucophora fusca new species.

9. Leucophora sociata (Meigen).

10. Leucophora maculata (Stein).

11. Leucophora unilineata (Zetterstedt).

12. Leucophora alhiseta (von Eoser).

(Plate XVIII)

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

'V

LEUCOPHOEA

362

Journal New York Entomological Society

[VoL. XLVIII

Figure 13.
Figure 14.
Figure 15.
Figure 16.
Figure 17.
Figure 18.

Figure 19.
Figure 20.

Plate XIX

Ventral aspect of fifth abdominal sternum in male
Leucophora alhiseta (von Eoser).

Leucophora marylandica (Malloch).

Leucophora sociata (Meigen).

Leucophora unilineata (Zetterstedt) .

Leucophora maculata (Stein).

Leucophora fusca new species.

Ovipositor of Leucophora maculata (Stein)
Ventral aspect.

Dorsal aspect.

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

(Plate XIX)

LEUCOPHORA

364

Journal New York Entomological Society [Vol. XLVIII

Plate XX

Male copulatory appendages of Proboscimyia
Figures 21, 23, 25. Proltoscimyia siphonina Bigot.

Figures 22, 24, 26. Froljoscimyia trevis new species.

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

(Plate XX)

LEUCOPHORA

A RECLASSIFICATION OF THE TRIBE OBRIINI
OF LECONTE (COLEOPTERA, CERAMBYCIDiE)

By E. Gorton Linsley
University of California

The genera treated below were grouped together by LeConte
(1873) under the tribal name Obriini. The same arrangement
was maintained by LeConte and Horn (1883) and Leng (1886).
Recent workers have usually divided the genera into two widely
separated tribes and in some cases this has resulted in the plac-
ing of very closely related genera in different groups. In the
writer ^s opinion the complex relationships of these genera can
best be expressed by the recognition of several tribes as follows:

Key to Tribes

1. Elytra entire, neither abbreviated nor attenuated 2

Elytra either greatly abbreviated or attenuated 4

2. Anterior coxal cavities closed posteriorly; mesosternnm with intercoxal

process parallel-sided; pronotum with sides angulate or tuberculate;
second segment of antennae much less than one-half as long as third;
abdomen of female with first sternite as long as remaining segments
together, sternites 2-5 greatly modified; ultimate segment of palpi

cylindrical 3

Anterior coxal cavities open posteriorly; mesosternnm with intercoxal
process triangular; pronotum with sides rounded; second segment of
antennae one-half as long as third; abdomen of female with first seg-
ment shorter than remaining together, sternites 2-5 unmodified; ulti-
mate segment of palpi dilated Eumichthini

3. Anterior coxae conical, exserted, nearly contiguous, cavities not angulated

externally; intermediate coxal cavities not open to epimera; episterna
of metathorax divided by a longitudinal furrow; pronotum without

dorsal tubercles Obriini

Anterior coxae globular, separated, cavities angulated externally; inter-
mediate coxal cavities open to epimera; episterna of metathorax with-
out a longitudinal furrow; pronotum with dorsal tubercles.

Hyboderini

4. Anterior coxal cavities open posteriorly; elytra less than one-half as long

as abdomen, apices rounded or truncate, never attenuated or subulate;
intercoxal process of mesosternum narrow, triangular, truncate behind ;
antennae longer than the body in the male; posterior wings without a

post-cubital vein Molorchini

Anterior coxal cavities closed posteriorly; elytra more than one-half as

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Journal New York Entomological Society [Vol. XLVIII

long as abdomen, apices attenuated, subulate; intercoxal process of
mesosternum broad, flat, emarginate behind; antennge shorter than the
body in both sexes; posterior wings with a single postcubital vein.

Stenopterini

Tribe Eumichthini Linsley, new tribe

Head short; eyes finely granulated, deeply emarginate; antennae with
second segment elongate, about one-half as long as third segment; palpi
short, last segment triangularly dilated; ligula membranous. Pronotmn
rounded at sides, without lateral tubercles, dorsal callosities, if present,
feeble; anterior coxae globular, cavities angulate externally, open posteriorly;
intermediate coxal cavities open to epimera, intercoxal process triangular;
posterior wings with a single post-cubital vein; legs moderate in length,
femora clavate, tibial spurs short. Abdomen of female with first sternite
shorter than the following segments together, sternites two to five unmodified.

This tribe is necessary for two genera which, in the absence
of females, were placed in the Obriini by LeConte and Horn.
In the Aurivillins (1912) and Leng (1920) catalogues one of
the genera, Eumichthus, is listed in the Obriini, the other, Poe-
cilohrium, in the Molorchini. They appear to me to be equally
out of place in either tribe. From the Obriini they differ in
the shape and structure of the anterior coxas and their cavities,
the form of the intercoxal process of the mesosternum, the
elongated second segment of the antennge, the dilated palpi, and
the entire, unmodified abdomen of the female. From the Molor-
chini they may be distinguished by the entire elytra, less promi-
nent and more globular anterior coxae, elongate second segment
of the antennae, laterally rounded pronotum, distinct stridnlatory
area of the metanotnm, and dilated palpi.

Two genera occur in North America. They may be separated
as follows :

Third segment of antennsB shorter than fourth; last segment of palpi nar-
rowly triangular; first two segments of anterior tarsi tumid in male;

integument non-metallic Eumichthus

Third segment of antennae longer than fourth; last segment of palpi broadly
triangular; first two segments of anterior tarsi slender in both sexes;
integument metallic : Poecilobrium

Genus Eumichthus LeConte

Eumichthus LeConte, 1873, Smithson. Misc. Coll., XI, 264: 190,
265 : 305 ; LeConte and Horn, 1883, Smithson. Misc. Coll.,
XXVI, 507 : 291 ; Leng, 1886, Entom. Amer., 2 : 27.

Dec., 1940]

Linsley: Obriini

369

Eumichtus, Aurivilliiis, 1912, Coleopt. Catal., 31 : 138.

Integument dull. Head with front declivous ; antennge
slightly longer than the body in the male, third segment not
longer than fourth segment ; antennal tubercles scarcely elevated,
intervening area convex; clypeus with a small impression on
each side at base; palpi with last segment narrowly triangular.
Pronotum without any dorsal callosities ; elytra maculate ; legs
moderately short, femora very strongly clubbed, anterior tarsi
of male with first two segments swollen, of female slender.

Genotype: Eumichthus oedipus LeConte (by single reference).

The type and only known species of this genus occurs on the
Pacific coast of North America. It bears a strong superficial
resemblance to the unrelated Phymatodes decussatus (LeConte).

Genus Poecilobrium Horn

Pcecilohrium Horn, 1883, Smithson. Misc. Coll. XXVI, 507 : 291 ;

Horn, 1883, Trans. Am. Ent. Soc., 10 : xi ; Leng, 1886, Entom.

Amer., 2: 27.

Callimus, LeConte, 1873, Smithson. Misc. Coll., XI, 265 : 305.

Integument shining, metallic. Head with front subvertical ;
antennas shorter than the body in both sexes, third segment
longer than fourth segment; antennal tubercles slightly ele-
vated, intervening area feebly concave; clypeus with a large
impression on each side at base ; palpi with last segment broadly
triangular. Pronotum with feeble dorsal callosities; elytra con-
colorous; legs short, femora moderately strongly clubbed; an-
terior tarsi with first two segments slender in both sexes.

Genotype: Callimus chalyheus LeConte (by single reference).

This monotypic genus is based on a small, brilliant, metallic-
blue species which occurs along the Pacific coast of North
America.

Tribe Obriini Thomson

C. G. Thomson, 1859, Skand. Coleopt., 1 : 150, Ohriina.

C. G. Thomson, 1866, Skand. Coleopt., 8: 40, Ohriina.

Lacordaire, 1869, Genera Coleopt., 8 : 360, Ohrionides.

Pascoe, 1869, Trans. Ent. Soc. Lond., 1869 : 550.

LeConte, 1873, Smithson. Misc. Coll., XI, 265 : 305, Ohria.

370

Journal New York Entomological Society [Vol. XLVIII

LeConte and Horn, 1883, Smithson. Misc. Coll., XXVI, 507:
290, Ohria.

Long, 1886, Entom. Amer., 2 : 27.

Gahan, 1906, Fauna Brit. India, Coleopt., 1 : 164.

Blatchley, 1910, Coleopt. Indiana, p. 1027.

Plavilstshikov, 1932, Best.-Tab. eur. Coleopt., 102: 62, Ohriina.

Gressitt, 1935, Ins. Matsnmnrana, 9 : 146.

Gressitt, 1939, Ling. Sci. Jour., 18: 10.

Head moderately short ; eyes large, coarsely granulated,
deeply emarginate; antennae with second segment short, very
much less than one-half as long as third segment ; palpi unequal,
last segment cylindrical ; ligula membranous. Pronotum with
sides protuberant or tuberculate ; anterior coxae conical, exserted,
nearly contiguous, cavities rounded externally, closed posteri-
orly ; intermediate coxal cavities not open to epimera, intercoxal
process parallel-sided ; metepisterna with a longitudinal furrow ;
wings with two post-cubital veins ; legs moderately long, femora
clavate and pedunculate. Abdomen with first sternite very long
in both sexes; first sternite of female as long as following seg-
ments together, second sternite deeply emarginate, densely
clothed with long, recurved hairs, third sternite very short,
fourth and fifth sternites longer.

The outstanding characters of this tribe are the conical, ex-
serted anterior coxee with their cavities rounded externally and
closed behind, the closed intermediate coxal cavities, and the
coarsely granulate eyes. This latter character is slightly vari-
able in Ohrium and in the Philippine and Formosan genus
Pseudiphra, the eyes are said to be finely facetted.

Genus Obrium Dejean

Ohrium Dejean, 1821, Catal. Coleopt., ed. I, 110; Curtis, 1825,
British Ent., 2 : 91 ; Latreille, 1829, in ; Cuvier, Regne Animal,
5 : 119 ; Serville, 1834, Ann. Soc. Ent. France, 3 : 93 ; LeConte,
1850, Jour. Acad. Nat. Sci. Phila., (2), 2: 21; C. G. Thomson,
1859, Skand. Coleopt., 1 : 151 ; Fairmaire*, 1864, Genera
Coleopt. Eur., 4: 179; J. Thomson, 1864, Systema Ceramb.,
440; C. G. Thomson, 1886, Skand. Coleopt. 8; 41; LeConte,
1873, Smithson. Misc. Coll., XI, 265 : 306 ; LeConte and Horn,

Dec., 1940]

Linsley: Obriini

371

1883, Smithson. Misc. Coll., XXII, 507 : 291 ; Long, 1886,
Ent. Amer., 2 : 27 ; Gahan, 1906, Fauna Brit. India, Coleopt.,
1 : 165 ; Blatchley, 1910, Coleopt. Indiana, p. 1027 ; Reitter,
1912, Fauna Germ. Kafer, 4: 29; Planet, 1924, Encycl. Ent.,
A, 2: 137; Portevin, 1927, Encycl. Ent. 2 (snppl.) : 22;
Scheerpeltz, 1930, Tierw. Mittelenr. Kafer, 5: (2) : 209; Ma-
tushita, 1933, Jour. Fac. Agr., Hokkaido Univ., 34: 305;
Gressitt, 1935, Ins. Matsnmurana, 9 : 145.

Head with front sloping anteriorly; antennae usually at least
one-third longer than the body in male, barely longer than body
in female, segments three and four subequal in length, basal
segments without long, apical hairs ; clypeus with a deep, arcuate
impression above; palpi short. Pronotum longer than broad,
constricted near base, narrowly so before apex; elytra parallel-
sided or slightly widened posteriorly, apex broadly rounded;
legs elongate, first segment of posterior tarsi as long as follow-
ing two together.

Genotype: Ceramhyx cantharinum Linnaeus (Curtis desig-
nation, 1825).

This genus reaches its greatest development in America but
is well represented in the Old World. Phyton Newman cannot
be separated on the basis of the characters used in the literature,
but it is possible that a careful study, in the light of the neo-
tropical fauna, may reveal other characters of generic or sub-
generic importance.

Tribe Hyboderini Linsley, new tribe

Head moderately short; eyes finely granulated; antennse with second seg-
ment short, much less than one-half as long as third segment ; palpi unequal,
last segment cylindrical; ligula membranous. Pronotum with sides and disk
tuberculate; anterior coxae globular, separated, cavities angulated externally,
open behind; intermediate coxal cavities open to epimera, intercoxal process
parallel-sided ; metepisterna without a longitudinal furrow ; wings with a
single post-cubital vein (two in Megobrium) ; legs moderately long, clavate.
Abdomen with first sternite elongate in both sexes ; first sternite of female as
long as following segments together, second sternite deeply emargiuate,
densely hairy, third sternite very short, fourth and fifth sternites longer.

This tribe is related to the Obriini with which it agrees in
the structure of the female abdomen. It differs markedly, how-
ever, in the finely granulate eyes, globular anterior coxae with

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Journal New York Entomological Society [Vol. XLVIII

their cavities widely angulated externally, open intermediate
coxal cavities, etc. Recent workers have placed the genera in
the Molorchini, a group from which they differ in the entire
elytra, the modified abdomen of the female, the parallel-sided
mesosternal process, and the presence of at least one post-cubital
vein in the wing. The North American genera differ as follows :

1. Pubescence sparse, depressed hairs few, flying hairs numerous ; punctation

coarse 2

- Pubescence short, dense, appressed, flying hairs absent; punctation fine.

Hyhodera

2. Intermediate coxae separated by at least their own diameters ; metepisterna

broad, parallel-sided ; elytral punctures coarse, distinct throughout,
elytral apices dehiscent, inner angle acute; posterior tibiae arcuate.

Lampropterus

- Intermediate coxae separated by much less than their own diameters; met-

episterna attenuated apically; elytral punctures becoming obsolete over
apical one-third, apices separately rounded; posterior tibiae straight... 3

3. Anterior coxae distinctly sej)arated; intercoxal process of mesosternum

less than twice as wide as that of prosternum; antennae of male dis-
tinctly shorter than body Pseudopilema

- Anterior coxae narrowly separated; intercoxal process of mesosternum

several times as wide as that of prosternum; antennae of male dis-
tinctly longer than body Megobrium

Genus Hybodera LeConte

Hyl)odera LeConte, 1873, Smithson. Misc. Coll., XI, 264; 191,
265: 306; LeConte and Horn, 1883, Smithson. Misc. Coll.,
XXVI, 507 : 291 ; Leng, 1886, Entom. Amer., 2 : 27.
Pubescence short, dense, appressed, without an intermixture
of flying hairs; punctation fine. Head with front short, de-
clivous; vertex canaliculate; basal suture of clypeus deeply im-
pressed; eyes deeply emarginate; antennae longer than the body
in the male, shorter than body in female; palpi short, last seg-
ment slightly oval. Pronotum a little longer than broad,
strongl}^ constricted anteriorly, less so posteriorly, sides obtusely
angulate, disk quadrituberculate ; prosternum distinctly separat-
ing coxae; intercoxal process of mesosternum broad, truncate;
metepisterna attenuated apically; elytra somewhat flattened,
parallel-sided, apex rounded ; legs with femora strongly clavate,
posterior tibiae straight; first segment of posterior tarsi longer
than following two together.

Dec., 1940]

Linsley: Obriini

373

Genotype: Hyhodera tuherculata LeConte (by single ref-
erence).

Only two species are known in this remarkable genus. Both
are confined to the Pacific coast of North America.

Genus Lampropterus Mulsant

Lampropterus Mulsant, 1863, Coleopt. France, Longie., Ed. 2,

214.

Callimus Mulsant, 1846, Coleopt. France, 4, Append; Thomson,
1860, Class Ceramb., p. 160 ; Thomson, 1864, Systema
Ceramb., p. 412; Lacordaire, 1869, Genera Coleopt., 8: 489;
LeConte and Horn, 1883, Smithson. Misc. Coll., XXVI, 507 :
291 ; Leng. 1886, Entom. Amer., 2 : 27, 29 ; Casey, 1912, Mem.
Coleopt., 3: 310; Keitter, 1912, Faun. Germ., Kafer, 4: 28,
Planet, 1924, Encycl. Ent., A, 2 : 133 ; Portevin, 1927, Encycl.
Ent., 2 (suppl.) : 22; Scheerpeltz, 1930, Tierw. Mitteleur.,
Kafer, 5 (2) : 209.

Pilema LeConte, 1873, Smithson. Misc. Coll. XI, 264; 191, 265:
306.

Callimellum Strand, 1928, Ent. Nachr. BL, 2 : 2, Plavilstshikov,
1932, Best.-Tab. eur. Coleopt, 102 : 81, 100.

Pubescence sparse, flying hairs numerous, depressed hairs few ;
punctation coarse. Head short, front declivous, vertex canalic-
ulate ; clypeus deeply impressed at base ; eyes deeply emargi-
nate ; antennas shorter than the body in both sexes ; palpi slender,
last segment cylindrical. Pronotum about as wide as long, sides
obtusely angulate, disk tuberculate; prosternum very narrowly
separating coxae; intercoxal process of mesosternum at least as
broad as coxae, truncate ; metepisterna broad, not greatly attenu-
ated apically; elytral punctation coarse, distinct throughout,
apices dehiscent, inner angle acute ; legs slender, femora mod-
erately clavate ; posterior tibiae curved, first segment of posterior
tarsi about as long as following two together.

Genotype : Necydalis femorata Germar.

Lampropterus occurs throughout the Palearctic Region but in
North America is represented by two species on the Pacific coast.

Genus Pseudopilema Linsley, new genus
Pubescence sparse, consisting mostly of flying hairs; punctation coarse.
Head short; front declivous, vertex canaliculate; last segment slightly oval.

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Journal New York Entomological Society [Vol. XLVIII

Proiiotum about as wide as long, constricted at base and apex, sides obtusely
angulate, disk tuberculate ; anterior coxae distinctly separated by pro-
sternum; intercoxal process of mesosternum less than twice as wide as that
of prosternum, much narrower than width of coxae; metepisterna attenuated
apically; elytra coarsely punctured at base, the punctures becoming obsolete
beyond middle, apices separately rounded; legs slender, femora moderately
clavate, posterior tibiae straight.

Genotype: Callimus Jioppingi Van Dyke.

This new genus is proposed for a species which differs from
Lampropterus in having the anterior and intermediate coxal cav-
ities narrowly separated, the metepisterna attenuated posteri-
orly, the elytral punctures becoming obsolete over the apical
one-third, and the posterior tibiae straight.

Genus Megobrinm Le Conte

Megohrium LeConte, 1873, Smithson. Misc. Coll., XI, 264: 192,
265 : 306 ; LeConte and Horn, 1883, Smithson. Misc. Coll.,
XXVI, 507 : 291 ; Leng, 1886, Entom. Amer., 2 : 27.

Pubescence sparse, flying hairs numerous, depressed hairs few ;
pnnctation coarse. Head with front short, sub vertical; vertex
distinctly concave between antennal bases ; eyes large, moderately
emarginate; antennae longer than the body in the male; palpi
short, last segment slightly oval, truncate at apex. Pronotum
longer than wide, broadly, equally constricted at base and apex,
lateral tubercles large, dorsal tubercles small but distinct;
prosternum narrowly separating coxae ; intercoxal process of
mesosternum several times as wide as that of prosternum ;
metepisterna attenuated apically; elytral punctation becoming
obsolete beyond middle, apices separately rounded, legs slender ;
femora not strongly clavate, posterior tibiae straight, first seg-
ment of posterior tarsi a little longer than following two
together.

Genotype : Megohrium edwardsi LeConte (by single reference).

In this genus there are two post-cubital veins in the wing as
in the true Obriini. The remaining characters, however, are
those of the Hyboderini. A single species is known, confined
to California.

Tribe Molorchini Thomson

C. G. Thomson, 1866, Skand. Coleopt., 8 : 43, Molorchina.

Lacordaire, 1869, Genera Coleopt., 8: 482, Molorchides.

Dec., 1940]

Linsley: Obriini

375

Gahan, 1906, Fauna Brit. India, Coleopt., 1 : 169.

Plavilstshikov, 1932, Best.-Tab. eur. Coleopt., 102 : 79, Molor-
china.

MatusMta, 1933, Jour. Fac. Agr., Hokkaido Univ., 34: 221.

Gressitt, 1939, Ling. Sci. Jour., 18 : 10.

Head short; eyes finely granulated; antennae with second seg-
ment small, much less than one-half as long as third segment;
palpi short, subequal, last segment not dilated; ligula mem-
branous. Pronotum with sides rounded, scarcely tuberculate;
anterior coxae globose, cavities angulate externally, open pos-
teriorly; intermediate coxal cavities open to epimera, intercoxal
process narrow, triangular, truncate behind; mesonotum with-
out a well-defined stridulatory area; elytra short, less than one-
half as long as abdomen, apices rounded or truncate; wings
without post-cubital veins ; legs moderately long, femora clavate.
Abdomen with sternites unequal, gradually diminishing in
length, not abnormally modified in female.

The tribe Molorchini differs from the Stenopterini, with which
it has been combined by recent workers, by having abbreviated
(not subulate) elytra which cover less than half of the abdomen,
posteriorly open anterior coxal cavities, no mesonotal stridula-
tory area, a narrow intercoxal process on the mesosternum, and
no post-cubital veins in the wing.

Genus Molorchus Fabricius

Molorchus Fabricius, 1792, Ent. Syst., 1: 356; Mulsant, 1839,
Coleopt. France, Longic., p. 107 ; J. Thomson, 1864, Systema
Ceramb., p. 411 ; C. G. Thomson, 1866, Skand. Coleopt., 8 :
43; Lacordaire, 1869, Genera Coleopt., 8: 486; LeConte, 1873,
Smithson. Misc. Coll., XI, 265 : 307 ; LeConte and Horn,
1883, Smithson. Misc. Coll., XXVI, 507: 292; Leng, 1886,
Entom. Amer., 2 : 27, 30 ; Gahan, 1906, Fauna Brit. India,
Coleopt., 1 : 169 ; Blatchley, 1910, Coleopt. Indiana, p. 1028 ;
Plavilstshikov, 1932, Best.-Tab. eur. Coleopt., 102 : 81, 93.
Heliomanes Newman, 1840, Entom., 1 : 17 ; LeConte, 1850, Jour.

Acad. Nat. Sci. Phila., (2) 2 : 21.

Glaphyra Newman, 1840, Entom., 1 : 19 ; LeConte, 1850, Jour.

Acad. Nat. Sci. Phila., (2) 2: 38.

Laphyra Newman, 1842, Entom., 1 : 418.

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Journal New York Entomological Society [Vol. XLVIII

Head with front short, vertical; vertex slightly concave be-
tween antennal bases, antennae much longer than body in male,
shorter than body in female, eleventh segment appendicnlate
or divided in male, simple in female; eyes deeply emarginate.
Pronotum at least as long as broad, constricted at base, feebly
so at apex, disk roughened or calloused; prosternum with inter-
coxal process very narrow; elytral apices dehiscent; legs elon-
gate, posterior pair attaining apex of abdomen in male; tarsi
slender, first segment of posterior pair as long as following two
together.

Genotype: Necydalis minor Linnaeus (Thomson designation,
1864).

Molorchus is dominant in the Holarctic region but is also
represented in the Indomalaysian and Australian faunas. Four
species occur in North America, one ranging from coast to coast,
two confined to the Pacific slope, and one described from the
West Indies.

Tribe Stenopterini LeConte

LeConte, 1863, Smithson. Misc. Coll., XI, '265 : 306, Stenopteri.

LeConte and Horn, 1883, Smithson. Misc. Coll., XXVI, 507 :
292, Stenopteri.

Blatchley, 1910, Coleopt. Indiana, p. 1028, Stenoptini.

Reitter, 1912, Faun. Germ., Kafer, 4: 28, Stenopterina.

Scheerpeltz, 1930, Tierw. Mitteleur., Kafer, 5 (2) : 209.

Head porrect; front large, oblique; antennse with second seg-
ment small, less than one-half as long as third segment; palpi
short, sub equal, last segment not dilated; ligula membranous.
Pronotum tuberculate at sides; anterior coxge globose, cavities
angulate externally, closed posteriorly; intermediate coxal cav-
ities open to epimera, intercoxal process broad, flat, emarginate
posteriorly; mesonotum with a large stridulatory area; elytra
elongate, subulate, covering more than half of the abdomen;
wings with a single post-cubital vein; legs long, femora clavate.
Abdomen with sternites unequal, gradually diminishing in
length, not abnormally modified in female.

A single genus, Callimoxys, represents this tribe in North
America.

Dec., 1940]

Linsley: Obriini

377

Genus Callimoxys Kraatz

Callimoxys Kraatz, 1863, Berl. Ent. Zeitschr., 7 : 105 ; Thomson,
1864, Systema Ceramb., p. 412; Fairmaire, 1864, Gen.
Coleopt. d’Eur., 4: 152; Lacordaire, 1869, Gen. Coleopt., 8:
489 ; LeConte, 1873, Smithson. Misc. Coll., XI, 265 : 307 ;
LeConte and Horn, 1883, Smithson. Misc. Coll., XXVI, 507 :
292 ; Leng, 1885, Entom. Amer., 2 : 27 ; Reitter, 1912, Faun.
Germ. Kafer, 4 : 28 ; Plavilistshikov, 1932, Best.-Tab. eur.
Coleopt., 102; 82, 106.

Head elongate ; front short, declivous ; vertex deeply impressed
between antennal bases; antenna shorter than the body in both
sexes, eleven segmented ; eyes deeply emarginate. Pronotum
longer than broad, narrower in front, widest behind middle,
sides tumid, disk tuberculate; anterior coxae separated by pro-
sternum; intercoxal process of mesosternum broad; elytra with
apices attenuated, subulate; legs elongate, posterior pair attain-
ing apex of abdomen in male, femora suddenly clavate, posterior
tibiae curved inwards, outer face armed with two rows of acute
tubercles.

Genotype: Stenopterus gracilis Brulle (by single reference).

Three species comprise this genus, one in southern Europe, two
in North America.

STUDIES ON THE BIOLOGY OF THE EPHEMER-
OPTERA. II. THE NUPTIAL FLIGHT^

By Herman T. Spieth
College of the City of New York

The spectacular nuptial flight of the ephemerids early at-
tracted the attention of scientists. Swammerdam (1675) re-
corded the emergence, nuptial flight, and oviposition of Palin-
genia longicauda on the rivers of Holland. He misunderstood
the character of the nuptial flight and thought fertilization took
place after the eggs had been deposited in the water. Reaumur
(1742) observed the nuptial flight of Ephoron virgo at Paris.
Both Swammerdam and Reaumur studied species whose flights
take place at dusk or after darkness has fallen, and thus they had
great difficulty in seeing the actual behavior of the insects.
When Collinson (1746) observed the flight of Ephemera vulgata
at Winchester, England, he noted that copulation took place in
the air and saw the typical up-and-down movements of the in-
dividuals. In 1750, he reported the observations of John
Bartram on an ephemerid species from the Schuylkill River,
Philadelphia. Bartram had seen a typical flight and also con-
cluded that copulation was achieved in the air. Charles de Geer
(1755 and 1771) observed Ephemera vulgata in Sweden, and
saw that the male copulated with the female by placing himself
beneath her and then bending his abdomen upwards. Because
of the short time that the copulatory act lasted, he was not able to
ascertain where the male grasped the female with his fore legs.
Eaton (1883) has given a brief account of the behavior of the
ephemerids during the nuptial flight, and since then many work-
ers have noted certain characteristic and interesting aspects of
the flights of these insects. Morgan (1913), Needham (1927),
and Needham, Traver, Hsu, etc. (1935) have dealt with the
subject at some length. There are, however, phases of the prob-
lem that have not been elucidated ; to present these data properly,
it is necessary to review the problem in its entirety.

1 1 wish to thank the authorities of the American Museum of Natural His-
tory and especially Dr. Frank E. Lutz for making available space in which to
conduct research work.

380

Journal New York Entomological Society [Vol. xlviii

THE NUPTIAL DANCE

On the basis of present information, all species of ephemerids,
except for possibly a few partlienogenic ones, engage in a nuptial
flight during their brief life as winged individuals. This flight
results in copulation and the insemination of the female.
Although there are variations and modifications to the flight, as
will be noted later, it typically follows a rather standard pattern.
Such a standard flight pattern can be observed in genera as di-
verse and distantly related as Baetisca, Hexagenia, Siphlonurus,
Ephemerella, and Baetis.

Briefly, the general pattern of this phenomenon, in which the
males play the dominant role, is as follows : After preliminary
patrolling, a number of males gather near or over a body of
water, into which the females will ultimately oviposit, and com-
mence an attractive and characteristic up-and-down dance. Some-
times the aggregation consists of countless individuals; at other
times it is made up of only a few males. Now and then a soli-
tary male can be seen in flight. The upward motion of the
dance is accomplished by the individual flying upward and for-
ward with strong, steady wing strokes. Meanwhile the fore legs
are stretched forward, the straightened body is elevated anteri-
orly and slopes downward posteriorly; the caudal cerci are
parallel with each other and extend posteriorly along the longi-
tudinal axis of the body. Having propelled himself upward and
forward for a brief time, he then ceases flying and, due to the
force of gravity, floats earthward with the wings extended up-
ward and somewhat outward. At the same time much of the
body, especially the thorax, comes to be almost parallel with the
earth while the posterior end of the abdomen is bent slightly
upwards. The long cerci extend divergently backwards and up-
wards. These cerci are always divergent and never parallel dur-
ing the downward movement. The fore legs are kept extended
forward and are bent slightly upwards. Except possibly for the
diverging of the cerci, the position assumed in the downward
flight is entirely passive and due to air resistance and gravity
rather than to any activity on the part of the individual. The
long cerci and fore legs seem to serve as balancers and stabilizers
in this passive downward movement. It is to be noted that the

Dec., 1940]

Spieth : Ephemeroptera

381

fluted, fan-like character of the wings is helpful in maintaining
the individual on an even keel. As Needham, Traver, Hsu, etc.
(1935, p. 103) have pointed out, the specimens often sway from
side to side as they passively descend. This is due to the effect
of the wind currents upon the inactive, lightly balanced indi-
viduals as they float earthward.

In such a nuptial dance there is no sign of timing among the
insects. Thus some are ascending while others are descending,
and so a constant passing and re-passing takes place.

The insects always face into the wind if there is any air move-
ment. If the air currents are strong and gusty, then the indi-
viduals are often blown away or down into the surrounding vege-
tation. In a moderate or gentle breeze, however, the forward
component of the upward flight just about equals the backward
component of the downward drop and the individual rises and
falls repeatedly in about the same area. If there is a slight
breeze or if air movements are completely lacking, then the
downward drop is vertical or almost so and the individual moves
forward with each upward flight. After a number of upward
and forward flights and downward drops, the insect makes a
quick reversal and faces in the opposite direction. There then
occurs a repetition of the ups and downs. Thus, invariably the
nuptial swarm is maintained over a relatively small area and
all the individuals fly in parallel directions.

Ephemerids can reverse their direction of flight with almost
lightning-like rapidity, can fly upward, downward, forward, and,
as is recorded below, they can hover; but they are apparently
incapable of lateral flight movements.

After the dance of the males has continued for a time, indi-
vidual females come flying into the swarm. Unlike the males,
they fly on an even keel, parallel to the ground, but usually in
the same direction as the males. The combined methods of flight
of the two sexes result in the female flying above a male that is
dancing up and down. The male recognizes the female and on
his up-flight approaches her from below, seizes her and the pair
immediately joins in copulation. De Geer (1755) was the first
to report that the male seizes the female from below, but he
did not see how it is accomplished. The feat is accomplished

382

Journal New York Entomological Society [Vol. XLVIII

because of the male’s ability to supinate his entire fore tarsi
and bend them backwards and thus grasp the pronotnm of the
female with his tarsal claws. Considerable speculation has
arisen as to the exact spot on the female that the male grasps.
Needham (1927) and Needham, Traver, Hsu, etc. (1935, fig. 13)
indicate that the mesonotum at the wing roots is the chosen spot.
All other workers have indicated the pronotum. My own ob-
servations suggest that the posteromedial edge of the pronotum
is the point of attachment.

Almost simultaneously the male bends the end of his abdomen
upward and forward and grasps the eighth abdominal segment
of the female with his genital forceps. At the same time the
penes are inserted into the genital ducts of the female which open
on the conjunctival membrane between the seventh and eighth
sternites. In such a position the head of the male lies under
the mesothorax of the female. Thus the thorax of the male
extends posterior to that of the female and the male’s abdomen
is bent acutely upward and forward.

The united pair continues to fly but, with few exceptions, its
combined flight powers are not sufficient to support them for
long and they graduall}^ descend toward the ground. Generally
the two separate before they reach the substratum over which
they have been flying. Thus the copulatory act is restricted to
a few seconds. In Siphloplecton hasale, however, not only does
the united pair continue its combined flight but it is able to
avoid other males that attempt to displace the copulating male.
Kepeated timings on this species have shown that even here
copulation lasts only from one to one and one-half minutes.

Sometimes pairs will remain in copula even after they have
descended to the substratum. De Geer (1771) observed pairs
of Ephemera vulgata upon the vegetation. Needham, Traver,
Hsu, etc. (1935) relate that many pairs of Siphlonurus sp.?
were observed to light on a parked car. Eaton (1883) observed
a pair of the genus Ecdyonurus remaining in copula for at least
‘‘six or seven minutes after they had come to rest.” Pairs of
Siphloplecton l)asale will remain in copula when captured in an
insect net. Needham (1927) found a pair of Cinygmula mimus
in copula even after the insects had been killed in a cyanide jar.

Dec., 1940]

Spieth : Ephemeroptera

383

A figure of this pair has been published but is not typical in
that the male is too far forward with respect to the female, his
abdomen is not sufficiently bent, and the fore tarsal claws reach
to the middle of the mesothorax instead of to the posterior edge
of the prothorax. A much more accurate picture of copulation
is given in ‘'The Biology of Mayflies” (1935, fig. 13) except
that the fore legs of the male are not delineated correctly.

VARIATIONS OP THE NUPTIAL FLIGHT

There are variations to the typical flight described above that
should be noted. Eaton (1883, p. 9) writes that Heptagenia
and its allies maintain a hovering position so that they are
“locally designated in the valley of the Axe (Devon) as Yellow
Uprights.” Cook (1940) observed such a behavior in Stenonema
vicarium. I have seen it repeatedly in various members of the
genus Stenonema. In such cases the flight varies from the typical
in that the male, instead of moving up and down, hovers on an
even level, a dance accomplished by the insect assuming the fol-
lowing position : the body is held rigid and straight with the
longitudinal axis almost perpendicular to the ground. The fore
legs are extended along the axis of the body. The caudal cerci
are held widely divergent and almost parallel to the ground, i.e.,
almost at right angles to the longitudinal axis of the body, while
the wings are rotated forward. An individual when flying in
this manner is reminiscent of a feeding humming bird. In such
a position the male can hover in one place or move forward and
parallel to the substratum at a slow rate. It can not fly side-
ways or reverse its flight. In order to achieve a reversal of
direction, the individual tips forward abruptly and flies in
normal manner for a short distance, i.e., a foot or so, reverses
and then reassumes the hovering flight. What factor or factors
are responsible for the individual assuming the hovering type of
flight are obscure. Apparently only some members of the fam-
ily Heptageniidae are capable of this variation. It is to be noted
that such a flight pattern is assumed only when there is no wind.

DOWNWARD FLIGHT MOVEMENTS

Not always is the downward drop of the male in the nuptial
dance passive. Sometimes he actually flys downward. Need-

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Journal New York Entomological Society [Vol. XLVIII

ham, Traver, Hsu, etc. (1935, p. 103) report that Baetis flavis-
irigia makes the upward part of the flight with great rapidity
while the downward passive drop gives the impression of a
dogbane seed floating earthward. If, however, the nuptial flight
of this species is conducted in a brisk breeze, then the dowmward
drop is accomplished by vigorous movements of the wings and
it is just as rapid as the upward movement, and the individual
keeps rising and falling in an almost vertical line. So far as
my observations go, such downward wing-movements are re-
stricted to small, light, fragile species. It has not been observed
in large, heavy-bodied species.

SUBSTRATUM COPULATION

Practically all species of ephemerids mate in the air, although
Eaton (1883, p. 10) reports that Plethogenesia papuana mates
on the surface of the water. In this species the fore legs of
the males, because of the abbreviated tarsi, are short. A similar
condition is found in the species of the genera Mortogenesia and
Anagenesia. It is possible that the species of all three of these
specialized genera may mate on the surface of the water.

Occasionally species that normally mate in the air will attempt
to copulate while on a Arm substratum. Reaumur (1742) ob-
served that the male individuals of Ephoron album, when at-
tracted to land near his lantern, would attempt to copulate with
females with whom they came in contact.

While watching a pair of Ephemera guttulata on the under-
side of a Platanus occidentalis leaf, I observed that the male
brought himself parallel and close beside the female, grasped
her abdomen with his genital forceps and proceeded to copulate
in the usual manner. In this instance, the fore legs of the male
were not used and both individuals remained attached to the
leaf.

SEMI-NOCTURNAL AND NOCTURNAL SPECIES

Species of genera such as Ephoron, Campsurus, and Palin-
genia emerge just at darkness or even during the night. Since
the meso- and metathoracic legs of the females and the males of
Campsurus are either absent (Campsurus), or so weak (Epho-
ron and Palingenia) that they can not support the individuals.

Dec., 1940]

Spieth: Ephemeroptera

385

they mate immediately; the females oviposit, and then the indi-
viduals of both sexes die. The total life span of the winged
stages in such species is from one to four hours. The nymphs
of these are negatively phototactic and positively thigmotactic
burrowing forms that normally never come in contact with the
light. Thus a complete generation may exist without the indi-
viduals seeing daylight. Such species do not indulge in a typical
nuptial dance. The males fly rapidly back and forth over the
body of water from which they have emerged. The flight is
parallel or almost so with the water and without the up-and-down
motion. Tientsuu (1935) reports that males

have an undulating flight. Every so often the individuals
quickly reverse their direction of flight. This, plus the rapid
beating of the wings, gives these forms the appearance of being
in a great hurry. Just how the males And the females in the
total or almost total darkness, I have never been able to observe
exactly. All these males have undivided compound eyes that
are relatively small. Perhaps their vision is extremely acute,
but fragmentary observations indicate that they may establish
contact by tactile rather than visual means. In this connection
it should be noted that these species usually emerge in great
numbers. This plus the fact that the nuptial flight is held close
to the surface of the water insure a relatively large chance for
actual contact between individuals. The method of copulation
seems to be the same as in all other species.

In these species, the female remains during her entire winged
existence a subimago.

COMPOUND EYES OF DIURNAL SPECIES

Except for the semi-nocturnal or nocturnal species, the ephe-
erid males have large ej^es that are greatly expanded on the
dorsal surface. In many the eyes are actually divided into a
large dorsal and a smaller ventro-lateral portion. This condi-
tion reaches its extreme development in Baetis and its relatives
where the dorsal part is carried upward on a stalk and the visual
elements face dorsally while the small ventral portion is more
rounded and the visual elements face outward somewhat as in
the normal insect eye. Even in those species in which the eye
is not completely divided morphologically (such as Hexagenia,

386

Journal New York Entomological Society [Vol. XLVIII

Stenonema, Siplilonuriis, etc.), it is easily seen that there is a
functional division.

Repeated observations indicate that the dorsal part of the eye
is used by the male to recognize the female as she flies over him
while he is engaged in the nuptial dance. In the morpholog-
ically divided eye types where the dorsal part faces upward {i.e.,
Baetis, Pseudocloeon, etc.), the upward component of the nuptial
dance is almost vertical. Conversely those species which do not
have the eyes separated morphologically and in which the upper
part is only slightly larger than the lower part {i.e., Siphlonurus,
Hexagenia, Stenonema, etc.), the upward movement of the nup-
tial dance is not vertical but forms about a forty-flve degree
angle with the substratum.

The focal distance of the upper part of the eye apparently is
rather short. The male seizes only those females that come
within a short distance of him. The males normally are able to
identify the females of their own species, but often they make
mistakes, if other insects are flying in the same immediate area.
I have seen Stenonema fuscum males repeatedly attempt to copu-
late with caddis and stone flies. A Cinygmula mimus male
was captured while trying to mate with a large chironomid.
Tills may explain Venour’s observation (1906), upon E. dancia
males mating with female subimagoes that had just emerged and
were forced to fly through the nuptial dance in order to reach
the surrounding vegetation. Ide (1930), has also observed
male imagoes of Callibaetis americanus mate with female sub-
imagoes of the same species.

The lateroventral part of the eye is employed by all species
to prevent the individual from coming into close proximity with
large objects or protuberances of the substratum. The ephem-
erids, especially the males during the nuptial flight, are ex-
tremely alert and have a rather keen vision. Anyone who has
collected ephemerids can testify that while flying they avoid with
great consistency any object that projects from the background.
Thus if a bridge crosses a stream over which they are engaged
in nuptial flight, they will invariably avoid it, rising when they
approach it so that they maintain the same distance from it that
they do from the surface of the water. When a collector places

Dec., 1940]

Spieth: Ephemeroptera

387

himself upon the bridge, they fly still higher over the spot where
he is standing. This holds true whether he moves or remains
perfectly still.

TIME OF FLIGHT

As indicated above, most species are diurnal. These species
mate during the day or at twilight, the exact time depending
upon environmental and seasonal differences as well as upon
specific pecularities.

In general those small species having hyaline wings and males
in which the abdominal segments 2-7 are hyaline or semi-hyaline
mate during the middle of the day and only rarely at twilight.
Thus species of Baetis, Callibaetis, Pseudocloeon, Cloeon, Cen-
troptilium, Paraleptophlebia, Habrophlebia and Habrophleboides
are often seen swarming at various times in broad daylight. I
have a record of B. moffati mating at seven o’clock in the
morning.

Large, dark, heavy-bodied, conspicuous species such as Hexa-
genia. Ephemera, Potamanthus, Isonychia and most members of
the family Heptageniidse usually swarm from just at sundown
to darkness.

No hard and fast rules can be drawn for any species because
environmental effects play a large part in the timing of the
nuptial dance as can be seen by the following typical observation.

While collecting on Spring Creek in Yellowstone Park, Wyo-
ming, July 7, 1936, a thunder shower of small dimensions de-
veloped about five o’clock in the afternoon. The spot where I
was collecting was on the edge of the storm area and only a
few raindrops fell at this point. Individuals of both sexes of
Ephemera simulans appeared and engaged in a nuptial flight.
In a few minutes the storm disappeared and the individuals
immediately dispersed and were not to be seen until they reap-
peared just at sundown about two hours later. Repeatedly I
have observed, in many diverse genera, that the appearance of
a storm with its attendant drop in temperature and light in-
tensity will stimulate adult ephemerids to activity. Perhaps
the relative humidity is also important since the ephemerids
are rather sensitive to desiccation.

388

Journal New York Entomological Society [Vol. xlviii

A combination of factors — latitude, elevation, season, tem-
perature, humidity, light intensity, and atmospheric pressure —
rather than any single one is probably responsible for stimulat-
ing the imagoes to leave their resting place on the surrounding
vegetation and indulge in the nuptial dance. Thus the species
that normally mate at twilight will sometimes be found mating
in the middle of the day.

PLACE OF NUPTIAL FLIGHT

Although no perfect rule can be drawn, members of each genus
are fairly consistent in the locality selected for the nuptial dance.
Ephoron and Campsurus mate over the water; Ephemera over
the water or close by; Hexagenia along the edge of the shore;
Siphloplecton, Csenis, Choroterpes, Stenonema, Iron, Heptagenia,
Cinygmula, and Rhithrogena usually but not always engage in
their nuptials over the surface of the water. I have, however,
observed exceptions in all of these genera.

In comparison, I have never seen Blasturus, Trieorythus or
Ephemerella, with the sole exception of the early spring E. suh-
varia, swarming over the water. These three genera all choose
open areas a distance from the stream for their dance.

Small, clear- winged species of Thraulus, Habrophlebia, Habro-
phleboides, Baetis, Cloeoii, Pseudocloeon, and many species of
Paraleptophlebia choose small openings in the trees or bushes
along the edge of a body of water. Only seldom are they seen
over the water.

Species of Isonychia and Siphlonurus choose almost any spot :
sometimes over the water, sometimes over a grassy meadow,
sometimes in the tree-tops. Individuals of a single species may
be found in different localities on the same day at the same
time.

Ephemerids, both males and females, often mistake wet roads
for bodies of water, and forms which normally mate over water
will often be found dancing and ovipositing over a road.

The height at which the dance takes place is also extremely
variable. Generally when a strong wind is blowing, the indi-
viduals are close to the ground. Under normal conditions most
species dance from five to thirty feet above the substratum. A

Dec., 1940J

Spieth: Ephemeroptera

389

few, such as Ephemerella, usually conduct their dance high in
the air, about thirty to one hundred feet above the ground.

FATE OF THE MATED INDIVIDUALS

Copulation achieved, the female flies off to deposit her eggs.
The male, being polygamous, returns to the dance. This dance
may continue for several hours. For species that mate at twi-
light, darkness is the limiting factor to the dance of the males.
In species of Campsurus and Ephoron the nuptial dance lasts
until the individuals are exhausted. Apparently the same thing
happens to numerous individuals of other species.

The males of most species, if they have not mated or are not
exhausted, return at the end of the dance to the vegetation and
await the dance of the following day. Careful watching of a
dance will always show that a large percentage of the males fly
back into the surrounding vegetation. Since they are unable to
feed in the winged state, they must live upon stored energy. A
still greater danger is that of desiccation. I have often seen
males fly rapidly from the vegetation and drop down upon the
surface of the water with wings outspread. After remaining in
this position for a few seconds, they would then fly back into the
vegetation. As is well known the adult ephemerid’s body sur-
face is hydrofugic. Experiment has shown, however, that the
area of the degenerate mouthparts is not hydrofugic in the
imagoes and that individuals are capable of imbibing water.
Apparently the males referred to above return to the water for
a drink.

In any case, the winged life span of an ephemerid is relatively
short and has only one function; the fertilization and ovipo-
sition of the ova.

CONCLUSIONS

1. All ephemerids, with the exception of a few parthenogenic
forms, as adults engage in a nuptial dance which insures the
males copulating with the females.

2. This dance and the copulatory movements follow a rather
standard pattern for all species of the order. This is another
evidence that the order is monophyletic.

390

Journal New York Entomological Society [Vol. XLVIII

3. There are a few modifications to the standard dance pattern.
These (hovering, horizontal and vertical) are modifications to
meet specialized conditions.

4. The dance may take place at any time of day or night
depending upon the species and environmental conditions.
Within limits, however, each species has a preferred time for
mating.

5. Various genera usually choose specific types of localities to
engage in the dance.

6. The males of many species live several days and engage in
several nuptial dances. They apparently counteract desiccation
by their ability to imbibe water.

SELECTED BIBLIOCEAPHY

1675. Swammerdam, Jan, Epliemeri vita, of afbeeldingli van’s mensclien
leven, vertoont in de wonderbaarelycke histoire van bet vliegentende
een-daglilevent Haft of Oever-aas, etc.

1742. Eeaumur, E. a. F., Memoires pour servir a 1 ’histoire des insects.
Tome 6, pp. 457-522.

1746. CoLLiNSON, P., Philos. Trans. Eoy. Soc. London (abridged), vol. 9,
p. 290.

1750. COLLINSON, P., Philos. Trans. Eoy. Soc. London (abridged), vol. 10,

p. 28.

1775. deGeer, C., Mem. Sav. Etr. Acad. Paris, Tome 2, pp. 461-469.

1771. deGeer, C., Memoires pour 1 ’histoire des insects. Tome 2, pt. 2, pp.
617-660.

1883. Eaton, A. E., Eevisional Mono. Eec. Ephemeridae, pt. 1, pp. 8-11.
1906. Venour, S., Ent. Mo. Mag., vol. 42, p. 258.

1913. Morgan, A, H., Ann. Ent. Soc. Amer., vol. 6, pp. 371-413.

1927. Needham, J. G., Canad. Ent., vol. 59, pp. 133-136.

1930. iDE, F. P., Canad. Ent., vol. 62, p. 227.

1935. Needham, Traver, Hsu, etc.. Biology of Mayflies, pp. 102-105.

1935. Tientsuu, L., Suomen Hyonteistieteellinen Aikokauskirja, vol. 1, no.
1, pp. 3-23.

1940. Cook, H., Ent. News, vol. 51, no. 1, pp. 12-14.

A NOTE ON REARING THE BROOD OF POLISTES
FUSCATUS, FABRICIUS (HYMENOPTERA—
VESPIDiE)

On several occasions during the past season, it has been my
fortune to collect several nests of Polistes fuscatus, Pabricius.
As it was expedient to kill off all the adults in collecting the nests,
the brood had no means of securing nourishment. It was decided
to attempt to rear the existing brood, which ranged from eggs to
nearly matured pupae.

The larvae required both a liquid and solid diet. The liquids
were administered to each larva through a small glass pipette.
A drop of juice or water slightly smaller than the head of the
larva was discovered to be sufficient for one drink ; naturally this
volume would vary with the size of the larva. All larvae greedily
accepted tap water and would occasionally accept a dilute aque-
ous solution of glucose or saccharose. Upon occasion the smaller
larvae would accept the offerings of pasteurized milk but this was
not consumed with any avidity.

The solid diet was prepared by chopping small bits of meat
into fragments about one half of one cubic millimeter in volume
and then making a paste out of the mass by the addition of water.
This kept the meats juicy and permitted the larvae to handle the
particles more easily. Forceps with medium size jaws were used
in administering the solid foods ; as much food as would adhere
to the forceps through capillarity was sufficient to feed one large
larva. Raw and cooked beef, lamb and chicken, prepared as
above, were readily eaten as were liverwurst and ham (spiced).

Two or three feedings and as many drinks every day were
enough to satiate the greedy larvse and permitted them to grow
at a reasonable rate.

The absence of trophallactic relations with adults seemed not
to have an effect on the development of the brood, although at
all times the larger larvae would produce their trophallactic secre-
tions when touched upon the head. These larvae would indulge
in autotrophallaxis by imbibing their own secretions, inde-
pendently of how well they were fed. This imbibition seems

392

Journal New York Entomological Society [Vol. xlviii

necessary for tlie maintenance of nest cleanliness as the paper
would soon become sodden and moldy if the secretion were not
removed ; it therefore seems that autotrophallaxis might be a
practical habit rather than an indication of weakness in the
colony.

To gain access into the deep cells in the middle of the nests, it
was found expedient to tear away the paper walls of the cells.
When the larvae in these cells began to pupate they could not
produce enough silk to build up their compartments. Indeed,
they had no instinct for repairing the nest. One larva even tried
to build a cocoon cap attached only to the head of an adjacent
larva.

To circumvent this difficulty, two types of artificial cell were
made. One was a block of wood bored with holes 5 mm. in
diameter and 16 mm. in depth; the other was a set of 5 mm.
diameter paper tubes. When the large larvse were observed to
move their heads about in the circular motion characteristic of
the pre-pup ating Polistes, or when they would accept no further
food, they were removed from the original nest and placed in
either the wooden block or the paper tubes. These served as a
satisfactory substitute for the nest as the larvae spun their white
silk pupal caps over the apertures and settled down to pupation.
Perfectly normal adults emerged in due time.

Adults that emerged prior to the pupation of the last larva
were removed that they might not interfere with the experiments.

The span of life of the larvae could be varied at will by increas-
ing or decreasing the feeding. Fifteen days to six weeks were
spent in the larval stage by individuals that were respectively
overfed and nearly starved. The average pupation period was
nineteen days with the temperature varying from 66 to 87
degrees F.

The only real difficulty encountered in rearing the brood was
in feeding the smaller larvae whose capacity was so small that
even the smallest portions offered them could not be completely
consumed. This created a problem in nest cleanliness, so the
excess had to be carefully removed after each feeding.

Another important consideration in the health of the brood
was sufficient ventilation. In closed or poorly ventilated con-

Dec., 1940]

Gaul: Polistes

393

tainers, the formation of molds caused a high mortality among
both larvae and pupae. Nests fully exposed on a shelf or table
remained free from molds although precautions were necessarily
taken against the predatory activities of ants.

The inverted position of the nest seemed not to interfere with
the more or less normal development of the young.

By virtue of the few precautions mentioned, it was found prac-
ticable to rear adult Polistes from eggs, thus entirely supplanting
the natural relations between adult and brood. Although the
trick of rearing Polistes is of no great consequence, the possibility
of supplanting the adults may lead to further investigations on
their social conduct. — Albro Tilton Gaul.

BOOK NOTICE

Introducing Insects, A Book for Beginners. By James Needham,
with illustrations by Ellen Edmonson. The Jaques Cattell
Press, Lancaster, Pennsylvania, 1940. 7^x5^ inches, vi +

129 pp., illus. $1.50 postpaid.

This is a slim, attractively bound, well printed volume by a
well-known entomologist whose writings always command re-
spect, and attention. In it. Dr. Needham refrains from “human-
izing” his subjects and also from treating them in journalistic
style, which methods are commonly used by writers in attempt-
ing to popularize insects. Instead he introduces the insects as
they really are, in simple language free of technicalities, and his
examples are all species that one meets within everyday life in
the home, in our possessions, in our food, and in our rambles in
fields and along roadsides.

In addition to sections on butterflies, dragonflies, common
caterpillars, leaf-eating beetles, mosquitoes, insects that eat our
food, etc., there are sections devoted to collecting and rearing
insects, and to the importance and pleasure of studying them.

It is a good book for the young or old beginner in entomology
and for the citizen of any age who is inquisitive enough to want
to know something reliable about the creatures with which he
comes in contact at various times during his lifetime, and the
numerous illustrations in the text add to its merit. — H. B. W.

394

Journal New York Entomological Society [Vol. XLVIII

An Introduction to Entomology. By John Henry Comstock,
Late Professor of Entomology and Invertebrate Zoology in
Cornell University. Ninth edition, revised. Ithaca, New
York, Comstock Publishing Company, Inc., 1940. x 6
inches, xix + 1064 pp., Frontis., 1228 Pigs. $5.00.

The appearance of the ninth edition of this work of acknowl-
edged excellence makes superfluous any praise that I might be-
stow upon it. And so this notice is merely an announcement of
its availability in a revised form and of its recognition of the
interest in biological control by its revision and extended discus-
sion of certain superfamilies of the Hymenoptera. Although
entitled An Introduction to Entomology, it is really a compre-
hensive general treatise involving various ramifications of the
external and internal anatomy of insects, of metamorphosis, and
of classification, with numerous analytical tables for the deter-
mination of families. In this latest edition, new discoveries and
opinions have been included.

Its publication reminds me that the only book I ever borrowed
and failed to return was a copy of A Manual for the Study of
Insects, by John Henry Comstock and Anna Botsford Comstock,
2nd edition, published in 1897. I borrowed it in 1900 and still
have it. I always meant to return it and never felt right about
keeping it so long, but at the time it was indispensable to me,
and I could not spare it. When I was ready to return it, the
original owner was no longer alive.

The present work, I am sure, is just as necessary to present day
entomologists, but I recommend its purchase rather than borrow-
ing and keeping it for forty years. — H. B. W.

BOOK REVIEW

Fauna of British India, Dipt era Yl, Calliphoridce. By B.
Senior- White, D. Aubertin and J. Smart (London, March 28,
1940).

This beautiful volume of 288 pages, 152 figures and 1 map is
an important contribution to the subject of the Oriental Mus-
coidea, but unfortunately it is marred by numerous errors. It
appears from the preface that Senior- White is mainly responsi-
ble for the system of classification adopted including generic
scope, as well as for terms and names employed in the volume.
It is much to be deplored that uniformity in taxonomy and termi-
nology is still a postponed consummation. Still more to be de-
plored is the current refusal to accept the priority principle in
nomenclature.

As to taxonomy : Bengalia and Pollenia and their allies are not
nearly so closely related to Calliphora as is Chrysomya, yet they
are included in the Calliphorinas while Chrysomya is recognized
as the type of a separate subfamily. The Chrysomya contingent
merits no more than tribal rank. Both bare and ciliate remigia
occur in the small tribe Mesembrinellini, which shows that this
character is not even of tribal value.

Tainanina is included in the Calliphorinse, but it belongs un-
mistakably in the very distinct family Melanophoridag.

Chsetoptiliopsis is also included in the Calliphorinse. Accord-
ing to its author, it does not belong even in this superfamily but
is referable to the Macquartiini in the superfamily Oestroidea.
Baranov’s reference implies a well developed infrascutellum in
Chsetoptiliopsis. Both the original description and the one given
here indicate the correctness of his reference.

Elephantoloemus is abundantly distinct from Booponus gener-
ically by having markedly different head and both belong with
Bengalia tribally. Both the Bengalia and the Pollenia groups
belong in the family Rhiniid^.

Numerous valid genera are sunk as synonyms in the volume,
but they will eventually be recognized on their genetic index (see
MM, IV, 269).

396

Journal New York Entomological Society [Vol. XLVIII

As to terminology ; The great desirability of a uniform English
terminology securely based on morphology led the reviewer to
make the most careful selection of terms possible for the anatomic
parts of the fly and the early stages both externally and inter-
nally. It is given in MM, I & II. The earliest and most ap-
propriate terms are there fully elucidated to prevent confusion
and error. A quite different terminology is employed in the
present volume with the apology that, while it has practically
no morphological value, it was chosen for convenience and on the
grounds of usage. It must not be forgotten that a sound and
lasting terminology can be based only on a valid interpretation
of homology, which in turn can be based only on morphology.
Few exceptions to this rule are permissible and even these are
questionable (MM, II, 63).

Evidently the student of myiology must still for a time learn
the separate terminologies of all the workers publishing not only
in his own language but in all other languages employed. This
is a great hardship, causing great loss of time and frequently
much confusion if not actual error. Further, why use a whole
descriptive clause often repeated, as in the subfamily key on page
26 and elsewhere, to avoid saying “greater ampulla”? Obvi-
ously because the user did not know that the part has a distinc-
tive name. In same key, a four-line sentence could have been far
more intelligibly expressed by saying “posthumeral bristle” or
just “PH” present or absent. Besides, the sentence says what
it does not mean. It means the posterior (not anterior) two
bristles !

As to nomenclature : The earliest generic name used in the
volume is Sarcaphaga (1826), yet the Sarcophaginae are included
in the family Calliphoridae, derived from Calliphora (1830),
ignoring priority in derivation of group names above the genus.
Sarcophaginae should be Stephanostomatinae, derived from
Stephanostoma (1794—1802), which much antedates Sarcophaga.
There is at present a mistaken movement afoot to ignore the
priority principle in derivation of group names above the genus,
but it can not prevail in the long run. Future nomenclature can
find stability only in observing this principle. Arbitrary enact-
ment to the contrary can only cause confusion and loss of time.

Dec., 1940]

Townsend: Book Eeviews

397

Further, the original spelling of names must be adhered to in
order to avoid confusion and emendations must be treated as
synonyms.

Errors in the volume require correction as below, by pagina-
tion. Note that Gt means genotype ; Ht, holotype ; At, allotype ;
Coll, collection; MM, Manual of Myiology; AMNH, Annals &
Magazine of Natural History.

Pp. 28-29. In 1931 Townsend (AMNH, 10, VIII, 375) gave
Cynomya fortis Wlk. as Gt of Hypopygiopsis on Ht from
Borneo described in 1857. Years later Walker described Pyrel-
lia fumipennis 2 from New Guinea, being evidently same species.
His Silhomyia fumipennis J from Singapore, described in 1857,
is doubtfully the same.

Pp. 32-33. Gt of Calliphora is voniitoria L. as shown by
Desvoidy’s description.

Pp. 37-38. Tricycleopsis paradoxa VII is Bengaliini, while
Pseudo calliphora semifulva Mil. is Calliphorini. Ht J of latter
is in Amsterdam.

Pp. 41-42. Hemipyrellia stands on its Gt curriei TT. from
Mount Coffee, Liberia, which is evidently a distinct species from
fernandica Mq. from Fernando Po. H. orient alis TT. (1927) is
distinct on color of antennae and palpi.

Pp. 46 & 53. Agoracrites was validated by Aubertin (1933),
Lucilia porphyrina Wlk. being referred to it. Mount Ophir,
alias Talaman, is in Sumatra.

P. 70. Ht 2 of Caiusa indica Sf. is in Paris and At is in
Milan (MM, V, 72).

P. 72. C. duhiosa VII, is distinct, Ht 2 iii DEI.

Pp. 78-81. Larva of Booponus and that of Elephantoloemus
are certainly (not probably) the cause of the sores, the two
differing markedly in parts attacked.

P. 82. Chaetoptiliopsis, as already noted, does not belong in
the Muscoidea.

P. 88. Ht 2 of Musca jejuna F. is from Bengal and in Copen-
hagen F. Coll (AMNH, 10, VIII, 371).

P. 91. Ht2 of Bengalia lateralis Mq. is in Paris (MM, V, 67).

P. 101. Ht of Musca varicolor F. is in Copenhagen F. Coll
(MM, V, 70).

398

Journal New York Entomological Society [Vol. XLVIII

P. 102-103 & 106. Catapicephala splendens Mq. is a synonym
of Musca micans P., Ht in Copenhagen P. Coll from East
Indies. Trongia viridis TT. from Siam is same (AMNH, 10,
VIII, 375).

Pp. 108-110. Tainanina grisella Vll. is a distinct species, the
squamal hairs brown. Belongs in Melanophoridae, as already
noted. Pignre 53 is upside down.

Pp. 111-114. Coquillett’s designation (1910) of cognata M.
as Gt of Melinda is invalid because not originally included.
Townsend’s designation (1916) of ccerulea KD. is valid and
equals cognata M. The synonymic name ccerulea M. is pre-
occupied by Wiedmann (1819). The Sumatran species given as
Melinda are Neomelinda, to which may also belong M. leucocera
RD. from Australia.

Pp. 119-132. P. townsendi is a synonym of Ocrisia testacea
TT. Strange disposition to put all the tribe Polleniini except
Wilhelmina in one genus.

P. 133. Ht 5 of W. nepenthicola Vll. is in Amsterdam. The
genus and species are Vll.

P. 135. Rondani (Arch. Zool. Modena, III, 27, 1863) said
^‘Marginalis del Pabricius, che forma il tipo del Genere Chry-
somya. ’ ’ This is invalid as a Gt designation, since this name was
not originally included. Coquillett (1910) merely repeated this.
Townsend (1916) made the first valid designation as regalis RD,
which equals marginalis W. (not P.). Paracompsomyia and
Pycnosoma are synonyms (MM, V, 124). Compsomyia Gt is
Musca megacephala P., Callitroga B is synonym of Compsomyia
(MM, V, 126). Somomya R equals Calliphora (MM, V, 141).
The other genera are valid.

P. 137. M. marginalis W. Ht 5 is in Copenhagen Westermann
Coll (MM, V, 124).

P. 138. M. megacephala P. type locality Guinea was error for
East Indies.

P. 149. There is no doubt of the complete absence of Rhiniini
in the Americas.

P. 151 & 157-158. Strongyloneura is abundantly distinct from
Isomyia, as shown by the characters given for latter’s Gt delec-
tans Wlk. On his two visits in 1928 to the British Museum, the

Dec., 1940]

Townsend: Book Eeviews

399

reviewer was told that Ht of same was lost. Gt of Thelychseta
BB. is chaly'hea BB. from Borneo. Tachina viridaurea W. from
the East Indies belongs to Bhyncomya, Ht 2 in Copenhagen
(AMNH, 10, VIII, 372) ; no in Copenhagen and no types in
Vienna. Musca viridaurea W. from Java is congeneric with
chaly'hea BB. but not conspecific. Musca munda W. from Java
is also distinct and the description under 105 is of this species,
which is apparently referable to Strongyloneuropsis.

Pp. 152-154. Description 100 is of Eucosmina vittigera MU.
(not of Strongyloneuropsis malayensis TT.). Figure 74 of J'
genitalia is said to have been drawn from the type of vittigera,
but the only types of that species are females.

P. 163. Referring to footnote, S. viridana TT. types agree
with the description in the “parafacial hairs partly or wholly
blackish. ’ ’

P. 168. The seven species with J Ht can all be determined in
the female.

P. 172. Description 119 is of ftdvicornis Big. Dictya cenea P.
is a distinct species, Ht 2 Copenhagen F. Coll (AMNH, 10,
VIII, 372). Synamphoneuropsis viridis TT. from Nepal frontier
(Sukhwani) is distinct.

Pp. 176-177. Eucosmina, given on nonexistent J' type page
152 as synonym of S. malayensis TT., is here given again as
synonym of Alikangiella on the actual 2 types. The descriptions
under 100 and 123 are exactly the same except a slip on midtibial
bristling. It seems that two of the authors disposed of Eucos-
mina without either one knowing what the other had done and
both dispositions got published in the volume ! The genus is
distinct from Alikangiella as well as from Strongyloneura.

P. 179. Metalliopsis setosa TT. with bare prosternum and 5R
widely open is made conspecific with Chlorrliynchomyia clausa
TT. with bristled prosternum and closed 5R, the two being very
distinct generically. Ht 2 of C. clausa is not in the Berlin Mu-
seum but in the DEI in Berlin-Dahlem.

P. 183. Gt of Rhyncomya is Musca ruficeps F. B. callopis
Lw. does not belong. But Tachina viridaurea W. belongs.

P. 187. Alikangia Gt is pulchella Vll. Genus distinct from
Borbororhinia.

400 Journal New York Entomological Society [Vol. XLVIII

P. 191. Musca lunata P. Ht J' is in Copenhagen F Coll (MM,
V, 108).

P. 192. Musca discolor F. Ht J' is in Copenhagen F Coll
(MM, V, 98).

P. 197. Idia xanthogaster W. Ht J is in Copenhagen Wester-
mann Coll (MM, V, 101).

P. 199. Idia mandarina W. Ht & At J are in Copenhagen
Trentepohl Coll (MM, V, 100).

P. 204. Rhinia testacea KD. from Mauritius is a synonym of
Idia apicalis W. from Tenerife. Ht J' of B. glossina K,. is in
Genoa (MM, V, 105).

P. 206. Chlororhinia viridis TT. is distinct from exempta
Wlk. latter being a lowland species while former is a highland
species. In the key the word ‘Hrons” is mistakenly used for
frontalia.

P. 208. Gt of Bl^esoxipha is laticornis M. that of Ravinia is
striata F. and that of Helicobia is rapax Wlk. Glaucosarcophaga
hnahi TT. is wrongly credited to Parker.

P. 225. Species 164 is Ravinia striata F.

P. 231. Enderlein’s J' of Lioproctia doleschalli JT, deter-
mined by him as Sarcophaga aurifrons Dll. has the propleura
bare and the midtibise thickly long villous, besides possessing 1
PA. Evidently the species here described is distinct in lacking
PA and having bare male midtibiee. Whether it is the same as
DoleschalPs Ht is not certain. Johnston is twice printed
Johnson.

P. 276. Species 211 is Stephanostoma stepliano stoma Lz.
(MM, VI, 70-74).

P. 277. Sarcophila R. is synonym of Agria RD., Gt latifrons
Fll. Leucomyia is very distinct generically.

P. 279. Musca affinis Fll. is Gt of Pseudosarcophaga. Agria
hutsoni SW does not belong to latter genus.

P. 281. Tachina nuba W. is evidently a Disjunctio. The
maggots have been used in surgery! (MM, V, 190).

Practically all the errors and omissions in the volume could
have been avoided by consulting the Annals & Magazine of
Natural History, series 10, VIII, 371-378 (1931) and the Manual
of Myiology, I, II, V, VI (1934-1938).

Charles H. T. Townsend

A PROMISING FUNGOUS PATHOGEN OF ADULT
JAPANESE BEETLES (POPILLIA
JAPONICA)

The New Jersey Department of Agriculture has actively con-
ducted Japanese beetle suppression work since 1927. During this
period opportunity of every description has been available to
evaluate the comparative benefits of the weapons of artificial and
natural control which are now at our disposal. In a report issued
by this Department in 1932, a statement was made that a com-
municable disease among adults would probably prove to be the
most vulnerable channel of Japanese beetle suppression. The
control recommendations even as they apply today, consist largely
of the use of repellents rather than chemicals which have a de-
structive effect on the beetle population. The performance of
Japanese beetle traps in New Jersey has not been sufficiently con-
vincing to justify the continuation of experimentation on this
phase of artificial Japanese beetle suppression.

This Department, in view of its conviction that a communicable
disease among adults would make serious inroads in the Japa-
nese beetle population, employed an entomologist for a period of
two months during the summer of 1933 and stationed him in the
most heavily infested county in the state. His job consisted of
examining piles of trapped Japanese beetles, taking samples
therefrom and making careful examination of these samples to
ascertain the presence of symptoms of bacterial or fungous dis-
eases. These observations did not yield any positive findings.
Since 1932 our suppression crews have been instructed to give
special attention to the appearance of dead and living adult Japa-
nese beetles with particular reference to the appearance of disease
symptoms. To date our field observations have not encountered
any such indications of a pathogenic organism being destructive
among the adults.

In October, 1939, a list of a local library accessions included a
reference to a paper which appeared a few months earlier in the
Canadian Journal of Research, the title of which was ‘‘Pathogen-
icity of Beauveria hassiana (Bals.) VuilL, on Colorado potato

402

Journal New York Entomological Society [Vol. XLVIII

beetle larvse. ’’ The author of this paper, a staff member of the
Canadian Department of Agriculture, was the recipient of a
letter from this Department requesting a copy of the publication
on this subject and also, if possible, a culture of the fungus.
Upon the completion of the necessary arrangements with the
office of Foreign Plant Quarantines in Washington a permit was
procured for the introduction of this culture and it was accord-
ingly sent and received in Trenton November 27, 1939.

According to the individual who supplied this culture this
strain of Beauveria hassiana is an isolate from cultures made
from infected Colorado potato beetle larvae, discovered in a potato
field at Fredericton, New Brunswick, Canada. Its pathogenicity
on the larvae of the Colorado potato beetle has been established.

During December, 1939, and January, 1940, a number of tests
were conducted at the White Horse Laboratory of this Depart-
ment to determine whether or not the Canadian strain of Beau-
veria bassiana was capable of infecting third instar larvae of the
Japanese beetle {Popillia japonica). Spores of the fungus were
applied dry to some beetle larvae, while aqueous spore suspensions
were applied to other groups of larvae. A number of beetle larvae
were also confined in tins with soil heavily inoculated with the
Beauveria spores. The rate of incidence of the fungous disease
was very low in these tests, and it was concluded that the fungus
probably had little potentiality as a control measure for larvae of
the Japanese beetle.

In July and August, 1940, experimentation was resumed using
the adult Japanese beetle, and exposing these to infection by the
fungous spores. When the beetles were dusted with the spores
a very high rate of infection was obtained, often 100 per cent.
Beetles allowed to feed on plant leaves which had been sprayed
with a dilute aqueous suspension of the spores were also attacked
by the fungus, the rate of infection approximating 75 per cent in
a number of tests.

Initially healthy beetles, kept in close association with beetles
previously exposed to infection, were found to contract the dis-
ease, the rate of incidence varying from 20 to 70 per cent in the
initially unexposed beetles.

In a test performed outdoors by spraying food plants of the

Dec., 1940]

Eex: Beauveria

403

beetle with an aqueous spore suspensiooi an indeterminate rate of
infection occurred, a number of dead, typically attacked beetles
being later found under the sprayed plants. Some beetles re-
moved from the sprayed plants after having been observed to
feed, subsequently died of the fungus.

Other beetles, first dusted with the fungous spores, and then
placed in outside screened cages began dying in four days, and
mortality was practically complete within ten days. Healthy
beetles were then introduced into these cages, and later, large
numbers of these died, exhibiting the typical external growth of
Beauveria.

The spores of Beauveria hassiana may be produced cheaply and
in large quantities upon a variety of culture media. The pre-
liminary tests of infection in adult beetles indicate that the fun-
gus may be a promising new natural control agency for the Japa-
nese beetle. — Edgar G. Rex, New Jersey Department of Agricul-
ture, Trenton, N. J.

BOOK NOTICE

Insect Transmission of Plant Diseases. By Julian Gilbert Leach,
Ph.D., Professor of Plant Pathology and Head of the De-
partment of Plant Pathology and Bacteriology, West Vir-
ginia University; Formerly Professor of Plant Pathology,
University of Minnesota. First edition. 8vo., cloth, 615 pp.,
238 illus., McGraw-Hill Publications in the Agricultural
Sciences, N. Y., McGraw-Hill Book Co., 1940. $6.00.

That certain insects have an important part in the spread and
development of many plant diseases has become more and more
keenly realized during the past two or three decades as constantly
increasing evidence has appeared in the literature of plant pathol-
ogy. Since much of this evidence has been recorded only in
widely scattered publications, and has never hitherto been ade-
quately summarized or coordinated, it would appear that there
is a definite need for this book. Particularly is this true, since
the author has endeavored to bring together in one publication
all of the more important contributions in this field of study, and
has made special effort to evaluate and interpret the evidence in
the light of the more recent advances in entomology and plant
pathology. In presenting the material, an effort has been made

404

Journal New York Entomological Society [Vol. XLVIII

to be brief. No attempt has been made to discuss all diseases
transmitted by insects or all insects that transmit diseases. Em-
phasis has been placed on those which have been most thoroughly
investigated or which may be best suited to illustrate the various
principles of insect transmission. Then too, in discussion of a
disease only those facts relevant to insect transmission have been
included and much information of general interest in plant pa-
thology has been purposely omitted. In numerous cases, how-
ever, suitable references to such information have been given.
In like manner, no complete technical descriptions of the insect
vectors have been included since information of this kind can
be readily obtained from standard entomological texts or from
special references cited. In addition to very full and complete
introductory matter, the various chapter subdivisions of the book
include discussion of such subjects as the interrelationships of
plants and insects; symbiosis between insects and microorgan-
isms and its significance in plant pathology; the relation of
insects to the spread and development of plant diseases; plant
diseases caused by toxicogenic insects; insects and bacterial dis-
eases; insects and fungus diseases; insects and virus diseases;
insects and phytopathogenic Protozoa; mites, nematodes, and
other small animals as vectors of plant diseases; the anatomy
and physiology of plants in relation to infection and insect
vectors ; the anatomy and physiology of insects in relation to the
transmission of plant diseases ; the Inocula of plant pathogens in
relation to insect transmission; the feeding and breeding habits
of insects in relation to the transmission of plant diseases ; insect
transmission of animal diseases compared with insect transmis-
sion of plant diseases ; and, methods useful in the study of insect
transmission of plant diseases. No general bibliography has been
given, but following each chapter there appears a list of refer-
ences, at times of considerable length, pertaining to its subject
matter. All people concerned with plant life, its cultivation
and protection will find this book a useful source of informa-
tion.— J. S. W.

THE TYPE OF THE GENUS PYRRHOPYGE (LEPI-
DOPTERA— HESPERIIDiE)

By W. H. Evans

There has been a great deal of confusion regarding the type of
the genus Pyrrhopyge and the identity of certain species of the
genus described by Linnaeus and other authors. In order to
clear up the matter it is necessary to summarize the relevant
literature.

(1) Linnaeus (1758 : Syst. Nat. 10th edition) described on page
485 under the generic group Papilio Pleheius TlrMcola {=Hes-
periidce) :

(a) ‘‘Phidias. 164. P. P. alis rotundatis atris nitentibus
margine albis, ore anoque rubris. M.L.U. Pet Gaz t. 43.
f. 15. Habitat in Indiis. ” The description relates to a
black insect with white cilia, palpi and anus red. The
figure bears no relation to the description : it is of a moth
with a pale apex to the forewing and a broad pale margin
to the hindwing.

(b) “Bixce. 165. P. P. alis rotundatus fuscis, basi virenti-
bus posticis subtus fascia alba. Merian t. 44. Pet Gaz t.
32. f5. Habitat in America.” The description is of a
dark insect with a white band on the hindwing underside.
The Merian figure indicates such an insect, with a red
head, palpi and anus : the white band is basal and extends
to the end of the cell. The Pet. Gaz. figure depicts an
insect with large hyaline spots on the forewing: it does
not correspond with the description and represents the
insect known as Epargyreus tityrus Fab.

(c) On page 487. “Acastus 186. Papilio Barhams. Alis
rotundatus albis : maculis quinque transversis apicibusque
fuscis : subtus flavescentibus : In Indiis. ’ ’ The insect is
a pierid, which seems to have been overlooked in the
“Lepidopterorum Catalogus. ”

(2) Clerck (1759: leones) published the following figures of
insects :

(a) Plate 42, fig. 4. “Papilio hixee’^ on the plate, depicting

406

Journal New York Entomological Society [Vol. xlviii

a well known West African insect with a white band on
the hindwing underside, centrally situated, not basally as
in the Merian figure, quoted by Linnaeus.

(b) Plate 44, figs. 1. 2. Unnamed on the plate and called
‘^Papilio phidias^^ in the index. It depicts an insect with
a very narrow, broken, white band on the hindwing under-
side : with a red h^ad, palpi and anus.

(c) Plate 44, figs. 3. 4. Unnamed on the plate or in the index.
It depicts an insect with a red margin to the hindwing
underside and : it is correctly known as harcastus Sepp.

(3) Linnaeus (1764: Mus. Ludovicae Ulricas ) referred to

(a) page 334. No. 152. pliidias, quoting Clerck plate 44 figs
1. 2. 3. 4. He admits the Pet. Gaz., figure to be a Phalmna.
He specifies a white band on the hindwing underside,
which he says sometimes extends to the upperwing : he adds
that the female has a red edge (harcastus). It will be
seen that he had departed from his original description
and has included three other species under the name
phidias.

(b) page 335. No. 153. hixce, quoting Clerck plate 42 fig.
4 as well as Merian plate 44. He adds that the Pet. Gaz.
figure he previously quoted is of an insect from Carolina
with hyaline spots on the forewing and a white band on
the hindwing underside (tityrus).

(4) Linnaeus (1767 : Syst. Nat. 12th edition 1/2 : page 795 made
the following additions to the 10th edition.

^‘Phidias. 263. Mus. Lud. Ulr. 334. Clerck ic. t. 44. f. 1.
2. 3. 4. Bixce. 264. Mus. Lud. Ulr. 335. Clerck ic. t. 42.
f. 4.”

(5) Cramer (Pap. Exot.) described and figured the following:

(a) 1775. Vol. I. plate 41, C.D. Papilio P. U. acastus Nov:
Surinam. An insect with a yellow edge to the hindwing
underside. His name falls as a homonym of acastus Linn-
aeus, see (1) above and the correct name is venezuelce
Scudder.

(b) 1779. Vol. Ill, plate 199, C.D. Papilio P. U. hixce Linn:
Guiana. The white band on the hindwing underside ex-
tends to the upperwing and represents a different insect to

Dec., 1940]

Evans: Hesperiid^

407

that figured by Merian or figured by Clerck : it was named
fluminis by Butler in 1872.

(c) 1779. VoL III, plate 199, E. Papilio P. U. acastus Cra-
mer : Surinam. An insect with a red edge to the hind-
wing underside: it is harcastus Sepp.

(d) 1779. Vol. Ill, plate 244 A.B. Papilio P. U. pkidias
Linn: China, Bengal and Surinam. It is not any of the
insects called phidias by Linnaeus, but the very different
Pyrrhochalcia iphis Drury.

(6) Fabricius (1793 : Ent. Syst. 3 : pages 344 to 347) includes :

(a) hixce Lin: considered to be a variety of tityrus Fab.

(b) zeleucus Nov: black with white cilia, head and anus red:
“Indiis”; figured in Jones icones (unpublished), vol. 6,
plate 25, fig. 2. It is the true phidias Lin., which he mis-
identified.

(c) mcenas Fab. 1787 : black with white cilia, palpi and anus
red, a white band on the hindwing underside : in America :
Mus. Dr. Hunter. It is the true hixce Lin., which he
misidentified.

(d) phidias Lin: placed as being synonymous with acastus
Cram.

(7) Donovan (1800: Insects of India etc.) figured zeleucus
Fab. on plate 51, fig. 3, corresponding with Fabricius’ descrip-
tion, excepting that the legs are dark red.

(8) Hiibner (1819: Verzeichniss : 103) introduced the genus
Pyrrhopyge for :

1077. hixce Linn : Syst. Pap. 264. Cramer 199 C. D.

1078. hyperici Hiibner: not described till 1823 (Zutrage 2:

16).

1079. phidias Linn: Syst. Pap. 263. acastus Cramer 41 C, D

and 199, E.

1080. amyclas Cramer: 199, E. 1081. arinas Cramer: 100, D.

(9) Swainson (1820: Zool. 111. 1/1: plate 33) figures zeleucus
Fab. assigning a new generic name Tamyrias: the figure cor-
responds with Fabricius’ description and the figure in Jones
Icones, with black and not the red legs of Donovan’s figure.

(10) Latreille (1824: Enc. Meth. 9: 732-740) includes under
Hesperia:

408

Journal New York Entomological Society [Vol. XLVIII

zeleucus Fab., with thasns Cramer (quite a distinct species)
as a synonym.

phidias Lin. : with acastus Cramer as a synonym.

Mxcb Lin. : with the white band on the hindwing underside as
basal and not central as in Clerck’s figure.

acastus Fab : with apastus Cramer and various other quite
distinct species as synonyms.

(11) Doubleday and Westwood (1852: Diurn Lep. Hi 609
and 515) included:

Pyrrhopyga thasus Cram. = zeleucus Fab.

mcenas Fab. = hixm Cram.
phidias Lin. = acastus Cram. 199 and 41.

Ismene chalyhe Nov. = hixce Lin.?, but not the hixce of Merian
plate 44.

(12) Wallengren (1858: Kon. Vet. Akad. Forh. 15: 2) intro-
duced the generic name Pachyrhopala for phidias Lin.

(13) Felder (1867: Keise Novara Zool. 2) uses the generic
name Tamyris and not Pyrrhopyge.

(14) Herrich Schaffer (1869: Corr.-Blatt. Regensburg 23:
164-166) in his Prodromus included under Pyrrhopyga:

zeleucus Swainson (unmarked on hindwing underside).

hixce Cramer (white band on hindwing underside).

acastus Cramer (yellow edge on hindwing underside) : 41 C.

phidias Lin. (red edge on hindwing underside).

mcenas Fab. : unknown to him.

(15) Scudder (1872: Fourth A. R. Peabody Acad. Sci. : 167)
listed the genus thus :

Pyrrhopyga Hubner.

Type Papilio hixce Linnaeus.

1. affinis HS. 2. venezuelce Nov. 3. chalyhea Nov.

(16) Butler (1872: Cist. Ent. 1: 176) realizing that hixce
Cramer was a different species from hixce Lin., renamed the
former fluminis.

(17) Scudder (1875: Proc. Amer. Acad. Arts & Sci., Boston,
10: 261) substituted hyperici Hubner as the type of Pyrrhopyge,
on the ground that phidias (hixce) had already been taken as
the type of Pachyrhopala Wallengren, which genus was a
synonym of Tamyris Swainson, type zeleucus Fab.

Dec., 1940]

Evans: Hesperiid^

409

(18) Mabille (1878: Ent. Belg. 21: 13). Gen. Insectorum
1903. Pyrrhopygince revision with Boullet 1908 : An. Sci. Nat.
9/7) came to the following conclnsions:

(a) Pyrrhopyge hixcB Cram nec Linnaeus.

(b) Pyrrhopyge phidias Lin.: Clerck’s plate 44, figs. 1. 2.:
= mcenas Fab.

(c) Pyrrhopyge zeleucus Fab.: on Donovan’s figure assigned
a red pectus, an entirely different species to the true
zeleucus.

(d) Mysoria Venezuela Scudder, acastus Cramer and harcastus
Sepp. {= phidias Fab.) as three separate species.

(e) Rhopalocampta hixce Lin. from W. Africa.

(19) Plotz (1879: Stett. Ent. Zeit. 40: 179, 533, 535. 1884:

Id 45: 65) dealt with the various species as follows:

(a) Pyrrhopyge phidias Lin. = mcenas Fab. and hixce HS.

(b) Pyrrhopyge fluminis Butler = hixm Cramer.

(c) Pyrrhopyge zeleucus Fab. : as described by Fab.

(d) Pyrrhopyge harcastus Sepp. = phidias Clerck plate 44,
figs. 3. 4.

(e) Pyrrhopyge acastus Cramer 41 C, B- phidias Lin. 1764.

(f) Ismene hixce Lin.: W. Africa.

(20) Aurivillius (1882: Kong Svenska Vet. — Ak. Handl.
19/5: 61, 121, 123) in his analysis of the species described by
Linnaeus.

(a) acastus Lin. is probably a Pierid.

(b) phidias Lin. is the species figured by Clerck on plate 44,

fig. 1. 2.

(c) hixce Lin. is to be taken as figured by Clerck on plate 42,
fig. 4 and Merian’s figure refers to phidias.

(21) Watson (1893: PZS : 11) follows Scudder (1875 not
1872) in taking hyperici Hiibner as the type of Pyrrhopyge.
He places hixce Lin. in the genus Rhopalocampta Wallengren
and erects the genus Mysoria with acastus Cramer as the type.

(22) Godman and Salvin (1893: Biol. Centr. Amer. Rhop. 2:
246, 247) considered hyperici to be the type of Pyrrhopyge; they
follow Aurivillius regarding phidias Lin. and Mabille regarding
zeleucus Fab.

(23) Seitz (1921: Macrolepidoptera) follows Aurivillius re-
garding phidias and hixce Lin. ; Mabille regarding zeleucus Fab.,

410

Journal New York Entomological Society [Vol. xlviii

gives hixce Cramer {-mcenas Fab.) as a separate species and
treats acastus Cramer and venezuelm Scndder as separate sub-
species.

(24) Lindsey (1925: Ann. Ent. Soc. Amer. 18: 99) asserts
that Sciidder’s first type selection must be regarded as valid,
i.e., tixcB Lin. : he adds that Mx(b and hyperici are congeneric.

(25) Bell (1931: Jour. N. Y. Ent. Soc. 39: 420, et seq. 1933:
Id 41: 268) deals with the genera and species as follows:

(a) Pyrrhopyge type hyperici Hiibner.

(b) zeleucns Fab.: cannot recognise.

(c) phidias hin. - mcenas Fab.: with a narrower white band
on hindwing underside.

(d) fluminis Butler = Cramer, nec Linnaeus, which is an
African species.

(e) Mysoria pallens M.ah. = acastios Cramer a homonym of
acastus Lin : pallens is a subspecies of venezuelce Scud.

In accordance with the International Rules for Zoological
nomenclature hixce Lin. must be taken as the type of Pyrrhopyge.
In any case hyperici Hiibner cannot be used, as that species
had not been described at the time that Pyrrhopyge was in-
troduced. If Aurivillius’ determination of the identity of hixce
Lin. is correct, it follows then that the generic name Pyrrhopyge
must be used in replacement of the name Coeliades Hiibner
{- Rhopalocampta Wallengren). Tamyris Swainson type zele-
ucus Fab. would have to be used for the American species
usually included under Pyrrhopyge.

The identity of any scientific name, such as hixce, must, how-
ever, be considered in respect of the original description: quali-
fications subsequently published can only be regarded as an aid
to the elucidation of the author’s original intention and must
be rejected if they conflict therewith. The description given by
Linnaeus in 1758 agrees with the Merian figure and not with the
Pettifer figure he quotes: so the Merian figure must be taken
as representing the type of hixce Lin. The Clerck figure rep-
resents an entirely different insect from W. Africa, which
Linnaeus in 1764 quite incorrectly considered to be conspecific
with the insect figured by Merian. Aurivillius’ determination
must therefore be rejected.

Bell (see No. 25 above) in his comprehensive revision of the

Dkc., 1940]

Evans: Hesperiid^

411

genus Pyrrhopyge brought to light that there was more than
one species exactly resembling hixce Linn., as here determined,
but differing markedly in the genitalia. It is considered that
the name should be applied to the species, which is in the B.M.
from Surinam, the type locality, with genitalia as figured by
Bell for Pyrrhopyge latifasciata Butler (1931: Jour. N. Y. Ent.
Soc. 39: 485). Actually latifasciata is a form with a wider
white band on the hindwing underside : in hixoe the band extends
to the end of the cell but not beyond. Mcenas Fab. is consid-
ered to be a synonym of hixce Lin.

A new name is required for Mxce Clerck nec Lin. and Coeliades
bixana Nov. is hereby assigned : it is sufficiently defined by Evans
(1937: Cat. Afric. Hesp. : 11) and a male specimen in the
British Museum from the Gold Coast has been marked as the
holotype.

The identity of phidias Lin. must also be considered in respect
of the original description. The Pettifer figure must be re-
jected as not agreeing with the description. The Clerck figures
subsequently quoted by Linnaeus must also be rejected : the
original description makes no mention of either the white basal
band of phidias Clerck or of the yellow edge of the second set
of figures. The figure given by Cramer does not agree with
Linnaeus’ description. The description of zeleucus Fab. and
the figure thereof in Jones’ leones does agree with the original
description of phidias Lin. It is considered therefore that
zeleucus Fab. should be regarded as a synonym of phidias Lin.
Here again Bell has shown that there is more than one species
exactly similar to phidias Lin., as here determined, but differing
markedly in the genitalia. It is considered that the name should
be applied to the species described by Bell as Pyrrhopyge wil-
liamsi (1931: Jour. N. Y. Ent. Soc. 39: 430). As pointed out
above (No. 18) Mabille was incorrect in his determination of
zeleucus Fab.

Bell (see No. 25 above) is quite correct in his action regard-
ing acastus Cramer : the name must be abandoned as a homonym.
It is considered that the oldest name for the collective species is
Mysoria harcastus Sepp. of which harcastus Sepp (= acastus
Cramer 1779 and verbena Butler), pallens Mabille {-acastus
Cramer 1775) and venezuelce Scudder are forms or subspecies.

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A NEW BRENTHIS FROM ALASKA (LEPIDOPTERA,
NYMPHALID^)

By Alexander B. Klots
College of the City of New York

Breuthis (Clossiana) aphirape denali, new subspecies.

While the writer feels a certain hesitation at adding another
snbspecific name in aphirape to the already rather long list, the
form named here is so utterly distinct from any other members
of the species that there is no doubt of its validity. From other
subspecies of aphirape in North America, denali may be differen-
tiated as follows :

Male — Upper side: Ground-color pale yellow-brown, the black markings
fine and clear-cut. The ground-color is lighter than that of any other apM-
rape subspecies except laddi Klots (Snowy Eange, Wyo.) and caelestis Hem-
ming (Colorado). In the fineness and sharpness of the black markings
denali likewise resembles laddi and caelestis most closely, for the other North
American subspecies all have heavier markings with considerable suffusion,
as well as a deeper orange-brown or red-brown ground-color. In the basal
and anal regions of the hind wings denali has slightly more fuscous dusting,
and is more hairy, than laddi and caelestis.

Female — Upper side: The ground-color is somewhat duller and the black
markings heavier and more suffused than in the males. Compared with
females of other North American subspecies, denali females average paler
with lighter markings and less fuscous suffusion than those of any subspecies
other than laddi and caelestis; their marking are slightly more suffused, and
the basal dusting of the hind wings more extensive, than in these two latter.

Both sexes — ^Under side: The fuscous or black markings, repeated from
the pattern of the upper side, are slightly heavier in the females than in the
males. In other respects the females show no essential differences from the
males. In both sexes the fuscous or black markings are more reduced than
in any other North American subspecies, including laddi and caelestis.

The ground color is much paler than in any other North American sub-
species, having a very “washed out“ appearance. On the hind wings the
post-basal and median rows of darker spots are of a light, yellow-brown color,
much lighter than in any other North American subspecies except caelestis.
The submarginal area which, in all other North American subspecies is largely
filled in with dark orange-brown or red-brown in cells Ei, Eg, Mi, Cui, and Cin,
at least outwardly, is in denali almost entirely of the very light, yellowish
ground-color with only a slight brownish tinge in the costo-distal corner of
cell El. The row of round, ocellate spots in this submarginal area is very
much obscured, being almost entirely faded out in some specimens. The
marginal and the irregular sub-median rows of light-colored spots are of a
pale, yellowish-white with only an occasional trace of pearly luster.

414

JouKNAL New York Entomological Society [Vol. XLVIII

Denali is almost the most distinct subspecies of aphirape in
North America, the extremely pale, “washed out” appearance
of the under side of the wings sufficing for its recognition at a
glance. Of the 43 specimens of the type lot, only two show suffi-
cient dark coloring beneath to be in any way confnsable with any
other North American subspecies; and the resemblance of these
is to caelestis, the Colorado subspecies, rather than to nichollm or
triclaris, the closest subspecies geographically.

Two specimens from Atlin, B. C., are intermediate between
denali and nichollm. They are not to be considered as representa-
tive denali, or as part of the type lot.

Leiissler’s reference (Bull. Brooklyn Ent. Soc., 1935, 30: 52)
to a male and a female of aphirape from the Mackenzie Delta
under the name of alticola Barnes & McDunnongh (a homonym,
= caelestis Hemming) is inadmissible, as pointed out by the pres-
ent writer (Journ. N. Y. Ent. Soc., 1937, 45: 325). The pale
coloration which led Lenssler to use the name alticola for these
specimens is here seen to be characteristic of denali. It is there-
fore quite probable that the Mackenzie Delta specimens should
be referred to denali or, at least, that they represent a transitional
population between denali and triclaris.

The name of this new subspecies is the Indian name for Mount
McKinley.

Types : Holotype J', July 25, 1931, and allotype 5? J^Jy, 1931,
McKinley National Park, coll, by Frank Morand, ex Guilder
Collection.

Paratypes : 21 the same locality and collector, July, 1931 ;
1 the same locality and collector, 24 July, 1931 ; 1 the same
locality and collector, 25 July, 1931 ; 1 the same locality and col-
lector, 28 July, 1931 ; 4 the same locality and collector, July,
1931 ; 1 2, the same locality and collector, 24 July, 1931 ; 3
and 1 2, the same locality, coll, by G. P. Engelhart, 3500 ft. alti-
tude, 5 July, 1938; 1 J' and 1 $, the same locality, coll, by D.
Fraser, 29 July, 1930; 1 Cantwell (S. of Fairbanks) Alaska,
coll, by D. G. Nicholls, 14 July, 1937 ; 3 ,c?c? and 1 ?, Alfred Creek
Camp, Alaska, coll, by R. A. Pope, 15 July, 1922 ; 1 Alfred
Creek Camp, Alaska, coll, by R. A. Pope, 21 July, 1922.

The holotype, the allotype and most of the paratypes are in the
American Museum of Natural History. Paratypes are in the
collections of Cyril dos Passos and the author.

INDEX TO NAMES OF INSECTS AND PLANTS IN
VOLUME XLVIII

Generic names begin with capital letters. New genera, subgenera, species,
subspecies, and new names are printed in italics.

Achsetoneura
samiae, 261
Aconopliora

hrunnea, 281
erecta, 280
Alchisme

laticornis, 277
pinguicornis, 277
spinosa, 278
Amastris

peruviana, 286

Ants, relation to Japanese beetle
and parasites, 85
Apteroloma

tenuicorne, 233
Arotes

amoenus, 155

Butterflies, list of New Jersey, 47
Beauveria

bassiana, 401
Benthis

aphirape

denali, 413

Cacama

crepitans, 117
CalliphoridaB, 395
Carterocephalus
palaemon

mesapano, 244

Ceresa

grisescens, 283
luteimaculata, 282
Chiomara

mithrax, 116
salma, 116
Clidophelps

pallida, 118

Ephemera

guttulata, 384
vulgata, 382
Ephoron

album, 384
Erythrandra
picipes, 140
Eumichthini, 368
Euphoron

ladogensis, 385

Gonomyia

macswaini, 115
remota

ohtusistyla, 115

Heliothis

obsoleta, 305
Helius

distinervis, 112
Hyboderini, 371
Hypopelta

scrofa, 143

Insect losses, 195

Insects, of Lenina minor, 320

Ischnothrix

patens, 113

Japanese beetle

relation of ants to, 85
parasites, relation of ants to, 85
Johnsonia

borealis, 140

Laccoprosopa

sarcophagina, 138
Lemna minor, insects of, 320
Leucophora

albiseta, 342

415

416

Journal New York Entomological Society [Vol. xlviii

annexa, 355
fusca, 353
johnsoni, 343
maculata, 347
marylandica, 344
obtusa, 350
sociata, 346
unilineata, 348, 352
Limonia

paucilohata, 111
Lipsothrix

neotropica, 114
Lycoderes

luteus, 275

Macronichia

aurata, 136
confundens, 136
Metoposarcophaga
importuna, 144
Micrutalis

nigromarginata, 289
Myllocerus

castaneus, 318

Neoplatypedia

constricta, 125
Neotipula, 105
Nicropliorus

guttulus, 235
investigator, 235
vespilloides, 234
Nisoniades

selma, 116

Obriini, 367
Okanagana
bella, 121
calif ornica, 122
fumipennis, 124
luteobasalis, 122
rubrovenosa, 120
schafferi, 124
striatipes, 123
synodica, 124
utahensis, 123
vanduzeei, 121

Okanagodes

gracilis, 119
Opelousia

obscura, 137
Oppiopsis

sheldoni, 139
Opsodexia

abdominalis, 138
bicolor, 137
Orimarga

melampodia, 112

Palingenia

longicauda, 379
Paragargara
nigra, 279
Platanus

occidentalis, 384
Platypedia

mohavensis, 125
putnami

lutea, 125
Pletliogenesia

papuana, 384
Polistes

canadensis

mexicanus, 10
satan, 11
varieties, 7
weheri, 12
wheeleri, 12
exclamans

louisianus, 14
varieties, 13
fuscatus, 391

connectens, 21
hunteri, 23
maritimus, 25
montanus, 26
neotropicus, 28
varieties, 17
major

haJceri, 15
varieties, 15

Popillia

japonica, 85, 401

DEC., 1940]

Index

417

Proboscimyia
irevis, 357
siphonina, 356
Pseudogaurax
anchora, 260
Pseudopilema, 373
Pyrrhopyge, 405

Ehopalocera of New Jersey, 47

Sarcofahrtia

ravinia, 141
Sarcophaga

aculeata, 224
alcedo, 224
aldrichi, 224
atlanis, 225
bisetosa, 218
cimbicis, 219
laakei, 221
latisetosa, 221
latisterna, 219
1 ’herminieri, 221
misera

exuberans, 216
liarpax, 216
sarracenioides, 217
pusiola, 222
stimulans, 222
sueta, 223
Silpha

bituberosa, 233

opaca, 233
trituberculata, 233
Siphloplecton
basale, 382
Sitophilus

granarius, 37
oryzae, 37
Spilocryptus

extrematus, 262
Stictolobus

nitidus, 284
marginatus, 285
Stylocentrus

rubrinigris, 276

Thrasymedes

virescens, 288
Tibicinoides

cupreosparsus, 125
mercedita, 125
Tipula

browniana, 107
pectinella, 106
scriptella, 109
ctcnopyga, 110

Trox

unistriatus, 303
Tynelia

nigra, 287

Wohlfahrtia
vigil, 142

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