y ii! wy Sena
roe |

ol

LE
oN

; ‘y x 'INetOR | i
@) ake oe eS aise ;
prey, ih: Le ak Rica ” wy i & ll
i 4s : yoy “oe ‘ 4, i :
“iy, Mel Tey Mal
eg A " ae Yi 5 HE _

aaa
fe)
a

<y : SS
pnp e

“ (@) iE
S HH, Sy j
be

RK all
| i:

ob, Nets
, *
&

Oy >

Ss, NG
ss 3
he | Hii fle qt
WN. 7,
nal

i

“p

x
ibn l

4p \ | ye
We

eu %
Se

mee %,

P 3
ie)
Hs oy
\"
iit

4
4

s, “ fay E = “> 4 v Hi i"
ane % 3 Th

ae

WwW

Al
wet !
0, al a
Ze it
: “a am | i | x | Ye) 7

a“? py,
ph a
, ic a
I

“>

TY.

0s ~ ’

- 47

ta Hi Ln Le Vii x un
yar ae

; Ai ARS)
Bret a vA
Lit) ae :
LP a) e: i] |
Aba 4
ry ee

vibe ¥

(cage he

i Me oy nay
| : ve fel
} TR ee
: ue Weer ih

Again 6s vy Pew hy iy alts Hae ae ti jap a

or Bay, 1 nin A Cn nate aes hon Bae RP an
se av a Te 4 0 Sean eat - Kony
A A a a ¥ oy ;
i ee) epee : , a vie 7 ia |

io iis e Aer ay hat at vd ? Wy ) LP

ee MAI: Yi 9 ahh, Hienie, hiatt

f ets 4 wt) Meret Went: fs : AMY ea a Li a te vy

Hina Le Cae Hh es on Ran lag Aha
any: , aah en: Aish,” phen em eat

Bn tevin | he

fi ed Viava | . ace ee

A he Rh f
nn © f
hy} }
7?
y an i) { Vie
aro | rf an
ay tae e f
7 rl a : eyk i i

~
VY \ ~~ 54°
Q \_\ a a
v lo’,
us ae
{,_// f &
CA

ANNALS

OF

The Entomological Society of America

V¢
VOLUME Xi. 19:19
Se v
f ;
, JAN 26 19
Xare
EDITORIAL BOARD Se
HERBERT OSBORN, Managing Editor,
CoLUMBUS, OHIO.
lL. O. HOWARD, WM. A. RILEY,
WASHINGTON, D. C. MINNEAPOLIS, MINN.
VERNON L. KELLOGG, FRANK E. LUTZ,
STANFORD UNIVERSITY, CALIF. NEw YORK CIty, N. Y.
WM. M. WHEELER, WM. S. MARSHALL,
Boston, MAss. MADISON, WIS.
E. M. WALKER J. G. SANDERS,
TORONTO, CANADA. HARRISBURG, PA.

PUBLISHED QUARTERLY BY THE SOCIETY
COLUMBUS, OHIO

CONTENTS OF VOLUME XII.

PAGE
Van Dyke, Epwin C.—The Distribution of Insects in Western North America.. 1

Brues, C. T.—A New Chalcid-fly Parasitic on the Australian Bull-dog Ant.. 13
ALEXANDER, C. P.—Notes on the Crane-flies of the Hawaiian Islands

(CTE reel ri 2x 28°] D Wyo RN one ee Pa Nr CR ee 25
PAEL, Hi. C©—The Californian Species of Malthodes......................--- 31
HERRICK, GLENN W. and DETWEILER, JOHN D.—Notes on the Repugnatorial

(Clandsyotm Certain \oLtodonideCatenpillans se. oem cicraiee eee eer 44
MosuHer, Epna—Notes on the Immature Stages of the Deltometopus rufipes

Melss(Coleopterayeucnemidae) nw. scr onde eee ce ee se eres 49
ALpricH, J. M.—Proceedings of the Baltimore Meeting...................... 57
Dietz, W. G.—The Streptocera Group of the Dipterous Genus Tipula,

ETT) COIS ee res ory od xylan MMe MRCS oe aha Sebayele ap ataicttets wohl hears 85
CLAASEN, P. W.—Life-History and Biological Notes on Chlaenius Impuncti-

HEROS, SDs mere Oo tReet eee ihe a ences <a ROR aera a ea rt 95
PLLA oh obbenties of the’Genus Prax... .......-0---s60seee eevee ne 103
(SAHAN ‘Some Chalcid Wasps Reared from Cecidomyid Galls.......... 159
Milman ene ropress of Scolia manilae Ashm.........-....:22+--25--++-5: 171
WHEELER, WitttAM M.—The Ants of the Genus Metapone.................. sie
Rucxes, eraeec-—Notes on the Male Genital System in Certain Lepidoptera. .192
WeEtcH, Paut S.—The Aquatic Adaptations of Pyrausta penitalis Grt.

SE SESISIERLEIP ED) [ino Aree eo 213
FRACKER, S. B.—Chariesterus and its Neotropical Relatives (Coreidae

EXCECEOD LCL) Merete ean errr rye, eee ricky Soe Se ae cen ae each oe 227
SANDERS, J. G. and DELonc, D. M.—Eight New Jassids from the Eastern

WiraiGeRS GAC SMe pewter ase ee tent crit go. cote areal oe ee ate Seas Prey Sas beets apes Sd ats 231
CHAMBERLIN, RALPH V.—New Western Spiders................2202020 eee sees 239
Wacker, E. M.—The Terminal Abdominal Structures of Orthopteroid

Insects arAGe My OGeNeDICHS TUG 4.6 seu mito saree ecieci elem eiclardy oeesveetia ec y- 267
ALEXANDER, C. P.—Undescribed Species of Japanese Crane-flies.............. 327

Braun, ANNETTE F.—Wing Structure of Lepidoptera and the Phylogenetic

and Taxonomic Value of Certain Persistent Trichopterous Characters... .349
Dickerson, E. L. and Weiss, Harry B.—The Life-History and Early Stages

of Platymetopius hyalinua Osb., a Japanese Leaf-hopper in New Jersey. . .369
PiatH, O. E.—The Prevalence of Phormia azurea Fallen (Larva Parasitic‘on

Nestling Birds) in the Puget Sound Region and Data on Two Unde-

Sctibedunies Ob Cimilar biabita a me mane. owe: ce ee eee e erence se ale
TOWNSEND, C. H. T.—Description of the New Species of Phormia.............379
ALpRICH, J. M.—Description of a New Species of Hylemyia.. ae, .. .080

Kino, J. L.—Notes on the Biology of the Carabid Ae Petiends
Se plemib a ane Me MACKIUS. cori ieee meiciis sg se ace whee sess es Hoe Mew om 382

- December, 1919, number mailed December 31, 1919.

-*
¢ Ca
7. es ay ?
q - -
¢ . Pa é bt i
,. 7 *%
7 ia
f os ;
:
} 4 ey i.
2 =. .
v- : ' 7
“' g! }
ae 7 hee 4
.
7 i
-
wweltieas ¥

DATES OF ISSUE. | “IgE

March, 1919, number mailed ‘April 26, 1919.
June, 1919, number mailed July 2, 1919. sd
September, 1919, number mailed October 21, 1919.

ay

“VERNON L. KELLOGG,

ANNALS

OF

MARCH, 1919

AS eye EDITORIAL BOARD
‘| HERBERT OSBORN, Managing Editor,
COLUMBUS, OHIO,

LL: oO. HOWARD, WM. A. RILEY,

WASHINGTON, Dy: C

STANFORD UNIVERSITY, CALIF.

WM. M. WHEELER,
Boston, Mass.

E, M. WALKER,
+; TORONTO, CANADA.

PUBLISHED QUARTERLY BY THE SOCIETY

fey COLUMBUS, OHIO

FRANK E. LUTZ,
NEw YorxK City, N. -Y.

WM. S.,. MARSHALL,
MaAnDIson, WIs.

J. G. SANDERS,
‘HARRISBURG, Pa.

Number 1.

. The Entomological Society of America

+

MINNEAPOLIS, MINN.

Entered as second class matter April 11, 1908, at the Post Office at Columbus, Ohio,

> under the Act_ of Congress of March 3, 1879,

The Entomological Society of America
Founded 1906.

OFFICERS 1919.

%

President
James G. NEEDHAM. f : ‘ : 7) Sbthaca, IN SYs

First Vice-President
Panes WGPOLsOM er oe (GM : ‘ : . . Urbana, Tl.

Second Vice-President
R..V. CHAMBERLIN. ; ; : . . Cambridge, Mass.
Managing Editor Annals
HERBERT OSBORN : i : a s Columbus, Ohio
Secretary-Treasurer
‘J. M. AtpricH . : . National Museum, Washington, D. C.

Executive Commitiee

THE OFFICERS

AND
We Bs wn ARTHUR GIBSON,

F.E
G.'A. DEAN, G. C. CRAMPTON.

Committee on Nomenclature

Pe Pel LET, Si DIME BAT COCKERELL, NATHAN BANKS.

Thomas Say Foundation

NATHAN BANKS, A. D: MacGIrLivRay, P. P. CALVERT,
_ E. B. WILLIAMSON, —

J. M. Atpricu, Editor, E. D. Batt, Treasurer.

Committee on National Museum.

T. D. A. CocKERELL, Chairman,

HERBERT OsBorN, Ww. BARNES, Wn. M. WHEELER
James G. NEEDHAM.

ANNALS

OF

The Entomological Society of America

Volume XII MARCH, 1919 Number |

THE DISTRIBUTION OF INSECTS IN WESTERN
NORTH AMERICA.*

By Epwin C. VAN Dyke,
University of California, Berkeley, Cal.

The fauna of the western part of North America has long
been recognized as possessing many characteristics which have
differentiated it from that found elsewhere in North America.
This is particularly the case as regards the insects and Le Contet
as early as 1859 called attention to this. Other entomologists
in more recent years have mentioned this fact and have cited
many cases to show its peculiarity. The insect fauna as a
whole, however, has not been studied in regard to this point,
in the manner that it has merited.

Many years ago, I became interested in the subject and
I have kept up my interest ever since and have availed myself of
all opportunities that would enable me to gain information
bearing uponit. I have studied all orders of insects to a certain
extent, but most of my conclusions have been based upon a
close study of the Coleoptera, particularly of the wing ‘ss and
less mobile groups, such as certain of the Carabide, the Silphide,
the Tenebrionide, the Otiorychine, and so forth. The other
Coleoptera were used as checks as were in fact other groups of
insects, and all other information bearing upon the subject of
distribution in the territory under study, freely drawn upon
for purposes of guidance.

* A revision of the paper read August 5, 1915, at the University of California,
before the Summer Meeting of the Entomological Society of America,

t The Coleoptera of Kansas and Eastern New Mexico, by John L. LeConte,.
M. D., Smith. Miss. Contrib. to Knowledge, Vol. XI (1859),

2 Annals Entomological Society of America [Vol. XII,

As a result of these studies, I have come to believe that
though the West Coast fauna is to a degree complex, it is yet
made up of a definite number of elements which can be distinctly
separated. The insect fauna as we now know it, has been
derived from older faunas and these were either northern or
southern in their origin. Each of these primary portions was
in its turn composed of elements which had come into their
present territory at different times and along different roads.
Certain of these had remained pure and are at present restricted
to definite areas, while others had either wholly or partially
invaded regions already occupied by other elements and become
mixed with them. These facts are of course in general in

\'

7 Yancouveran

Bee Sierran

tee

reat Basin

a

1 a

i ———

way ise S)
daa (iS 2 SS

Mte. of Northeast ——

moetly Canadian, i ——=
l —

Fig. 1. Map showing the Faunal Areas of the Pacific States of North America.

1919} Distribution of Insects 3

keeping with those worked out by Merriam* and other mam-
malogists and ornithologists and substantiated by the botanists.
Certain of the detailed results secured have, however, been
found to be at variance with those found by others or to have
previously not been worked out.

In general the insects which are considered to be of northern
origin now occupy the northern and more mountainous portion
of the territory. The greater portion of this, such as that
including the lower levels of the Aleutian Islands, particularly
the eastern ones, the southern margin of the Alaska Peninsula,
southeastern Alaska, the western part of British Columbia
as from the Selkirks west, Washington and Oregon west of the
Cascade Mountains, and the wet and cool coastal strip of
California, extending as far south as middle Monterey County,
now supports a fauna which I have called the Vancouveran.
The name Transitional as applied to this fauna, though generally
used by previous workers, I consider misleading. The fauna
is a pure one, in fact one of the purest in North America. It
contains many peculiar groups of insects, some almost in their
entirety, and is especially rich in such families as the Carabide,
Staphylinide, Elateride and certain groups which are more or
less closely associated with the coniferous forests. The following
are good representatives: the genus Omus, the subgenus
Brennus of Scaphinotus, the impunctate division of Pterostichus,
Zacotus, Metrius, Promecognathus and the American representa-
tives of Trigonurus. Rosalia funebris Mots. is also a char-
acteristic member of this fauna, but it extends considerably
beyond its confines. The fauna has existed in approximately
its present territory from the Tertiary period and no doubt
from well back in that period, being in fact a Tertiary fauna
which has come down to us in a pure state. Many of the
species that it contains have probably not changed during this
long series of years but others have, some having become
broken up into races which, becoming more and more isolated,
have formed new species, subspecies, and so forth. Asa result,
the fauna is not the same throughout its extent. It gradually
divides itself up into several subfaunas.

* The Geographical Distribution of Life in North America with Special
Reference to Mammalia, by C. Hart Merriam, Proceed. Biol. Soc. Wash. Vol. VII,
April 18, 1892.

4 Annals Entomological Society of America [Vol. XII,

The first of these occupies the following regions: the coastal
belt of southeast Alaska and British Columbia with the adjacent
islands, the Puget Sound Basin of Washington and the Willa-
mette Valley of Oregon, and extends fully up to the five thousand
foot level on the Cascades. This contains the Vancouveran
fauna in its purest form and with its species but little modified
throughout its extent. Characteristic Coleoptera of this area
are Omus dejeani Reiche, Cychrus tuberculatus Harr., Carabus
taedatus Fab., in its typical form, Nebria mannerheimi Fisch.,
Pterostichus herculaneus Mann. and Pterostichus validus De}.,
and Miscodera insignis Mann.

Next we have a subfauna that might be called the Pacific
Maritime. This occupies that very wet region to the west of
the Olympic and Coast Ranges of Washington and with sec-
ondary modifications taking place chiefly in northwestern
Oregon, in southwestern Oregon and northwestern California,
in Mendocino County, California, and south of the Russian
River in California, extends down the coast as far as middle
Monterey County, California. Within this maritime area, the
fauna as far as the species are concerned, is quite similar to that
found in the pure Vancouveran, but the form and appearance
of many of the species has changed. Melanism is markedly
evident here and is to be noted in such widely separated families
as the Carabide, the Elateride, the Cantharide (Lampyride), the
Scarabaeide, and the Rhynchophora. Increase in size is also
to be noted with regard to many of the species. To illustrate
this, I will cite but a few examples. In the Carabidae, in the
genus Scaphinotus, we have the species S. angulatus Harr.
which is of a metallic purple color in those specimens dwelling
in the Puget Sound Basin and the Willamette Valley, and
absolutely black in those which are found in the wet coast
belt, though otherwise unmodified. In southwestern Oregon
and northwestern California, a related but distinctly different
species, SS. behrenst Roesck, is found replacing the above.
Scaphinotus angusticollis Fisch. in the pure Vancouveran region
from southeastern Alaska to the eastern side of the: Willamette
Valley, is typically of a ferruginous color. West of the Olympics
and Coast Range of Washington and the Willamette river in
Oregon, it is absolutely black. In Washington, it is structurally
the same as the typical form; in western Oregon, larger and with

1919] Distribution of Insects 5

the alternate elytral intervals more reduced or irregular; in
southwestern Oregon and northwestern California, it occurs
with a more angulate prothorax, still more modified elytra,
and with the male tarsi somewhat changed; while in the southern
part of its range, in Mendocino and northern Sonoma County,
California, it appears as a much larger insect and much modified
in every regard. In Ptlerostichus amethystinus Dej. we have a
species with bluish elytra which extends from Alaska to middle
Mendocino County, California. South of this, it is replaced
by P. scutellaris Lec., a species which has absolutely the same
habitat and only differs from the preceding by being all black.
It extends into Monterey County, California. Among the
Elateride, we have numbers of species which like Athous
ferruginosus Esch., have a black phase within the southern
maritime area. In the genus Silis, the species and varieties
found along the coast and in the high mountains of the Pacific ©
Coast are mostly black, whereas in the interior lowlands they are
yellow. Among the Rhynchophora, we have one, Rhynchites
bicolor Fab., one of the most widely distributed weevils in
North America which has its only known black phase in the
middle coastal belt of Oregon.

A second and more greatly modified portion of the Van-
couveran fauna is that which inhabits a strip of country that
starts in the west central part of Oregon, runs south and south-
east, including all of the moderately elevated mountains in
Southern Oregon and Northern California, and extends along
the western flanks of the southern Cascades and Sierras into
Southern California and with breaks, also into the San Pedro
Matir of Lower California. This subfauna I would call the
Sierran. It occupies much of the same country as that covered
by the western yellow pine, and though it receives many species
from those faunas found immediately above and below it, is in
no sense a transitional one as often called. It is a distinct
subfauna and a direct offshoot of the Vancouveran. Its species
are almost always either like those to be found in the pure
Vancouveran or are derivatives of the same. In the northern
area of the Sierran subfauna, certain species show a strong
tendency to break up into many weak races which are more or
less limited to various small areas. This unstableness is no
doubt partly due to climatic and partly to topographical
conditions, for the country is much broken up, there being many

6 Annals Entomological Society of America [Vol. XII,

small ranges with variously exposed slopes. In the region
extending from Mt. Shasta south to the American river, some
of the offshoots of the primitive species have segregated them-
selves into rather distinct species and subspecies, serving
thereby to define this as a rather distinct subregion and one
with a fairly well defined secondary fauna. In the territory
running from the American river south to the Merced river, we
find marked off in a similar way another secondary faunal
area; in the southern Sierras, quite another; while in the San
Bernardino Mts. there is still another. These secondary
faunas, though not well defined by the majority of the Sierran
species, are yet very distinctly outlined by others, as for
instance, by certain species of Omus, Scaphinotus, Pterostichus
and Pleocoma.

Two of the North American faunas most closely related to
the Vancouveran and two, which like it, are most likely rem-
nants of the same northern Tertiary fauna, are one small fauna
confined to parts of the mountains of western Idaho, the Coeur
d’Alene and Moscow; and the fauna of the upper levels of the
southern Appalachian mountains, the so-called Alleghanian.
These two are relict and endemic faunas and possess as would be
expected, many species or related species 1n common. In the
Idaho fauna, we have certain species like Pterostichus sphodrinus
Lec. which are closely related to those found in the Vancouveran
like Pterostichus ovicollis Schaef.; and others as Scaphinotus
relictus Horn and H. merkeli Horn that have their nearest
relatives in Scaphinotus imperfectus Horn, S. debilis Lec. and
S. incompletus Schz. of the southern Alleghanies. In Pteros-
tichus we have a peculiar and small group of species that have
more or less prognathous mandibles and are quite subterranean
in habit. On the Pacific Coast we have three, P. caligans Horn,
being the best known, one in Idaho, and P. grandiceps Ch. and
P. rostratus Neum. in the Appalachian Mountains. All of the
above mentioned species are so highly specialized that they
cannot be anything but relicts. They could not possibly have
migrated to their present abode subsequent to the Glacial
Period, though like the last mentioned, they might have
extended their territory.

In the Cascades and Sierras, at elevations immediately
above those occupied by the Vancouveran and its offshoot,
the Sierran, we have a fauna that is more alpine in its nature.

1919] Distribution of Insects Zh

The lower belt of that fauna, that found within the fir zone, is
generally spoken of as the Canadian, that immediately above
it and in the more barren treeless areas as the Hudsonian, and
that on the tops of the high peaks as the Arctic. These faunas
are in reality marginal or fringing faunas. They fringe the snow
fields of the mountains and the barren wastes of the north,
advancing and retreating with them, and have done so not
only during the present period, but undoubtedly during the
entire portion of the Great Ice Age, and I believe even before
that were only to be found in the cooler areas of the north.
During the period of greatest ice development, the Pleistocene,
they were. driven down to the lower levels and forced much
farther south than they were before. With the decline of this
period, they retreated both upwards and northward. At the
more southern limits of their distribution their retreat was often
prevented by breaks in the ranges or by the lowness of the
mountains so that their continuity was interrupted leaving as
we find today little islands of fauna here and there restricted
to the more elevated parts of the high mountains.

The first of these, the Canadian, is in the west almost entirely
a forest loving fauna. Where it comes in contact with the
Vancouveran and Sierran, it blends to some extent with them
and as a result is often hard to separate. It has also borrowed
many species from these as.they have from it. It commences
in the north as a continuation on to the mountains of that
extensive lowland fauna which populates the vast areas of
western Canada. Then it continues south along the mountains,
at first as a rather broad belt, later as a much narrower one,
and wedged in between the Vancouveran and Sierran, and the
Hudsonian on the west slopes; and the Great Basin fauna, a
portion of the so-called Upper Sonoran, and the Hudsonian
when the mountains are sufficiently high for that, on the east
side. Its forest types of Coleoptera are fairly characteristic,
such as many of its Elateride and Cerambycide, but its Carabide
are less so. Pterostichus protractus Lec. and Platynus bogemannt.
Gyll. are, however, quite distinctive of it in the more southern
Cascades and the Sierras.

The Hudsonian is in most places a very narrow zone, but
it is very distinct. In Alaska, it is found on the uplands of the
Aleutian Islands, on the mountains and the north side of the
Alaska Peninsula, to quite an extent about the Kenai Peninsula,

8 Annals Entomological Society of America {Vol. XII,

and thence south along the flanks of the ranges which fringe
the Coast. In British Columbia it is more inland and on the
higher mountains and it continues as such along the Cascades of
Washington and Oregon and the Sierras of California, but
gradually ascends as it extends southwards. In the Cascades,
as on Mt. Rainier, it is found at about seven thousand feet, on
Mt. Shasta at eight thousand, in the Lake Tahoe region near
nine thousand, while in the southern Sierras it is well above
ten thousand feet. A few of its more widely distributed and
characteristic beetles are Nebria sahlbergt Fisch., Bembidium
incertum Mots., Amara erratica Sturm, and Cryptohypnus
bicolor Esch. Besides these, there are certain others which
are only to be found in the mountains of the West but which,
because of certain peculiarities of distribution throw so much
light upon the problem that they are worthy of being especially
mentioned. Such is Nebria trifaria Lec. a large and very
attractive black species. This is undoubtedly a glacial relict,
for it is now only to be found high up near the snow fields on
the Olympic Range, Mt. Rainier and Mt. Jefferson of the
Cascades, and the Rockies of Colorado. The rare N. ingens
Horn of the high southern Sierras, which differs from the
preceding chiefly in having very much rounded humeri and
atrophied wings, is, I am convinced, but a degenerate offshoot
of the preceding which was forced south and later entirely
separated from the parent stock. Nebria ovipennis Lec. and
its three associated species are also of importance from the same
viewpoint. They are all of moderate size, entirely apterous,
and with elliptical elytra, hence extremely specialized and
dependent upon their particular environment. The most
northern, N. kincaidi Schw., an entirely metallic species of a
purplish copper color, has been found close to the coast at
Farragut Bay, Alaska, and near the snow fields on Glacier
Peak and Mt. Rainier, Washington. A second, N. columbiana
Casey, which resembles the preceding, but has only the elytra
metallic, was described from British Columbia, but has also
been found on Glacier Peak and Mt. Rainier, Washington, and
Mt. Jefferson, Oregon. The third, N. ovipennis Lec., which is
but an offshoot of the preceding and differs only in color, being
entirely brownish or piceous, in other words more heavily
pigmented, is to be found on the higher peaks of the Lake
Tahoe region in California and above 10,500 feet in the southern

1919} Distribution of Insects 9

Sierras. The fourth species, N. diversa Lec. is of a yellowish
color and is confined to the sea coasts of Washington and
Oregon. The only close relatives that these four have are
certain species found in the more eastern part of the Himalayas
and in northeastern Asia. The last beetle to be mentioned in
this connection shows another peculiarity of distribution due
to the retreating ice. This is Pterostichus brunneus De}j., a
species first described from Sitka on Baranoff Island, later
found on Orcus Island near the mouth of Puget Sound and no
doubt occurring on other islands in the same general region,
and on the mainland only high up near the timber line as in
the Selkirks of British Columbia, on Glacier Peak and Mt.
Rainier in Washington, and on Mt. Jefferson in Oregon. On
the islands, it was left stranded, but by adaptation was able
to persist, while on the mainland it could preserve its natural
environment by merely retreating with the ice to the higher
levels.

The faunas of southern origin are to be found in their purest
state only in the more southern part of our territory though
derivatives of the same do in certain regions extend quite far
to the northward. The best known of these is the Sonoran
which, when considered in its strictest sense, may be said to
occupy all those hot and more or less barren uplands in northern
Mexico and the semi deserts and drier regions of our own
Southwest, with extensions into western Texas, southern New
Mexico and Arizona, and the more desert parts of southeastern
California. The fauna of the Colorado Desert as well as that
of its more upland extension, the Mojave Desert, is typically
Sonoran. Certain elements of this also extend more west-
ward along our southern border to the coast at San Diego and
from the Mojave through the hot Walker Basin into the
southern San Joaquin Valley. Here it is to be found mainly
on the west side of the valley as in western Kern County and
in an attenuated form in southern Monterey County. Some
of its most characteristic beetles are among the wingless Tene-
brionide, such as in the genera Edrotes, Triorophus, Zopherus
. and Asida, and in the wingless Otiorychine like Ophrastes and
Eupagoderes, and the genus Monilema of the Cerambycide.

A derivative of the Sonoran fauna which is generally spoken
of as the upper Sonoran, though it is more accurately defined
as the Great Basin fauna, extends throughout the entire area

10 Annals Entomological Society of America [Vol. XII,

between the Rocky Mountains and the Sierra Nevadas, and
thus includes northern Arizona, California east of the Sierras,
Nevada, eastern Oregon and eastern Washington, Utah, parts
of Wyoming, the lowlands of Idaho, and reaches its northern
limit in the Okanagan Valley in eastern British Columbia.
Certain portions of this fauna also break through the mountain
barriers on the west and thus extend themselves. South
of Mt. Whitney a portion passes through the Walker Basin
in company with the more typical Sonoran and extends into
the San Joaquin Valley. Another portion runs westward from
Modoc and Lassen Counties and passing north of Mt. Shasta,
invades the northwestern part of Siskiyou County. The
insects of this fauna are generally derivatives of the true
Sonoran so not sharply differentiated. The jeyunas group of
the genus Platynus, Agrilus walsinghamt Cr., and the hairy
group of Eleodes are perhaps as characteristic as are any
Coleoptera that we have in the subfauna.

That other southern fauna, the one which comprises the
greater portion of the insect population of the southern part
of California, is not a derivative of the present Sonoran. Itisa
fauna which has come to us directly from the south through
Lower California and presumably in earlier times from lands
farther to the south. It is very old and very distinct, having
many genera and the bulk of its species totally different from
those of the Sonoran. It came into California long before the
Sonoran did and consequently is more thoroughly established
in southern and middle California. ‘The species are now some
of the most characteristic within the state and the fauna as a
whole is so dominantly Californian that it might be called the
Californian fauna. It is now to be found not only throughout
all of Southern California west of the San Bernardino and
Sierra Madre Ranges, but along the coast to San Francisco,
throughout the more southern portion of the Coast Range, and
the greater part of the San Joaquin and Sacramento Valleys,
extending as far north as Shasta County. In the drier parts
of the state, it is therefore even more dominant than the
Sonoran and it sends many of its characteristic forms well
within that claimed by the Vancouveran and Sierran as in the
foothill regions of the more northern Coast Range and the
Sierra Nevada itself, thus sharing equally with the faunas of
northern origin, the possession of the land. At one time, the

1919] Distribution of Insects Lit

northern species extended much farther south than at present
and in general were more in evidence in the south than now,
but within recent times they have undoubtedly both retreated
and decreased in numbers in the southern areas. The southern
forms have, on,the contrary, been doing the opposite so that
they have gradually supplanted the preceding. This, therefore,
accounts for the fact that there are islands of northern forms
within the territory occupied by southern forms. Wherever
southern forms have run north, they are always to be found
connected with their basic stock in the south, no matter how
far north they have gone and no matter at what time they
advanced. Some of the characteristic species of the Californian
fauna are the members of the dilatatus group of Anisodactylus
in the Carabide; Dystaxia, Schizopus and Glyptoscelimor pha,
in the Buprestide; Ipochus fasciatus Lec. in the Cerambycide;
Phloeodes, Coelus, Eulabis, Nycotporis, Cibdelis and Coniotis, in
Tenebrionide; and Trigonscuta and Rhigopsts in the Otiorychine.

Our west coast fauna we thus find has the bulk of its species
of insects included within the very old Vancouveran fauna and
the equally old Californian fauna, two faunas that are restricted
to the Pacific Coast and that have passed through the Pleisto-
cene without much injury to themselves. Supplementing these,
are the several marginal faunas of the mountains and the
desert faunas of the southeast. The Vancouveran, like those
found in the mountains of western Idaho, and in the southern
Alleghanies, is a relict fauna, a remnant of a more or less upland
fauna which was widely distributed throughout the more
northern parts of North America during Tertiary times. It is
the largest remnant of the three and is only surpassed among -
similar fauna by that of the Japano-Manchurian region. The
Californian is quite isolated though it shows a strong relation-
ship to that found in the more barren parts of northern Chili
and Peru. The marginal faunas on our mountains link up our
territory with that to the north and northeast of us and the
Sonoran does the same with regard to the country to the south-
east. Certain of our peculiarities can also be indicated by
considering the subject from a negative standpoint. For
instance, we find that very little of the Neotropical or tropical
fauna of South America, has reached us, whereas quite a
noticeable amount has found its way into eastern North
America. Dzabrotica soror Lec. is one of our few derivatives

12 Annals Entomological Society of America [Vol. XII,

from that. We also lack the most characteristic elements of
those faunas, the Austro-Riparian and Carolinian, which are
such a feature of the eastern part of North America. Thus
we have but a few Scarabaeide and few Chrysomelidé as com-
pared with the East; no representative of Lucanus, Copris,
Onthophagus, Anomala and but few of Phyllophaga (Lachnosterna)
from among the Lamellicorns; but a weak representation of
Melanotus, an Elaterid genus rich in species in the eastern
states; and not a single species of Evarthrus, Pasimachus and
Dicaelus from among the Carabidz. Hence we may say that
the fauna of western North America as a whole when judged
from the standpoint of its insects, is a most distinct one, yet
one which can be linked with the faunas of the rest of the
world.

A NEW CHALCID-FLY PARASITIC ON THE AUSTRALIAN
BULL-DOG ANT.*

By CuHARLEs T. BRUEs.

Several decades ago, Professor August Forelt found in
-cocoons of the large Australian bull-dog ant, Myrmecia forficata
Fabr., two specimens of a fine Eucharid. He recognized them
as parasites of the ant larve and sent them to the English
hymenopterist, Peter Cameron, who described them as Eucharis
myrmicie.{ This was the first record of a Eucharid parasite
of an ant, although several other genera have been reared
subsequently from other ants and it seems likely that all of the
Eucharide are ant-parasites.

Very recently Dr. R. J. Tillyard, of Hornsby, New South
Wales, bred from a cocoon of Myrmecia gulosa another Eucharid
which he sent to Prof. W. M. Wheeler, who has published
accounts of the habits of Orasema and several other genera.§

The species reared by Dr. Tillyard is quite distinct from
Eucharis myrmicie, although I suspect that both may belong
to the same genus. Cameron’s species is undoubtedly not an
Eucharis as that genus is at present restricted and it may
perhaps be a Psilogaster to which I believe the new species is
referable. Psilogaster was first proposed for an Egyptian
species by Blanchard and recently another one from Abyssinia
has been more carefully described and figured by Reichen-
spergert as P. fraudulentus. In addition to these, Dr. Wm. M.
Mann tells me that he has a new species which he hopes soon
to describe, taken by him in the Solomon Islands. The
Abyssinian form occurs with Pheidole megacephala. Both of
the African species have 11-jointed antenne as does a Tasmanian
Eucharid described by Walker as Psilogaster pallipes. Recently
Girault has added one from Australia with 10-jointed antenne,
and has proposed the genus Parapsilogaster for another with
12-jointed antennz. Since it is a difficult matter to say

* Contribution from the Entomological Laboratory of the Bussey Institution,
Harvard University, No. 152.

+t Am. Soc. Ent. Belgique, Vol. 20, p. 8. (1890).

t Mem. & Proc. Manchester Lit. & Philos. Soc., Vol. 4, p. 187. (1891).

§ Bull. American Mus. Nat. Hist., Vol. 23, pp. 1-98 (1907).
* Zool. Jahrb. Abth. f. Syst., Vol. 53, pp. 185-218. (1913).

13

14 Annals Entomological Society of America [Vol. XII,

whether the apical part of the antenna (“‘club’’) consists of one,
two or three separate joints, descriptions without figures may
be misleading on this point. The species bred by Dr. Tillyard
from Myrmecia differs considerably from P. fraudulentus
in both the adult and pupa and as they occur with such different
ants they may quite likely prove to be generically distinct.
In accordance with the large size of Myrmecia, it is more than
twice the size of the Abyssinian parasite of Pheidole and by
reason of its striking color pattern may be very readily recog-
nized.

Psilogaster fasciiventris sp. nov.

Imago. (Plate \T1Figsi2)ae Crengih (25) Sam elead
and thorax greenish bronze, less greenish and with purple
reflections on the propodeum; abdomen piceous, with trans-
verse yellowish bands, orange yellow on the second segment
and lighter yellow on the following segments; band of second
segment narrowly interrupted on the median line and curving
forward on the sides to near the apex of the petiole, separated
by its own width from the apical margin of the segment; those
of the following segments of similar form, but not interrupted
medially, that of the fifth widened medially and angularly
extended forward; petiole black with a purple cast; antenne
ferruginous, scape lighter; mandibles light brown; coxz black,
the front ones bronzed; femora fulvous; tibiae and tarsi pale
yellow, last tarsal joint black at apex; wings hyaline at base,
brown elsewhere, more deeply so anteriorly near the middle.
Head much narrower than the thorax, fully three times as broad
as thick; posterior margin slightly concave; face below the
antenne transversely striate, the stria continuing over the
cheeks and upward behind the eyes and across the occiput;
occipital margin raised; ocelli in a nearly straight line, the
posterior ones nearly as far from one another as from the eye-
margin; antennal basin transversely striated; eyes bare; malar
space nearly as long as the eye; antennz 11-jointed, scape
twice as long as thick; pedicel very short; first flagellar joint
as long as the scape and pedicel together, over three times as
long as broad; following joints growing shorter, the seventh
and eighth less than twice as long as thick; apical one longer.
Thorax above coarsely rugose-reticulate; parapsidal grooves
impressed only on posterior half; scutellum without distinct

1919] Chalcid-fly Parasitic on Ant 15

median furrow or depression; separated from the post scutellum
by a shallow groove, the two together acutely convex at apex,
but without teeth or projections. Propodeum flattened and
elevated medially, the sides of the central part convexly elevated
at the sides, lateral to this with a deep groove extending to the
hind coxa separating the lateral part which is acutely convex
above near the root of the hind wing. Pleurez sculptured like
the mesonotum. Petiole of abdomen as long as the slope of
the propodeum; second segment twice as long as the third and
fourth together; following very short, not visible from above;
apical margins of the second and third segments deeply excised
medially. Tarsal claws simple. Submarginal vein thickened
on its apical third; marginal half as long as the submarginal
and fully four times as long as the short, stout, nearly per-
pendicular stigmal vein; postmarginal extending about half-way
to tip of wing, but gradually evanescent at tip.

Type from Hornsby, New South Wales, in the writer’s
collection, bred from cocoon of Myrmecia gulosa by Dr. R. J.
Tillyard.

Male. A pupa (Plate II, Fig. 2) shows twelve distinctly
separate antennal joints and is, I suspect a male, and the apical
joints are longer, more slender and of approximately equal
length. In the female, if the lengthened and constricted 11th
joint were regarded as two joints, the apical one would be much
shorter than the penultimate.

PLANIDIUM, OR FirST LARVAL STAGE.

There are no specimens of this instar, but the cast skin
of one was found still attached to the mature larva described
below. This exuvium was firmly attached to the under surface
of the thorax just behind the head and I have been able to
remove it almost intact. When mounted in balsam it gives
some idea of the structure of the planidium which proves to be
very similar to that of Perilampus as described by Smith.*

Compared with Smith’s figure (1912, p. 46, fig. 26 d), of
the engorged planidium it appears to have eleven sclerites
behind the head, while Perilampus has twelve. The apical
plates are very small, however, and quite likely the number
is the same in both genera. The head is separated from the

* Bull. U. S. Dept. Agric., Bur. Ent., Tech. Ser. No. 19, pt. IV, pp. 33-69
(1912) and Psyche, Vol. 24, pp. 63-68. (1917).

16 Annals Entomological Society of America [Vol. XII,

rest of the dorsal integument in the cast skin, showing that
ecdysis must be accomplished by the formation of a dorsal
rent behind the head. The thoracic and abdominal plates,
except the last three are widely separated by the enormously
distended membrane, so that they appear as widely separated
bars like those of a much swollen termite queen. The first or .
prothoracic plate is broader than the others and crescentic,
being deeply emarginate in front. The three following are
narrower and but slightly curved; the following become gradu-
ally smaller to the minute apical one. At each side of each
plate is a more or less triangular, paler extension, separated
by a fine clear line; these probably represent pleural plates;
they are similar to those of Perilampus in having the tip pro-
longed into a long bristle-shaped point, but the posterior edge
bears several short teeth instead of bristles. Between the
plates and on the ventral surface, the membrane bears some
bristly hairs as in Perilampus, but the arrangement of these
cannot be made out clearly in the specimen. I find no trace
of the caudal bristles present in Perilampus, but these may
easily have been lost in ecdysis.

Larva. Full-grown, length 12 mm. There is a single
larva, evidently full-grown and ready to molt into the pupal
condition as the appendages of the pupa are to be seen through
the skin. The body is rather stout (Plate I, Fig. 1) nearly
straight and broadest near the posterior end. At the anterior
end there is a large vesicular swelling which extends ventrally,
behind it on the ventral side lies another smaller swelling
which evidently represents the head and mouthparts, although
no definite structures can be made out. Dorsally, and to a less
degree on the sides and below, the surface of the body is undu-
lating, indicating eleven segments, including the anterior
vesicle previously referred to. Of these segments the three
anterior ones (thoracic) are larger than the basal abdominal
ones, although the last several abdominal segments increase in
size till the sixth is the largest body segment; beyond the
segments become rapidly smaller and bent down ventrally;
finally at the apex on the ventral side of the body is a minute
tubercle which evidently is a twelfth segment. The surface
of the whole body is clothed with minute, colorless, sparsely
placed hairs which undoubtedly function as exudate organs.

1919| Chalcid-fly Parasitic on Ant 17

Pupa. (Plate II, Figs 1, 2,3). Length 15 mm. Elongate
and slender, with the abdomen not noticeably swollen and not
curved downward, the whole body with vesicular swellings.
The head is bent under the thorax so as to be scarcely visible
from above, but its projections extend forward as a central
bifurcate, transversely wrinkled papilla arid another one at each
side projecting forward and outward. The mesonotum bears
a small rounded tubercle at each side of the anterior edge,
an acute one near each tegula and a widely separated pair of
sub-acute ones on the disc before the scutellum; otherwise the
integument is simple except for a raised, longitudinally wrinkled
area between the anterior pair of tubercles. There are no tuber-
cles above the propodeum and petiole like those figured by
Reichensperger (loc. cit.) for P. fraudulentus.

Abdomen narrowly oval, broadest near the anterior end,
composed of five apparent large segments and a short apical
one, followed by a portion which appears to be retractile and
composed of several fleshy segments, terminated by a polliciform
projection. Dorsally the five large segments are separated
by sharply elevated transverse ridges, each interrupted narrowly
along the median line. At the median end each ts elevated as a
tooth, again as a more conspicuous and longer tooth at the
lateral end, which is well down on the side of the body; midway
between there is another tooth. Between the large teeth,
especially near the dorsal line, there are more or less distinct
minute denticulations. The ridges between all segments are
very nearly of the same size, except that the lateral tooth-like
projections of the first are much larger. The ventral surface
.is clearly separated by a slight ridge and groove just inside the
intersegmental teeth and is less convex than the dorsal surface;
only the second to fifth transverse ridges are clearly indicated;
they are continuous and much less conspicuous than the dorsal
ones; the first is faintly visible and bears an anteriorly directed
tooth at its middle. Seen from below, the dorsal surface of
the short sixth segment extends further down than the others
leaving only a narrow ventral part which is conically elevated;
in addition the apex of the fifth ventral segment bears a pair
of closely approximated tubercles at apex; following retractile.
segments without distinctly separated dorsal and ventral
surfaces.

Described from one perfect and several damaged specimens.

Rees
or

18 Annals Entomological Society of America [Vol. XII,

The very great similarity of the planidium stage of Psilo-
gaster to that of Perilampus is extremely interesting and sug-
gests a close affinity between the Eucharide and Perilampidz
as has already been pointed out by Smith from a comparison
of the planidia of Perilampus and Orasema. In fact, this stage
of Psilogaster fasciiventris 1s almost as close to that of Perilampus
hyalinus as the latter is to ‘“‘Perilampus, species a’’ figured
by Smith. In this case it may of course be possible that
“species a’’ is not a true Perilampus as Smith did not succeed
in rearing the adult. The host relations are very different
in the case of the two families so far as is known; all bred
Eucharids have been found to be parasites of ants while Peri-
lampus has been bred from a diverse series of hosts, including
Lepidoptera, Coleoptera, Neuroptera and some parasitic
Hymenoptera and Diptera.

The later larval and especially the pupal stages are much
more highly modified in the Eucharids than in Perilampus,
although in the larve there seems to be much diversity in this
respect. In the case of the larva, that of Psilogaster is com-
paratively simple, without conspicuous exudate organs, although
it is well supphed with minute surface hairs which no doubt
function as secretory organs of this type. Orasema possesses
numerous, highly developed exudate tubercles arranged seg-
mentally on both the thorax and abdomen. On the other hand,
Perilampus shows similar conspicuous tubercles and projections,
although it seems improbable in view of the host relations in
this case that these can function as true exudate organs. In
the pupa, that of Perilampus is simpler, although with small
intersegmented ridges on the abdomen. In Orasema the pupa .
exhibits well developed transverse welts between the abdominal
segments as well as some on the head. The climax appears in
Psilogaster where the exudate organs are extremely large and
elaborate as described in the present account.

There would seem to be two possible reasons for the presence
of the highly developed exudate-like organs on the Perilampus
larva. It might be that Perilampus is derived from Eucharid-
like ancestors in which these organs actually functioned for
’ the elaboration of an exudate, but this seems highly improbable
in view of the much more specialized habits and structure of the
Eucharide. On the other hand, it may be that the integu-
mental modifications of Perilampus really do function as exudate

1919] Chalcid-fly Parasitic on Ant 19

organs. But if such were the case it seems impossible to point
out any way in which they might bear any relation to the host.
It is of course possible that neither of the foregoing suppositions
is true, and that the apparent exudatoria or blood spaces between
the body and integument are simply adaptations to assist
in ecdysis. Such can hardly be the case however, as they do
not exist in other parasitic Hymenoptera with habits similar
to those of Perilampus and in which ecdysis must be undergone
under the same conditions.

None of these suppositions seems adequately to explain
the structure of the Perilampus larva and pupa, and unless the
exudatoria are functionless their presence must be due to some
other cause. It is barely possible that they may bear a relation
to the host other than those enumerated above.

It is known that the larve or many if not all externally
feeding parasitic Hymenoptera secrete a salivary liquid which is
injected into the host. This contains digestive enzymes which
act upon the tissues to induce extra-intestinal digestion and a
consequent liquefaction of the food material which is then more
readily ingested. In ants at least, as has been pointed out by
Wheeler (1918) there are undoubtedly cases where such secre-
tions have both a digestive and exudatorial function. With this
in mind it is possible that the secretion which is actually pro-
duced on the surface of the body of the Perilampus larva
(vide Smith 1912, p. 50) may have a digestive function. With
this hypothesis there is one difficulty; only a small part of this
secretion could possibly enter the wound caused by the feeding
larva in the host.

Of all these conflicting suppositions none appears satis-
factorily to explain the conditions as they occur in Perilampus,
and it is perhaps more likely that the exudatoria may have
been present in the common ancestors of both Perilampide and
Eucharide. If such be true, they have persisted in both
families and have in the Eucharide assumed a secondary
function-as their exudates have proved to be attractive to the
ants upon which they are parasitic. We may then believe that
they. :were originally either secretory or excretory organs,
or those’ assisting in some way in the process of ecdysis

Tubércular excrescences of somewhat similar appearances
are to be seen in many larval insects of very diverse groups,
é. Bx in, many caterpillars, in certain parasitic beetles, in one

20 Annals Entomological Society of America [Vol. XII,

genus of bees (Allodape) and in some ants. In connection with
some of these, Professor Wheeler has suggested to me that they
may be of assistance in eliminating the large amounts of water
ingested by rapidly growing insect larve. Quite recently
Roubaud* has figured (p. 15) larve of the remarkable Rhip-
iphorid beetle Macrosiagon ferruginea which are external
parasites of the wasp, Synagris. In these larve the body is
covered with tubercles much like those of Psilogaster and
Perilampus. Roubaud also found in the larva of a Braconid
parasite occurring in the nests of Rhynchium, transverse welts of
apparently similar nature (/. c., p. 35, Fig. 14). In this case the
Braconid larve (Allodorus major) feed not upon the wasp
larve, but upon one of the caterpillars used for provisioning
the nest. They develop at an unusually rapid rate, and on
this account may easily require additional facilities for excreting
water, particularly as they possess a closed alimentary tract
and must excrete all excess water through the skin. The
habits of a somewhat closely related genus of Braconide,
Chelonus, have been described by Pierce and Holloway in
America.t This species is an internal parasite and undergoes
a much slower development, the larva requiring about three
weeks to mature. These writers do not describe the larva,
but it seems probable that they are not tuberculate, as such a
peculiarity would undoubtedly have been mentioned. It
seems probable, therefore, that at least one form related to
Allodorus, but developing more slowly, lacks the welts present
in Allodorus. This would appear to lend color to the suggestion
made above, that the welted or tuberculated integument may
function in excreting excess water.

* Ann. Sci. Nat. Zool. 1916 Recherches biologiques sur les guépes solitaires
et sociales d’ Afrique. ;

t+ Notes on the Biology of Chelonus texanus Cress., Journ. Econ. Entom.,
Vol. 5, pp. 425-428. (1912).

1919]

Chalcid-fly Parasitic on Ant

EXPLANATION OF PLATES.

PLATE 1. Psilogaster fasciiventris sp. nov.

Fig. 1. Mature larva, lateral view.
Fig. 2. Female imago.

PLATE II. Psilogaster fasciiventris sp. nov.

Fig. 1. Pupa, lateral view.
Fig. 2. Pupa, ventral view.
Fig. 3. Pupa, dorsal view.

21

ANNALS E.S. A. VOL. XII, PLATE I.

Fig. 2 i4

C. T. Brues

ANNALS E.S. A. VoL. XII, PLATE I!

C. T. Brues

eee

at a
a Se eee). Meese eee

: i
i
7
'
,
A Aare
;
i 1
th =. .
t A
i
y
y : a

NOTES ON THE CRANE-FLIES OF THE HAWAIIAN
ISLANDS (Tipulidz, Diptera).

By Cuarves P. ALEXANDER, Urbana, Illinois.

During the past few years some extensive collections of
Hawaiian crane-flies have been received from Dr. James F.
Illingworth and Mr. O. H. Swezey, to both of whom I am
indebted for many favors. The records of these collections,
together with a few additional specimens from other sources,
are herein recorded to supplement our scanty knowledge of
the Hawaiian Tipulide. A few general statements of the
faunal constituents may be given.

As might be expected, the crane-fly fauna of the Islands
is strongly endemic. The dominant genus is Dicranomyia
with about a dozen species, all of which are apparently con-
fined to the Islands. The remaining genera, Libnotes, Sty-
ringomyia, Gonomyia and Trimicra, are represented each by a
single species, all, with the exception of Styringomyia didyma,
apparently being endemic. This last mentioned species breeds
in decaying organic matter, such as manure, and has probably
been spread over the Australasian islands through the agency
of man. The entire lack of the larger, vigorous Tipuline forms
is striking, but entirely in agreement with the fauna of many
other oceanic islands, where the Tipuline are entirely lacking
and the dominant genera are Dicranomyia and related groups,
(Seychelles, Guam, Fiji, etc.)

The following transcription of a letter received Bgoanel Ds
Illingworth will give a clear idea of the conditions under which
Hawaiian crane-flies are found. Under date of March 6, 1917,
he writes as follows:

“Last week end I spent in the Waianae Mountains, which are at
the far end of the Island. The range is much older than the mountains
near Honolulu (Koolau) and the native insect fauna appears to be more
abundant.

‘““T have never seen crane-flies more abundant, but I fear they
belong to a few species. I followed along the stream which had many
falls—every wet cavern was filled with them. I tried sweeping the
grass along the banks but with no success. One only gets crane-flies
in the mountains here and it is a rather difficult matter to get to the
places where they are abundant. One or two common species come to
the lamps in the low lands—these I have sent before.”

25

26 Annals Entomological Society of America {[Vol. XII,

Since the appearance of the Fauna Hawaiiensis (1901) in
which Grimshaw describes practically all of the Hawaiian
crane-flies, a new species, Dicranomyia foliocuniculator, has
been described by Swezey. This latter species is of extreme
interest because of the leaf-mining habits of its larvee—a habitat
that is unique in the family so far as at present known.

The types of the new species, with the exception of Dicrano-
myia swezeyt, are in the writer’s collection; +his latter type
and paratypes of the other species were returned to Mr. Swezey.

FAMILY TIPULIDA.
SuB-FamILy LIMNOBIINZ.
Genus Libnotes Westwood.
Libnotes perkinsi (Grimsh.)

This handsome crane-fly is common and apparently well
distributed throughout the Islands. It was described as a
Limnobia but the reference to Libnotes seems more nearly correct.
The following additional records from Oahu are at hand:

Honolulu, April 10, 1914 (O. H. Swezey).

Nunanu Valley, May 4, 1915 (A. H. Case).

The exotic genera that center around Limnobia and Dicrano-
myta include members which run very close to the above groups.
In their typical condition these genera (such as Libnotes,
Dapanoptera, Pertpheroptera, etc.) are well defined, but the
presence of less clearly differentiated species tends to break down
all distinctions. Speiser (1909) described three African species
of Limnobia (Limonia) that are, in my opinion, referrable to
two other genera, Libnotes, (rhizosema and oresitropha) and
Rhipidia (miosema). Other authorities, as Bergroth, Edwards,
etc., include the species of Dicranomyia that possess a long
subcosta in the genus Limnobia, disregarding all other features
of organization, and despite the fact that Osten Sacken in his
classic monograph of a half century ago clearly differentiated
these two groups of Dicranomyia. In determining the generic
limits of this division of flies, other features of structure must be
considered, and the entire burden not placed on a single char-
acter. When this is done, the habitus of the insect and the
smaller details of structure that are perceivable but difficult
of definition, will aid in separating various groups. It is admit-
tedly true that in many such cases the reference of species to
genera becomes largely a matter of personal opinion. The

1919] Crane-fly of Hawaiian Islands

LS)
“J

suggestion has been made that it would perhaps be better to
consider these diverse Limnobiine genera as lesser groups, such
as sub-genera, but in my opinion no benefit would be derived
from such an action, since it would be exactly as difficult to
separate and define these sub-genera. These various exotic
genera are probably valid, but Goniodineura van der Wulp is
doubtfully distinct from Libnotes.

Genus Dicranomyia Stephens.
Dicranomyia stygipennis, sp. n.

Coloration dark brown throughout, including the wings.

Male: Length, 6-7.5 mm.; wing, 7.5-8 mm.

Female: Length, 6.4-8.5 mm.; wing, 7.8-9 mm.

Rostrum and palpi dark brown. Antenne dark brown throughout,
the flagellar segments elongate-oval. Head dark brown.

Thorax dark brown, the pronotum a little yellowish. Legs dark
brown, the extreme base of the femora a little paler. Halteres dark
brownish black throughout. Wings with a strong brown suffusion;
stigma rather distinct, of a somewhat darker brown. Venation: Sc
ending about opposite the origin of the sector, Sco close to the tip of
Sa; Rs about twice the length of the basal deflection of Ris;
cell 1st Mz closed; basal deflection of Cm, at the fork of M.

Abdomen dark brown. Hypopygium of male with the ninth
pleurite small, only about four-tenths the length of the ventral pleural
lobe; ventral lobes large, fleshy, the basal inner angle suffused inwardly,
near its base with two powerful spines; dorsal lobes short, stout, slightly
curved, the tip of each produced into a slender, blackened spine.

Holotype, &, Waianae Mountains, Oahu, altitude 2000
feet, March 5, 1917 (J. F. Illingworth).

Allotype, 9°, with the type.

Paratypes. Very numerous specimens of both sexes, from
the type locality; Kaupo, Maui, altitude 2000 to 4000 feet,
January 1, 1915 (H. T. Osborn); Haleakala, Maui, altitude
9000 feet, September 27, 1914 (H. T. Osborn).

This common fly is notable by its dark coloration. I would
have identified it as being D. brunnea Grimsh. (1901) except
for the discrepancy in the coloration of the wings. In any case,
Grimshaw’s name is preoccupied by D. brunnea Doane (Journ.
Ent. N. Y. Soc., Vol. 8, p. 184; 1900), a fact that was pointed
out by the author several years ago (Psyche, Vol. 18, p. 194;
1911).

Dicranomyia grimshawi, nom. n.

New name for D. apicalis Grimsh. (1901), not D. apicalis
(Wied.) (1828).

28 Annals Entomological Society of America [Vol. XII,

Two males from Olaa, Hawaii, altitude 2500 feet (W. H.
Ashmead), in the collection of the U.S. Nat. Museum.

Dicranomyia hawaiiensis Grimsh.

This beautiful Dicranomyia is widely distributed in the
Islands. The following additional records of distribution are
available. Island of Oahu: Honolulu, February 7, 1913
(O. H. Swezey); Palolo, February 8, 1914 (O. H. Swezey);
Olympus, September 8, 1912 (O. H. Swezey); Waianae Moun-
tains, altitude 2000 feet, March 5, 1917 (J. F. Illingworth).

Dicranomyia foliocuniculator Swez.

This recently described species is apparently common and
widely distributed in the Islands. As stated before, it is unique
in the leaf-mining habits of the larva, a habitat that will prob-
ably be found to be shared by other species of the Islands,
especially D. jacobus, D. swezeyi, etc., forms that are undoubtedly
related to D. foliocuniculator.

Numerous specimens are available from the Waianae
Mountains, Oahu, altitude 2000 feet, March 5, 1917 (J. F.
Illingworth) associated with D. hawaziensis, D. stygipennis, etc.
This locality has been mentioned in the introductory paragraph
of ‘this article. Waimano, Oahu, January. 5), 1913, (©) ae
Swezey). Iao Valley, Maui, altitude 700 feet, January 3, 1914
(J. F. Illingworth).

Dicranomyia jacobus, sp. n.

Size larger than in foliocuniculator (length of the male over 4 mm.);
coloration of the body darker brown; wings with a decided brownish
tinge.

Male: Length, 4.3-4.4 mm.; wing, 5.2-5.5 mm.

Female: Length, 5.1-6.5 mm.;° wing, 5.8—6.2 mm.

Rostrum and palpi dark brownish black. Antenne rather short,
dark brownish black, the flagellar segments short, suboval to rounded,
the apical segments more elongated. Head dark brown.

Neck rather elongated. Thorax very high and gibbous, dark brown,
without stripes (in alcohol). Halteres rather short, the knobs large,
the stem pale, the knobs darker, brown. Legs with the coxe and tro-
chanters brown; remainder of the legs rather dark brown. Wings
light brown, the stigma indistinct, only a little darker than the remain-
der of the wings; veins dark brown. Venation: Sc short, Sc, ending
about opposite or slightly beyond the origin of Rs; Sc. removed from the
tip of Sc, for a distance about equal to cross-vein 7; Rs long, somewhat
arcuated, from. two to three times as long as the basal deflection of
Rais; cell 1st Mz large, as long as the veins issuing from it; basal deflec-
tion of Cu, just before, at, or even slightly beyond, the fork of M.

1919] Crane-fly of Hawaiian Islands 29

Abdomen dark brownish black, the sternites a little paler. Hypo-
pygium with the ninth tergite almost straight across the caudal margin;
pleurites rather short, shorter than the apical lobes; ventral inner
side of the pleurite produced entad into a complex lobe which is
expanded at its apex and bears at the inner margin a lateral cylindrical
lobe with long hairs. Dorsal apical appendage a chitinized rod almost
straight basally, toward the tip slightly bent inward and acutely pointed;
ventral apical appendage a large fleshy lobe whose inner margin sub-
apically is produced into a long, slender, cylindrical, chitinized point,
rather blunt at the apex and here with a few hairs; at its base on the
outer side with two sharp chitinized spines which are directed back-
ward. Anal tube short, very broad, projecting slightly beyond the
caudal margin of the ninth tergite, its apex feebly concave. Penis
guard elongate, narrowed toward the apex which is bent strongly
ventrad.

Holotype, &, Iao Valley, Island of Maui; altitude 700 feet;
January 3, 1914 (J. F. Illingworth).

Allotype, 2, topotypic.

Paratopotypes, 23’s, 2 Q’s.

‘This insect is respectfully dedicated to Dr. James F. Illing-
worth, to whom I am indebted for many favors.

Dicranomyia swezeyi, sp. n.

Coloration pale yellow throughout; wings yellow; cell 1st Mz open
by the atrophy of m.

Female: Length, 2.8-3 mm.; wing, 3.7-3.8 mm.

Rostrum and palpi pale yellowish brown. Antennz short, pale
brown. Eyes black. Head light yellow.

Thorax pale yellow, the prescutum a little brownish. Pleura
sparsely dusted with gray. Legs with the coxe pale; femora pale
brownish yellow, the tips brown; tibiae and tarsi yellowish brown.
Halteres pale. Wings pale yellow, the stigma pale brown; veins
yellow. Venation: Sc short, ending before the origin of Rs; Rs mod-
erately elongated, about one and one-half the length of the deflection of
Riis; deflection of R14; strongly arcuated at origin; cell /st M2 large,
subquadrate, indistinctly open by the atrophy of m; basal deflection of
Cu, just before the fork of M.

Abdomen brownish yellow, dorsal valves of the ovipositor mod-
erately elongated and quite strongly arcuated.

Holotype, &, Olympus, Oahu, September 8, 1912 (O. H.
Swezey).

Allotype, 2, with the type.

This interesting little species is dedicated to its collector,
Mr. O. H. Swezey, to whom I am indebted for many favors.
It is apparently related to D. foliocuniculator Swez., but is
readily distinguished by its very small size, uniform pallid
coloration, and the open cell 1st Me.

30 Annals Entomological Society of America [Vol. XII,

Genus Styringomyia Loew.
Styringomyia didyma Grimsh.
Specimens are in the collection from Honolulu taken in
May, 1914. It is apparently well distributed throughout parts
of Australasia.

Genus Trimicra Osten Sacken.
Trimicra lateralis Grimsh. )
Specimens from Honolulu, April 10, 1914 (O. H. Swezey).

Genus Gonomyia Meigen.

Gonomyia (Leiponeura) hawaiiensis, sp. n.

Coloration dark brown; antennze dark brown, the first flagellar
segment light yellow; thoracic pleura striped with brown and white;
wings with a pale brown tinge, longitudinal veins long and slender.

Female: Length, 3.2-3.4 mm.; wing 3.2-3.5 mm.

Rostrum and palpi dark brown. Antennz dark brown, the first
segment of the flagellum yellow. Head gray.

Mesonotal prescutum dark brown, the extreme lateral margin
pale; remainder of the thoracic notum dark brown. Pleura white, with
a dorsal dark brown longitudinal stripe; a ventral brown stripe occu-
pying the sternum. Legs with the coxe yellowish; trochanters and
femora brown, the apices of the latter and the tibiz a little darker;
tarsi brownish yellow. Halteres dark brown, the knobs conspicuously
light yellow. Wings pale brown, iridescent, the stigma indistinctly
darker brown; veins brown. Venation: Sc short, ending before the
origin of the sector; Rs long, gently arcuated; basal deflection of Rijs
very. short; longitudinal veins beyond the cord elongated; basal
deflection of Cu, at or before the fork of M.

Abdomen dark brown, the ovipositor with the dorsal valves very
long and slender, slightly up-curved, dark brown at the base, horn
yellow beyond.

Holotype, 9, Koolau Mountains, Oahu, altitude 1500
feet, February, 1917 (J. F. Illingworth). Found on a wet
bank.

Paratypes, 9's, Wailuke, Oahu, January, 1915. (O08 H.
Swezey); Tantalus, Oahu, October 15, 1911 (O. H. Swezey);
Palolo, Oahu, February 8, 1914 (O. H. Swezey).

The type of the subgenus Lezponeurt Skuse (Proc. Linn. Soc.
New South Wales, ser. 2, vol. 4, p. 795; 1889) is Gonomyia
gracilis (Skuse); the name is homonymous with Gonomyia
gracilis (Zett.)(1838) and is here renamed Gonomyia (Leiponeura)
skuset, nom. n.

THE CALIFORNIAN SPECIES OF MALTHODES.
ByataCaWALL,

The presence of several species of Malthodes among the
things which Dr. Blaisdell has asked me to study in connection
with his forthcoming paper on the Coleoptera of Humboldt
County, California, offers the opportunity of reviewing the
Californian species of this genus in the Doctor’s and my own
collections, a very- large proportion of which are as yet unde-
scribed. ”

These delicate little things have been generally neglected
by collectors, and such cabinet specimens as one runs across
are too often ruined in mounting. When carefully collected
and mounted, they are wonderfully interesting little creatures
and offer an almost unparalleled wealth of sexual variation
in the formation of the terminal abdominal segments of the
male. The females are on the other hand rather monotonous
in their uniformity, and are rarely distinguishable specifically
except as accompanied by males. In the males themselves
there are few taxonomic characters outside of the ventral sexual
modifications, and this together with the fact that there is
little use in attempting to give comparative measurements in
insects having such fragile structure and soft integuments, is a
sufficient reason for the brevity of the following descriptions,
which indeed might about as well have been made still shorter.

Very little has been said in the descriptions about the
females. These almost invariably differ from the males by the
smaller, less prominent eyes and consequently narrower head,
which is usually scarcely as wide as the thorax; by the shorter
antennz, with joints two to four more nearly equal in length, or
at least with the third joint not longer than the second; by the
rather more transverse thorax, and sometimes somewhat
shorter elytra. The last ventral segment in this sex is rather
deeply incised in the same general manner in all the species
here treated.

The remarkable modifications of the abdominal apex in
the male are often difficult to see in their entirety, but the form
of the sixth and seventh ventrals is usually visible, and these

31

32 Annals Entomological Society of America [Vol. XII,

have therefore been selected for representation in the accom-
panying plate. These sketches are not drawn to any given scale,
and are not to be regarded as absolutely accurate as regards
relative dimensions of parts, but they are sufficiently so for the
purpose they are designed to serve. A certain degree of
variation has indeed been observed in these dimensions in one
or two species, quite independent of any extension or contraction
of the parts, which though sometimes noticeable, is less common
than one would expect.

In Le Conte’s latest treatment of the Lampyrida* there is
‘given a table of our species of Malthodes, of which three—
laticollis, fragilis and fusculus—are accredited to California.
The type of fragilis is from the Atlantic Coast region, and an
examination of the Le Conte specimens some years ago con-
vinced me that the Californian form so referred was not the
same. This latter did not appear to be identical with any of
the new species herein described. This leaves only two Le-
Contean species described from the state, one of which, fusculus,
I do not recognize in the material at hand. To these two species
must be added the M. ligulifer described by Bergrotht from
Monterey, California, and said to be related by its male sexual
characters to spado Lec. The prothorax of ligulifer is said to be
‘‘nigro-piceus, anguste luteo-circumlimbatus,’’ which does not
apply normally to any Californian species known to me, and is
only approached by some forms of basalis, near which it may
be listed for the present; the terminal ventral segments of the
male are, however, not defined with sufficient exactness to
enable me to include it in the following table.

* Trans. Am. Ent. Soc. 1881, p. 60.
t Ann. Soc. Ent. France (Bull. Ent. CCIII) 1889.

1919] Californian Species of Malthodes 30

TABLE OF SPECIES.

Seventh ventral segment of male not produced, sixth broadly, not deeply
emarginate, fifth broadly angulate emarginate.
eimale ap LEROUSr icine (it ao cache «ie < rs vil bine < OAL iu RRR eee a reflexus
HER SIRE UA NVA GC CLMERR AN CRRA Pe Ss iscd the .c oon ays esd Soe Sy Le Roe laticollis
Seventh ventral segment of male more or less strongly produced, sixth very
broadly and deeply emarginate, fifth not emarginate.
Seventh ventral of male rounded at tip, size large.............. magister
Seventh ventral of male emarginate, notched or forked at tip.......... 2
Seventh ventral of male not or scarcely more than twice as long as wide,
never projecting beyond the sixth a distance as great as the length of
EASY TEV RU SIee Sex o eaete eet ai or OER eevee NOE IE EIELOT A oa eas echo. Bn Shoe em Aan
Seventh ventral segment of male much more than twice (usually three to
five times) as long as wide, and projecting beyond the sixth a distance
equal to or greater than the length of the latter.......................::- 4
Seventh ventral of male deeply incised at tip, eyes very large, separated
anteriorly by not more than their own horizontal width as viewed from

THERETO tee Se aerenes hac etaeh cee orc earn tele aban ood GR Atousy otto nte eae Ose Ie vigilans
Seventh ventral of male broadly notched or emarginate at tip.
Antennzeswathytworsasal joimtsmyellowe cc. dace cis cis neem = lee basalis

Antenne entirely fuscous or piceous.
Eyes large and prominent, the head conspicuously wider than the

thorax thoraxmyellows cert iok acters Ae cee ole Sine sete ee rake mitificus
Eyes relatively small, head not very much wider than the thorax,
Mew AL Ce RED ICCOUS te cree rier weaermcrts item tS. mea. e ce toctoceze Rice a piceolus
Seventh ventral of male with apical notch not deeper than the width of the
SERIMEMen atts HART OWESUIPALUS <4. sos cnc. cdcekece surm acltd ele ohne ea ale ee 5
Seventh ventral of male more deeply forked than the minimum width of
(ENS GEESE se x cabs bro ID Aicich Brenton testa cua SAG a cree on ete AM ee 6
Apex of seventh ventral of male concavely beveled inferiorly, as seen in
profile.
Seventh ventral of male nearly straight in profile............... mollis
Seventh ventral of male distinctly ascending in profile, the tips more
Giveroinolas wiewed: vertically 222 se.. ach eked ee ts eee appendiculatus

Apex of seventh ventral of male simple.

Fifth (?) dorsal segment (co) with a strongly produced overhanging
process which is horizontally dilated and rounded at tip, broadly
arched as viewed in profile; seventh ventral not bisinuate...obductus

Dorsal segments not produced apically, seventh ventral of male

StronplympisintatesineprOllens er peia ie cranes ror cieenia: bicurvatus
Ascending portion of seventh ventral straight (in profile) and lying between
long, slender, rectangularly bent dorsal processes............. complicatus

Ascending portion of seventh ventral straight, tip beveled, no slender
dorsal processes, ventral segments 3 to 5 silvery sericeous at
SACHIN: | othe dod oMON GH Ao O MC TORO EO CT RICIES ao oon Reet ae sericetventris

Ascending portion of seventh ventral sinuous or bent.

Emargination of sixth ventral (co) not truncate at bottom, seventh of
nearly uniform width, forks parallel.
Head (co) much wider than the thorax, eyes large, separated in

frontwoy, abouvinern Own Giametereesss sscesnectes ts suss 4. vapidus
Head (co) narrower eyes smaller and separated on the front by
more than 1/4 times their own diameter.................... fusculus:

Emargination of sixth ventral truncate at bottom.
Seventh ventral sinuate apically in profile, less widely and deeply
ova asl od Us open OO GOS 6 EO Ob OT SA Oe eee DS SN CACC naam tularensis
Seventh ventral apically bent, more widely and deeply forked. .visceratus

34 Annals Entomological Society of America {[Vol. XII,

Malthodes reflexus sp. nov.

Piceous, prothorax not paler. Antennz piceous, basal joint not
appreciably paler, about reaching the elytral tips in the male, a little
shorter in the female, joints 2-4 subequal, the third scarcely or barely
as long as the second, the fourth only slightly if at all longer than the
second, intermediate joints two and one-half times as long as wide in
the male, twice as long as wide in the female. Head subequal in width
to the thorax in the male, obviously narrower than the thorax in the
female; eyes not very prominent, the width of the front between them
fully three times that of the eye. Prothorax quite strongly transverse,
sides slightly convergent from the anterior angles to the base, the
margins more strongly reflexed than usual; disk bitmpressed at middle,
surface distinctly, moderately closely punctate. Elytra normal in the
male, in the female much shorter, less than twice as long as wide, the
wings entirely wanting, and the abdomen projecting beyond the elytra
a distance as great as the elytral length. Length about 2 mm.

Abdominal sexual characters. The seventh ventral of the male
is not produced, the genital appendages are similar in type to those
of laticollis, the lateral processes broader basally, rapidly narrowing to
the acuminate tip, which is slightly curved or everted, central piece not
keeled beneath, but angularly prominent at tip superiorly, the fila-
mentous process inferior in position; last ventral of female acutely
rather deeply incised.

California. Middle coast region. 46, 59, sent by Dr.
Blaisdell.

Specific localities are: Marin Co. (type o); Fairfax, Marin
€o.,. V-7-11- Vine” Hull, “Contras Costa ~Co:. > Vila7-05- Niles
Canon, V-16 (W. M. Giffard).

Very easily recognizable by the entirely piceous color,
broad distinctly punctate thorax with entire side margin
reflexed, simple, non-produced seventh ventral of male, and
apterous female.

Malthodes laticollis Lec.

Piceous or piceo-testaceous, head darker, thorax yellow: with the
front angles more or less infuscate. Antennze fuscous, basal joint paler,
as long as the body in the male, second and third joints equal, fourth
just visibly longer, intermediate joints about three times as long as
wide. Head slightly wider than the thorax in the male, eyes mod-
erately large, their longest diameter not much less than the minimum
width of the front between them; head in the female a little narrower
than the thorax, the eyes notably smaller. Thorax strongly transverse,
a little more so in the female as usual, widest at the front angles, which
are reflexo-incrassate, sides nearly straight and parallel posteriorly, the
margin distinctly reflexed throughout; surface evidently but finely

1919] Californian Species of Mathodes aD

punctate, disk biimpressed at middle, sometimes only feebly so. Length
about 2144 mm. |

Abdominal sexual characters. Seventh ventral of male simple and
not produced; genital appendages short, consisting of two piceous
slender lateral processes, hooked (abruptly bent at right angles and
acuminate) at tip; a central piece, pale in color, broad at base, with a
very thin laminate keel beneath and a curved terminal filamentous
process.

California. Described from Santa Cruz Island. Specimens
in my collection are from Riverside, Pomona, Pasadena, Mt.
Wilson and San Bernardino Mts., May, July.

The abdominal characters of the male as given by Le Conte
in his synopsis of the Lampyride (Trans. Am. Ent. Soc., 1881,
p. 60), do not seem.to apply to the species here described, but
the fact remains that there are no males in the Le Conte col-
lection. Two females only represent the species, the first
(on the label) being the original Santa Cruz Island type, while
the second—a later acquisition—is evidently a different species.
Specimens of the species I have described above were carefully
compared with the Santa Cruz type and are to all appearances
identical.

Malthodes magister sp. nov.

Piceo-testaceous, head darker, thorax fulvotestaceous with a blackish
discal spot in the type which is nearly obsolete in a second example.
Antenne entirely fuscous, passing the elytral tips in the male; joints
2-4 distinctly increasing in length, the intermediate joints fully four
times as long as wide. Head (<7) distinctly wider than the thorax, eyes
only moderate in size, separated in front by more than twice their own
width as seen from the front. Thorax but slightly wider than long, the
front angles a little prominent and reflexed, sides behind the angles
narrowly margined, surface finely sparsely punctulate. Length 4.5 to
o mm.

Abdominal sexual characters. Male: Sixth ventral deeply, broadly
emarginate; seventh moderately produced, tip gradually narrowed and
rounded, entire. Within the seventh ventral is a broad tongue-like
process which is emarginate at tip. The terminal dorsal segments
project considerably beyond the ventral at apex. Female not known.

California. Humboldt Co. The type collected at Blair’s
Ranch, Redwood Cr., Sept. 6, by H. S. Barber; the second
example at Green Point, June 4, by Dr. F. E. Blaisdell. This is
the largest species yet discovered in our fauna and is quite
distinct in the male sexual characters.

36 Annals Entomological Society of America |Vol. SL,

Malthodes vigilans sp. nov.

Piceo-testaceous, head darker, thorax yellow. Antennz fuscous,
slightly longer than the entire body in the male, joints 2—4 increasing
in length, the third however, only slightly longer than the second;
intermediate joints (co) about five times as long as wide, (9) twice as
long as wide. Head much wider than the thorax in the male, scarcely at
all so in the female; eyes (c’) very large and prominent, separated in
front by but little more than their own width. Thorax moderately
transverse, a little more so in the female as usual, sides parallel, feebly
margined, the margin just perceptibly thicker at the front angles,
surface scarcely visibly punctate. Length about 3 mm.

Abdominal sexual characters. Male with sixth ventral deeply,
broadly emarginate, seventh rather short and broad, projecting beyond
the sixth a distance less than the length of the latter, the tip deeply
narrowly incised. Last dorsal not projecting beyond the last ventral.
Female as usual.

California. San Bernardino Mts., July; 2°, 19, collected
by the writer. The type is a male.

Malthodes basalis sp. nov.

Fuscous, front anteriorly yellow, prothorax varying from yellow
with faint diffuse fuscous discal cloud, to nearly evenly fuscous with the
extreme basal and apical edge yellow. Antenne fuscous with basal two
joints yellow, as long as the body in the male, joints two to four increas-
ing slightly in length, intermediate joints about three times as long as
wide. Head (co) slightly wider than the thorax, the eyes only mod-
erately prominent, separated in front by twice their own width or
slightly less. Thorax moderately transverse, sides parallel, narrowly
margined, the front angles scarcely more prominent. Length 2 to 3 mm.

Abdominal sexual characters. Male: sixth ventral deeply emarginate;
seventh broad, nearly straight in profile, extending beyond the sixth
a distance subequal to the length of the latter, the tip broadly emar-
ginate; last dorsal not produced.

California. Yosemite, June 16 (W. M. Giffard), sent by
Dr. Blaisdell; Licking Fork, Mokelumne River, June, (Blais-
dell); Lake Tahoe, July 17-21.

Nevada. Ormsby Co., July (Baker).

Fifteen examples are before me. The type is a Yosemite
male.

The bright yellow basal two joints of the antenne will
probably serve to separate this from the other California species.

1919] Californian Species of Malthodes al

Malthodes mitificus sp. nov.

Fuscotestaceous, thorax yellow. Antenne longer than the body
in the male, entirely fuscous, joints two to four distinctly increasing in
length, intermediate joints about four times as long as wide. Head (0)
conspicuously wider than the thorax; eyes large and prominent, sep-
arated anteriorly by a distance not much greater than their own diam-
eter. Prothorax moderately transverse, sides parallel, narrowly mar-
gined, front angles slightly thickened but scarcely more prominent;
surface shining, minutely sparsely punctulate. Length about 3 mm.

Abdominal sexual characters. Male: sixth ventral deeply emar-
ginate, seventh broad, extending beyond the sixth by about the length
of the latter, gradually narrowed apically, the tip broadly notched; last
dorsal segment not produced.

California. Mokelumne Hill. 5, sent by Dr. Blaisdell.
This species should be easily recognizable by the tabular
characters. The ventral sexual structure is nearly as in basalis.

Malthodes piceolus sp. nov.

_ Piceous, elytra, body beneath and legs. more or less piceotestaceous
as usual. Prothorax entirely piceous except for paler basal and apical
edge; mouth yellowish in some examples, not so in others. Antenne
entirely dark, fully as long as the entire body in the male, in which sex
joints two to four are evidently increasing in length, and the inter-
mediate joints fully four times as long as wide. Head a little wider
than the prothorax in the male, eyes not very large, separated ante-
riorly by rather more than twice their own width as viewed from the
front. Prothorax only moderately transverse, sides parallel or slightly
convergent behind, margin narrow, slightly thickened at the front
angles, surface moderately shining and very finely punctate. Length
(head deflexed) 2 mm. to tip of elytra.

Abdominal sexual characters. Male: sixth ventral deeply emarginate,
seventh broad, nearly parallel, passing the apex of the sixth by a dis-
tance rather less than the length of the latter, -tip broadly not deeply
emarginate, a little deflexed at its extremity as seen in profile.

California. Green Point, Humboldt Co., June 4-7 (Blais-
dell). The type is a male.

Malthodes mollis sp. nov.

Fuscotestaceous, head blackish, thorax rufotestaceous, frequently
with a more or less evident diffuse darker discal shade. Antenne about
as long as the body in the male, entirely piceous, second and third joints
subequal, the fourth slightly longer; intermediate joints three to three
and one-half times as long as wide. Head a little wider than the thorax
in the male, the eyes not very prominent. Thorax moderately trans-
verse, sides parallel, margin fine, not appreciably thickened at the
front angles. Length about 214 mm. to the tips of the elytra.

38 Annals Entomological Society of America |Vol. XII,

Abdominal sexual characters. Male: sixth ventral deeply emar-
ginate, seventh rather broad, strongly produced, passing the sixth by
nearly twice the length of the latter, gradually somewhat narrowed
apically, the tip notched or narrowly emarginate, medially grooved or
impressed and appendiculate each side near the base of the emargina-
tion; last dorsal not produced, broadly rounded at tip.

California. Marin Co., numerous specimens from Muir
Woods, IV-23; Lagunitas, IV-25, and ‘‘Marin Co,” (type <),
all sent by Dr. Blaisdell.

Var. longipennis var. nov.

Two examples (o', @) from Eureka, Cal. (H. S. Barber), are
provisionally separated as a variety of the preceding. In the male the
elytra seem slightly longer than in the typical form, or any other form
known to me; the antenne are a little paler basally, the seventh
ventral of the male narrower, not medially impressed beneath at tip,
the appendix therefore not double, but continuous from side to side.

Malthodes appendiculatus sp. nov.

Fuscous, thorax yellow. Antennz (c’) attaining the tips of the
wings, fuscous, basal joints paler; third joint slightly or scarcely
longer than the second, fourth evidently longer, intermediate joints
four times as long as wide. Head distinctly wider than the thorax,
eyes rather large, separated anteriorly by a distance less than twice
their width as viewed from the front. Thorax moderately transverse,
sides just perceptibly converging posteriorly, margin fine, scarcely
thickened at front angles. Length 314 mm. to tip of wings.

Abdominal sexual characters. Male: sixth ventral deeply broadly
emarginate; seventh rather narrow, passing the sixth by nearly twice
the length of the latter, gradually narrowed for three-fourths its length,
then widening, ascending and forkéd at tip, each fork appendiculate
beneath or emarginate at tip as viewed in profile; last dorsal not
produced, broadly rounded at apex.

California. ‘‘ Mts. near Claremont”’ (Baker); Pasadena.
A single male from each locality, the first named being the
type.

This species is rather closely allied to mollis, but differs
appreciably in the seventh abdominal segment of the male,
which is here narrower, curved upward at its extremity, the
forks more divergent. The eyes are a little more prominent,
the thorax feebly narrowed posteriorly, and there are some
other slight differences which may however be constant. The
genital appendages also appear to be different, but these are
not very clearly visible in mollis. In appendiculatus there is a
long curved central ligula narrower than the seventh ventral,

1919| Californian Species of Malthodes 39

above which it lies, horizontally flattened and notched at tip,
and on either side of this a very slender straight needle like
process.

Malthodes obductus sp. nov.

Brownish testaceous, thorax yellowish, disc clouded with fuscous.
Antenne as long as the body in the male, fuscous, the basal joints
paler; joints two and three subequal, four a little longer, intermediate
joints about three and one-half times as long as wide. Head obviously
wider than the thorax in the male, the eyes only moderately prominent.
Thorax moderately transverse, sides straight and parallel or very
nearly so, side margins fine, not thickened at the front angles. Length
2 mm. to tip of elytra.

Abdominal sexual characters. Male: fifth dorsal segment strongly
produced, arched, overhanging the terminal segments, broader at tip,
which is truncate with rounded angles, strongly carinate beneath
except at apex, which is concave; sixth ventral deeply, broadly emar-
ginate; seventh strongly produced, deeply notched at apex, in profile
bent upward a little at about apical third, the tip again horizontal.

California. Green Point, Humboldt Co., June 4-7 (Blais-
dell); Washington, Seattle (Prof. O. B. Johnson).

The type is a male from the first named locality.

In the typical form the apical notch of the seventh ventral
is as deep as the terminal width of the segment. In the Seattle
males the segment is more deeply cleft, the forks more diverging
and the tip of the produced fifth dorsal is a little different.
These may represent a closely allied species, but as the sexual
modifications are of the same type throughout, I prefer to
consider it a varietal form.

Malthodes bicurvatus sp. nov.

Fuscotestaceous, thorax entirely yellow, head piceous. -Antenne
as long as the body in the male, fuscous, scarcely paler basally in fully’
colored specimens; third joint slightly or scarcely longer than the
second, fourth distinctly longer than the third, intermediate joints:
about four times as long as wide. Head evidently but not greatly
wider than the prothorax, eyes only moderately prominent. Thorax:
moderately transverse, sides straight, finely margined, front angles
not thickened. Length (head deflexed) about 2 mm. to tips of elytra.

Abdominal sexual characters. Male: last visible dorsal segment
broadly truncate, the lateral angles produced downward and backward
in a rather broad process which is dilated and rounded at tip. Sixth
ventral broadly deeply emarginate, the curvature becoming more nar-
rowly parabolic at the bottom of the emargination; seventh ventral
narrow, elongate, notched at tip, bisinuate in profile, with a short
tooth-like process at middle of upper side.

40 Annals Entomological Society of America |Vol. XII,

California. Mokelumne Hill, April (Blaisdell), type 0;
Adams Springs, Lake Co., June 18 (Blaisdell).

Malthodes complicatus sp. nov.

Fuscotestaceous, head piceous, thorax entirely yellow or with the
extreme margin at apical angles infuscate. Antennz longer than the
body in the male, entirely fuscous, second and third joints subequal,
fourth evidently longer, intermediate joints more than four times as
long as wide. Head (co) wider than the prothorax, eyes moderately
prominent. Prothorax moderately transverse, sides parallel and nar-
rowly margined, margin not appreciably thickened at the front angles.
Length (head deflexed) 214 mm. to tips of elytra.

Abdominal sexual characters. Male: last (2) dorsal with a slender
descending lateral process, which is abruptly bent at right angles, the
terminal portion horizontal. Sixth ventral deeply, broadly emar-
ginate, the bottom of the emargination truncate; seventh ventral
received between the descending dorsal processes, elongate, narrow,
forked rather deeply at tip, obliquely ascending in profile in its apical
two-thirds.

California. Pomona, *:March.< 25; (type. o'); Pasadena;
Claremont (Baker); Santa Barbara, Feb. 4; Ojai Valley,
March 5.

Superficially closely resembling the preceding species, but
very distinct by the sexual characters.

Malthodes sericeiventris sp. nov.

Fuscotestaceous, head piceous, thorax entirely clear yellow.
Antenne as long as the body in the male, entirely fuscous, joints two to
four visibly increasing in length, intermediate joints nearly five times as
long as wide. Head (male type) notably wider than the thorax, with
prominent eyes; less wide in a second male. Thorax moderately trans-
verse, sides straight and nearly parallel, side margins fine, not appre-
ciably thickened at the front angles. Length (head deflexed) 244 mm.

Abdominal sexual characters. Male: ventral segments 3 to 5 silvery
sericeous at middle throughout their length; sixth deeply emarginate;
seventh elongate, narrow, deeply divergently forked apically, in profile
obliquely ascending in about its apical half, the tips of the forks
thickened and concavely beveled.

California. Pomona, April 25-May 20;26%, 29;type co.

The sericeous ventral area of the male is quite unique and has
the appearance of an excessively short and dense silvery pile,
but it is not resolvable, as such by a 14 inch triplet, and is
probably due to surface sculpture.

1919] Californian Species of Malthodes 41

Malthodes vapidus sp. nov.

Fuscotestaceous, head piceous, thorax yellow. Antenne entirely
fuscous, as long as the entire body in the male; joints two to four
increasing in length, intermediate joints four times as long as wide.
Head much wider than the thorax, the eyes large and prominent,
separated anteriorly by not more than their own diameters. Thorax
moderately transverse, sides straight and nearly parallel, side margins
very fine, not thickened appreciably at the front angles. Length (head
deflexed) about 2 mm. to tips of elytra.

Abdominal sexual characters. Male: last dorsals not produced;
sixth ventral deeply broadly emarginate; seventh ventral rather wide,
parallel sided, deeply widely divided at apex, sinuately ascending in
profile.

California. Lake Tahoe, July 17-21, ’97 (type &).

Malthodes fusculus Lec. Proc. Acad. Nat. Sci. Phila. V, p. 346.

Le Conte’s description, Synopsis of Lampyridz, Trans.
Am. Ent. Soc., IX, 1881, is as follows:

“Last ventral (oc) bent in a sinuate manner obliquely upwards and
more deeply nicked at tip than in fragilis; prothorax transverse, finely
margined, tinged with testaceous; head black, eyes large, prominent,
antenne long, slender, fourth and following joints longer than the
second or third, o' two-thirds, 2 one-half as long as the body. Length
Som. Cal’

A variety is mentioned with the prothorax ‘‘yellow testa-
ceous 9.”

While probably most closely related to vapidus, this species
is at once separable by the much smaller eyes and consequently
narrower head, which is not very much wider than the pro-
thorax. The last ventral is forked in a similar manner, but the
emargination is much less deep than in vapidus, the depth
being subequal to the width of the segment at the bottom
of the emargination, while in vapidus it is much deeper than the
segmental width at this point. In fusculus the forks comprise
not more than one-fourth the entire length of the segment, in
vapidus about one-half the length of the segment.

The type was taken at or in the vicinity of San Francisco.

Malthodes tularensis sp. nov.

Fuscous, thorax yellowish, more or less tinged with fuscous,
especially along the side margins. Antennz entirely fuscous, as long
as the body in the male; third joint scarcely longer than the second,
fourth evidently longer, intermediate joints four times as long as wide.

42 Annals Entomological Society of America |Vol. XII,

Head evidently wider than the thorax, eyes moderately prominent.
Thorax quite strongly transverse, sides straight and parallel, rather
strongly margined. Length (head deflexed) 2 iv; mm. to tips of elytra.

Abdominal sexual characters. Male: sixth ventral very deeply,
broadly emarginate, the emargination truncate at bottom, seventh
narrow, very elongate, polished, yellow, very sparsely pubescent,
gradually narrowed for two-thirds its length, then widened and deeply
forked; as viewed in profile, horizontal in rather more than basal half,
then rather abruptly sinuately ascending; last dorsal not produced.

California. Tulare Co., June 25,.2¢1; 19. Type o-

Malthodes visceratus sp. nov.

Fuscous or fuscotestaceous, thorax yellow with side margins more or
less infuscate. Antennz entirely fuscous, as long as the body in the
male, third joint but slightly longer than the second, fourth a little
longer, intermediate joints fully four times as long as wide. Head (0)
not much wider than the thorax, narrower than the thorax in the female.
Thorax strongly transverse in both sexes, sides nearly straight and
parallel, strongly margined, the margin thickened at front angles.
Length 2 to 244 mm.

Abdominal sexual characters. Male: sixth ventral broadly, deeply
emarginate, the emargination truncate at bottom; seventh elongate,
deeply, broadly divided at apex, the forks slender, strongly up-curved
basally in profile, then bent suddenly in horizontal direction; sixth (?)
dorsal acutely projecting downward at sides to enclose the last ventral.

California. _ Mill Valley, III-3-’07 (type #); Mt. Tamalpais,
V-1-’13; Green Point, Humboldt Co., VI-5-’'16; Mariposa
Co., VI-16 (W. M. Giffard), all from Dr. Blaisdell.

EXPLANATION OF PLATE III.

All figures show the outline of the sixth and seventh ventral segments of the
male, and beneath them in each case, except Fig. 1, the seventh ventral as viewed
in profile. In complicatus, Fig. 11, is shown also one of the downward projecting
dorsal appendages, between which the seventh ventral lies.

Fig. 1. Terminal ventral segments of Malthodes reflexus and laticollis.
Rigs. 2: Same of magister.

i onee os vigilans.

Fig. 4. . basalis.

1D We “ mitificus.

Fig. 6. “ piceolus.

Meier fc “mollis.

Fig. 8. “ appendiculatus.
Big. 19) “ obductus.

Fig. 10. “ bicurvatus.
Fig. 11. “ complicatus.
Fig. 12. “ sericeiventris.
Fig. 13. “ vapidus.

Fig. 14. “tularensis.

Fig. 15. “_ visceratus.

VOL. XII, PLATE III.

ANNALS E.S. A.

oY
sls
Ly

v
.
-

H.C. Fall

NOTES ON THE REPUGNATORIAL GLANDS OF
CERTAIN NOTODONTID CATERPILLARS.

GLENN W. HERRICK and JoHN D, DETWILER.

In the course of a study which the writers are making of the
life histories and habits of the red-humped apple caterpillar
(Schizura concinna) and the yellow-necked caterpillar (Datana
ministra), the junior author noted that while handling the
caterpillars of the former species, especially when they were
pinched with forceps, a considerable amount of moisture
would suddenly appear in an area about the body of the larva
as it lay in the hand or rested upon the table. Curiosity as to
the source of the moisture was aroused and the following
experiments were made by the writers to determine its origin,
nature, and method of dispersion.

The presence of repugnatorial eversible glands in Notodontid
caterpillars has been known since the time of DeGeer, and
Packard* gives a list of nine species in which the glands have
been found. S. concinna appears in the list and Packard
remarks as follows concerning it.

‘While examining the very gaily colored and heavily spined
caterpillars of Schizura concinna, I observed that when a fully
grown one was roughly seized with the forceps or fingers it sent out
a shower of spray from each side of the prothoracic segment,
exactly like that of Cerura and Macrurocampa.’’ Several years
previously a note from C. S. Denhamf appeared in Insect Life’
in which he said that in rearing a brood of these caterpillars he
‘‘discovered that they had the power to emit quite a quantity
of strong hydrochloric acid strong enough to be decidedly
corrosive to the skin and easily perceptible in the atmosphere.”’

In investigating the source and nature of the moisture which
appeared whenever the Schizura larve were handled at all
roughly it soon became evident that these caterpillars possessed
considerable amounts of the liquid and that they were able to
eject it to a surprisingly long distance and over a comparatively
large area. In determining the habits and behavior of the
different individuals of Schizura concinna in ejecting the liquid

+r. Ne Yo Entesoc., Vole IM pps l0=12 71895:
t Insect Life, Vol. I, p. 148, 1888.

44

1919] Repugnatorial Glands 45

the caterpillars were placed on large sheets of blue litmus paper
and were then irritated at different points on the body by pinch-
ing them with forceps. It was difficult to see the spray on
account of its fine misty quality. Occasionally, by getting the
caterpillar in just the right position between the observer and
the rays of light, one could see a tiny jet or cloud of mist. The
effect on litmus paper, however, was instantaneous and con-
spicuous, for every drop that touched the paper changed it to a
characteristic red color. Moreover, the area covered by the
spray and the distance to which it could be ejected were easily
and accurately determined. The following experiments and
observations on S. concinna illustrate the action of the glands.

A full-grown larva was carefully placed near the center of a
large sheet of litmus paper. While in this position it was pinched
with tweezers on the fifth abdominal segment. At once a
fine invisible spray was thrown out around the body in a
posterior direction. An area surrounding the body about
3 inches wide and 5 inches long was colored solidly red, while
about the lateral and posterior borders of this solidly colored
area was a field covered with drops of various sizes. From
right to left the farthest drops were 5% inches apart.

The larva was pinched again near the thorax, the main
pressure being exerted on the right side of the body. There
was an immediate spray toward the right and in a posterior
direction. The farthest drop posteriorly was 71% inches from
the caudal end of the abdomen, while the farthest drop laterally
was 514 inches from the body. Figure 1 shows the pattern of
the spray on the litmus paper.

An attempt was made to determine the direction of the
spray in relation to the point of irritation. When irritated on
the right side and the head left free, the head is thrown to the
right and the spray is directed largely to the right side of the
body. When irritated on the left side the movement of the
head is in that direction and the spray is directed mostly to the
left. When irritated near the posterior end of the abdomen
there is a decided tendency for the spray to be thrown in a
posterior direction. When pinched just behind the hump on
the second thoracic segment all of the secretion was projected
backward and to the left. In case of this particularly vigorous
larva the drops were thrown approximately 7 inches distant
in a lateral direction from the head and in a posterior direction

46 Annals Entomological Society of America [Vol. XII,

to the distance of 74% inches. Again the larva was touched
lightly on the right side of the body. The head was immediately
turned to the right whereupon a sudden pinch with forceps
caused the liquid to be projected in a fine spray almost entirely
on the right side of the body. In all, this larva was irritated
seven distinct and successive times. It responded to each of
the first six irritations by an ejection of liquid although in a
lesser quantity toward the end, while no response was obtained
from the seventh irritation. Evidently the supply became
exhausted or the larva became fatigued and indifferent to the
irritation.

Sse]
oy

Fig. 1. Spray pattern (1% size), reproduced from litmus paper, of one ejection_of
liquid by a vigorous larva. C, position of caterpillar.

1919] Repugnatorial Glands 47

Since the direction of the spray appeared to correspond to
the direction in which the head was free to move we obtained a
fresh larva and held the head so that there was practically no
movement to the right or left. In response to stimulus under
these conditions the larva was still able to throw a spray of
the liquid to the distance of from 6 to 7 inches, but wholly in a
posterior direction over the entire body. This experiment was
repeated with another larva with similar results. It appears
evident that the caterpillar can direct the liquid backward to a
distance of several inches even though the head be prevented
from free movement to the right or left. Moreover, experiments
show that the larva can project the spray in an anterior direction.

The caterpillars under observation were too far advanced to
determine whether the larve possess functional glands in all of
the earlier instars. A late brood, however, was found in which
the larvze were in the next to the last instar. One of these
larve, after what seemed to be a greater degree of irritation
than usual, did eject a considerable quantity of the fluid, the
farthest drop falling at a distance from the body of about 314
inches. We hope to be able, next season, to make further
investigations of this phase of the subject.

The opening of the gland is a narrow transverse slit on the
ventral side of the prothorax close to the cephalic border of
this segment... The gland is sack-shaped and situated within
the thoracic cavity. It appears to be quite similar to the
corresponding gland in Cerura vinula described by Klemen-
siewicz.* Further detailed description of this gland and its
morphology will be presented by the Junior author in a later
paper.

Regarding the chemical nature of the secretion it is sufficient
to say here that it gives a marked acid reaction with blue litmus
paper and that it has a pungent and characteristic odor that
immediately suggests acetic acid. The subject is under careful
investigation in conjunction with Professor E. M. Chamot of
the chemical department, the results of which will be given in a
later paper. Suffice it to say, the investigations indicate that
the secretion is formic acid. Poulton} has already shown that
the secretions from the similar gland in Cerura vinula is formic

* Verhandlungen d. Zool. Bot. Gesellsch in Wien. Vol. 32, pp. 468-474, 1882.
{ Trans. Lon. Ent., Soc. for 1886, p. 157.

48 Annals Entomological Society of America [Vol. XII,

acid. The effect of the fluid on the skin is such as to cause
burning and itching similar to the effect of the common plant
known as ‘‘nettles.’’ However, this effect is not so noticeable
unless the quantity falling on the hand is of considerable
amount. ;

It is hoped that opportunity may be found next year for
further observations and experirments on these glands in other
members of the genus.

The authors desire to express their acknowledgment to
Miss Ellen Edmonson for her careful reproduction of the
spray pattern in Figure 1.

NOTES ON THE IMMATURE STAGES OF DELTOME-
TOPUS RUFIPES MELS. (Coleoptera, Eucnemidze).*

EpNA MOSHER.

Larve of the family Eucnemide have been somewhat of a
puzzle to entomologists since the first description of a larva
of Fornax by Coquerel in 1856. The first American species
were described by Osten Sacken, from larve collected by Dr.
Horn and an unnamed species from his own collection. The
larvee are found in dead wood, some of them where it is still
very firm, others, including the species under discussion, prefer
wood which is more decayed. Although living in the wood,
they differ very materially from the ordinary type of wood-
bcrang ‘nsects for they have no powerful mandibles such as
these insects are provided with. Instead the head strongly
resembles that of the leaf-mining larvae without having even as
efficient mandibles as they. Schiodte quotes Coquerel as saying
that they are “‘without perceptible organs of the mouth—nay
without a buccal orifice.’’ It has been doubted by some authors,
including Schiodte, that they were able to make their own
burrows and were suspected of living in burrows made by
other insects. The same authors also differed as to the food of.
such larve, considering their lack of mouthparts, and Schiodte
seemed certain that they must feed on xylophagous larve and
pup, since it was quite evident they could not feed on wood.
Some have thought they lived on the sap or “juices of the
wood.”’ Little has been added to our knowledge of the larve
or pupz since Osten Sacken’s paper was written. Since these
larvee seemed to be such an entomological puzzle, I have been
much interested in collecting them from time to time in the
last five years. Altogether three species have been collected,
but only one or two specimens each time, and all my attempts.
at rearing them have been unsuccessful. Mr. H. G. Crawford,
however, found one of these species very abundant at Guelph,
Ontario and succeeded in rearing a number of adults. I am
greatly indebted to him for placing all of his material at my
disposal, and also to Mr. Charles Dury, of Cincinnati, who
identified the species for me as Deltometopus rufipes Mels.

* Contributions from the Department of Zoology and Entomology, Ohio State.
University, No. 56.

49

50 Annals Entomological Society of America [Vol. XII,

LARVA: The larva is depressed, somewhat moniliform, pale
yellow in color, with dark brown on the head, prothorax and
ninth abdominal segment. When full-grown it averages 14—16
mm. in length, or when fully expanded, as much as 18 mm.
The head is subtriangular in outline, heavily chitinized and
nearly all dark brown as shown by the stippled portions in
Figs. 1 and 2. The pale lines seen on either side of the head
capsule (Figs. 1, 2, 3, 4) may indicate sutures, as the head
capsule will break along these lines when softened. If they
are sutures, their homology is doubtful; but as they are furrows
on the ectal surface and ridges on the ental they are perhaps
for the attachment of muscles. The head is much thinner
cephalad and laterad of these lines. Along the cephalic margin
of the head capsule are two small triangular projections and
four similar, but larger ones, along the lateral margin, making
it serrate. The lateral margin also has a prominent blunt
projection caudad of the serrations. The head is inserted into
the prothorax and may move laterad until this blunt projection
strikes the dark brown protuberance on the prothorax. The
head may also be moved dorsad and ventrad, and a prominent
projection like a condyle on the meson of each caudal margin
seems to facilitate these various movements and also prevents
the head from being shoved farther back into the prothorax.
The very dark line extending caudad from each cephalic pro-
jection (Figs. 3 and 4, t) marks the attachment of a strongly
chitinized bar to each surface, which is probably the tentorium.
Between these two bars at the cephalic end of the head is an
opening, the entrance into the buccal cavity. There are no
ocelli present. The head capsule is sparsely covered with very
minute seta. The appendages of the head are all retractile.
‘Their location is indicated in Fig. 3, where they are shown
extruded. Normally only the extreme tips of the mandibles
are visible and then only under high power, as all of these
structures are exceedingly minute. What I have considered
the antenna (Fig. 5) looks very like a maxilla but does not
seem to be in any way connected with the buccal cavity. Its
median projection at the distal end seems to be covered with
sensory structures and the lateral projection ends in at least
three finger-like projections. The structures articulated to the
tentorial bar are certainly the mandibles (Figs. 6 and 7, md).
Each moves in a horizontal plane and has one very long tendon.

1919] Deltometopus Rufipes Melts al

Mesad of these are a group of structures which seem to be
maxille and labium but these can only be seen distinctly under
the oil immersion lens. The median portion, which is pointed at
the end (Figs. 3 and 8, lb) has a longer, sharp projection on
either side which appears to be two-segmented. The dorsal
surface of this structure (Fig. 8) shows a long tube connected
with the median projection which I have traced back into the
prothorax and may be the alimentary canal, although it is very
small and the salivary duct should open here. The tube seems
to have a chitinous lining, however. Laterad of these structures
and slightly dorsad of them are what appear to be maxille.
The distal end bears a cluster of projections which appear to be
arranged in a circle. These structures are only a little less
strongly chitinized than the mandibles. There has been so little
morphological work done on this type of head that one is rather
at a loss to account for their peculiar structure and extremely
doubtful as to the homology of parts.

The entire surface of the thorax and abdomen is very finely
striate, except when otherwise indicated, and all the body seg-
ments are very similar, except that the thoracic segments are
shorter in proportion to their width.than those of the abdomen.
There are no traces of legs or prolegs. Each segment except the
caudal one has some very distinct areas which appear velvety
under a low power lens as if densely covered with fine sete.
On the dorsal surface of the prothorax this area is triangular and
located near the caudal part of the segment but on the other
thoracic segments it is cephalic and somewhat triradiate. The
areas are ovate on the abdominal segments and located near the
cephalic margin. The areas are the same shape on the ventral
surface except that the median projection is lacking on the
mesothorax and metathorax. These areas are really covered
with scales as shown much enlarged in Fig. 11, and a few fine
setz are also present. Caudad of these velvety areas on the
mesothorax and metathorax, on both surfaces, are much coarser
striations than on the remainder of the segment. Near the
caudal margin of each body segment except the prothorax and
the last abdominal segment on both surfaces, and the eighth
segment on the dorsal surface, is a clear, smooth area clearly
outlined by a fine, slightly elevated, brown, chitinous ring.
These are almost semicircular in outline on all segments except
the ventral surface of the eighth abdominal, where it is much

52 Annals Entomological Society of America [Vol. XII,

smaller. The ninth abdominal segment is apparently the last
one, although the portion surrounding the anal opening probably
represents the tenth segment. It is outlined and almost covered
with coarse, triangular spines. The remainder of the ninth
segment is sparsely covered with very minute sete. On both
surfaces the heavily chitinized caudal portion is punctate. At
the caudal end of this segment are two minute spines. The
spiracles are easily visible on the lateral part of the mesothorax
and first eight abdominal segments (Fig. 9). In the spiracular
region are a number of structures which seem to be olfactory
pores. The arrangement is shown on one segment in Fig. 9 and
several very much enlarged in Fig. 10. These seem to be all of
the simple type as shown in McIndoo’s Fig. 19. A few sensory
pores were also located on the head, and what appeared
to be a compound one with several openings was found on
each side of the ventral surface of the prothorax. These show as
a round white spot on the large, brown area on each side of the
meson. Nearly all of the body segments have a small, lateral
projection which seems to aid in locomotion, as the larva moves
on either side as well as on either dorsal or ventral surface.
Pura: The pupa (Figs. 12 and 13) is very much like that of
an elaterid and does not differ in essential details from any
other coleopterous pupa. The entire body, including appen-
dages, is densely covered with very fine pubescence. In dorsal
view (Fig. 12) a small portion of the head is visible and the
thoracic and nine abdominal segments clearly defined. A deep
furrow between the thorax and abdomen allows for considerable
movement. The caudo-lateral angles of the prothorax are
produced for at least one-fourth the total length of the segment.
The first eight abdominal segments bear spiracles but those of
the first, and sometimes those of the eighth, are concealed. The
lateral margin of each abdominal segment is more flattened than
the remainder and separated from it by a slight furrow as indi-
cated by dotted lines. There are many long, soft sete on the
various body segments as indicated in the figure, the row on
the base of the mesothoracic wings being quite prominent. In
ventral view (Fig. 13) it will be seen that none of the head
sutures are visible, the antenne are closely approximated and
mandibles, maxilla and labial palpi are easily distinguished.
The prosternal spine and its groove on the mesothorax are
easily seen and all of the appendages are quite normal in

1919] Deltometopus Rufipes Melts 53

arrangement. A portion of the seventh segment is elevated and
extends as a flap over a portion of the eighth when the body is
curved. The elevated portion containing the anal opening is
probably the tenth segment. Figs. 12 and 138 show a female
pupa. The arrangement of the caudal segments is somewhat
different in the male as shown in Fig. 14. The length of the
female averages 8 mm., the male 6 mm. The color is white
until the color of the developing beetle begins to show when it
appears to be a dull brown.

This species apparently takes three years to mature, as
larve of three different sizes have been taken in the same log
in autumn. The larve are certainly able to bore into wood.
They seem to use the serrate margin of the head to assist
progress and probably hold themselves firmly in a partially-
formed burrow by the lateral protuberances on the segments,
especially the strongly chitinized one on the prothorax. The
burrows are very small and not easily detected, but some of the
wood where they were found did not need much boring as the
larve could almost push their way along. As to their food, I
cannot state positively, but they have always refused any
insect larve placed in the cages, and have lived for at least
eight consecutive months on nothing but wood. In the case
of larve feeding in very firm wood there never seems to be any
““sawdust”’ or frass about, but very frequently a clear liquid is
seen to be expelled from the anus. Even such facts as have
been discovered about the mouth-parts will not entirely solve
the problem.

LITERATURE CITED.

Coquerel, C. Annals de la Soc. Entom. Vol. IV, 3e Serie.

Horn, G. H. Proc. Ent. Soc. Phil. 1 : 43.

McIndoo, N. E. Journ. of Morphology, 31 : 113-132.

Osten Sacken, C. R. Proc. Ent. Soc. Phil. 1 : 112-115.

Schiodte, J. C. Naturhistorisk Tidsskrift, Ser. 3, Vol. 3. (translation, Ann. and
Mag. Nat. Hist., Ser. 3 : 173-212.)

o4

Fig.
Fig.

Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.

Fig.
Fig.
Fig.
Fig.
Fig.

Annals Entomological Society of America [Vol. XII,

EXPLANATION OF PLATE IV.

Dorsal view of larva with all the segments extended to their full length,
as when the larva is in motion.

Ventral view of larva, with some of the segments retracted as they
appear when the larva is inactive and in most alcoholic specimens,
(drawn to slightly smaller scale).

Dorsal view of head showing mouth-parts extruded.

Ventral view of head showing mouth-parts retracted.

Dorsal view of antenna and tendons.

Mandible and tendons.

Showing articulation of mandible.

Dorsal view of labium and maxille.

Lateral view of fourth abdominal segment showing location of spiracle
and sensory pores.

Several sensory pores greatly enlarged.

Arrangement of scales on ‘‘velvety’’ areas.

Dorsal view of female pupa.

Ventral view of female pupa.

Ventral view of caudal segments of male pupa (without setae).

ANNALS E.S. A. VoL. XII, PLATE IV.

Edna Mosher

RESOLUTIONS ON THE DEATH OF
DR. S. W. WILLISTON.

Dr. Samuel Wendell Williston, who was an early member
and fellow of this society and was elected an Honorary Fellow
in 1916, died in Chicago, Ill., August 30, 1918. He was born in
Boston, Massachusetts, July 10, 1852, and had achieved an
almost unique position in American Science for the present
period in that he had attained distinction in at least three
branches of biology.

His interest in entomology was centered mainly in the
Diptera in which group he was an ardent worker, an extensive
contributor and his influence probably second to no other
American entomologist.

Aside from his standing in American entomology his host of
friends in all branches of science will remember him most
affectionately as a man of the finest human qualities and as a
striking example of the possibilities of achievement open to the
American youth of high ideals and devotion to science.

The Entomological Society of America desires to place on
permanent record its appreciation of his earnest and hearty
support of the society, his many and valuable contributions to
entomology, his standing as a representative of the best in
American science and his loyal friendship and manhood.

Committee: HERBERT OSBORN,

TE. PARKS,
Jeu DaAvis:

SAMUEL WENDELL WILLISTON

Honorary Fellow, Entomological Society of America

Courtesy of Sigma XI Quarterly

PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF
AMERICA.

The Thirteenth Annual Meeting of the Entomological
Society of America was called to order by the Secretary in
Room 9, Gilman Hall, Johns Hopkins University, Baltimore,
Md., at 1:20 Pp. mM., December 26, 1918.

The President and Vice-Presidents being absent, Dr. T. J.
Headlee was elected President pro tem. He appointed the
following committees:

AUDITING: H. A. Gossard, G. A. Dean, F. M. Gibson.

NOMINATING: W. E. Britton, W. A. Riley, J. M. Aldrich.

RESOLUTIONS: Herbert Osborn, T. H. Parks, J. J. Davis.

The Chair also appointed the following temporary members
of the Executive Committee, to replace absent ones: Jas. G.
Needham, E. P. Felt, J. G. Sanders, A. F. Burgess, E. D. Ball.
The meeting then adjourned until 9:00 A. m., December 27.

December 27, 1918.—The meeting was called to order at
9:00 A. M. by President pro tem Headlee, in the same room
as before.

The Executive Committee submitted the reports of officers,
and the following report of their own:

Your Executive Committee has elected within the year the following
members:

On May 1:

Pror. Mario Bezzi, Turin, Italy.

E. H. BLacKMorg, Victoria, B. C.
E. L. CHAMBERS, Seville, Ohio.

T. N. HusBett, Ann Arbor, Mich.
R. F. Hussey, Ann Arbor, Mich.

On December 26:

O. G. Bascock, Dallas, Texas.
A. J. BASINGER, Columbus, Ohio.
H. M. BrunprettT, Houston, Tex.

A. N. CAUDELL, Washington, D. C.

A. G. DusTANn, Fredericton, N. B.

J. E. Ever, Camp Sherman, Ohio.

W. O. HoLuistEer, West Lafayette,
Ind.

A. C. Kinsey, Forest Hills, Mass.

H. P. Kiorz, Woodmere, Long Id.

Wo. M. MAnn, Washington, D. C.

J. B. PARKER, Washington, D. C.
T. H. Parks, Columbus, Ohio.

FRED W. Poos, JR., Charlottesville, Va.

57

Jos. C. OUELLET, Outremont, Que.
LADISLAV SASKO, Chicago.

FRANK N. WALLACE, Indianapolis, Ind.
Wo. LUNDBECk, Copenhagen, Denmark.

Tuos. E. B. Pore, Milwaukee, Wis.

Max H. RuUHMANN, Vernon, B. C.

RAYMOND C. SHANNON, (Life Member),
Washington, D.C.

RatpH M. SmitH, Twin Falls, Idaho.

OLIVER I. Snapp, Agr’! College, Miss.

Wo. B. TuRNER, Hagerstown, Md.

R. H. VAN ZWALENBURG, Hagerstown,
Md.

Don C. WARREN, AUBURN, Ala.

L. P. WEHRLE, Ithaca, N. Y.

A. D. WHEDON, West Chester, Pa.

Total, 32.

58 Annals Entomological Society of America [Vol. XII,

We have re-elected Herbert Osborn as Managing Editor of the
ANNALS, and have elected the following members of the Editorial
Board: J. G. Sanders, Wm. M. Wheeler, E. M. Walker, to serve for
three years. T. D. A. Cockerell has resigned.

We have re-elected Nathan Banks and A. D. MacGillivray to succeed.
themselves in the Thomas Say Foundation.

We beg to submit the following recommendations:

First, that in future a time limit of fifteen minutes be placed on
papers in our programs, subject to extension by vote of the Society at
time of reading.

Second, that one year’s dues be remitted to each of our members
who has served in the present war in the army or navy of the United
States or an allied power; provided, that if such service in any case has
been for a longer period than one year, then the remission shall extend
to the full period.

Third, that the dues of M. C. Tanquary be remitted for the three
years in which he was engaged in Arctic exploration as a member of the
Crocker Land Expedition, as a recognition of his gratuitous services to
science.

T. J. HEADLEE, Chairman.

The report was adopted, including the recommendations.
The accompanying reports were as follows:

REPORT OF THE SECRETARY.

The year’s record is distinctly encouraging. Our membership has
almost held its own in spite of war; our permanent funds have nearly
doubled, including noteworthy gifts from two of our oldest and most
honored members; our current income is but slightly below that of
last year, which was the largest in our history; and the ANNALS has
withstood the rising cost of printing and maintained its high reputation.

The following members have died since the last report:
S. W. Wirutston, Honorary Fellow.
FREDERICK KNaB, Fellow and charter member.
W. HacurE Harrincton, charter member.
C. A. Hart, charter member.
Tuomas P. Lucas.
VERNON KING, aviator, killed in action.
V. A. E. DAEcKE, charter member.

The following have resigned:

H. E. CRAMPTON F. E. WATSON
M. M. Hicu M. A. YOTHERS
H. E. ScHRADIECK S. MARCOVITCH

Thirty-five have been dropped for non-payment of dues.
On Dec. 23rd the membership was in the following classes, as far as
the payment of dues is concerned:

@

1919] Proceedings of the Baltimore Meeting 59

(American Members, including Hawaiian.)

Hyatt ClO stegl OS Bree ne eos eM fer cicay- os oho cv oneiedeee Bingen 432
Onvinesiomonenyeatin: te sci. oe cccis chee anne 63
Owine fom twolyeanrs Ommore...... 50.5. -nades nee: 46
(Foreign)
Raidetorul GS ta men my emer: cial e s,s oc a ee ee eet 12
Owing fomone Ormloneyears:..., 0... 5.4. see eee 20
“TNOWIEN | ad) el oh Ae at aS Se nC ote a 6 Bie.c 4 573
Pleckedsthisumeetine. aay. se. a5 vsco ale see eee 21
Actualipresent membership:........5...s2sseee: 594

One year ago I reported 386 members fully paid up, which was
probably the highest record for the Society at the time; we now have
(counting foreign) 444, which shows how splendidly the members have
rallied to our support in war time.

In sending out the annual statements, the Secretary took occasion to
suggest that members, nearly all of whom in the United States are now
holding bonds of the government, might deposit a fifty-dollar bond in
payment for life membership. Four members did this, and two hon-
orary fellows, J. H. Comstock and E. A. Schwarz, each donated a bond
of the same amount. Other members indicated an intention, and some
a hope, of becoming life members in the near future. It is desirable to
keep this matter before the attention of members until after the next loan
is subscribed. The result has been encouraging, but should be larger.*

In October, when much of the business of the country was depressed
by the wave of influenza, the Secretary received a circular from the
Permanent Secretary of the American Association for the Advancement
of Science, in which it was announced that the plans of the Association
for the Baltimore meeting were much embarrassed, by reason of an
unexpected shortage of meeting rooms in Baltimore, the prevailing
influenza epidemic, the general war conditions, and the railroad and
hotel congestion. This circular suggested that affiliated societies cut
down their meetings to the transaction of necessary business and the
consideration of problems urgently important for the winning of the
war. In view of the fact that the Association of Economic Entomol-
ogists, our sister society, might be expected to take care of urgent
entomological problems, the Secretary, in transmitting copies of the
circular to members of our Executive Committee, took occasion to rec-
ommend the abandonment of our program as our logical duty in the
premises. The Executive Committee immediately agreed, and announce-
ment was made of this change from our regular policy. Later develop-
ments, especially the armistice and the cessation of influenza, so far
relieved the situation that I believe we all wish we were to have a program.
The decision of the Executive Committee, however, should be viewed from
the standpoint of mid-October. No precedent has been established,
and the suggestions adopted by our present business session will without
doubt operate to give us a better program for next year than we have
usually had. Respectfully submitted,

J. M. Avpricu, Secretary.

* Since the meeting E. D. Ball, G. C. Crampton and J. F. Illingworth have
become life members.

%
60 Annals Entomological Society of America [Vol. XII,

TREASURER’S REPORT.
CURRENT ACCOUNT.

RECEIPTS.
Balance.on anc @Amnall seyexala O10) bere $ 303.99
Dues of Members. eee or er oe ee ees 914.56
From! Manacine th ditto of Arata sian se sn ees spore trees 485.89
Interest ion CuTnrenity account aeser eric mite mice tiie rerienerreneice 8.49
Exchangevomichecks: see ae ee sere eee Re Aisne 40
TOGA sis cache Nad See Re eae EIS LL ORO:
DISBURSEMENTS.
Annalssior December, Giles a ete ee eee sores $ 303.51
Eneraving tor Aamalsine. 5. ce Sash Gus eo Ae er sjacte sn iter cere tesa 100.27
Clérical worl omAnmals.t:c2 of SW. oak Panivc oP See oe oes 46.80
Printing); Secretary: s) office: t,.0 7 Savane Ne tel. Teac eee a see eee esen: 27.25
Postagevandstationeny, Sames.cs5 5c ceeeep eee eee eaeeee 23.00
Clericalrassistance samen. 52 a:.0 4c, aiauoces 7 ocuas sense aces ees 37.00
One membership diplotmilan aaa eae oe ses cis cee eine cee .25
Balance in banks (excluding $2.74 in permanent fund)........... 1,171.25
Mota sone Y, Pats ate eats Breas bay SER) Stns ee hs Pavesi ees ate $1,713.33

Our present large balance is due to the fact that we have not yet
received bills for the last four numbers of the ANNALS.

PERMANENT FUuNDs.

Onvhand last‘reporta(Annalss Del 102).ia2 ca. 2 cece es sees ss PoLOnUO
Kugene Amandius Schwarz, gibtyss.-2 4. 52a.0a-42605- 8855s qe are ee 50.00
Johmblcnry; Comstock: cuts. nee aise icine cee neeoe 50.00
\iiane 10s ID Enis. betes seavesomloferlabhoys -o5u5 pd ooanopsouanosudeneondencoe 50.00
Howard Notman elite membership.).>..45.5.4.2.2.02009o08058 a. 50.00
Rv€aShannons dhifemembérshipis .. 6.8 ssc. s. oasis cdo saben 50.00
TEM Mdrich, lifesmembership, 2. scene aasen aera ane ek 50.00
Interest. Cleveland’ inust-Cox .5. 525%. acto eee ee Seen: LTO2
IniteresStion"pondse a sowwe pecan tere cote ee eae ee ee $3.70
Wess accruédiinterest- paides2: ).2.. 65.55.05 ssh. ase eser 1.60
—— 2.10
Increase in value of War Savings Stamps since purchase.......... 40,
WOT Ae Ss sc, SAPO Pee hank Cie are a ae tee $586.58

SECURITIES HELD. |
Government Bonds—

First Liberty Loan converted—No. 414,847................... $ 50.00
Second Liberty Loan converted—No. 1,784,246............... 50.00
Third Liberty Loan:

ING 2 DSS OGD ile cae chi tesrtns c ieita, chats, axa dens toate auarelnetel die eee 50.00
INO FZ ODS OB OIE te tte cect Ue. n Tecal af ANN AUN ecie iiete eles sae ae 100.00
INOUE TS 7 las bestia Rigs cine nbinidiabis of Sra ord lea unis SRE ate 100.00

Fourth Liberty Loan:
No ASGlasi0024 5 scsi cae ols aia Sa A aes roe stone oe cee 50.00
IN Koy 151] ee: Sh ee ee ee ir rane OL Ci am ec mae 50.00
ING: 490454 oe yon ck easel os; . ciate os Hee cra ee Regs ge ties ae 100.00
Total 85 meen eat teste Bee eee ed aed oe ceo
War Savings Stamps, present value, eight Stamps, at $4.28........ 33.84
Cash to balance, carried imicurrent) account... ..s4.ceese+ es see oe oe 2.74

1919} Proceedings of the Baltimore Meeting 61

All the permanent funds, it will be noted, have been invested in
government securities, except a balance too small to buy a War Savings
Stamp.

Respectfully submitted,

J. M. Atpricu, Treasurer.

REPORT OF THE MANAGING EDITOR.

I am happy to report that in spite of the unsettled conditions of the
year we have been able to carry the ANNALS through without, we
believe, any very vital loss. Aside from delays in the issue of the June
and September numbers due to labor conditions, our schedule has been
maintained and the volume of 450 pages with an interesting variety of
contributions will compare favorably with preceding volumes.

A number of authors deserve our thanks for their generosity in
contributing toward the cost of using plates and the printers deserve
credit for untiring efforts in the face of many difficulties.

The receipts of this office show a substantial gain over those of last
year. The number of outside subscriptions has been somewhat increased
and the sale of back numbers and reprints with contributions toward
plates giving a favorable balance.

With the return of normal conditions and with a goodly lot of papers
already in hand for the next volume, I believe we may feel assured of a
good showing for the coming year.

RECEIPTs.
SUDSCHPLIONS VEE Sram ee ie Melee, Sonera eds $275.88
IRNGTOSCIANS Cages a A ak cP ee SO eats eRe SO a See 138.30
Ba ckeaValemnesermy a etcnsrt seston zis fiieiecar. ep HAE Ele ae 93 66
PING tellers nice Dare PEIN Sa AN ane Seca! $507 . 84
DISBURSEMENTS.
Syichany os) Guarcl 2A (OL IDs syolnaseeege eae me code bobo meen oe $ 19.70
Workewrapping Annalsic).. sh sicas cccess es ers dec oe 2.25
IBATGALOMMTEASLETA tah Gy 7s ictroees ae Seeisiceeteeh ernest 485.89
MROtaeey era er eae tees oe Ss ee a Ah A IE oe. $507.84

Respectfully submitted,
HERBERT OSBORN, Managing Editor.

REPORT OF TREASURER OF THOMAS SAY FOUNDATION.
(Copy not on hand at time of going to press.)

This concluded the reports of officers. The reports of the
Resolutions, Auditing and Nominating Committees were then
presented, as follows:

62 Annals Entomological Society of America [Vol. XII,

REPORT OF THE COMMITTEE ON RESOLUTIONS.

Your Committee on Resolutions begs to report as follows:

That the Society recognizes with a special sense of loss the death of
our lamented honorary fellow, Dr. S$. W. Williston, and recommend that
a special memorial be prepared for publication in the ANNALS.

That we learn with sorrow of the loss by death of our fellow, Fred-
erick Knab, and of charter members, Chas. A. Hart, W. Hague Harring-
ton, V. A. E. Daecke, and members Vernon P. King and Thomas P.
Lucas.

That we desire to accord special praise to those of our members
who have courageously served in the war and with profound sorrow the
deaths of those who have lost their lives in this service.

That we extend the thanks of the Society to the officers of Johns
Hopkins University for the privileges afforded in connection with
meeting places and other favors during the present meeting.

HERBERT OSBORN,
TEL PARKS,
Je Jo DaAvis.

REPORT OF AUDITING COMMITTEE.
Dec: 26; 1918:

Your Committee has examined the records, vouchers and property
reported by Secretary-Treasurer J. M. Aldrich and Editor Herbert
Osborn of the ANNALS, and find the same to be correct and properly kept.

H. A. GossaArD,
Gro. A. DEAN,
Frank M. GIBsSon.

REPORT OF NOMINATING COMMITTEE.

The* Nominating Committee beg leave to nominate the following
officers for the coming year:
President—Jas. G. NEEDHAM.
First Vice-President—]. W. FoLsom.
Second Vice-President—R. V. CHAMBERLIN.
Secretary-Treasurer—J]. M. ALDRICH.
Additional Members Executive Committee—F. E. Lutz, ARTHUR
Gipson, G. A. DEAN, G. C. CRAMPTON.
Respectfully submitted,

H. A. GOSSARD,
W. A. RILEY,
J. M. ALpRIcH.

1919} Proceedings of the. Baltimore Meeting 63

On motion, the report of the Nominating Committee was
adopted, and the Secretary instructed to cast the ballot of the
Society for the officers nominated; which being done, the
President pro tem declared them duly elected.

Under general business, Dr. Headlee moved that it is the
sense of the meeting that each annual program should have one
session devoted to a symposium on some general entomological
topic. Seconded and carried.

Professor W. A. Riley proposed an amendment to the
Constitution, which was allowed to lie upon the table until the
next annual meeting, as required by the Constitution.

At 10:05 a. M. the meeting adjourned.

J. M. ALDRICH, Secretary.

b it : j nabs Va tN oom hy yi ro) yy oB q ;

i

i 4

8.54 4 Usk
-
Li ’ ,
'
ri; 5
i ‘ Ve b () Vihar

; ThE ewer
" ie ey ? he ri. Ts
*

na pried ‘tin Benet spi Nag SL bute we
21 Sage je EShea9 Bf ahs

Vageh t ‘ ae
; Ad ogite han a

cet est gat iM wip at a se s

fa one ane

ares) ic wr.

4 i "Beant —

a zAIRIOOe ol a sas Te: Te

5
a ‘ |
z ~
’
co
«
cr
“
)
ww :
x
. _
© 1s, -
!
= !
- | 2 :
y
,
’
4 re
=
*
:
i
“a é
i - : i
{ .
ro
uf Loe i
“
’ |
%
4
‘ ¥ J
ra
i
ik sf
»
oo ‘
H '
+ by if -
in ty
“ x : 7 ‘
aa ¥ et th Gy al
4 ‘> i 1 at sh

NOTICE TO MEMBERS AND CONTRIBUTORS.
\

The Annals of the Entomological Society of America, pub-
lished by the Society quarterly, includes the Proceedings of the
Annual meetings and such papers as may be selected by the
Editorial Board.

Papers may be submitted to any member of the Editorial
Board and should be as nearly as possible in the form desired as
final, preferably typewritten, and illustrations must be finished
complete ready for reproduction. Plates must not exceed 5x 7
inches unless intended to fold. In general, papers to be accepted
must be original, complete and previously unpublished and,
except in connection with the proceedings, it will not be the
policy to publish preliminary announcements or notes. Authors
will be allowed fifty reprints gratis and additional copies at cost
to the Society.

The Managing Editor is provided with the most recent
address of all paid-up members on record in the Secretary’s
office for mailing the numbers of the Annals and members failing
to receive their numbers should present their complaint to the
Managing Editor within four months from the date of the mail-
ing of the issue. After that time the numbers will be furnished
only at the regular published rate.

The regular annual subscription price for the ANNALS is in
the United States, Cuba, Porto Rico, Hawaii. and Mexico, $3.00;
Canada, $3.50; other countries, $4.00. Checks, drafts or money
order should be! drawn payable to ANNALS ENTOMOLOGICAL
SOCIETY: OF AMERICA, and addressed to HERBERT OSBORN,
State University, Columbus, Ohio, U.S. A.

Requests for information as to membership and the annual
dues of members may be sent to the Secretary-Treasurer,
J. M. ALtpricu, National Museum, Washington, D. C.

Communications relating to the ANNALS, and all orders for
separate copies or reprints should be addressed to
HERBERT OSBORN, Managing Editor,
ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA,
State University, Columbus, Ohio.

CONTENTS OF THIS NUMBER.

VAN Dyxk#, Epwin C.—The Distribution of Insects in

Western North America (6a. 2; iL ay oe RE aor i
BRUES, CHARLES T.—A New Chalcid-Ay Parasitic on
the Australian Bull- dog Amt eo at ae 13

ALEXANDER, CHARLES P.—Notes on the Crane-flies-

of the Hawaiian Islands (Tipulide, Diptera): .-.. 25 |

Fait, H. C_—The Californian Species of Malthodes... 31

HERRICK, GLENN W., and DETWILER, JoHN D.—
Notes on the Repugnatorial Glands of Certain
Notodontid’Caterpitars so) hs SF en ee 44

MosHER, Epna—Notes on the Immature Stages of
Deltometopus Rufipes Mels. (Coleoptera, Eucne-
BTU) ho NS RN Se yi oo ae ty eo A ga 49

- ALDRICH, J. M.—Proceedings of Baltimore Meeting... 57

: Price List of Publications.
Annals, Vols. I, II, III, IV, V, VI, VII, VIII, IX, X and XI complete, each. $3.00

Annals, Separate Parts except as below, each..........00 ccc ccc cece ceetess 1.00
Annaits, Vols, EF and ‘IT, Part 3, eden wey ae RO ee ae len ae SOOO tte
2 Annals, Vol AV, Part-4) each. 220 0 So te Br a) Se ies Sea Win canes 1.50 y

BACK VOLUMES

Of the ANNALS OF THE ENnTOMOLOGICAL SociETY oF AMERICA may
be secured from the office of the Managing Editor and new members
of the Society who may wish to complete a set are advised to secure
the earlier volumes while there is still a supply on hand and the price
is kept at the original subscription rate.

Address HERBERT OsBorn, Managing Editor,
ANNALS ENTOMOLOGICAL SOCIETY OF AMERICA,
State University, Columbus, Ohio.

fi Number 2.

Volume XII. |

aa ANNALS
“: The Entomological Society of America

em “ MY
PO solan (ng e/a
FS

aes ) JUNE, 1919 /s#

EDITORIAL BOARD

\

HERBERT OSBORN, Managing Editor,
CoLUMBUS, OHIO,

LO. HOWARD, WM. A, RILEY,
; WASHINGTON, D. C, MINNEAPOLIS, MINN.

VERNON L. KELLOGG, FRANK E. LUTZ,
STANFORD UNIVERSITY, CALIF. NEw YorE City, N. Y.

WM. M. WHEELER, WM. S. MARSHALL,
LR Boston, Mass. Mapison, WIs.

“> B, M) WALKER, J. G. SANDERS,
ue TORONTO, CANADA. HARRISBURG, PA.

a" PUBLISHED QUARTERLY BY THE SOCIETY
COLUMBUS, OHIO

Mi : Entered as second class matter April 11, 1908, at the Post Office at- Columbus, Ohio,
! under the Act of Congress of March 3, 1879.

Haven ide eed
Bree y. a rere Ghee oe ea i .

The Entomological Society of America
Founded 1906.

OFFICERS 1919.

President
James G. NEEDHAM d ; : : .« vdthaea; NY.

First Vice-President
James W. Fotsom . : 4 : 5 : . Urbana, Il.

Second Vice-President |
R. V. CHAMBERLIN . 5 : ; : . Cambridge, Mass.

Managing Editor Annals
HERBERT OSBORN. ; i : } . . Columbus, Ohio

Secretary-Treasurer
J. M. Atpricw : . National Museum, Washington, D. C.

Executive Commitiee

THE OFFICERS

AND
SUEZ; ARTHUR GIBSON,

F.E
G. A. DEAN, G. C. CRAMPTON.

Commitiee on Nomenclature

E. P. Fett, T. D. A. CockERELL, NATHAN Banks.

| Thomas Say Foundation
NATHAN BANKS, | A. D. MAcCGILLIVRAY, P. P. CALVERT;

E. B. WILLIAMSON,
J. M. Atpricu, Editor, E. D. Batt, Treasurer.

Committee on National Museum

T. D. A. CocKERELL, Chairman,

HERBERT OSBORN, Wm. BARNES, . Wu. M. WHEELER;
James G. NEEDHAM.

77x\\s0Nlan Inoys

a
Ve SU
\ W, /
Nee “ona Mysev™ Na ‘
ANNALS Tp. oa ha

OF

The Entomological Society of America

Volume XII JUNE. 1919 Number 2

THE OLFACTORY SENSE OF LEPIDOPTEROUS LARVZ.

N. E. McInpboo, Ph. D.,
Insect Physiologist, Deciduous Fruit Insect Investigations,
Bureau of Entomology, Washington, D. C.

INTRODUCTION.

A thorough investigation of the above subject will undoubt-
edly throw some light upon how plants attract insects, because
as yet no one has shown experimentally that lepidopterous
larve can smell and the olfactory organs have never been
described. Whenever it is proved that insect larve respond to
chemical stimuli, and whenever organs suitable for the reception
of these stimuli have been found, then we can intelligently ask
why do the cotton caterpillar (Alabama argillacea Hibr.) and
cotton boll weevil (Anthonomus grandis Boh.) feed exclusively
on the cotton plant?; or why does the silkworm (Bombyx mori
L.) feed almost exclusively on mulberry-tree leaves?. Many
more similar examples could be given by any entomologist, but
why is it that some larve are very selective in regard to their
food while others show little or no preference between members
of a large list of plants. Also, how does a female lepidopteron
distinguish the best or only suitable host plant for her progeny
so that she can deposit her eggs on this particular plant? The
only plausible answer for all of these questions is to suppose
that plants, as well as animals, emit odors, and that insects in
searching for food,. either for themselves or for their young
progeny, are guided by the odors emitted by the plants. In
order that the odors of plants may differ, it may also be assumed
that the various chemical constituents of plants emit particular
odors and that the odor emitted from a plant may be a com-
bination of all the odors from the various constituents, or pos-
sibly one odor might be so strong that it masks all the others.

65

. . 1
\ 1

\3 4

66 Annals Entomological Society of America [Vol. XII,

Upon this hypothesis, the odors from plants would vary accord-
ing to the number, combination and quantitative percentages of
the various constituents. Reasoning along this line of thought,
we may be able to explain why a few insects have only one host
plant; why many have a preferred host plant, but will eat
other allied plants; and why others eat a large number of
plants. If we could positively answer the above questions, we
might be able to devise practical methods for the control of
certain insects, as by trap baits, etc.

The present investigation will show that lepidopterous
larve respond to chemical stimuli and that they have organs
suitable for receiving these stimuli; and the following review of
the literature will indicate how insects are attracted by certain
host plants.

Verschaffelt (1910), experimenting with the cabbage-butter-
fly larve, Pieris brassice and P. rape, found that they are very
fond of the cultivated species of the Cruciferee and that in cap-
tivity they ate leaves from 15 indigenous species, representing
14 genera, of the same family. They did not eat all of these
species, however, equally readily, and refused to eat species
belonging to 17 other families, but did attack Trope@olum
and Reseda which belong to two other families. Chemists have
determined that all of the plants eaten by the above larve
contain mustard oils. To determine whether these larve
could be induced to eat leaves not ordinarily attacked, leaves
of Apios tuberosa were smeared with the juice from the leaves
of a crucifer (Bunias orientalis); these leaves were at once
attacked and in a short time devoured. Wheat flour and corn
starch, which when dry or moistened with water are rejected
by both Pieris-larve, are eaten with avidity when soaked with
Bunias-juice. The larve behave in a similar manner toward
filter paper saturated with Bunias-juice.

To ascertain the exact constituent in the plants and juices
which attracted the larve, Verschaffelt wet leaves of Apzos
tuberosa and Rosa (two species not containing mustard oil) with
a fairly strong solution of pure sinigrin (potassium myronate),
the glucoside of black mustard; these leaves were eagerly eaten
by the larve. He says: ‘It is clear that Pzeris-caterpillars seek
out various mustard oils, just like the various glucosides derived
from them, They are clearly attracted by the whole group of
substances.”

1919] McIndoo: Olfactory Sense 67

vd

RESPONSE TO CHEMICAL STIMULI.

While feeding silkworms it is easy to determine that they
can smell; this is done by placing fresh mulberry-tree leaves
near them so that they can neither see nor touch the leaves;
immediately the silkworms move their heads, work their
mouth parts and begin crawling toward the leaves, and some-
times a hungry silkworm for several minutes will follow a leaf
dragged slowly just in front of its head and the writer believes
that the larva is guided solely by means of the odors emitted
from the leaf rather than by seeing the leaf. Silkworms may also
be induced to eat other leaves which they will not ordinarily
even “‘taste.’’ This was proved by the following test: Leaves
from peach trees, cherry trees, plum trees, and apple trees were
dipped into the juice from mulberry-tree leaves and were then
fed wet to silkworms; all of these leaves were eaten, although
some of them apparently were not really relished. When other
leaves of the same trees were dipped into water, the silkworms
did not even “‘taste’’ them.

To test the responses to other chemical stimuli the various
larve used were confined in small observation cases and the
sources of the odors were usually kept in small vials which were
-held directly beneath the larva in the case. The following larve
were used in the experiments: Tent caterpillars, fall web-
worms, tussock-moth larve, army worms and larve of a.
butterfly (Papilio polyxenes). The following sources of odors
were employed and the average reaction time of the above
larve to them are: Oil of peppermint, 14.6 seconds; oil of
thyme, 9.5 seconds; oil of wintergreen, 17.6 seconds; dried
leaves of pennyroyal, 20.1 seconds; dried leaves of spearmint,
22.9 seconds; wild cherry-tree leaves, 42.5 seconds; fresh grass,
19.1 seconds, old honey and comb, 51.5 seconds; protruded
thoracic glands of above Papilio larve, 22.8 seconds; and as a
ccontrol—a clean and empty vial, 60 seconds (totally negative).
The details pertaining to these experiments are as follows:

In each set of experiments 10 larve were used, one larva
being confined in a case. As a rule, the more the larve were
handled the more satisfactorily they responded to odors, but
generally speaking these larve were the most unfavorable
insects that the writer has ever used for testing the responses to
odors. When placed in the experimental cases, all of them,

68 Annals Entomological Society of America [Vol. XII,

except the army worms, had the habit of crawling into the
corners and lying there more or less dormant and in such a
position would never respond to odors; therefore, before testing
a larva it was first necessary to see that it was lying flat on the
bottom of the case, and in order to insure, if possible, a response
to another odor after an interval of 15 minutes, it was usually
touched with a pencil to ‘“‘awaken it from its stupor.’”’ Caterpil-
lars in their webs rarely responded to the odors used. The fol-
lowing records include only the first responses and their reaction
times:

MEDIUM SIZED TENT CATERPILLARS (Malacosoma americana Fab.).

Oil of peppermint:
6 moved head slightly. 1 raised head and crawled away.
1 moved away slowly. 1 moved head and worked mouth parts.
1 raised head quickly.
Reaction time 3 to 15 seconds, average 7.5.seconds.

Oil of thyme:
6 moved head slightly. 1 raised head and moved away slowly.
3 moved away slowly.
Reaction time 3 to 15 seconds, average 5.4 seconds.

Oil of wintergreen:
5 moved head slightly. 2 raised head and moved away slowly.
2 showed no response. 1 moved away slowly.
Reaction time 4 to 60 seconds, average 20.6 seconds. In all cases 60
seconds was regarded totally negative.

Dried leaves of pennyroyal:
5 moved away slowly. 1 moved head sidewise.
3 moved head slightly. 1 showed no response.
Reaction time 38 to 60 seconds, average 22.2 seconds.

Dried leaves of spearmint (odor very weak):
5 showed no response. 2 moved head slightly.
2 moved away slowly. 1 moved leg back and forth.
Reaction time 5 to 60 seconds, average 38.3 seconds.

Wild cherry tree leaves cut into small pieces (their favorite food):
4 showed no response. 1 moved away slowly.
4 moved head slightly. 1 raised head slowly.
Reaction time 5 to 60 seconds, average 42.5 seconds.

Old honey and comb (odor very faint):
6 showed no response. 4 moved head slightly.
Reaction time 20 to 60 seconds, average 51.5 seconds.

Empty and odorless vial (used as a control):
10 showed no response.

SMALL Fatt WEBWoRMS (Hyphantria cunea Dru.)
Oil of peppermint:

3 moved slightly. 1 moved head sidewise.
2 raised head and thorax quickly. 1 turned around.
2 worked mouth parts. 1 raised head.

Reaction time 2 to 40 seconds, average 11.8 seconds.

1919] McIndoo: Olfactory Sense
Oil of thyme:
4 raised head quickly. 1 moved away quickly.
3 moved head and worked 1 raised thorax quickly.
mouth parts.
1 moved slightly.
Reaction time 2 to 10 seconds, average 4.5 seconds.
Oil of wintergreen:
3 moved head slightly. 1 worked mouth parts.
2 raised head quickly. 1 moved head sidewise.
2 moved away quickly. 1 raised head and thorax slowly.
Reaction time 2 to 30 seconds, average 7.5 seconds.
Dried leaves of pennyroyal:
5 moved slightly. 1 moved head slightly.
2 raised head and thorax slowly. 1 moved head and thorax sidewise and
1 moved away slowly. wire-screen bottom of case.
Reaction time 2 to 25 seconds, average 7.4 seconds.
Dried leaves of spearmint (odor very weak):
4 moved slightly. 2 moved away quickly.
2 raised head quickly. 1 moved head slightly.
1 moved slightly.
Reaction time 3 to 15 seconds, average 6.4 seconds.
LARGE Tussock-MotH LARVAE (Hemerocampa leucostigma S. and A.)
Oi! of peppermint:
6 moved slightly. 1 showed no response.
2 moved away. 1 raised head and thorax quickly.
Reaction time 2 to 60 seconds, average 17.8 seconds.
Oil of thyme: '
$8 moved slightly. 1 showed no response.
1 raised head.
Reaction time 2 to 60 seconds, average 13.1 seconds.
Oil of wintergreen:
3 raised head. 1 moved head sidewise.
2 moved slightly. 1 moved away slowly.
2 jerked head backward. 1 showed no response.
Reaction time 2 to 60 seconds, average 16.3 seconds.
Dried leaves of pennyroyal:
4 moved away slowly. 2 raised head.
3 moved slightly. 1 showed no response.
Reaction time 2 to 60 seconds, average 23.2 seconds.
Dried leaves of spearmint (odor very weak):
6 moved away. 2 showed no response.
2 moved slightly.
Reaction time 5 to 60 seconds, average 30 seconds.
Protruded thoracic glands of larvae (Papilio polyxenes).
5 moved slightly. 2 showed no response.
2 moved away slowly. 1 raised head.
Reaction time 2 to 60 seconds, average 22.8 seconds.
LARGE ArMy Worms (Cirphis wnipuncta Haw.)
Oil of peppermint:
5 raised head quickly. 3 raised head quickly and waved

2 raised head ‘slowly. sidewise.
Reaction time 2 to 5 seconds, average 2.9 seconds.
Oil of thyme:
4 raised head quickly and moved 3 raised head quickly.
sidewise. 3 raised head slowly.
Reaction time 2 to 5 seconds, average 2.5 seconds.

69

bit

it

70 Annals Entomological Society of America [Vol. XII,

Oil of wintergreen:
5 raised head quickly and moved 2 raised head slowly.
it sidewise. 1 raised head quickly.
2 raised head quickly and moved
backward.
Reaction time 2 to 10 seconds, average 2.9 seconds.

Dried leaves of pennyroyal:
8 raised head slowly. 1 moved away slowly.
1 raised head quickly.
Reaction time 2 to 10 seconds, average 4.2 seconds.

Dried leaves of spearmint (odor very weak):
6 raised head slowly. 1 raised head slowly and waved it
3 raised head quickly. sidewise.
Reaction time 2 to 4 seconds, average 2.7 seconds.

Fresh grass in vial (food of these larve):

2 raised head slowly. 2 moved backward slowly.

2 moved slightly. 1 moved forward quickly.

2 showed no response. 1 raised head and bit screen-wire bottom
of case.

Reaction time 2 to 60 seconds, average 19.1 seconds.

ButTERFLY LARVAE (Papilio polyxenes Fab.)

Oil of peppermint:
5 raised head. 5 showed no response.
Reaction time 3 to 60 seconds, average 33.2 seconds.

Oil of thyme:
5 raised head. 2 showed no response.
2 moved quickly. 1 moved caudal end of body.
Reaction time 2 to 60 seconds, average 22.1 seconds.

Oil of wintergreen:
5 moved slightly. 5 showed no response.
Reaction time 7 to 60 seconds, average 40.7 seconds.

Dried leaves of pennyroyal:
4 showed no response. 3 moved slightly.
3 raised head.
Reaction time 10 to 60 seconds, average 43.5 seconds.

Dried leaves of spearmint (odor very weak):
5 showed no response. 2 moved slightly.
3 raised head.
Reaction time 3 to 60 seconds, average 37.8 seconds.

In comparing the preceding reaction times, no conclusion
can be drawn in regard to the comparative sensitiveness of the
five species tested, because the responses depended more on the
behavior of the larva tested than on its ability to perceive
chemical stimuli. Of the five species used, only the army
worms were favorable for experimental purposes; these were
usually active, never failed to respond when tested, and the
responses were generally quick and pronounced ones. Of the
five species used, the butterfly larve were the most sluggish
and their reaction times are the slowest of all.

1919] McIndoo: Olfactory Sense al

MORPHOLOGY OF OLFACTORY PORES OR PUNCTURES.

The preceding pages show that lepidopterous larve respond
to chemical stimuli, and the following pages will show that
these larve have organs suitable for the reception of chemical
stimuli, but no experiments were performed to determine the
function of these organs which were first called olfactory pores
by the writer (1914a). The same type of organs seems to be
common to all adult insects and the writer has proved exper-
imentally that they receive chemical stimuli in Hymenoptera
(1914b) and Coleoptera (1915). They have also proved to be
common to all of the coleopterous and lepidopterous larve yet
examined by the writer, and now for several years systematists
have known them in lepidopterous larve as ‘‘punctures.’’ The
writer (1918) has recently described the external and internal
anatomy of them in a coleopterous larva of the ‘fig-eater,’
Allorhina (Cotinis) nitida L.

1. Disposition of Pores or Punctures.

In making a comparative study of the distribution and
number of these pores in lepidopterous larva, 30 species,
belonging to 28 genera and representing 20 families have been
used. Most of these were collected by the writer and the
remainder were kindly furnished by Mr. Carl Heinrich who
identified all of them. With a few exceptions, the writer has
adopted Mr. Heinrich’s (1916, 1918) nomenclature given to
most of the punctures present on the head capsule and at the
suggestions of Messrs. Heinrich and Busck he has formulated
new names for those pores not already named. The various
parts of the anatomy on which these pores occur were verified by
Dr. Adam Boving; the three foregoing mentioned men also
belong to the Bureau of Entomology.

In most cases only one specimen of each species was exam-
ined, and consequently, owing to unfavorable mounts and the
lack of sufficient material, the total number of pores recorded
‘can not be regarded as accurate. Since the army worm was
the most favorable material at hand, several specimens of it
were treated with caustic potash and the pores on it have been
studied and drawn in detail.

The description of them will be given first and then will
follow a brief account of those in other species.

72 Annals Entomological Society of America [Vol. XII,

Army Worm: Upon examining the head capsules of lepidop-
terous larve under a low-power lens, several minute circular
light spots were seen; these spots resemble hair sockets from
which the hairs have been removed, but former studies dealing
with similar spots at once suggested that they might be the
olfactory pores, so common to adult insects and coleopterous
larve. Upon examining them under a high-power lens, it was
soon observed that their external anatomy is different from
that of hair sockets and sections through them proved that they
are really olfactory pores.

Fics. 1 ANpD 2.—Disposition of the pores or punctures and setz on the head of a
small army worm (Cirphis wnipuncta Haw.), only the pores being here named,
xX 20. Fig. 1, dorsal view and Fig. 2, ventral view. Frontal pore (Fa);
adfrontal pore (Adfa); anterior pore (Aa); posterior pores a (Pa) and b (Pd);
lateral pore (La); ultraposterior pores a (UPa) and 6b (UPb); labral pore
(Lra); mandibular pores a (Mda) and 6 (Mdb); maxillary pores a (Mxa), c (Mxc),
d (Mxd), e (Mxe), and f to 7 (Mxf to 7); labial pores a to c (Lba to c); antennal
pore (Anta); genal pore (Ga); ocellar pore (Oa); subocellar pores a (SOQa),
b (SOb) and ¢ (SOc); secondary sete (4d; and Az); and Nagel’s so-called
olfactory pegs on antenne (THr).

Referring to Figures 1 and 2, it is seen that they are always
paired and they are located and named as follows: On the front:
the frontal pore (Fa); on the adfrontal piece: the adfrontal
pore (Adfa); on the dorsal sureface of the epicranium: the
anterior pore (Aa), posterior pores a (Pa) and b (Pb), lateral
pore (La), and ultraposterior pores a (UPa) and b (UPd); on
the ventral surface of the epicranium: the genal pore (Ga),

1919] McIndoo: Olfactory Sense hes

ocellar pore (Oa), and subcellar pores a (SOa), 6 (SOb) and
c (SOc); on the dorsal and ventral surfaces, respectively, of the
mandible: the mandibular pores a (Mda) and 6 (Mdb); on the
ventral surface of the labium: the labial pores a, b and c (Lba
to c); and on the dorsal surface of the labrum: the labral pore
(Lra). Relative to the maxilla they are as follows: On the
ventral surface of the stipes: the maxillary pore a (Mxa); on
the dorsal surfaces of the palpiger, first segment of the maxillary
palpus and maxillary lobe, respectively: the maxillary pores
b,c and d (Fig. 8, Mxbd to d); on the ventral surfaces of the first,
second and third segments of the maxillary palpus, respectively:
the maxillary pores e (Fig. 2, Mxe) and f to7 (Mxf to7). On
the ventral surface of the second or terminal segment of the
antenna, near the proximal end at the outer side, les the
antennal pore (Anta). ,

Relative to the thorax and abdomen, the pores are located
and may be named as follows: On the anterior surface of the
femur: the femoral pores a and b (Fig. 3, Fea and Feb); on the
posterior surface of the femur: the femoral pore c (Fig. 4, Fec);
on the posterior surfaces of the tibia and tarsus, respectively:
the tibial pore (Fig. 4, Toa and tarsal pore (Ta); on the tergum
of the prothorax: the thoracic pores a, b and c (Fig. 5, TJa to
c); on the tergum of the last abdominal segment: a single pair
of pores, the abdominal pores (not drawn); and on the dorsal
and ventral surfaces, respectively, of the anal proleg: the
anal-proleg pores a and 0 (Figs. 6 and 7, APa and APD).

Relative to the pores on the head of the army worm, the
subocellar pores (Fig. 2, SOb and SOc) and mandibular ones
(Mdb) are the most difficult to be found. While the former lie
in almost transparent chitin, the latter one lies in very dark
chitin; all of these, as well as the pair (Fig. 1, Mda) on the
dorsal side of the mandibles, seem to have been overlooked by
other observers. The ultraposterior ones (Fig. 1, UPa.and
UPb) are easily mistaken for the secondary sete (A; and Az)
and can only be distinguished from them by aid of a high-power
lens. Perhaps all of the pores on the labium and maxille have
been seen by systematists, but it seems that the third labial
pair (Figs. 2 and 20, Ldc), and maxillary pores b, g to 7 (Figs.
8 and 9, Mxb, g to7) have never been drawn. As far as known
to the writer, those on the legs, last abdominal segment and on
the anal prolegs are here reported for the first time.

74 Annals Entomological Society of America [Vol. XII,

The location and total number of the pores on various
parts of the army worm are as follows: front, 2; adfrontal
piece, 2; epicranium, 22; antennz, 2; mandibles, 4; labium, 6;
labrum, 2: maxilla, 20>" trochanters, 1S: tibia 62 tarsi, 16:
prothorax, 6; last abdominal segment, 2; and anal prolegs, 4;
making 102 pores in all.

Fics. 3 To 7.—Disposition of pores or punctures on leg, first thoracic segment and
on anal proleg of a small army worm, X 20. Figs. 3 and 4, anterior and
posterior surfaces, respectively, of left prothoracic leg. Fig. 5, most of tergum
of prothorax; the shield or strongly pigmented stripes are more heavily shaded
than are the lightly pigmented portions. Figs. 6 and 7, dorsal and ventral
surfaces, respectively, of same anal proleg. Femoral pores a (Fea), b (Feb)
and ¢ (Fec); tibial pore (Tba); tarsal pore (Ta); thoracic pores a (TIa), b (TIb)
and c TIc); anal-proleg pores a (A Pa) and 6 (A Pb); and secondary seta (A3).

Other species. The integuments of other species examined
were not studied as critically as were those of the army worm,
and consequently, some of the pores have certainly been over-
looked, particularly the ones most difficult to be found. Some
of the preparations had to be bleached with chlorine gas before
they could be studied, but most of them were so light in color
that it was difficult to find some of the pores, and the butterfly

1919} McIndoo: Olfactory Sense Fi

larvee were the least favorable of all, owing to their almost
total lack of pigment and to the fact that the. hairs on the
epicranium arise from large tubercles.

The following table gives the larger variations concerning
the disposition of these pores and now the smaller variations
may be briefly stated. For sake of brevity, instead of using
the long scientific names of the larvae examined, the species will
be numbered from 1 to 30, and those interested in associating
the names of the species with the variations described may do
so by referring to the names and numbers of the species in the
table on page 76. For each of the 30 species examined, the
pores are constant in number on the following parts of the
integument: Front, 2; each antenna, 1; each stipes, 1; each
maxillary lobe, 1; each second segment of the maxillary palpus,
1; and each third or terminal segment of the maxillary palpus, 4.
The number of pores found on the epicranium (including the
adfrontal piece) varies as follows: On each of 14 species,
OV pores (NOS 95," 6,700; 12°14 6-18, 21,°24, 25,29, 30);
on each of 4 species, 12 pores (Nos. 2, 4, 5, 22); on each of 9
species, 14 pores (Nos. 9; 11, 13, 15, 19, 20, 23, 27, 28); on
each of 2 species, 18 pores (Nos. 7, 26); and on 1 species, 24
pores (No. 8, the army worm). .

If all of the pores on each epicranium had been found, per-
haps 20 would be a common number for nearly all of the
species. One pore was found on each mandible of 29 species,
but had this appendage been examined more carefully two
pores might have been found on each mandible as already shown
for the army worm. The number of pores found on the labium
varies as follows: On each of 12 species, 4 pores (Nos. 1, 12, 13,
15-21, 23, 26); and on each of 18 species, 6 pores (Nos. 2-11, 14,
22, 24, 25, 27-30). The number of pores found on the labrum
varies as follows: On each of 5 species, 2 pores (Nos. 7-10, 16);
on each of 11 species, 4 pores (Nos. 6, 11, 14, 17-19, 21, 23,
25-27); and on each of 14 species, 6 pores (Nos. 1-5, 12, 13, 15,
20, 22, 24, 28-30). The first segment of each maxillary palpus
has 1 pore, except 2 were found on each one of 4 species (Nos.
3-5, 8). Three pores were found on each femur, except 2 on
each one of 10 species (Nos. 14, 15, 17-19, 21, 22, 24, 25, 29).
One pore was found on each tibia, except 2 pores on each of
1 tibia of 2 species (Nos. 9, 14) and 3 pores on 1 tibia of 1
species (No. 30). One pore was found on each tarsus, except

TABLE I.
Disposition of the pores or punctured found on lepidopterous larve.

FAMILY
AND
NUMBER AND NAME OF SPECIES

SPHINGIDEZ—
1. Phlegothontius sexta Joh..........
2. Ceratomia catalpe Bvd...........
SATURNIDA—
OS PANOMLENUSIVONE AID a ee
ARCTHDA—
4. Hyphantria cunea Dru............
NoctuIpbzA—
5. A patela(Acronycta)americanaHarr
6. Prodenia ornithogalli Guen........
eM CLIZORS Oe, ma timcens Aeee VaR ence ae
8. Cirphis unipuncta Haw...........
NoTODONTIDA—
9. Datana integerrima G.& R......
10! Datana nunisira Wr. ss). a 2..
LIPARIDA—
11. Hemerocampa leucostigma S.& A.
LASICAMPIDA—
12. Malacosoma americana Fab......
BOMBYCIDA—
IB JO MONGE TOI Wi csidaaosoncborsoe
GEOMETRIDZ—
14. Alsophila pometaria Harr.......
Ps yCHID4:—
15. Thyridopteryx
ephemereformis Haw.........
CocHLIDIIDA—
16. Sibine stumulea Clem............
MEGALOPYGIDA—
We bagoaerspaiaeack pees ae.
18. Megalopyge opercularis S. & A...
ZYGENIDE—
19. Harrisina americana Guér-Mén. .
PYRALIDZA—
20. Diatrea saccharalis Fab.........
ile Achrova erisellawWaler see sak
22. Dioryctria abettella D. & S.......
OLETHREUTIDA—
23. Laspeyresia pomonella L........
24. Laspeyresia molesta Busck.......
YPONOMEUTIDA—
Doe Attevarauned ses uC sae eee
GELECHIIDA—
26. Pectinophora gossypiella Saund..
BLASTOBASIDA—
27. Valentinia glandulella Riley.....
PAPILIONIDA—
2Se Papiliorpolyxenesalins sa asiee cele
PIERIDZ—
PY) SAVIO HEE Ngoc onobooscde ume
NYMPHALIDE—
30. Basilarchia archippus Cram.....

Variation. sjcren eas eee ee

NUMBER OF PORES ON
Total
HEAD THORAX ABDOMEN ee
eer r of
Head First ast S€s"| pores
ae append-| Legs | thoracic pee found
ages segment prolegs
12 30 30 72
14 32 30 76
12 36 30 78
14 34 30 78
14 34 30 sae 1 79
12 30 30 aff Foe a2,
20 28 30 6 2 86
26 34 30 6 6 102
16 28 31 3 78
12 28 30 2 72
16 30 30 76
12 30 30 2 2 76
16 30 30 3 79
12 30 19 61
16 30 18 64
12 26 30 68
12 28 24 64
12 28 24 64
16 28 24 68
16 30 30 4 ae 80
12 28 24 Me he nate 64
14 32 24 4 1 75
16 28 30 , 3 1 78
12 32 24 2 ee 70
12 30 24 66
20 28 30 2 80
16 30 30 2 78
16 32 30 78
12 32 24 68
12 32 32 76
12-26 | 26-36 | 18-32 0-6 0-6 61-102

1919] McIndoo: Olfactory Sense fi

none on the tarsi of 2 species (Nos. 14, 15). The number of
pores found on the first thoracic segment varies as follows:
Six on each of 2 species (Nos. 7, 8); 4 on each of 2 species
(Nos. 20, 22); 3 on each of 2 species (Nos. 9, 23); and 2 on each
of 5 species (Nos. 10, 12, 24, 26, 27). Two pores were found on
each of the last abdominal segment of 3 species (Nos. 7, 8, 13).
The number of pores on both anal prolegs varies as follows:
One for each of 3 species (Nos. 5, 22, 23); 2 for each of 2 species
(Nos. 12, 13); and 4 for the army worm (No. 8).

The total number of pores found varies from 61'to 102, but
no conclusion in regard to the comparative sensitiveness of the
various species can be drawn, owing to the fact that only one of
the species listed was critically studied.

In these examinations no attention has been paid to the
size or age of the larva being examined and at first thought one
might think that the disposition of the pores would vary
according to the instars, but Mr. Busck informs me that he has
found no such variations.

Discussion. According to the earlier papers concerning
lepidopterous larve, some of the entomologists have observed
the more conspicuous pores on the heads of these larve; they
have called these organs sensory pits and punctures, but knew
nothing about their internal anatomy. Within the past few
years, systematists have been making comparative studies of
the setze and punctures present on the integuments of lepidop-
terous larve and have used these characters successfully for
classifying the larve.

Forbes (1910) seems to have presented the first comparative
paper on this subject. He appears to have found some punctures
on all of the larvae examined; he has mapped the frontal
punctures on 33 species and adfrontal ones on 28 species, but
represents only a few of those present on the epicranium and
mouth parts. The same author (1911) mapped 3 pairs of punc-
tures on the labrum each of 4 sphingids.

Tsou (1914) mapped 2 pairs of punctures on the dorsal sur-
face of the prothorax of two genera.

Fracker (1915) mapped the frontal and adfrontal punctures
of several species.

Heinrich (1916) named and mapped most of the punctures
present on the head capsule of a micro-lepidopteron, and the

78 Annals Entomological Society of America [Vol. XII,

same author (1918) continued this study using the genus
Opostega.

Busck (1917) mapped most of the punctures present on the
head capsules of the pink bollworm (Pectinophora gossypiella
Saund.) and the scavenger bollworm (Pyroderces rileyi Wals.).
He found 22 punctures on each capsule, while the present
writer found 20 on the former capsule; in all probability there
are 26 on each capsule.

Wi

2I a2 os) "54 25 27 28

Fics. 8 ro 28.—Disposition and external view of pores or punctures on same army
worm as represented in Figs. 1 to 7. Figs. 8and 9, disposition of pores on

dorsal and ventral surfaces, respectively, of left maxilla, X 50. Maxillary
pores a to 7 (Mxa to 7). Figs. 10 to 28, external structures of pores and sete,
x 320. Fig. 10, ventral surface of second and third segments of maxillary
palpus, showing maxillary pores f to 7 (Mxf to 7) and tactile hairs (THr) at
tip of palpus. Fig. 11, tibial pore; Fig. 12, thoracic pore c; Fig. 138, abdominal
pore; Fig. 14, anal-proleg pore b; Figs. 15 and 16, maxillary pores d and 3,
respectively; Fig. 17, labral pore; Fig. 18, mandibular pore a; Figs. 19 and 20,
labial pores a to c, drawn in proper relation to each other; Fig. 21, frontal
pore; Fig. 22, adfrontal pore; Fig. 23, lateral pore; Fig. 24, subocellar pore c;
Fig. 25, ultraposterior pore b; Fig. 26, antennal pore; Fig. 27, secondary seta
(Fig. 5, As) on prothorax; and Fig. 28, secondary seta (Fig. 1, A2) on head.
Pore aperture (Ap); pore border (B); and pore wall (W).

2. External Anatomy of Pores or Punctures.

Under a high-power lens it is observed that a pore usually
has a dark border (Fig. 21, B) which may be round, oblong or
occasionally almost diamond-shaped, and it may show indica-
tions of radial streaks. Inside the border lies the wall (W)
which is usually dark and heavy; it is the most conspicuous
part of the organ and may be round or oblong. Inside the wall
the chitin is lighter in color and at the center may be seen an

1919] McIndoo: Olfactory Sense 79

aperture (Ap) which appears as a transparent spot; the aper-
ture is a minute opening passing through the thin chitin inside
the wall.

Sometimes it is almost impossible to distinguish a secondary
seta from a pore, but in almost all cases the chitin inside the
wall of such a seta is lighter in color than that inside the wall of
a pore, and this fact may help to distinguish a seta from a pore
in case the seta has been pulled out; this comparison may be
seen by referring to Figures 25 and 28, both of which structures
come from the same region on the epicranium.

In size the pores do not vary greatly, as may be observed by
looking at Figures 10 to 28. The smallest one (Fig. 10, Mx7)
lies on the terminal segment of the maxillary palpus and the
largest one (Fig. 14) in the army worm lies on the anal proleg;
the former is scarcely discernible and seldom has a border. The
one (Fig. 10, Mxz) nearest it is always slit-shaped and algo in
some species can scarcely be seen. The other two (Mxg and
Mxh) on this segment are easily seen in good mounts, and have
distinct borders.

3. Internal Anatomy of Pores or Punctures.

A reference to Figures 29 to 48 shows that the internal
structure of these organs is like that of those in other insects,
and consequently only a brief description of the various parts
of them will suffice here. Lying in the thick hypodermis
(Fig. 29, Hyp) is the large sense cell (SC) whose peripheral end
(SF) passes through the pore canal (Cam), pierces the chitinous
cone (Fig. 30, Con) and then stops in’the bottom of the pore
aperture (Ap) where it seems to come in direct contact with the
external air. The chitin covering the cone may be dome-
shaped, with the dome either lying in a depression (Fig. 29) or
elevated above the surrounding chitin(Fig. 33); or it may form
a depression whose bottom is pierced by the pore aperture
(Fig. 36); or it may lie on the same level with the surrounding
chitin (Fig. 31).

Since the chitin was so thick, not a single section showed all
the details of one of these organs, because the microtome knife
never passed properly through the structure; but Figures 29
and 30 combined give a good idea of their anatomy, which is
very different from the internal anatomy of a large hair (Fig. 34)
which is formed by a large hair-mother cell (HrMC) sending

80 Annals Entomological Society of America [Vol. XII,

forth processes through the pore canal to the base of the hair
(Hr). Should both the base of the hair and the hair-mother cell
be missing, the hair socket itself in sections serves well to
distinguish this structure from a pore.

To determine whether all of the structures recorded as pores
are really pores or hair sockets, thin sections made through the
various parts of the integuments bearing these structures were

Fics. 29 to 48.—Sections, showing internal anatomy of pores or punctures and of
one hair from various lepidopterous larve, X 500. Figs. 29, 31 to 34 are pores
from Ceratomia catalpe; Fig. 29, being from epicranium; Fig. 31, from antenna;
Fig. 32, from palpiger; Fig. 33, from labium; and Fig. 34 is internal structure
of a large hair on epicranium. Fig. 80 is a pore from labrum of Telea poly-
phemus and Fig. 35, two pores from labrum of silkworm (Bombyx mori). Figs.
36 to 38, maxillary pores f (Maf), g (Mxg), 7 (Wxj) and 7 (Mx1), respectively,
of tomato worm Phlegothontius sexta). Fig. 39, pore from labrum and Fig. 40,
pore from antenna of cabbage-butterfly larva (Pontia rape). Figs. 41 to 48,
sections from material, treated with caustic potash. Fig. 41, pore from
terminal segment of maxillary palpus of codling moth (Laspeyresia pomonella).
Figs. 42 to 48, pores from army worm (Cirphis unipuncta); Fig. 42, from tibia;
Fig. 43, from antenna; Fig. 44, from maxillary lobe; Fig. 45, from femur; Fig. 46,
from mandible; Fig. 47, from front; and Fig. 48, from anal proleg. Sense
cell (SC); hypodermis (Hyp); sensory fiber (SF); pore canal (Can); chitinous
cone (Con); pore aperture (Ap); hair-mother cell (HrMC); and base of
hair (Hr).

carefully studied; in every case the suspected pore proved to
be a real pore. Since most of these are so scattered, they are
easily overlooked in sections made from fixed material, but '
were found much more easily in sections made from material
treated with caustic potash (Figs. 41 to 48).

1919} McIndoo: Olfactory Sense 81

Nagel (1894) called the small hairs at the tip of the antennz
(Fig. 1, THr) of a lepidopterous larva olfactory pegs, but in
the opinion of the present writer they are nothing more than
tactile hairs. Sections through them show that they are not
true hairs, because they do not arise from sockets, but each one
is nevertheless innervated; the larger ones seem to be provided
with sense cell groups (Fig. 49, SCG), while the smaller ones
(Fig. 50, THr) arising from the dome seem to be provided with

Fics. 49 anp 50.—Longitudinal sections, showing internal anatomy of antenna
of tomato worm. Fig. 49, the larger type of one of Nagel’s so-called olfactory
pegs at tip of antenna, X 500; sense-cell group (SCG). Fig. 50 shows how well
antenna is innervated; semidiagrammatic, X 100. Hypodermis (Hyp); nerve
(NV); antennal pore (Anta); probably hair-mother cell (HrMC); sense cell
(SC), connected with pore; sense cell (SCi), connected with peg; smaller
type of one of Nagel’s so-called olfactory pegs (T'Hr) arising from dome;
and base of large true hair (Hr).

single sense cells (SCi). The large hair (Hr) at the tip of the
antenna is a true hair, but it does not appear to be sensory,
although lying at its base there are one or two large cells (Hr MC)
which resemble hair-mother cells more than sense cells.

One author draws the antennz of lepidopterous larve as if
they were composed of 3 or 4 segments, but sections show
only two distinct segments in each, although sometimes the
basal one is so folded that indications of two more segments
are visible, as shown in Figure 50, which also gives a good idea
of how well the antenna is innervated.

82 Annals Entomological Society of America [Vol. XII,

Fics. 51 AND 52.—Longitudinal sections, showing internal anatomy of maxillary
, & palpus. Fig. 51 shows sensory fibers (SF) running to 3 of hairs (THr) at tip
of palpus of Ceratomia catalpe, X 500. Fig. 52 shows how well palpus is
innervated; semidiagrammatic, X 190. Sense cells (SCi), connected with
hairs (THr); sense cells (SC), connected with pores; maxillary pore f (Wxf);
fe" and nerve (JN).

[{' | The terminal segment of the maxillary palpus ends bluntly
(Fig. 53) and the ue is provided with 8 or 9 minute pseudo-
hairs (Figs. 51 and 52, THr), each of which seems to be inner-
vated by a single sense reall GSC;).. These sense cells lie in a group
at the proximal end of the segment near another group of sense
cells (SC) which evidently belong to the pores. Figures 52 and
53,show how. well the maxillary palpus is innervated; a portion
of ‘the terminal segment in Figure 53 1s shown in perspective.

S

ee ———

— SSS

53

Fic. 53.—Schematic drawing of maxillary palpus of tomato worm, showing innerva-
tion of maxillary pores f to 7 (Mxf to 7) and tactile hairs (JHr); a portion of
terminal segment is in perspective.

1919] McIndoo: Olfactory Sense 83

SUMMARY.

To determine whether lepidopterous larve respond to chem-
ical stimuli, tent caterpillars, fall webworms, tussock-moth
larve, army worms and larve of Papilio polyxenes were tested
by using the following sources of odors: Oils of peppermint,
thyme and wintergreen, dried leaves of pennyroyal and spear-
mint, wild cherry-tree leaves, fresh grass, old honey and comb,
and the protruded thoracic glands of the above Papilio larve.
The larve usually responded to the exhalations from these sub-
stances, but the average reaction times obtained seemed to
depend more on the degree of sluggishness of the larve than on
their sensitiveness to odors.

Organs, called olfactory pores by the writer, but known as
punctures to systematists, were found widely distributed on the
head capsule, head appendages, legs, dorsal surfaces of the
prothorax and last abdominal segment, and on the anal prolegs.
It is believed that a few of those on the head and all of those
found on the legs and abdomen are here reported for the first
time. Their internal structure is like that of those in adult
insects and coleopterous larve, and consequently are well
adapted to receive chemical stimuli, because their sensory
fibers running from the sense cells pass into the minute pores
or punctures and, seem to come in direct contact with the
external air. No experiments, however, were performed to ,
determine their function.

Verschaffelt determined experimentally that cabbage-butter-
fly larvee are attracted by the various mustard oils contained in
the host plants, and this explains why these larve refuse plants
not containing such oils; he also thinks that the larve smell the
odors from the mustard oils before they begin to eat the food.
If we knew more about the chemotaxis of insects, we might be
able to devise practical methods for the control of certain
insects, as by trap baits, etc.

%

84 Annals Entomological Society of America [Vol. XII,

LITERATURE CITED.

Busck, August. 1917. The pink bollworm, Pectinophora gossypiella. Jour.
Agr. Research, v. 9, no. 10, p. 343-370, 7 figs., 6 pls.

Forbes, W. T. M. 1910. A structural study of some caterpillars. Ann. Ent.

Soc. Amer., v. 3, p. 94-1438, 11 pls.

— . 1911. A structural study of the caterpillars—II. The Sphingide,

Ibid., v. 4, p. 261-282, 2 pls.

Fracker, S. B. 1915. The classification of lepidopterous larve. II]. Biol.
Monograph, v. 2, no. 1, 169 p., 10 pls.

Heinrich, Carl. 1916. On the taxonomic value of some larval characters in the
Lepidoptera. Proc. Ent. Soc. Wash., v. 18, no. 3, p. 154-164, 2 figs., 1 pl.
—————. 1918. On the lepidopterous genus Apostega and its larval affinities,

Ibid., v. 20, no. 2, p. 27-34, 4 pls.

McIndoo, N. E. 1914a. The olfactory sense of the honey bee. Jour. Exp. Zool.,
v. 16, no. 3, p. 265-346, 24 figs.

—. 1914b. The olfactory sense of Hymenoptera. Proc. Phila. Acad.
Nat.:Sci., v. 66, p. 294-341, 3 figs., 2 pls.

————. 1915. The olfactory sense of Coleoptera. Biol. Bul., v. 28, no. 6,
p. 407-460, 3 figs., 2 pls.

—————. 1918. The olfactory organs of a coleopterous larva. Jour. Morph.,
v. 31, no. 1, p. 113-1381, 33 figs.

Nagel, W. A. 1894. Yergleichend physiologische und anatomische Untersuch-
ungen ttber den Geruchs und Geschmacksinn und ihre Organe. Bibliotheca
Zoologica, Heft 18. Die Schmetterlingsraupen, p. 113-115 with fig. 32.

Tsou, Y. H. 1914. Homology of the body sete of some lepidopterous larve.
Trans. Amer. Mic. Soc., v. 33, p. 223-260, 4 pls.

Verschaffelt, Ed. 1910. The cause determining the selection of food in some
herbivorous insects. Proc. Sec. Sci. Royal Acad. Sci., (Kon. Akad. van
Wetenschappen) Amsterdam, v. 13, pt. 1, p. 536-542.

THE ee GROUP OF THE DIPTEROUS GENUS
TIPULA LINNAEUS.

We. G. Dietz, M.D:

A natural group of species, which briefly may be characterized
as follows: Wings approximately unicolorous, stigma distinct,
the antestigmal spot always extends into cell lst M2, but is
interrupted in the outer end of cell R in most of the species.
The eighth sternite of the male is prolonged and narrowed
posteriorly, except devia. The hypopygium is of moderate
size. .The ninth tergite, generally small, is emarginate in the
middle, except fragmentata. The pleural suture is distinct,
except devia—the pleurite prolonged into a process,, generally
long and twisted, hornlike, rarely into an acutely pointed
cone, derbyi, or simply tipped by a sharp point, devia. The
Oovipositor is very short, the upper valves generally rounded or
oval, in some species ending in a downward curved point,
giving thus a beaklike appearance; the lower valves scaphoidal.
In all the specimens in my collection, the posterior end of the
abdomen, especially the males, is curved upward and sometimes
backwards towards the abdominal tergum. Whether this
also obtains in the living insect I have no means to know.

The species, below medium in size, are yellowish to testaceous
in color, with more or less lustre. They resemble each other
very closely and the genital structure of the males must in
almost all cases be depended upon for differentiation.

KEY TO THE SPECIES.

1. Joints of the antennal flagellum unicolorous. Western species.............. 2
Flagellar joints bicolored. Eastern United States; aberrant form.
Ae Suerenelats God SNES Peele G Aen BF i aN eet aren sO anne ae devia spec. n.

2. Rhemimth tergite of the male distinctly emarginate....-..........+.......+.0
Not as in the alternative. ........... Oe LC Oe ea ee Ome fragmentata spec. n.
Se LNeEMleLnd spLOCESSHLONnOeMODMlKee. 4.22 8 sary Stowe ce cosa scces ee) eae ke: i
The pleurite prolonged into a pointed, conelike process....derbyi Doane
4. The pleural process long, twisted, more or less curved, glabrous SE TSE 5)

The process shorter, approximately straight, stiletto- like, hairy,
od Bota OU NOHO IE OS BOS Be SOIC aE One pela atenor monochroma Spencer
HEETOCESSICONSPICUOLISIVECURVEUser LE tla: - hide sins uetisee cine ca tack ema een Sac. « 6
Processsapproximatelya Straight»... .. 26 «Ate adne ses trypetophora spec. n.
6. The ninth tergite with a deep, oval emargination, the sides prolonged into
converging processes, the inner margin with a conspicuous tooth.
ng GS EARS 5 IG aN: 3 PRG MeSIneS aia 9 ie sn drs raed ee ara opisthocera spec. n

INOifaSta DOCH Ee peer EP ke Terrence PA te nea Say istad s «(ecuns eben 7
7. Process shorter, viewed laterally, much wider in basal half...mutica spec. n.
ING ERaSwilgn Oe aalbenilabivien scorn = cece cae GuPoe te cite eo chafeveens esos = disidele wis eels 8
8. Antennal flagellum—except basal joint—dark brown....... streptocera Doane
Antennz entirely yellowish................ var. streptocera pallidocera var. n.

\
\

\

S6 Annals Entomological Society of America [Vol. XII,

Tipula devia sp. nov.

Yellow. Antennz yellow, flagellar joints beyond the third, brown
at the base. Thoracic stripes obsolete. Pleurite of hypopygium
tipped with a sharp point.

Male, length 13 mullimeters, wing 14.5 millimeters.

Head entirely, together with mouth-parts and palpi, yellow;
last joint of palpi a trifle longer than the three preceding joints
together. Antenne yellow, flagellar joints beyond the third brownish
at the base; joints cylindrical, basal enlargement very slight, the sete
about as long as the respective joints. Thorax concolorous, dorsum
subtranslucent, stripes obsolete, pleura with a faint whitish sheen,
coxee and trochanters yellow, rest of legs lost. Halteres pale yellow,
club infuscate. Wings light gray, costal cells yellow, stigma’ light
fuscous, veins C,S C, R and Cu, yellow, the other veins brown; the
antestigmal spot strongly marked, extending through cell lst M2 into
base of cell M3. As far as I can perceive, vein M1 and M2 alone are
distinctly setigerous, the sete rather long.

Abdomen yellow, darker posteriorly, the eighth sternite very little
prolonged, the posterior margin with a median notch and a slight
emargination on each side; a dense brush of golden-yellow, bristly hair
arises from beneath the margin and a stout bristle, curved outward
from the lateral angle. Hypopygium (Pl. V, Figs. 1 and 2) testa-
ceous. Ninth tergite markedly narrowed posteriorly, deeply emarginate,
the upper surface impressed mesially, the sides of the tergite prolonged
into an acute process, denticulate within. Ninth sternite broadly
emarginate, base of the emargination subangulate. Pleural suture
obsolete, the pleurite projects as an acute point. Outer apical append-
ages large, hatchet-shaped, projecting inward, lower appendages large,
sausage-shaped bodies, coarsely hairy and projecting downward into
the emargination of the ninth sternite.

Holotype, o. Plummers Island, Maryland, June 28th,
1914 (W. F. McAtee).

A distinct, but somewhat aberrant species in this group.
The absence of the pleural suture, the pleurite merely projecting
into an acute point and the scarcely prolonged eighth sternite
is at variance with the other species included in this paper.
It is the only species with bicolored antennal flagellum and the
only one from the Eastern United States. The type and
only specimen is defective, an antenna, one wing and all the
legs wanting.

NoOTE:—Since the above has been: written, three more
specimens of this species have turned up in my collection.
A male, Southern Pines, North Carolina, June, 1910, (A. H.
Mance); a male and female specimen, Black Mountain, North
Carolina, July, 1912, (Wm. Bentermueller).

1919 Dietz: Streptocera Group of Tipula, 87

Allotype, @, has the ovipositor elongated, testaceous and
highly polished. The upper valves long and slender, pointed.
The lower valves nearly four-fifths the length of the upper
valves, obtuse at the apex.

Tipula fragmentata sp. n.

Reddish-brown with some lustre. Thoracic stripes distinct; ninth
tergite rounded posteriorly, with a subtriangular prolongation from
beneath the posterior margin. Pleural prolongation slender, twisted,
strongly inclined proximad.

Male, length 11 millimeters, wing 13.5 millimeters.

Head dark, margin of orbits whitish. Frontal prolongation rather
long, reddish-brown, shining, beset with black hairs. Nasus short,
obtuse, with a tuft of white hair at the apex. Palpi brown. Scape and
first flagellar joint of the antenne pale yellow, second flagellar joint
fuscous, remainder of antennee wanting.

Thoracic stripes reddish-brown, with some lustre, separated by five
yellowish lines of the ground color; the median stripe divided by a
fine line. Pleura fuscous, dull. Scutum uniformly dark fuscous with a
faint lustre. Scutel yellowish and like the yellowish-fuscous postnotum,
shining. Legs slender, yellowish, femora yellow, apices fuscous; tibia
and tarsi yellowish-fuscous.: Halters yellowish, knob wanting. Wings
grayish, stigma fuscous, the antestigmal spot faint interrupted in
cell Ist, M2; a hyaline spot beyond the stigma.

Abdomen pale testaceous, shining, with median dorsal and lateral
stripes, the latter somewhat interrupted; the median stripe does not
extend upon the posterior tergites. Hypopygium (Plate XXX, Figs.
3 and 4) yellowish, pleural processes brownish. Ninth tergite rounded
posteriorly and from beneath its posterior margin extends a sub-
triangular process or prolongation. Ninth sternite deeply emarginate,
the emarginations almost entirely canceled by the narrowed end of
the prolonged eighth sternite, the latter with a brush of hair at its apex.
Pleural processes twisted, broad at the base, strongly attenuated and
twisted outwardly, inclined towards each other. The outer apical
appendage digitiform, extends obliquely inward.

Holotype, o, bears a label marked California (Bradley),
ype CiU,

The specimen is in the collection of Mr. C. P. Alexander.
It was received in alcohol and practically dismembered. In its
hypopyginal characters, however, more especially the rounded
ninth tergite, it is at once differentiated from all the other
species. In its distinct thoracic stripes it resembles streptocera,*
trypetophora and opisthocera.

* A specimen, labeled T. streptocera, received from Prof. Doane, and another
from my collection, determined by him, have the thoracic stripes nearly obsolete.
The same is also the case in nearly all the specimens of this species in my col-

lection.

88 Annals Entomological Society of America [Vol. XII,

Tipula monochroma spec. n.

Yellowish. Antennal flagellum unicolorous, fuscous. Thoracic
dorsum brownish-yellow, dull. Prolongation of pleurite straight,
stilletto-like, hairy.

Male. Length 11 millimeters; wing 12.5 millimeters.

Head concolorous; front and occiput gray, with a fine, dark median
line. Frontal prolongation long, shining and like the nasus above, beset
with short, blackish hairs. Palpi sordid yellow, last joint infuscate and
scarcely longer than the preceding joints together. Antenne moderate,
the three basal joints yellowish, the remainder fuscous; segments sub-
cylindrical, the basal enlargement blackish, each with a whorl of black,
moderately long setz.

Thoracic dorsum brownish-yellow; pronotal scutem darker about
the middle. Praescutal stripes ill-defined by fine, paler lines. Scutum,
scutellum and postnotum semiopaque, slightly more infuscate and
somewhat sericeous. Pleura pale yellowish, sericeous. Legs yellowish,
outer tarsal joints infuscate, the whole leg beset with coarse, short
black hairs. Halters pale, club infuscate. Wings pale grayish; costal
cells very pale yellowish; stigma pale, slightly infuscate. The ante-
stigmal spot inconspicuous, extends through cell Ist M2 to base of
cell M3, and sending a narrow, pale streak to the wing margin. The
veins included in the antestigmal spot are whitish. Veins R and Sc.
yellow, the other veins dark brown and delicate. The veins beyond
the cord are indistinctly margined with subhyaline, The R, s, and the
veins beyond the cord, except those limiting cell 1st M2, setulose. :

Abdomen darker yellow, almost testaceous, shining; an ill-defined,
more or less interrupted, fuscous dorsal stripe; tergites three to seven
with a fuscous spot near the anterior angle and from which extends an
indistinct streak along the lateral margin of the tergite. Venter with
median blackish stripe, somewhat interrupted and obsolete beyond the
fourth sternite. The eighth sternite projecting, narrowed posteriorly,
the apical margin nearly truncate, with a brush of stiff, yellowish-
fuscous, incurved bristles. Hypopygium (Pl. V, Figs. 7 and 8)
dark testaceous, medium sized. The ninth tergite broad, slightly
narrowed posteriorly, the free margin with small V-shaped, median
notch, emarginate each side, the apical angles curved downwards.
Ninth sternite with a rather deep, V-shaped emargination, from which
projects a pencil of stiff bristles, lateral angles with a small appendage,
beset with bristles. Pleural suture distinct. The pleural -process
has the shape of a stilletto, beset with hairs. The outer appendages
are yellowish-white, membranous, bilaminate, somewhat protuberant
and acuminate; the inner lamina beset with bristles, directed inwardly.

Holotype, o, Chimney Gulch, Golden, Colorado, (E. J.
Oslar).

Paratype, o’, Silver City, Nevada, June 3rd, 1913.

The paratype is paler than the holotype; the dark markings
of the abdomen, with the exception of the lateral spots, nearly
obsolete. The hairy stiletto-shaped pleural process of the
male will readily distinguish this species.

1919] Dietz: Streptocera Group of Tipula 89

Tipula trypetophora spec. n.

Yellowish, testaceous, shining. Antennal flagellum unicolorous.
Thoracic stripes distinct. Wings faintly gray, veins beyond the cord
broadly margined with hyaline. Pleural process conspicuous, long,
twisted, nearly straight.

Male. Length, 13.5 millimeters; wings, 13.5 millimeters.

Head testaceous. Frontal prolongation and rostrum beset with short,
black hairs. Palpifuscous, the last joint scarcely as long as the preceding
joints together. Front and occiput fuscous with an obscure, median
darker line; the space surrounding the antennal insertion somewhat
paler. Antenne of moderate length; scapal joints pale yellow, first
flagellar joint pale yellow at base, infuscate apically, the remaining
joints fuscous, cylindrical, basal enlargement with the usual whorl
of sete, the latter shorter than the respective segments; pubescense
whitish, very short; the last antennal joints rather strongly attenuate.

Thorax concolorous, shining. Pronotum scarcely darker in the
middle. Praescutal stripes brown, well defined, the median stripe
very little narrowed posteriorly, the lateral stripes abbreviated in front,
extended upon the scutum, leaving a narrow external and internal
margin of the ground color. Scutel and postnotum concolorous.
Pleura with a pale sheen. Legs yellow, femora somewhat infuscate
at the apex; tibiae darker yellow; tarsi fuscous; tibize and metatarsi
subequal; pilosity very dense, short, blackish. Halters sordid yellow,
club infuscate. Wings grayish-hyaline, costal cells with a faint, yellow-
ish tinge, stigma pale yellowish-fuscous. Veins brown, those beyond
the cord broadly margined with hyaline. Cell Ist M2 fully two and
one half longer than wide. The antestigmal spot more or less inter-
rupted, extends into cell Ist M2, Rs. and veins beyond the cord, except
Cu, setulose.

Abdomen concolorous; an ill-defined dorsal stripe, more distinct
posteriorly; sides of tergites three to eight somewhat infuscate, three to
five with a dark fuscous spot near the anterior angle. Venter with ill-
defined, median darker stripe. The eighth sternite strongly narrowed
and prolonged posteriorly, the apical angles with a small, triangular
appendage; two narrow processes project from the middle of the posterior
margin, the latter without a brush of hair. Hypopygium (Pl. V
Figs. 9 and 10) reddish-testaceous and rather large. The ninth tergite
short, with narrow median incision, the apex of each side curved dorsad
in the form of a small, obtuse tubercle, which is concave and shining
on its posterior face. Ninth sternite short, broadly emarginate; pleural
suture complete. Pleural process conspicuous, twisted at the base
and within the latter arises a sharp, conical process; the direction of the
process is nearly straight, caudad; apical appendages inconspicuous.

Female. Length 12.5 millimeters; wing, 13.5 millimeters. Flagellar
joints two and three yellowish towards the apex, remainder of flagellum
light brown. The lateral abdominal stripe distinct from tergite three to
eight. Ovipositor short, concolorous, viewed laterally upper valves
almost circular, lower valves elliptic, the upper margin ending in a
very fine, spine-like prolongation. :

90 Annals Entomological Society of America [Vol. XII,

Holotype, o&, Victoria, British Columbia, July 6th, 1912.

Allotype, @, topotypic.

Paratypes, 1 &, 69 9’s, topotypic.

This species very closely resembles opisthocera. The wings
are a trifle paler, likewise the stigma. The very different
construction of the ninth tergite of the male, absolutely differ-
entiates the two species. One male paratype has the thoracic
stripes much less distinct.

Tipula opisthocera spec. n.

Testaceous; antennal flagellum unicolorous, fuscous. Thoracic
stripes distinct. Pleural process conspicuous, strongly curved.

Male. Length, 12 millimeters; wing, 15 millimeters.

Head concolorous; frontal prolongation long, clothed with blackish
hair, rather long on the nasus. Palpi brownish, the fourth joint darker
and but little longer than the three preceding joints together. Face
yellow. Antenne slender, scapal joints yellow, first flagellar joint yel-
lowish fuscous, joints subcylindrical, basal enlargement moderate with
a whorl of rather long, black bristles; pubescense very fine, short, whitish.
Front and occiput very dark fuscous, except near the base.

Thorax concolorous, shining. Praescutal stripes distinct, brown,
rather broad and indistinctly margined with darker brown, the median
stripe scarcely narrowed posteriorly, the lateral stripes much abbreviated
in front, continued upon the scutum, where they leave but a narrow
lateral stripe and a very narrow margin within, of the ground color.
Scutellum and postnotum concolorous, lightly touched with fuscous.
Pleura pale testaceous, with a faint, pale sheen. Legs yellowish-brown,
densely clothed with brown hair; coxa and base of femora paler, yellow;
apices of femora and tibiz fuscous, the latter longer than the metatarsi;
tarsi fuscous. Halters concolorous, paler at base, club infuscate.
Wings grayish with a fuscous tinge; vein M and all the veins beyond
the cord broadly margined with hyaline; costal cells a very faint yellow;
stigma brown; the antestigmal spot extends into the base of cell M3, is
faint beyond cell R1; a hyaline spot beyond the stigma. Rs, vein M
and the apical veins except those limiting cell Ist M2, strongly setigerous.

Abdomen testaceous, paler towards the base; a fuscous dorsal
stripe extends from the second tergite to the end of the seventh, becoming
more diffused posteriorly; tergum indistinctly margined with fuscous,
more accentuated in a dark spot on the anterior angles of tergites three
to seven, a faint median ventral stripe. The eighth sternite clasping
cephalad, narrowed and somewhat prolonged posteriorly, the caudal
margin rather deeply emarginate, with a triangular appendage at each
angle of the emargination, the latter filled with a whitish membrane,
which has the appearance as though it consisted* of matted hair.
Hypopygium (Pl. V, Figs. 11 and 12) reddish-brown; ninth tergite
rather narrow, directed obliquely dorsad, somewhat turgid, with a
very deep oval emargination, the lateral angles prolonged, turgid,

1919} Dietz: Streptocera Group of Tipula 91

inclined inwardly, the inner margin with a large obtuse tooth and
several smaller denticles. Ninth sternite very short, with slight
medium notch. Pleural process long, somewhat flattened, twisted
and appearing like an extended S. Apical appendages small and
inconspicuous.

Female. Length, 12 millimeters; wing, 13.5 millimeters.

Antenne shorter and more slender. Dorsal stripe of abdomen broader
and more diffused posteriorly; the lateral stripes broad, strongly marked,
tergites three to seven without lateral spots. Eighth segment entirely
fuscous. Ovipositor dark testaceous, the upper valves short, quadrate-
oval, dark brown; lower valves paler, very small, scaphoidal.

Holotype, o&, Sonoma County, California, April 20th, 1914.

Allotype, 92, topotypic.

Paratypes, 40'c"’s, 3 2 9’s, topotypic. :

A very distinct species. The ninth tergite of the male,
with its deep, oval emargination, the sides thereof prolonged
and strongly incurved with a large, obtuse point on its inner
margin, distinguish this from the other species. The thoracic
stripes are pronounced in all my specimens.

Tipula mutica spec. n.

Yellowish testaceous, shining. Antennal flagellum unicolorous.
Thoracic stripes almost effaced. Pleural process twisted, curved,
broad in basal portion.

Male. Length 12 millimeters; wing, 14 millimeters.

Head. Frontal prolongation and rostrum reddish-brown, with short,
black hair. Palpi dark fuscous, joints two to four whitish at the base, the
fourth joint scarcely as long as the preceding joints combined. Front
and occiput grayish-fuscous, whitish along the upper orbital margin,
a dark brown fronto-occipital stripe. Antenne of moderate length,
slender; first three joints pale yellow, remaining joints fuscous; basal
enlargement of segments blackish with the usual whorl of sete, the
latter much shorter than the respective segments; pubescense fine,
whitish.

Thorax concolorous, shining, the stripes faintly indicated, a shade
darker than the ground color. Scutel and postnotum finely sericeous.
Pleurz pale yellow, faintly sericeous. Legs yellow, femora and tibiz
infuscate at the apex; the short, dense, black pilosity gives the legs a
darker appearance; tibize distinctly longer than the metatarsi; tarsi
fuscous. Halters pale fuscous, lighter towards the base, club dark
fuscous. Wings pale gray, costal cells with a pale, yellow tinge, stigma
pale yellowish-brown. Cell 1st M2 about twice as long as wide, veins
brown, those beyond the cord broadly margined sub-hyaline, veins
M and Cu less distinctly margined. The antestigmal spot is rather
faint and does not extend beyond cell lst M2; a faint hyaline spot beyond
the stigma. Rs, veins M, Cu, A; and apical veins setulose.

92 Annals Entomological Society of America [Vol. XII,

Abdomen testaceous, shining; a well marked dorsal stripe extends
from the first to the eighth tergite, a lateral fuscous stripe more or
less interrupted, extends from the third to the sixth tergite, more
accentuated in a darker spot near the anterior angle of the respective
segments. Eighth sternite prolonged and narrowed posteriorly, the
posterior margin with a broad, rounded emargination which is filled
with a whitish membrane; each angle of the emargination bears a large,
triangular appendage, the inner margin of which has a brush of short,
dense, golden-yellow hair. Hypopygium (Pl. V, Figs. 13 and 14)
reddish, testaceous. The ninth tergite about as long as wide, broadly
and deeply emarginate, the lateral angles prolonged as digitiform
processes, slightly bent downward at the apex; the upper surface of the
tergite with a median, longitudinal carina; viewed laterally, the upper
surface appears subangulate about the middle. Ninth sternite short,
with a median acute incision. Pleural suture complete, pleural process
conspicuous, curved upward and markedly broadened just beyond the
base, convoluted, then extending directly caudad and slightly bent
outwardly. The outer apical appendages small, ovoidal, fringed with
hair; the inner appendages a broad perpendicular lamina, extending
above into an acutely pointed and strongly chitinized process; the
lower margin strongly bearded with yellow, bristly hair.

Holotype, o&, Somona County, California, May 21, 1914.

Paratypes, 4 2 9’s, topotypic.

The formation of the hypopygium alone must be depended
on for differentiation from allied species.

Tipula streptocera-pallidocera var. n.

A specimen in my collection received from Prof. Doane and
bearing the label “‘L. streptocera,’’ agrees in all details with
the description and other specimens of this species, received
or determined by its author, except that the antenne are
entirely pale yellowish. The thoracic stripes are ill defined.

Holotype, <&, Keyport, Washington, July, 1905, (R. W.
Doane).

1919]

Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.

a ge CR Ee ee

some

Dietz: Streptocera Group of Tipula 93

EXPLANATION OF PLATE V.

Hypopygia.
Dorsal aspect of T. devia.
Lateral aspect of 7. dewa.
Dorsal aspect of T. fragmentata.
Lateral aspect of 7. fragmentata.
Dorsal aspect of T. derby.
Lateral aspect of T. derbyt.
Dorsal aspect of T. monochroma, without apical appendages.
Lateral aspect of T. monochroma.
Dorsal aspect of T. trypetophora.
Lateral aspect of T. trypetophora.
Dorsal aspect of T. opisthocera.
Lateral aspect of T°. opisthocera.
Dorsal aspect of T. mutica, without apical appendages.
Lateral aspect of T. mutica.
Dorsal aspect of 7. streptocera, without apical appendages.
Lateral aspect of T. streptocera.

—

WO

VoL. XII, PLATE V.

LIFE HISTORY AND BIOLOGICAL NOTES ON
CHLAENIUS IMPUNCTIFRONS SAY.
(COLEOPTERA, CARABID Z).*

P. W. CLAASSEN.

In the Proceedings of the Entomological Society of Wash-
ington I, 22, 1884, C. V. Riley says: ‘‘ Perhaps one of the most
interesting discoveries of the year 1884 is the mode of oviposi-
tion in some of our Carabide. From the terrestrial habits of
most of our species one would expect that the eggs are depos-
ited within the ground, and such may yet prove to be the case
with many; but I have proved by actual breeding from eggs
to the imago that it is not so with Chlenius impunctifrons, and
have strong proof that Chlenius estivus, Scarites subterraneus
and the genera Dicelus and Galerita share with that species its
singular mode of oviposition. The remarkable and unexpected
fact, in insects so essentially terrestrial, is that the eggs are laid
singly on the leaves of trees and shrubs and encased in a cov-
ering of mud or clay. I had often observed these little convex
mud cells on the under side of leaves while collecting along the
Mississippi in Missouri in years gone by, and was puzzled to
make out their real nature. In May and June, 1883, while col-
lecting on the Virginia side of the Potomac, I found these clay
cells tolerably common and, fortunately, fresh, each containing
a large soft white egg. That year I obtained larve, but only
during the past year were any of these reared to the imago.”’

The above account is not accompanied by any figures or
descriptions, and as far as I am aware, the life history of
Chlenius impunctifrons has not been written hitherto.

On May 23, 1918, while collecting in the cat-tail marshes
around Ithaca, New York, I observed a little ball of mud on the
upper surface of a leaf of Typha. The mud ball occurred near
the tip of the leaf, four to five feet above the surface of the
ground. I took the leaf with the mud ball into the laboratory
and upon breaking it open found a glistening white egg inside.
Plate VI, Fig. 5. The egg was put into a tin salve box for rearing.
The following day I found two more of the same kind of mud

* Contribution from the Department of Entomology, Cornell University.

95

'
i ‘

t

96 Annals Entomological Society of America [Vol. XII,

cells, each on a Typha leaf near the tip. These two specimens
also were brought to the laboratory and placed in the tin box
with the other egg.

On May 28th the first egg had hatched. The young larva
had opened one of the other mud balls and devoured the egg
within. The third mud ball was then removed to another tin
box. The first young larva died two days later, May 30th.
That left only one egg of the lot. This egg hatched on the first
of June, six days after it had been brought into the laboratory.
The young larva was transferred to another clean tin salve box
containing moist, sterilized sand.

Various kinds of food were offered to the young larva; larve
and pupe of Lymnecia phragmitella Stainton, larvee of Nonagria
oblonga Grote, Arsilonche albovenosa Goeze, and Arzama obliqua
Walk. Flies and fresh meat were also offered to the larva. At
first the young larva fed somewhat on flies and on larve and
pupe of L. phragmitella, but it showed a decided preference for
the little larvee of A. obliqua.

During the second and third instar all other food was refused
except larvee of A. obliqua,

All the above mentioned Lepidoptera are inhabitants of
Typha. A. albovenosa is an incidental surface feeder on the
leaves; L. phragmitella lives in the heads of Typha, while
A. obliqua and N. oblonga both begin their larval activities as
leaf miners, later becoming solitary stem borers.

It is possible that the larve of Chlenius impunctifrons feed
altogether on the larve of A. obliqua. The former hatch about
the same time that the larve of the latter species leave the
mines of the leaves of Typha to become stem borers. Thus
while they are exposed they might easily fall prey to the Carabid
larve.

On June 3, 11:00 A. M., the larva cast its first skin, the first
instar having lasted three days. June 6, at noon, the larva
molted again. The third instar larva had a voracious appetite
and consumed many of the little A. obliqua larve. Five days
later, June 11, the larva had tunneled into the moist sand and
hollowed out a little cell. In this cell it was lying on its side as
if ready to pupate. Having been disturbed the larva came to
the surface of the sand and remained there, lying on its back.
Whenever the box was opened it made efforts to turn over
but was unable to do so. When turned over the larva crawled

~
~
--

8.
1919] Claassen: Chlaenius Impunctifrons 7 97

pe

about a little but soon fell over on its back again. In this
condition the larva remained on top of the sand for sixteen
days without taking any food. On June 27th the larva pupated.
The pupa remained quietly on its back till July 4th, when the
adult emerged. The adult was rather weak. The legs, especially
the metathoracic legs, were not fully developed. The adult
specimen was determined by Dr. J. C. Bradley.

TABLE OF INSTARS.
LENGTH OF STAGE

ES OurG un Ceeeee ole tee sire ies i ING EE 772 Ee ee et Ee APA Sires oY ey
Bp eet chedewartyrser ae cs er: amen cere eno one ee Mee Stave edays place
PIGS GAOME ot coe dialed og a Hae oD CINA Clon co Saanom oad Be Memeo e a ol MoRCRENNae rh miaHOLAiASL
pecondemolita cry ate ache nee uel eee aoe eee eee ine SeCONGmnShanisicdanse
IPXERO EN Ola, oe eet Ae eee ime eeeeee ase ee ening imStareoieday.se
Emergence of adult.......... uli pe: ee Re ee eee eee eee Pupal period 7 days.

DESCRIPTION OF STAGES.

MUD CELL.
(Plate VI, Fig. 5).

The little mud ball which encloses the egg is made of fine gray sand.
It is placed singly on the upper surface of a Typha leaf six to eight
inches from the tip. The mud ball is oblong oval, somewhat depressed.
It measures 3.43 mm. in length, 2.28 mm. in width, and 1.75 mm. in
thickness.
EGG.
(Plate VI, Fig. 1).

In the center of the mud cell occurs the egg. It is oblong oval, of a
glistening white color, with rather large faint reticulations. It measures
2.28 mm. in length and 1 mm. in its greatest diameter.

FIRST INSTAR LARVA.
(Plate VI, Fig. 3).

Color. General color of larva dark brownish black, except in the
less chitinized places which are grayish white. Head yellowish brown
with a dark brown area in front between the epicraneal sutures. Antenne
and maxillary palpi dark brown, with a narrow yellow-brown ring at
the joints.

Form. The body is elongate, tapering to both ends, but more
posteriorly. The ninth abdominal segment bears two processes which
equal the combined length of the pro- and mesothorax. The anal
tube projects posterio-ventrad from the ninth abdominal segment.

Head. Head, excluding mouthparts and appendages, wider than
long, narrowed behind the eyes. Dorsal surface slightly convex, with
two short oblique grooves terminating at the lateral margin. Ventral
surface more convex. Edge of labrum roughly toothed, with one larger
tooth on each outer angle of the anterior margin. Surface of head smooth

98 Annals Entomological Society of America [Vol. XII,

with few hairs. Ocelli six, on a raised portion back of the antenne.
Antenne four-jointed, the first and fourth joint about equally long; the
second joint two-thirds the length of the first; the third joint slightly
longer than first, with a crook above the middle. Two or three bristles
originate from this outward crook or elbow, and one from the inner
side. Fourth, or apical joint, with four bristles at the tip.

Mandibles long curved, with a strong tooth below the middle,
directed inward and slightly downward. <A small bunch of hairs near
the inner base of mandibles.

Maxille with a long stout basal joint, bearing numerous hairs,
especially on the inner and upper surface. Outer palpus four-jointed;
first joint as wide as long, second joint somewhat more slender and at
least twice as long; third joint slender and a little shorter than second;
fourth joint as long as first, but very slender. Inner palpus two-jointed,
the basal joint stouter and a little longer than second. Just inside
the inner palpus is another small joint tipped with a bristle.

Mentum about as wide as long. Anterior margin produced to a
point bearing two hairs. Palpus with basal joint little longer than
broad; second joint little longer than first and tapering greatly.

Prothorax narrowing anteriorly, a third wider than long. Surface
finely punctate.

Meso- and Metathorax each twice as wide as long.

Abdomen of nine true segments, gradually tapering posteriorly.
The ninth segment bears the anal tube and the two posterior processes.
These processes each bear five long stout hairs, three about equidistant
along the process and two at the tip.

Legs long with stout hairs and two claws at the tip.

Measurements of Cast Skin. Length from tip of mandibles to tip
of caudal processes, 7.43 mm.; greatest width across thorax, 1.26 mm.

SECOND AND THIRD INSTARS.

The second and third instar larvee do not differ greatly in structure
from the first except in size and proportions. The first instar 1s more
spindle shaped. The body of the second and third instar is more
flattened.

SECOND INSTAR LARVA.
(Plate VI, Fig. 7).

The second instar larva differs from the first in the following points.

Antenne. First and third joint of equal length. Second and fourth
joint subequal, each a little shorter than first and third joint.

Palpi of mentum with basal joint twice as long as broad; second joint
as long as first.

Prothorax about twice as wide as long.

Meso- and Metathorax each more than twice as wide as long.

Caudal processes with four long hairs; one about half way back,
another three-quarters back, and two at the tip.

Measurements of Cast Skin. Length from tip of mandibles to tip of
caudal processes, 10.28 mm.; greatest width across thorax, 2.25 mm.

1919] | Claassen: Chlaenius Impunctifrons’ 99

THIRD INSTAR LARVA.
(Plate VI, Fig. 4).

Antenne. First, second and third joints of.equal lengths. Fourth
joint about two-thirds the length of ‘the other segments.

Prothorax less than twice as wide as long, i. e., the width equals
one and three-quarters the length.

Caudal processes as in the second instar.

Measurements of Cast Skin. Length from tip of mandibles to tip
of caudal processes, 14.57 mm.; greatest width across throax, 3.43 mm.

PWIPAS
(Plate VI, Fig. 2).

The pupa is of a creamy white color. The eyes are the first to show
a mottled brown color. The mandibles also become chitinized early.

Ventral view. Head bent under so as to show a small portion of the
prothorax. Labrum and clypeus together a third longer than broad,
emarginate in front. Mandibles long with a notch and tooth at the
apex. Maxillary and labial palpi plainly visible. Front of head with
four setae arranged as shown in figure. Antenne eleven jointed, the
tips almost reaching the tips of the metathoracic wings. Prothoracic
legs extending to a point half way between the tips of the maxillary
palpi and the tips of the antenne. Mesothoracic legs underneath the
antenne, extending slightly beyond the tips of the antenne. Meta-
thoracic legs visible only at the tips where they project from beneath
the metathoracic wings. Elytra show the longitudinal striz plainly.
The elytra extend about two-thirds the length of the metathoracic
wings. Wings with a hump on the lateral margin. They extend back
to a point one-third across the fourth abdominal segment. Abdomen
with segments two to nine visible. Segment two with a small hair-
tipped tubercle on each lateral margin. Segments three, four and five
each with lateral processes as shown in the figure. Segments seven and
eight smooth. Segment nine with two short posterior tubercles.

Dorsal view. Dorsally the pupa presents a hairy appearance.
Long hairs occur on all but the last three abdominal segments. On the
thorax the hairs are shorter and occur more scatteringly. On segments
one to six, of the abdomen, the hairs are long and are arranged in two
groups on each segment; one group on each side of the median line.
The hairs are arranged in two more or less distinct rows, thirteen hairs
in each lateral group, or twenty-six long hairs on each segment.

Measurements of the living pupa. Length, 10.28 mm.; greatest
width of abdomen not including the lateral processes, 4.57 mm.

ADULT.
(Plate VI, Fig. 6).

The adult which emerged from the pupa in the laboratory measured
13.7 mm. in length and 4.7 mm. in its greatest width. The original
description of the adult is found in the Transactions of the American
Philosophie Society II, 64, 1823.

100 Annals Entomological Society of America [Vol. XII,

EXPLANATION OF PLATE VI.

Chlaenius impunctifrons Say.

Fig. 1. Egg, length 2.28 mm.

Fig. 2. Pupa, ventral view, length 10.28 mm.

Fig. 3. Larva, first instar, length 7.438 mm.

Fig. 4. Larva, third instar, length 14.57 mm.

Fig. 5. Mud cell on leaf. Opened to expose egg inside.

Fig. 6. Adult beetle, length 13.7 mm.
Fig. 7. Larva, second instar, length 10.28 mm.

Figures 2, 3, 4, 6 and 7 all drawn to the same scale.

P. W. Claassen, Del.

VOL. XII, PLaTE VI.

ANNALS E.S. A.

P. W. Claassen

ROBBERFLIES OF THE GENUS ERAX.*
Sass Si INES

The species of this genus are flies of medium to large size.
Their larve, so far as known, live in the soil and are predaceous,
feeding on various forms of animal life present in such situations.
The adults prey on many kinds of insect life, and with other
genera of the same family may be considered as having the
same relation among insects as hawks among birds.

Scopoli proposed Erax as a generic name in 1763 and
included under it fourteen species. No genotype was proposed
until 1910, when Coquillett designated one of Scopoli’s species,
barbatus, as the type. As matters stand at the present time
all the species which Scopoli placed under Evax are located in
other well established genera and the genus in his sense is left
without a single representative.

In 1838 Macquart used Erax as a generic name. He gave
Scopoli credit for the genus, but the characterization he used
differs to such an extent that an entirely new group of species
was admitted, and some of these have long been known as
typical species of Erax. It is evident therefore, that oppor-
tunity for change of generic name is presented, but no action is
taken because I am satisfied to use Erax in Macquart’s sense
and thus obviate the changes which another interpretation
would involve. Since no-genotype has been named for the
genus in Macquart’s sense, Erax rufibarbis Macquart is desig-
nated as the type of Erax. It is the twenty-second species in
Macquart’s arrangement in Dipt. Exot., I, 2, 107-119.

As an attempt has been made to associate and study all the
North American species of the genus it was necessary to pro-
cure the loan of much material and to receive aid from various
sources. A generous response has made it possible for me to do
much more than could have been accomplished otherwise, and
my appreciation is most cordially extended to the following
institutions and students for services rendered:

Kansas University, Academy of Natural Sciences, National
Museum, Carnegie Museum, Museum of Comparative Zoology,
American Museum of Natural History, Biological Survey,
Charles W. Johnson, J. M. Aldrich, Frederick Knab, Nathan

_*Contribution from the Department of Zoology and Entomology, Ohio State
University. No. 57.

103

.
,

'
’

“Ae 7
~~

104 Annals Entomological Society of America [Vol. XII,

Banks, J. C. Bradley, Charles’Schaetier, R..H. Beamer, C> P:
Alexander, F. R. Cole and A. L. Lovett. Much material has
been procured also by purchase or exchange from others who
have expressed interest in the work.

Having at my disposal an extensive collection of the species
of Erax from a wide range of territory in North America, a
careful attempt has been made to properly identify the species
concerned. Types have been studied in all cases where such
were available. In some instances descriptions had to be used
entirely and since some of these descriptions, of older authors
especially, are brief and indefinite it is not impossible that
oversights have occurred. The external male genitalia are used
much in designating species. The vestiture of these organs
obstructs one’s view at times, but the dorsal part is less obscured
than the ventral, so the former has been outlined more care-
fully, and is the part to which students are referred as an aid to
specific determination.

It has not been possible to reach a conclusion in regard to
the identity of some of the species so the following names are
not included in the descriptive part of the paper:

Erax comatus Bellardi was not identified in the material
studied. It belongs to the carinatus group and is described as
25 millimeters in length.

Erax aper Walker very likely belongs to the carinatus group
also but it was not identified. It was described from Mexico
and may be one of the species mentioned in the paper but its
description is indefinite.

I consider Erax pumilus Walker and Erax disjunctus Williston
the same as Eristicus nigripes Bellardi. This species 1s consid-
ered in Trans. Am. Ent. Soc. XLIII, 297, under Ezcherax
nigripes. ‘The insect is not mentioned either by Osten Sacken
or Williston in their works on Central American Diptera.

Erax pavidus Williston, from Mexico, appears to be well
characterized but it was not identified in the material studied.

Erax lascivus Wiedemann, Asilus amarynceus Walker,
Asilus flavofasciatus Wiedemann, Asilus macularis Weidemann
and Asilus antiphon Walker, most likely are all species of Erax.
They have been reported from North America and most of them
from South America also. Records of their occurrences are
meager and they have not been identified in recent years.

1919} ‘ Hine: Genus Erax 105

LIST OF NORTH AMERICAN SPECIES OF ERAX. WITH SYNONYMY.

The barbatus Group.

armatus new species.
barbatus Fabricius, Syst. Antl., 169, (Dasypogon), 1805.
pogonias Wiedemann, Dipt. Exot., 198, (Asilus), 1821; Auss. Zw. I, 460,
(Asilus), 1828.
albibarbis Macquart, Dipt. Exot. I, 2, 118, 1838.
cinerascens Bellardi, Saggio, II, 39, tab. II, fig. 10, 1861.
tricolor Bellardi, Saggio, II, 40, tab. II, fig. 12, 1861.
furax Williston, Trans. Am. Ent. Soc. XII, 67, 1885.
bicolor Bellardi, Saggio II, 47, 1861.
bimaculatus Bellardi, Saggio II, 45, tab. II, fig. 11, 1861.
cingulatus Bellardi, Saggio, II, 41, 1861.
grandis new species.
leucocomus Williston, Trans. Am. Ent. Soc. XII, 69, 1885.
quadrimaculatus Bellardi, Saggio, II, 44, tab. II, fig. 13, 1861.
sagax Williston, Biol. Centr. Am., Dipt., I, 324, 1901.
tagax Williston, Trans. Am. Ent. Soc. XII, 65, 1885.
similis Williston, Trans. Am. Ent. Soc. XII, 68, 1885.
willistoni new species.
zonalis new species.

The aridus group.

aridus Williston, North Am. Fauna, No. 7, 254, 1893.
harveyi new species.

prattii new species.

snowi new species.

subpilosus Schaeffer, Jr. N. Y. Ent. Soc., XXIV, 67, 1916.

The anomalus group.

anomalus Bellardi, Saggio, II, 32, tab. II, fig. 7, 1861.
candidus Coquillett, Can. Ent. XXV, 176, (Efferia), 1893.
pernicis Coquillett, Can. Ent. XXV, 175, (Efferia), 1893.

The aestuans group.

aestuans Linneaus, Syst. Nat. XII, II, 1007, (Asilus), 1767.
aestuans Fabricius, Syst. Antl., 164, (Dasypogon), 1805.
niger Wiedemann, Dipt. Exot., 196; Auss. Zw., I, 460, (Asilus), 1821.
macrolabis Wiedemann, Auss. Zw., I, 458, (Asilus), 1828.
aestuans Macquart, Hist. Nat. Dipt., I, 312, (Asilus), 1834.
mcisuralis Macquart, Dipt. Exot., I, 2, 117, 1839.
bastardi Macquart, Dipt. Exot., I, 2, ina plate 9, fig. 7, 1839.
tubtalis Macquart, Dipt. Exot., I, 2, 118, 1839.

affnis Bellardi, Saggio, II, 41, 1861.
marginatus Bellardi, Saggio, II, 46, 1861.

apicalis Wiedemann, Dipt. Exot., 191 (Asilus), 1821; Auss. Zw., I, 448, (Asilus),

1828.

vicinus Macquart, Dipt. Exot., Supl. I, 85, 1846.
aurimystaceus new species.
belfragei new species.
femoratus Macquart, Dipt. Exot. I, 115, 1838.
fulvibarbis Macquart, Dipt. Exot. , Supl. 3, 28, tab. 2, fig. 18, 1848.
haloesus Walker, List II, 405, (Asilus), 1849: List Valls 624, 1855.
kansensis new species.
loewii Bellardi, Saggio, Appendix, 21, tab. III, fig. 17, 1862.
dolichogaster Williston, Biol. Centr. Am., Dipt. I, 326, tab. 6, fig. 6, 1901.
mexicanus new species.

106 Annals Entomological Society of America [Vol. XII,

nigrimystaceus Macquart, Dipt. Exot., Supl. 2, 1847.

nigritarsis new species.

portoricensis new species.

rufitibia Macquart, Dipt. Exot., Supl. 3, 42, tab. II, fig. 11, 1848.
fortts Walker, List. VII, 623, 1855.

slosson# new species.

stylatus Fabricius, Syst. Ent. 795, (Asilus), 1775.
haitensis Macquart, Dipt. Exot., Supl. 3, 28, tab. IT, fig. 10, 1848.
caudex Walker, List II, 404, (Asilus), 1849.
invarius Walker, Ins. Saund., Dipt. I, 131, 1851.

*tabescens Banks, new species.

The carinatus group.

carinatus Bellardi, Saggio, II, 36, tab. II, fig. 9, 1861.
comatus Bellardi, Saggio, II, 34, 1861. Not identified in my material.
concinnatus Williston, Biol. Cent. Am., Dipt.. I, 323, 1901.
costalis Williston, Trans. Am. Ent. Soc., XII, 64, 1885.
cressoni new species.
jubatus Williston, Trans. Am. Ent. Soc., XII, 66, 1885.
prolificus Osten Sacken, Biol. Cent. Am., Dipt. I, 202, 1887.
latrunculus Williston, Trans. Am. Ent. Soc., XII, 67, 1885.
parvulus Bellardi, Saggio, II, 35, tab. II, fig. 8, 1861.
subcupreus Schaeffer, Jr. N. Y. Ent. Soc., XXIV, 66, 1916.
unicolor Bellardi, Saggio, II, 37, 1861.

The tuberculatus group.

productus new species.
spiniventris new species.
tuberculatus Coquillett, Jr. N. Y. Ent. Soc., XII, 34, 1904.

The rufibarbis group.

bicaudatus new species.
rufibarbis Macquart, Dipt. Exot., I, 2, 116, 1838.
aestuans Wiedemann, Dipt. Exot., 200, (Asilus), 1821, (not Linneaus); Auss.
Zw., I, 467, (Asilus), 1828.
completus Macquart, Dipt. Exot., I, 2, 117, tab. IX, fig. 9, 1838.
dascyllus Walker, List II, 401, (Asilus), 1849. :
virginianus van der Wulp, Tijdschr. v. Entom. XXV, 109, tab. X, figs. 5 and 6,
(Proctacanthus), 1882.
ravus Coquillett, Canadian Entomologist, XXV, 176, (Efferia), 1893.

The stramineus group.

argentifrons Hine, Ohio Naturalist, XI, 308, 1911.
argyrogaster Macquart, Dipt. Exot., supl. 1, 84, 1846.
argyrosoma Hine, Ohio Naturalist, XI, 310, 1911.
auripilus Hine, Ohio Jr. Science, XVII, 22, 1916.
aurivestitus new species.

californicus Schaeffer, Jr. N. Y. Ent. Soc. XXIV, 67, 1916.
canus Hine, Ohio. Jr. Sci. XVII, 22, 1916.

coquillettii new species.

dubius Williston, Trans. Am. Ent. Soc., XII, 64, 1885.
eximeus Bellardi, Saggio, II, 38, 1861.

inflatus Hine, Ohio Naturalist, XI, 310, 1911.

* Among a number of species received from the Museum of Comparative
Zoology were two species which Dr. N. Banks had named and the descriptions
in manuscript were turned over to me. The two species are Erax tabesuns and
Erax texanus, and Dr. Banks is credited with these, although they are published
for the first time in this paper.

1919] Hine: Genus Erax 107

interruptus Macquart, Hist. Nat. I, 310, (Asilus), 1834.

maculatus Macquart, Dipt. Exot., I, 2, 111, tab. IX, fig. 6, 1838.
lateralis Macquart, Dipt. Exot., I, 2, 116, 1838.

ambiguus Macquart, Dipt. Exot., supl. 1, 84, 1846.

villosus Bellardi, Saggio, II, 49, (Eristicus), 1861.

nemoralis Hine, Ohio Naturalist, XI, 311, 1911.

pallidulus Hine, Ohio Naturalist, XI, 309, 1911.

pilosus new species.

plenus Hine, Ohio Jr. Sci., XVII, 21, 1916.

rapax Osten Sacken, Biol. Centr. Am., Dipt., I, 201, 1887.
splendens Williston, Biol. Cent. Am., Dipt., I, 325, tab. VI, fig. 5, 1901.
stramineus Williston, Trans. Am. Ent. Soc., XII, 68, 1885.
texanus Banks, new species.

triton Osten Sacken, Biol. Centr. Amer., Dipt., I, 200, 1887.
truncatus Hine, Ohio Naturalist, XI, 309, 1911.

varipes Williston, Trans. Am. Ent. Soc., XII, 71, 1885.

bo

KEY TO GROUPS OF ERAX.
Furcation of the third vein distinctly before the base of the second posterior

cellimating the Second) submarginal cell long. 2.:... 2.02.00. 0050.2.. 40.0. 2
Furcation of the third vein opposite or bevond the base of the second posterior
Cates Serer tee) See RRM ioA RI ANG SRE Stine HREPRS oh crnePeeR eas wera 4
Three submarginal cells. ............. Ler spe A cdeste th Sota The anomalus group
A WORSUD Ina LeUT alae ell Sin wet Kamen sites crak sa eteivk: Sie ioc cies, oe Sic weiats Me wl aides vad 3

Rear of thorax and. scutellum with abundance of bristles, male abdominal
segments six and seven silvery, female with tip of oviduct from above
IROHROAHE Ds tote Ina auth estat lS dtaed auth ast Ge Act anc ge ee The rufibarbis group

Male abdomen usua!ly with several segments silvery, female with tip of ovi-
duct from above never divided. Male, except in a very few cases, with
one or more abdominal segments bearing long white hair parted at the

muiddlevandudinected outward. <..))..4..9602 sls oe The stramineus group
Posterior branch of the third vein curved backward at tip, meeting the costa
ator fMeyond! the apex Of Che* Win: 3.0.5. ,000nn. The aestuans group
Posterior branch of the third vein curved forward, meeting the costa plainly
WELOKe mui CkanexTOmulenWwalloraa sien e NAC AMA nt hesemninu team aamie me emi eecme Sao oe 5
Mesothorax compressed anteriorly and with a conspicuous crest of erect
haimswon) phe waiatd=-dorsal Wine a: 6.08 ss. dee ee The carinatus group

Mesothorax broadly rounded above and without a conspicuous crest of hairs. .6
Mesothorax with numerous bristly hairs which are unusually long, scutellum
conspicuously hairy and with numerous marginal bristles. .The aridus group
Mesothorax with very short hair anteriorly, a few rather prominent bristles
posteriorly. Scutellum with short hair on the disc and usually with not

More wuan halt arr«dozenibristlesiom 1S maneine. 9.. 26.4. eee dees ace ote. 7
Male with ventral prominences on abdominal segments four, five and six,
Sinclil@slender species bcs. is oes ekoeb cs The tuberculatus group

Male abdomen without ventral prominences usually large or medium sized,
LODUSUMSPECIES spout cS oe airois ice eutiocans <a viev sid UB eheate alg The barbatus group

108 Annals Entomological Society of America [Vol. XII,

The barbatus Group.

The members of this group have a broad mesothorax, quite evenly
curved, short-haired all over, especially anterior to the transverse
suture. Only a few prominent bristles above the attachment of the
wings and posteriorly. Scutellum short-haired on the disc, and usually
with not more than half a dozen bristles on the margin, although
occasional specimens may have one or two more than this number, in
most cases there are less. Branching of the third vein never anterior
to the base of the second posterior cell, occasionally it is opposite the
base of this cell, but in nearly all cases it is plainly beyond, making the
second submarginal cell short. The posterior branch of the third vein
bends forward, and reaches the costa distinctly before the apex of the
wing.

In some species the male has the costa thickened and expanded on
the anterior margin, somewhat before the apex, but in other species
this is not the case.

A species which is typical of this group is what in recent
years has been called albibarbis, but here it is considered the
same as barbatus which is an older name.-as indicated in the list
of species on another page.

1. Hypopygium small or of normal size, medium sized to small species, usually
of slender build, male costa not at all or only very shghtly expanded,
female with each abdominal segment black, with posterior gray border
Ofvamloust widthsraccOncdingetOnSpeCiGS amar si aii eee ie eae 2

Hypopygium distinctly enlarged, large to very large species, male costa
rather conspicuously thickened and expanded, female abdomen variously
marked, but usually the segments are not black with posterior gray
borders, although this is the:casein at least two Speciesa.....4-4-m-e 5

Hypopygium distinctly enlarged, costa not at all thickened and expanded,
mystax straw yellow in both sexes, bristles of the thorax and scutellum
pale, abdomen uniformly yellowish gray pollinose, male with the apical
abdominal segments partly or wholly silvery........ leucocomus Williston

2. Gray bands of the abdomen regular and wide, approaching the width of the
black intervals between them, hypopygium red............ zonatus n. sp.

Hypopygium black, gray abdominal bands irregular or very narrow......« 3

3. Mystax pale, sometimes with a few black bristles on upper margin,

barbatus Fabr.

Mystax largely black with pale hairs and bristles intermixed............. 4

4. Sixth and seventh segments of male abdomen silvery, pale bands of the
abdomen brownish gray, with front margin more or less irregular,

cingulatus Bellardi
Fifth, sixth and seventh segments of male abdomen silvery; pale bands of

the abdomen silvery, regular and narrow............. willistoni n. sp.

5. Male hind tibiae with an abnormal NEES: Pe teak orl tc ok AOA eA 6
Male hind tibiae normal: o.oo. o-h cero ae eoer o eaoeee a eeeie 8

65) Wines quite distinctly browne qe ue oer eee ee sagax Williston
Wangs ny allies a. 8) esc cs aie eiete ots mS OE te ee 7

a. Apical third votithe male tibia enlarzedie sss eee cee. tagax Williston
Apical half or more of male tibia enlarged................. armatus Hine

8. Male seventh abdominal segment entirely silvery dorsally; female fifth
abdominal sesmentimot Silvery... ))- ae eee eee eee Eee eee 9

Male seventh abdominal segment clear black on the greater part of the
dorsum; female fifth abdominal segment broadly silvery on each side,
with a black stripe on the middle of the dorsum:...................... 10

1919} Hine: Genus Erax 109

OmaWanesihyalinesnopunusually lange... .... 1. sae ae bicolor Bellardi
Wings brownish, especially toward the apex; very large species. .grandis n. sp.
10. Mystax straw yellow with a few black bristles above, wings pale yellowish

[NCO 5 Joop ooSo oo SOL Ome Ho pen COMO e Goce occce bimaculatus Bellardi
Mystax black with a few pale bristles intermixed; wings fuscous, more
apparentacowardsttumerapexse.s....-....+s.eearee quadrimaculatus Bellardi

Erax barbatus Fabricius.

Length 13 to 20 millimeters. Male. All vestiture of the head white
to pale yellowish. Quite often a few black bristles in the mystax and
occasionally on the palpi. Thorax pale pollinose, not always of the
same shade, markings very obscure, anteriorly with short pile variable
in color, posteriorly with a number of black pale bristles. Scutellum
with sparse pale pile on the disc and a few black or pale bristles on the
margin. Femora black in ground color obscured by white pollen and
hair, apex of each tibia and whole of tarsi largely black, tibia otherwise
pale yellowish, bristles of the legs variable from black to pale, wings
slightly -yellowish, nearly hyaline, costa not dilated, anterior branch of
the third vein with a distinct stump variable in length in different
specimens, basal section almost in direct line with the cross vein at
the base of second posterior cell. Abdomen dorsally, first and second
segments pale pollinose, usually a rather small rounded blackish spot
on each side of the second; third, fourth and fifth largely black with
posterior margins and often median stripe pale pollinose; sixth and
seventh silvery. Hypopygium black. See Figures 2 and 8.

Female colored like the male, abdominal segments six and seven
colored like the preceding in most cases, but sometimes the segments
are nearly wholly gray pollinose, oviduct shining black.

This species is the most widely distributed of any of the
genus. It ranges from Guatemala and Lower California to
Washington and New England and often is the first of the
genus taken in any locality within its range.

The uniform color of the thorax without apparent markings
and the two rows of black spots on the abdomen make its
determination easy. Although the two longitudinal rows of
black abdominal spots are present in most cases there is varia-
tion, some specimens having these spots much reduced while
others have them enlarged until nearly the whole abdomen is
black. The two basal segments, however, most always are
uniformly gray and the other segments have at least a narrow
middorsal stripe and posterior border on each, the same color.

The possibility of more than one species in the material
used has been considered but it does not appear possible to
find characters for satisfactory separation, so it is preferable
to include all in a single species.

110) 0 Annals Entomological Society of America [Vol. XII,

Erax bicolor Bellard1.

Total length 21 to 26 millimeters; vestiture of the head largely
yellowish white, some black bristles in the mystax; bristles of the palpi
almost wholly black. Whole body nearly uniformly gray pollinose,
mid-dorsal stripe of the thorax quite plainly black and divided length-
wise by a gray interval. In some specimens rather obscure darker
spots on some of the abdominal segments. Wings hyaline costal
border dilated, furcation of the third vein plainly beyond the base of
the second posterior cell, basal three-fourths or more of the tibia pale
yellowish, otherwise legs black with black bristles. Sixth and seventh
abdominal segments silvery, hypopygium large and black. See Figure 9.

Female colored like the male, except abdominal segments six and
seven are velvety black. Oviduct black, about as long as segments
four to seven.

Habitat: Mexico, Texas, Mississippi, New Mexico and
Arizona. Twenty-four specimens.

Bellardi described the species from a female and mentioned
the black sixth and seventh abdominal segments which seems to
be a peculiarity of this species. It is more evident in some
specimens than others.

Erax willistoni n. sp.

Male. Total length 22 millimeters. Very dark colored species;
mystax black with a few pale hairs intermixed, beard white, palpi
with black bristly hairs. Thorax thinly gray pollinose, mid-dorsal
stripe and irregular spots on either side black, with black hair and
bristles, scutellum gray pollinose with pale hairs on the disc and about
eight black bristles on the margin. Wings nearly hyaline, very slightly
tinged with brown on apical half; costa somewhat dilated, branching
of the third vein beyond the base of the second posterior cell, stump
very short. Legs with black bristles and pale hairs, extreme apexes
of femora, tibize and tarsi entirely dark reddish brown, remainder of
femora black. Abdominal segments two, three and four black above,
narrow posterior margin and sides gray pollinose, segments five, six
and seven silvery, hypopygium elongate, emarginate at apex, black
with black vestiture. See Figure 7.

Female. Total length 23 millimeters. Colored like the male with
segments five, six and seven like those that precede them, ovipositor
black, as long as the last four abdominal segments.

Type male and allotype, on the same pin, collected at
Williams, Arizona, July 21, by H. Barber. Deposited in the
U.S. National Museum. Five other specimens from Colorado,
Arizona and Chihuahua, Mexico, by Tucker, Barber and
Townsend.

1919] Hine: Genus Erax 111

This species looks like estuans but the wing venation, color
of the legs and the male genitalia characterize it. It is the
only species of its group with the femora reddish brown at the
apexes although some specimens do not show the character
very plainly.

Erax cingulatus Bellardi.

Total length 20 to 25 millimeters. Male. Mystax largely black,
but intermixed with pale yellowish hairs; beard pale yellowish; palpi
with numerous black bristly hairs. Much of the thorax and abdomen
brownish pollinose, thorax with a wide mid-dorsal stripe and spots on
either side black, vestiture mostly black, but a few pale hairs posteriorly,
scutellum and extending to the posterior part of the mesothorax ash
gray, conspicuously contrasting in color with the rest of the thorax,
wings tinted with brownish, costa very slightly dilated, furcation of the
third vein distinctly beyond the base of second posterior cell, stump
shorter than the basal section, three-fourths or more of tibiz from bases,
pale yellowish, otherwise legs black, except some of the basal tibial
segments which are dark brown. Some of the bristles of the legs are
pale in some specimens, but most of them are black, abdominal segments
two, three, four and five black anteriorly and pale brownish pollinose
posteriorly, the divisions between the colors irregular and not sharply
defined, segments six and seven silvery. Hypopygium black with
black vestiture, rather large. Figure 34.

Female colored like the male except abdominal segments six and
seven are like those that precede them. Ovipositor black, slightly
longer than the last three abdominal segments.

Seven specimens from Cuernavaca, Mexico, collected by
DD: bh. Crawtord.

The ash-colored scutellum and space before it contrasting
with the rest of the thorax is characteristic of this species, but
only well preserved specimens show it. There are only two to
four black bristles on the margin of the scutellum.

Erax grandis n. sp.

Male. Total length 33 millimeters. Mystax composed of numerous
yellowish gray hairs and black bristles, beard pale yellowish gray,
palpi with black and pale hairs, thorax gray pollinose with unusual
markings, scutellum with black hairs on its disc and four black bristles
on its margin, femora, apexes of tibize and tarsi black, remainder of
tibie pale yellow, wings slightly brownish, costa distinctly dilated,
furcation of third vein plainly beyond the base of the second posterior
cell, stump very short, two, three, four and five abdominal segments
black above, with narrow posterior margin and sides gray, segments
six and seven silvery. Hypopygium large, black with black vestiture.

112 Annals Entomological Society of America [Vol. XII,

Female. Total length 31 millimeters with tip of ovipositor broken
off, probably 34 or 35 millimeters when complete, colored like the male,
but segments six and seven not silvery, stump of the anterior branch of
the third vein much longer than in the other sex.

Type: Male and allotype, from Round Mountain, Texas,
deposited in the museum of the~Philadelphia Academy of
Natural Sciences.

This is the largest Erax I have ever seen. It suggests bicolor
but aside from its being much larger the costa is more strongly
dilated, the disc of the scutellum has black hair instead of
white, and the male genitalia are different.

Erax zonatus n. sp.

Male. Total length 16 millimeters. Vestiture of the head largely
white, several black bristles in the mystax and numerous black hairs
on the palpi. Tibiz red except at extreme apex, tarsi largely red, but
somewhat darker than the tibize and furnished with numerous black
bristles, femora black. Thorax quite uniformly yellowish gray pollinose
with the usual marking very obscurely shown. Wings very slightly
tinged with yellowish, nearly hyaline, costa just a little dilated, branching
of the third vein plainly beyond the base of second posterior cell, stump
slightly longer than the basal section. Abdomen with four black bands
as follows: One on the middle of the second segment and one on each
anterior margin of segments three, four and five. These black bands
all are wider than the gray bands following them. Segments six and
seven silvery. Hypopygium red, prominent ventrally, more so in
appearance because of a ventral tuft of chestnut brown hairs.

Female. Total length 18 millimeters. Colored like the male except
the sixth abdominal segment is banded with black like five, and seven
is black with a very narrow gray posterior margin, ovipositor black,
about as long as abdominal segments five, six and seven.

Type: Male and allotype from Southern Arizona, F. H.
Snow, August, 1902. Collection of the author. Ten other spec-
imens, some of them from the collection of the Academy of
Natural Sciences, Philadelphia. Habitat, San Antonio Canyon,
California, Southern Arizona and Alamogordo, New Mexico.

Easily known from all species of its group by the zonate
appearance of the abdomen, and the prominent array. of
ventral hairs on the hypopygium, most conspicuous basally.

Erax armatus Hine.

There are three species of Erax with the hind tibize modified in the
male. All are larger than the average. In the male the furcation of
the third vein is beyond the base of the second posterior cell, while in
the females it is usually about opposite the base of that cell. Stump

1919} Hine: Genus Erax £13

of the anterior branch always present. The females are easily associated
with the males, but have no very good characters for distinguishing
them from others of the same sex. The wings are hyaline in armatus,
at least the yellowish tinge is hardly evident. The tibia of the male
shows an angular prominence posteriorly, somewhat nearer its base
than its apex, the female has the first five abdominal segments rather
densely gray pollinose, with rounded black spots above on two, three,
four and five; segment six is nearly all black above and seven is entirely
black. Femora, tarsi and apexes of tibiz black, basal three-fourths or
more of each tibia pale yellow. Total length 30 to 32 millimeters.

Ten specimens from Davis Mountains (F. M. Gaige)
Cotulla (F. C. Pratt) and Uvalde, Texas. Taken in June and

July:

Erax sagax Williston.

Total length 25 to 27 millimeters. Wing wide and conspicuously
light brown all over in both sexes; male with the hind tibia suddenly
enlarged beyond the middle of its length; both sexes with silvery
markings on the abdomen. In the male, segments two and three are
narrowly silvery on the sides, four, five and six are largely silvery, but
four has a rather large black triangular spot above and five has a narrow
black marking above anteriorly, six entirely silvery, seven black above
with narrow sides silvery. :

Female with segments two, three, four and five silvery at the sides;
this color is extended dorsally on the hind margin of three and on the
whole of four and five, but 1n no case does it meet above so there is a
conspicuous black space left dorsally on each segment; six and seven
nearly entirely black. The femora are red apically in both sexes, and
posteriorly in the female, in the specimens at hand.

Three specimens from Guadalajara, Mexico, June 24, 1903,
collected by McClendon.

Erax tagax Williston.

Total length 28 to 30 millimeters. Wing very pale brownish.
Male hind tibia with a distinct prominence posteriorly near distal third
and a conspicuous patch of black pile anteriorly just opposite and
extending to apex. Abdominal segments of the male two, three, four
and five black with narrow sides and posterior margin silvery white
pollinose, six wholly silvery, seven mostly silvery, but with posterior
margin above dark. Female, segments two, three, four, five and six
black, with sides and narrow posterior margin white pollinose, seven
black, oviduct black not much longer than abdominal segments six and
seven. Femora black in the male and mostly redish posteriorly in
the female in the specimens at hand.

A dozen specimens from Arizona.

114 Annals Entomological Society of America [Vol. XII,

There is no record that Erax similis has been recognized
since it was described by Williston and we have good reason to
believe that it is the female of tagax.

Erax leucocomus Williston.

A large pale species with bristles and hairs of head, thorax and
abdomen all pale yellowish; legs usually with black femora and black
bristles throughout; wings delicate yellowish, veins quite distinctly
yellow on the basal part. Total length 25 to 30 millimeters. Male
wing with the furcation of the third vein beyond the base of the second
posterior cell, stump about as long as the basal section, costa not
dilated in the least. Hypopygium large from dorsal view gradually
widened toward the tip, where it 1s much wider than in other species
of its group. The abdomen in both sexes is pale yellowish gray without
markings, but somewhat changeable according to the direction from
which it is viewed. In the male, segments six and seven may be said
to be silvery and in the female segment seven is somewhat darker than
the others. Oviduct black. Nearly as long as the last three abdominal
segments.

Ten specimens from Western. Kansas, collected by F. X.
Williams and one from Dallas, Texas, taken September 5, 1905,
by F. C. Bishop.

The species is very distinct and readily recognized but ap-
pears to be rare and I had two species questionably referred to |
leucocomus before I procured it. It would not be strange if some
specimens would have some black bristles on the body. It is
well adapted to life on the sand on account of its color.

Erax quadrimaculatus Bellardi.

Total length 28 to 34 millimeters. Mystax rather sparse, composed
largely of black bristles with a few pale ones and pale hairs intermixed,
beard nearly white. Whole body black and appearing darker than
other species of Erax. Thorax very sparsely gray pollinose with black
bristles and hairs, scutellum with short black hairs on the disc and two
to four black bristles on the margin; femora, tarsi and apexes of the tibiz
black, remainder of tibize pale yellow, wings uniformly fuscous all over,
costa only slightly dilated in the male, furcation of the third vein clearly
beyond the base of the second posterior cell. In the male abdominal
segments two, three, four and five with sides narrowly gray pollinose,
extended dorsally on the posterior margin of each, but not uniting
across the dorsum; segment six silvery on each side, widest before
leaving a black dorsal triangular spot with its base on the posterior
margin; segment seven similar, but the silvery areas much reduced and
the black consequently extended. Female segments two, three and four
as in the male, five conspicuously silvery on each side, black on the
mid-dorsum; six and seven velvety black, oviduct black, hardly as long
as abdominal segments five, six and seven.

1919} Hine: Genus Erax dek5

Several specimens from Colombia, South America, from the
Carnegie Museum of Pittsburgh. Other specimens from Mexico.

Erax bimaculatus Bellardi.

Total length 25 to 28 millimeters. Mystax and beard pale yellow,
the former with a few black bristles above; thorax brown pollinose
with the usual markings; wings yellowish, costa of male distinctly
dilated, femora black, remainder of legs yellowish red with black
bristles. Abdomen colored much as in quadrimaculatus in the male seg-
ments four and five entirely silvery, or at most with a very narrow mid-
dorsal stripe, segment seven silvery on front margin and on the sides, black
otherwise, female with abdominal segments two, three and four white
pollinose on the sides, fifth white with the exception of a mid-dorsal
stripe, sixth and seventh velvety black, oviduct shining black.

The yellow mystax and beard readily separates this species
from quadrimaculatus. The two species are entirely distinct
although similarly colored.

‘Several specimens from Guadalajara, Mexico, collected by
McClendon, and from Cuernavace, Mexico, collected by D. L.
Crawford.

The aridus Group.

A few species from the more or less arid regions of western United
States are much like members of the stramineus group, but the branching
of the third vein is almost exactly opposite the base of the second
posterior cell. The following key may be used to separate the species.

emai stamesiinawayiell@wen aes lAreder Gkunine 2 ASLI cattle th Mache tite Sci whadlehe Sheed 2
INL Siac AVENE ha Sets og o eawed RlaiD eee ee hens SEMA s ERR ete Eee ee eee ere ne rer 3

2. Male abdominal segments six and seven, silvery.............. harveyi n. sp.
Male abdominal segments four, five, six and seven silvery. Four more or less
Dlackyonsthesmicd=dorsalilimetes sy. Varia cis seo Actes shige ce ys ss snowi n. sp.

3. Hypopygium from dorsal view plainly widened towards apex where it is dis-
TiNChiyer wider timane AGM DASEL jer Jn aclooe pies © atic as scares aot prattii n. sp.
Hypopygium not or but very slightly wider at apex than at base........... 4

4. Bristles of the body and legs nearly all white, male first and second abdominal
SECIMENESIO AC Kmeninyeys ar his cy aici: ashore cpa einne subpilosus Schaeffer

Male first and second abdominal segments white hairy..... aridus Williston

Erax harveyi n. sp.

Male. Total length 18 millimeters. Mystax, beard, ocellar and
occipito-orbital bristles pale straw yellow, palpi black and yellow
haired, anterior part of dorsum of mesothorax black haired, remainder
of thorax with pale vestiture, scutellum with numerous pale yellow
marginal bristles; wings hyaline, furcation of the third vein exactly
opposite the base of the second posterior cell, stump about as long as
the basal section; legs faded reddish, mostly with pale vestiture, but
an occasional black bristle, femora quite distinctly darker than the
other parts. Abdomen pale haired all over, general color dark, thinly

116 Annals Entomological Society of America [Vol. XII,

gray pollinose, more dense in some places than others giving a mottled
appearance from some views; sixth and seventh segments silvery,
hypopygium rather small, dark colored from dorsal view, sides nearly
parallel, angular at extreme apex.

Female. Total length 21 millimeters. Colored like the male.
Sixth and seventh abdominal segments, from side view, wedge shaped
at tip, from dorsal view not furcate apically.

Type: Male and allotype, from Vernon, B. C., August 11
and 15, 1904. Collected by the late Captain R. Valentine Har-
vey for whom the species is named.

This species suggests bicaudatus but the structure of the
oviduct is very different and the second submarginal cell is
shorter.

Erax snowi n. sp.

Male. Total length 22 millimeters. All the bristles of the head
except four bristles near the ocelli pale yellowish tinged, thoracic
dorsum and disc of scutellum black hairy, bristles near the wing roots
and on the margin of the scutellum pale yellowish, pleuree with pale
hair, legs largely black with abundant long pale hair and black and
pale bristles, basal half of each front and middle tibia and basal fourth
of each hind tibia reddish, although not conspicuously so. Wings
faintly yellowish hyaline, furcation of the third vein nearly opposite
the base of the second posterior cell, stump about the length of the
basal section, first three abdominal segments mostly black, thinly
yellowish pollinose laterally, with black hair above and pale yellowish
and rather long hairs at the sides; fourth segment narrow black before
and on the middle of the. dorsum, white pollinose and long white haired
laterally, fifty, sixth and seventh segments silvery and with a few short
white hairs.. Hypopygium black, mostly with pale vestiture, of medium
Size.

Female. Size and coloration very nearly as in the male. Abdomen
entirely yellowish gray pollinose-and pale haired, from some views
more or less shining; oviduct black, scarcely as long as the last three
abdominal segments.

Type: Male and allotype, from Clark County, Kansas, 1962
feet elevation, taken in May by F. H. Snow, for whom the
species is named. In the author’s collection. More than fifty
other specimens from Kansas, Colorado, New Mexico and
Texas, show some variation in size, and a little in color of
bristles, but may be said to be quite uniform.

Some specimens have more black bristles on the posterior
part of the mesonotum and on the margin of the scutellum than
the: sty pe:

The color of the abdomen distinguishes the species from
all others except plenus which has very different male genitalia.

1919] Hine: Genus Erax LL

Erax aridus Williston.

The female type of aridus isin the National Museum. There does
not appear to be any male specimens at hand. The species is peculiar
in being very hairy and bristly and Williston described it as belonging
near latrunculus, but it surely is very distinct from that species. True
there is a row of rather long black hairs in the middle of the anterior
part of the dorsum of the mesothorax, but the thorax is not compressed
and it is almost sure that when the male of aridus is finally located
it will not have a dilated costa which is always present in the carinatus
group. Palpi with black hair, vestiture of remainder of head pale,
mesothoracic dorsum with black hair, scutellum with light hair, bristles
near the wing bases pale, pleuree pale haired, legs with abundance of
long pale hairs, and mostly. pale bristles, legs black, base of each tibia
with just a suggestion of reddish, wings hyaline, furcation of the third
vein nearly opposite the base of the second posterior cell, stump dis-
tinctly longer than the basal section, abdomen black, gray pollinose,
more plainly so laterally and on the narrow posterior border of each
segment. Oviduct black, short, only a little longer than the last two
abdominal segments.

The short oviduct readily separates aridus from what is
considered as the female of subpilosus a nearly related species.
For a time the possibility of the type of swbpilosus being the
male of aridus was considered but there seems to be good reason
for separating them as distinct. Length about 25 millimeters.

Williston’s type was taken by the Death Valley Expedition
in April, 1891.

Erax prattii n. sp.

Male. Total length 16 millimeters. Mystax and beard white,
palpi with black bristles, front mostly with pale hairs, several bristles
near the ocelli, some black and the others yellow, occipito-orbital bristles
pale yellowish. Bristles and hairs of the mesothoracic dorsum black,
pleurze white hairy; legs abundantly long white hairy and with black
and pale bristles, femora black, tibize and tarsi mostly pale brownish;
wings hyaline, costa not at all dilated, branching of the third vein
almost exactly opposite the base of the second posterior cell, stump
scarcely as long as the basal section. Abdominal segments one to five
inclusive black, each with a narrow gray border of uniform width,
segments six and seven silvery. Hypopygium from dorsal view grad-
ually widened from base to apex, where it is nearly truncate and dis-
tinctly wider than in most other species. See Figure 13.

Male type from Laredo, Texas, November 25, 1905. Col-
lected by F. C. Pratt, from whom the specimen was procured.
Collection of the author.

The size and the width of hypopygium make it easy to deter-
mine this species.

118 Annals Entomological Society of America [Vol. XII,

Erax subpilosus Shaeffer.

Male. Length 22 millimeters. Head grayish-white pollinose,
beard and mystax white, occipito-orbital bristles white. Thorax
black, faintly grayish-brown pollinose, mid-dorsal stripe faint; dorsum
sparsely covered with short, black hairs, which are longer apically.
Scutellum with black hairs and bristles, some of which are white apically.
Pleurz faintly pollinose, covered not densely with moderate long white
hairs. Abdomen black, first to third segments with black hairs, longer
at sides, fourth segment with long white hairs intermixed with black
hairs, which are directed outwardly; fifth segment white pollinose and
with shorter white hairs, sixth and seventh segments densely white polli-
nose; apical margins of segments two to five grayish pollinose. Venter
grayish-white pollinose, covered with long, white hairs. Hypopygium
black, clothed with shorter white hairs. Wings pure hyaline. Legs
entirely black, clothed with shorter and longer white hairs. Furcation
of the third vein almost directly opposite the base of the second posterior
cell, stump rather short. i

Female. Total length 22 to 27 millimeters. Colored much as in
the male, abdomen black, narrowly gray pollinose on the sides and
posterior margin of each segment, hair of the abdomen largely white,
but some black, especially on the dorsum. Oviduct rather long,
longer than the last three abdominal segments.

Type: Male, in the Brooklyn Museum of Arts and Sciences,
is labeled Beaver Creek Hills, Beaver County, Utah. Other
specimens at hand from Reno, Nevada, New Mexico and Cal-
ifornia. The California specimens are at the Museum of
Comparative Zoology at Cambridge, Mass., and are from the
collections of Loew and Osten Sacken. Loew gave the manu-
script name lencotrichus to the species.

The predominance of pure white bristles of the legs is a
distinctive character and easily separates it from most related
forms. The bristles of the feet especially practically all are
white.

Williston’s arzdus is much like subpilosus 1n general appear-
ance but the latter has a much longer oviduct. The color of
the vestiture of the abdomen is more or less variable in sub-
pilosus.

1919} Hine: Genus Erax 119

The anomalus Group.

Three species belong to the group with three submarginal cells
normally. They are all rather light colored and inhabitants of more or
less arid regions. Coquillett has proposed the genus Efferta for the
reception of these species, but usually it is considered a synonym of
Erax. The furcation of the third vein occurs about midway between
the base of the second posterior cell and the anterior cross vein. Costa
not dilated.

1. Bristles of the palpi, scutellum and upper part of the occiput, black,

anomalus Bellardi
Bristles of the mystax, palpi, upper part of the occiput and usually of the
SOUL ta MNCL OMae wipe rr Tyee ret acetate alec rae in wks vk 3, Vie cy 4 sys arte ML CR OPER 2

Male abdomen with white hairs parted at the middle and directed outward.
candidus Coquillett
Male abdomen gray pollinose and without white hair parted at the middle
aniducimec ted! OUiWand emais.. Sets cst sie mi srs teie in sae eare kira cts aA = pernicis Coquillett

to

Erax anomalus Bell.

Male. Total length 18 to 26 millimeters. Mystax varying from
white to pale yellowish, beard white, palpi sometimes clothed with
white hair and sometimes with black, thorax with white and black hair
and black bristles, scutellum with white pile on its disc and a row of
black bristles on its margin. The color of hair and bristles appear to
be somewhat variable, however; wings hyaline. Femora dark, usually
black, anteriorly and reddish posteriorly, tibia reddish, darker near
apex, tarsi darker than the tibia. Abdomen largely silvery pollinose,
segments two to six with long silvery hair parted at the middle and
directed outward. Hypopygium shown from side in Figure 10. Wing
. shown in Figure 5.

Female. Abdomen above with a prominent black triangle on each
segment with its base on the anterior margin and its apex not quite
reaching the posterior border, otherwise gray pollinose. Some variation
in the extent of the black occurs however. Oviduct almost as long as
abdominal segments five, six and seven.

Range: Kansas to Texas and Mexico and westward.

Erax candidus Coquillett.

Total length 15 to 30 millimeters. Body in both sexes white
pollinose, mystax and beard silky white, palpi with white hair, femora,
apexes of tibiz, and tarsi black, basal three-fourths of each tibia some-
times pale yellow, sometimes darker. Legs everywhere with white
hair and an occasional black bristle. Male abdominal segments two
to four with white hair parted at middle and directed outward, segments
six and seven without such hair. Hypopygium shown from the side
in Figure 11.

‘Range: Kansas, New Mexico, Arizona and California.

120 Annals Entomological Society of America [Vol. XII,

Erax pernicis Coquillett.

Total length 15 to 20 millimeters. Known from candidus by being
less distinctly white pollinose, the male is devoid of long hair on the
abdomen. The species averages smaller and the hypopygium is
different. See Figure 12.

Specimens from Arizona and California.

The aestuans Group.

The species of this group have been divided into two parts for the
purpose of the present paper. So far only one species has been found
to be common to both the North American continent and to the West
Indies. This being the case the two faunas are separated and a key
given for each in order to facilitate determination.

The species falling in the group are recognized at once by the
peculiar posterior branch of the third vein which, instead of curving
forward at its apex, as in other groups, curves backward and reaches
the costa near or beyond the apex of the wing.. The branching of the
third vein occurs beyond the base of the second posterior cell. The
following key is offered as an aid in separating the species of the North
American Continent.

iY Wegscentirely.shiningublack../ Sayin. ok Wee eee ae ata alo ae eee Ree 2
AtHleast thes tibiae distinethymmedtac) a3. se acre Comin ee ee ee 4

Za Winesphyaline imalercosta, mormnmalle sane a) ereier en ames ...Slossone n. sp.
Wings distinctly pale brownish, more pronounced on apical half; male costa
plainby expanded e272 Se ocr) le co iets cece tte rae ate grea nA oe MO 3

3. Total length, 20 millimeters or over, hypopygium longer than abdominal
Sienamleyolcy ine Syb< Bhalel Gavel, oavoncaseauuoohsslsudsoccronse loewii Bellardi

Total length not over 15 millimeters, hypopygium tumid....mexicanus n. sp.
4. Hypopygium from side view, most prominent apically on upper half, small

SPECIES! Waa, eed seas he ees, artim cre ein ngs cee eee tice retain rea eee oe ae 5
Hypopygium most prominent at middle apically, medium to large species. . .6

Dea Miy stax yellow: joists ton po er TRO ee aE eee aurimystaceus n. sp.
Miystax black with) gray, airs autermixed:- oe. a acnseer cae belfragei n. sp.

6.) shemora,” red mapically:.. sim isec cc. Ie Ee ore ne affinis eae

Remora blacks 2 hema) tie alsearcah nue tera ae bb Ue ee ene Gferere OE TOE RRR Cree aoe
Costa more or less dilated in the male, usually large robust species. Niystax
black) intermixed swath uwhaterhainses aeee mae Gece ee een eine ee ere eae
Costa not at all dilated, medium sized slender species. Mystax with ee

bristles above and pale yellow hairs below. Three species difficult to
Separate. | Refer tomeures ofhypopyeiums:-e sacs eee meee eeees 10

8. Hypopygium unusually large, conspicuously pilose below (Cuban species,
one record from Florida), mystax with a few black bristles. .stylatus Fabr.
Hypopygium of normal size. Mystax composed mostly of black bristles... .9

9. Total length 25 millimeters or more, the largest species of the aestuans

s~I

BTOUP Ace cmv te east pore Cee CRITE eee femoratus Macquart

Total length 23 millimeters or less, tibiae dark brown, general body color
blacker’ than, qusuall 54). dec 40 et seee Ce eee aestuans Linn.

10. Hypopygium with a tuft of dense pile below at apex. ..tabescens Banks, n. sp.
lalyAorojonigeaiohag! \yraelareybliy) GUO A), Wb, aod oka auosdEnedloneunecnogeuecooanoo‘t 11

11. Apical part of hypopygium narrowed and curved downward, with a tooth-
like; prominenceraboOve™ ane setae ner are apicalis Wiedemann

Apical part of hypopygium nearly as wide as the preceding part,
kansensis n. sp.

1919] Hine: Genus Erax 121

Erax belfragei n. sp.

Male. Total length 10 millimeters. Mystax largely black though
with some gray intermixed; palpi with black pile, beard gray, thorax
gray pollinose with the usual black markings, hair and bristles mostly
black, although some gray intermixed on posterior part and on the
scutellum. Wings slightly fumose, a little more intense toward the
apex, costal border not dilated in the least, stump on anterior branch
of the third vein not quite as long as the connecting vein, perhaps the
length of this stump will be variable however. Femora entirely black,
narrow apexes of tibiz and entire tarsi black. Abdomen dark, largely
gray pollinose, dorsum of segments two to four black before the narrow
posterior gray borders, segments five, six and seven silvery; genitalia
elongate, sides nearly parallel, widened somewhat on posterior third
and notched at tip. Figure 21.

Female colored like the male, stump of anterior branch of the third
vein fully as long as abdominal segments four to seven inclusive.

?

Type: Male and allotype bearing the label ‘Texas,’
Belfrage. Inthe U.S. National Museum.

Most closely related to aurimystaceus but the color and male
genitalia are clearly distinctive.

Erax slosson@ n. sp.

Male. Total length 14 millimeters. Mystax prominent, mostly
white, but with a number of black bristles in the upper part, palpi
with black pile, beard white, dorsum of the mesothorax with long black
hair and bristles, scutellum with white hairs on its disc and a few black
bristles on its margin. Wings hyaline, very short, much shorter than
the connecting vein; legs entirely shining black, clothed with white
and black hair and black bristles. Abdominal segments two,’ three
and four black before the narrow, gray pollinose, hind-margins, dorsally.
Narrow posterior margin of four, all of five, six and seven silvery;
genitalia shining black, with black hair, widened from the side gradually
to middle, then narrowed gradually and with a rather large extension
apically partly cut off.by incisions above and below. See Figure 25.

Male type from Jacksonville, Florida, collected by Mrs.
Anna T. Slosson and forwarded by C. W. Johnson.

Very distinct from other species of its group from Eastern
United States on account of its entirely shining black legs.
The male genitalia distinguish it readily.

Erax affinis Bellardi.

Total length 17 millimeters. Mystax black and white mixed,
palpi with black and white hairs and bristles, beard white, thorax
yellowish brown pollinose with the usual black markings; scutellum
white pilose with black or white marginal bristles; legs variable to some
extent, femora usually black on basal three-fourths and red at apex,

122 Annals Entomological Society of America [Vol. XII,

but nearly all red in some specimens, although all I have seen have at
least a trace of black on the front side of each; tibia red, darker apically,
tarsi dark red with black bristles. Wings yellowish, slightly more
intense toward apex, male costa slightly dilated. Abdomen dark,
gray pollinose on venter and sides. Male with segments six and seven
silvery, genitalia shining black with an extension apically attached below
and directed upward. See Figure 28.

Female colored like the male, except femora are more extensively
red than in the other sex in some of the specimens studied.

The females taken in the same locality as the males, as well
as other females before me, agree with Bellardi’s description of
marginatus. He described affinis from a male and marginatus
from a female. It appears that they are sexes of the same
species.

Specimens from Mexico and from Bonda, Columbia, S. A.

Erax kansensis n. sp.

Total length 17 millimeters. Mystax largely pale straw yellow,
but with some black bristles intermixed, palpi clothed with black hair,
beard nearly white, though with a tinge of straw color. Thorax above
with black and white pile and some black bristles; wings hyaline, a
tinge of brownish in the apical part of the marginal cell, costa not
dilated, anterior branch of the third vein with only an indication of a
stump; femora, apexes of tibize and entire tarsi black, basal three-fourths
or more of tibiz bright yellow; abdomen dark in ground color, largely
thinly gray pollinose, seventh segment just slightly silvery. Male
genitalia narrow; not long, narrowed suddenly and bearing at the
tip an angular extension. See Figure 19.

Female colored like the male, genitalia rather wide and somewhat
longer than abdominal segments five, six and seven.

Type: Male and allotype, from Clark County, Kansas,
elevation 1962 feet. F. H. Snow. In the author’s collection.
Easily identified by the male genitalia. Most closely related
to apicalis.

Erax aurimystaceus n. sp.

Length of male 13 millimeters. Mystax rather large, golden yellow,
beard paler yellow. Thoracic dorsum with numerous rather long black
hairs and some pale yellowish hairs intermixed on the posterior part
where they are distinctly longer than on anterior part. Scutellum with
numerous pale hairs on the disc and two black bristles apically. Wings
uniformly very pale yellowish hyaline, femora entirely deep black,
tibiae bright yellow with extreme apexes black, tarsi darker, but mostly
reddish, with numerous black bristles. Legs clothed with fine pale
hairs and black bristles. Abdomen yellowish gray pollinose, segments
two to five each with a large quadrate black spot above, segments six

1919} Hine: Genus Erax 1s

and seven silvery. Hypopygium shining black with abundance of
fine black hair; from side view notched at tip and quite different from
other species of its group. Refer to Figure 22.

Holotype: Male from Clark County, Kansas, F. H. Snow.
Several males and females from Clark, Ellis, Grove and Ness
Counties, Kansas, mostly taken in July.

The female differs from the male only in sexual characters.
The oviduct is distinctly shorter and stouter than in aestuans
and apicalis, being only about as long as the last three abdominal
segments.

The species is entirely distinct and easily recognized by its
small size, yellow mystax and male and female genitalia. The
bristles at the apex of the scuteHum are subject to slight varia-

tion in number and in some specimens are golden yellow instead
of black.

Erax mexicanus n. sp.

Male. Total length 12 millimeters. Mystax black above, pale
vellow below, palpi with black bristles, beard pale yellowish, wings
nearly uniform brownish, costal margin very slightly dilated, stump
of anterior branch of the third vein much shorter than the connecting
vein; legs black in all their parts. Abdomen black with yellow hairs,
segments five, six and seven silvery, venter of last abdominal produced
backward the length of more than two abdominal segments and with
several black hairs at the tip. Genitalia black, shining, tumid, suddenly
narrowed before the apex, produced into a distinct tubercle. See
Figure 27.

Female colored like the male; stump of the anterior branch of the
third vein a little longer than in the, male, but not much longer than
the basal section. Genitalia somewhat longer than segments five, six
and seven. |

Male type and allotype from Tapachula, Chiapas, collected
by D. H. Crawford. Three other males from the same locality.
One male is somewhat larger than the type and has only abdom-
inal segments six and seven silvery.

The small size, black legs and peculiar male genitalia serve
to distinguish the species.

Erax femoratus Macquart.

Total length, male and female, 26 to 30 millimeters. Mystax
largely black, but with white hairs intermixed, beard pale, palpi black
hairy, occipito-orbital bristles black; mesothoracic dorsum before with
very short black hair, several strong black bristles above the roots of
the wings and before the scutellum; scutellum with fine white hairs

124 Annals Entomological Society of America |Vol. XII,

on the disc and several strong black bristles on its margin, femora black,
tibiae and tarsi dark red, apices of tibize and tarsi darker in some speci-
mens than others; wings hyaline, male costa slightly dilated. Male
abdomen, first four segments gray on the sides and narrow hind margin;
segments five, six and seven silvery, hypopygium black and largely
black hairy. Female abdomen with each segment gray on sides and
narrow posterior border, oviduct nearly as long as the last four abdominal
segments.

From Slidell, Louisiana, and from Florida.

Much like @stuans and there is some doubt in regard to its
distinctness from that species, but the large size and robust
form give it quite a different appearance. The specimens I
collected at Slidell, Louisiana, were taken from the trunks of
trees:

Erax aestuans Linne.

Variable in size. Usually ranging from 18 to 22 millimeters, but
specimens as small as 14 millimeters and others as large as 28 are taken
occasionally, especially females if measurement is taken to the tip of the
oviduct.

The body usually is more hairy than in other species of its group.
Mystax composed of black bristles and gray hairs, dorsum of the
thorax with numerous black hairs and bristles on the posterior part,
scutellum with an apical row from six to ten black bristles, wings
slightly tinged with brownish, legs black with bases of tibia rather dark
brown, abdomen black with a narrow gray posterior border to each
segment. The male has from two to four apical abdominal segments
silvery. Figure 17 shows the form of hypopygium from lateral view.
The oviduct is slender, nearly as long as the last four abdominal segments
and usually turned to one side or the other in dry specimens. Figure 4.

Habitat: Widely distributed over north-eastern North
America, ranging to Florida, New Mexico and Wyoming. A
very abundant species over much of its range.

The rather pronounced hairiness of the body, especially the
posterior part of the dorsum of the thorax where the hairs are
unusually long and plentiful, the row of bristles on the scu-
tellum, the brown and not yellow tibiz and form of hypopygium
rather easily distinguishes this species from others of its group.

Variation in size and the extent of the silvery color of the
male abdomen has given an excuse for a number of synonyms
of the species. Thus macrolabis is used for the specimens
having four segments silvery. I have observed this character
often and find it appearing so irregularly that it can hardly be

1919] Hine: Genus Erax 12!

considered specific. Wiedemann’s niger appears to be a rather
large female of aestuans. Some Georgia specimens before me
measure 26 millimeters and agree with the description of niger.

Erax apicalis Wiedemann.

Size variable. Males and females before me are from 14 to 22
millimeters in total length; average specimens are about 17 millimeters
long. Mystax white with only a few black bristles above, beard silky
white. Dorsum of the thorax with very short black pile anteriorly anda
sparse arrangement of longer white hairs and black bristles posteriorly,
scutellum with rather long white hair on its disc and two black bristles
on its apical margin. Wings hyaline, slightly infurcated near the
apex. Femora, tarsi and apexes of tibia black, remainder of tibiz
bright yellow. Abdomen gray pollinose with a large quadrate black
spot on the dorsum of each segment. Male with the last two abdominal
segments silvery but often the anterior one of the two is more or less
encroached upon by a black spot. Female with oviduct shining black
and near the length of the last four abdominal segments. Lateral
view of the hypopygium shown in Figure 18.

The species is southern in range. Specimens from Decatur
County, Georgia, collected by Bradley, and from southern
Pines, North Carolina, by Manee.

The trim appearance of the species is notable. The hairs of
all parts of the body are shorter and sparser than in @stuans;
normally there are only two black bristles at the apex of the
scutellum, although some variation may be encountered, and
the tibiz are bright yellow instead of brown.

Wiedemann ascribes the species to North America, but it is
presumed his specimen was from the southern part of the
country for other species described near it are from Savannah,
Georgia. Prof. Comstock is the only recent writer to mention
the observation of sepcimens when he records their preying
upon larve of Aletia argillocea in Alabama.

Macquart described vicinus from Galveston, Texas, and the
description suggests that it is the same as apicalis.

Erax loewii Bellardi.

Total length 19 millimeters. Mystax black and pale intermixed;
beard gray with just.a tinge of yellowish, palpi with black hair. Thorax
largely gray pollinose with the usual black markings, wings nearly
hyaline basally, apical half yellowish, male costa a little dilated, stump
of the anterior branch of the third vein short. Legs entirely black,
posterior of tibia and tarsal segments golden pilose. Male abdomen

126 Annals Entomological Society of America [Vol. XII,

dark, segment six on posterior corners and hind margin and segment
seven silvery above; genitalia black, regular, with a small rounded
prominence apically. Figure 20.

Female colored like the male, stump of the anterior branch of the
third vein rather long, longer than the basal section, genitalia somewhat
longer than abdominal segments five, six and seven.

Specimens from Mexico, Tapachula, Chiapas, Cordoba,
Morales, Guatemala, and from Onaca, Columbia.

There are only a few species of Evax with entirely black legs,
so this one is rather easily identified. Williston’s clear descrip-
tion of dolichogaster lends a feeling of satisfaction to the student
who has it for determination and Bellardi’s figure of loewit is
very good. That the two names refer to the same species is quite
evident. Neither Williston or Osten Sacken recognized lewit
in Biologia Centrali Americana.

Erax tabescens Banks n. sp.

Male. Total length 18 millimeters. Similar to Erax aestuans
in having the hind part of third vein ending beyond wing tip, and the
fork beyond the base of the second posterior cell. The mystax is black,
with some white hairs, palpi black haired, head and thorax also with
black hairs. The venter is gray with dark median streaks, the sides
and hind margins of the segments above pale, the lateral margin of the
sixth segment broadly silvery, the seventh wholly silvery. The abdomen
is more slender than in Erax aestuans. The hypopygium is rather
smaller, but noticeably more slender than in Erax aestauns and is
clothed with black hairs; the legs are black, the tibia, except tip, pale
yellowish, the hair of femora and tibiz is almost wholly white (mostly
black in Erax aestuans). See Figures 16 and 26.

From Florida (Loew collection and his manuscript name.)

It appears that only one species of the genus Evax outside
of the estuans group has been reported from the West Indies.
Both Bigot and Williston report rufibarbis from Cuba, but it
seems that this species is only occasionally taken. The species
of these islands are much alike in general appearance and some
of them are confused with others in literature, although not
much has been written aside from the original descriptions.

What is known now as estuans Linne has a large number of
synonyms as may be seen by reference to the list of species on a
previous page, and besides it has been customary to identify
related species with this one because of the close relationship of
members in the group. Study of much material from various
parts of the country seems to throw some doubt on the cor-

‘

1919] Hine: Genus Erax 124

rectness of the extensive distribution of @stuans which is given
as occurring from New England to Guiana, including records
for the West Indies.

Seven species are recognized in the material at hand from the
West Indies, mainly based on a study of male genitalia, and
none of them are regarded as the common estuans Linne of the
Eastern United States. The following key will aid in
separating them.

1. Mystax pale; scutellum with white pile and bristles......................4: 2

Mystax in large part black, scutellum with black bristles.................. 4

RM VANES UCUXS UT CUERVO OWUHIAPY nicl = fc cise ee alice Slacha si ceibetl cies ws cle ae nigritarsis n. sp.

ITB S eae vy ili Ce PN eee abe ich ts eh Sette ea PENS coerce tae. pig Eee aes Sis iclcea} «5, Dds hae se Seen ae 3

DLV POpy Camm OngceamdemanrrOwi nw ao ees 2 sur ole «seis a las portoricensis n. sp.
lslyiooroderbian, Slovory fhalall qabiaaklola suet ooo mean Le opoaue een oe rufitibia Macq.

4. Male costa distinctly thickened and dilated beyond the tip of the auxiliary

WETS Nba Ohi EuAKOUlhiranetaKohiSial erste 6 Goloma od bln Seddon caeteen a Uenulor ioe san 5

Male costa not thickened and dilated, wings nearly hyaline................ 6

5. Hypopygium much enlarged, rather large robust species....... stylatus Fabr.
Hypopygium small, species 16 millimeters in total length nigrimystaceus Macq.

6. Hypopygium short and tumid, shining black all over....... rufitibia Save
LS Ry iaayonrseatiniant GIMOMEANT DNs fo bes cores cee atta Looe ae ane oO aoe © mee eee

i. Large species, hypopygium much elongate............... fulvibarbis Macey,

Medium sized species, hypopygium only shghtly elongate....haloesus W. K.

Erax stylatus Fabricius.

Total length, male 20 to 26 millimeters, female, 18 to 28 millimeters.
Mystax composed of numerous black bristles intermixed with fine
white hair, beard white, palpi with white hairs, but mixed in is an
occasional black one, in some specimens more than others. Femora
entirely black, tibiz yellow, each very narrowly black at apex, tibize
somewhat darker than the tibiz and with numerous black bristles;
wings distinctly brown.

Male genitalia large, fully as long as abdominal segments five, six
and seven. See Figure 32. Costal border of the wing distinctly
dilated beyond the tip of the auxiliary vein. Usually no stump on
anterior branch of the third vein, although in some cases there is a very
short one.

Female genitalia shining black, about as long as segments four to
seven, wings almost hyaline, costal border of wing not dilated, anterior
branch of the third vein with a distinct stump.

Specimens from several places in Cuba, from Porto Rico,
and one male from extreme southern Florida, indicating a wide
range. The Florida specimen is peculiar in having the whole
abdomen from the apex of the second segment to the genitalia
silvery, but I can find no other differences.

Although stylatus has not been recognized by most authors,
Wiedmann’s figure should help in its determination. With more

128 Annals Entomological Society of America [Vol. XII,

than thirty very well preserved specimens for study the lit-
erature has been consulted fully and the above synonymy
worked out. I have no doubt the males and females before me
are the sexes of the same species. They seem to combine with
characters mentioned by Fabricius, Macquart and Walker who,
as a usual thing, seem to have had each a single specimen in
hand. Thus the descriptions of stylatus and invarius were taken
from females, while those of caudex and haitensis were of males.
The name caudex, as Walker indicates, was used because of the
‘‘very large’’ genitalia. Macquart’s figure of the wing of
haitensis shows the distinctly dilated costal border which is
shown in the males before me. The large size Walker gives for
invarious, together with other characters, is good proof of its
synonymy with stylatus. These statements are emphasized
when it is pointed out that among no less than eight West
Indian species this is the only one that combines the dilated
costa and large male genitalia, while only one other has a
slightly dilated costa and one a somewhat enlarged hypopygium.
Most likely Van der Wulp was in error when he reported
stylatus from Wisconsin. The descriptions given by all the
author’s mentioned in this connection are more or less deficient
and one could easily apply them to several species of the genus.

Erax rufitibia Macquart.

Total length 18 millimeters. Rather dark species, thorax and
abdomen largely gray pollinose. Mystax black with a few white hairs
intermixed, beard white, palpi black haired, hairs and bristles of the
thorax all black; wing slightly fumose, no costal dilation in the male,
anterior branch of the third vein with a stump which is about equal in
length to the basal section. Femora black, tibia and tarsi almost
wholly red. Sixth and seventh abdominal segments silvery in the male,
genitalia of male short, somewhat tumid. Figure 30.

Females agrees with the male except in sexual characteristics.

From Bahamas and Cuba.

Erax portoricensis n. sp.

Total length 20 millimeters. Mvystax, hairs of the palpi and beard
white, an occasional black hair in the mystax, dorsum of the thorax
largely rusty brown pollinose, wings pale brown, nearly hyaline, costal
margin not dilated, stump of the anterior branch of the third vein
equal in length to the basal section, femora and tips of tibiz black,
basal three-fourths of tibize yellowish brown, tarsi dark reddish brown,
nearly black, with abundance of black bristles. Abdomen darker

1919] Hine: Genus Erax 129

dorsally, segments six and seven silvery. Male genitalia elongate,
rather narrow, apical part narrowed, then rather suddenly widened,
angular at apex. See Figure 24.

Female colored like the male, but tibia more plainly reddish brown,
genitalia hardly as long as segments four to seven inclusive.

Type: Male from Ensenada, Porto Rico, June 14 to 19,
1915, and allotype from the same place at the same date,
property of American Museum of Natural History.

The male is easily recognized by its peculiar genitalia.

Erax nigritarsis n. sp.

Total length 18 millimeters. Mystax, beard and hairs of the palpi
all light colored. Thorax in large part gray pollinose, mid-dorsal space
and some spots on either side of mesothorax black on account of the
very sparse pollen with which they are covered. Wing hyaline, only a
tinge of yellowish, costal border only very slightly dilated, so slight that
it is hardly evident, stump of the anterior branch of the third vein
very short, much shorter than the basal section; femora black, tibiae
yellow to near the apex, tips of tibiae and tarsi intensely black, ventral
side of metatarsal segments and some other segments golden pilose.
Abdomen gray pollinose, darkest dorsally, segments six and seven
silvery, genitalia short, somewhat tumid from side view, narrowed
from the middle towards apex where it is truncate. See Figure 29.

Female colored like the male in all respects, genitalia shining black,
approximately as long as abdominal segments four to seven inclusive.

Type: Male and allotype from Holguin, Cuba, December
19, 1904, collected by H. S. Parish. In the author’s collection,
a male and four females from various Cuban localities in
American Museum of Natural History.

The pale mystax, tibial hairs, beard and black tarsi, with the
structural characters described are distinctive for the species.

Erax fulvibarbis Macquart.

Total length 25 millimeters. A dark appearing species. Much
of the thorax and parts of the head and abdomen rusty brown pollinose;
mystax composed of numerous black bristles with a tuft of rather coarse
white hairs below, beard white, palpi black haired; femora all black,
each femur, especially the posterior, has a longitudinal row of six or
seven rather strong black bristles on the outer side, beginning well
towards the base and extending to near the apex. Most species have
not more than four of these bristles. Tibiz light brown with the apex
of each distinctly black, tibia black; wings tinged with brownish,
front costal border not dilated in the least, anterior branch of the third
vein with a very short stump. Abdomen dark, segments two and three
each, with a silvery triangular marking on each side posteriorly, segments
six and seven silvery dorsally, lateral view of male genitalia shown in
Figure 23.

130 Annals Entomological Society of America [Vol. XII,

A male specimen from San Domingo. This species agrees
exceedingly well with Macquart’s description of fulvibarbis
and comes from the type locality. The row of six or more
bristles on outside of hind femur easily suggests the species.

Erax haloesus Walker.

Total length 18 millimeters. Mystax white with numerous heavy
black bristles, beard white, scutellum with white hairs on its disc and
about four rather slender black bristles on its margin; tibiae bright
reddish yellow, except distally, tips of tibize and tarsi black, although
the latter may be partly dark reddish brown, costa not thickened, wings
dilute yellowish hyaline. See Figure 36. Hypopygium elongate,
distinctly red at tip.

A male specimen from Jamaica furnished by C. W. Johnson
by whom it was identified. The hypopygium enables this species
to be determined readily.

Erax nigrimystaceus Macquart.

Total length 17 millimeters. Mystax largely black with pale yellow-
ish hairs intermixed, palpi with black hairs, beard straw yellow, thorax
dark, gray pollinose, bristles and hairs black, many black upright
hairs on the scutellum, wings distinctly yellowish, a little more intensely
so toward apex, costal border plainly, though not strikingly, dilated,
stump of the anterior branch of the third vein very short, not half
the length of the connecting vein; entire femora, apexes of tibiz and
tarsi clear black, basal three-fourths or more of tibiae brown. Abdomen
dark, partially gray pollinose, segments six and seven silvery.
Hypopygium black, not tumid, gradually widened from the base to
beyond middle then slightly narrowed and truncate at apex with two
minute emarginations, one dorsal and the other ventral. See Figure 31.

Male from Dominica, West Indies, November, 1903.
Furnished by Chas. W. Johnson.

The carinatus Group.

In Western United States and extending into Mexico and Central
America there are several species of the genus Erax that are distinct
from the others on account of the following characters:

The dorsum of the mesothorax is more convex than usual, with a
distinct crest of erect hairs on the middle of the dorsum, beginning just
behind the head and extending back to beyond half the distance to the
scutellum. Branching of the third vein beyond the base of the second
posterior cell; stump rudimentary or altogether absent in the male,
present and somewhat variable in length in the female; costa in the
male always thickened and expanded but much more evident in some

1919] Hine: Genus Erax Pol

species than others. Ten species of this group are at hand and at least
one other is characterized sufficiently so that it should be determined
easily when specimens are procured.

1. Vestiture of the scutellum, aside from the black marginal bristles, black or

ie) lone eerr te ta ae ee! 2), oo. Pa Te cc PNT 2
Vestiture of the scutellum, aside from the black marginal bristles. white... .4

Are VeStibunerOtmpnesscutelllum lol acleers wpe ccs ces 2 1nc.+ osteo ee jubatus Williston
Vestiture of the scutellum composed of black bristles on the margin and yellow

In ahe (Gi LANG bl Cts Ae bk al a Re Rr ls! pce ae

3. Male with the last four abdominal segments silvery. .... concinnatus Williston
Male with the last two abdominal segments silvery....... carinatus Bellardi

a.) Khemord ced apically and sometimes posteriorly..............-...sssn.bcoe.D
teri oy a a DS SIG eh is es OA Ee i ne cl oe I Be Sol eS) Fa > 6

Second to fifth abdominal segments of the male with long white hair parted
at the middle and directed outward, abdomen wholly gray pollinose,

unicolor Bellardi

Abdomen without hair parted at the middle and directed outward. basal

Scoments ol uneralodomentiblacks, .) sss mate ase foe parvulus Bellardi

6. Robust species, mane-like crest of the anterior part of the mesothorax,

; especially thick and long, male costa very prominently thickened,

subcupreus Schaeffer

Slender species, mane-like crest normal, male costa only slightly thickened. . .7

Male abdomen with only the last two or three segments entirely silvery,

without prominent white hair parted at the middle and directed outward;

“I

other segments with black markings................. latrunculus Williston
Male abdomen with at least four or five segments silvery, and with long white
hainparted at the muddle and* directed outward. . 2)... 9.2 92..00) 122. 8

8. Hypopygium divided apically with both parts of nearly equal length,
cressoni n. sp.
Hypopygium emarginate apically with the upper part much shorter than the
PONV tae Pema te Sep Bien Oe ON ep Ae ieopat ners. aks Salah Pcs t costalis Williston

Erax carinatus Bellardi.

Total length 18 millimeters. Body yellowish gray pollinose, hairs
and bristles black and pale yellow. Mystax yellow with a few black
bristles, beard paler yellow than the mystax, ocellar bristles black,
hair on scutellum and in front of it straw yellow, a few black bristles
on the scutellar margin; femora black, tibiz and tarsi largely red,
bristles black. Abdomen with short, sparse, yellowish hair, segments
six and seven silvery. See Figure 58.

Male from Guerrero, Mexico.

Erax concinnatus Williston.

Total length 16 millimeters. Mystax and beard pale yellow, the
former mixed with black bristles, palpi black haired. Disc of scutellum
and rear of mesothorax yellow haired, margin of scutellum with a few
black bristles. Femora entirely black, tibiz and tarsi yellow, apex of
tibia and each tarsus darker. Wings yellowish hyaline. Abdominal
segment one and two and base of three black dorsally, sides of first
three segments, apex of three and all of four to seven inclusive, silvery,
hypopygium black, of medium size. See Figure 57.

Male from Guerrero, Mexico.

1382 Annals Entomological Society of America |Vol. XII,

Erax parvulus Bellardi.

Total length 13 to 15 millimeters. Mystax and beard white, the
former with black bristles intermixed. Disc of scutellum and rear of
mesothorax white haired. Margin of scutellum with a few black bristles.
Wings hyaline. Femora, except at extreme apex, black, apexes of
femora, tibia and tarsi red. Male, first three abdominal segments
dorsally black, narrow posterior margins of two and three and all of
four, five, six and seven silvery. Hypopygium black. See Figure 55.
Female abdomen black, narrow sides and hind margin of each segment
gray pollinose, oviduct slightly longer than the last three abdominal
segments.

Male and female from Guerrero, Mexico.

Erax unicolor Bellardi.

Total length 16 to 19 millimeters. Mystax pale yellow with a few
black bristles, beard pale yellow, palpi black haired, mesothorax with
black hairs and bristles, scutellum with white hairs and a few black
bristles on its margin, femora black to near the tips, apexes of femora
and tibie and tarsi red. Abdominal segment ane and base of two gray,
apex of two, and whole of segments three to seven, inclusive, silvery;
segments two, three, four and five with long white hair parted at the
middle and directed outward. Hypopygium black, emarginate at the
tip. See Figure 60.

Female colored much as in the male, abdominal segments two to
seven each with a dark triangle dorsally, oviduct about as long as the
last three abdominal segments.

A male and female from Amatitlan, Guatemala, and five
males and females from Tehauntepec, Mexico, (Sumichrast).

Erax latrunculus Williston.

Total length 15 to 18 millimeters. Mystax white with a very few
black bristles, beard white, rear of the mesothorax and disc of the
scutellum with white hair, margin of the scutellum with several black
bristles, femora all black, tibiz and tarsi red. . Male, abdominal seg-
ments one to five each with a black spot above, large on segments two,
three and four, small on five. Sides and hind margin of two to five
inclusive and all of six and seven silvery. In one specimen six is all
silvery, lacking the dorsal black spot. Hypopygium shining black,
mostly with pale hairs. See Figure 56.

Female, abdominal segments one to seven inclusive each with a
black spot above, sides and hind margin of each segment densely
gray pollinose. Oviduct somewhat shorter than the last three abdominal
segments combined.

Male and female from Arizona. This description is taken
from specimens from Cornell University. Professor Bradley

1919] Hine: Genus Erax 133

considers them as belonging to the material which Williston
had when he wrote the original description although they are
not marked as types.

Erax jubatus Williston.

Total length 18 to 23 millimeters. A large, black, robust species,
plainly the largest of the group. Mystax composed of numerous long
bristly hairs which are mixed white and black in about equal numbers,
beard silky white, palpi black haired, hairs and bristles of the thorax
largely black, numerous long black bristly hairs on the scutellum and
in front of it, although in some specimens white ones are intermixed.
Femora black, tibiz and tarsi dark red. Abdomen largely black,
narrow posterior margin of the segments gray. Male, abdominal
segments six and seven silvery, hypopygium noticeably large. See
Figures 3 and 54. Costal margin of the wing prominently expanded
and there may be a very short stump on the anterior branch of the third
vein.

Female, usually a distinct stump on the anterior branch of the third
vein; oviduct somewhat longer than the last three abdominal segments.

Forty specimens of both sexes from Northern Mexico, Ari-
zona, New Mexico and Colorado.

A study of the variations exhibited in the specimens at hand
suggest strongly that prolificus Osten Sacken, is a synonym of
jubatus. The type locality for jubatus is New Mexico and that
of prolificus is Northern Sonora, Mexico. Specimens vary
much in some particulars, many have white hairs on the scu-
tellum and before it, the bristles of the body are mostly black,
but yellow and brownish ones occur, the fifth abdominal seg-
ment of the male is almost wholly black, with a rounded black
spot, or entirely silvery.

The large size, generally dark color, and enlarged hypo-
pygium suggest the species. However, there is much difference
between the largest and smallest specimens.

Erax subcupreus Schaeffer.

Total length, male and female, 15 to 20 millimeters. Larger and
more robust than costalis. Mystax rather long and dense, composed of
black and white bristles and hairs in nearly equal proportion. Mane-like
crest of hairs on the middle of the dorsum of the thorax conspicuous,
black and reaching well beyond the transverse suture where it merges
into a widened area of black and white hairs and bristles which reaches
to the scutellum. Scutellum densely white haired and with several
black marginal bristles, wings hyaline, femora black, tibiz and tarsi
very dark reddish. Male abdomen quite densely haired, first five or

134 Annals Entomological Society of America [Vol. XII,

six segments with long white hair parted at the middle and directed
outward. Hypopygium black with black hair. See Figure 52

Female abdominal segments all black above, sides and narrow hind
margin of each gray pollinose, oviduct about as long as the last four
abdominal segments. Fresh specimens have the body more or less
metallic, which accounts for Schaeffer’s name.

Specimens from Montana, Colorado, New Mexico, Arizona
and Nevada.

The, robust form and white-haired scutellum are character-
istic for the species.

Erax cressoni n. sp.

Male. Total length 14 millimeters. Mystax and beard white,
palpi black haired, mane-like row of hairs on the middle of the meso-
thoracic dorsum black and composed of numerous short hairs and an
occasional long one, space in front of the scutellum with scattered black
bristles and short white hairs. Scutellum with numerous short white
hairs on the disc and four long dark bristles on the margin; femora
black, tibiz and tarsi red; wings hyaline, branching of the third vein
far beyond the base of the second posterior cell, anterior branch with
scarcely a suggestion of a stump. Abdomen, first segment white
haired, second to fifth segments with white hair parted at the middle
and directed outward, hair shorter on the fifth segment than on the
others, first segment, anterior part of second and a small triangle
anteriorly on dorsum of third showing dark from above, otherwise '
abdominal segments silvery white. Hypopygium dark, pale hairy,
superior part notched at apex, part above notch narrow, but of same
length as part below. See Figure 50.

Female. Length 14 millimeters, like the male in most respects;
anterior branch of the third vein with a stump about equal in length
to the basal section, each abdominal segment with a triangular spot
which is not as densely pollinose as the other parts, oviduct shining
black, about as long as the last three abdominal segments.

Type: Male and allotype from Alamogordo, New Mexico,
in the collection of the Academy of Natural Sciences of Phila-
delphia; other specimens from E] Paso, Texas.

Named for E. T. Cresson, Jr., who sent me the specimens.

Erax costalis Williston.

Total length, male and female, 14 to 16 millimeters. Mystax
white with some black bristles intermixed, scutellum with abundance
of white hair and about four black marginal bristles, wings hyaline,
femora black, tibiz and tarsi mostly dark red. Male, abdomen with
segments two, three and four with long white hair parted at the middle
and directed outward, first three segments largely black dorsally,
apex of three and all of segments four to seven inclusive silvery, hypo-

1919] Hine: Genus Erax 135

pygium black with white hair. See Figure 53. Female, abdomen
with each segment marked with black dorsally, sides and posterior
margin of each segment white pollinose, oviduct somewhat longer than
the last three abdominal segments.

Specimens from Montana, Wyoming and Colorado.

There is some doubt as to the proper identification of
costalis, but the information available indicates that this
species is the one for which Williston used the name in his key,
Trans. Am. Ent. Soc., XII, 64. No description was ever wtriten,
and Aldrich did not give it in his catalogue.

The tuberculatus Group.

The males of this group are at once known by the presence of a
tooth-like prominence on the venter of each of abdominal segments
four, five and six, and costa entirely normal. Branching of the third vein
distinctly beyond the base of the second posterior cell. The species
are slender and of rather small size.

1. Hypopygium and ventral tooth-like process of the abdomen red,

spiniventris n. sp.
Hypopygium and ventral tooth-like processes black........................ 2
Ventral tooth-like processes of the abdomen all acute at apex,

tuberculatus Coquillett
First two ventral tooth-like processes knobbed at the apex... .productus n. sp.

NO

Erax tuberculatus Coquillett.

Total length 15 to 17 millimeters. Body gray pollinose and white
hairy, ocellar bristles and some bristles in lower part of mystax black,
palpi black hairy; femora, apices of tibia and tarsi black, tibia otherwise
red. Male with venter of abdominal segments four, five and six each
with an acute, black, tooth-like process produced downward. The
presence of these ventral spines will put the male into its group at once,
wings hyaline, branching of the third vein distinctly beyond the base
of the second posterior cell, stump usually shorter than the basal section.

There is no very good way to characterize the females, but the small
size and the forking of the third vein distinctly beyond the base of the
second posterior cell are suggestive of the group. The oviduct is only
slightly longer than abdominal segments six and seven in tuberculatus,
but longer in other species of its group.

The type is in the United States National Museum. Other

specimens from Davis Mountains, Texas (F. M. Gaige), and
from West Fork, Texas (J. E. Scherer), taken in June.

Erax spiniventris n. sp.

Male. Total length 12 millimeters. Vestiture of the body largely
white, ocellar bristles black, bristles of the legs largely white, but now
and then a black one is intermixed; tibia, except the extreme apex of

.

136 Annals Entomological Society of America |Vol. XII,

each red, otherwise legs black, wings hyaline, branching of the third
vein beyond the base of the second posterior cell, stump very short.
Abdomen largely gray pollinose, venter of segment four and five each
with a distinct red prominence, narrowed basally and enlarged apically,
with the main part of the enlargement directed posteriorly; sixth segment
with a narrow pronounced red tooth-hke appendage pointing almost
directly downward and acute at apex. Hypopygium red in color,
from side view, widest at basal third and gradually narrowed toward
apex, where it is almost truncate.

Type: Male from Santa Rita Mountains, Arizona, 5,000 to
8,000 feet, July, (F. H. Snow). In the author’s collection.

The form and color of the hypopygium and ventral abdom-
inal prominences easily characterize the male of the species.

Erax productus n. sp.

Male. Total length 16 millimeters. Body largely white pilose
and gray pollinose, palpi black hairy, basal two-thirds of tibiz red,
legs otherwise black, and with black bristles; wings pale yellowish,
hyaline, branching of the third vein beyond the base of the second
posterior cell, stump very short. Abdominal segments four and five
ventrally, each with a short blunt prominence, segment six with a large
acute prominence directed downward. Hypopygium, from side view, of
nearly uniform width, rather long, apically each outer corner prominently
produced.

Female. Of similar size and color as the male; oviduct fully as
long as the last three abdominal segments.

Type: Male and allotype from Flinn Springs, Lakeside,
Cal., August 9, 1917. E. G. Holt, collector. Collection of the
author. Other specimens from Spring Valley and Poway,
August 10 and 14, by the same collector.

This species is of nearly the same size as tuberculatus, but
the ventral abdominal appendages and hypopygium are quite
different and easily characterize the male.

The rufibarbis Group.

The females of this group differ from all other species by having
the last segment of the oviduct divided at the tip. This character is
easily seen from dorsal view. The mesothorax and scutellum is unus-
ually bristly, the third vein branches beyond the middle of the distance
between the base of the second posterior cell and the small cross vein.
The male has abdominal segments six and seven silvery. The oviduct
is rather short and rigid and the hypopygium is rather small. Mystax
without or with very few black bristles.

1. Mystax yellow, body and wings dark colored............ rufibarbis Macquart
Mystax nearly white, body gray pollinose, wings hyaline... .bicaudatus n. sp.

1919] Hine: Genus Erax 137

Erax rufibarbis Macquart.

Male. Total length 16 to 30 millimeters. Mystax and beard
distinct reddish yellow, palpi black and clothed with black hair, color
of the body dark brown pollinose, thorax with black hair and bristles,
scutellum with black hairs on the disc and numerous black bristles,
irregularly arranged, on the margin. Legs mostly black with numerous
dark and reddish hairs and black bristles, tibia dark reddish on basal
half or more, wings brown, many specimens have the margins of the
veins more intensely colored, costa not dilated, branching of the third vein
nearly opposite the base of the second posterior cell. Abdominal
segments six and seven silvery, hypopygium shining black.

Female colored like the male and just as variable in size; oviduct
shining black, about as long as abdominal segments six and seven,
abruptly narrowed before the apex, furcate from dorsal view.

. The species has a wide range in Eastern North America,

including the West Indies. Normally colored specimens are at
hand from as far west as the Dakotas and central Texas.
Farther west specimens are lighter in color and indications are
that rufibarbis more or less eeacually approaches bicaudatus
in that region.

A study of venation in Erax reveals some variations which
are useful in grouping species. There is a small group of
species that has three submarginal cells normally and there are
some species outside this group where now and then a specimen
has three submarginal cells in only one or in both wings. Thus
anomalus, candidus and pernicis may be characterized as a
group by the presence of the extra cell while rufibarbis and
bicaudatus have an occasional specimen only with this char-
acter. An estimate of material at hand indicates that about
two per cent of these species are abnormal in this respect. The
type of ravus in the U. S. National Museum is an abnormal
rufibarbis and from much material studied, results indicate that
scarcely any doubt exists that completus is the same, thus adding
two more to the long list of synonyms of our common Erax.

Macquart’s figure of the wing of his completus shows the
forking on the third vein opposite the cross-vein at the base of
the second posterior cell, and not midway between the latter
cross-vein, and the small cross-vein as is the case in all the
species of Hrax having three submarginal cells normally. This
fact and his statement regarding the male ‘‘Abdomine segmentis
duobus apicalibus niveis,’’ which is distinctly so in regard to
rufibarbis but not of any of the anomalus group, are the con-

138 Annals Entomological Society of America [Vol. XII,

vincing facts in support of the conclusion reached in regard to
completus.

There is some doubt about Aszus dascyllus of Walker but
some things in the description suggests rufibarbis and the great
amount of collecting in Massachusetts, the type locality, up to
the present has not revealed any other possibilities, so have
used my best judgment. Walker himself transferred dascyllus
to Erax later when he had a better understanding of the genus.

Van der Wulp’s figure of the female of his Proctocanthus
virginianus is an excellent outline of Erax rufibarbis. The tip of
the oviduct is quite suggestive and it is not right for Procta-
canthus at all. The size is right and the description is not bad.

Erax bicaudatus n. sp.

Male. Total length 24 millimeters. Mystax and beard white,
palpi black with black hair. Thorax largely with white hairs and
bristles. Mid-dorsum before the suture with a row of rather long black
hairs, which is suggestive of the mane in the jubatus group, although
quite different, for it is made up of numerous hairs somewhat sparsely
placed over a rather wide strip and does not form a dense comb as in
the group mentioned. Well preserved specimens show it to best
advantage. Scutellum with black hair on the disc and numerous white
bristles placed irregularly on the margin. Wings hyaline, costa not
dilated, branching of third vein a little before the base of the second
posterior cell, stump of the anterior branch somewhat longer than the
basal section. Legs largely black with numerous white hairs and black
bristles, tibia reddish on basal third, especially the anterior and middle
pairs. Abdomen dark, partly gray pollinose with much white hair
and dorsally with some short black hair, segments six and seven silvery.
Hypopygium medium sized, black, reddish in part, clothed mostly
with white hair.

Female. Total length 25 millimeters. Colored like the male and
differing only in sexual characters, oviduct about as long as abdominal
segments six and seven, viewed from side narrowed abruptly before
apex with a backward projecting point which from dorsal view is seen
to be furcate.

Male type from Montclair, Colorado, July 30, 1898, and
allotype from Morrison, Colorado, May 20, 1898. Both
localities are near Denver.

Numerous other specimens of both sexes from a _ wide
range, varying in size from 17 to 31 millimeters, and slightly
in color, some specimens appearing much blacker than others.

The structure of the oviduct will separate the female from
all species of its genus except rufibarbis, of which bicaudatus

1919] Hine: Genus Erax 139

appears to be the western representative. Over much of its
range the species is very differently colored from rufibarbis,
but the two approach each other in some localities. An
occasional specimen has three submarginal cells either in one
or both wings. The furcation of the third vein averages
slightly further forward in bicaudatus than in rufibarbis, but
this is variable within limits in both species. Specimens have
been sent in under no less than half a dozen different specific
names.

The stramineus Group.

One of the largest groups of the genus Erax is characterized by
having the furcation of the third vein distinctly anterior to the base of
the second posterior cell. In most all the species the mesothorax has
many bristles posteriorly and the scutellum has many bristles on its
margin and often numerous long hairs on its disc. Practically all the
species are western or southern in distribution.

1. Furcation of the third vein at or before the middle of the distance between

the base of the second posterior cell and the small cross vein............ 2
Furcation of the third vein distinctly beyond the middle of the distance
between the base of the second posterior cell and the small cross-vein....18

2. Male abdomen without long hair parted at middle and directed outward
ONG Mes OLAITOLESESTECMUS Nara eye teie ek ies oe See ees eh s See ae tenes 3

Male abdomen with long hair parted at the middle and directed outward
BipeB ee MOLG Se OMICILSE 20. cer aed Mile aiid binls = Os Se. n> Ketel nie xe 5

3. Hypopygium from side view plainly divided at tip, oviduct conical,
interruptus Macquart

Hypopygium from side view not divided.. 4

4. Hypopygium from side view narrowest at apex, tibiae basally bright reddish
CIC WA Reedy MINE eRe Ces Pen AE ged texanus Banks, n. sp.
Hypopygium from side view not narrowed at apex, tibiae basally dark reddish
OURO VINE LS ce We seh Ba hear Reece Cae & ae ee oe californicus Shaeffer

mE INSURE CULO EArt ean Sete. BEET) Sed oe fo) Pe aa en NGI eh a teat 6
LN Sf SEBS OraPO BSS TENG iia ne akc Aaa Se a Rage Oat as SANT Mee peed Tete tae ees 11
Opebae climictoriar mele Tea. tic Sein bla. aie ov ee keel Melee & Skieat nk Yee td 7
en SEIS OVID ay ds Pech coil UAE aoc ied Ng oe OS A a 9

7. Male third abdominal segment only with long white hair parted at the
muddlesandedimectedvoubwardt) Jes. .5. 59). sacs seer aurivestitus n. sp.

Male abdomen with at least four segments with long white hair parted at
He MG Ces GITeCLeGMOUUWArG sc. cf) fo. won gets antec as woe eace cena ee 8

8. Abdomen white haired to base. Hypopygium with a rather conspicuous
LUTNOL MaleinhemOriyaat rapemse a. ¢ hess. Uae. ete. Adi canus Hine

Abdomen, especially the male, conspicuously darker on first and second
segments than on the following ones, no tuft of hairs at apex of Hypopygium,
dubius Williston

9. Body conspicuously dark colored, male with abdominal segments four, five

Ana sicily, | whipernamedie: A khGs / kris, 5 clashed e716 9% splendens Williston

Body largely yellowish gray pollinose, male with abdominal segments three

‘to seven inclusive white haired.............................-....- se 10

10. Usually not to exceed 18 millimeters in total length..... stramineus Willis.
Specimens 20 millimeters or over in LIGSay EAE, Sr ceed Seebeh he rapax Osten Sacken

11. Mystax composed of black and white bristles and hairs intermixed. Fel,

Ria irimeve KCl paaletaslentJe) uel taba Aah oity chet Mid? Alb Gaica das vy bebe eee ee 13

140 Annals Entomological Society of America [Vol. XII,

12. Hypopygium from dorsal view conspicuously widened at apex. .inflatus Hine

Jeli jovojongemhunonts reQOimn WyPKOVSMISL 5 soo ne dacpeyouonoapebeeacuak eximius Bellardi
13. Bristles and hair of body and legs largely pale. Hypopygium from dorsal
view distinctly widened! atrapexwe ns )s2 ss. . ore eee varipes Williston

Bristles of the legs largely black. Hypopygium not widened apically..... 14

14. Hypopygium with a rather conspicuous tuft of hair below at apex, furcation
of the third vein very near the middle of the distance between base of
second posterior cell and small cross vein.................. canus Hine
Hypopygium without conspicuous tuft of hair apically, furcation of the third
vein plainly before the middle of the distance between the base of the

second posterior cell and the small cross-vein,...........-...-i-+5-->: 15

15. Ey popycium@ieplainly notehedionyapicaliborden pene ere eee tee 16
Hypopyzium not notched on apical border’. 0.02.5) ees eee eee 17

16. Thorax yellowish pollinose, first two segments of male abdomen colored
likes the wthve raccsd is, eee Siac cae heel ce eae ee argentifrons Hine
Thorax gray pollinose, first two male abdominal segments colored like the
following’ Ones: as oc aoe ae See ee cel ne see argyrosoma Hine

iia Ey popyciumeLguncateray apexiare aan see eae ec en ieieee truncatus Hine
Eby popy sium roundedratapexomee sek roee ao ater pallidulus Hine

18. Femora largely red, especially posteriorly........, ode Hen Se thchep he MOP PEE 19
Remoracentineliys Dla kes 3.002. pis a pereatvoncikayersfere scat orate ant oe Ae eee ae eee 20

19. Male abdominal segments two, three and four with black markings,
argyrogaster Macquart
Male abdominal segments two, three, four, five and six with black markings,
triton Osten Sacken
20. Mystax white, scutellum with long white hair and bristles................ 21
Mystax yellow, scutellum with yellow or black hair and bristles.......... 22
21. Very small species, 12 millimeters; mesothorax posteriorly white haired,
pilosus n. sp.
Larger species, not under 15 millimeters, mesonotum posteriorly black
IVE 5 as edn yma te wees cuca SS See REECE oe eae coquillettii n. sp.
22. Mystax bright yellow, scutellum with long yellow hair and bristles,
auripilus Hine

Mystax pale yellow, scutellum with black hair and bristles........ .23
23. Male, first three abdominal segments without silvery markings, female
UiauiKormanlhye yelltohwe jorOlhiNONS: Gass oooacnopenaaceosauccsoudnoe plenus Hine

Male, only first two abdominal segments without silvery markings; female,
abdomen with a series of large black markings dorsally, one to each seg-
'00Xc) 11 ieee a ea REE eS AN teen Ae etary AA ein id 6 15. nemoralis Hine

Erax stramineus Williston.

Total length 15 to 18 millimeters. Mystax, beard and hairs of the
palpi pale but distinctly yellow, bristles of the front and occipito-
orbital bristles sometimes black, but they may be yellow and in most
specimens examined at least part of them are pale yellow. Dorsum
of the mesothorax with black hairs and bristles in many specimens,
but in some the bristles posteriorly, at least are pale; scutellum with a
row of many long black or pale bristles on the margin. Wings hyaline,
narrow, furcation of the third vein distinctly before the middle of the
distance between the anterior cross-vein and the base of the second
posterior cell, stump usually near twice as long as the basal section.
Legs nearly entirely black in some specimens, tibia reddish in others.
In many species of the genus the tibia normally are widely red basally
and narrowly black apically. Here it may be said that the normal
color of the legs is uniformly black and the variation toward red appears
to be due to the lack of varying amounts of the dark pigment, thus

1919] Hine: Genus Erax 141

giving more or less the appearance of a faded condition to the tibiz
and even to the other parts of the legs. Legs with much pale yellow
hair and black and some pale bristles. Male. First and second
abdominal segments gray pollinose with rather sparse pale hairs,
segments three, four and five with prominent white pile parted at the
middle and directed outward; segment six similar, but pile much less
prominent; segment seven silvery pollinose with short and scattered
white hairs. Hypopygium, from dorsal view, rather narrow, sides
nearly parallel to apical third then gradually widened; from side view,
rather long of nearly the same width throughout and with a prominent
extension apically. See Figures 1 and 33. Female, abdomen nearly
uniformly gray pollinose, oviduct shining black, somewhat longer
than abdominal segments five, six and seven.

Several specimens of both sexes from Montana and
Wyoming.

It is quite nearly related to rapax, as Osten Sacken stated
when he described the latter. The smaller size and the fact
that the entire third abdominal segment is silvery seem to
distinguish stramineus. A quantity of material, including
the two species, is at hand from the territory extending from
Montana to Guadalajara, Mexico, and I find it difficult to
make a satisfactory separation from all localities, but this is
easy to do with specimens from the northern and southern
limits of the area indicated.

Erax dubius Williston.

There is some doubt of the status of this species, for Williston
gave the name in a key in Volume 12, Transaction of the
American Entomological Society, page 64, but did not give
a description in full. It is reasonably sure, however, that his
short diagnosis of Erax n. sp., on page 68 of the same paper,
refers to dubius. A distinct effort has been made to properly
identify the species. By correspondence with C. P. Alexander,
of the University of Kansas, I find that the specimen which
Williston mentions from Washington has been lost, but a
specimen from Arizona, very likely that one referred to in the
diagnosis, is labelled as the type in the museum at Lawrence.
A study of this proves it to be very near stramineus, but with
black femora and red tibiz and tarsi. The color of the body
is exactly that of stramineus and the form of the hypopygium
is the same. Williston described stramineus as having entirely
black legs, but the fact is that-in a long series of specimens

142 Annals Entomological Society of America [Vol. XII,

variations occur even to the extent of having the tibize and tarsi
in large measure reddish. It seems best, after considering
the facts to regard dubius as no more than a variety of stramineus
if not a synonym.

Several specimens from Wyoming agree with the type
specimen of dubius.

Erax rapax Osten Sacken.

Total length 20 to 25 millimeters. Mystax, beard and palpal
hairs decidedly yellow, frontal and occipito-orbital bristles mostly black.
Bristles and hairs of the mesothoracic dorsum largely black, margin of
the scutellum with a row of many long bristles which are viarable
between black and yellow in different specimens. Wings yellowish
hyaline, furcation of the third vein much nearer the small cross than
to the base of the second posterior cell, stump distinctly longer than the
basal section. Legs black with an abundance of yellow hair, bristles
black, but an occasional pale one intermixed. As in stramineus the legs
vary in color and many specimens have the bases of the tibia reddish
while others have the legs generally paler throughout. Male abdomen,
segments one, two and anterior part of three dark, nearly black, re-
mainder of three and all of four and five silvery and with silvery hairs
parted at the middle and directed outward, six similar but hairs shorter,
seven silvery pollinose. Hypopygium black with mostly black pile,
although more or less yellow pile is likely to be present. Ventrally
this pile is longer and more conspicuous than elsewhere. See Figure
41. Female generally less pilose than the male, abdomen gray pollinose,
oviduct shining black, somewhat longer than abdominal segments
five, six and seven.

Common in southwestern United States and Mexico.
The type locality is Northern Sonora.
-_ . This species is difficult to separate from stramineus in the
northern part of its range.

Erax aurivestitus n. sp.

Male. Total length 22 millimeters. All the vestiture of the head,
except the bristles in the region of the ocelli and one or two occipito-
orbital bristles on each side, distinctly yellow; head, thorax and base
of abdomen yellow pollinose, dorsum of the mesothorax with black
hairs and bristles, pleuree mostly with black hairs and bristles, but with
some pale ones intermixed, coxz with yellow bristles, other parts of
legs mostly with abundance of yellow hair and some black bristles,
femora black, in ground color, tibia and tarsi dark reddish, wings
yellowish hyaline, furcation of the third vein distinctly before the
middle of the distance between the anterior cross vein and the base
of the second posterior cell, stump longer than the basal section. First
and second abdominal segments dark in color with mostly dark hairs

1919] Hine: Genus Erax 143

- both above and beneath; third segment with a peculiar band of long
hair, silvery above and darker beneath, extending entirely around it
and parted on the mid-dorsum and also on the mid-venter and in each
case directed outward; the hairs on the anterior part of the segment
are directed somewhat backward and those on the posterior are directed
forward to about the same extent; narrow posterior margin of the third
segment and segments four to seven inclusive with but very little pile
but conspicuously silvery pollinose. Hypopygium black and almost
entirely black pilose, but with a little yellow pile basally and also
apically; black pile especially long and prominent on sides and venter;
dorsally of nearly uniform width, but wider and shorter than in sagax;
from side view a very prominent extension at apex. Figure 42.

Male type from Morelos, Mexico, collected by D. C. Craw-
ford, in the author’s collection.

The peculiar third abdominal segment and the hypopygium
characterize this species.

Erax splendens Williston.

Total length 22 to 25 millimeters. Mystax and beard bright
yellow, hair and bristles of the front and the occipito-orbital bristles
black, thorax rather sparsely brown pollinose and with black hair,
wings yellowish, furcation of the third vein not far from the anterior
cross vein, stump a little longer than the basal section. Front legs
with dense yellow hair anteriorly on coxe, and ventrally on femora,
tibize and tarsi, but there is much black hair on the appendages also;
middle ahd hind legs mostly black haired, but there is some variation
in a series of specimens. Legs entirely shining black in ground color.
Male abdomen black with black vestiture, except segments four, five
and six, which are silvery. See Figure 39. Female, abdomen dark,
segments two to six inclusive yellowish gray pollinose with rather small
irregular markings dorsally. Oviduct shining black, hardly as long as
the last three abdominal segments.

Several specimens of both sexes from Guadalajara, Mexico,
collected by McClendon.

The very dark color of the body of this species with the
bright yellow mystax and beard makes its identification easy. ’

. Erax truncatus Hine.

Thorax yellowish-brown, above, abdomen gray, first four segments
with long white hairs, legs black with the exception of the basal part of
each tibia which is light reddish. Total length 22 to 30 millimeters.

Mystax and beard white, ocellar, occipito-orbital and a transverse
row of bristles on the dorsum of the prothorax black, palpi with black
and white hairs intermixed, dorsum of the thorax and the scutellum
with many black hairs and bristles, but there are some white ones
intermixed, wings hyaline.

144 Annals Entomological Society of America [Vol. XII,

First segment of the male abdomen with long white hairs on each
side, second, third and fourth segments with long silvery hair parted
at the middle and directed outward, fifth, sixth and seventh segments
silvery white pollinose, but without long hair; hypopygium rather
large, from dorsal view about as wide as the last segment of the abdomen,
from side view most prominent near middle above and cut off at tip
so as to give a truncate appearance. Figure 45.

First seven segments of the female abdomen silvery white; oviduct
slender, shining black, about seven millimeters in length.

Several specimens from the Huachuca Mountains, Arizona,
July 28, 1907.

The large compact hypopygium of the male and the long
oviduct of the female give this species a distinct appearance
which makes its separation from others easy.

Erax pallidulus Hine.

A pale colored species with black legs and hyaline wings. The
male has the hypopygium small and, from dorsal view, very narrow.
Total length 18 to 28 millimeters. Mystax very pale yellowish, beard
white, palpi black with white hair, occipito-orbital and ocellar bristles
mostly black; thorax dorsally pale yellowish gray with short black hair
anteriorly and black and white bristles and hairs posteriorly, scutellum
with pale hairs and bristles, legs black, except bases of tibize, which are
pale, wings hyaline.. Male abdomen silver white, first four segments
with long white hair, two, three and four with hair parted at the middle
and directed outward, five, six and seven without long hair, hypopygium
small, black, narrowed toward apex where, from lateral view, it appears
nearly evenly rounded. Figure 44.

Female colored like the male. Oviduct black, only slightly longer
than the last two abdominal segments.

Three male specimens and a female from Albuquerque,
New Mexico, collected by J. R. Watson, and one female from
Colorado.

The short hypopygium and oviduct are characteristic of
this species.

Erax argyrosoma Hine.

Body nearly uniformly white all over, mid-dorsal stripe of the
thorax not plainly marked. Length 23 to 25 millimeters. Mystax
and beard white, palpi black with white hair, some of the occipito-
orbital bristles black and some white; legs black except the basal parts
of the tibiz, which are yellowish haired, wings hyaline; anterior part
of the dorsum of the thorax with short black hair; posterior part and the
scutellum with black and white bristles and hairs. First four abdominal
segments of the male with long white hair; on two, three and four,
this is parted at the middle and directed outward, segments five, six

1919} Hine: Genus Erax 145

and seven silver white, but without long hair, hypopygium black with
short white hair, somewhat notched at the apex with the lower part
extending into a prominence. Figure 40.

Female abdomen gray pollinose, oviduct shining black, four milli-
meters in length.

Taken by J. R. Watson near Albuquerque, New Mexico.
More than a dozen specimens from other localities in New
Mexico and western Texas.

Erax auripilus Hine.

A medium sized species characterized by abundance of rather bright
yellow hair on all the parts of the body.

Male. Total length 22 millimeters. All the hairs on the various
parts of the head yellow, sternum and sides of the thorax with yellow
hair, dorsum with black hair, wings clear hyaline, furcation of the third
vein distinctly before the base of the second posterior cell and with a
long appendage, costa not enlarged near the tip of the auxiliary vein,
posterior branch of the third vein meets the costa distinctly before the
tip of the wings; legs with yellow hair and black bristles, in most part
black in ground color, but the basal half or more of each tibia is bright
yellow; abdomen with yellow hair, segments two to five black above
with light colored lateral and hind margins, segments six and seven
silvery white, hypopygium of medium size, shining black in ground
color and clothed with yellow hair. Figure 47.

One male, taken at Clifton, Texas, May 29, 1907, by E. B.
Williamson. A male in the collection of the Academy of
Natural Sciences was taken at Round Mountain, Texas.
Several females in the U. S. National Museum from different
localities in Texas.

Erax canus Hine.

Male. Total length 24 millimeters. General body color rather
hoary white produced by white pollen and hair, dorsum of the thorax
with black hair and bristles, legs with black bristles, mystax and beard
pale yellow, bristles of the front black, palpi black with pale yellow hair
and bristles, wings hyaline, costa uniform throughout, furcation of the
third vein distinctly before the base of the second posterior cell and
with a distinct appendage, the posterior branch of the third vein bends
forward to meet the costa plainly before the apex of the wings, legs
black with the basal third of each tibia reddish; abdomen uniformly
hoary white, segments two, three and four on the dorsum with long white
hair parted at the middle and directed outward; hypopygium of medium
size, black and clothed with white hair. Figure 38.

Female colored like the male, but abdominal segments two, three
and four devoid of the peculiar long hair described for the male. Oviduct
shining black, slightly longer than abdominal segments six and seven
combined. :

146 Annals Entomological Society of America [Vol. XII,

Male and several females from Claremont, California, sent
in by Carl F. Baker; three males in the U. S. National Museum
from Los Angeles County, California, Collection Coquillett.
The smallest female measures only 17 millimeters in total
length.

Erax eximeus Bellardi.

Total length 17 millimeters. Mystax pale with some black bristles
intermixed, beard white, palpi black haired, ocellar and occipito-
orbital bristles black. Thorax with black hairs and bristles, scutellum
with several black, marginal bristles; femora reddish posteriorly and
at apex, black anteriorly, tibiae and tarsi largely red with mostly white
hairs and black bristles, wings very slightly smoky hyaline, branching
of the third vein midway between the small cross vein and the base of
the second posterior cell, stump only very slightly longer than the
basal section. Abdomen largely gray, first segment, anterior two-
thirds of second and the anterior parts of segments three and four black,
posterior part of the second and nearly all of the third and fourth with
long hair parted at the middle and directed outward; segments five,
six and seven with much shorter hair. Hypopygium dark red, black
in places. See Figure 46.

A male specimen from Cuernavaca, Mexico, collected by
Dt. Crawtord.

This species does not answer in detail to eximeus, but goes
there in Bellardi’s Key. It is the only Erax I have with the
apexes of the femora red following black.

Erax inflatus Hine.

A dark colored species with the mystax composed of black and gray
hairs intermixed, wings hyaline, slightly fumose at apices, legs black
with the exception of the extreme bases of the tibize which are yellowish
red. Length of the males 20 to 25 millimeters; of the females 22 to 26
millimeters.

Front yellowish-gray pollinose, antennz black, first two segments
clothed with gray hair, occipito-orbital bristles and ocellar bristles
black, mystax composed of black and gray hairs intermixed, beard
silky white, palpi black with black hairs; thorax brownish-gray pollinose
with a dark mid-dorsal stripe abbreviated posteriorly, clothed with
gray and black hairs and bristles existing in different proportions in
different specimens; legs black except the extreme apices of the tibia,
which are yellowish-red, furnished with white hairs and black bristles
and some golden pile on the underside of some of the segments; wings
hyaline, slightly darkened at apex.

Male abdomen with the apex of the second segment and all of the
segments from three to seven inclusive silver white, segments two and
three with long white hair parted at the middle and directed outward,

1919] Hine: Genus Erax 147

four and five shows this arrangement somewhat, but the hairs are
short; hypopygium clothed mostly with white hair, enlarged at apical
half until it is nearly twice as wide as the seventh abdominal segment.

Female abdomen with each segment white pollinose at sides and
apex, otherwise black above, oviduct black, scarcely five millimeters
in length, equivalent to the last three abdominal segments.

Twenty specimens received from F. Grinnel, Jr., and taken
in Los Angeles County, California. A very distinct species
on account of the male hypopygium which appears as if inflated
and is nearly twice as wide as the seventh abdominal segment.
Males and females from Los Angeles, Kern and San Bernardino
counties, California, Coquillet Collection.

Erax nemoralis Hine.

A dark colored species with yellowish mystax and fumose wings.
Femora, tarsi and apices of the tibize black, bases of tibia reddish-
brown. Length, male, about 25 millimeters; female, to the tip of the
oviduct, 24 to 27 millimeters. Face and front covered with yellowish
dust, mystax and beard pale yellow, occipito-orbital and ocellar bristles
black, as are most all of the hairs and bristles of the front, palpi black,
furnished with many black hairs which often are intermixed with pale
yellow ones. Prothorax mostly clothed with pale hairs, remainder of
thorax with many black hairs and bristles, but these often are reduced
by the presence of greater or less numbers of pale ones; mid-dorsal
stripe dark and well marked, abbreviated behind and divided anteriorly,
on either side the markings are in the form of ill-defined spots caused
by the difference in intensity of the rust-colored dust which gives the
thorax its peculiar color; legs black, except the bases of the tibiae, which
are reddish-brown, clothed with black bristles and pale hairs of different
lengths, the shorter ones recumbent, some of the segments inwardly,
more especially the metatarsi and front tibia, clothed with golden
recumbent pile.

In the male abdominal segments one, two and base of three dark,
largely clothed with black hair, apex of three and all of four, with the
exception of a small black triangle on each anteriorly, white with long
white hair parted in the middle and directed outward, five and six
silver white with very short hair, remainder of abdomen black, with
black hair, however, in some specimens part of seven is whitish and
there may be a few pale hairs on the hypopygium. -Figure 51.

In the female the segments of the abdomen are gray on the sides
and hind margin, otherwise black above, but the latter color is not well
defined, especially if viewed with a lense; oviduct about six millimeters
in length, equivalent to the last four abdominal segments.

Several specimens of both sexes procured in a brushy
woodland at New Roads, Louisiana, July 15, 1905. The
specimens were captured while resting near or on the ground.

148 Annals Entomological Society of America [Vol. XII,

It is a predaceous insect of possible value on account of its
size. Other specimens from Texas and from Falls Church,
Virginia.

Erax argentifrons Hine.

Much like rapax. Front white pollinose, mystax white, legs with
white hair. Length 18 to 23 millimeters.

Palpi black with white hair, occipito-orbital and ocellar bristles black,
antenne black, first two segments with white hair, beard white. Thorax
yellowish-brown with the usual mid-dorsal stripe darker, hairs of sides
almost uniformly pale, of dorsum variable between pale yellowish and
black; wings hyaline, legs black, except the extreme bases of the tibia,
which are reddish-yellow, clothed with pale hairs and black bristles.
First two segments of the male abdomen colored like the thorax, seg-
ments three to seven inclusive silver white, apex of two, all of three and
four with long white hair parted at the middle and directed outward,
hypopygium much narrower than in stramineus, clothed with black and
white hair and distinctly notched at the apex. Figure 35.

Female abdomen uniformly yellowish pollinose and clothed with
pale hairs, oviduct about four millimeters in length, equivalent to the
last three abdominal segments.

Specimens of both sexes taken in Clark County, Kansas,
by Dr. F. H. Snow.

As has been stated, the species has much the appearance
of rapax, but the somewhat stouter form, the white mystax
and beard and much slenderer hypopygium designate it as
wholly distinct from that species.

Erax plenus Hine.

A large, robust species, varying in length from 23 to 30 millimeters.
Male. Total length 27 millimeters, antenne black, style nearly twice
as long as the third segment, palpi black and clothed with pale yellow
hair, face and cheeks with abundance of pale yellowish hairs, ocellar
bristles and several bristles on the upper part of the occiput black.
Thorax yellow pollinose with most of the hairs and bristles black;
wings with a very pale yellowish tinge, costa not thickened near the
tip of the auxiliary vein, furcation of the third far before the base of the
second posterior cell and with a distinct appendage, posterior branch of
the third vein reaches the costa distinctly before the tip of the wing;
legs clothed with pale yellowish hairs and black bristles, black, except
the tibia which is largely reddish. First three abdominal segments
dark, mostly with black hair above and white hair beneath, four white
with long white hair parted in the middle and directed outward; five,
six and seven white, hypopygium dark in color, short and soméwhat
tumid. Figure 48. .

1919} Hine: Genus Erax 149

Female unusually robust for an Erax. Abdomen, except the
oviduct, uniformly pale yellowish pollinose, oviduct shining black, about
as long as abdominal segments five, six and seven combined. Otherwise
colored as in the male.

Several specimens of each sex from, Douglas County,
Kansas, 900 feet elevation (F. H. Snow). From Onaga,
Kansas, and from Osborne County, Kansas, 1557 feet elevation,
collected August 3, 1912, (F. X. Williams), Ardmore, Indian
Hermtory, (C R.~Jones);,) Plano, Texas, {@ucker);..Waco,
Texas, (Belfrage), and other localities in Texas.

A male from Onaga, Kansas, taken August 20, 1901, is like
the other males, except that abdominal segment seven is
black instead of silver white. This gives the specimen quite a
different appearance, but since similar variations have been
observed in other species of the genus it is not considered
specific here.

Erax coquillettii n. sp.

Male. Total length 16 millimeters. Mystax and beard white,
ocellar tubercle and posterior orbits above with black bristles, meso-
thorax with black hair and bristles. Scutellum with abundance of
long white hair on the disc and numerous mostly white bristles on the
margin, wings hyaline, furcation of the third vein distinctly anterior
to the vein which closes the discal cell, but plainly beyond the middle
of the distance between this vein and the anterior cross-vein, stump
of the anterior branch much longer than the basal section. Femora
black, tibize yellowish red with the extreme apexes dark, tibiz dark
red, legs with white hair and black and white bristles. Abdominal
segments one, two and basal part of three black in ground color and
only sparsely hairy, apical part of three and all of four and five densely
silver-white and white hairy, parted at the middle and directed outward,
six and seven silvery with very short hair. Hypopygium from dorsal
view narrow on basal third, much widened apically where the width
is nearly double that at the base. See Figure 14.

Female colored like the male. Abdomen shining black, each segment
gray pollinose posteriorly, giving a distinct banded effect. Oviduct
shining black, about as long as the last four abdominal segments.

Type male and allotype from San Diego County, California,
collected by Coquillett in April. In the United States National
Museum. Several other specimens of both sexes with the same
data.

The inflated appearance of the apical half of the male
genitalia, the nearly naked first and second abdominal segments
which are black in the male, and the banded abdomen of the
female suggests the species.

150 Annals Entomological Society of America |Vol. XII,

Erax pilosus n. sp.

Male. Total length 11 millimeters. Hair and bristles of the whole
body white, except about four black bristles above the base of each wing,
numerous white bristles on the margin of the scutellum; wings delicate
hyaline, narrow, the third vein branches near the middle of the distance
between the anterior cross vein and the apex of the discal cell, stump
on the anterior branch much shorter than the basal section. Femora
all black, remainder of legs red with extreme apexes of tibize darkened.
Abdomen, except the first segment and anterior part of the second, with
abundant silver-white hair parted at the middle and directed outward.
Hypopygium from dorsal view of uniform width for almost basal half,
then gradually widened and rounded at tip; from side view nearly
uniformly convex dorsally and concave ventrally. See Figure 15.

Male type from El Paso, Texas, April 5, 1902. Collection
Academy of Natural Sciences, Philadelphia.

The small size, delicate appearance, male genitalia and
furry abdomen are characteristic of the species. I know of no
Erax where the long white hair of the male abdomen extends
to the tip of the seventh segment as in this one.

Erax californicus Schaeffer. |

Male. Total length 24 millimeters. Vestiture of the head white,
with the exception of the ocellar and occipito-orbital bristles which are
black; mesothorax largely white hairy, but some black bristles near
the wing bases and some short black hair on the anterior part of the
dorsum. Legs long white hairy and with black bristles, femora black,
tibiz and tarsi largely red; wings hyaline, furcation of the third vein
very near the middle of the distance between the small cross vein and
the base of the second posterior cell, stump slightly longer than the
basal section. Abdomen black, largely covered with white pollen and
everywhere with white hair, longer on two, three and four and directed
outward. Hypopygium from the side of nearly uniform width, rather
obliquely truncate at apex, with a rounded prominence above and a
tuft of white hair pointing backward below. See Figure 6.

A female from the same locality is colored like the male. Oviduct
fully as long as the last three abdominal segments.

The most characteristic thing about the species is the
absence in the male of distinct silvery abdominal segments,
contrasting with others that are not silvery. The whole
abdomen is white pollinose, but in no place is this so dense
that it may be said to be silvery as in most species.

More than 20 specimens of both sexes ranging from Washing-
ton and Montana to California and Nevada, vary in size from
18 to 26 millimeters and show quite an amount of variation
in the color of the tibiz and tarsi.

1919] Hine: Genus Erax Tot

The type, which Mr. Schaeffer kindly sent me for study,
is in Brooklyn, and was collected in Shasta County, California,

Erax texanus Banks n. sp.

Male. Total length 23 miullimeters. Head white-haired; palpi
black haired, ocellar and upper orbital bristles black. Thorax with
short black hair, pleura white haired. Abdomen without parted hair,
the basal segment with long white hair on the sides and beneath, and
the apical parts of the segments whitish pollinose, sixth and seventh
silvery, sixth blackish at base. Abdomen rather long and slender,
the hypopygium much narrower than the last segment, long and slender,
black haired. Femora black, tibiz reddish on basal part, black beyond,
tarsi almost black, femur black haired above, beneath and the tibia
white haired, except that the latter has dark hair near the tip. Wings
with costa normal, third vein with end before wing tip, the fork very
much before the base of the second posterior cell. Figure 43.

From Texas (Lafr.). The hypopygium is much longer than
in Erax vartipes.

Erax triton Osten Sacken.

Face yellowish-pollinose; facial tubercle prominent, with a tuft of
- pale yellowish hairs and bristles; palpi beset with black bristles, mixed
with yellow ones; cheeks with soft, whitish hair; upper occipital orbit
with a row of stiff black bristles; basal joints of the antenne reddish,
the third joint darker; arista much longer than the third joint. Thorax
with a well-marked, broad, dark brown dorso-central stripe, its median
line feebly marked, reddish; sides of the dorsum yellowish, with slight:
brassy reflections; pleurze brownish-yellow, with soft pale hairs; scutellum
yellowish-gray pollinose, with black macrochaete. Halteres reddish-
yellow. Abdomen, male, its prevailing color, an impure, somewhat
silvery, white, with a row of black triangles, the triangles gradually
diminishing in size from segment two to segment five, segment six
with only a brownish line in the middle; segment one blackish, with
some grayish pollen on the sides; forceps rather large, elongate, reddish-
brown, beset with paler hairs and with a brush of hairs on the underside,
in shape like those of E. anomalus. Female, the black triangles of nearly
equal size on segments two to four, occupying the whole middle of the
segment and touching the hind margin with the apex; the sides of the
segments filled by triangles of an impure silvery-white; on segments six
and seven the lateral margins only whitish, the middle occupied by an
opaque square of brownish-black; segment five forms the transition
in both sexes, but principally in the male; the silvery parts of segments
two to four beset with sparse white hairs, combed outwards; ovipositor
comparatively short, equal to about two or two and a half of the pre-
ceding segments; legs deep reddish, with the usual appressed pubescence
or whitish hairs and long, soft, whitish or yellowish hairs (especially
in the male) and black bristles; femora black on the underside. Wings

152 Annals Entomological Society of America (Vol. XII,

with a slight yellowish-brown tinge; second submarginal cell distinctly
appendiculate, rather long, its proximal end reaching considerably
beyond the proximal end of the second posterior cell; no incrassation
of the costa in the male. Length 23 to 25 millimeters; without ovi-
positor 22 to 23 millimeters, with it about 26 millimeters.

Erax argyrogaster Macquart.

Total length 22 millimeters. Vestiture of the head pale yellowish,
except the palpe bristles are black, as are some of the occipito-orbital
bristles, legs largely red, femora black beneath only, bristles black;
wings yellowish, branching of the third vein just beyond the middle
of the distance between the small cross-vein and the base of the second
posterior cell, stump about the length of the basal section. Abdomen
largely white, first segment dark, second with a large black triangle,
including more than half of the anterior part of the segment, third with
a smaller black triangle, fourth and each following segments with a
narrow black mid-dorsal line which on five, six and seven is almost
obsolete. Hypopygium elongate, narrow and reddish in color.

One male from Tehauntepec, Mexico, (Sumichrast). This
agrees well with Macquart’s description. I have not sufficient
material to make a comparative study of argyrogaster and
triton, but they appear to be much alike and perhaps the same
species.

Erax varipes Williston.

Total length 22 to 28 millimeters. A rather large and robust ash-
colored species with pale bristles everywhere. There may be some black
in front of the scutellum and on the legs and.in some specimens the
bristles of the legs are largely black, although in the material studied
at least a few white bristles appear scattered irregularly. Hair and
bristles of the whole head entirely pale, short hair on anterior part of
mesothorax black, scutellum with numerous pale bristles on its margin,
wings hyaline, furcation of the third vein much nearer the anterior
cross-vein than the apex of the discal cell, making the second submarginal
cell very long. Legs variable in color, sometimes largely reddish, but
usually the femora dorsally and the tibize basally are reddish with the
other parts dark. Abdomen nearly uniformly gray and rather short
pilose in both sexes; in the male there is not much indication of the pile
parting at the middle and pointing outward as in most species with
which varipes appears to be most nearly related. From dorsal view
male genitalia dark reddish, basally about as wide as the seventh abdomi-
nal segment, sides parallel to near the middle, then rather rapidly
widened and rounded apically as in Figure 59. For lateral view see
Figure 49. Oviduct mostly black, slightly longer than abdominal
segments six and seven.

1919} Hine: Genus Erax 153

Specimens from Western Kansas and from Arizona.
The dorsal red stripe of each femur and the male genitalia
are very characteristic of the species.

Erax interruptus Macquart.

Length 22 to 27 millimeters. General color brownish, mystax pale
with a few black bristles above, palpi with black bristles, beard pale,
thorax largely gray pollinose. with a wide, dark mid-dorsal stripe nar-
rowly divided before. Scutellum with many black bristles on the
margin; wings reddish hyaline, legs largely reddish brown, under sides
of femora and tibiz dark, nearly black. Abdomen nearly black in
ground color, yellowish-gray pollinose spots on the hind corners of the
segments, not meeting at the middle on segments one to four, segments
five with the posterior margin pollinose, segments six and seven wholly
pollinose in the male and each with a small black triangle with the base
posterior, in the female.

This common species is easily known by the furcate male
genitalia and the conical oviduct, not compressed as in other
species of the genus and about as long as abdominal segments
six and seven. See Figure 37.

It is widely distributed South, reaching from Coast to Coast,
from the latitude of Kansas to Gautemala and even further
south. It was one of the early North American species
described and has had several names applied to it as may be
seen in the list of synonyms on a previous page.

154 Annals Entomological Society of America [Vol. XII,

EXPLANATION OF PLATES VII, VIII AND IX.

Wing of Erax:
» 1, stramineus; 2, barbatus; 8, jubatus; 4, aestuans; 5, anomalus; 6, californicus.

Hypopygium of Erax, dorsal view:
13, prattit; 14, coquilletti; 15, pilosus; 16, tabescens; 59, varipes.

Hypopygium of Erax, lateral view:

7, willistoni; 8, barbatus; 9, bicolor; 10, anomalus; 11, candidus; 12, pernicis;
17, @stuans; 18, apicalis; 19, kansensis; 20, loewit; 21, belfragei; 22, auri-
mystaceus; 28, fulvtbarbis; 24, portoricensis; 25, slossone; 26, tabescens;
27, mexicanus; 28, affinis; 29, nigritarsis; 30, rufittbia; 31, nigrimystaceus;
32, stylatus; 338, stramineus; 34, cingulatus; 35, argentifrons; 36, haloesus;
37, interruptus; 38, canus; 39, splendens; 40, argyrosoma; 41, rapax; 42. auri-
vestitus; 48, texanus; 44, pallidulus; 45, truncatus; 46, eximeus; 47, auripilus;
48, plenus; 49, varipes; 50, cressont; 51, nemoralis; 52, subcupreus; 58, costalis;
54, jubatus; 55, parvulus; 56, latrunculus; 57, concinnatus; 58, carinatus;
60, wnicolor.

Vou. XII, PLATE VIII.

ANNALS E.S. A.

inp)
N

ANNALS E.S. A. VOL. XII, PLATE IX.

4| os es
42 4

56
Foi
on oy 60

1
4
°
A!
'
i
i
i
i
ai
ni

ie

s

me a,

; } f | t f
‘ at maf hi = -:
A. < ae a
men 9s: “ay lat
Fa ;

Pa reow s

i i Pi eS
a Yi uh oo aan
mv sth RNS ba hy i ie

‘ ‘an ae
J ‘
L
w Ye i
-~ \ 7
i di
‘ \ -
y
t if ey
iy (]
uy i "
: fe
i
7
q
t - : I nh ae A
- at fo oe gh ,
L : al L "
‘1 *y, *: Ke WaT | ah
a ix: ¥ ; ce ]
bi Tig?
i | THIN ie Vast
7 4 1 Ou -
. .
> a ay ogy i

SOME CHALCID-WASPS REARED FROM
CECIDOMYID GALLS.

By A. B. GAHAN,
Entomological Assistant, U.S. Department of Agriculture, Bureau of Entomology.

All except two of the species of Chalcidoidea described
and listed in this paper are stated by the collectors, Mr. V. L.
Wildermuth and Mr. E. G. Smyth, to have been reared from
galls of the alfalfa gall-midge, Asphondylia websteri Felt. The
two exceptions were reared from unidentified gall-making
Cecidomyids, one being from a gall on Pluchea borealis and
the other on Suaeda sp.

Of the species reared from galls of Asphondylia websteri
the exact host relations of some are unknown. Some are
undoubtedly primary parasites of the midge, while others are
believed to be secondary upon that host. The clover seed-
chalcid, Bruchophagus funebris Howard, was reared from some
of the galls along with the parasites and it is possible that
some of the parasites may have had this seed-chalcid for host.
Mr. Wildermuth states that all specimens reared under Tempe
Nos. 2223, 2224 and 2251 were reared from dissected Asphon-
dylia webstert material and are therefore positively known
to be either primary or secondary parasites of the midge. The
material under Tempe No. 2264 was reared from collected galls
of Asphondylia websteri placed in vials, and hence the host
relations of this material can only be surmised. It was in the
lot of material under this number that Bruchophagus funebris
was found and it is probable as will be pointed out under
discussion of the species that at least one species of parasite,
Tetrastichus bruchophagi, came from this host.

Family EURYTOMID.

Eurytoma medicaginis new species.

Female. Length 2mm. Head and thorax with strong umbilicate-
punctate sculpture and thinly clothed with short silvery white hairs.
Antenne rather short; scape not reaching the front ocellus; pedicel
conical, narrower and scarcely as long as the first funicle joint; ring-
joint small; funicle five-jointed, the joints, except the first, subquadrate;
the first joint slightly longer than broad, the fifth slightly broader than

159

160 | Annals Entomological Society of America [Vol eur

long; club three-jointed, a little broader than the funicle, and not
quite equal in length to the three preceding funicle joints combined,
the basal joint subquadrate; second joint as long as the basal; third joint
conical; ocelli in a very low triangle, the postocellar line more than
twice the length of the ocellocular line; occiput immargined above,
cheeks margined posteriorly; dorsal portion of pronotum very nearly
as broad as the mesonotum and distinctly shorter than the mesoscutum;
mesepisternum granularly rugulose, the mesepimeron indistinctly,
longitudinally striate; propodeum very nearly perpendicular to the
apex of scutellum, broadly but very shallowly impressed medially, the
impressed area granularly rugulose; remainder of propodeum more
coarsely rugose; marginal and postmarginal veins practically equal,
the stigmal about three-fourths the marginal; abdominal petiole nearly
twice as long as the hind coxe, rather slender, flattened and nearlv
smooth above, longitudinally striate at sides and beneath, and joinin,;
the main part of abdomen ventrally, or, at least, at the antero-ventr 1
angle; body of abdomen subglobose and ending in a short sharp pc's,
strongly compressed from the sides, very strongly arched above © :e
attachment of petiole as viewed from the sides, a little longer than e
head and thorax combined and viewed laterally not one-third lo er
than thick dorso-ventrally; fifth segment the largest, approxime -ly
twice the length of fourth on the median dorsal line; ovipositor slig sly
exposed at tip; hind coxe granularly rugulose like the middle of
propodeum. Black; flagellum dark brown; scape brownish testaceous;
coxee and femora black; apices of femora and all tibize yellowish brown;
all tarsi pale yellowish, claws brown; wings hyaline, venation brownish.

Male. Length 1.35 mm. Scape somewhat swollen beneath, reach-
ing to the front ocellus; pedicel globose; between pedicel and first funicle
one small ring-joint; funicle five-jointed, the joints, except the fifth,
narrowed at apex into a distinct neck, each joint about twice as long as
the thickened portion is thick and bearing hairs about twice as long as
the length of segment; neck on apex of first segment shorter than on
the other segments and the thickened portion longer; fifth segment not
narrowed at apex, about twice as long as thick, with the hairs shorter
‘than on the other segments; between each of joints two, three, four, and
five of the funicle is what appears to be a very small transverse ring-
joint, (visible only in balsam-mounted specimens); club two-jointed, the
joints subequal in length, the apical joint ending in a very short incon-
spicuous nipple-like point; the propodeum is usually nearly uniformly
reticulately rugose all over, the sculpture of the depressed area not
markedly different from the rest of the surface; abdominal petiole
longer than the hind coxe, but not twice as long, slender, and weakly
shagreened, not striate; body of abdomen small globose, hardly more
than half as long as the thorax. Otherwise similar to the female, but
with all tibia mostly blackish-brown.

Type locality, Tempe, Arizona. Type, Cat. No. 22295,
U.S. Nat. Mus.
Host: Asphondylia websteri Felt.

1919] Gahan: Chalcid-Wasps 161

Type, allotype and seven paratypes reared by V. L. Wilder-
muth from galls of the alfalfa midge and recorded under Webster
No. 7268, Tempe No. 2251. Also a large series of specimens
from the same host and locality reared by Wildermuth under
Tempe No. 2264.

Antenna of type female on Hymenoptera Slide No. 781, and
antenne of a male and a female paratype on Hymenoptera
plgewNo: 762,00. Nat. Mus. ’

Bruchophagus funebris Howard.

Specimens of this seed-chalcid were reared by Mr. Wilder-
muth from galls of Asphondylia webstert at Tempe, Arizona,
under Tempe No. 2264. Its presence in these galls is undoubt-
edly accounted for by the fact that it not infrequently happens
that developed seeds of alfalfa are present within the gall formed
by the midge and these seeds afford opportunity for the develop-
ment of the Bruchophagus.

Family CALLIMOMID.

Callimome asphondyliz, new species.

Female. Length 2.25 mm. ‘Head and thorax shagreened above;
face below antennz less strongly sculptured than the vertex; antennz
rather short and thick; funicle seven-jointed, the funicle joints subequal
in length, but increasing slightly in breadth toward the apex, and all
joints broader than long, the first only slightly so, the last about half as
long as thick; club three-jointed, not as long as the three preceding
funicle joints; ocelli in a low triangle, the lateral ocellus separated
from eye-margin by about the long diameter of the ocellus; parapsidal
grooves complete, but weakly impressed posteriorly; propodeum without
carine and uniformly shagreened, the sculpture not as strong as on
scutellum; hind coxe laterally sculptured like the scutellum; hind femora
outwardly sculptured about like the propodeum; wings sparsely ciliated,
the cilia more or less arranged in rows; stigmal vein very short, post-
marginal also short; abdomen about equal to the thorax in length, the
first tergite smooth, other tergites weakly sculptured, more strongly
so at sides; ovipositor extending beyond the apex of abdomen about one
and one-third times the length of abdomen. Color, dull blackish
green, the head and thorax sparsely clothed with silvery white hairs;
eyes pale reddish; face, propodeum, hind femora and the abdomen
more strongly metallic than dorsum of thorax; sides of thorax and hind
cox bluish; antennal scape and all tibiz and tarsi brownish testaceous;
flagellum, median and front femora, tarsal claws and ovipositor sheaths
brownish black; wings hyaline; venation pale brownish.

Male. Length 2 mm. Tibiz all dark brown. Otherwise agrees
with the female except the general color is perhaps a little darker,

162 Annals Entomological Society of America [Vol. XII,

Type locality, Tempe, Arizona. Type, Cat. No. 22296,
igo Nat. Wins:
Host, Asphondylia websteri Felt.

Six females and two males reared by V. L. Wildermuth
from galls of the alfalfa midge under Webster No. 7268, Tempe
No. 2251. May be a secondary parasite of Asphondylia, but
is more likely primary.

Syntomaspis medicaginis, new species.

Closely resembles Syntomaspis thalassimus Crosby, but differs
by having the antennal scape pale testaceous, the face more deeply
concave and more hairy, the mesoscutum and scutellum more deeply
and closely sculptured and therefore less shining, the axille a little
more widely separated and the tibiz not so dark.

Female. Length 2mm. Face more than ordinarily deeply concave,
rather strongly sculptured and closely set with silvery hairs; lateral
ocelli separated from the eye-margin by about the diameter of an
ocellus or a little less; first joint of funicle a little broader than long,
following joints decreasing gradually in length outwardly; club barely
longer than the two preceding funicle joints; mesoscutum, axille and
scutellum deeply, closely reticulate-punctate, subopaque; cross furrow
on scutellum distinct, the surface behind the cross-furrow reticulately
sculptured but not quite as deeply and finely so as the rest of scutellum;
propodeum weakly reticulated, shining, with a few foveolate punctures
on the anterior margin but without carine, the spiracles elliptical;
discal ciliation of the forewing arranged in distinct rows’ and confined
to the area distad of a line running obliquely basad from the stigmal
vein to the posterior margin; the base of wing and a broad wedge-
shaped area immediately beneath and extending the whole length of
marginal vein entirely bare or with only a very few very weak cilia;
in the apical portion of the wing a distinct row of cilia runs from the
stigmal vein in a broad curve to the apical middle of wing; abdomen
very slightly longer than the thorax, distinctly though not deeply
reticulated, shining, the first tergite nearly smooth, tergites one to three
emarginate at the middle; ovipositor exserted the length of thorax and
abdomen. Head, thorax, abdomen, coxe and ‘all femora metallic
green, thoracic pleura and the coxe strongly tinged with purplish or
blackish; antennal pedicel and flagellum brownish-black; antennal
scape, all tibiz and tarsi and the ovipositor brownish-testaceous; eyes
pale reddish; wings hyaline, venation pale yellowish.

Male. Length 1.7 mm. Agrees with the female except that the
scape is somewhat metallic, the dorsum of thorax is slightly more
shining, the cross furrow on scutellum less distinct, the tibize except
at apex are all blackish with more or less of a metallic lustre, and the
abdomen is much shorter and smaller than the thorax.

1919} Gahan: Chalcid-Wasps 163

Type locality, Tempe, Arizona. Type, Cat. No. 22297,
Unjo. Nat. Mus:

Host, Asphondylia websteri Felt.

Type female reared by V. L. Wildermuth from galls of the
alfalfa midge and recorded under Webster No. 7268, Tempe
No. 2264. One paratype with the same data except Tempe
No. 2251. Allotype, four male paratypes and five female
paratypes reared by E. G. Smyth at Tempe from the galls
of the same insect and recorded under the same Webster
number. Wing of a female paratype mounted on a slide.

Syntomaspis umbilicata, new species.

In the sculpture of the thorax resembles elegantissima Ashmead,
and warreni Cockerell, but is easily distinguished from both by the
much shorter funicle joints, metallic colored femora and the smaller
size.

Female. Length 2.5 mm. Head viewed from in front about as
long as broad; face strongly sculptured with large shallow punctures;
antennal depression deep and nearly smooth within, the antennz
separated at base by a carinate ridge; postocellar line equal to twice
the ocellocular line, the lateral ocelli not farther from the eye-margin
than the diameter of an ocellus; antennal pedicel conical, about equal
in length to the first funicle joint; ring-joint a little more than twice
as broad as long; first funicle joint nearly quadrate, following joints
diminishing very gradually in length, the-last about two-thirds as long
as broad; club three-jointed, not broader than the funicle and about as
long as the three preceding funicle joints; pronotum rugulose; meso-
scutum with large, closely placed umbilicate punctures; surface of
the scutellum before the distinct cross-furrow sculptured like the
mesoscutum, behind the cross-furrow finely reticulately sculptured;
propodeum polished, without carinz and with a row of large punctures
along the anterior margin; ciliation of the forewing similar to that
figured for Syntomas pis medicaginis (ante), but not so distinctly arranged
in rows, the row running from stigmal vein to apex of wing present, but
very poorly defined and the wedge-shaped hairless area behind the
marginal vein somewhat more restricted, there being three or four
coarse cilia basad of the stigmal vein in the apex of the wedge; hind
coxze about twice as long as thick with the dorso-posterior angle weakly
carinately margined; abdomen about as long as the thorax; first, second,
and third tergites emarginate medially, dorsally mostly polished; the
second and third at sides, and the fourth entirely, distinctly reticulate;
ovipositor exserted the length of the body; color of head and thorax above
dull coppery green; propodeum polished metallic green; pleura, coxz
and all femora metallic greenish-black or bluish in some lights; first
three tergites above steel-blue, sides of the abdomen and dorsum
beyond the third tergite brassy-green; antennal pedicel and flagellum

164 Annals Entomological Society of America [Vol. XII,

brownish-black, more or less metallic; scape, all tibize, and tarsi reddish
testaceous; ovipositor sheaths black; wings hyaline, venation pale
yellowish.

Male. Length 2 mm. Antennal pedicel a little shorter than the
first funicle joint; first funicle joint slightly broader than long and not
longer than the second; joints beyond the second very slightly diminish-
ing in length; sculpture of scutellum like that of female but the cross-
furrow not distinctly impressed; propodeum faintly rugulose with a
very weak median carina; abdomen a little shorter than the thorax, the
tergites not distinctly emarginate medially; scape dark above, reddish
testaceous beneath; all tibiz brownish-black; tarsi pale yellowish.
Otherwise like the female.

Type: locality, Tempe, Arizona. Type, Cat. No. 22298;
U.S. Nat. Mus.

Female type, male allotype and two male paratypes reared
by Mr. V. L. Wildermuth from galls of a Cecidomyiid on
Suaeda species and recorded under Tempe No. 2741.

Family PTEROMALID:.

Pseudocatolaccus americanus, new species.

This species is apparently very similar to Pseudocatolaccus as phon-
dylie Masi, Boll. Lab. Zool. Agr., Portici, 3, 1908, p. 189 (—Pteromalus
polyphagus Foerster, according to Kurdjumov, Rev. Russe d’Entom.
18, 1918, p. 7) differing principally from the description of that*species
in its smaller size, somewhat shorter malar space, slightly shorter
antennal joints and the more acute second and third teeth of the
mandibles.

Female. Length 2.56 mm. Head and thorax strongly reticulate-
punctate, the sculpture of mesonotum somewhat stronger than that
of the head. Head fully four times as broad as long, as viewed from
above; ocelli in a low triangle, postccellar line a little shorter than the
ocellocular line; as viewed from in front the head is distinctly broader
than long, antennz inserted above the lower extremity of eyes, malar
space shorter than the height of eye, clypeus irregularly striate, its
anterior margin sinuate; cheeks at base of mandibles distinctly concave;
mandibles fcour-toothed, the outer tooth acute, two median teeth
subacute, inner tooth blunt at apex; antennal scape cylindrical and not
reaching the anterior ocellus; pedicel conical and longer than thick;
three ring-joints strongly transverse, about equal in length but increasing
in breadth from first to third; funicle five-jointed, the first joint about
one-third longer than broad, following joints decreasing slightly in
length, the fifth subquadrate or very slightly broader than long; club
shghtly thicker than the funicle, three-jointed, and about twice as
long as the fifth funicle joint; mesoscutum much broader than long, the
parapsidal grooves weakly impressed on the anterior half; scutellum

1919] Gahan: Chalcid-Wasps 165

as long or a little longer than the mesoscutum; propodeum short, mostly
sculptureless with a weak median carina’and strong lateral folds, the
spiracles elliptical and close to the anterior margin of propodeum;
wings hyaline, the area before the submarginal vein with a few cilia
apically; behind the submarginal bare; ciliation of the rest of wing rather
sparse; stigmal and postmarginal veins subequal and each about four-
fifths the length of marginal; abdomen about as long as head and thorax,
conic-ovate and somewhat narrower than the thorax. Head and thorax
aeneous, scape pale testaceous; flagellum dark brown; mandibles brown-
ish; coxee concolorous with the thorax; all femora brownish-black; all
tibize brownish except at apex which is pale yellowish like the tarsi;
apical joint of all tarsi dark; abdomen metallic blue-green at base,
bronzy black beyond.

Male. Length 2 mm. Scape reaching the front ocellus, antennze
with two ring-joints and a six-jointed funicle, the funicle joints subequal
and slightly longer than broad; ocellocular and postocellar lines about
equal; abdomen not as long as the head and thorax; color of head and
thorax metallic blue-green; all femora dark brown; all tibiz and tarsi
pale yellowish, the apical tarsal joint dark; otherwise like the female.

Type locality, Tempe, Arizona. Type, Cat. No. 22299,
U. S. Nat. Mus.

Host, Asphondylia websteri Felt.

Described from eight females and one male reared by
V. L. Wildermuth from the alfalfa midge under Webster No.
7268, Tempe No. 2251. Also one female and one male from
the same locality and host reared by Mr. Wildermuth under
Tempe No. 2224.

Mouthparts and antenna of a female paratype mounted on
a slide (Hymenoptera Slide No. 783, U. S. Nat. Mus.) Other
type material mounted on card points.

This species, according to Mr. Wildermuth, is known to be
a true parasite of Asphondylia websteri Felt.

Trimeromicrus maculatus Gahan.

One specimen of this species, reared under Tempe No.
2223 is, according to Mr. Wildermuth, known to have been a
true secondary parasite of Asphondylia webstert, having been
dissected out of that host and then reared to maturity. Other
specimens of the species. were reared under Tempe No. 2264
from galls of Asphondylia webstert along with a number of other
hymenopterous parasites and including several specimens of
Bruchophagus funebris.

166 Annals Entomological Society of America |Vol. XII,

Trimeromicrus maculatus was originally recorded as a
parasite in alfalfa seed-pods infested with Bruchophagus funebris,
and subsequently proven by Mr. T. D. Urbahns to be a primary
parasite upon that host. The record under Tempe No. 2223
is interesting as showing that this species may develop upon
parasitic forms of Hymenoptera as well as upon the plant-
feeding Bruchophagus funebris. In the case of the record under
Tempe No. 2264 it is impossible to say whether the specimens
of this species were parasitic upon Bruchophagus, which was
shown to be present in the Asphondylia galls, or whether it
was actually present in its role of a secondary parasite of
Asphondylia.

Family EULOPHIDA.

Tetrastichus sobrius, new species.

Female. Length1.8mm. Face reticulately shagreened, with two
longitudinal rows of shallow punctures between the antenna and eye-
margin; antennal pedicel slightly shorter than the first funicle joint;
first funicle joint the longest, joint two very slightly longer than three,
joint three one and one-half times as long as broad and approximately
three-fifths the length of the first; club short, pointed ovate, obscurely
three-jointed, the first suture distinct but shallow, the second suture
subobsolete; thorax finely lineolated above, the mesoscutum with a
distinct complete median groove, scutellum without a median groove,
but with the dorso-lateral grooves complete and distinct; propodeum
very short, without any carinze and very faintly reticulately sculptured,
nearly smooth; mesopleura practically smooth, the prepectus distinctly
shagreened; basal joint of hind tarsi a little shorter or no longer than
the second joint; submarginal vein of the forewing with at least four or
five bristles above; abdomen about one and two-thirds times as long as
head and thorax, as broad as the thorax and pointed at apex, the tergites
very faintly sculptured. Head, thorax, abdomen, and all femora
brownish-black; flagellum brownish; scape, pedicel, all tibiz and tarsi
pale testaceous; an obscure transverse spot in front of the anterior
ocellus, also, pale; metanotum medially, more or less testaceous; wings
hyaline, the venation testaceous.

Male. Length 1.3mm. Scape of antenna a little thicker than the
pedicel and about twice as long; pedicel conical, a little longer than
thick; ring-joints very small, the number not discernible; funicle four-
jointed, the first joint as broad as long, somewhat shorter and broader
than the pedicel, second about one and two-thirds the length of the
first; third and fourth joints equal and each a little longer than the
second; club plainly three-jointed, about as long as the two preceding
funicle joints combined, the apical joint produced at apex into a sharp
point; each of the four funicle joints with a half-whorl of hairs above

1919] Gahan: Chalcid-Wasps 167

near the base, the hairs three to four times as long as the joint bearing
them; basal joint of club with a half-whorl of similar long hairs above
near middle of joint and another half-whorl beneath and a little more
basad; abdomen not longer than the head and thorax, narrower than
the thorax and not pointed at apex. Antenne entirely pale testaceous;
abdomen with a large pale, nearly white spot at base above. Other
characters as in the female.

Type locality, Tempe, Arizona. Type, Cat. No. 22300,
U.S. Nat. Mus.

Three specimens, one female and two males, reared by Mr.
Wildermuth from galls of the alfalfa gall midge, Asphondylia
webstert Felt, under Webster No. 7268, Tempe No. 2264. Also
two females and two males reared by E. G. Smyth from the
same source. Antenna of female paratype and male allotype
on a slide.

The host relations of this species are unknown to the writer.
It may be either a primary or secondary parasite of Asphondylia
or it may possibly have been parasitic upon Bruchophagus
funebris, specimens of which were reared from the same lot of
galls.

Tetrastichus bruchophagi Gahan.

Specimens of this species were received from Mr. Wildermuth
under Tempe No. 2264, having been reared together with several
other species of Hymenoptera, including Bruchophagus funebris,
from galls of Asphondylia webstert Felt, collected in the field
and placed in vials for rearing parasites. The species is known
to be a primary parasite of Bruchophagus funebris and may
have been present in the Asphondylia gall in this role. On the
other hand, it is not unlikely that this species, like 77vzmero-
micrus maculatus, may have developed as a secondary parasite
of Asphondylia using some one of the primary parasites of the
Cecidomyid as host.

Paragaleospomyia gallicola, new species.

It is very close to P. eja Girault, but differs in having the abdomen
more slender, and the hind tibiz black only on the apical half.

Female. Length 2 mm. Head, viewed from in front, slightly
broader than long, the ventral margin nearly straight, cheeks slightly
.convex in profile; antennal ‘depression deep and broad with a shallow
subquadrate extension below the base of antennz; ocelli in an obtuse
triangle, the postocellar line about twice the ocellocular; face and cheeks

168 Annals Entomological Society of America [Vol. XII,

finely and weakly reticulated, the frons and vertex with similar but
stronger sculpture; antennze separated at base by a distance about
equal to that between the base of antenna and the eye margin; scape
slender, flattened outwardly; pedicel three times as long as thick, about
equal in length to the first funicle joint and the ring-joints combined;
four distinct, transverse, ring-joints; funicle three-jointed, the first
joint the longest and about twice as long as thick; second joint distinctly,
though not a great deal longer than broad and a little longer than the
third; third joint subquadrate; club solid, and very slightly longer than
the two preceding funicle joints combined; pronotum and mesoscutum
strongly shagreened, parapsidal grooves deep, praescutum without
trace of a median groove; scutellum more weakly sculptured than the
mesoscutum, lineolate-reticulate with the paired dorsal grooves distinct,
but without a median groove; propodeum rather long, distinctly, though
not deeply, reticulate-punctate, slightly shining, with a distinct median
carina; mesepisternum and prepectus sculptured lke the mesoscutum;
mesepimeron mostly polished, its dorso-posterior margin reticulated;
hind coxz strongly reticulate-punctate; abdomen as long as the head
and thorax, slightly narrower than the thorax, pointed at apex, the first
and basal half of second tergites polished, the apical half of second and
the following tergites distinctly reticulate-punctate; second tergite a
little shorter than the first; third and fourth large, subequal, and each
about as long as the first. Black; flagellum brownish testaceous;
scape, anterior tibia, basal half and narrow apical portion of median
and hind tibiz testaceous; all tarsi pale yellowish, the apical joint black;
wings hyaline, venation testaceous.

Male. Length 1.25 mm. Antennal scape with a slight keel-like
expansion on apical half beneath; three transverse ring-joints; funicle
four-jointed, the first joint small and subquadrate or very slightly
broader than long, second joint one and one-half times as long as broad,
third very slightly longer than broad, the fourth quadrate; club solid,
nearly as long as the three preceding funicle joints and distinctly thicker
than the funicle; abdomen subpetiolate, about as long as the thorax, the
first tergite a little longer than the second, smooth; second and third
tergites sculptured, the third about twice as long as the second, tergites
beyond the third very short. Scape black, flagellum testaceous.
Otherwise like the female.

Type locality, Tempe, Arizona, , Type; Cat. No. 22301,
U.S. Nat. Mus.

Two females and six males reared by V. L. Wildermuth
from Cecidomyid stem-galls on Pluchea borealis and recorded
under Tempe No. 2742.

Galeopsomopsis transcarinatus, new species.

Very similar to G. multisulcatus Girault, but may be distinguished
by the fact that the first funicle joint is the longest of the funicle joints,
the true metanotum is smaller, not quite so strongly sculptured, with the

1919} Gahan: Chalcid-Wasps 169

median carina less strongly developed, the submarginal vein has four
to six erect hairs above instead of two, and the propodeum is slightly
different.

Female. Length 2mm. Antennal pedicel conical, a little shorter
than the first funicle joint; ring-joints transverse and apparently three
in number; first joint of funicle slightly longer than the second and
fully twice as long as thick, second a little longer than the third, the
latter ovoid and not one and one-half times as long as broad; club
not quite as long as two preceding joints, ovate, distinctly three-jointed
and terminating in a short spine, the joints subequal in length, but joints
two and three narrowing rapidly toward apex; antennal depression deep
and triangular; malar furrow basally represented by a deep rounded
fovea; ocelli large, the lateral ocellus separated from the eye margin
by distinctly less than the long diameter of an ocellus; a deep groove
runs anteriorly from each lateral ocellus, the two grooves converging in
front of the anterior ocellus; face laterad of the antennal depression and
the vertex strongly sculptured; mesoscutum and scutellum lineolately
sculptured; praescutum with a faintly indicated non-impressed median
line, and two or three rows of pimple-like rugosities along the margins
of parapsidal grooves; scutellum apparently with only an apical pair of
bristles; grooves of the scutellum not deeply impressed, the sublateral
pair with a few pimple-like rugosities similar to those of praescutum;
metanotum faintly sculptured with an indistinct median carina; pro-
podeum strongly sculptured all over, with a distinct median longitudinal
carina and a strong transverse carina, which is more or less interrupted
medially; the surface in front of the transverse carina shagreened;
behind the transverse carina also shagreened but divided by several
‘more or less distinct longitudinal carinze; marginal vein longer than the
submarginal, the stigmal about one-third the marginal and scarcely
at all thickened at apex; abdomen subsessile, conic-ovate, and strongly
shagreened above. Color metallic blue-green; wings hyaline; antenne,,
tegulz, wing veins and the legs for the most part pale testaceous; all
coxze concolorous with the thorax, all femora; except apically, blackish
brown.

Male. Length 1.6 mm. Scape very slightly expanded beneath
on apical two-thirds; apparently but one ring-joint, and this very
obscure; funicle five-jointed, the first funicle joint very slightly shorter
than the pedicel and a little longer than broad; second joint about one
and one-half times the first; third and fourth joints subequal and twice
as long as the first; fifth funicle joint longer than the first, but shorter
than the second; all funicle joints narrower at apex than base, and
bearing near base above a half-whorl of hairs approximately twice as
long as the joints; the fifth joint has in addition a half-whorl beneath
and nearer the base than the dorsal whorl which is located near the mid-
dle of segment; club two-jointed, slender, the joints subequal in length,
the apical one tapering to a sharp point at apex; the basal joint of club.
is provided with a whorl of long hairs which are not as long as those of
the funicle joints; ocelli large and narrowly separated from the eye

{ s
\
%

~~

170 Annals Entomological Society of America (Vol. XII,

margin; propodeum with the transverse carina weak, especially medially;
abdomen not longer than the thorax, parallel-sided or nearly, not de-
pressed above and much narrower than the thorax. Otherwise like
the female.

Type locality, DYempe, Anigora.| ype, Cat. Nei 22305;
U.S. Nat. Mus.

Host, Asphondylia websteri Felt.

Type, allotype and six paratypes reared by Mr. V. L.
Wildermuth from galls of the alfalfa midge and recorded under
Webster No. 7268, Tempe No. 2251. Also ten specimens with
the same data under Tempe No. 2264. All specimens on card
points; antenne from a female and a male paratype mounted
on a slide.

Whether the species is a primary or secondary parasite
of Asphondylia the writer is unable to state.

THE PROGRESS OF SCOLIA MANILZ ASHM.
IN HAWAII.

By F. Muir,
H.S. P. A. Experiment Station, Honolulu.

In a former article* the introduction of Scolia manile into,
and its establishment in, the Hawaiian Islands was described.
At that time the wasp was well established but‘ its host,
Anomala orientalis, was very abundant and doing great damage.

As an example of the difference between the conditions
then and now the following comparisons are of interest. In
1917 from one-twentieth of an acre, 3500 Anomala grubs were
collected from between the rows without digging around the
roots of the sugar cane. Now in the same locality but over a
larger area only four grubs could be found by following the
plow and searching. In another locality 104 eggs, 181 grubs,
7 pup and 10 adults were collected from around the roots of
one stool of sugar cane, now in the same locality one can only
find a stray specimen after considerable search.

The parasite has extended its range beyond the area infested
by Anomala ordentalis, and is now working on Adoretus tenut-
maculatus, a beetle common all over the archipelago.

The reduction of the Anomala is due solely to the action of
the Scolia as all other factors are the same as previous to the
introduction of the Scolia, when the Anomala was constantly
increasing and spreading.

The phenomenal reduction of A. orientalis has raised the
question as to the possibility of its eventual extinction. Should
the Scolia maintain itself upon the Adoretus but show a great
predilection for the Anomala such a thing may happen. The
final outcome is being watched with great interest.

*Ann. Ent. Soc. Am., X, p. 207 (1917).

17I

NOTICE OF PROPOSED AMENDMENT TO THE
CONSTITUTION OF THE ENTOMOLOGICAL
SOCIETY OF AMERICA.

By Wm. A. RILEY.

ARTICLE V—SEc. 3. The six additional members
of the Executive Committee shall be elected for a
term of three years, two members retiring each year.
The six members first elected under this rule shall
determine by lot their respective terms of service.

172

-NOTICE TO MEMBERS AND CONTRIBUTORS.

The Annals of the Entomological Society of America, pub-
lished by the Society quarterly, includes the Proceedings of the
Annual meetings and such papers as may be selected by the
Editorial Board.

Papers may be submitted to any member of the Editorial
Board and should be as nearly as possible in the form desired as
final, preferably typewritten, and illustrations must be finished
complete ready for reproduction. Plates must not exceed 5x7
inches unless intended tofold. In general, papers to be accepted
must be original, complete and previously unpublished and,
except in connection with the proceedings, it will not be the
policy to publish preliminary announcements or notes. Authors
will be allowed fifty reprints gratis and additional copies at cost
to the Society.

The Managing Editor is provided with the most recent
address of all paid-up members on record in the Secretary’s
office for mailing the numbers of the Annals and members failing
to receive their numbers should present their complaint to the
Managing Editor within four months from the date of the mail-
ing of the issue. After that time the numbers will be furnished
only at the regular published rate.

The regular annual subscription price for the ANNALS is in
the United States, Cuba, Porto Rico, Hawaii and Mexico, $3.00;
Canada, $3.50; other countries, $4.00. Checks, drafts or money
order should be drawn payable to ANNALS ENTOMOLOGICAL
SociETY OF AMERICA, and addressed to HERBERT OSBORN,
State University, Columbus, Ohio, U.S. A.

Requests for information as to membership and the annual
dues of members may be sent to the Secretary-Treasurer,
J. M. Atpricu, National Museum, Washington, D. C.

Communications relating to the ANNALS, and all orders for
separate copies or reprints should be addressed to
HERBERT OSBORN, Managing Editor,
ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA,
State University, Columbus, Ohio.

CONTENTS OF THIS NUMBER.

McInpdoo, N. E.—The Olfactory Sense of Lepid-

opterous Harve ricco Tes Oak lal Me 65
Dietz, W. G.—The Streptocera Group of the Dipterous
Genus Tipula: Lintceus. ool ie oe Cherie wee 85

CLAASSEN, P. W.—Life History and Biological Notes
on Chlaenius Impunctifrons Say. (Coleoptera,

Carabidae) oc widis cncin bw bracin piece alate aioe! leis 95
Hing, Jas. S.—Robberflies of the Genus Erax.....--- 103
GAHAN, A. B.—Some peaune ius Reared From

Wecidomyid: Galtsiy age Seon ae Sra tae 159
Muir, F.—The Progress of Scolia Manile Ashm. in

Hasta aE OR ap O'R GE ae CNC, Ce a laa 171

Price List of Publications.
Annals, Vols. I, II, III, IV, V, VI, VII, VIII, IX, X and XI complete, each. $3.00

Annals, Separate Parts except as below, each.........cceceecseccececeeces 1.00
Annals, Volsiiland EjsPart 3; Cael 6 cis, Ai eke oie eibarerecw cides eatem nels .50
Annals Volo iV 4 Part: 45 Cacia ioe iy Wiank crcsis elon siete mises eielshe Cpeceisiniemioleale 1.50

BACK VOLUMES

Of the ANNALS OF THE ENTOMOLOGICAL SocIETY OF AMERICA may
be secured from the office of the Managing Editor and new members
of the Society who may wish to complete a set are advised to secure
the earlier volumes while there is still a supply on hand and the price
is kept at the original subscription rate.

Address HerBERT OsBorn, Managing Editor,
ANNALS ENTOMOLOGICAL SOCIETY OF AMERICA,
State University, Columbus, Ohio.

}

A
i
*

*

BROS Chai cea eee Number. 3.

ANNALS

oF

‘ os The Entomological Society of Amenca

Ae

SEPTEMBER, 1919

, fe <
yc ee | EDITORIAL BOARD fe
es * \ ; (ie 7 A
HERBERT OSBORN, Managing Editor, \. 4
CoLUMBUS, OHIO, Sk?
_L. O. HOWARD, WM. A. RILEY,
WASHINGTON, D. C. MINNEAPOLIS, MINN.
VERNON L. KELLOGG, ; FRANK B, LUTZ,
i STANFORD UNIVERSITY, CALIF. NEw YorrE City, N. Y.
WM. M. WHEELER, WM. S. MARSHALL,
Boston, Mass. MADISON, WIs.
E. M. WALKER, J. G. SANDERS,
TORONTO, CANADA. HARRISBURG, Pa.

in .

PUBLISHED QUARTERLY BY THE SOCIETY
e COLUMBUS, OHIO

Entered as second class matter Apri! 11, 1908, at the Post Office at Columbus, Ohio,
under the Act of Congress of March 3, 1879.

The Entomological Society of America
Founded 1906.

s

OFFICERS 1919.

President : |
James G. NEEDHAM . Se iy i - Ithaca, N. Y.

First Vice-President
Justus W.Fotsom .... : : ° ° . Urbana, Ill.

Second Vice-President |
R. V. CHAMBERLIN . : : a hist Xe . Cambridge, Mass.

| Managing Editor Annals
HERBERT OSBORN. : TS Ghas ; . Columbus, Ohio

Secretary-Treasurer

J. M. AtpricH . .- ., National Museum, Washington, D.C.

Executive Commitiee

‘ THe OFFICERS

AND
. LUTZ, ARTHUR GIBSON,

F.E
G, A. DEAN, G. C. CRAMPTON.

Committee on Nomenclature
E. P. FEtt, T. D. A: CocKkERELL, NATHAN BANKs.

Thomas S ay’ Foundation

NATHAN BANKS, A. D. MacGItLitvray, PLP: CALVERT,
E. B. WILLIAMSON,
J. M. Arpricu, Editor, E. D. Batt, Treasurer.

Committee on National Museum

T. D. A. CocKERELL, Chairman,

HERBERT OSBORN, Wm. Barnes, Wa. M. WHEELER;
James G. NEEDHAM.

ANNALS

OF

The Entomological Society of America’

Volume XII SEPTEMBER, 1919 Number 3

THE ANTS OF THE GENUS METAPONE FOREL.*

By WiLL1AM Morton WHEELER.

The singular genus Metapone was established by Forel, in
1911}, for an ant from Ceylon, M. greeni, characterized by a
peculiar Ponerine habitus (resemblance to Cylindromyrmex and
Simopone), scrobed head, supposedly 11-jointed antennz in the
male as well as in the worker and female, one-jointed maxillary
and three-jointed labial palpi, terminally spinose or dentate
middle and hind tibiz and metatarsi and what he regarded as
an unusually slender, strongly segmented, non-tuberculate
larva, with long sparse hairs, and stout mandibles, and pupating
without spinning a cocoon. He says of this genus, which he
made the type of a new tribe, Metaponini: ‘‘I regard it as
constituting a special section, which I place provisionally
among the Ponerine and which I call Promyrmicine. Perhaps
later it will be necessary to transfer it to the Myrmicine.
I reserve my opinion in regard to this matter,”’ etc.

A year later, Emeryf examined M. greenz and its larva more
critically and found that alcoholic specimens of the latter when
properly softened and expanded had the usual shape of body,
head and mouthparts of the Myrmicine larva and were furnished
with long, serially arranged, hooked, dorsal hairs unlike any
known Ponerine larve, but like many larval Myrmicine. He
concluded that Metapone is a true Myrmicine ant and says: “It
seems to me that the comparison of Metapone with Cylindro-
myrmex and Simopone on the one hand and Sima on the other,

* Contribution from the Entomological Laboratory of the Bussey Institution,
Harvard University. No. 159.

{ Sur le genre Metapone n. g. nouveau groupe des Formicides et sur quelques

autres formes nouvelles. Rev. Suisse Zool. 19, 1911, pp. 445-451, 1 Pl.
} Etudes sur les Myrmicinae. Ann. Soc. Ent. Belg. 56, 1912, pp. 94-105, 5 Figs.

173

174 Annals Entomological Society of America [Vol. XII,

as formulated by Forel should be reversed, that Metapone
should be regarded as belonging to the subfamily Myrmicinz
and that the resemblance of the genus to Pseudomyrma and
Sima is due to genetic relationship and not merely to con-
vergent adaptation. I nevertheless accept Forel’s section
Promyrmicine but as a division of the Myrmicine and extend
its scope to include two tribes, the Metaponini and Pseudo-
myrmicini.’”’ Apparently Emery reached this conclusion as a
result of regarding Metapone as closely related to the African
Pachysima aethiops Smith and especially to P. latifrons Emery.

As Forel’s knowledge of the male and female of M. greeni
was restricted to a study of pupal specimens, he could give no
account of the venation of the anterior wings. In 1913,
however, he described a second species, M. sauter1, from a
female specimen taken in Formosa* and figured the fore wing,
which has an open radial cell and a single cubital cell.
This led him to incline to Emery’s view and to suggest a
resemblance: between Metapone and the Myrmicine genus
Liomyrmex.

In 1915 Forel described a third species of Metapone from
Queensland, M. mjébergit, and I have since described two
species, M. bakeri, from a female specimen taken in the Philip-
pinest and M. hewitt1, from male specimens taken in Borneo§.
The latter species showed that the male Metapone really has
12-jointed antennz and that Forel had evidently overlooked the
second funicular joint in the male of greenz. Still more recently
I have found two undescribed species, one from New South
Wales and one from Queensland, among material sent me
for study by the Museum of South Australia. Thus the
genus Metapone, as at present known, comprises seven species.

Concerning the habits of these ants very little has been
recorded. Mr. E. E. Green took the specimens of the type
species ‘‘from galleries in a decayed branch, which was also
infested by two species of termites.’’ And Forel adds: “It
lives, therefore, like Cylindromyrmex, in wood, with termites.

* H. Sauter’s Formosa-Ausbeute. Formicidae II. Arch. f. Naturg. 79, 1918,
pp. 183-202, 1 Fig.
{+ Results of Dr. E. Mjéberg’s Swedish Scientific Expeditions in Australia
1910-1913, Ark. f. Zool. 9, 1915, pp. 1-119, 3 Pls. and 6 text Figs.
Four New and Interesting Ants from the Mountains of Borneo and Luzon.
Proc. New Eng. Zool. Club 6, 1916, pp. 9-18, 4 Figs.
§ The Ants of Borneo. Bull. Mus. Comp. Zool. 1919, 63, pp 43-147

1919} Wheeler: The Genus Metapone 175

We cannot deduce from this either that it is or is not a termi-
tophagous species, but that it is seems very probable. At any
rate, it inhabits dead branches like many termites and numerous
tropical ants (Leptogenys mucronata, Pseudomyrma_ kiinckeli,
etc.) ”’

Although none of the specimens of Metapone, except those
of M. greent, was accompanied by notes on their habits, I deem
it very probable that all of the species form small colonies and
live in the dead twigs and branches of trees near or actually
in the galleries of termites, presumably of the genus Calotermes
and prey on these insects or their young. That this is the
case is indicated by the cylindrical shape of the body and the
fact that the workers of some of the species (greeni, mjébergi
and ftillyardi) are colored much like the termitophagous ants
of the genera Aéromyrma and Paedalgus, while the females of
others (sautert and Jee), in having the head and thorax black
and the gaster red, strikingly resemble the females of the
termitophagous genera Carebara and Ephebomyrmex.

“Returning to the question of the affinities of Metapone,
I may say that I am unable to accept Forel’s original view
and that I find Emery’s in part unacceptable. Forel’s view is
precisely stated in the two following quotations.

At page 448 he says: ‘‘The general form of the three sexes
(of Metapone greenz) is cylindrical and altogether recalls the
facies of the species of Cylindromyrmex. The analogy with
C. meinerti Forel is very striking, notwithstanding the clypeus,
which is absolutely different, and the postpetiole. The clypeus
has a certain resemblance to that of Szmopone in which it also
terminates behind with an arcuate suture, but much less trans-
verse and passing more posteriorly. In Simopone the frontal
carinz are also less separated. A certain distant resemblance
to the species of Sima and Pseudomyrma is probably due to the
convergence of an arboricolous life in cylindrical vegetable
cavities. The relationship with the Ponerine genus Cylindro-
myrmex (C. meinertt) is certainly real and not due solely to
the phenomena of convergence (mandibles, flat eyes placed
behind the scrobes, antenne, legs, petiole, sculpture, etc.)”’
And at page 452 he says: ‘‘In my opinion they (the Meta-
ponini) should: be placed directly between the Cylindro-
myrmicini and the Melissotarsini.’’ The Cylindromyrmicini

176 Annals Entomological Society of America [Vol. XII,

are now placed by Emery with the Cerapachyi among the
primitive Ponerine, whereas the Melissotarsus and Rhopalo-
mastix are now regarded by both Emery and Forel as Myrmi-
cine, and Forel would place them at the head of this subfamily
‘‘as being the most closely allied to the Ponerine.”’

I admit that Metapone closely resembles the Ponerine genus
Cylindromyrmex, especially in the cylindrical shape of the body
and head and the scrobes for the accommodation of the antenne.
Moreover, the habits of Cylindromyrmex and of the allied genus
Simopone seem to be much the same as those of Metapone.
According to Mayr*, Cylindromyrmex striatus Mayr (C. brast-
liensis Emery) was collected by Hetschko in Santa Catharina,
Brazil, ‘‘in wood in the galleries of a termite,’’ and Arnoldf
recently described Szimopone marleyi from three specimens
taken by Marley at Durban ‘in hollow stems of the. castor oil
plant.’’ A closer comparison of Metapone with Cylindromyrmex
and Simopone, however, shows that the resemblances are merely
superficial or convergently adaptive to this very peculiarity of
habitat. The long, cylindrical body in various genera of wood-
inhabiting ants is no more an, indication of genetic affinities
than is the similar shape so frequently and strikingly exhibited
by various families of wood-boring Coleoptera (Ipidz, Cleride,
Bostrichidea, Lymexylonide, some Erotylide, Trogositide,
Buprestide, Cerambycide, Lucanide, etc.) As long ago as
1891 Forel seems to have been deceived by. these superficial
characters when he was writing his original account of the
genus Simopone.t He there stated that ‘‘the resemblance of
the genera Cylindromyrmex and Simopone to the genus Sima
Roger, which belongs to the Myrmicides, is not a fact of con-
vergence or of mimicry, but seems to me to be due to real
affinities, notwithstanding the difference in the form of the
pedicel. The genus Szmopone, especially, with its very pro-
nounced abdominal constriction, seems almost to form a
transition to the Myrmicides, notwithstanding its Ponerid
sting and pygidium.”’ Here, again, the resemblance of Cylin-
dromyrmex and Simopone to Tetraponera (Sima auct.) is man-
ifestly due merely to similarity of habit, for the species of

* Sidamerikanische Formiciden. Verh. zool. bot. Ges. Wien 37, 1887, p. 546.

+ A Monograph of the Formicidae of South Africa I. Ponerinae and Dorylinz
Ann. S. Afr. Mus. 14, 1915, p. 21.

} In Grandidier’s Histoire Phys. Nat. Polit. Madagascar 20, 1891, p. 141.

1919] Wheeler: The Genus Metapone ik gr

Tetraponera and of the allied genus Pseudomyrma live in
cylindrical cavities or galleries in the branches or twigs of
plants.

While I am of Emery’s opinion that the species of Metapone
are true Myrmicine ants and show no particular relationship to
the Ponerine, I do not agree with Emery in accepting Forel’s
unfortunate term ‘‘Promyrmicine’’ or in associating the
Metaponini as the first tribe of the Myrmicine with the Pseudo-
myrmicini in a section under that name. It seems to me that
Emery is too much influenced by Forel’s prepossessions. There
is, in fact, little or nothing that is primitive or ancestral about
Metapone, but much that is highly specialized and secondary,
e. g., the shape of the antenne and especially the reduction of
the number of antennal joints in the male and female phases,
the peculiar reduction of the palpal joints, the simplified vena-
tion of the fore wings, the peculiar structure of the petiole,
postpetiole and legs, the vestigial condition of the eyes in the
worker, etc. The larva is not only purely Myrmicine, but
quite unlike that of Pseudomyrma, Tetraponera and Pachysima,
as may be seen by comparison of Emery’s figure (here repro-
duced as Fig. 2) with those in one of my recent papers.* Such
study as I have been able to make of four species of the genus
Metapone convinces me that it is an abberrant and highly
specialized, though probably ancient genus of Myrmicinae,
neither primitive nor ancestral, without special affinities to the
tribe Cylindromyrmicini or other Ponerinze and moreover not
even closely related to the tribes Peudomyrmicini. It should, of
course, constitute an independent tribe, Metaponini, as Forel
and Emery maintain, but its position among the other tribes
of the Myrmicine is not easily determined. It might be placed
provisionally between Emery’s Melissotarsini and his Stereo-
myrmicini, which have 11-jointed antennz in both male and
female phases.

The following key will help in separating the six Metapone
species of which the worker or female is known. WM. hewittz,
known only from the male, is redescribed below.

* A Study of Some Ant Larvae, with a Consideration of the Origin and Meaning
of the Social Habit Among Insects. Proc. Amer. Phil. Soc. 57, 1918, pp. 295-348,
12 Figs.

178 Annals Entomological Society of America [Vol. XII,

i)

Key To METAPONE SPECIES.

Clypeus with a narrow, projecting, rectangular anterior lobe, transversely |
truncated in front, concave at the sides, with acute tooth-like corners. . . .2.

Clypeus searcely projecting anteriorly, without distinct lobe, bluntly or

obscurely; \bidentate sya. Sheps: soci ee ea © atcha eee ee esomeree 5.
Petiole broader than long, its posterior corners blunt, flattened and lobular;
postpetiole transversely elliptical or sub-elliptical; color brown.......... 3.

Petiole much longer than broad, its posterior corners produced as somewhat
diverging teeth which are longer than broad. at their bases; postpetiole
subrectangular; color black, with red or brown gaster..............01...-. 4

Mandibles 4-toothed; mesoépinotal suture very distinct and impressed; pos-
terior corners of epinotum not swollen; sculpture of head and thorax rather
coarse; brownish red, with castaneous gaster, female darker; length 5.9-8.9
PONE e Pins easils itah kB SEN Ge eat pia en orcad aR Reta erectus ee greent Forel.

Mandibles 5-toothed; mesoépinotal suture indistinct; posterior corners of
epinotum swollen; sculpture of head and thorax finer; pale brown, head

darker: “length a7. GO) Prince. mor ete ecu mae era eee ee mjobergt Forel.
Body not greatly flattened; length 10.5-11.1 mm.; petiole and postpetiole
longitudinally striate; black, with dark brown gaster....... sautert Forel.

Body much flattened; length 6.5-7 mm.; petiole transversely striate, post-
petiole smooth; apical mandibular teeth smaller than the basal; brown
black withiterrcuginous redigasten a). .-t5 an ace ieee ie le@ sp. nov.

Mandibles 5-toothed; posterior clypeal suture distinct; head and thorax longi-
tudinally striate; petiole broadly and feebly excised behind; legs short,
femora very broad; head and thorax castaneous; gaster and legs ferrugineous;
lenigith, ‘sb SOe nama? jee AG nome an cite eisiers ben oir oe eae Gem neOe tullyardt sp. nov.

Mandibles 4-toothed; posterior clypeal suture obsolete; body very smooth and
shining; petiole longer, deeply and angularly excised behind; legs longer,
with narrower femora; black, with dark reddish brown legs; length 6.4
PYVELD ee Pa zate ths yee fe pistes pees aie fe Rese ook iS Oe, HO tere bakeri Wheeler.

1919] Wheeler: The Genus Metapone 179:

1. Metapone greeni Forel.

Forel, Rev. Suisse Zool. 19, 1911, p. 449, Pl. 14, 8 9 o and larva; Emery, Ann.
Soc. Ent. Belg. 56, 1912, p. 95, Fig. 1, larva.

(Figs. 1 and 2.)

Worker. Length 5.9-8.9 mm.

Head subrectangular, nearly one and one-half times as long as
broad, with subparallel sides and moderately concave posterior border.
Eyes small, elongate, in the largest individuals with about 15 ommatidia
in their longest and about 10 ommatidia in their shortest diameter.

Pesan: \

AY.

Si
&
O

C.

Fig. 1. Metapone greeni Forel (after Forel). a, worker; b, head of same from above;
c, antenna of same; d, scutellum of male from above.

There are three ocellar pits, but no ocelli in the largest workers. Man-
dibles 4-toothed, with large basal lobe. Clypeus large, with subrec-
tangular anterior lobe. Scape slightly surpassing the middle transverse
diameter of the head, flattened and apically much dilated; funiculus
much dilated and flattened at the tip, joints 2-6 at least three times as
broad as long, 8 and 9 decidedly broader than long, the apical joint
longer than broad and much longer than 8 and 9 together. Thoracic
dorsum oblong, scarcely broader in front than behind, somewhat
flattened above, submarginate on the sides. Promesonotal suture very
indistinct, mesoépinotal suture distinct and impressed. Pro- and
mesonotum subequal, each one and one-half times as broad as long,

180 Annals Entomological Society of America [Vol. XII,

together one and one-fourth times as long as broad; base of epinotum
rather longer than broad, passing through an abrupt curve into the
short, subvertical declivity. Petiole subcuboidal, rather flat above,
widened behind, a little broader than long, its posterior border above
deeply and broadly excised, its anterior and posterior surfaces truncate,
the dorsal somewhat thicker than the ventral portion. Anteriorly
it has a short peduncle and its ventral surface has in front a large,
longitudinal rectangular, translucent lobe, followed by a strong tooth.
Postpetiole broader and shorter than the petiole, one and three-fourths
times as broad as long, rounded cuboidal, bearing on its ventral side a
large tooth, followed by a welt. Gaster oval; sting long and stout.
Femora greatly dilated, the posterior pairs scarcely one-third longer
than broad; tibia and metatarsi short and broad.

Fig. 2. Metapone greent Forel. Larva (after Emery). a, lateral view; b, head of
same; md, mandible; mx, maxilla.

Shining; head and thorax regularly and not coarsely longitudinally
striated, with scattered, effaced punctures; the remainder of the body
smooth, with fine, scattered, piligerous punctures.

Erect hairs yellowish red, short, very sparse on the body, almost.
lacking on the thorax, more abundant on the tibiz and _ scapes.
Pubescence reddish, sparse, very distinct on the gaster and pedicel,
- but scarce elsewhere.

Deep brownish red; gaster deep chestnut brown, sometimes blackish,
with red tip. Small workers sometimes paler (tmmature?).

Female (mature pupa). Length 9.5 mm.

Eyes very large, occupying nearly one-half the sides of the head,
moderately convex. Mesonotum and scutellum flat, small, together
forming scarcely half the thoracic dorsum. In other respects like the
worker, but with the head a little longer. Dark brown, nearly black.

1919} Wheeler: The Genus Metapone 181

Male (mature pupa). Length 7 mm.

Mandibles small, tridentate. Clypeus very large, convex in front,
flattened behind. Head slightly longer than broad; eyes occupying
more than half its sides. Frontal carine parallel, as far apart as
they are from the lateral borders of the head. Front truncated anteriorly
and separated from the preocellar space by a transverse carina con-
necting the frontal carinz. Antennal scrobes small, marginate behind
and laterally. Antennal scape very short, scarcely longer than broad;
first funicular joint very small, globular, broader than long, the remain-
ing joints cylindrical, the more basal a little broader than long, but
increasing gradually in length, the last twice as long as broad. Thorax
as broad as the head, rather flat. Mesonotum with Mayrian furrows;
scutellum behind with two short, flat, horizontal teeth. Epinotum
subcuboidal, its base longer than its declivity. Petiole and_ post-
petiole as in the worker, but the former much broader than long, more
rounded and not emarginate behind, with a small anteroventral tooth,
the postpetiole unarmed below. Gaster elongate. Legs of the ordinary
form, not dilated.

Head and thorax opaque or subopaque; longitudinally rugose, with
reticulate interrugal spaces; sides of thorax striated. Clypeus coarsely
and transversely rugose. Pedicel and anterior half of gaster densely
reticulate-punctate and opaque or subopaque; pesterior half of gaster
becoming more shining. Postpetiole longitudinally rugose above and
coarsely rugose on the sides like the posterior portion of the epinotum.

Erect hairs very oblique on the appendages, nearly appressed, almost
absent and very short on the scapes. Pubescence somewhat more
abundant than in the worker.

Black; margins of gastric segments, genital valves, tarsi, tibie and
scapes sordid yellow or reddish; funiculi brown.

Type-locality: Peradenyia, Ceylon (E. E. Green).

2. Metapone mjoebergi Forel.
ForelpArior fe Zool 9 etOlla. po0N.o: Ol

Worker. Length 7.6 mm.

Somewhat smaller than greeni. Mandibles 5-toothed, their outer
margin feebly convex and their basal lobe smaller. Anterior lobe of
clypeus much shorter than in greeni and sauteri, but also rather
rectangular. Head rectangular, about one-third longer than broad,
not broader behind than at the base of the frontal carinz, only anteriorly
somewhat narrower, with nearly straight sides and feebly concave
posterior border (in greeni broader behind and more convex laterally,
longer in sauteri). Scape and funiculus more slender than in greeni,
about as in sauteri. The very small, perfectly flat eyes have only a few
indistinct ommatidia and lie as in greeni below the hind end of the
scrobes, but are much smaller. Whole head in profile appearing as
if very obliquely truncated anteriorly, 1. e., with the anterior half of the
clypeus much more abruptly declivous than in greeni and sauteri so

182 Annals Entomological Society of America [Vol, XII,

that the clypeus does not seem to stand away from the mandibles.
Thorax very feebly convex, two and one-fourth times as long as broad,
With parallel sides. Anterior corners of pronotum obtuse; posterior
corners of the concavely emarginate epinotum protruding and swollen,
a condition not seen in greeni. Promesonotal suture absent, mesoépinotal
suture feebly indicated (deeply impressed in greeni). Petiole much
broader and shorter than in greent, as broad as the epinotum, not quite
twice as broad as long, its anterior border feebly, its posterior border
deeply concave, narrower in front than behind, where its angles extend
out as flat, lobular projections. It is truncated anteriorly and posteriorly
and even somewhat concave in front, and bears ventrally only one
strong longitudinal lobe. Postpetiole not broader than the posterior
border of the petiole, scarcely one and one-half times broader than long,
rounded on all sides, only about half as broad as the gaster, with two
teeth below. Legs of the same shape and quite as broad as in green,
the tibia shorter and stouter than in sautert.

Sculpture of body, especially of the head, somewhat weaker and
more finely longitudinally striate than in greeni; petiole and gaster
distinctly more densely punctate. Mandibles nearly smooth, feebly
striate and sparsely punctate.

The gaster and thorax exhibit a very short, fine, rather abundant,
erect pilosity, which partly passes over into the pubescence and is
sparser on the posterior portion of the head. There are only a few long
hairs.

Pale brown, head dark brown; appendages brownish yellow.

Female (dealated). Length 7.9 mm.

Searcely distinguishable from the worker, only much darker, brown-
ish black, and with more distinct sutures outlining the mesonotum and
scutellum. Hind corners of epinotum less protuberant, eyes large,
ocelli present. Postpetiole somewhat broader than in the worker.

Type-locality: Malanda, Queensland (E. Mjoberg).

3. Metapone sauteri Forel.
Forel, Arch. f. Naturg. 79, 1918, p. 189, Fig. @.

Female. Length 10.5-11.1 mm.

Head rectangular, at least one and one-half times as long as broad,
with straight sides and feebly concave posterior border, Eyes much as
in greeni. Mandibles much longer and broader, with straight 5-6-toothed
terminal border and longer and broader basal lobe. Clypeus with
more distinct posterior suture, its anterior rectangular lobe much longer
than in greeni, with perfectly straight anterior border, concave sides and
acute corners. Frontal groove distinct. Antennz very similar to
those of greeni, but somewhat longer and narrower, especially the scape.
Thorax decidedly narrower than the head, but otherwise as in greent, the
epinotum, -however, much longer, nearly one and one-half times as long
as broad, more strongly convex and less strongly marginate on the
sides. Petiole also longer, much longer than broad, posteriorly with

1919} Wheeler: The Genus Metapone 183

two longer and more slender teeth, which are longer than broad at their
bases. Postpetiole more rectangular than in greeni, nearly as long as
broad. Gaster much broader, nearly twice as broad as the postpetiole,
first segment nearly three times as long as the postpetiole. Lobes and
teeth on the ventral surface of the petiole and postpetiole precisely as in
greeni, also the teeth on the posterior tibia and metatarsi, but the
femora, especially the anterior pair, twice as long as broad. Wings
with a single cubital cell, a discoidal and a tong radial cell, the latter
slightly open at the tip; radial and cubital veins separating at the tip
of the cubital cell.

Longitudinal striation of the head and thorax much as in greeni,
but somewhat coarser, the head less punctate. Petiole and postpetiole
not only punctate as in greeni, but also sharply longitudinally striate;
the striz on the petiole arcuate behind. Gaster shining, sparsely
punctate and feebly and finely coriaceous.

Hairs sparse, reddish yellow, both erect and appressed on the
body; appressed hairs on the head and thorax very scarce.

Black; antenne and gaster dark reddish brown; tibiz and tarsi
reddish; femora and coxe brownish. Wings brownish, with brown
veins and pterostigma.

Type-locality: Sokutsu, Banshoryo District, Formosa (Hans
Sauter).

4. Metapone lez sp. nov.
(Figs. 3 and 4.)

Female. Length 6.5-7 mm.

Head one and one-half times as long as broad, distinctly narrower
at the anterior corners than behind, the sides feebly convex and the
posterior border nearly straight; the upper surface very flat, in profile
about two and one-half times as long as high. Mandibles convex,
their basal lobes large, their terminal borders with a series of five teeth,
smallest apically and largest basally. Clypeus flat, with a long pro-
jecting median lobe, which has a straight, transverse anterior border,
concave sides and acute corners. Eyes rather large, but evidently
smaller than in greeni and sauteri. Ocelli small, near the occipital
border. There is a longitudinal, impressed line, bordered on each
side by a ruga, representing the frontal groove, extending from the
posterior border of the clypeus to the anterior ocellus and growing
deeper posteriorly. Antenne rather stout, flattened; scape elliptical,
somewhat more than twice as long as broad; funiculus with 3-jointed
club, first joint as long as broad, joints 2-7 transverse, broader than
long, terminal joint about as long as the two preceding joints together.
Thorax elongate, subhexagonal, fully two and one-half times longer
than broad, very flat above, four times as long as high, broader through
the wing insertions where it is as broad as the head. Humeri of pro-
notum bluntly angular; mesonotum slightly concave behind, as broad
as long, with the scutellum on the same level as the base of the epi-
notum, which is longer than broad, narrower behind than in front,

184 Annals Entomological Society of America [Vol. XII,

with straight sides and broadly and feebly excised posterior border.
In profile the declivity is vertical and only about one-third as long as
the base into which it passes through an abrupt curve. Petiole shaped
like a mammalian vertebra, seen from above only half as broad as the
epinotum at its base, one and one-third times as long as broad, with
convex rounded anterior and slightly concave lateral and posterior
borders, the posterior corners produced as two long blunt, diverging
teeth and the middle of the posterior border above elevated as a low

Fig. 3. Metapone lee sp. nov. Female from above, with wings removed.

angular}tubercle. In profile the anterior and posterior surfaces are
truncated, the anterior and lateral surfaces concave, the ventral surface
with two small angular, widely separated projections. Postpetiole
from above nearly square, a little broader than long, a lttle broader
than the petiole through its posterior teeth, feebly convex above,
sharply truncated anteriorly, ventrally with a stout angular projection
followed by a prominent transverse welt. Gaster narrow, twice as
broad as the postpetiole, two and one-half times as long as broad,
flattened dorsoventrally, the first segment as long as broad. Legs
short, shaped much as in greeni, with apically dentate posterior tibiz

1919] © Wheeler: The Genus Metapone 185

and metatarsi. Wings short (5 mm. long), with the same venation as
in bakeri, the radial cell being slightly open at its tip and the radial and
cubital veins fused for a short distance beyond the tip of the cubital
cell.

Shining; mandibles smooth, finely and sparsely punctate. Clypeus,
head and thorax longitudinally striate; the striz on the clypeus fine, on
the upper surface of the head separated by smooth areas, especially
on the sides of the front. On the sides of the head, however, and in the
scrobal cavities the striz are very sharp and regular, and curve around
the antennal insertions. Neck transversely reticulate-rugulose. Rugz

ov,

Tram)

S

SS

a

Fig. 4. Metapone lee sp.nov. Female, lateral view.

on the pro- and mesonotum and scutellum sharp and regular, inter-
spersed with a few punctures, rugee on the base of the epinotum very
fine, diverging behind, the surface also with coarse scattered punctures.
Sides of thorax obliquely and finely striate. Petiole very finely and
densely transversely striate above and also with a few coarse, scattered
punctures. Postpetiole and gaster smooth and shining, with very fine,
sparse, piligerous punctures. Legs shining.

Hairs yellowish, short, sparse, bristly, erect or suberect, of uneven
length, longest on the postpetiole and gaster; pubescence coarse, very
sparse on the thorax, more distinct on the gaster. Hairs on the legs
short and sparse, more or less oblique.

186 Annals Entomological Society of America |Vol. XII,

Brownish black; mandibles, antenne, clypeus and anterior lobes of
frontal carine deep red; gaster and legs paler, ferruginous red; sides of
thorax and lower surface of petiole and postpetiole somewhat castaneous.
Wings brownish hyaline, with black pterostigma and brown veins.

Described from 22 specimens taken by Mr. A. M. Lea on
Mt. Tambourine, Queensland. (Museum of South Australia).

This species is most closely related to sauter1, but is much
smaller and is very distinct in the extremely flat, narrow body,
differently sculptured petiole and postpetiole, paler gaster, etc.

Sq”
[oN
lo
(e@ VBEIN
( )
>=
\
\
| \
\.
NS .
Pe 54

\<

a
Fig. 5. Metapone bakeri Wheeler. Female.
a, lateral view; b, head, dorsal view; c, petiole and postpetiole.

.

5. Metapone bakeri Wheeler.

Wheeler, Proc. New Eng. Zool. Club 6, 1916, p. 10, Fig. 1. @.
(Figab)

Female. Length 6.4 mm.

Head less than one and one-fourth times as long as broad. Mandi-
bles short and convex, 4-toothed, with small basal lobe. _Clypeus
convex, not marked off by a suture behind, its median lobe short and
broad, with a blunt, tooth-like projection on each side. Antennz
with very distinct 3-jointed club, the two basal joints of which together
equal the terminal joint. Thorax narrower than the head, nearly
three times as long as high, its sides submarginate above; pronotum with
subangular humeri, mesonotum feebly convex, together with the scutel-

1919] Wheeler: The Genus Metapone 187

lum broadly elliptical, longer than broad. Petiole from above a little
longer than broad, broader behind than in front, marginate on all sides,
with straight anterior and lateral borders, the posterior border deeply
and somewhat angularly excised; in profile anvil-shaped, fully one and
one-half times as high as long, its ventral surface with two large blunt
angular projections. Postpetiole transversely elliptical, nearly twice
as broad as long, ventrally with a blunt projection in front and a small
tooth behind. Gaster small, suboblong. Legs short, but the femora
much narrower than in the other species. Wings rather short, the
radial cell narrowly open at the tip, the radial and cubital veins fused
for a short distance beyond the apex of the cubital cell.

Extremely smooth and shining throughout, with sparse, very incon-
spicuous, piligerous punctures on the dorsal surface; posterior portion
of antennal scrobes densely, longitudinally striate.

Hairs whitish, short, sparse, erect, nearly lacking on the pleure,
most conspicuous on the gaster and legs.

Black; mandibles tinged with red; antenne, fore legs, tibia, tarsi,
tips and bases of middle and hind tibiz reddish castaneous. Wings
feebly infuscated, especially along the anteroapical margin; veins
resin-colored; pterostigma dark brown.

Type-locality: Mt. Banahao, Luzon Island, Philippines
(Brok, ©. Fr. Baker).

This species differs from all the others in its highly polished,
non-striate head and thorax, dark color, more slender legs,
smaller size, etc. In my original description and figure the
radial cell of the fore wing is represented as closed at the tip.
It is really narrowly open. I have corrected the error in the
figure as here reproduced.

6. Metapone tillyardi sp. nov.
(Fig. 6.)

Worker. Length 5.5-6 mm.

Head subrectangular, about oné and one-fourth times as long as
broad, a little broader behind than in front, with straight sides and very
feebly concave posterior border. Eyes very small and indistinct,
flat, at the lower posterior border of the scrobes. Clypeus rectangular,
evenly convex anteroposteriorly, continuing the feeble convexity of
the head, a little narrower in front than behind, with straight sides,
separated behind from the head by a distinct suture, its anterior border
scarcely produced as a lobe, bluntly bidentate in the middle. Each side
of the clypeus overarches a cavity confluent with the scrobe and becomes
continuous with the frontal carina which as in the other species extends
to about the posterior fourth of the head. There is a distinct impressed
line representing the frontal groove and extending to the posterior fifth
of the head. Mandibles stout, their external borders rather straight,
the apical border oblique, with five teeth, those at the apex largest,

188: Annals Entomological Society of America [Vol. XII,

the basal lobe small. Antenne short, flattened, the scape elliptical,
scarcely more than twice as long as broad; funiculus with distinct
3-jointed club, first jomt longer than broad, joints 2—7 small, transverse,
terminal joint a little longer than the two preceding joints together.
Thorax regularly oblong, two and one-half times as long as broad, a
little narrower than the head, with straight parallel sides, blunt anterior
corners and concave posterior border, representing the boundary between
the base and declivity of the epinotum which is square. The dorsal
surface is flattened, bluntly submarginate on the sides, the pleure

Fig. 6. Metapone tillyardi sp. nov. Worker. a, dorsal; 6, lateral view.

concave. There is an extremely faint trace of a promesonotal suture;
mesoépinotal suture distinct, rather sharply impressed, especially on
the sides. In profile the thorax is about two and one-half times as
‘long as high, vertically truncated anteriorly and posteriorly, the latter
truncation being the epinotal declivity, the sides of which are marginate
but not swollen above. Petiole from above as broad as the epinotum
and twice as broad as long, narrower in front than behind, with straight
anterior and lateral borders and broadly excised posterior border, the
posterior corners flattened and lobe-like; in profile the segment is trun-
cated anteriorly and posteriorly, with the anterior and lateral surfaces
strongly concave, the ventral surface bearing a single stout, triangular

1919] Wheeler: The Genus Metapone 189

projection. Postpetiole a little narrower than the petiole, transversely
elliptical, twice as broad as long and more than half as broad as the
first gastric segment, ventrally with two blunt transverse projections.
Gaster short, one and two-thirds as long as broad, first segment some-
what shorter than broad. Legs very short, flattened; femora very
broad, elliptical, tips of middle and hind tibiz and metatarsi dentate
as in the other species.

Shining; mandibles striato-punctate, basally smoother and sparsely
punctate. Clypeus, head and thorax regularly and rather finely
longitudinally striate; the posterior fourth of the head smooth and very
shining. Interspersed with the strie on the head are a few coarse
punctures, arranged in rows, but becoming sparse on the occiput.
The striz on the pro- and mesonotum are somewhat finer and more
regular than those on the head, on the base of the epinotum they are
still finer and merge posteriorly into lines of punctures diverging to
the posterior corners and into scattered punctures on the declivity.
Sides of thorax obliquely striate. Petiole, post-petiole and gaster
punctate, the gaster more finely. Legs smooth and shining, with very
fine, sparse punctures.

Hairs yellow, in the form of sparse pubescence on the head and
thorax, becoming conspicuously longer, more abundant and suberect
on the gaster and especially on the sides of the petiole and postpetiole.
There are also a few short erect hairs on the head and thorax. Legs
with very short, rather bristly hairs, oblique on the tibia.

Rich castaneous brown; mandibular teeth and lateral borders of
clypeus and anterior border of head black; antenne a little paler brown;
gaster and legs ferrugineous.

Described from 10 workers taken at Dorrigo, New South
Wales (Museum of South Australia).

This species, which I dedicate to my friend, the eminent
Australian entomologist, Dr. R. J. Tillyard, resembles bakeri
most closely in the structure of the head and clypeus, but in
color and sculpture it is more like greent and mjébergi, though
sufficiently distinct from either.

7. Metapone hewitti Wheeler.
Wheeler, Bull. Mus. Comp. Zool. 1919, 63 p. 620”.
(Fig. 7.)

Male. Length 6-7 mm.

Body long and slender. Head as broad as long, evenly convex and
rounded behind, without posterior corners; cheeks very short; eyes.
moderately large, but not very convex; ocelli rather small. Mandibles.
small, but well developed, their external borders slightly sinuate basally,
convex at the tips, apical and basal borders distinct, subequal, the
former with four subequal teeth. Clypeus large, convex, somewhat
broader than long, slightly depressed or flattened posteriorly. Front

ee

199 Annals Entomological Society of America [Vol. XII,

truncated anteriorly, with a transverse crest or carina separating it
from the preocellar space and connecting the frontal carinze, which are
prominent, nearly straight, subparallel and as far apart as they are
from the lateral borders of the head. Posteriorly each carina curves
forward medially of the eye as a distinct ridge and terminates opposite
its anterior end, thus enclosing a small, shallow, elliptical scrobe.about
the base of the antenna. Antenne 12-jointed; scape very smqall,
about twice as long as broad; first funicular joint also very small,
broader than long, subglobular; second joint considerably larger but fis
broader than long, the remaining nine joints stout, cylindrical,, dis-
tinctly longer than broad, increasing somewhat in length distally,
terminal joint nearly as long as the two preceding joints together, with
tapering tip. Thorax long, narrower than the head through the eyes.

Fig. 7. Metapone hewitt Wheeler. a, male in profile; b, head, dorsal view; c, fore-
wing; d, scutellum, dorsal view; e, antenna.

Pronotum well-developed, truncated in front; mesonotum and scutellum
somewhat flattened above, the former with distinct Mayrian furrows,
the latter on each side near its posterior border with a peculiar blunt,
spatulate spine, slightly curved inward at the tip. Epinotum longer
than broad; subrectangular from above, its base horizontal and twice
as long as the vertical declivity into which it passes through an abrupt
curve, the sides of the base and of the declivity above coarsely and
rather irregularly marginate. Petiole with a short, stout peduncle
anteriorly, and a thick cuboidal node, which is a little longer than broad
and slightly higher in front than behind, with truncated anterior and
posterior and rounded dorsal and lateral surfaces. Seen in profile its ventral
amargin is slightly bisinuate, with a small triangular tooth at the anterior
end of the peduncle. Postpetiole distinctly broader than the petiole
and .broader than long, transversely elliptical, in profile truncated
anteriorly, convex and rounded above, its ventral border unarmed,
nearly straight. Gaster elongate elliptical, with straight anterior

‘ '

1919} Wheeler: The Genus Metapone 191

border and tapering tip. Genitalia completely retracted; cerci appar-
ently absent; pygidium and hypopygium short and pointed. Legs
short, of the usual simple form, without the tibial and metatarsal teeth
or spines of the worker and female; spurs of the middle and hind tibiz
simple, blunt at the tip. Tarsal claws very small, strongly curved,
nonpectinated. Wings very short (4.5 mm.), with a well-developed
discoidal cell, a single cubital cell and the radial cell slightly open at
the tip. Pterostigma large and conspicuous.

Subopaque; mandibles opaque, longitudinally rugose and very finely
punctate. Head reticulate-rugose, the clypeus more coarsely, though
not transversely asin greent. Front behind its anterior truncation with
regular longitudinal rugee converging to the anterior ocellus. Antennal
scrobes less distinctly longitudinally rugose. Upper surface of meso-
notum and scutellum and sides of thorax sharply and regularly longi-
tudinally rugose, on the mesonotum with elongate, shallow foveolze
in the narrow interrugal spaces. Dorsal surface of epinotum, including
the upper portion of the declivity with extremely coarse reticulate ruge,
some of which are clearly transverse. Petiole above less coarsely and
even more irregularly rugose. Postpetiole and gaster very finely and
densely punctate, with superimposed small, sparse and very regular
piligerous punctures.

Hairs grayish brown, rather abundant, erect on the head, thorax
and petiole, mostly subappressed or oblique on the postpetiole, gaster
and legs. Antennal funiculi with very short fine hairs, or pubescence.
Wings minutely hairy.

Black; mandibles, antenne, legs and tip of gaster reddish brown, the
tarsi slightly paler. Wings grayish hyaline with slightly infuscated
tips and anterior margin; veins sharply defined, brown; pterostigma
dark brown.

Type-locality: Kuching, Borneo (John Hewitt).

The four males from which this species was described were
taken by Mr. Hewitt in 1908 and though they have been in my
collection ever since, I was unable to assign them to any genus
till Forel published his description and figures of Metapone
greent. ‘Though closely related to the male of this species,
hewittt differs nevertheless in having 4-toothed mandibles, a
different sculpture, especially on the clypeus, and very different
spines on the scutellum. The antenne are certainly 12-jointed
as shown in the accompanying figure. Forel evidently over-
looked the second funicular joint in the mature pupal males of
greent which he examined. There is a possibility, of course,
that hewittt may be the male of sauterz or bakeri, though it
seems to be too small to be the male of the former and. too
opaque to belong to the latter species.

NOTES ON THE MALE GENITAL SYSTEM IN CERTAIN
LEPIDOPTERA.*

By HERBERT RuckeEs, A. M.

INTRODUCTION.

Changes going on in organisms, whether studied grossly
or minutely, have always been interesting subjects and many
times astonishing truths have been revealed, where they were
least expected to occur.

A great deal of the work carried on in entomological
histology, especially under the heading of metamorphosis, has
been relative to the muscular system, alimentary tract, adipose
tissue, etc., while little attention has been given to other equally
important systems. This is true of the reproductive system;
so for this reason a study of the morphological and histological
metamorphosis of the male genital tract in certain Lepidoptera
was attempted.

The study was worked out under the direction of Professor
W. A. Riley, to whom the writer is indebted for his prolonged
patience and valuable criticism, and for the use of prepared
slides. The writer also wishes to thank Professor Lester W.
Sharp for his advices regarding the subject of spermatogenesis
and for the use of his slides of maturation mitosis in insects.
To Mr. John R. Eyer thanks are also due for use of his valuable
slides of the genitalia of a large number of Lepidoptera.

The Lepidoptera were chosen to be worked on, for the meta-
morphosis could be readily controlled, by artificial means; and
besides, this group shows apparently a greater diversity in the
form of the genital system than most other orders. The
Saturniide were chosen because when the study was first
begun, they were most abundant.

Studies in the morphology and internal anatomy of insects
date back to the earliest zoologists. Dissection of forms,
however, did not properly commence till the advent of the
microscope, so that we find the first worker on the genital
system to be Malpighi (1669) whose treatise on the internal

* Contribution from the Entomological Laboratory of Cornell University.
This paper is part of a thesis (by title, On the Metamorphosis of the Genital

System in the Saturniidae) presented as part requirement for the degree of Master
of Arts.

192

1919] Ruckes: Male Genital System in Lepidoptera 193

structure of Bombyx mori Linn. is now a classic. From that
date onward we find little concerning the genital system till,
in 1815, Herold produced his paper on the metamorphosis of
Pieris brassice Linn., including other systems besides the
reproductive. This paper formed the basis for subsequent
work on the genital tract for it discussed the peculiar migration
of the testes to a median plane and, there, complete fusion into
a spherical body. It has been proved that this condition is
by no means typical of all Lepidoptera, being representative
of only the higher Rhophalocera. From the time of Herold’s
work till 1865 seems to be a more or less stagnant period for
work of this sort. In 1865 Bessels published a few observations.
From the morphological point of view the greatest contributions
have been given by Chlodkovsky (1880-1913). Though short,
these papers are thorough and explicit, explaining the evolution
of the Lepidopterous genital tracts and their genealogical
relationship to those of other orders. Since 1884, histological
and cytological methods, combined with a continued increase
of our knowledge of chromosomes and chromosome reduction,
have led to the production of a vast amount of work and a large
number of observations on all groups of insects. A great deal
of this work has been restricted to the study of the maturation
of sex cells, regardless of the structure and function of the
surrounding tissues. Those doing the most comprehensive
general studies since 1884 are LaValette St. George, Verson,
Blatter,, Berlese, Toyama and at a recent date (1913) Zick.
The views and criticisms of these authors will be discussed as
their particular points come up.

METHODS AND MATERIAL.

As above stated the Saturniide furnished the basis of the
following study, though they were by no means the only forms
observed. The following is a list of species that were used:

LEPIDOPTERA. ORTHOPTERA.
Philosamia cynthia Drury. Dissoteira carolina Linn.
Samia cecropia Linn.

Callosamia promethia Drury. HEMIPTERA.
Thelea polyphemus Cramer. Anasa tristis DeGeer.
Pieris (Pontia) rape Linn. Euschistus fissilis Uhler.
Anosia plexippus Linn. E. variolarius Stal.

Vanessa antiopa Linn. Nezara viridula Stal.

194! “Annals Entomological Society of America [Vol. XIE)!

By far the greatest part of the work was done on P. cynthia}:
since it was most readily available. \ S. cecropia offered the best ’
material to follow out morphological changes—being the largest’
form. | Material was procured at Ithaca and New York City: : \

For various ends, various technique was used. The material ’
for ‘spermatogenesis was killed and fixed in Benda’s fluid,’
Zenkers, Gilson’ S’ Or « Herman’s mixtures. Pereyni’s reagent’
was! found to ‘be very good for synaptic stages of mitosis. A’
new mixture, formulated as follows, fixed ‘“‘ring”’ stages better!
than any of the more common solutions: 0

220 copper sulphate... scene 3 parts

POO, platinum ehlornd sao eee 2 parts

Acetic Feels Dura AS Cena a Se ee at on .... 6-10 drops to every 100 c. ¢.
Water (distilled oto) G82 Scere 10 parts

This solution is ae a nuclear fixer. For the study "
the tissues other than the germ cells, Gilson’s or Herman’s
mixtures were employed.

Of the stains, Weigert’s copper haemotoxylin was found
best, this staining cytoplasm with as much exactness as the
nucleoplasm. Mallory’s connective tissue stain was indis-
pensable for the muscular and fibrous coats of the various
organs, especially the coats of the testis. Delafield’s haemo-
toxylin counterstained with picro-fuchsin was found to be very
good for determining the presence of a cuticula covering.

In making dissections, especially during the larval and early
pupal periods, when thé organs of the genital system are very
delicate and very difficult to see, the preparations were lightly’
stained with Delafield’s haemotoxylin, the stain being dropped
on the dissection after the water had been poured out of the
pan. In preparations made in this way, the finer organs like
the vasa deferentia, ligaments, and: nerves, could be followed
along their course. These structures stain deeply, while the
fat lobules do not. :

In order to make a more detailed study of the nerves invest-
ing the caudal portion of the abdomen, specimens were injected
with 1% Grutbler’s ‘“‘B-X”’ methylene blue in a .9% salt solution ‘
and allowed to remain so treated, in a living condition for from :
twelve to twenty-four hours. .The earlier stages were found to.
take up the stain more readily than the adults. In adults,
thirty-six hours were required to have this intra vitam method
take effect. See

1919] Ruckes: Male Genital System in Lepidoptera 195

PARTS OF THE ADULT MALE GENITAL SYSTEM.

In insects the male reproductive system consists of a pair of
testes, a vas deferens, leading from each testis into a set of
ducts that vary greatly in different groups of insects (Cf.
Berlese Gli Insettt, Figure 1087), and a penis. The testes
themselves are composed of testicular tubes. These tubes may
be numerous or few; in the Lepidoptera there are four; the
tubes also may be separated (Hepialus, Fig. 1A) or they may be
fused (Philosamia, Fig. 1. B and E, Fig. 2), and in some
cases the two testes may approach the median plane and fuse .
Pieris, Fig. 1 D). In all cases the testicular tubes are homo-
logues of the ovarian tubes. In the Saturniide the testes lie
in a latero-dorsal position, close to the alimentary canal, just
under the fifth and sixth segments of the abdomen. They are
reniform in shape, the convex side of each, called the apical region,
being mesad; the concave region known as the hilum lies ectad,
and is the junction of the vas deferens and the four testicular
tubes (Fig. 3). The vasa deferentia are long irregular tubes,
extending caudad to the eighth segment, where each passes
through the loop-hole of the tenth abdominal nerve (Fig. 4, Vd).
After passing through the nerve each turns cephalad to the
seventh segment where it unites with the seminal vesicle.
The seminal vesicles we may consider as distended portions of
the vasa deferentia, each joined anteriorly to a long convoluted
tube known as the accessory gland; posteriorly they unite to
form the upper end of the ejaculatory duct: The ejaculatory
duct is really in two parts; the upper end which in early pupal
stages may readily be seen to consist of two approximated
short tubes (prolongations of the seminal vesicles) is known in
the adult as the ductus ejaculatorius duplex; the posterior and
longer portion, which is an invagination of the ectoderm (it
bears a cuticula or chitinous lining), is known as the ductus
ejaculatorius simplex, and terminates in a very muscular “‘bulb,”’
the bulbus ejaculatorius at the base of the penis. This last named
organ is the caudal extremity of the entire genital system. It is
wholly chitinous, bearing various types of prolongations, and
is connected with the body wall by means of muscles. It is
capable of extrusion during copulation. (The adult system is
figured in Fig. 1E).

196 Annals Entomological Society of America |Vol. XII,

Of the parts of the reproductive system, the germ cells, the
ductus ejaculatorius simplex, and the penis are formed from the
ectoderm. The remaining organs, and the muscular covering
of the ductus ejaculatorius simplex are derived from the
mesoderm.

ECTODERM . MESODERM
Germ cells Testicular coats
({Verson’s Cell) ? (Verson’s Cell, or Follicula repithelium)?
Epithelium of ejaculatory duct simplex | Accessory glands
Penis Vasa deferentia

““Ejaculatory duct duplex’”’
Muscles of all the above.
Muscles of the ejaculatory duct simplex

The origin of the various organs, and their constituent
tissues has been a problem of great importance in insect
embryology. It is not the purpose of this paper to deal with
the various views set forth by embryologists, but to confine the
remarks to post-embryonic development.

THE NERVES.

The nerves innervating the reproductive organs are com-
plicated and deserve special mention. Their development and
changes occurring in them during the pupal and adult periods
are subjects that well deserve detailed study. Some interesting
features are as follows.

In the early pupal stage (Fig. 4), the seventh abdominal
ganglion is closely approximated to the eighth plus tenth ganglia,
near the caudal portion of the seventh abdominal sternite.
Three pairs of nerves arise from the seventh ganglion; these are
the seventh anterior, going to the seventh spiracles, the seventh
posterior going to the body tissue, and the seventh splanchnic
or sympathetic, also innervating the seventh spiracles. The
posterior nerve is somewhat reduced (vestigial); this is fre-
quently the case in Lepidoptera.

The caudal ganglion, which apparently is a fusion or con-
densation of the eighth, ninth and tenth ganglia, gives rise to
three or four pairs of nerves. The most cephalic of these is the
eighth sympathetic, which innervates the vestigial eighth
spiracle. The next nerve has two branches, a long heavy one
(undoubtedly the eighth anterior) passes to the eighth spiracle,

1919] Ruckes: Male Genital System in Lepidoptera 197

while the shorter and weaker branch (the eighth posterior)
extends into the body tissue. In Samia cecropia, the ninth
nerve arises separately from the ganglion. In Philosamia
cynthia, it is joined for a distance with the tenth nerve. In
either case, this nerve has two branches, a long anterior branch,
which extends caudad to the juncture of the seminal vesicles,
-and the vasa deferentia, innervating this juncture, and forming
a prominent plexus there. This may be called the vesicular
plexus (Vp. Fig. 4). The shorter branch is homologous with
the shorter branches of the seventh and eighth nerves and is no
doubt the ninth posterior. The tenth nerve is very heavy and
long. Itis the most interesting of all; the strong anterior branch
extends caudad, to the region of the vesicular plexus, where it
forms a prominent loop hole, through which the vas deferens
passes; this nerve continues and gives rise to delicate branches
that form what may be called the caudal plexus (Cp. Fig. 4),
innervating the tissue that later becomes part of the genital
appendages. From each loop hole a delicate branch extends
entad, and unites with the vesicular plexus. The tenth posterior
nerve passes caudad, and supplies the same tissue that the
caudal plexus innervates.

What changes take place during the transformation stages
have not been studied. The caudal plexus no doubt forms the
nerve supply for the genitalia, and the vesicular plexus probably
controls the muscles of the ducts.

HISTOLOGY, OF THE TESTIS:

As previously stated, each testis consists of four chambers, or
testicular tubes, combined, and surrounded by a capsule.
Regardless of the form of a testis, from the histological aspect,
one may consider it as consisting of two main parts, an ecto-
dermal part of the sex cells themselves, and a mesodermal portion
comprising the follicular epithelium and the various coats that
make up the wall of the testis. A study of the sex cells involves
observations on their origin, growth, maturation and trans-
formation, topics that come under the heading of sperma-
togenesis and which are not to be treated in this paper. (Cf.
Pauline Dederer, Biol. Bull., Vol. 13—Spermatogenesis in P.
cynthia). The author wishes to deal solely with the mesodermal
tissue. This may readily be considered under two headings,

198 Annals Entomological Society of America [|Vol. XII,

the follicular epithelium and the testicular envelopes. It is
simpler to treat, first, the envelopes, and then consider
the epithelium, since the latter needs some explanation.

In studying the testicular chambers, one finds that their
lining consists of a thin structureless membrane, which is the
basement membrane (Bm.—Figs. 5,6, 7), of the follicular
epithelium; the distribution and function of the follicular
epithelium will be explained later. Outside of the basement
membrane is a prominent wall, which in the Lepidoptera, in
general, consists of two heavy layers, an inner testicular tube
coat (Ttc.—Fig. 5, 6, 7) and an outer capsular coat (Cc.—Fig.
5, 6, 7). Besides these on the periphery of the capsular coat
is a thin layer of material that is apparently chitinous in nature
and is probably due to the degeneration of trachea, which are
very abundant on the testis and ramify freely over its surface.

The Testicular Tube Coat (Figs. 5, 6, 7). Of the two layers
forming the wall of the testis, the inner one, or testicular tube
coat, is the more interesting. The coat not only surrounds the
entire testis but sends three partitions or septz toward the
hilum, forming the four chambers or testicular tubes. The cells
composing this layer are very peculiar. They are of meso-
dermal origin, and assume the appearance of true connective
tissue. They render the coat loose in its texture and fibrous
in its nature. In most cases (in Lepidoptera) the cells are
separated from one another, leaving large intercellular spaces
in which delicate cyloplasmic fibres, from the periphery of the
cells; ramify and anastomose. These two features give the
coat its loose and fibrous appearance. The shape of the cells
varies according to the species of insect studied. In P. cynthia,
they are elongated and spindle shaped, in C. promethia, they are
angular, less elongated, closer and have fewer cytoplasmic
prolongations. The cytoplasm in all cases is drawn out and
fibrous in nature. What is more interesting than the position
of the cells as to shape and texture, is their content. They are
apparently storage tissues, either for reserve materials or.
for products of katabolism. In the case of the Saturnids they
have a prominent fat content (Fd.—Fig. 5, 6). The fat
globules show very plainly and beautifully in osmic acid
preparations. When stained with Mallory’s C. T. stain, they
stand out as brilliant orange or orange-red droplets in the
pinkish cytoplasm. In Pieris, Lycaena, and species in which

1919]... Ruckes: Male Genital System in Lepidoptera 199

the testes are colored, this coat is the repository for the pigment
granules (Pg.—Fig. 7), which are believed to be waste products
of urate character. Fine purplish and brown granules can be
readily observes in small bits of this coat tern. from the testis
of Pieris larva and viewed in toto.

*Through. the interstices of this tissue, trachez having
passed through the capsular coat, branch and. ramify. These
tracheze also pass through the basement membrane into the
chambers of the testis, there to resolve themselves into tracheoles
which lie in close relationship with the groups of spermatocysts.
This condition is to be expected since the sex cells are rapidly
growing bodies, and consequently need relatively large amounts
of oxygen for metabolism. .Choldkovsky (1880), stated that
the tracheze never penetrate the basement membrane, and enter
the follicular chambers. Koschevnikoff (1891) disproved this
statement for Apis mellifica, and the present writer has found
that each testicular chamber has small masses of tracheoles in
it, which can be traced back to the tracheze passing through the
basement membrane, usually the membrane lying aPELONUARE
to the septe.

The Capsular Coat. Less interesting than the testicular
tube coat is the capsular coat, surrounding the entire testis,
and in some cases easily separated from its inner neighbor.
Usually about the same in thickness as the inner coat, this outer
layer of cells shows a marked contrast, in view of the nature of
the cytoplasm and the compactness of the tissue. The layer
stains poorly, but on close examination, it may be seen that
the cytoplasm is very spongy and granular. The cells are
usually cuboidal, and very closely placed. They may be only
one layer of cells, as in C. promethia and Pieris rapae. The
nuclei are usually spherical and centrally placed. There is
no pigment or fat distributed in the outer coat.

As mentioned above, over the periphery of the capsular coat
lies a layer of structureless material that has all evidence of
being chitin or of chitinous origin. It is flakey, tough and
readily stains with iron haemotoxylin, picro-fuchsin, and aniline
blue. It originates probably through the degeneration of the
tracheze which are abundant over the surface of the testis.
In breaking down, the chitin composing the tenidia becomes
chemically changed, assumes a plastic consistency, and flows

200 Annals Entomological Society of America [Vol. XII,

together to make a homogeneous cover over the entire reproduc-
tive gland. This layer may readily be called the ¢tracheal
membrane (Tm., Figs. 5, 6, 7).

The Follicular Epithelium. So much for the coats of the
testis. Each of the four tubes in each testis, is the male
homologue of each ovarian tube. Therefore there must be a
follicular epithelium. Previously it was stated that the lining
of each chamber consisted of a delicate basement membrane.
The basement membrane in all epithelia is on the ental periphery
of the cells. Hence, the follicular epithelium must be some-
where inside of the follicular chamber. Where is it and what
is its nature? That is the question that has puzzled a great
number of workers on this subject.

Verson (1889) described a peculiar cell occurring in the
apical region of each testicular chamber of certain insects.
Other authors (La Valette St. George, Spichardt, Tischomiroff)
have studied this cell (called Verson’s cell, -Spichardt’s cell,
la cellule geante, the apical cell, etc.), and have given various
interpretations as to its structure and function.

Balbiani (1866) is probably the first to have noticed this
peculiar structure; he considered it a mother cell for the sperma-
togonia. Spichardt (1886) regarded the element in question
as the primordal germ cell, forming nuclei at its periphery.

Verson, in studying Bombyx mori, explained that this peculiar
gigantic cell, produced protoplasmic extensions, fibrous in
nature, that formed a complete meshwork throughout the
testicular chamber. In the meshes of this net, the sperma-
togonia are at first scattered, then they become collected, and
finally a spermatocyst or a group of germ cells surrounded by a
capsule, is formed.

Verson believed, however, that the giant cell was a primordal
spermatogonium. Cholodkovsky (1894) upheld the view
of Verson.

Toyama (1894), in studying Bombyx mori, believed that
there occurred, in the apical portion of each chamber, an
invagination of the testicular wall, carrying with it an enlarged
and specialized cell, which later lost its connection with the
coat of the testis. This became the stellate cell, described
by Verson. It had no genetic relationship with the sperma-
togonia.

1919] | Ruckes: Male Genital System in Lepidoptera 201

LaValette St. George (1897) believed that Verson’s cell was
simply a transformed spermatogonium (hence, a sister and
not a mother-cell to primary germ cells), giving forth pro-
longations that became the spermatocyst capsules.

Tichomiroff (1898) probably added the most to the studies
of the apical cell. He definitely proved it to give rise to the
capsules of the spermatocysts; it had no genetic relationship
with the sex cells, it gave off nuclei to the capsules, and was in
nature connective tissue.

Truly, to try to understand the significance of Verson’s
cells, from simply a study of the testes of the Lepidoptera,
would be very difficult and confusing. By making comparisons
with the structures of the testes of other insects, the true
homology of the apical cell may be understood.

In most insects there is a thin, delicate layer of cells lining
~ the testicular chamber, this layer, it has definitely been shown,
gives rise to the spermatocyst capsule, by sending into the
interior of the chambers, delicate protoplasmic prolongations
that envelop groups of spermatogonia. This layer, frequently
called the cystogenous tissue, is nothing more or less than the
follicular epithelium of the testis, and comparable in all respects
with the follicular epithelium of the ovarian. tubes. In the
Lepidoptera the state of affairs is so modified as to present an
apparent absence of follicular spithelium. Comparative anatomy
shows, however, that the capsular tissue in one case arises from
a definite epithelium, while in the other it arises from a single
enlarged apical cell. Homologizing, then, Verson’s cell (Fig. 7),
must be a localization and reduction of the follicular epithelium,
playing the same role, 1.e., that of nutrition, as does the epithelium
inthe ovary. Due to this special disposition of the epithelium
it has lost its characteristic form and assumed a condition
somewhat fibrous and similar to connective tissue.

Most investigators have missed the significance of Verson’s
cell, because they did not study it in stages early enough.
It no doubt arises from the mesoderm, as does the follicular
epithelium in the female, and only during embryonic develop-
ment can its disposition be thoroughly understood.

In the larval stages, after all the spermatogomia have been
gathered into spermatocysts, the apical cell disintegrates, for:
it is no longer needed. The capsule of each spermatocyst can
readily offer all the nutriment that is needed by the growing
germ cells (Fig. 8).

202 Annals Entomological Society of America [|Vol. XII,

THE HISTOLOGY OF THE GENITAL TUBES.

A. VAS DEFERENS.

From each testis the vas deferens extends caudad to the
seminal vesicles. In larval and early pupal stages these
efferent ducts are straightish, delicate, almost invisible tubes.
They look like very fine transluscent threads leading to the
little papilla in the eighth segment, that later becomes the
remaining portion of the genital tract. During the larval and
early pupal stages, the vas deferens is uniform in diameter and
possesses a prominent lumen, which is regular (i. e., without
ridges) initscontour. During these stages the cellular structure
is represented by Figure 9. The cells are short cylindrical or
columnar, possess a rather uniformly staining cytoplasm,
and show a prominent central nucleus. The chromatin of
these cells occurs in irregular masses, staining prominently.
These masses are rather large. The outer covering of the
efferent duct may be seen as a fibrous coat that later becomes a
layer of circular muscle.

As the vas deferens grows in length, it becomes convoluted;
coincident with its growth in length, there is a corresponding
increase in. diameter. Cell division takes place very rapidly,
during the mid-pupal period. In Figures 10, 11 and 12 we
have represented the changes that occur during this time. At
first the cells simply grow in length (Fig. 8), becoming tall-
columnar; the nuclei are still very prominent and the chromatin
remains in irregular masses. The cytoplasm begins to become
somewhat less dense near the tip of the cells, and begins to show
a vacuolated appearance. At first it was thought that this
condition was an artifact due to improper fixation. Its con-
stancy led to the abandonment of the idea. The outer coat
becomes heavier.

A little later stage will show the conditions represented in
Figure 11. Marked changes are beginning. Some of the cells
remain stunted or are suppressed in their development. Their
neighbors continue to elongate and we have at first a slight
irregularity in the contour of the lumen. Parallel with this
change, the cytoplasm becomes definitely vacuolate at the tips
of the cells, and a change takes place.in the nucleus. The
chromatin begins to break up into smaller particles and these

1919] Ruckes: Male Genital System in Lepidoptera 203

have a tendency to round off. Whether this change has
anything to do with the secretory activity of the cells the
author is not prepared to say. The activity of the nucleolus
was not studied. At any rate, there is a very evident change in
the character of the chromatin.

Continuing a step further, we get a condition represented by
Figure 12. The rapidly elongating cells crowd over one
another, and we have the formation of a little nest-like mass of
cells. The function of the nidus in the epithelium of the
digestive tract is to replace cells, as they are cast off into the
lumen. As far as was determined in the vas deferens, there is
no migration of the cells into the lumen, and therefore these
groups of cells should be called pseudonidi. The vacuolization
continues, and the chromatin now appears as large granules.
The nuclei remain round or oval and centrally placed.

There is a gradual, but very prominent growth of the cells
in the vas deferens from now on till we reach the end of the
pupal period when several changes take place rather rapidly.
The pseudonidi become very prominent, and the contour of
the lumen becomes markedly rigid, as figured in Fig. 16. The
muscle layer has been formed and outside of it is the peritoneal
membrane. The most striking feature of this stage is the
structure of the epithelial cells (Fig. 12). They have become
totally vacuolate, and throughout the cytoplasm occur promi-
nent, deep staining particles that are evidently secretion
droplets; where these particles come from has not been
determined. The nuclei migrate to the periphery of the cell
and become quite large, but the chromatin appears in very fine
granules, producing a condition that does not differentiate the
nucelus very well from the rest of the cell. The nuclei assume
the shape of the cell. Between the muscle layer and the cells
is a prominent basement membrane.

The condition of the cells at the time of the emergence of the
imago may be represented by Fig. 14. The number of secretion
granules is reduced, but we have in the lumen of the duct, a
whitish liquid containing also an abundance of sperms. This
is the seminal fluid and has been produced by the cells of vas
deferens. The lower portions of the cells show a peculiar
fibrous condition of the cytoplasm. This is very difficult
to illustrate. The tips still show prominent vacuoles.

2()4 Annals Entomological Society of America [Vol. XII,

After copulation, all the secretion droplets have passed from
the cytoplasm and this becomes still more fibrous in its nature.
The vacuoles persist at the tips. (Fig. 15).

The above described metamorphosis takes place in the
upper two-fifths of the efferent duct. The lower two-fifths
is somewhat similar, though the lumen remains somewhat
larger, and the pseudonidi are more prominent. The middle
fifth of the tube remains as it was in its larval, or early pupal
condition, that of a thin, uniform duct, with cells cuboidal and
without pseudonidi. Secretion takes place in the upper portion
and in the lower portion of the vas deferens, but not in the
middle part.

B. THE SEMINAL VESICLES.

These organs may be considered as simply distensions of the
vasa deferentia; their epithelia have the same growth tendency,
i. e., to form pseudonidi. However, the development of the
pseudonidi is not carried as far as in the case of the efferent
ducts. At the cephalic and caudal ends of the vesicles, the
epithelium loses its irregular character and assumes a strict
columnar condition, where it graduates at one end into the
epithelium of the accessory glands and at the other into that of
the ejaculatory duct. The vesicles are chambers for the
retention of the semen and are not centers of secretion. This
latter process is localized in the vasa deferentia.

Nussbaum (1882) originally considered the vesicles, as well
as the accessory glands, ejaculatory duct and penis, as
derivatives of the ectoderm. Others, who also have made
extensive studies of the reproductive system, believe the
vesicles to be of mesodermal origin. At any rate, they are so
similar to the vasa deferentia, and sufficiently unlike the other
organs, to be called distentions of the former, and therefore
of a mesodermal nature.

Kochevnikow (1891) found in the seminal vesicles of the
honey bee, two layers of muscle, an inner circular one and an
outer longitudinal. He could find none on the vasa deferentia.
In the Saturniide, there is one layer of muscle, this is circular,
and is a continuation of that found on the vas deferens (Fig. 17).

1919] Ruckes: Male Genital System in Lepidoptera 205

Cy thy ACCESSORY GEANDS.

In the accessory glands, the cells do not become con-
spicuously glandular, as might be expected from the terminology
of these organs.

The cells remain columnar in character and do not form
pseudonidi. They do not become vacuolate, in the same sense
that the cells of the vas deferens become vacuolate. The
cytoplasm presents a very spongy appearance, that is present
in very young cells of the glands. At first this condition might
easily be taken for a general vacuolization. Comparisons with
other types of cells show that it is quite different, however.
Many authors believe that there is a secretion of these organs,
and that the secretion causes the formation of spermatophores.
This may be true. Secretion is not as evident as in the cells of
the glandular part of the vas deferens, and proceeds in a differ-
ent manner.

The accessory glands differ from the organs previously
discussed, in the possession of a single longitudinal layer of
muscle. This in early stages of the development of the repro-
ductive system may be continuous with the muscle of the upper
end of the ejaculatory duct. This has not been definitely
determined.

Koschevnikow (1891) found three layers of muscle in the
glands of Apis; an inner longitudinal layer which may not
extend the entire length of the tubes, a central circular and an
outer longitudinal.

In some insects the accessory glands may actually be
ectodermal in origin. Many authors do not believe so, however.
If they were ectodermal, there would be a tendency toward the
formation of an intima, which, as far as has been determined,
does not exist in them.

D. THE EJACULATORY DUCT.

As stated previously, the ejaculatory duct consists of two
parts, at least it is so considered by several authors, prominent
among whom is Schroeder. The upper part is divided into two
tubes, the ductus ejaculatorius duplex (of Schroeder), which
are short and connect at their cephalic end with the seminal
vesicles. These tubes are probably simply caudal prolonga-

206 Annals Entomological Society of America |Vol. XII,

tions of the seminal vesicles. Their epithelium is somewhat
similar to that of the vesicles and their musculature is the same.

Passing caudad to the union of this so-called ductus ejacula-
torius duplex, we find we have a change in the musculature,
which now takes on a longitudinal course. (Fig. 19). For
quite a distance the epithelium retains its uniform columnar
appearance. The longitudinal muscle is characteristic of the
ductus ejaculatorius simplex, which is the caudal portion
leading to the base of the penis. Undoubtedly this duct is
ectodermal in origin. It is an invagination of the hypodermis
and bears, for more than one-fifth of its length, a prominent
intima. Above the caudal fifth the chitinous lining becomes
thin and indistinct and 1s wanting at the juncture of the ductus
ejaculatorius simplex and ‘“‘duplex.”’

The longitudinal muscle, characteristic of the upper end
of the single tube, has added to it, a prominent layer of circular
muscle about half way along the duct (Fig. 20). The
circular muscle appears outside of the longitudinal. Toward
the caudal extremity the musculature becomes very uneven and
very prominent, till about the so-called “‘bulbus ejaculatorius,”’
it is very heavy, and interwoven. A third, outer longitudinal
layer is added (Fig. 21). Koschevnikow could not find
muscle in the ejaculatory duct of Apis mellifica. This seems
very strange, in view of the function of this organ.

The epithelium of the ejaculatory duct goes through a
metamorphosis comparable with that of the vasa deferentia,
1. e., it forms pseudonidi, but these are covered over with the
intima. In the so-called “ejaculatory bulb,” the epithelium is
actually folded, assuming the appearance of the epithelium of
the proventriculus.

What by some authors is called the ejaculatory bulb is in
reality a continuation of the simplex duct, with abundant
muscle, with no enlargement, but a reduction of its lumen.
Its function is to force the seminal fluid through the chitinous
penis.

CONCLUSION.

(1) The Saturniid testis belongs to Chlodkovsky’s second
(larval) type, consisting of four testicular tubes combined in a
common capsule.

(2) The greatest changes take place during the pupal stage.

1919] Ruckes: Male Genital System in Lepidoptera 207

(3) The inner coat, of the testis and not the outer one, is
fatty in nature.

(4) Verson’s cell is a modified follicular epithelium, giving
rise to the spermatocyst capsules.

(5) The cells of the vas deferens are the source of the
seminal fluid, the accessory glands and seminal vesicles are
storehouses.

(6) The accessory glands have a longitudinal layer of
muscle; the seminal vesicles a circular layer, the upper end of
the Saculaten, duct has longitudinal muscles, the lower portion
has added a circular and an outer longitudinal layer as well.

(7) There is a definite and complicated innervation of the
internal genital organs.

BIBLIOGRAPHY.

1. Berlese, A. 1909. Gli Insetti. Milan.

2. Bessels, E. 1857. Studien tber die Entwicklung der Sexualdrtisen bei den
Lepidopteren. Zeit. f. Wiss. Zool. Vol. XVII.

3. Blatter, P. 1892. Sur l’histologie des organes annexes de l'appareil male chez
la Periplaneta orientalis. Comptes Rendus CXV. 1332-1334.

4. Blatter, P. 1897. Etude sur la structure histologique des glandes annexes de
l'appareil male de l’hydrophile. Arch. Anat. Micro. Vol. I. 384-910.

5. Bordas, L. 1900. Recherches sur les organes reproducteurs males des Cole-
opteres. Ann. Sc. Nat. Zool. Vol. 2. 283-448.

6. Cholodkovsky, N. N. 1880. Uber die Hoden der Schmetterlinge. Zool. Anz.
Vol. TIT. 115-117, 214-915:

7. Cholodkovsky, N. N. 1884. Uber die Hoden der Lepidopteren. Zool. Anz.
Vol. VII. 564-568.

8. Cholodkovsky, N. N. 1892. Zur Kenntnis der Mannerlichen Geschlecht-
sorgane der Dipteren. Zool. Anz. XV. 178-180.

9. Cholodkovsky, N. N. 1905. Uber den Bau des Dipterenhodens. Zeit. f, wiss.
Zool. Vol. XXXII. 389-410.

10. Cholodkovsky, N. N. 1913. Zur Kenntniss der Trichopteren und Lepidopteren
Hodens. Zool. Anz. Vol. XLII.

11. Demokidoff, K. 1902. Zur Kenntniss des Baues des Insectenhodens. Zool.
Anz. Vol. XXV. 575-578.

12. Gruenberg, K. 1903. Untersuchung tber die Keim und Nahrzellen in der
Hoden und Ovarien der Lepidopteren. Zeit. f. wiss. Zool. Vol. LXXIV.
327-3895.

13. Hasse, E. 1887. Die Vorfahren der Insecten. Abhand. Gesell. Isis. Dresden.
Vol. XI.

14. Herold, M. J. D. 1815. Entwicklungsgeschichte der Schmetterlinge, anat-
omisch u. physiologisch bearbeitet. Cassel. w. Marburg.

15. Henneguy, F. M. 1904. Les Insectes. Paris.

16. Kellogg, V. L. 1907. Sex differentiation in larval insects. Biol. Bull. Vol.
XXII.

17. Koschevnikow, G. 1891. Zur Anztomie der mannlichen Geschlechtsorgane der
Honigbiene. Zool. Anz. Vol. XIV.

18. Landois, H. 1863. Uber die Verbindung der Hoden mit dem Ruckengefass bei
der Insekten. Zeit. f. Wiss. Zool. Vol. XIII. 316-18.

Annals Entomological Society of America |Vol. XII,

Lecaillon, A. 1902. Sur le testicule d’Anurida maritima. Bull. Soc, Ent.
France. 64-67.

Lecaillon, A. 1902. Sur la disposition, la structure et le fonctionnement de
l'appareil reproducteur male des Collemboles. Bull. Soc. Philom. Vol. IV.
99-103.

Lowne, B. a 1895. Anatomy of the Blow Fly. Vol. II. p. 660. London.

Malpighi, M. 1669. Dissertatio epistolica de Bombyce, Societati regiae
Londini ad scientiam naturalem promovendam institutae dicata. London.

Meves, F. 1907. Uber Centralkérper in Mannerlichen Geschlechtszellen von
Schmetterlingen. Anat. Anz. Vol. XIV.

Nusbaum, J. 1882. Zur Entwicklungsgeschichte der Ausfthrungsgange der
Sexualdriisen bei der Insecten. Zool. Anz. Vol. V. 637-643.

Packard, A. S. 1898. Textbook of Entomology. New York.

Peterson, W. 1900. Beitrage zur Morphologie der Lepidopteren. Mem. Acad.
St. Petersbourg. Vol. 1X

Peterson, W. 1904. Die Morphologie der Generations-organs der Schmetter-
linge und ihre Bedeutung fiir die Artbilding. Mem. Acad. St. Petersbourg.
Vol. XVI.

Peterson, A. 1912. Anatomy of the Tomato Worm Larva Protoprace car-
olina. Ann. Ent. Soc. Amer. Vol. V. 246-49.

Riley, W. A. 1915. Laboratory Outlines in Entomological Histology.

Saint-George, LaValette. 1897. Zur Samen und Eibildung beim Seiden-
spinnen (B. mori). Arch. Mikr. Anat. Vol. I.

Schneider, A. 1883. Uber die Entwicklung der Geschlechtsorgane der Insek-
ten. Zool. Beitrage v. A. Schneider. Vol. I.

Schroeder, Chr. 1912. Handbuch der Entomologie. Jena.

Severin, H. H. P. and H.C. M. 1908. Internal organs of reproduction of the
male saw-fly. Cimbex americana Leach. Ann. Ent. Soc. Amer. Vol. I.
196-206.

DeSintey, R. 1900. Homologation du testicule chez les Phasmes. Bull. Soc.
Ento. de France. Vol.

DeSintey, R. 1901. Recherches sur la biologie des Phasmes. La Cellule, 1901.

Spichardt, C. 1886. Beitrag zur Entwickelung der mannlichen genitalen und
ihrer Ausftthrgange bei Lepidopteren. Ver. d. Nat. Vereins zu Bonn. 43 Jahrg.

Stitz, H. 1903. Zum Genitalapparat der Lepidopteren. Zool. Anz. Vol.
XXVII.

Tichomiroff, A. 1880. Bau der Sexualdriisen und Entwicklung der Sexual-
producte bei Bombyx mort. Zool. Anz. Vol. III.

Tichomiroff, A. 1898. Zur Anatomie des Insectenhodens. Zool. Anz. Vol.
XXII.

Toyama, N. K. 1894. On the spermatogenesis of the silk worm. Bull. Coll.
Agri. Imp. Univ. Japan. Vol. II. }

Verson, E. 1889. Zur Spermatogenesis. Zool. Anz. Vol. XII.

Verson, E. 1889. La Spermatogenesi nel Bombyx mori. Padova.

Verson, E. F. and Bisson, E. 1895. Development postembryonnaire des
organes sexual accessoires chez le male du Bombyx mort. Arch. Ital. de
Biol. Vol. XXIV.

Verson, E. 1899. Sullufficio della cellula gigante nei follicoli testicolari
degli insetti. Atti. Ist. Venet. Vol. LVII.

Verson, E. 1904. Zur Entwicklungsgeschichte der mannlichen Geschlects-
anhange bei Insecten. Zool. Anz. Vol. XXVII.

Verson, E. 1911. Uber die Versonsche Zelle der Autoren in den Hodenfachern
der Lepidopteren. Zool. Anz. Vol. XX XVIII.

Zick, Karl. 1911. Beitrage zur Kenntniss postembryonalen Entwicklungs-
geschichte der Genitalorgane bei Lepidopteren. Zeit. f. wiss. Zool. Vol.
xX<CViEE

1919]

Ruckes: Male Genital System in Lepidoptera 209

EXPLANATION OF PLATES

.- A, B, C, D. Chlodkovsky’s four types of Lepidopterous testes.

All diagramatic.
E, adult genital system in Samia cecropia, semi-diagramatic. *& 8. °
Sagittal section of testis, showing chambers, hilum, and two coats.

Note the continuity of the basement mem-
brane of the chamber with that of the epithelium of the vas deferens.

The relation of the nervous and reproductive systems in an early pupal

Detail of apical region of a chamber in the testis of Pieris rape, showing
The black parti-
cles in the cytoplasm of the apical cell are portions of the parent
(probably formed by amitosis) migrating to the fibrous
prolongations to become the nuclei of the spermatocyst capsule.

PLATE X.
Fig. 1
A, Hepialus; B, Saturnia; C, Lycaena; D, Pieris.
Bip: 2:
Diagramatic.
Fig. 38. Detail of hilum of testis.
Camera lucida drawing.
PiaTE XI.
Fig. 4.
stage of Philosamia cynthia.
Fig. 5. Detail of coats of testis of Callosamia promethia.
Fig. 6. Detail of coats of testis of Philosamia cynthia.
lier tle
Verson’s cell giving rise to spermatocyst capsules.
nucleus,
Fig. 8S. Adult (sperm stage) spermatocyst.

PLATE XII.

Figs. 9-15. Growth of the epithelium of vas deferens from larval (or early pupal)

stage (9) to the late pupa, (13) when the cytoplasm contains secretion
droplets, to the adult stage (15), when the secretion droglops disappear
and the tips of the cells vacuolize.

A, Trans-section of ejaculatory duct, near ae union Ww ith the ‘‘ductus

Fig. 16. Trans-section of vas deferens, showing general contour oF -spithelium,

Fig. 17. Epithelium of upper end of seminal vesicle.: //

Fig. 18. Epithelium of accessory glands. /

Fig. 19.
ejaculatorius duplex.’ B, Epithelium of same.

Fig. 20. Trans-section of ejaculatory -duct, midway along its length. Note
the addition of: circular muscle.

Fig. 21. Trans-section of ejaculatory duct near ejaculatory

“bulb.’’ Note the

addition of muscles and the presence of an intima.

Each scale line is equivalent to 100 micra.

ABBREVIATIONS.

Ac.—Apical cell (Verson’s cell)—

Follicular epithelium.
Ag.—Accessory glands.
Be.—Ejaculatory “‘bulb.’
Bm.—Basement membrane.
Cce.—Capsular coat.
Cm.—Circular muscles.
Cp.—Caudal plexus.

Ded.—‘‘Duetus ejaculatorius duplex.”
Des.—Ductus ejaculatorious simplex.

Epi.—Epithelium (of genital ducts).
Fd.—Fat droplets.

I.—Intima.

Lm.—Longitudinal muscles.
Lvd.—Lumen of vas deferens.
P.—Penis.

Pg.—Pigment granules.
Sd.—Secretion droplets.
Sp.—Spiracle.

Spe.—Spermatocyst capsule (follicular
epithelium).

Spg.—Spermatogonia.

Spn.—Nucleus of spermatocyst capsule.

Spv.—Vestigial spiracle.

Sv.—Seminal vesicles.

T.— Testis.
Tm.—Tracheal membrane.
Tr.—Trachee.

Ttc.—Testicular tube coat.
Vd.—Vas deferens.
Vp.—Vesicular plexus.

For Fia. 4.

7, 8, 9, 10—Abdominal nerves.
A—Anterior.
P—Posterior.
S—Sympathetic.

VoL. XII, PLATE X,

ANNALS E. S. A.

eee ae

--—Vd

Ded

2

He

:
y
“N
DS
A
“\

Herbert Ruckes.

ANNALS E.S. 4A, VOL. XII, PLATE XI.

Herbert Ruckes.

} ,

Vou. XII, PLATE XII.

ANNALS E. S. A.

TY

VirbertPruches cat

Herbert Ruckes.

THE AQUATIC ADAPTATIONS OF PYRAUSTA PENITALIS
GRT. (LEPIDOPTERA).*

PAUL S. WELCH.

INTRODUCTION.

The unique and ingenious methods whereby aquatic animals
have solved their problems of maintenance are always. interest-
ing, particularly in a group having an immediate terrestrial
ancestry, as is the case with aquatic insects. It is a. well-
known fact that among insects all graduations between the
terrestrial and aquatic species can be found, often within: one
family. Similar transitions are now known to exist within a
single species.. Some of these transitional forms, while retaining
certain terrestrial. habits .and potentialities, have. progressed
far in the development of aquatic adaptations and solve the
problems of the new environment with remarkable ease. - While
Lepidoptera :are preéminently terrestrial, there exists a very
small, heteregeneous group of species which presents. aquatic
adaptations: of : peculiar interest: . The climax ..in:, aquatic
adjustment for this group is probably represented by the gilled
caterpillars of (:Nymphula. (Welch,. 1916). Pyrausta: penitalis
Grt., on-whichithis.paper is. based; presents a striking. case, not
only of transition from terrestrial to aquatic habit, but: also
of a development of physiological adaptations which parallel
closely those Su Ssh in certain other unrelated sor ae aaa
(Welch, 1914)...

The field data used in this ‘paper were Beate ania the
summer of 1917, while the writer was a member of the Ohio
State University Lake Laboratory staff. In the protected
coves of Sandusky Bay, Lake Erie, particularly Beimiller’s
Cove, Nelumbo lutea grows abundantly and during July, 1917,
there appeared upon this plant large numbers of the caterpillars
and pupz of Pyrusta penitalis. The abundance of material
and the constant presence of practically all degrees of develop-
ment made possible a wide range of observations in a relatively
short time. Since the writer has spent but the one season at

_ *Contribution from the Zoological Laboratory of the University of
Michigan.

2X3

j
'
. H
: '

214 ‘Annals Entomological Society of America [Vol. XII,

the Lake Laboratory, this report is incomplete in many respects,
but the data contained herein are presented with the hope that
they may serve as an added basis for future work.

- Several papers already published contain information on this
insect, but in all of them the data are meager and fragmentary,
the sum total representing a very incomplete account. Smith
(1890a; 1890b) discussed it under the name Botis nelumbialis,
giving account of the imago, mature larva, and pupa. Later,
Hart (1895) described in more detail the stages of the life-
history and presented a few biological features of the larva and
pupa. Recently, Chittenden (1918) recorded some interesting
terrestrial activities of this species.

Cogquillett (1880) described what he thought to be the larva
of ‘‘ Botis penitalis,’”’ stating that it fed on Indian hemp (A pocy-
num cannabinum). Chittenden (1918, p. 454), however, points
out a certain disagreement of Coquillett’s description with the
true larya of penitalis and suggests the probability that some
other species was under observation. Dr. J. McDunnough, ina
recent letter to the writer, confirms the suspected error in this
record and states that the A pocynum feeder is the allied futilalis
Led. and that he has bred the latter in abundance at Decatur,
Illinois. 7 :

The writer wishes to express his indebtedness to Dr. J.
MecDunnough for confirmation of the identification of this
SPe€Cies.

THE LIFE-CYCLE.

In the above-mentioned papers are found accounts of the
full-grown larva, the pupa, and the imago and further mention
is not required here. However, the eggs and place of oviposition
have heretofore remained undescribed. The writer, after
much searching, finally found, on July 28, a single egg-mass on
the upper side of a Nelumbo leaf, which, when brought into the
laboratory and kept in an aquarium yielded larve that were
unquestionably those of penztalis. This mass was amber-
colored, and approximately circular in contour, having a
diameter of 3 mm. About sixty eggs were present, surrounded
by an amber-colored matrix, and closely set together. On
hatching, the egg-shells lost the amber color and assumed a
dirty grey appearance. All of the eggs developed normally
and larvae emerged, but since the exact time of the oviposition

1919] Welch: P yrausta Penitalis

of the mass was unknown, the length of the egg-stage could not
be determined. The young larve, at the time of hatching,
were about 1.5 mm. long. They immediately began feeding
upon the leaf in the vicinity of the egg-mass, consuming the.
upper surface in the characteristic manner. The surface
webbing, described later, appeared very soon, subsequent
development and activities continuing in the fashion normal
for the species.

The difficulty of finding the egg-masses is quite puzzling
since the writer examined hundreds of Nelumbo leaves and the
amber color of the egg-mass against the green background of
the leaf, if that be the usual place of oviposition, should make
detection rather easy. Furthermore, the writer observed
dozens of leaves each bearing very small larve which, it would
seem, must have emerged from eggs deposited very near by,
but no indications of old egg-masses were detected.

Chittenden found a considerable number of these larve
boring into the canes of raspberries and likewise, to some
extent, in the stalks of corn. However, he states (p. 455), that
‘‘there is no evidence that the species feeds on either healthy
corn or raspberry, although it feeds on the pith to.a considerable
extent, but on the contrary develops chiefly upon lotus and other
aquatic or semiaquatic plants and enters cornstalks and the
cut ends of raspberry canes chiefly as a retreat for passing the
winter and for subsequent transformation.” ‘This statement is
not entirely clear. It seems to imply that corn and raspberry
may not be affected unless in some unhealthy condition. It
also appears to suggest that these larve pass their early develop-
ment in aquatic situations and then some of them migrate
to certain terrestrial plants in order to find a hibernation place.
Chittenden has tried to formulate ‘‘an approximate life history”’
from the fragmentary bits of available information and sug-
gested that the larve constitute the hibernation stage; that the
adults begin emerging in March; that larve appear in due course
of time and feed until late in August when they complete their
growth, seek a convenient winter retreat and hibernate. Such
a statement suggests only one generation per year.

Certain observations of the writer do not seem to corroborate
the above-mentioned inferences. It was not possible to
determine the hibernation stage and it may be that some of the
larvz, in late autumn, do migrate to shore and find hibernation

216 Annals Entomological Society of America |Vol. XII,

quarters in connection with other plants. However, Nelumbo
often occurs somewhat remote from shore and while, as will be
shown later, these larve are efficient swimmers, it seems rather
doubtful if such is the only method. Since pupz are found in
the upper end of the Nelumbo petioles during the summer, the
writer is inclined to suspect that hibernation, in the aquatic
situations, may occur as pup# in cocoons within the old food
plants. Circumstantial evidence also seems to indicate more
than one generation per year. During the first week in July,
larvee of all sizes as well as pupa were found and from that
time to early August, when the writer’s observations ceased,
adults were emerging continually from the Nelumbo petioles.
Quite a number of such pupz were reared in aquaria in the
laboratory. Since these immature forms occurred in connection
with the fresh léaves and petioles and were of all sizes and ages,
the evidence seems clear that they were not overwintering
individuals. Furthermore, the emergence of adults throughout
the most of July (and possibly to some extent later) and the
finding of eggs:‘on July 28th, suggests a midsummer brood.
Therefore, the writer is inclined to believe that-in the region
of Lake Eriethe species is at least double-brooded and that
hibernation may occur in connection with the aquatic feod-
plants.’ ce ak sls tk Si aa vere a SEDER EE AT sae
ADAPTATIONS TO AQUATIC. SURROUNDINGS.

‘The chief aquatic adaptations manifested in this species are
related to the’ problems of food-getting, locomotion, and
respiration—problems which are always of vital importance in
the adjustment of an animal of terrestrial origin and organiza-
tion to partial or complete aquatic life. While this species
thrives under the conditions of its aquatic surroundings, it is
interesting to note that the complete, original terrestrial
organization has apparently been retained, that no structural
features of aquatic importance have appeared, and that the
adjustments to the water are very largely, if not entirely,
physiological, consisting of habits and activities which result
in the favorable solution of the problems involved. The
physiological adjustments of this species to water offer a
very close and interesting parallel to those of Bellura melanopyga
Grote (Welch, 1914) a species which is remote from Pyrausta
penttalis in kinship, but which has solved its major problems

~]

1919] Welch: Pyrausta Penitalts ai

of locomotion, nutrition, protection, and respiration not by
structural developments, but by modified activities remarkably
similar to those of P. penitalis.

FEEDING HABITS.

Food Plants—In the Sandusky Bay region, P. penitalis
was found only on Nelumbo lutea. Other plants in the same
habitats and on nearby shores were examined from time to
time, but appeared to be unaffected by this insect. However,
it is evidently a general feeder, since other writers have reported
it on Polygonum incarnatum, Polygonum hydropiperoides, Eupa-
tortum (species not given), and Nelumbo nucifera (‘Egyptian
lotus’’). An erroneous record of Apocynum cannibinum was
mentioned on a foregoing page.

Leaf-feeding Period.—Two well-defined periods can be
recognized in the feeding activities of the larve, (1) an early
one, the leaf-feeding period, and (2) a later period, the petiole
period. ‘These two periods were given bare mention by Smith
(1890a) and Hart (1896, pp. 180-181), but the details and
adaptations have not been described.

After hatching, the young larve feed upon the outer surface
of the Nelumbo leaf, removing the upper epidermis and a con-
siderable part of the underlying chlorophyll-bearing tissue.
The feeding areas are irregular and vary in size, depending upon
the amount of feeding done. They often become confluent,
forming large irregular areas. When first made, they are light
greenish, but soon change to brown and in time the whole
leaf, if heavily infested, assumes that color.

While any part of the upper leaf surface is subject to attack,
feeding at the periphery differs, in certain distinct respects, from
that within the periphery, and likewise involves distinct
differences in accompanying adaptations.. These activities will
be described in the reverse order of mention.

Leaves bearing young larve almost invariably show, when
examined critically, numerous, very fine strands of silk all over
the upper surface, marking the routes of the wandering larve.
This apparent wastefulness in silk production is perhaps the ©
first form of aquatic adaptation to appear in the life-cycle since
the silken thread continually anchors the tiny larva to the leaf
during its search for an appropriate feeding place and aids in
preventing it being washed off the leaf by waves or other surface

218 Annals Entomological Society of America [Vol. TE

disturbances.. When active feeding begins, the silk formation
becomes a more noticeable and important feature. Feeding
areas are surface excavations, usually with a depth greater than
the diameter of the caterpillar, across which is spun a fine,
somewhat substantial silk web, and under which the larva works
and rests. Frequently, the threads are tied across from one
radiating leaf vein to the adjacent one with sufficient tension
to pull them slightly towards each other, thus producing a
depression beneath the web and providing a retreat for the
larva. These nets are of varying extent, but since they are a
constant accompanying feature of surface feeding, an attempt
was made to determine their significance. That they play
some real part in the life of the larve is manifested by the fact
that while individuals occasionally leave the web-covered
tunnels for a certain amount of wandering, they almost
invariably return to the original ones or preempt others which
happen to be empty. Since the leaves of Nelumbo rest upon the
surface of the water, wave-action, even of a slight degree, often
inundates them and, unless protected in some way, the young
caterpillars would be constantly exposed to the serious menace
of being washed off. A series of experiments showed that the
silk webbing constitutes an efficient safeguard against dis-
lodgment by waves. When the larve are ensconced under
these webs, a vigorous dragging and threshing of the leaf
through the water in a manner more severe than the effect
of a heavy storm in the protected coves, fails in most cases, to
wash them away. Possibly, these webs may serve, also, as
a partial protection against certain enemies and excessive light,
but according to the evidence at hand, its chief réle is that of
an important adaptation to an ever present feature of the
aquatic environment.

Peripheral feeding and accompanying silk formation differ
in character from the above-described activities. For varying
distances the thin edge of the leaf is folded over and tied by
strands of silk which extend from the upturned edge to the upper
leaf surface. Varying amounts of silk are used in this activity.
Instances were observed where the web was thick and brownish,
resembling, to some extent, a sort of cocoon. Often, however,
it is a fine, thin, dainty webbing through which the larve are
plainly visible. Feeding invariably occurs under these webs.
Peripheral feeding is a very common habit nearly every infested

1919} Welch: Pyrausta Penitalis 219

leaf showing evidence of it, and occasional leaves are affected
only in this way. Sometimes, the entire margin is turned up
and tied over. Funnels formed in this fashion are effective
retreats and provide efficiently against the larve being washed
away by wave-action. Sometimes, the turned over edge collects
a small quantity of water, and the larve may be partly sub-
merged for a time, although no inconvenience seems to be
suffered. Since complete filling of the peripheral tunnels with
water is possible, the larva may, in such an emergency, escape
drowning by migration to the petiole junction—a region more
elevated, normally, than the margins and more likely to be free
from water. The rare occurrence of pupzw in these upturned
margins will be mentioned later. Whether the leaf-feeding
period is confined to certain instars was not determined.
Exuvize from small larve were frequently seen, particularly
under the webbings. Field observations on hundreds of larve
showed that the surface feeders include individuals up to about
14 mm. in length. It thus appears that more than one instar
are included in this period.

Petiole Period.—When a certain growth is attained (appar-
ently about 14 mm.), a distinct change in the feeding habit
occurs. Surface feeding under webbings 1s abandoned and the
larva begins to tunnel lengthwise into the upper end of the
petiole, boring through from the upper surface. By what
means the petiole-leaf junction is located with striking exactness
is a matter of conjecture. Instances were observed in which
wandering larve encountered an uninfested leaf, located
immediately the petiole junction and began excavation. Some-
times, the larva bores into one of the chief veins of the leaf,
tunneling down it to the petiole. Occasional failure to make
exact connection with the end of the petiole, resulting merely
in a hole through the leaf, is compensated for by constructing
an entrance into the side of the petiole jist beneath the leaf.
A surface net is often constructed over the petiole-leaf junction
beneath which the larva works in forming the burrow. When
disturbed, it may leave this shelter, wander about over the
leaf for a time, later returning to the web or seeking refuge
under old nets at the periphery.

The burrow varies somewhat but usually the length is about
23-70 mm. and the diameter a little greater than that of the
larva itself. Its entrance is often surrounded by an accumula-

220 Annals Entomological Society of America |Vol. XII,

tion of brownish excrement, the burrow being kept compara-
tively free from such accumulations. Although most, if not all,
of the material excavated from the petiole is passed through the
digestive tract of the larva, the principal function of the burrow
is other than that of a place to feed since the total length does
not exceed double that of the full-grown larva—a condition
which would not prevail if 1t were a true feeding burrow, as in
the case of Bellura melanopyga (Welch, 1914, p. 102) where
burrow length increases with occupancy. Not only is the
tunnel always short, but also, once formed, it becomes brown
inside, and thereafter shows no evidence of further excavation.

After a burrow long enough to contain the whole larva is
constructed, the caterpillar turns about and feeds mostly on
the upper surface of the leaf just outside the entrance. Larve
are frequently observed on the upper leaf surface with only
one or two posterior body segments remaining in the burrow.
A very slight disturbance, however, causes rapid return. Feed-
ing in this position usually results in elongate surface etchings
which radiate from the petiole junction. Sometimes the surface
within easy reach from the burrow is completely denuded. It
thus appears that the chief function of the burrow is to afford
a safe retreat from which surface feeding may be accomplished.
This accounts for the fact that an examination of a large number
of inhabited petioles showed the head of the larva almost
invariably directed towards the leaf. As will appear later,
the burrow serves also as a pupation chamber.

At no time was more than one larva found in a burrow, the
same being true for the pupe. Occasionally, a nearly mature
individual may, for some unknown reason, desert its burrow and
wander about over the Nelumbo leaves, ultimately constructing
a new burrow in some uninfested leaf or else preémpting an
already formed one which it may chance to find unoccupied.
Instances were observed where such a _ wandering larva
attempted to enter a burrow already inhabited, whereupon the
occupant and the invader engaged in active combat at the
entrance, interlocking mandibles, twisting heads, and pushing
each other about in a vigorous manner, until the former suc-
ceeded in repulsing the latter, or until the invader forced an
entrance, displacing the original occupant.

Effect on Food-plant.—In proportion to the numbers of
larve present, the leaf of the food-plant loses its chlorophyll-

1919} Welch: Pyrausta Penitalis 2K

bearing tissue and ultimately shows signs of deterioration. The
upper surface shows discoloration, loses its waterproof character,
and, in the Sandusky Bay region, usually becomes covered with
chironomid (Diptera) larve, which form their mud cases all
over the exposed surface. Burrow construction partly severs
the connection of the leaf with the rootstalk and often results
in the ultimate deterioration of the floating part, even in the
absence of surface feeding. Since the work of a single larva
may lead to leaf deterioration and since it is known that, when
present, the buds and seed-capsules are also attacked, it is
evident that P. penitalis is a serious enemy of Nelumbo. The
ability of the older larve to migrate from leaf to leaf, as described
later, also increases their menace to the food-plant.

LOCOMOTION.

Crawling.—Change of position on the food-plant is accom-
plished by the ordinary methods of crawling. These larve
are sufficiently active to enable them, where Nelumbo leaves are
contiguous, to wander widely from the hatching place.

Swimming.—One of the noteworthy adjustments to the
aquatic surroundings appears in a well-developed form of
surface swimming. Larve removed from petiole tunnels and
dropped lightly on the water remain supported on the surface.
After a few initial squirming movements, active swimming is
begun and, as long as nothing interferes with the surface position
of the larva, progression is distinctly efficient. Swimming
movements consist of an alternate, horizontal, whip-like
motion of the posterior half of the body. The sequence of
motions is as follows: The posterior end of the body is flexed
horizontally on one side into a position in which the caudal
end is directed cephalad; then, with a strong, quick sweep, as
if released from considerable lateral tension, it returns to the
normal position, stopping there for a brief interval; then
a similar flexure and return is made on the opposite side.
This whip-like motion, first on one side and then on the other,
resembles the action of a straight steel spring fixed at one end,
bent laterally, suddenly released and stopped at the median
position without further vibration; then bent and released on
the opposite side. This strong, vigorous return of the posterior
end to its normal position constitutes an effective stroke against
the water and propels the animal at a goodly rate. So strong

222 Annals Entomological Society of America |Vol. XII,

is this lateral beat against the water that it swings the anterior
end of the larva in the opposite direction, the succeeding stroke
swinging it in the reverse direction, thus making the course
described by the head a distinct zig-zag, characterized by
abrupt rhythmic changes in the lines of progression. While
the swimming activities of this larva resemble in many respects,
those of the larva of Bellura melanopyga (Welch, 1914, pp.
110-112), there is a distinct difference in the swimming move-
ments, since in the latter they are of the graceful sinuous type
and lack entirely the jerky character of the former.

As mentioned above, these larve swim in a surface depres-
sion. They have an integument which resists wetting and
enables them to utilize the surface film, but experiments showed
that if these caterpillars break entirely through it they sink
slowly, performing in vain the swimming movements described
above. They seem to have no ability whatever to swim in
water and individuals which become completely submerged
ultimately go to the bottom. The fate of such larve is not
known. It is possible that some may return to the surface by
crawling up the stems and petioles of floating and emergent
vegetation, since experiments showed that they can withstand
continued submergence for at least one and one-half hours.

Sometimes a larva gets inverted on the surface film and
has great difficulty in righting itself. The swimming activities
are continued, however, in the regular way, progress being
made until contact with some plant or floating object enables
the caterpillar to regain its normal position.

Ordinarily, larva seem somewhat unwilling to leave a
supporting leaf and take to water, but when sufficiently dis-
turbed they will voluntarily do so. No doubt new retreats
and new feeding surfaces are sought in that way also.

RESPIRATION.

Larval Stages.—As mentioned before, the larva has developed
no structural adaptations in connection with its aquatic rela-
tions. So far as respiration is concerned, it is a typical air-
breather and requires direct exposure to the atmosphere.
Throughout the whole larval existence, protection against
complete submergence is afforded. Certain features of the
food-plant aid materially in accomplishing this end. The
exposed surface of the leaf is waterproof, due, according to

1919| Welch: Pyrausta Penitalis 223

Pond (1918, p. 181), to many very delicate hairs which enclose
an envelope of air, preventing the water from actually touching
the epidermis. Unless the leaf has suffered considerable
damage, water thrown upon its upper surface rolls off imme-
diately in silvery masses. Some water is occasionally trapped
on the surface by upturned margins, but it also fails to wet
the surface, and rolls off at the first opportunity. Furthermore,
the region of the petiole junction 1s normally somewhat higher
than the surrounding area, thus producing a sloping surface
which easily sheds off any water falling upon it. These two
features of the plant are effective not only in keeping the
exposed surface above water, but in keeping water away from
the burrow. Surface feeding ultimately makes the leaf unten-
able by the removal of the waterproofing provision and the
subsequent wetting and submergence of the surface. Except
for short accidental periods of submergence, contact of these
larve with the water is very slight and offers no obstacle
to their obligatory holopneustic respiration. Surface swimming
also permits dissemination without interference with the
usual respiration activities.

All of the burrows which came under the writer’s observa-
tion seemed remarkably free from water, in spite of the fact
that at least a part of each fully constructed burrow is below
the water-level outside. In contrast to the larva of Bellura
melanopyga which lives in a water filled burrow in the petiole
of the yellow waterlily and secures oxygen only when at the
top, the larva of P. penitalis seems to occupy a burrow which
is free of standing water and ordinary spiracular respiration is
possible at all times.

Pupal Stage.-—Although the exact oxygen demands of the
pupa are not known, it is fair to assume that they differ but
little from other similar pupe formed in terrestrial environ-
ments since the structural features of the respiratory system
are the same. Continued submergence of a pupa removed
from its silken covering results fatally, indicating that it also
must have a certain protection from water. In connection
with the field observations, the writer examined many cocoons,
none of which contained water. In fact, the pupa appears to be
well provided for in this regard. When the full-grown larva
begins preparations for pupation, the first activity is the forma-
tion of cream-colored, circular, concavo-convex, very closely

224 Annals Entomological Society of America |Vol. XII,

woven silk cap or plug at the upper end of the tunnel, com-
pletely closing it. This plug is quite firm in texture and has a
thickness of about 0.7-0.8 mm. over most of its area. At the
periphery, where it makes contact with the side of the burrow,’
it is reduced in thickness to an extremely fine membrane, thus
providing for the emergence of the imago. The greater part of
the burrow is then lined with a strong, silken cocoon, in the
upper end of which the pupa is formed. This cocoon is con-
nected with the above-mentioned cap by means of a vertical
curtain of silk. The closing plug and complete cocoon seem to
be effective in excluding the water and make it possible for this
type of pupa to successfully pass its quiescent period in a position
slightly below the water line. That thisis an aquatic adaptation
is indicated by Chittenden’s observation (1918, p. 456), that
under the terrestrial conditions of raspberry plants a ‘“‘small
amount of silk is used in the construction of these hibernating
chambers, and.a little is usually to be found at either end’’—a
description which in no way applies to the same activities of the
larva in the Nelumbo petiole. The versatility of this larva in
becoming adjusted to different surroundings is manifested by
the fact that the writer found a few instances of pupation in
the rolled up edges of the Nelumbo leaf—instances in which
a strong, silken cocoon was formed, but no hint of any special
features such as the lid-like cap to the petiole burrow. Nothing
can be stated at present concerning the nature of the stimuli
or sets of stimuli which are operative in impelling the larva to’
produce an entirely different cocoon for each of the three known
pupation conditions, viz., the petiole burrow, the rolled leaf
margin, and the stems of certain terrestrial plants. The fact,
however, certainly emphasized the special adjustments which
have been developed in connection with the aquatic conditions
and which have been superimposed upon a persistent ancestral
habit.

While it seems evident that the closing cap at the entrance
to the tunnel and the underlying, strong, silken cocoon provide
a certain protection against drowning, they are not sufficient
to safeguard the pupa from another danger—the larve of its
own species. It often happens that some larva, wandering
on the Nelumbo leaves, eats away the closing cap of a pupal
chamber, penetrates the upper end of the cocoon, destroys the
upper portion of the pupa, and ensconces itself in the re-opened

1919] Welch: Pyrausta Penitalis 225

tunnel. Whether the tissues of the pupa were actually used as
food or whether the tunnel making instinct of the invading larva
caused it to excavate anything which obstructed passage was
not determined.

SUMMARY.

1. Pyrausta penitalis Grt. (Pyralidz) occurs abundantly in
certain protected situations about Lake Erie in connection with
its favorite food plant, Nelumbo lutea. It belongs to that
extremely limited but interesting, heterogeneous group of
Lepidoptera, which has made progress in the development of
aquatic adjustments.

2. The life-cycle is imperfectly known. Evidence indicates
at least two generations per year. Larve and pupe develop in
connection with Nelumbo, and eggs, described for the first time,
occur on the same plant. Previous writers report relations not
only to certain other aquatic plants, but also to two strictly
terrestrialones. Dependable data on the hibernation are lacking.

3. While successfully adjusted to aquatic surroundings,
this species has retained the ability to utilize terrestrial con-
ditions, at least to a limited extent. The complete, original,
terrestrial organization has apparently been retained also, so
that the aquatic adaptations manifested are very largely, if not
exclusively, physiological.

4. Surface feeding by young larve occurs under silken

webs so constructed as to afford adequate protection against
dislodgment by wave-action, an ever present menace. The
upturned, silk bound margins of the leaf function similarly.
Older larve construct short petiole tunnels which function
only as places for retreat and protection, from which a different
type of surface feeding is performed.
5. The older larve possess a well-developed, efficient form
of surface swimming consisting of an alternate, horizontal,
whip-like motion of the posterior portion of the body. They
have, however, no ability to swim under the surface and, when
completely submerged, sink to the bottom.

6. While an older larva can withstand constant submergence
for about one and one-half hours, its typical holopneustic
tracheation necessitates access to the atmosphere. Protection
from prolonged submergence is provided in part by the water-

{
a , ‘
- ‘

226 Annals Entomological Society of America [Vol. XU,

proof character of the upper epidermis of the Nelumbo leaf and
the elevated position of the petiole junction. Surface swimming
permits dissemination without serious interference with normal
respiration.

7. Pupation occurs in the petiole burrow below the water-
level outside and since submergence is fatal, protection is pro-
vided not only by the walls of the petiole and the formation of a
firm, silken cocoon, but also by the construction of a special
closing device—a cream-colored, circular, concavo-convex cap—
at the top of the tunnel excluding the water, but providing for
the ultimate emergence of the adult. Occasional pupation in
the upturned leaf margin is accompanied only by simple cocoon
formation, but pupation in terrestrial plants, according to
certain writers, appears to be nearly devoid of such provision.

LITERATURE CITED.

Chittenden, F. H. 1918. The Lotus Borer. Journ. Ec. Ent., 11:453-457. 1 pl.
Ragnillstty ie W. 1880. On the Early Stages of Some Moths. Can. Ent.,
Hart, C. A. 1896. On the Entomology of the Illinois River and Adjacent Waters.
Bull. Ill. State Lab. Nat. Hist., 4:149-273. 15 pl.
Pond, R. W. 1918. The Larger Aquatic Vegetation. Fresh-water Biology, by
Ward, H. B., and Whipple, G. C., pp. 178-209. 19 fig.
Sua ae An Enemy to the Egyptian Lotus. Garden and Forest,
1890b. A ee Species of Botis. Entomologica Americana, 6:88-90. 2 fig.
Welch, P. S. 1914. Habits of the Larva of Bellura melanopyga Grote (Lepid-
optera). Biol. Bull., 27:97-114. 1 pl.
1916. Contribution to the Biology of Certain Aquatic Lepidoptera. Ann.
Ent. Soc. Am., 9:159-190. 3 pl.

CHARIESTERUS AND ITS NEOTROPICAL RELATIVES
(COREIDZ HETEROPTERA).

S. B. Fracker, Madison, Wis.

The Squash-bug family contains an unusually large pro-
portion of insects of bizarre form and possessing strange struc-
tures. Expanded or curved tibia, swollen and spinose femors,
and flattened or clavate antennal joints seem to be the rule
rather than the exception.

Among the more modest of the peculiar species are those of
the tribe Chariesterini, a group whose members possess a
flattened spatulate expansion of the third antennal segment
and obliquely truncate, usually spinose, antenniferous tubercles.
Chariesterus antennator, the most northern member of the
tribe, is well known from New Jersey and Jowa to the Gulf
of Mexico. A new species from California is described below,
whose peculiarly shaped pronotum makes it one of the most
remarkable of the Coreide. :

‘The tribe is confined to the western hemisphere and includes
three genera: Plapigus, Staluptus, and Chariesterus.

Plapigus now contains seven or eight described species, all
South American except circumcinctus Stal from Mexico. This is
a black form, with margins of pronotum, corium, and abdomen
flavescent, a distinct callosity behind the eye, tibize with
testaceous annulus, and expansion of third antennal segment
longer than broad. The internal angle of each antenniferous
tubercle is acute, and the first segment of the antennz sub-
clavate; the pronotal margins are entire, and the humeri each
armed with a single long slender spine.

Staluptus marginalis Burmeister, according to the original
description, is black, with unarmed posterior femora, which are
fulvous at base, scarcely dilated third antennal segment, and
brown venter and femora. It was described from Mexico.

-The species of Chariesterus, seven in number, are more
widely distributed than other members of the tribe, ranging
throughout both continents. In this genus the first antennal
segment is parallel-sided, though sometimes slightly broader

227

\ ‘

ied

‘

‘

\ a '
’

‘ 1

228 Annals Entomological Society of America |Vol. XII,

toward the tip. The callosity behind the eye, instead of being
wanting, as Stal states, is merely usually smooth without acute
points, although a few specimens of C. antennator from Kansas
and Nebraska are at hand in which this tubercle is distinctly
muricate.

The following synopsis is based on specimens in the col-
lection of Dr. E. D. Ball, Mr. William J. Gerhard, of Chicago;
Mr. H. G. Barber, Mr. Carl J. Drake, the Wisconsin Agricul-
tural Experiment Station, and the Milwaukee Museum, except
for C. armatus and C. albiventris, for whose positions reliance
was placed on descriptions. The distribution of each species
is indicated in the synopsis for the sake of brevity.

KEY TO THE SPECIES OF Chartesterus.

a. Antenne dilated equally on both sides of third segment.
b. Dilation of third antennal segment only three-tenths as wide as
long; legs and antenne ferrugineous; size 9 x 1.2 mm. (Antilles).
gracilicornis Stal.
bb. Dilation of third antennal segment at least four-tenths as wide as
segment is long; legs and antenne black; body above brown,
below albo-farinaceous; length 11 mm. (Texas and Mexico).
albiventris Burm.
aa. Antenne with third segment more angularly dilated on lower side; length
over 10 mm.
b. Lateral margins of pronotum armed with distinct tubercles in front
of humeri; first antennal segment armed with distinct acute
tubercles, larger near base.

c. Pronotum with expanded, elevated lobes at each lateral angle,
armed with four or five acute processes of similar size; head
with prominent, multispinose tubercle behind each eye.
(Calafornia) feds cack Axe he Bera o ceraeusitigees eemcrer ie balli n. sp.

ec. Pronotal angles not expanded, armed with one large tooth
preceded by several smaller ones; head with tubercles behind
eye smooth or slightly muricate. (United States, east

Of the RockysMountains) eee =e or ae ee antennator Fabr.

bb. Lateral margins of pronotum unarmed in front of humeral spine;
first antennal segment armed with acute tubercles at base only
(or none); subtropical or tropical species.

c. Body castaneous, margins and angles of pronotum darker;
dilation of third antennal segment not notched; venter
refescent; dorsum of abdomen rosaceous; spines of pronotum
remarkably long and slender. (Texas to Panama).

cuspidatus Dist.
ec. Body: testaceous, pronotal margins concolorous with or paler
than disc; dilation of third antennal segment notched; venter
with black spots.
d. Dorsum of abdomen black. (Mexico)........... moestus Burm.
dd. Dorsum of abdomen subsanguineous, base and apex black.
(South America) oer Jee aeoccci ser crac armatus Thunbg.

i

, .
1919] £ Fracker: Chariesterus * 229

ste

Chariesterus balli new species. (See Figure).

Head quadrate, armed on the dise with two longitudinal rows of
two or three vertical spines each; prominent multispinose tubercles
caudodorsad of each eye; antenniferous tubercles spinose at both
obtuse lateral and acute mesal angles. Antenne with first segment

Fig. 1. Chartesterus ballin. sp.

stronglygtriquetrous, lateral carine with basal two-thirds armed with
numerous acute tubercles, mesal carinze unarmed except at base; second
segment much more slender, shorter, compressed and slightly expanded
near the middle; third segment longer than fourth, strongly expanded
throughout three-fourths of the length, four-tenths as wide as long.
Rostrum scarcely attaining posterior coxe.

4
'

230 Annals Entomological Society of America {|Vol. XII,

Pronotum more elongate than in C. antennator, with each lateral angle
elevated, expanded, irregularly armed with three to five large subequal
teeth, anterior and posterior lateral margins with several smaller teeth;
posterior angles each with flat triangular tooth. Scutellum unarmed.
Mesosternum and metasternum, in part, deeply sulcate. Posterior
tibize shorter than femora. All femora armed with short spines near the
apex.

Color black, marked with dark ferrugineous. Head black above,
ferrugineous beneath, with a rufescent vitta below each eye, rugose;
antenniferous tubercles wide, adjacent, black. Antennz black, immacu-
late, except that second segment is ferrugineous; rostrum scarcely
attaining posterior coxe.

Pronotum and scutellum black, pubescent, and very rugose, former
with narrow, scarcely perceptible median line paler and slightly sulcate.
Corium ferrugineous. Membrane sooty black. Venter and _ legs
dusky ferrugineous.

Cotypes: Two males, captured at Cabazon, California, by
Dr. E. D. Ball, and now in his collection.

In the presence of spinose callosities behind the eyes, this
species approaches Plapigus, while the parallel sided basal
antennal segments and the shape of the pronotum indicate
Chariesterus as the proper genus.

C. antennator is the only species which is likely to be con-
fused with balli as all the others possess a single long spine at the
humeral angles. The species here described is more slender
than antennator, a deeper black in color, and with the lateral
angles of the pronotum expanded and elevated and armed
with four or five large teeth. The tubercle behind the eye is
more prominently spinose than in any other species of the
genus.

EIGHT NEW “JASSIDS” FROM THE EASTERN
UNITED STATES.

Family Cicadellidae (Hemiptera~-Homoptera)

J. G. SANDERS and D. M. DELOonG, Harrisburg, Pa.

The mountain regions of Pennsylvania and certain less
elevated sections of New Jersey, and those portions of Maryland
and Virginia between the Chesapeake Bay and the ocean,
have received but scant attention from collectors of Homoptera
if existing records are dependable. Although relatively little
collecting has been done to date, we are able to describe seven
new species of distinct interest, while withholding other appar-
ently new forms for more material and better data.

These new species are described from Pennsylvania, one
from New Jersey and three from the Maryland-Virginia
peninsula; with one exception, all collected by the senior
author during the past two years.

Especially interesting and beautiful is the new Phlepsius
collected near South Orange, N. J., by Mr. Harry B. Weiss,
from ‘‘Hercules Club” (Aralia spinosa); and also the very
distinct and beautifully marked Deltocephalus from grasses
on the mountain top near Port Matilda in central Pennsylvania,
taken by the senior author.

Platymetopius rubellus n. sp.
Plate XIII, Figs. 1, 2, 3, 4.

_ Form of cuprescens with long pointed vertex, but smaller and with
reddish color above and on face. Female, 4.75; male, 4.25.

Vertex longitudinally concave, a little more than twice as long
as width between the eyes, margins slightly concave, either side to a
very acute apex. Pronotum short and broad. Not quite two-thirds
as long as vertex, and more than twice as broad as long. Elytra rather
short and broad, broadly rounded at apex. From side view, the front
is concave and very acutely angled with vertex. Face long, frons
extremely narrow, clypeus constricted just above middle, expanded to
rounding apex.

Color: Generally reddish brown, shading toward infuscated apices of
elytra. Vertex with median apical line and a line either side extending
along margin half way to eyes, and then inward on disc, with faint traces
toward median basal portion, pale. Median line deep and black on

231

;
c; | 2

\ 4 ‘ea
Annals Entomological Society of America |Vol. XII,

bo
(Su)
bo

basal half. Faint traces of vittee on pronotum and scutellum; the latter
with apex and point at middle of either side, white. Median impressed
transverse line, black. Elytra reddish brown with veins darker, super-
numerary costal veinlets and vermiculate markings brownish, heavier
at apex. Apex with a white band. Areoles few and distinct; two
along suture on posterior half of clavus, a row along anterior portion of
apical cells and posterior portion of anteapical cells, and one at anterior
end of first and second anteapical cells. Front and face uniform
reddish brown, often flecked with reddish, and infuscated at apex of
vertex, angular line long and very distinct, irregular, dark margin
above and below. Eyes red.

Genitalia: Last ventral segment of female with wide sloping to
rather prominent angles, then slightly concavely rounded to produced
median truncated lobe. Whole segment dark bordered. Male valve
broad and roundingly produced to a distinctly angled apex twice as
long as preceding segment, which is concavely angled on posterior
margin. Plates broad at base, wider than valve, gradually narrowed
to sharp tips.

Described from two females and one male from Battle
Point, Va., June 22, 1918, and’ a female from Parksley,;Vax
June 21, 1918. Collected by the senior author.

Platymetopius collaris n. sp.
Plate Sau ericsH oe Ones

A blunt-headed pale species with pronotum and scutellum largely
black. Length, Female, 5 mm.; Male, 4.5 mm.

Vertex nearly twice as long as width between the eyes. Posterior
angle of head flaring. Pronotum strongly arcuate anteriorly, truncate
posteriorly, obliquely cut to outer flaring angles. Elytra broadly
rounded at tip. Approximately ten supernumerary veinlets along
costal margin.

Color: Vertex dull pale orange, elongated median spot at apex,
irregular spot at either side ivory white, between these and extending
posteriorly, irregular lines and washing of brown merging and fading
posteriorly. Median posterior spot white. Line behind eyes black.
Pronotum dull yellow, median two thirds black or dark brown, with
pale median line. Scutellum before transverse suture black or dull
brown, apex bright yellow. Elytra pale to buff, apical cells clouded
and with variable tiny vermiculate markings. Areoles few, white, two
in clavus, one at base of each apical cell, and at apex of each anteapical
cell. Supernumerary costal and apical veinlets, black or brown.
Frons pale shading to dull orange on the cheeks. Pale line extending
from tip of apex across eyes in well marked specimens. White “v”
below apex indistinct. Venter mottled with gray and brown; pale
orange on plates.

e
,

1919} Sanders and DeLong: Eight New “ Jassids”’ 250

Genitalia: Female last ventral segment strongly and evenly pro-
duced to a blunt angle. Male valve large, half as large as plates,
strongly produced to a blunt angle. Plates evenly narrowed to blunt
tips.

Described from four females and one male swept from
Vaccinium sp. on the mountain at Penfield, Clearfield Co.,
Pa., August 24, 1918, by the senior author. Type and para-
type in authors’ collections.

Deltocephalus delector n. sp.
Plate XIII, Figs. 8, 9, 10.

Similar in form and general appearance to bilineatus and marginatus,
but much smaller, really a minute species with distinct markings.
Length, male, 3 mm.

_ Head well produced, with apex distinctly pointed, as long as pro-
notum, about one-fourth longer on middle than width between the eyes.
Pronotum about twice as wide as long, strongly arcuate in front, lateral
margins very short, rounded. Elytra with clavus not reticulate and
with middle anteapical cell strongly constricted at middle.

Color: Vertex creamy white, with two broad tawny bands extending
from base to apex, where they converge and form two black triangular
spots either side of white tip, and extend over margin. Behind these
in each band a ‘‘v”’ shaped black spot inclined inwardly. A narrow
black curved line extends from eye around ocellus on margin toward
apex. Median white band distinctly narrowed to apex. Pronotum
white, with four longitudinal tawny bands, one behind either eye, and
the central two are extensions of the bands on vertex, continuing across
scutellum. Elytra tawny with costal, sutural and apical margins and
veins, white. A spot in distal portion of inner apical cell, the costal
veinlets of the apical and anteapical cells, and a curved band on inner
border of the apical white margin, black. Face pale, a spot on the outer
margin of lore, one above antenna, and inner line of antennal pit,
brown. Beneath pale yellow, male plates with inner apical third black,
but white margined.

Genitalia: Male valve short and broad, with nearly truncate apex.
Plates three times length of valve, gradually sloping to broad truncated
apices, the outer margins set with coarse spines.

Described from one male specimen swept from grasses by the
senior author on the top of mountain between Port Matilda
and Phillipsburg, Pa., August 23, 1918. A second specimen
was taken, but escaped.

This species is very distinct from others of the bzlineatus
group, on account of its minute size and markings.

234 Annals Entomological Society of America |Vol. XII,

Deltocephalus latidens n. sp.

Closely resembling misellus, but slightly larger and more robust,
with color pattern more vague and with distinct genitalia. Length
3) Tati,

Vertex: Flat, slightly longer than width between the eyes, width
across eyes almost equaling combined length of vertex and pronotum.
Pronotum short, more than twice as broad as long, anterior margin well
produced betaveen the eyes. Elytra a little more than twice as long as
broad, barely exceeded by abdomen. Face strongly inflated.

Color: Buff irregularly marked with pale orange. Vertex with
traces of the four quadrants found in misellus usually appearing as a
broken, broad transverse band between anterior margins of the eyes,
and two indistinct triangular spots approaching apex, pale orange.
Pronotum with traces of six longitudinal pale orange bands. Scutellum
pale. Elytra generally darker with pale veins and cells pale orange with
few dark markings. Two small spots along suture, one on middle of
clavus, one just behind anterior cross vein, posterior margin of inner

Dilatid

ens n. 8p. 2

anteapical and posterior margin of apical cells, brown. Face brown with
traces of about eight arcs. Beneath yellow, heavily marked with brown.
Ovipositor and broad tooth of last ventral segment in female brown.
Male plates brown.

Genitalia: Female last ventral segment more than twice as long as
preceding, posterior margin deeply, roundingly incised near lateral
margins two-thirds to base leaving a broad central truncated tooth one-
half width of entire segment produced beyond the lateral angles, which
appear as long, narrow spatulate processes. Male valve broadly
triangular, convex, twice as broad as long. Plates short and broad,
sloping almost uniformly to broadly rounded apices. Outer margins
set with fine white hairs.

Described from eleven females and eight males collected in
1916, at Amery, Wis., August 10; Cameron, Wis., August 7;
Merrillam, Wis., August 5; Taylors Falls, Minn., August 16;
Trout Lake, Wis., September 9; and Sturgeon Bay, Wis.,
July 19, 1914, by the authors.

This little species resembles misellus so closely in general
appearance that the two have been confused. The female
genital characters, however, are very distinct.

Ou

1919] Sanders and DeLong: Eight New ‘‘ Jassids”’ 23

Phlepsius tenuifrons n. sp. - \°
Plate Sal, Figs. ii, 12, is. 14:

Form of ramosus, but shorter, more robust, head more foliaceous
and with distinct genitalia. Length 7 mm.

Vertex well produced, almost entirely concave to sharp, foliaceous
apex. More than twice as long as broad, almost twice as long at
middle as next the eye. From side view almost entire vertex thin,
foliaceous, and upturned. Pronotum short and broad, almost two and
one-half times as wide as long. Elytra short and broad, flaring. Face
yery broad, and rapidly narrowed to base of clypeus. Upper portion
very concave or almost angled to thin vertex.

Color: Vertex and pronotum pale with numerous brown irrorations.
Median portion of vertex with an irregular brown longitudinal line.
Face uniformly irrorate, with pale spot at upper angles of lore. Elytra
milky white, but very heavily marked with broken ramose lines; veins
brown. Central portion more closely inscribed with brown, and
forming a somewhat obscure saddle across’ the middle, having a sug-
gestion of paler area toward base of elytra. Beneath umiform brown.

Genitalia: Last ventral segment of female about three and one-half
times the length of preceding segment. Gradually rounded either
side from half its length to posterior margin, forming rounding lobes
instead of prominent angles. Then concavely arcuate to a broad
produced median lobe, which is keeled, and narrowly notched at middle.
Male valve more than three times as broad as long, roundingly angled
at apex. Plates broad at base, then rather suddenly narrowed at
about half their length and produced to blunt apices, a little longer
than combined width at base. Pygofers very broad at base, convexly
produced and sharply incurved toward tips.

Described from a pair collected at Greensburg, Pa. September
3, 1901, and September 2, 1902. Collected by Mr. M. Wirtner.

The short, broad body with sharp foliaceous head. well
distinguishes this species from its allies.

Phlepsius tinctorius n. sp.
Plate XIII, Figs. 15, 16, 17, 18.

A moderately robust, round-headed species with bright orange
markings on vertex and scutellum. Female, 6 mm. and male 5 mm.
long.

Vertex about three-fourths of width between eyes, very obtusely
angled, almost evenly rounded and parallel margined, with apex narrowly
but decidedly inflated before a distinct depression. Pronotum short
and broad, truncate posteriorly with distinct flaring lateral angles
broader than head. Scutellum transversely concave. Elytra broad
with rounded tips. Face broad and flat with clypeus constricted at
base and broadly truncate at apex.

236 Annals Entomological Society of America |Vol. XII,

Color: Vertex ivory white with small irregular black spot either
side of apex; an irregular semicircular black spot above each ocellus
vaguely connected by transverse wavy line following depression.
Median suture black; on either side a large quadrangular orange spot
posteriorly margined with irregular black lines. Pronotum tawny
anteriorly, shading to darker, generally marked with vermicular brown
lines. Scutellum bright orange with apex and triangular spot midway
of lateral margins ivory white. Transverse suture with central spot
and median line, and irregular lines extending anteriorly from lateral
white spots, black. Elytra milky to smoky with pale tawny areas,
uniformly marked with broken irregular lines; tip of clavus, margin |
of apex, spots of outer apical cell and costal cross-vein, black. Face
black with close irregular white or tawny irrorations. Antenne black,
with pits and apices of basal segments white. Venter generally black
or dark brown.

Genitalia: Female segment longer than preceding and smoothly
truncate. Male valve half as long as preceding segment, broadly
and bluntly angled; plates long triangular, evenly tapering to posterior
third with long curved terete tips.

Described from eight females and one male collected by
H. B. Weiss from ‘‘Aralia spinosa” at Irvington and South
Orange, Essex Co., New Jersey, July 23 and August 1. Type
and paratypes in authors’ collections, and in New Jersey
State Museum Collection.

Chlorotettix productus n. sp.
Plate XIII, Figs. 19, 20.

Small wedge-shaped species with well rounded vertex, a little more
produced than in viridius. Female, length 5 mm.

Vertex roundingly produced, not angled, almost parallel margined;
slightly longer on middle than next the eye, about twice as wide as long.
Pronotum strongly and convexly produced, truncated posteriorly,
lateral margins well rounded. Front broad, inflated.

Color: Dull yellowish green without definite markings. Vertex
and frons shading to buff. Elytra greenish, subhyaline, distinctly
shiny. Paler below, tip of ovipositor and bristles, yellowish.

Genitalia: Female last ventral segment one-third longer than
preceding and longitudinally striated. Lateral angles roundingly
produced. Posterior margin concavely sinuate to produced median
third, slightly notched at middle. Pygofers with heavy bristles on
posterior half.

The produced last ventral female segment distinguishes this species
from others of the genus.

Described from three female specimens swept from a low
shrub at Battle Point, Virginia, June 23, 1918, by the senior
author. Type and paratypes in authors’ collections.

1919} Sanders and DeLong: Eight New ‘‘ Jassids”’ 2at

Chlorotettix fumidus n. sp.
Plate XIII, Figs. 21, 22, 23.

Resembling tergatus in size and color, but narrower, with head well
produced and angled. Length, female, 8 mm.; male, 7.5 mm.

Vertex produced, distinctly angled; one-half longer on middle than
next the eye, and about twice as wide as long. Pronotum very convex
anteriorly and almost truncate posteriorly, length one-half the width.
Second apical cell much narrower than in tergatus.

Color of vertex dull pale orange; face pale straw-yellow. Pronotum
and scutellum smoky green, elytra dark smoky. Body above black;
beneath pale yellow.

Genitalia: Last ventral segment of female about one-half longer
than preceding; obliquely produced to outer angle, with posterior
margin truncate or slightly concave to broad deep median notch two-
thirds distance to base. Male valve triangular, as wide as preceding
segment, apex rounded and posterior margins slightly sinuate. Plate
short and broad, convexly rounded to acute appressed tips.

Described from three female and two male specimens.
Type specimen, female, collected at Mt. Alto, Pa., August 24,
1918, by T. L. Guyton. Two females from Moscow, Tenn.,
collected by the junior author, June 23, 1915; one male from
Paris, Tenn., June 26, 1915, and one male from Speeceville,
Pa., July 22, 1917, by the senior‘author. Type and paratypes
in authors’ collections.

EXPLANATION OF PLATE XIII.

1.—Platymetopius rubellus n. sp., head (dorsal view).

2.—head (side view). 3.—o genitalia. 4——9Q genitalia.
5.—Platymetopius collaris n. sp., head.

6.—o genitalia. 7.—Q genitalia.
8.—Deltocephalus delector n. sp., head.

9—o genitalia. 10—elytron
11.—Phlepsius tenuifrons n. sp., head (side view).

12.—head (dorsal view.. 13.—o genitalia. 14.— 9 genitalia.
15.—Phlepsius tinctorius n. sp., head (dorsal view).

16.—head (side view). 17.—@ genitalia. 18.—< genitalia.
19.—Chlorotettix productus n. sp., head. 20.—Q genitalia.

21.—Chlorotettix fumidus n. sp., head.
22.—o' genitalia. 23.— 9 genitalia.

{
j

e
sal ’
ANNALS E.S. A. VoL. XII, PLATE XIII.

Sanders and DeLong.

NEW WESTERN SPIDERS

By Ratpo V. CHAMBERLIN.

Most of the new forms of spiders described below were
found in collections made by the writer in Utah in the latter
part of August and the early part of September, 1917. The
collections were made on Chalk Creek in the Uintah Mts.,
at elevations of from 7,300 to 10,000 ft. and at and near Fill-
more, Millard Co. In addition to the spiders from Utah, a
number of new forms from other localities noted while identi-
fying the collections mentioned are here described.

DICTYNID.

Amaurobius utahensis sp. nov.

Male—Carapace, sternum, labium, endites and legs yellow, without
any reddish tinge, the legs unmarked. Abdomen blackish, the anterior
middle region of dorsum obscure yellow, the posterior half with dark
marks which are, excepting the first ones, only vaguely separated
from each other. Anterior row of eyes procurved; medians close
together, but more than their diameter from the laterals. Posterior
eyes much smaller than the anterior ones, the medians slightly smaller
than the laterals, about twice their diameter apart and about twice
as far from the laterals. Area of median eyes rectangular, being as
wide behind as in front, longer than wide. Distal joints of palpus
heavier than in nevadensis. Tibial apophyses very similar to those of
the species mentioned; but the exterior one distally less uncate, the
median one basally broader and more narrowed distad, and the mesal
one obviously broader and shorter and differing particularly in being
united with or carried forward on a common base with the median
one much farther, the ridge uniting the two in nevadensis being slight
or obsolete.

Length, 11.5 mm. Length of cephalothorax, 6.1 mm. Length of
tib. + pat. I, 6.4 mm.; of tib. + pat: IV, 6.7 mm.

Type—M.. CC. Z. 414, Utah: Fillmore.

This species is nearest A. nevadensis Simon. It is a more
robust species, wholly lacking the reddish cast characteristic
of nevadensis. It differs chiefly otherwise in the details of
the male palpus as indicated above and also presents minor
differences in the eyes.

239

240) Annals Entomological Society of America [Vol. XII,

Amaurobius nomeus sp. nov.

Female—Carapace and sternum light yellowish brown, becoming
blackish in anterior portion of pars cephalica. Legs paler. Labium
and endites chestnut, pale across tips. Cheliceree black. Abdomen
above greyish yellow, with a narrow, median longitudinal dark mark
extending well caudad of the middle, where its point ends at the angle
of a transverse chevron line, which is followed by several others; in
front of the first chevron there are two or more lines running obliquely
caudoectad from the edge of the median mark. Sides of abdomen
darkened by numerous spots and streaks as is also the venter on which
they tend to form two longitudinal stripes and a narrower median line
separated by clear spaces. Anterior row of eyes procurved; median
eyes smaller than the laterals, their radius or a little more apart, and a
little more than their diameter from the laterals. Posterior row of
eyes longer than the anterior; eyes smaller than the anteriors; the
medians smaller than the laterals. Median eyes forming a trapezium
wider behind than in front in ratio of about 25:20 and wider behind
than long, the length scarcely exceeding the anterior width. Tibia I
armed beneath with two pairs of spines, none being present at distal
end; tibia with a single spine at distal end in addition to the two pairs.
Epigynum smooth, median interval narrow, the process ventrally
rounded. See Plate XIV, Fig. 1.

Male palpus is shown in Plate XIV, Fig. 2.

Length 11.2 mm. Length of cephalothorax, 5 mm. Length of
tib. + pat. I, 4.1 mm.; of tib. + pat. IV, 4.3 mm.

Type—M. C: Z, 416.

Utah: Uintah Mts., 10,000 ft.

This species has resemblance to A. benneti B and A. pictus S.
When placed alongside of these species the cephalothorax is
seen to be obviously more narrowed at the anterior end. It is
distinct in the details of epigynum and eye arrangement.

Dictyna uintana sp. nov.

Female—Carapace fuscous or nearly black, clothed with white hairs.
Sternum blackish. Legs yellow, distinctly annulate with black, the
annuli of tibia and metatarsus deepest, the femora with a submedian
and a distal annulus, patella with annulus at distal end, tibia with an
annulus at each end and metatarsus with an annulus at distal end and
a broader, but less distinct proximal one. Abdomen testaceous; at
base above with a long triangular to hour-glass shaped dark brown
mark followed by a number of chevron marks, of which the first is
connected by its apex with the basal mark; caudal end of dorsum
typically covered by a solid black area; sides of abdomen darkened
above by streaks and spots of brown; venter commonly showing an
often vague longitudinal median dark band. Anterior row. of eyes

i
1919] Chamberlin: Western Spiders 241

straight; median eyes decidedly smaller than laterals, scarcely their
diameter from the laterals, one half their diameter from each other and
about the same distance, or slightly farther, from the lower edge of
clypeus. Posterior row of eyes straight; median eyes obviously larger
than the anterior medians, once and a quarter or scarcely more their
diameter apart and near the same distance from the laterals; posterior
laterals equal to the anterior laterals. Area of median eyes a little
wider behind than in front. In front view the inner edges of the
cheliceree are obviously curved ectad over most of their length, as shown
in Plate XIV, Fig. 3; Epigynum as shown in Plate XIV, Fig. 4.

Male—Chelicerze with a low rounded tubercle or tooth at exterior
angle of base. Palpus with tibial apophysis short, but rather stout.
CE Plate X1Ve Pig. 5:

Type—M. C. Z. 405. XIV, Utah: Uintah Mts:; Chalk Creek,
elevation, 7,500 ft. Webs on boughs of pine trees.

Resembles sublata. The latter, among other differences,
has the eyes of the anterior row obviously more nearly
equidistant and the form of the epigynum different.

Dictyna coloradensis sp. nov.

Female—This species has to some extent been identified with
D. arundinaceoides Keys, but is not that species, which seems to be
really the D. sublata of Hentz. Carapace and sternum fuscous. Legs
yellowish or pale brown, without distinct markings or often with femora
darkened and tibiz and metatarsi annulate at distal end and with less
distinct annuli toward proximal end. Abdomen above with the usual
triangular mark at base; behind this two rows of annular markings more
or less connected by dark lines or chevrons between them. Anterior
row of eyes straight; median eyes a little smaller, separated by about
their diameter, very slightly closer to laterals. Clypeus rather low,
the height being between once and a half and twice the diameter of an
anterior median eye.

Posterior row of eyes straight; eyes equidistant. Area of median
eyes wider behind than in front. Epigynum shown in Plate XIV, Fig. 7.

Male—Darker than the female. Femora darkened and a distinct
dark annulus at distal end of tibiz and metatarsi and a less distinct
one proximally, or legs sometimes lacking these dark markings. Clypeus
much higher than in the female and median eyes smaller. Chelicerz
dentate at base toward ectal side as shown in Plate XIV, Fig. 6.
Process of tibia of palpus stout, of moderate length, distally bi-dentate.
Cf. Plate XIV, Fig. 8.

Length of female up to 3.5 mm.

Type—M..C. Z. 395. Colorado: Colorado Springs (R. V.
Chamberlin, coll.); Eldora; Pagosa Springs; Fort Collins, etc.
New Mexico: Beulah; Mesilla.

242 Annals Entomological Society of America |Vol. XII,

Dictyna bellans sp. nov.

Female—Carapace and sternum pale brown to yellow, sometimes
of slight reddish cast. Legs yellow. Abdomen grey, under the lens
showing the usual network of fine dark lines; a triangular dark spot
on base above followed by two or three pairs of large dark dots which
may be connected on each side. Venter immaculate grey. Anterior
median eyes a little smaller than the laterals. Posterior row of eyes
recurved. Area of median eyes as wide in front as behind. Epigynum
as shown in Plate XV, Fig. 4.

Male—The abdomen in the types is either wholly immaculate or
shows the dorsal markings only vaguely. The basal tibial process
of the male palpus is erect, moderately long, and shows but a single
distinct tooth or point at distal end. Toward anterior end of tibia
above is a second much shorter precess or tooth as shown in Plate XV,
Fig. 5. Process at base of chelicera long and curved. See Plate XV,
Fig. 3.

Length of female up to 2.8 mm.

Type—M. C. Z. 403. Mississippi (R. V. Chamberlin.)

Dictyna sociella sp. nov.

Female—Carapace fuscous, the pars cephalica a little paler. Sternum
dusky brown or fuscous. Legs yellow, some darker annuli faintly
traceable. Abdomen on basal half of dorsum with two connected
triangular dark marks and the usual line of more or less connected dark
spots on each side behind, elsewhere the usual fine network of dark
lines over a light ground. Anterior eyes nearly equidistant, the median
eyes only slightly smaller than the laterals. Posterior eyes also nearly
equidistant.. Area of median eyes a little wider behind than in front.
Epigynum, Plate XV, Fig. 8.

Length, 3.2 mm.

Type—M. C. Z. 412. Washington: Wawawai.
Distinguished particularly by the form of the epigynum and
by the eye relations.

Dictyna insolens sp. nov.

Male—Carapace and sternum fuscous. Legs yellow, somewhat
dusky, the tibiz and more especially the metatarsi obscurely annulate.
The narrow annulus at distal end of metatarsi most distinct. Abdomen
dorsally with a black triangular mark at base followed by a pair of dark
spots and then two heavy chevrons behind which is a clear area; sides
darkened; venter light. Anterior median eyes smaller than the laterals.
Posterior eyes equidistant. Area of median eyes a little wider behind
than in front. Process of tibia of male palpus very short and pro-
portionately wide with the anterior tooth much the larger. See further
Plate XV, Fig. 6. Chelicera with basal tooth slight, rounded.

Length, 2 mm.

Type—M. C. Z. 409. Washington: Olympia.

1919] Chamberlin: Western Spiders 243

Dictyna vincens sp. nov.

Female—Carapace and sternum fuscous. Legs yellow to pale
brown. Abdomen above at base with a black mark having caudal end
excised in an obtuse re-entrant angle and on each side the usual dark
lines radiating from its angles and edges; behind the middle first a pair
of widely separated dark spots and then typically two exceptionally
thick chevrons, which sometimes give the caudal end the appearance
of being almost solid black; sides often darkened; venter either uniform
or with a dusky or blackish band over its length, or with a black area
only in front of spinnerets. Anterior median eyes essentially equaling
the laterals, their diameter apart, closer to the laterals. Posterior
eyes equidistant. Area of median eyes as wide in front as behind.
Epigynum, Plate XV, Fig. 1.

Male—In what is regarded as the male of this species the coloration
is similar to that of the female though the general appearance of the
abdomen is darker and more uniform with the markings less distinct
and sometimes almost obliterated. The palpus is shown in Plate XV,
Bigs. 2.

Length of female, 3.25 mm.; of male, 2.5 mm.

Type—M. C. Z. 410. Washington: Olympia.

Dictyna olympiana sp. nov.

Male—This small form has the usual general coloration; the carapace
and sternum dusky brown; legs yellow; abdomen at base above with a
median dark line expanding into a triangular spot in front of middle,
this followed behind by the usual pairs of longitudinally connected
spots, the sides also dark. Anterior median eyes smaller than the
laterals and than the posterior medians. Area of median eyes clearly
wider behind than in front. The species is readily characterized by
the large size of the tarsus of the palpus and the form of the particularly
large palpal organ. The process of tibia is small, erect, and bidentate,
the teeth being equal. See further Plate XV, Fig. 7. Chelicera rather
small, slightly bulging at base in front, but without a true process or
tooth.

Length, 1.7 mm.

Type—M. C. Z. 408. Washington: Olympia.

Dictynoides gen. nov.

Pars cephalica broad, low, not gibbous. Anterior row of eyes
straight, the eyes equal or the medians smaller. Lateral eyes on each
side subcontiguous. Posterior row of eyes ‘straight, eyes equidistant
or the median farther apart than from the laterals. Area of median eyes
quadrate or trapeziform, wider than long. Clypeus with height much
exceeding the diameter of the anterior eyes. Cheliceree long, with lower
margin armed with two or three teeth. Labium broad, moderately
narrowed distad, extending a little beyond middle of endites. Endites
oblique, inclined. Cribellum entire. Legs without spines.

244 Annals Entomological Society of America |Vol. XII,

Genotype—D. arizonensts sp. nov.

Includes also Lathys trivittata Bks., known from New
Mexico. The genus differs from Lathys in its inclined endites
and much higher clypeus, and from Dictyna in its shorter
labium and lower, less convex, pars cephalica.

Dictynoides arizonensis sp. nov.

Female—Carapace rufous brown along each side, the median and
a marginal stripe on each side paler; along the pars cephalica and
converging caudad are three bands of white hair as in trivittata. Sternum
dusky yellow to nearly black. Legs yellow, not annulate. Abdomen
pale with some darker spots above; clothed with white hair. Anterior
row of eyes straight; eyes equidistant or the medians a little farther
from each other than from the laterals; the medians smaller than the
laterals. Height of clypeus more than twice the diameter of the anterior
median eyes, slanting conspicuously forward from above below. Posterior
row of eyes straight or nearly so, eyes equal in size and obviously larger
than the anterior median, nearly equidistant, the medians slightly
farther apart than from the laterals. Area of median eyes wider behind
than long (cir. 12:10) and obviously wider behind than in front, the
anterior width being equal to the length. Epigynum as shown in
Plate XVI, Fig. 1.

Length, 5 mm. Length of tib. + pat. I, 2.8 .mm.; tib. + pat. IV,
128) mam.

Type—M. C. Z. 3938. Arizona: Huachuca Mts.; Miller
Canyon, elevation 5,600 ft. (W.M. Wheeler, 17th Nov., 1910).

ULOBORID.

Uloborus utahensis sp. nov.

The types of this species, which are not quite fully mature females,
seem readily distinguishable by pronounced differences in coloration in
connection with structural features. Carapace and sternum black or
nearly so, with the usual clothing of white hair. Leg I with femur
and patella entirely black, the tibia black, excepting a narrow ring at
proximal end, and the metatarsus with a narrow black annulus at each
end and two broader ones in the intervening region, the tarsus dark
except at ends. Leg II has the femur black throughout, with no narrow
pale median annulus; metatarsus with two broad dark annuli or these
indistinctly separated, each close to its end of the joint. Femur III
black, with one pale annulus distad of middle and one less distinct
at proximal end; tibia with a pale annulus at proximal end and a sub-
median one; metatarsus pale at proximal end and at middle. Leg IV
with femus having two pale annuli of which the more proximal is com-
monly incomplete below; the other joints as in leg III. The abdomen
somewhat elongated, without any distinct humps above; dark, being
covered with a dense network of black lines, a brown median longi-

1919] Chamberlin: Western Spiders 245

tudinal black stripe above at base; venter blackish with a pale longi-
tudinal line on each side. Posterior row of eyes rather strongly recurved,
the median eyes much farther from each other than from the laterals.
Area of median eyes longer than wide, a little narrower in front than
behind.

Length, 3 mm. Length of cephalothorax, 1 mm. Length of tib.
Odie, lenlona math. TOL bibs -- pate LV, 1.13: nom.

Type—M. C. Z. 476. Utah: Fillmore.

DRASSIDéE.

Drassodes robinsoni sp. nov.

Female—Carapace and sternum yellow of slightly reddish-brown cast.
Legs yellow. Labium and endites brown, pale at tips. Abdomen
yellowish grey, dusky on sides, especially posteriorly; a weakly indicated
hestate outline from base to middle, with very faintly indicated chevron
lines traceable behind it. Cheliceree deep brown. Lower margin of
furrow of chelicera armed with two rather small teeth. Anterior
row of eyes procurved; the eyes subequal or the medians a little the
smaller; median eyes nearly their diameter apart, half as far from the
laterals. Posterior row of eyes procurved; median eyes their larger
radius apart, between once and a half their diameter apart from the
laterals. Tibia I and II armed beneath with a single spine between
middle and distal end. Posterior tibiz armed with a spine above at
base and with one distad of middle. Epigynum as shown in Plate XVI,
Fig. 2.

Length, 9.6 mm. Length of cephalothorax, 3.9 mm. Length of
tib + pat. 1, 3.9 mm.; of tib. + pat. IV, 3.3 mm.

Type—M. C. Z. 418. Utah: Fillmore.

Dedicated to Mr. Joseph Robinson, of Fillmore, who gave
enthusiastic assistance in the collecting of spiders in that
region.

Drassodes gosiutus sp. nov.

Male—This species belongs in the Japidosus group. It is char-
acterized among North American species in wholly lacking a tibial
apophysis on the male palpus and in having the lower margin of the
furrow of the chelicera bearing but a single, noduliform tooth. Carapace,
sternum and legs yellow, the abdomen light grey or whitish. Anterior
row of eyes procurved; median eyes larger than the laterals, about
three-fourths, or somewhat more, their diameter apart and their radius
from the laterals. Posterior row of eyes much longer than the anterior,
obviously procurved; median eyes much larger than the laterals,
subelliptic, somewhat angulate, only about their lesser radius apart,
more than their long diameter from the laterals. Tibiz I and II armed
beneath distad of the middle with a single spine. Metatarsi I and II
armed beneath its base with a single spine. The claw of the chelicera

246 Annals Entomological Society of America |Vol. XII,

long and somewhat sinuous. Upper margin of furrow of chelicera
armed with three teeth of which the most proximal is nearer the median
than is the most distal. Tarsus and bulb of male palpus proportionately
small and slender; the tarsus bearing two long spines on its ventral
face distad of the bulb. See Plate XVI, Fig. 3.

Length, 7.5 mm. Length of cephalothorax, 3.5 mm. Length of
tib. + pat. I, 4 mm.; of tib. + pat. IV, 4mm.

Type—M. C. Z. 389. Utah: Pullmore.

Drassodes melius sp. nov.

Female—Carapace and sternum pale testaceous, the legs more
yellowish. Abdomen above dark brownish grey with a somewhat
obscure, darker, median longitudinal stripe with vaguely indicated
chevron marks behind, the venter somewhat lighter, much so in front
of the epigastric furrow; the epigynum dark chestnut. Anterior row
of eyes procurved; eyes subequal; median eyes nearly their diameter
apart, a little less than their radius from the lateral. Posterior row of
eyes straight; eyes smaller than the anterior ones: and the medians
slightly smaller than the laterals; median eyes a little less than once
and a half their diameter apart and a little more than that distance
from the laterals. Area of median eyes quadrate, as wide in front as
behind. Lower margin of furrow of chelicera armed with two stout
teeth. Tibie I and II each armed beneath with a single submedian
spine. Posterior tibia not armed above at base. Epigynum as shown
in Plate XVI, Fig. 5.

Male—Palpal organ as shown in Plate XVI, Fig. 4

Length of female, 11 mm. Length of cephalothorax, 4.1 mm.
Length of tib. pat. I, 4mm.; of tib. pat. 1V, 5mm.

Length of male, 8.8 mm. Length of cephalothorax, 3.7 mm.
Length of tib. + pat. I, 5mm.; of tib. + pat. IV, also 5 mm.

Type—M. C. Z. 419. Utah: Uintah Mts., 10,000 ft.

Herpyllus oabus sp. nov.

Male—Carapace, legs and sternum yellow, without markings.
Endites whitish across tips. Abdomen yellowish grey, darker above
than beneath, without definite markings. Anterior row of eyes pro-
curved; eyes about their diameter from edge of clypeus; median eyes
larger than the laterals, scarcely their radius apart, closer to the laterals.
Posterior row of eyes slightly procurved; eyes somewhat smaller than
the anterior, the medians smaller than the laterals; median eyes nearly
once and a half their diameter apart, closer to the laterals. Tibia
I armed beneath with two pairs of spines, one pair basal and one sub-
median. Tibia II armed beneath with five spines, there being an unpaired
spine at base, a pair at middle, and a pair of smaller ones at the distal
end. Metatarsi I and II each armed beneath with two pairs of spines.
Male palpus as shown in Plate XVI, Fig. 6.

Length, 4.6 mm. Length of cephalothorax, 2.4 mm. Length of
tib. + pat. I, 2.4 mm.; of tib. + pat. IV, 2.8 mm.

Type—M. C: 2.386. Utah: (Billmore:

1919] Chamberlin: Western Spiders 247

Zelotes tuobus sp. nov.

Female—Carapace black, in part of a dull chestnut cast. Sternum
dense black. Legs black, of slightly brownish cast, excepting the tarsi,
which are paler. Endites blackish, paler across distal end. Abdomen
black or nearly so, a little paler, greyish black, beneath. Anterior row
of eyes strongly procurved, the median eyes lying above the line con-
necting the centers of the laterals; laterals fully twice the diameter of
the medians; the medians about their diameter apart, but very close
to the laterals. Posterior row of eyes. straight; median eyes a little
smaller than the laterals, about three-fourths their diameter apart,
a little farther from the laterals. Quadrangle of median eyes obviously
longer than wide, slightly wider behind than in front. Tibia I and II
unarmed beneath. Metatarsi I and II armed beneath at base with
one pair of spines. Posterior tibia not armed above. Epigynum as
shown in Plate XVI, Fig. 7.

Length, 9.5 mm. Length of cephalothorax, 3.9 mm. Length of
tib. + pat. I, 3.6 mm.; of tib. + pat. IV, 4.2 mm.

Type—M. C. Z. 387. Utah: Fillmore.

This species is very close to Z. ater (Hentz). The epigynum
is proportionately longer with the anterior pockets obviously
closer together and the inner lateral elevated edges also less
widely separated.

PHOLCIDE.

Psilochorus utahensis sp. nov.

Cephalothorax pale yellow. Legs darker yellow. Abdomen grey,
with some darker mottling. Posterior row of eyes straight; median eyes
nearly once and a half their diameter apart, contiguous with lateral
eyes, about their radius from anterior laterals and three-fourths their
diameter from anterior medians. Characterized by having femur I
in the female shorter than femur IV, the reverse of the condition in the
male. Process of chelicera in male large, arising at base and projecting
prominently forward and downward, at tip acutely pointed and curving
mesoventrad, (Plate XVII, Fig. 1). The species is characterized par-
ticularly by the details of structure of the male palpus. In this the femur
is slender from base to a little beyond middle, with the distal end
strongly enlarged; at base on ectal side an elongate tubercle or swelling
and a small process or tubercle from ventral edge at distal end. The
exterior process of the palpal organ straight, narrowest at middle, and
differing from that of pulullus and californie in appearing bifid at distal
end. (Plate XVII, Fig. 2).

Length of female, to4.4mm. Length of tib. + pat. L, 4.75 mm.; of
tib. 4+- pat. IV, 5 mm.

Type—M. C. Z. 451. Utah: Clear Lake; Fillmore; Fill-
more Canyon; Pine Canyon; et cetera.

248 Annals Entomological Society of America [Vol. XII,

LINYPHIIDAE.

Linyphia latescens sp. nov.

Female—Dusky yellow, darker at margin, the eyes edged with
black. Legs also yellow, the femora and patella dusky, the tibiz with
a broad annulus proximad of middle and a narrower, deeper one at
distal end, the metatarsi also similarly annulate. Sternum blackish.
Abdomen grey, with abdomen crossed, excepting at anterior end, by a
series of heavy blackish chevron marks connected by a more obscure
median longitudinal dark liné; sides dusky. Posterior row of eyes
straight, the median eyes a little less than their diameter from the
laterals, nearer to each other. Anterior row of eyes a little procurved,
the medians much smaller than the laterals, less than their diameter
apart, obviously more than their diameter from the laterals. Area of
median eyes as long as wide or very slightly longer.

Male—Coloration as in female. Palpus as shown in Plate XVII,
Fig. 3. Embolus coiled in a circle at distal end.

Length of female, 2.2 mm. Length of cephalothorax, .9 mm.
Length of tib. + pat. I, 1.36 mm.; of tib. + pat. IV, 1.25 mm.

Type—M. C. Z. 429. Utah: Uintah Mts.; Chalk Creek.

Linyphia orinoma sp. nov.

Female—Carapace, legs, cheliceree and endites yellow, without
markings. Sternum and labium dusky or blackish, the color of the
former appearing denser toward edges. Abdomen greyish yellow, with
a few scattered small silvery dots and sometimes a dusky area at base
of spinnerets on each side and occasionally also a dusky area in front of
this; with no trace of dorsal dark markings, such as occur in insignis.
Posterior row of eyes straight, the median eyes slightly nearer to each
other than to the laterals, separated by their diameter or a little more.
Anterior row of eyes straight; median eyes smaller than the laterals,
their radius or less apart, nearly their diameter from the laterals. The
epigynum projects in a straight, finger-like process, which is a little
clavately expanded distad and is rounded at the end, where it has
openings closer together than those of insignis; the dorsal surface just
proximad of the tip having a deep lunate impression with concavity
caudad, the form and position of this impression differing strongly
from that of insignis. (Plate XVII, Fig. 4).

Length, 3.25mm. Length of tib. + pat. I, 2.2 mm.

Type—M. C. Z. 427. Utah: Millard County; Clear Lake.
Also a not fully matured specimen in Uintah Mts. on Chalk
Creek.

Evidently close to insignis from which distinguished by the
uniform presence of silver-colored spots on the abdomen,
with absence of dorsal markings and particularly by the detailed
structure of the epigynum.

1919] Chamberlin: Western Spiders 249

Bathyphantes fillmoranus sp. nov.

Female—Carapace and sternum dusky yellow, the sternum the
darker. Legs yellow. Abdomen above white, broken into closely
arranged spots, a median dorsal line from which oblique lines on each
side extend in an anastomosis. Venter greyish, witha few scattered small
white dots. Posterior row of eyes straight; median eyes less than
their radius apart, and about three-fourths their diameter from the
laterals. Anterior row of eyes straight; medians smaller than laterals,
less than their radius apart, more than their radius but less than their
diameter from the laterals. Clypeus much narrower than length
of area of median eyes. Upper margin of furrow of chelicera with six
teeth, the lower margin with five much smaller ones. For epigynum
see Plate XVII, Fig. 5.

Length, 2.2 mm. Length of cephalothorax, 1 mm. Length of
tib. + pat. I, 1.25 mm.; of tib. + pat. IV, 1.13 mm.

Type— M. C. Z. 425. Utah: Fillmore Canyon.

Grammonota orites sp. nov.

Male—Cerapace dusky yellow-grey, with the eyes edged with black.
Sternum and labium dusky over yellowish to nearly solid black. Legs,
endites and chelicere yellow. Abdomen dusky over a paler background
to nearly solid black; dorsum without definite markings. Posterior
row of eyes slightly procurved, eyes equidistant, separated by nearly
their diameter. Anterior row of eyes straight; median eyes much smaller
than the laterals, hardly their diameter from the latter, nearer to each
other. Clypeus a little wider than the eye area. Pars cephalica rising
behind eye area as usual; crossed by a single median longitudinal series
of sete, which are longest just caudad of the eyes. Upper margin of
furrow of chelicera with four or five large teeth, the lower margin with
several much smaller ones. Tibial apophysis of palpus large, plate-like,
with mesal side produced forward as a one-pointed process which curves
ectad at tip. See Plate XVII, Fig. 8.

Female—The epigynum is shown in Plate XVII, Fig. 7.
Length, 2.3 mm.

aiype—M.,C.-Z; 440; Utah: Uintah Mts:; Chalk Creek,
7,500 ft.
Distinguished from other North American species by the

form of the tibial process of the male palpus and the structure
of the palpal organ.

250 _ Annals Entomological Society of America [Vol. XII,

Grammonota simplex sp. nov.

Male—A smaller and paler species than the preceding one from
which it is at once distinguishable in lacking the conspicuous tibial
apophysis of the palpus. Carapace dusky yellow with the darker
shield-shaped area at posterior end of pars cephalica; the sternum darker.
Legs yellow. Abdomen blackish to grey. Posterior row of eyes straight,
the eyes equidistant. Anterior row of eyes straight, eyes close together,
sometimes appearing nearly equidistant, but most commonly with the
medians nearer to each other than to the laterals; the medians smaller
than the laterals. Tibia of palpus wholly lacking an apophysis at
distal end above. See Plate XVIII, Fig. 1

Female—Epigynum as shown in Plate XVIII, Fig. 2

Length of male, 2mm. Length of cephalothorax, .9 mm. Length
of tib. + pat. I, .§ mm.

Type—M. C. Z. 432. Utah: Uintah Mts.; Chalk Creek,
Toa00 At:

Grammonota obesior sp. nov.

Female—Carapace comparatively low, yellowish of slightly orange
cast with a black marginal line on each side; eyes edged with black.
Sternum dusky. Abdomen blackish, under the lens showing numerous
minute light dots and some light lines at sides of dorsum. ” Byes as in
the preceding species. .Abdomen proportionately broader and higher
than in the preceding forms. Distinguishable by the characteristic
form of the epigynum which is represented in Plate XVII, Fig. 6.

Length, .2mm. Length of cephalothorax, .8 mm.; width, .66 mm.
Length of tib. + pat. I, .8 mm.; of tib. + pat. IV, .9mm.

Type—M. C. Z. 437. Utah: Uintah Mts.; Chalk Creek.

Grammonota fratrella sp. nov.

Female—A smaller species readily distinguishable by the form of the
epigynum. The carapace is slightly dusky over a dilute yellowish
background, the eyes and lateral borders narrowly edged with black.
Sternum very dusky, nearly black. Legs pale yellow. Abdomen
black or nearly so, without definite markings. Posterior row of eyes
straight; median eyes their diameter apart, slightly nearer to the lateral.
Anterior row of eyes a little recurved; median eyes much smaller than
the laterals, scarcely their radius apart, their diameter from the laterals.
Clypeus narrower than eye area. Epigynum as shown in Plate
XVII, Figs 3:

Length, 1.64 mm. Length of cephalothorax, .76 mm. Length of
tib. + pat. IV., .9 mm.

Type—M. C. Z. 443. Utah: Uintah Mts., 8,500 ft.

1819) Chamberlin: Western Spiders 251

Cornicularia monticolens sp. nov.

Female—Carapace dusky yellow to dusky chestnut, the lateral
margins solid black; eyes edged with black. Sternum more blackish
than carapace. Labium and endites also dusky. Legs yellow, often
dusky, with distal ends of coxze beneath often narrowly lined with black.
Abdomen blackish, without definite markings. Posterior row of eyes
straight or slightly recurved, the median eyes farther from each other
than from the laterals. Nearly twice their diameter apart. Anterior
row of eyes straight; median eyes much smaller than laterals, nearer
together, but well separated from the laterals. Area of median eyes
slightly wider than long. Clypeus moderately oblique, impressed
transversely below eyes, slightly higher than length of median eye area.
Epigynum as shown in Plate V, Fig. 4

Male—Anterior end of pars cephalica conspicuously elevated,
rounded, highest just back of anterior median eyes, bulging forward a
little over clypeus which is incurved above its lower edge. A long
auditory hair at distal end of tibia IV. In the palpus the tibial aphysis
is much more slender than ine. g. Cornicularis spiralis, the genotype, and
is slenderly uncate at tip. The embolus is strongly coiled in a flat
spiral, which is not in part covered as it is in spiralis, etc. See further,
Plate XVIII, Fig. 5.

Length of female, 1.7 mm. Length of cephalothorax, .77 mm.
Length of tib. + pat. I, .63 mm.; tib. + pat. IV, .7 mm.

Type—M. C. Z.444. Utah: Uintah Mts.; Chalk Creek.

This species in a group with the eastern C. spiralis Em.
and C. vigilax B., of Europe. It is strongly different from
those species in the form of the tibial apophysis of the male
and in the details of the eye relations, etc.

Diplocephalus wamotsus sp. nov.

Female—Carapace dusky yellow to dusky brown with the frontal
gibbosity paler, often whitish, the sternum somewhat darker than
carapace. Legs yellow. Abdomen grey. Posterior row of eyes more
procurved than typical, but the tangent to anterior edges of its median
eyes intersecting the posterior portion of the laterals; median eyes
nearly their radius apart, not quite twice as far from the laterals.
Anterior row of eyes slightly procurved; the medians much smaller
than the laterals, sub-contiguous with each other, but widely separated
from the laterals. Clypeus about as wide as length of median ocular
area. Area of median eyes much longer than wide. Upper margin
of furrow of chelicera armed. with three well separated, long teeth;
the margin with a close-set series of several very small teeth. Epigynum
as shown in Plate XVIII, Fig. 7

Male—Post-ocular pi iosity broad, with each antero-lateral corner
extended as a rounded protuberance, its median region and a narrower
inter-ocular region in front of it rather densely pilose: posterior median
eyes borne at base of its anterior face. Posterior row of eyes very
strongly procurved; the medians separated by nearly their diameter,

202 Annals Entomological Society of America |Vol. XII,

three times as far from the laterals. The clypeus obviously more
protuberant than in female, as usual. Tibia of palpus with anterior
end extended into a thin, cup-hke plate embracing the base of the tarsus
like a calyx and bearing a slender chitinous process from below its
distal end. See Plate XVIII, Fig. 6.

Length of male, 2.8 mm. Length of cephalothorax, 1.2 mm.;
width, .9 mm. Length of tib. + pat. I, 1 mm.; of tib. + pat. IV,
1.36 mm.

Type—M. C. Z. 446. Utah: Chalk Creek.

Wubana gen. nov.

Posterior row of eyes a little procurved; the median eyes farther
from each other than from the laterals. Anterior row of eyes slightly
procurved; eyes close together, the medians from a decidedly smaller
to nearly as large as the laterals; eyes equidistant or with the medians
nearer to each other than to the laterals. Area of median eyes trapezi-
form, much wider behind than in front, from a little longer than wide to
wider than long (some males). Clypeus as high as eye area. Thoracic
stria obscure. Pars cephalica in male rising behind the eye area;
piligerous in a median longitudinal stripe, with a brush of longer sete
just behind the eyes which may or may not be elevated on a distinct
process, one hair of this brush typically stouter, more spiniform than the
others. Sternum broad, extending between posterior coxe. Male
palpus with patella not enlarged; tibia prolonged in an elongate lobe
of ordinary texture along basal part of ectal side of the tarsus. Legs
long and slender, anterior tarsi shorter than metatarsi, and metatarsi
shorter than tibiz.

Genotype—W . drassoides (Emerton).

In addition to the genotype,. originally described under
Bolyphantes and also later referred to Sphecozone, the new form
described below and apparently also Bolyphantes pacificus
Banks, later referred to Nematogmus, belong in this genus.
The genus differs from Nematogmus in having the posterior
median eyes farther from each other than from the laterals
with the row at most but gently procurved, in its longer legs, in
lacking postocular impressions in the male, in the form of the
tibia of the male palpus and in not having the tarsus of the
latter deeply bifid at the apex. From Sphecozone the genus
differs, e. g., in having the median eyes of the posterior row
farther from each other than from the laterals, in not having
them borne upon the frontal lobe sometimes present in the male,
in lacking the notably long pedicle to the abdomen, and in the
form of the male palpus. I have taken the genotype, W.
drassoides, originally described from Mt. Carmel, in Con-
necticut and until now not since recorded from elsewhere, also
in the Uintah Mts., of Utah, at'an elevation of 7:500 ft.

bo
Or
Oo

1919} Chamberlin: Western Spiders

Wubana retrahens sp. nov.

Male—Cephalothorax and legs dull fulvous. Eyes edged with
black. Abdomen greyish; marked above as in W. drassoides with a
median longitudinal black line at base followed behind by several
transverse dark lines; sides and part of venter dusky. Eyes of anterior
row equidistant, the medians a little smaller than the laterals. Posterior
row of eyes gently procurved; medians scarcely their diameter apart,
nearer to the laterals. Area of median eyes longer than wide. Pili-
gerous stripe extending from caudal end of pars cephalica forward
between eyes, with no special process of elevation behind eyes beneath
the brush of longer hairs. Palpus as shown in Plate XVIII, Fig. 10.

Length near 2 mm.

Type—M. C. Z. 447. Utah: Fillmore.

The male is at once distinguishable from those of W.
drassoides and W. pacifica in lacking the postocular piligerous
eminence. It is notably different from W. drassoides in having
the posterior median eyes much closer together, with the area
of median eyes longer than wide instead of wider than long.
The structure of the male palpal organ is also a feature readily
distinguishing the species.

Hilaira uta sp. nov.

Female—Carapace, sternum and endites dusky over a light brown
ground, less often yellowish, the labium and chelicere darker. Legs
. dusky light brown to yellow. Abdomen grey, more or less dusky, with
no markings. Posterior row of eyes straight; median eyes their diameter
apart, once and a half as far from the laterals. Anterior row of eyes
substraight, being very slightly recurved; medians smaller than the
laterals, less than their radius apart, their diameter from the laterals.
Area of median eyes subequal in length and breadth or slightly longer
(11:10), narrower in front than behind in about ratio 9:10. Clypeus
in height equal to length of median eye area. Epigynum as shown
in Plate XVIII, Fig. 9.

Male—The palpal organ is distinctive. The spinulose submedian
lobe, or apophysis, of the bulb is a notable feature. See Plate XVIII,
Fig. 8.

Length of female, 3.8 mm. Length of cephalothorax, 1.8 mm.;
width, 1.3mm. Length of tibia + patella I, 1.4 mm.; of tibia + patella
IV, 1.7mm. Length of male, 2.8 mm.; of cephalothorax, 1.4 mm.

Type—M. C. Z. 448. Utah: Millard Co.; Clear Lake.

This species is readily recognizable by the form of the epi-
gynum in the female and of the palpal organ in the male.

254 Annals Entomological Society of America [Vol. XII,

ARGIOPID/E.

Aranea (Neoscona) utahana sp. nov.

Female—This species belongs with vertebrata McCook in the group
with longitudinal thoracic groove and having the anterior femora
armed beneath with a double series of numerous stout spines. It
differs from other forms of this group conspicuously in the form of the
abdomen, which is rather broadly triangular-oval instead of elongate
oval. The carapace grey of slightly yellowish cast marked with a con-
spicuous median longitudinal black band with lower border of pars
cephalica also black followed on pars thoracica by one or more obscure
dark spots. Sternum yellowish grey in the middle borders black.
Abdomen grey with a folium on the posterior half limited on each side
by a wavy black line edged with a pale line; just within the limiting line
of folium on each side is a straight dusky line, while on the median
region are two longitudinal dark lines close together; on each side of
folium are numerous dark lines running from its edge ventrad; each
anterolateral corner dark, the dark area crossed by fine vertical pale
lines; midventral region black, the stripe embracing at each border two
white spots, of which the anterior is the larger. Legs light yellow,
deeply banded with black. The femur of anterior legs with a black
submedian annulus and a broader distal one; the patella black, obscurely
divided by a pale median area; the tibia with a black annulus at each
end and one at the middle; and the metatarsus similarly triannulate.
The tibiz of legs III and IV lack the median black annulus as does
metatarsus III, while it is present on metatarsus IV. Hairs of body
in general whitish. The scape of the epigynum is in general similar
to that of vertebrata, but is evenly curved, not at all geniculate. See
Plate XIX, Figs. 1 and 2.

Length, 12 mm. Length of eine ge 8.5 mm.; width across base,

8.2 mm. Length of cephalothorax, 5.2 mm. Length Of tib: s-) patel,
7mm. or tib. ae pat. LV. mata:

os C. Z. 458. Utah:--Fillmore, Webs toundvin
sagebrush on desert flats west of the town. The color of the
spiders blends with the olive grey of the sage-brush.

Aranea tusigia sp. nov.

Male—Carapace light brown above, the lower part of sides blackish
brown with upper border of stripe serrate, a narrow marginal light stripe
below dark area on each side. Sternum black. Legs yellow, marked
with dark annuli; on femora a narrow annulus at base, a second broader
one distad of middle and a third one toward distal end, the extreme distal
end being of a reddish yellow cast. Tibiz black at proximal end, at
least above, and in a much broader annulus at distal end; metatarsi
also dark at both ends, more narrowly so at the proximal one; tarsi
darkened distally. Abdomen with folium above limited by a sinuous

1919] Chamberlin: Western Spiders 255

light line on each side, the folium back of middle solid black and at its
anterior end embracing a short median longitudinal stripe, which widens
caudad; venter with a broad black stripe behind epigynum in each
lateral edge of which is a yellowish spot; stdes marked by fine longi-
tudinal black lines or streaks. Thoracic furrow long, longitudinal,
with no transverse impressions. The first coxa has a stout hook at
distal end, while the second coxa is armed at base with a longer, stout,
conical and nearly straight process. The tibia of leg II is enlarged with
its mesal (anterior) surface convex; it is armed beneath with a double
‘series of long stout spines, the posterior series in the type consisting of
six spines and the anterior of three; the mesal or anterior surface in
addition is armed with numerous spines in two or partially in three series,
most of these spines being distad of the middle and these being much
shorter and stouter than the others, about seventeen spines in all on
the anterior surface. Palpus of male of characteristic form. See
Plate XIX, Fig. 3.

Length, 8 mm. Length of cephalothorax, 4.2 mm. Length of
tib. + pat. I, 5 mm.; of tib. + pat. IV, 4.2 mm.

dvpe—_ ML ©. 2, 46N. Utahe Chalke Creel.

CLUBIONIDA.

Clubiona orinoma sp. nov.

Female—Carapace dull yellow of slightly brownish cast. Sternum
and legs light yellow. Labium and endites brown. Chelicerze light
chestnut. Abdomen beneath light yellowish grey; dorsum reddist
brown over a yellow ground with a narrow median longitudinal reddish
line from base to behind middle, this pointed:at caudal end; sometimes
fine pale chevron lines traceable on posterior region. Lower margin
of furrow of chelicera armed with two moderately stout teeth with
sometimes distad of these a number of granular elevations. Anterior
row of eyes straight; eyes less than their radius from lower margin of
clypeus, equidistant, nearly their diameter apart. Posterior row of
eyes much longer than the anterior with eyes smaller; row straight;
median eyes two and a half times their diameter apart, twice their
diameter from the laterals. Area of median eyes much wider than
long, wider behind than in front in about the ratio 22:15. Tibia I and
II armed beneath with two pairs of spines, the corresponding metatarsi
each with a single basal pair. Epigynum as shown in Plate XIX, Fig. 4.

Length, 6.5 mm. Length of cephalothorax, 2.6 mm. Length of
tib. + pat. I, 2.2 mm.; of tib. + pat. IV, 2.2 mm.

Type—M. C. Z. 474.. Utah: Chalk Creek.

256 Annals Entomological Society of America |Vol. XII,

AGELENID.

Agelena mimoides sp. nov.

Male—Carapace with the usual broad median longitudinal dorsal
yellow stripe and yellow lateral stripes; a marginal dark line on each
side; the lateral dark bands rather narrow, with lower edge uneven;
eyes edged with black. Legs yellow, in type, with some vague darker
markings on femora, but with not distinct annuli. Sternum yellow, a
little deeper at the edges. Labium and endites darker, pale at tips.
Chelicere light chestnut. Abdomen beneath immaculate light greyish
yellow in middle region, an interrupted longitudinal dark line or series
of dark spots toward each side; dorsally a broad longitudinal yellow
stripe bordered with dark, the mesal edge of which is uneven or serrate
and enclosing a vague saggittate outline in basal half. Lower margin
of furrow of chelicera armed with two teeth as in californica and pacifica.
Anterior median eyes smaller than the laterals, less than their radius
apart and from the laterals. Posterior median eyes three-fourths or
scarcely more their diameter apart, their diameter from the laterals.
Tibia I armed beneath with three pairs of spines; tibia II with a single
spine at base and a submedian and apical pair. The palpus of the male
parallels in general structure that of A. pacifica. The most readily
noted differences are in the tibia and its processes. In the dorsal view
the excavation is seen to be more extensive, extending slightly farther
mesad; the limiting posterior ridge presents a distinct process above
which is always lacking in pacifica and in lateral view is seen to con-
tinue evenly out along the lateral apophysis, instead of showing an
abrupt interruption. Other differences are noticeable in the details
of the palpal organ. See Plate XIX, Figs. 5 and 6.

Female—The epigynum of the female is distinguished in having the
posterior region more widely elevated with the lateral spurs weaker and
more widely separated. .

Type—M. C. Z. 464. Utah: Fillmore Canyon.

Cicurina garrina sp. nov.

Female—Carapace, sternum, and legs pale brown, the legs darker
distally, the posterior pair somewhat chestnut. Labium, endites and
chelicerze chestnut. Abdomen immaculate grey. Eyes of anterior row
equidistant, the medians about five-sevenths the diameter of the
laterals. Posterior median eyes equal size to the anterior medians, the
posterior laterals scarcely smaller than the anterior laterals. Posterior
median eyes once and a half their diameter apart, their diameter from
the lateral on each side. Cheliceree strongly geniculate; lower margin
typically with three stout teeth followed by three much smaller ones.
Tibia I armed beneath with three pairs of spines, the anterior side with
three single spines of which the most proximal one is subventral. Meta-
tarsus I with three pairs of spines, the anterior face with two spines.
Tibia II armed beneath with 1-1-2 spines. Epigynum, Plate XIX,
Fig. 9.

1919] Chamberlin: Western Spiders PY

Male—The palpus very similar to that of C. arcuata, but differing
in details.

Length of female,9 mm. Length of cephalothorax, 3.2mm. Length
of tib. + pat. I, 3 mm.; of tib. + pat. IV, 3.1 mm.

Type—M. C. Z. 470.: Utah: - Chalk Creek, elevation
8,000 ft.

A species much resembling the Louisianan C. /udvoiciana
Simon in coloration, proportions and general structure. It
may be distinguished in having the posterior median eyes
much farther apart than from the laterals, instead of these
eyes being equidistant, in having three spines on the anterior
face of tibia I instead of but two; metarsus I with two spines
on anterior side, instead of but one, etc.

Cicurina utahana sp. nov.

Female—A small form with carapace and sternum yellow, the legs
more brownish especially distad. Carapace glabrous or nearly so.
Abdomen grey, immaculate, the epigynum light chestnut. Lower
margin of furrow of chelicera armed with four teeth, a minute fifth one
detectable on one side in the type. Posterior row of eyes straight‘
median eyes smaller than the laterals, nearly their diameter from the
latter, slightly farther from each other; posterior medians obviously
smaller than in arcuata and the spacing of the posterior eyes more
nearly equidistant. Epigynum, Plate XIX, Fig. 8.

Length, 5.2 mm. Length of cephalothorax, 2+mm. Length of
tib. + pat. I, 1.8 mm.

_Type—M. C. Z. 467. Utah: Millard Co.; Pine Canyon.

Cicurina utahana anderis subsp. nov.

Female—This form very much resembles in size and coloration the
typical C. utahana. The epigynum is also very similar, but is pro-
portionately smaller with the intermediate loop of the seminal duct on
each side rising only a little farther forward than the ectal
loop. The types differ from wutahana in the spining of the
anterior legs. Tibia I is armed beneath with three pairs of spines
instead of two pairs, with two spines on the anterior face. Tibia II
bears beneath three spines in line toward the caudal side with a fourth
one paired with the most distal of these at distal end while on the
anterior face is but a single spine; in utahana, on the other hand, there
are but two spines beneath toward the caudal side with one distal spine
not in line with these, while there are two spines on the anterior face.

Length, 6 mm. Length of cephalothorax, 2.1mm. Length of tib.
+ pat. 1, 1.8 mm.; of tib. + pat. IV, 2mm.

Type—M. C. Z. 469. New Mexico: San Geronimo.

258 Annals Entomological Society of America [Vol. XII,

Cicurina idahoana sp. nov.

Female—Carapace and legs yellowish brown. Abdomen cinerous,
without markings. Posterior row of eyes straight; eyes large, the
medians smaller than the laterals; median eyes separated from the
laterals by about their diameter, farther from each other. Anterior
row of eyes procurved; medians smaller than the laterals, their radius’
or less apart, nearer to the laterals. Area of median eyes wider behind
than in front; wider than long. Tibia I armed beneath with three pairs,
of spines and on the anterior face with three single spines of which two
more proximal ones are much nearer to each other as usual. Metatarsus
I armed beneath with three pairs of spines and with one spine on anterior
face proximad of middle. Tibia II and Metatarsus II also armed beneath,
each with three pairs of spines. The species is readily recognizable by
the structure of the epigynum (Plate XIX, Fig. 10).

Length, 7 mm. Length of cephalothorax, 3.1 mm. Length of
tib: spat. 1-3 mm: ob tba pat..L Vir. 2 mam.

Type—M. C. Z. 466. Idaho: Moscow Mts., J. A. Hyslop,
coll., 1910.

LYCOSIDE.

Pardosa utahensis sp. nov.

Female—A small form apparently closely related to P. emertoni
Chamb. It is a much darker species, though with the dorsal markings
corresponding. Carapace with integument brown, with a straight
black stripe along each side, a black marginal line and above it a heavier
black supramarginal line; eye region black; the light stripes clothed
densely with bright white hair, which is a little rufous about eyes.
Sternum black, excepting a pale median region, instead of pale yellow
with simply small marginal black dots. Legs dusky brown. Abdomen
with dorsum black along each side, leaving a pale, caudally narrowing
stripe over entire length, this enclosing anteriorly a dark hastate mark;
venter dark grey; epigynum dark chestnut; light areas clothed in life
with bright white hair. Anterior row of eyes much shorter than the
second, about equalling the distance from center to center of the eyes
of the latter; a little procurved; median eyes obviously more widely
separated than in emertoni, clearly more than their diameter apart and
less than their diameter from the laterals. Eyes of second row once
and a half their diameter apart. Anterior tibize and metatarsi armed
beneath as usual, the first two pairs of spines being very long and slender
with the first widely overlapping the second. Epigynum as shown in
Plate 3GLXe Bigs aie

Length, 6.5 mm, Length of cephalothorax, 2.9 mm. Length of
tib. + pat. 1, 3 mm.; of tib. + pat. IV, 3.6 mm.

Type—M. C. Z. 385. Utah: Chalk Creek, elevation 8,000
ft, August; L9ie7:

1919]

Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.

Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.

Fig.
Fig.
Fig.

Fig.
Fig.
Fig.
Fig.

Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.

Fig.
Fig.
Fig.
Fig.

el ees OM ht Ce

Care Capo oo TO)

Bo Se) Sf ee a Be GUE

mewn re

Chamberlin: Western Spiders 259

EXPLANATION OF PLATES.

PLATE XIV.

Epigynum of Amaurobius nomeus sp. nov.

Male palpus, dorsal view, of the same.

Right chelicera, ectal view, of Dictyna uintana sp. nov.

Epigynum of the same.

Right male palpus, ectal view, of the same.

Left chelicera, of male, ectal view, of Dictyna coloradensis sp. nov.
Epigynum of the same.

Right male palpus, ectal view, of the same.

PLATE XV.

Epigynum of Dictyna vincens sp. nov.

Right male palpus, ectal view, of the same.

Right chelicera, ectal view, of Dictyna bellans sp. nov.
Epigynum of the same.

Right male palpus, ectal view, of the same.

Right male palpus, ectal view, of Dictyna insolens sp. nov.
Right male palpus, ectal view, of Dictyna olympiana sp. nov.
Epigynum of Dictyna sociella sp. nov.

PLATE XVI.

Epigynum of Dictynoides arizonensis sp. nov.
Epigynum of Drassodes robinsoni sp. nov.

Left male palpus, ectal view, of Drassodes gosiutus sp. nov. (All hairs and
spines omitted as usual, excepting ventral distal spines of tarsus.)

Left male palpus, ectal view, of Drassodes melius sp. nov.
Epigynum of the same.

Left male palpus, ectal view, of Heryullus oabus sp. nov.
Epigynum of Zelotes tuobus sp. nov.

PLATE XVII.

Right male chelicera, ectal view, of Psilochorus utahensis sp. nov.
Right male palpus, ectal view, of the same.

Left male palpus, ectal view, of Linyphia latescens sp. nov.
Epigynum of Linyphia orinoma sp. nov.

Epigynum of Bathyphantes fillmorana sp. nov.

Epigynum of Grammonota obesior sp. nov.

Epigynum of Grammonota orites sp. nov.

Right male palpus, ectal view, of the same.

Right male palpus, dorsal view, of the same.

PLATE XVIII.

Left male palpus, ectal view, of Grammonota simplex sp. nov.
Epigynum of the same.

Epigynum of Grammonota fratrella sp. nov.

Epigynum of Cornicularia monticolens sp. nov.

260 Annals Entomological Society of America [Vol. XII,

Fig. 5. Left male palpus, ectal view, of the same.

Fig. 6. Left male palpus, ectal view, of Diplocephalus wamotsus sp. nov.

Fig. 7. Epigynum of the same.

Fig. 8. Left male palpus, ectal view, of Hilaira uta sp. nov.

Fig. 9. Epigynum of the same.

Fig. 10. Right male palpus, dorsal view, of Wubana retrahens sp. nov.
PLATE XIX.

Fig. 1. Epigynum, ventral view, of Aranea utahana sp. nov.

Fig. 2. Epigynum, lateral view, of the same.

Fig. 3. Left male palpus, ectal view, of Aranea tusigia sp. nov.

Fig. 4. Epigynum of Clubiona orinoma sp. nov.

Fig. 5. Right male palpus, ectal view, of A gelena mimoides sp. nov.

Fig. 6. Tarsus of right male palpus, dorsal view, of the same.

IQs Ze Epigynum of A gelena naevia Hentz, variety (Uintah Mts.)

Fig. 8. Epigynum of Cicurina utahana sp. nov.

Fig. 9. Epigynum of Cicurina garrina sp. nov.
Fig. 10. Epigynum of Cicurina idahoana sp. nov.
Fig. 11. Epigynum of Pardosa utahensis sp. nov.

ANNALS E. S. A. VoL. XII, PLATE XIV.

R. V. Chamberlin

ANNALS E. S. A. VoL. XII, PLATE KV.

R. V. Chamberlin

ANNALS E.S. A. VoL. XII, PLATE XVI.

R, V. Chamberlin

ANNALS E. S. A. VOL. XII, PLATE XVII.

R. V. Chamberlin

ANNALS E. S.A. VOL. XII, PLATE XVIII.

R. V. Chamberlin

“ANNALS E.S. A. VoL. XII, PLaTE XIX.

R. V. Chamberlin

NOTICE TO MEMBERS AND CONTRIBUTORS.

The Annals of the Entomological Society of America, pub-
lished by the Society quarterly, includes the Proceedings of the
Annual meetings and such papers as analy be selected by the
Editorial Board.

Papers may be submitted to any member of the Editorial
Board and should be as nearly as possible in the form desired as
final, preferably typewritten, and illustrations must be finished
complete ready for reproduction. Plates must not exceed 5x7
inches unless intended to fold. In general, papers to be accepted
must be original, complete and previously unpublished and,
except in connection with the proceedings, it will not be the
policy to publish preliminary announcements or notes. Authors
will be allowed fifty reprints gratis and additional copies at cost
to the Society.

The Managing Editor is provided with the most recent
address of all paid-up members on record in the Secretary’s
office for mailing the numbers of the Annals and members failing
to receive their numbers should present their complaint to the
Managing Editor within four months from the date of the mail-
ing of the issue. After that time the numbers will be furnished
only at the regular published rate.

The regular annual subscription price for the ANNALS is in
-the United States, Cuba, Porto Rico, Hawaii and Mexico, $3.00;
Canada, $3.50; other countries, $4.00. Checks, drafts or money
order should be drawn payable to ANNALS ENTOMOLOGICAL
SocIETY OF AMERICA, and addressed to HERBERT OSBORN,
State University, Columbus, Ohio, U.S. A.

Requests for information as to membership and the annual
dues of members may be sent to the Secretary-Treasurer,
J. M. Atpricu, National Museum, Washington, D. C.

Communications relating to the ANNALS, and all orders for
separate copies or reprints should be addressed to
, HERBERT OSBORN, Managing Editor,
ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA,
State University, Columbus, Ohio.

CONTENTS OF THIS NUMBER.

WHEELER, WILLIAM M.—The Ants of the Genus

-Metapone Forel............2- NANCE B hawt 8 173

RUCKES, -HERBERT.— Notes on the Male Genital
System in Certain Lepidoptera. ).- 220.222. 2/- ie ds) 4
WeEtLcH, PAauL §S.— The Aquatic Adaptations of
Pyrausta Penitalis Grt. (Lepidoptera)....-....-. 213
FRACKER, S. B.—Chariesterus and its ‘Neotropical |
Relatives (Coreide Heteroptera).-.--...----.-- 227
SanpeErs, J. G., and DeLonc, D. M.— Eight New _
‘‘Jassids’? from the Eastern United States. .--- Logit

CHAMBERLIN, RALPH V.—New Western Spiders.-.. 239

Price List of Publications.
Annals, Vols. I, II, III, IV, V, VI, VII, VII, IX, X and XI complete, each. $3.00

Annals, Separate Parts except as below, cach... ..... ec ee cee ceweeenes 1.00
Annals Vols! iandsDl Part 3 Canes. Si ase vied sea cs wheat uiciadare Sa earn atenns .50
Annals; Vols LV, “Part eaehe icc. PASS artic we be tee ee pihaienien wae aan 1.50

BACK VOLUMES

Of the ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA may
be secured from the office of the Managing Editor and new members
of the Society who may wish to complete a set are advised to secure
the earlier volumes while there is still a supply on hand and the price
fs kept at the original subscription rate.

Address HERBERT OsBorn, Managing Editor,
ANNALS ENTOMOLOGICAL SOCIETY OF AMERICA,
State University, Columbus, Ohio.

Sear
x

DECEMBER, 1919.

4

ie eee CORY
POT SORIA? lost

akan BOARD a Sepa
Oe ase ( JAN26 1990 )
i ee Reet ve "HERBERT OSBORN, Managing Editor, Nw is . ‘ie %
. : irate es a Wh ah cae ‘CoLumBus, OHIO, Rea ntne Sonar yose’>
sti O. ‘HOWARD, RAL 2, BO WM Ay RILEY, 3
Se hl WasHINGToN, DNC: . ee -Minnearouis, MINN.
wy y ‘
_ VERNON TEE LOeG, fe TL PRANK E. LUTZ, }
NORD UNIVERSITY, Canrr. NEw York City, N.Y.
wat M. WHEELER, a oe ‘WM. S. MARSHALL,
cOetORs Mass, Re at MADISON, WIS.
zB M, WALKER, ORD) GS eR ee SANDER Sy
girs {ROMANE CANADA. Pettis the _ HARRISBURG, PA.

PUBLISHED QUARTERLY BY THE SOCIETY
+ BN COLUMBUS, OHIO

_ Entered as aceouid class matter “April 11, 1908, at the Post Office at Columbus, Ohio,
under the Act of Congress of March 3, 1879.

The Entomological Society of America
Founded 1906.

OFFICERS 1919.

President
James G. NEEDHAM . ° ° ° : -, ‘dthaca, NAY

First Vice-President
Justus W. Fotsom . 5 oer te - $ . Urbana, Ill.

Second Vice-President
R. V. CHAMBERLIN . ‘ A 2 - + Cambridge, Mass.

Managing Editor Annals

HERBERT OSBORN . ! : : ° . Columbus, Ohio | |

Secretary-Treasurer
J. M. AtpricH . ; . National Museum, Washington, D. C.

Executive Commitiee

THE OFFICERS

AND
: Lor, ARTHUR GIBSON,

F.E
G. A. DEan, G. C. CRAMPTON.

Committee on Nomenclature
E. P. Fett, T. D. A. CocKERELL, NATHAN BANKS.

Thomas Say Foundation

NATHAN BANKS, A. D. MacGILtiivray, P. P. CALVERT,
E. B. WILLIAMSON,
J. M. Atpricu, Editor, E. D. BALL, Treasurer.

Committee on National Museum

T. D. A. CocKERELL, Chairman,

HERBERT OSBORN, Wma. BARNES, Wm. M. WHEELER,
James G. NEEDHAM, }

ANNALS

OF

The Entomological Society of America

Volume XII DECEMBER, 1919 Number 4

THE TERMINAL ABDOMINAL STRUCTURES OF
ORTHOPTEROID INSECTS: A
PHYLOGENETIC STUDY.

By E. M. Waker, Toronto, Ont.

INTRODUCTION.

There is still much difference of opinion concerning the
inter-relations of the various orders of insects, particularly
of the so-called “‘lower’’ orders, and even the question as to
what are the limits of these orders is by no means a matter of
general agreement.

If these problems are ever to be solved in a logical manner,
all the available data must be taken into account. The evidence
afforded by comparative anatomy, ontogeny and palaeontology,
or any facts bearing upon the subject, must all be fairly con-
sidered. Although much information has been accumulated
from these various sources, there are still important fields
which have received much less attention than they deserve.
In the field of external morphology the greatest advances
have been made in the study of wing-venation, which, thanks
to the classical labors of Comstock and eed now rests
upon a thoroughly sound basis; the mouth-parts have also
long been a favorite subject for investigation, while in com-
paratively recent years, good progress has been made in the
study of the thoracic and cervical sclerites, particularly by
Snodgrass and Crampton.

Our knowledge of the terminal abdominal structures,
especially the genitalia and associated parts, is still, however,
in a very unsatisfactory state. A constantly increasing value
is being attached to them by systematists in separating genera

267

268 | Annals Entomological Society of America [Vol. XII,

and species, but the specialist is seldom interested in these
structures except in so far as they afford good taxonomic
characters in the groups with which he is concerned, and does
not trouble himself to inquire into their homologies with the
corresponding parts in other orders. The natural result of
this is a multiplicity of terms and a great lack of unanimity
in their application.

The most fundamental upheaval of our generally accepted
ideas of insect classification that has occurred in comparatively
recent years is the system advocated by the eminent student
of fossil insects, Anton Handlirsch. This system, which was
first proposed in 1903,! and elaborated in 1908 in his monu-
mental work, ‘‘ Die fossilen Insekten,’’ is well known, and its
deviations from previously accepted views are due mainly
to the study of the fossil record. His division of the old class
Insecta (Hexapoda) into five classes—Collembola, Campo-
deoidea, Protura, Thysanura and Pterygogenea—is-not based
directly upon palaeontological evidence, but on general con-
siderations of structure; but the splitting up of the old order
Orthoptera is founded upon the actual fossil record, so that the
question as to whether or not the findings of comparative
morphology support his views becomes one of considerable
importance.

The present study of the genitalia and associated parts of
the groups commonly called Orthoptera is offered as a con-
tribution to this subject and, at the same time, an effort has
been made to clear up certain general questions on the homolo-
gies of the parts concerned.

Before proceeding with the discussion of the terminal
abdominal structures, it may be useful to give a brief summary
of Handlirsch’s views in so far as they relate to the origin and
relationships of the various groups still commonly known as
Orthoptera, i. e., the Blattide, Mantide, Phasmide, Acridide
and Acrydiide (Tettigide)*, together with such other groups as
may appear to be more or less closely related to them.

The earliest undoubted insect remains belong to the Car-
boniferous era. They are all winged insects of comparatively
large size, some of them very large. Eleven orders are recog-

1 Handlirsch, 1903. Zur Phylogenie der Hexapoden. Vorlaufige Mitteilung.
Sitzb. K. Akad. Wiss., Bd. 112, Heft 8, Abt. 1, pp. 716-738, Taf. 1.

* The Acrydiide of Most Authors.

|
: '
7

1919] Walker: Structure of Orthopteroid I: nsects 269

—_—

nized by Handlirsch as belonging to this period, only one of
which, the Blattoidea, is represented among the orders of the
present age, although four others, viz., the Protorthoptera,
Protoblattoidea, Protodonata and Protephemeroidea, appear
to be the direct forbears of the Orthoptera, Mantoidea, Odonata
and Plectoptera (Ephemerida) respectively.

A large proportion of the insects of this period are char-
acterized by their extremely generalized structure and are
regarded by Handlirsch as the groups from which all other
winged insects, or Pterygogenea, have descended. These are
the Palaeodictyoptera. Their two pairs of ample wings were
similar in size, form and venation, the latter being remarkably
like the hypothetical type on which the Comstock-Needham
system is founded.

The Protoblattoidea and Protorthoptera are independently
connected with the Palaeodictyoptera by forms which differ
very slightly from the latter, while, on the other hand, the most
primitive Blattoidea, such as Polvyctoblatta, grade almost
insensibly into the Protoblattoidea. The latter group consisted
of more elongate forms than the true Blattids, having a less
regularly elliptical outline, and usually a longer prothorax and
a more exposed and prognathous head. Some of them, at
least, had a well-developed exserted ovipositor.

The Protoblattoidea died out in the Permian, where the
first true Mantids appeared, these differing in venation very
little from the former group.

The Blattoidea are considered to be the forbears, not only
of their modern representatives, but also of the Isoptera,
Corrodentia (Psocide), Mallophaga and Siphunculata (Ano-
plura). None of these groups are known before the Tertiary
epoch, and the fossil record offers no clues as to their relation-
ships. Handlirsch is probably correct with respect to the
Isoptera, although they may well have arisen at a much earlier
age than the Cretaceous.

The Protorthoptera, which also persisted into the Permian,
embraced a considerable number of families and genera. They
were elongate forms, some with ambulatory legs and prognathous
phasmid-like heads, while others had saltatorial hind legs like
those of modern Orthoptera. An elongated ovipositor was
present in some, if not all, forms(e. g., Dieconeura arcuata
Scudd.) None possessed stridulatory organs.

270 Annals Entomological Society of America [Vol. XII,

-

No Orthopteroid insects are known from the Trias, the
insect record of which is very scanty, but true Orthoptera
appear in the Lias, belonging to several groups, some of which
were silent while others possessed stridulating organs.

Among the former were the Locustopside, which had
antenne and an ovipositor of the Tettigoniid type, but wing
venation more like that of the Acridoidea, and, like the latter,
lacked the stridulatory apparatus in the male tegmina. True
Acridoidea are known from the lower Tertiary, so they were
probably derived from the Locustopside during Cretaceous
times. The stridulating forms were in part, at least, true
Gryllide, and Handlirsch concludes that from primitive non-
stridulating saltatorial stock two branches arose, one leading
to the common ancestors of the Gryllide and Locustide
(Tettigoniide), the other giving rise to the Eleanide and Locust-
opsidz, from which latter the Acridioidea were evolved. He
regards the Tridactylide as probably derivatives of the
Elcanide, some of which, like Tridactylus, possessed peculiar
lobe-like swimming appendages on their hind tibia.

Other orders which Handlirsch assigns to the Orthopteran
stem are the Dermaptera, ‘‘ Diploglossata’’ and Thysanoptera.
These groups are unknown below the Tertiary epoch, and this
fact has evidently influenced Handlirsch’s judgment in his
attempt to find suitable ancestors for them in the Orthoptera
of the Cretaceous. :

The Ephemerida, Odonata and Plecoptera are considered to
have no direct relationship with each other or with other orders
except through their Palaeodictyopterous ancestors.

Since the publication of ‘‘Die fossilen Insekten,’’ two new
orders of insects have been discovered which must be considered
in any discussion of the phylogeny of the orthopteroid groups.
These are the Zoraptera, represented by a single genus, Zoro-
typus Silvestri (13)? containing five species, and the Gryllo-
blattoidea, likewise represented by one genus, Grylloblatta
Walker (14)? with a single species (G. campodeiformis). The
former group is compared by its author with the Isoptera and
Blattide and also with the Dermaptera (teste Caudell) and

2 Silvestri, Fil., Ballet. Lab. Zool. Gen. Agric. Portici, Vol. VII, pp. 193-209,
Figs. i-xiii (1913).
3 Walker, E. M., Can. Ent., Vol. XLVI, pp. 93-99, Pl. VI (1914).

1919] Walker: Structure of Orthopteroid Insects 271

Crampton ('15)* considers them as probably members of
the ‘‘ Panisoptera,’’ to which the first two named groups belong.
Caudell® likewise finds their nearest allies in the Isoptera.
Unfortunately, I have been unable to obtain specimens of
Zorotypus and can therefore add nothing to what has already
been written concerning it.

The systematic position of Grylloblatta has been discussed
in several papers by Crampton‘, as well as in the original
description by the present writer (loc. cit.). Its extraordinary
synthetic character is indicated by the variety of orders with
which it has been found to have important features in common.

As regards Prof. Crampton’s opinions on the relationships of
the Orthopteroid orders in general, a few words may be said
here. He has expressed views on this subject in several papers
(15, 716, °17, 718, ’19)7 and these have been modified somewhat
from time to time, especially with regard to the position of
Grylloblatta, which will be discussed later. His latest views
appeared in a paper entitled ‘‘Notes on the Phylogeny of the
Orthoptera,’’ and are summarized in his diagram on p. 48 (’19).?
They differ from those of Handlirsch mainly in the following
points:

1. The Isoptera, owing to the possession of certain primitive
characters not found in living Blattids and Mantids, are rep-
resented as arising, not directly from the Blattid branch, but
from the base of the common stem of the Blattide and Mantide,
or possibly somewhat farther along its path of development.

2. The Phasmide arose, not from Saltatorial Orthoptera
(Locustopside), but from near the base of the common
Orthopteran stem, a view which is supported particularly
by the presence of Plecopteroid characters in the primitive
Phasmid, Jimema californica.

a. . Phe Dermaptera are likewise not regarded as of
Orthopteran origin, but are separately derived from the common
stock, from which all the pterygote orders arose.

4 Crampton, G. C. Ent. News, Vol. X XVI, p. 3438 (1915).

5 Caudell, A. N. Can. Ent., Vol. L, p. 381 (1918).

6 Crampton, G. C., Ent. News, Vol. XXX, pp. 42-48, 64-65 (1919). (See also
following footnote.)

7 Crampton, G. C., Ent. News, Vol. XXVI, pp. 337-350, Pl. XIII (1915);
Ent. News, Vol. XXVIL: pp. 244-258, 297-307 (1916): Can. Ent., Vol. XLIX, pp.
213-217, Fig. 9 (1917); Ent. News, Vol. XXVIII, pp. 398-413, PI. XXVII (1917);
Journ. N. Y. Ent. Soc., Vol. XXV, pp. 225-237 (1917); Bull. Brooklyn Ent. Soc.,

Vol. XIII, pp. 49-68, Pls. II-VII (1918); Ent. News, Vol. XXX, pp. 42-48; 64-72
(1919).

272 Annals Entomological Society of America [Vol. XII,

4. The Dermaptera, Embiidina and Plecoptera are grouped
together more closely than is done by Handlirsch, who recog-
nized no near affinities among them.

In other respects Crampton’s diagram is not incompatible
with Handlirsch’s views, so far as it goes, but in regard to the
relationship of other orders not included among the ‘‘Orthop-
teroid’’ groups, but believed by Handlirsch to be derived from
Orthopteroid ancestors (including Blattoid and Protoblattoid
derivatives) his views are very different. These, however,
do not concern us here.

In general Crampton believes that the Plecoptera rather
than the Blattoidea most nearly represent among living insects
the ancestral stock from which the Orthoptera and Phasmoidea
have developed, while Grylloblatta has its closest affinities
among the Mantids, Embids and Dermaptera, and its line
of descent is therefore represented as coming from the ancestral
stock common to the Panisoptera and Panplecoptera. This
view differs somewhat from former views expressed by this same
author, in which he placed Grylloblatta in his super-order
‘‘Panorthoptera,’’ with the Orthoptera and Phasmoidea. The
position of this important annectant form will be further
discussed at a later stage.

THE TERMINAL ABDOMINAL STRUCTURES.

Two papers by Crampton (17 and ’18)® have recently
appeared, in which these structures in the more primitive
orders are discussed from the comparative standpoint: In
the earlier paper, which deals with the female, the author
states that ‘the neck and cervical structures furnish far more
definite characters for grouping these insects than the terminal
abdominal structures of the female do,”’ and in another paper
already cited (19, p. 64), he emphasizes the phylogenetic
importance of the former structures on account of their being
remarkably constant within an order or superorder and less
subject to such variations as depend upon changes of function.
While admitting the general truth of this statement, it should
be kept in mind that such characters as were present in the
common ancestors of all insects may be inherited by some of

8 Crampton, G. C., Jour. N. Y. Ent. Soc., Vol. XXV, No. 4, Pls. XVI, XVII
(1917); Bull. Brooklyn Ent. Soc., Vol. XIII, pp. 49-68, Pls. 2-7 (1918).

1919] Walker: Structure of Orthopteroid Insects 273

the members of any of the orders, and that they may therefore
be of little phylogenetic value except in determining the primitive
form of the structure concerned or in deciding which forms in a
particular group are its most primitive members. Specialized
characters, when the factor of convergence can be eliminated,
are often of more value than primitive ones, particularly in
complex structures where an opportunity for detailed com-
parison is present. For instance, the arrangement of the
mouth-parts of the Diptera or the Lepidoptera is so distinctive
that these structures alone serve as recognition marks of these
orders. It is largely this feature which renders wing-venation
so useful in phylogenetic studies of insects, and although I
should not attribute the same value to the genitalia, I do claim
that they are of great phylogenetic importance when studied
in detail; and in this connection I may point out that Prof.
Crampton’s studies of the thoracic and cervical sclerites are
very much more thorough than those of the genitalia. The
chitinous parts of the genitalia are in large measure internal
structures and unless their internal relations are carefully
investigated one is certain to be led to false deductions.

In another recent paper by Dr. A. G. Newell (’18)* the view
is held that the gonapophyses in both sexes represent three
pairs of serial appendages or limbs, belonging to the Sth, 9th and
10th abdominal segments; and an attempt is made to identify
these appendages in both sexes of all the orders. Although
a useful summary of the literature dealing with the subject
of insect’ genitalia is given, the investigation itself betrays a
lack of grasp of the fundamental principles involved, and the
facts of comparative morphology and development as given in
the bibliography cited appear to have been almost entirely
ignored.

The material on which the present study is based was
obtained from various sources. For the gift or loan of speci-
mens, indispensable to the work, I am especially indebted
to the following gentlemen, to whom I take pleasure in expressing
my most sincere thanks: Prof. G. C. Crampton, Dr. C. Gordon
Hewitt, Dr. N. Banks, Dr. L. O. Howard, Mr. Thos. E. Snyder,
Mr. Morgan Hebard and Mr. W. Downes.

9 Newell, Anna Grace, Annals Ent. Soc. Am., Vol. XI, No. 2, pp. 109-142,
Pls. IV-XVI.

Le)
~I
ws

Annals Entomological Society of America [Vol. XII,

PART I. THE TERMINAL ABDOMINAL STRUCTURES
OF THE FEMALE.

The female external genitalia of a typical generalized
Pterygote insect (e. g., Ceuthophilus, Figs. 1-4), consist of
the vulva or genital aperture, situated at or near the posterior
end of the eighth abdominal sternum’ and usually protected
by a backward prolongation of the latter (in some cases the
seventh sternum), the subgenital plate (st. 8); and three pairs
of processes, the gonapophyses or valvule, which co-operate to
form the ovipositor. These valvule are distinguished, from
their usual positions, as the ventral, dorsal and inner valvule
(valves) or the anterior, lateral and posterior gonapophyses,
respectively. The ventral valvule arise primitively from the
posterior margin of the eighth sternum, though often actually
from the intersternal membrane between segments eight and
nine; the dorsal and inner valvule from the ninth sternum,
primitively also from the posterior margin.

The ventral valvule (vv) consist of a shorter basal segment,
the basivalvula (Crampton, ’17)!° and a longer shaft. The
basivalvula (bs) is usually chitinized only ventrally or ventro-
laterally, if at all, the shaft externally, when a functional
structure, but becoming partly or entirely membranous when
the ovipositor is degenerate.

The dorsal valvule (vd) may be more or less distinctly
separable into a broad proximal portion and a longer, more
slender distal part, but there is no line of demarcation between
these parts and nothing comparable to the basivalvule; the
parts so designated by Crampton being in some cases the
lateral part of the ninth sternum (or valvifer, vide inf.)"4; in
other cases merely the basal part of the valvula itself,!? the
appearance of a suture being due to parts beneath showing
through the valve.

The inner valvule (vi) are enclosed by the other two pairs
and are usually the shortest pair. They commonly enclose
or roof over, the passage through which the eggs are passed out

10 Crampton, Journ. N. Y. Ent. Soc., Vol. XXV, p. 236 (1917). Also termed
“Basalstuck’’ (Van der Weele, Tijd. voor Ent., Deel XLIX, pp. 99-198, Pls. 1-3,
He and ‘‘basal plate’ (Walker, Univ. of Toronto Studies, Biol. Ser., No. 11,

i Crampton, Op, cit. Pl. Sova, Pics ae lepxeValle hiokelt2e

i2'Crampton, op, ecit., Pl. XVI, Figs: 156; Pil; SV Big 10:

1919] Walker: Structure of Orthopteroid Insects 26a

in oviposition. Very frequently the inner valves are connected
from the base distad to a varying extent by a fold of integument,
which may be termed the zntervalvular membrane (im). The
inner valvule are generally chitinized laterally and these
hardened parts or rami (rm) are connected by a strengthening
bar or pons valvularum (p) across the membrane, or the entire
fold may be chitinized dorsally. In the former case it is some-
times convenient to distinguish proximal and distal portions
of the rami according to their position in relation to the pons. —
Like the ventral valvule, the dorsal and inner pair may be

largely or wholly membranous when functionally degenerate.

Between the bases of the dorsal valvule and often closely
connected or even fused with them, is a median sclerite, the
superior intervalvula (sv). It bears a median vertical apodeme
for the attachment of important muscles connected with the
movements of the ovipositor, and its outer surface is continuous
below with the upper surface of the intervalvular membrane. In
some forms, such as Mantis and Stagmomantis, the rami of the
inner valves may be fused with this plate. Another median
sclerite, the inferior intervalvula (iv), is found on the ventral
surface of the base of the intervalvular membrane. With
the inferior intervalvula the rami of the inner valvule
are frequently connected, as in the Tettigoniide and Acridide,
and they are always connected more or less closely with a strong
ventral process from the base of each dorsal valvula, which may
be termed the inferior apophysis (iap). A similar but usually
larger process, the superior apophysis (sap) projects forward
into the hemoccele from the upper part of the base of the
valvula. Both pairs of apophyses serve for muscular attach-
ment.

The dorsal and ventral valvule meet laterally at their
bases, where they are both joined by a small plate, of more or
less triangular form, the valvifer (Crampton) (vf). This
sclerite represents the antero-lateral part of the ninth sternum
and its outer (upper) anterior angle is connected with an
apodeme in the form of a ridge following the constriction
between segments 8 and 9 (ap. 9, int. ap.). The ridge is gen-
erally continued along the front margin of the valvifer and also
often along the lateral margin (using these terms in a morpho-

13 The “‘épimerite’’ of Lacaze-Duthiers. Ann. Sc. Nat., 3 série, Zool., tome
Lipp, 207-251, Pls. 10; 15, 12) (1852):

276 Annals Entomological Society of America [Vol. XII,

logical sense), and near its lower end it is sometimes raised
into a prominent internal process (pap) for the attachment
of muscles. Inthe Acridoidea this process reaches an enormous
size, although the valvifer and ridge are greatly reduced or
absent.

Distal Valvular Connections.

The manner in which the valvule are connected beyond the
base varies greatly, depending upon differences in the methods
of oviposition. As a rule the dorsal and ventral valvule form
a sheath enclosing the inner valvule, the dorsal frequently
overlapping or roofing over both ventral and inner valvule.
Very frequently the inner and ventral valvule fit together
more or less firmly by a tongue-and-groove joint extending
along their adjacent outer edges for the greater part of their
length, and in some cases (most Tettigoniida) such a joint
also occurs between the dorsal and ventral valvule. In both
cases the ventral valvule bear the groove or grooves, the
other valvule the tongues. Where such connections exist
the only possible movements of the valvule in relation to one
another are sliding movements backwards and _ forwards.
These are best seen in the Tettigontide and doubtless also
occur in Grylloblatta. In the Gryllide, in which the inner
valvule are vestigial, the dorsal and ventral pair are immovably
united with one another at the apices and no movement takes
place between them. In the Acridide, Acrydiide and Tri-
dactylide (Ripipteryx), on the other hand, the dorsal and
ventral valvulz are free beyond their bases, and can be moved
in the sagittal plane like a pair of forceps, of which the dorsal
pair form one of the jaws, the ventral pair the other. In the
Phasmoidea, Mantoidea and Blattoidea, the valvule are more
or less flexible and apparently not functional in the forms
studied. There is no close connection between them beyond
the base, although there is usually an ineffective tongue-and-
groove engagement between the inner and ventral valvule.

Development of the Ovipositor.

The first indication of the ovipositor (Fig. 9) is generally
a pair of tubercles on the ninth sternum, sometimes on the hind
margin, but more frequently farther forward. These are soon
followed by two other pairs, one from the hind margin of the
eighth sternum, or just behind it, the other from the ninth

1919] Walker: Structure of Orthopteroid Insects 277

sternum (Fig. 10), between the first pair and sometimes a
little in front of them. The outer pair on segment 9 form the
dorsal valvule, the inner pair the inner valvule, while the
' pair from segment 8 become the ventral valvule. Where
styli occur, they may appear at an earlier stage than the papille,
which are destined to form the valvule, and invariably occupy
the same position as the outer papillae of the ninth sternum,
so that as these elongate into the valvule they carry the styli
with them as small apical appendages. The styli, in all Orthop-
teroid insects, ultimately disappear or become indistinguishable
from the apices of the dorsal valvule, but in many Odonata,
including all the Zygoptera and some of the Anisoptera
(Aeshnine and Petalurine of the Aeshnidz), well-developed
styli persist in the adult females as functional organs.

In early larval life, when the first rudiments of the ovipositor
appear, the abdominal segments are of relatively uniform size,
decreasing in length but little towards the caudal extremity.
In forms with a well-developed ovipositor (Orthoptera, Gryllo-
blattoidea, Mantoidea), however, as development proceeds,
the tenth segment becomes relatively smaller and the ninth
deeper, so that, when maturity is reached, the sternal region of
segment nine is shifted from a horizontal to a nearly vertical
position, thus bringing the bases of the dorsal and inner valvule
into the same transverse plane with those of the ventral valves.
An enlargement of the bases of the valvule usually accompanies
their increase in length and this brings about a crowding out,
as it were, of such portions of the ninth sternum as are not
occupied by the ovipositor itself. Thus, in the adult, the
lateral areas are represented merely by the valvifers, while the
median parts are the intervalvular membrane and the superior
and inferior intervalvule.

In contrast to the reduction of the ninth sternum, the eighth
sternum is usually prolonged caudad as the subgenital plate
covering the vulva and bases of the ventral valvule. In some
groups (Blattoidea, Mantoidea and Isoptera) it is, however,
much reduced and hidden by the greatly developed seventh
sternum, which replaces it as a subgenital plate. In such
cases the eighth sternum becomes more or less completely
dechitinized.

The important question as to the homologies of the valvule
will be deferred until after the consideration of both the male
and female structures in the various groups.

278 Annals Entomological Society of America [Vol. XII,

Orthoptera.
Tettigonoidea (Tettigoniide).

As the ovipositor and associated parts in this group reach
a high state of development and are at the same time of a
remarkably generalized type, it seems best to consider it first.

The ovipositor varies enormously in length, form and
armature, but with these features we need not be concerned.
It will suffice for our purpose to examine one of the more
primitive and one of the more specialized types.

Of the genera studied, we have found the most primitive
types of ovipositor in the cricket-like forms belonging to the
Stenopelmatine and Rhaphidophorine. The well-known “‘stone-
crickets’’ belonging to the genus Ceuthophilus, of the latter sub-
family, illustrate this type well.

In Ceuthophilus (e. g. C. lapidicola Burm.) (Figs. 1-4), the
ovipositor shows comparatively little tendency towards the
pronounced lateral compression or blade-like character of the
more typical green grasshoppers and katydids. It is sub-
cylindrical in the stouter basal part, becoming more com-
pressed distally. The dorsal valvule overlap the other two
pairs, completely concealing the inner valvule, and there is no
tongue-and-groove connection between the dorsal and ventral
valvule, such an engagement existing only between the ventral
and inner valvule. The ventral valvule are closely applied
together, meeting along the mid-ventral line. The basivalvulz
are distinct, though not heavily chitinized and are not wholly
concealed by the rather small subgenital plate. Between the
basivalvula, ninth tergite and base of the dorsal valvula, is
the subtriangular valvifer, the upper angle of which is articu-
lated with the ectal margin of the base of the dorsal valvula.
Its antero-dorsal and antero-ventral margins are raised inter-
nally into strong ridges, (Fig. 4), the latter being continuous
with the intertergal apodeme, which runs for a moderate
distance along the front margin of the ninth tergite and ter-
minates abruptly.

The inner edges of the dorsal valvule meet at a point near
the base, proximad of which they diverge again slightly, but
at the extreme base (excluding the superior apophyses) they are
closely united by the small triangular superior intervalvula,
which, as usual, projects beneath as a median vertical apodeme.

1919] Walker: Structure of Orthopteroid Insects 279

The superior apophyses are triangular processes of average
length. The inferior apophyses curve inwards to meet the
inferior intervalvula, which is a large, thin, sagittiform plate.

The inner valvule are much shorter than the other two
pairs and are somewhat widely overlapped by the dorsal
valvula. The rami .are connected across the well-developed
intervalvular membrane, a little beyond its middle, by the
broad pons valvularum, and are articulated at their bases
with the inferior intervalvula behind the inferior apophyses.
A portion of the proximal division of the rami is separated
from the remainder by a distinct joint. The inner valvule
are apparently supported by a strong chitinous process pro-
jecting inwards from the inner surface of the dorsal valvule
at point about opposite the posterior limit of the intervalvular
membrane.

The ninth and tenth segments are very short and the tenth
tergite, as in all the groups of true Orthoptera, does not form
a complete ring, there being a distinct though unchitinized
sternal area. The eighth spiracle occupies the pleural mem-
brane near the edge of the tergite.

The cerci are long and taper from a stout base to a slender
apex. They are unsegmented, but it may be noted that in
males of some species of the closely allied genus Pristoceuth-
ophilus (e. g. P. cercalis Caudell) a few small segments are
present at the apices of the cerci. Although this is very
probably a caenogenetic character, it is of interest in connection
with the other primitive characters met with in the subfamily
Rhaphidophorine.

As an example of the more specialized type of ovipositor
in the Tettigoniide, we may take Conocephalus fasciatus De
Geer (Figs. 5-8). The ovipositor of this species is straight and
sword-shaped, the valvule being more strongly compressed
and closely coherent than in Ceuthophilus. Both dorsal and
inner valvule engage the ventral valvule by tongue-and-groove ~
joints, the groove in each case running along the edge of the
ventral valvule. The basivalvula is not a distinct plate,
but there is a flattened ventral area, which is evidently its
equivalent. It is but little chitinized, except in its ventro-
lateral edge. The superior intervalvula is larger than in
Ceuthophilus and shows a distinct external pocket or invagina-
tion and a well-developed median apodeme, extending down-

280 Annals Entomological Society of America [Vol. XII,

wards and forwards. It articulates closely with the dorsal
valvule. The pons valvularum occupies the entire distal half
of the intervalvular membrane and is continued cephalad as a
slender median process, which meets the under side of the
superior intervalvula, but does not fuse with it. In some
species, such as Tettigonia- verrucivora, according to Dewitz
('75)44, there is an actual fusion with this plate. The inferior
intervalvula is a slender transverse bar, whose outer ends are
closely articulated with the rami of the inner valvule, and less
intimately with the inferior apophyses of the superior valvule,
which terminate beyond the articulation in an upwardly bent
spine. The dorsal apophyses are larger than in Ceuthophilus,
but not essentially different. The valvifer and intertergal
apodeme do not differ materially from those of Ceuthophilus.

The end segments are more elongate than in this genus,
the tenth segment and anal valves more prominent. ‘The cerci
are shorter and there is a distinct cercal basipodite.

The female genitalia of Ceuthophilus may be regarded as
somewhat more primitive than those of Conocephalus in the
following characters: The distinct basivalvule, the less com-
pressed ovipositor and the less complete cohesion of the valvulz
due to the absence of the tongue-and-groove joint between
the dorsal and ventral valvule. The more elongate and flexible
cerci might also be included, but it is in the males (vide postea)
that they show a decidedly more primitive form than-in Cono-
cephalus.

Finally, it may be added that in the Tettigonoidea in general
the supra-anal plate is not divided transversely into two
sclerites, as is commonly the case in the females of Acridioidea,
i. e., there is no separate eleventh tergite; and it is not always
sharply marked off from the tenth tergite. The paraprocts are
generally but little chitinized.

Grylloidea (Gryllide).

In this group, as in the preceding, the ovipositor is typically
long and slender, but, although sometimes compressed and
ensiform or falcate, it is usually cylindrical and more or less
enlarged apically, the enlarged part armed with teeth. In the
Gryllotalpine or mole-crickets the ovipositor has wholly dis-
appeared in adaptation to the subterranean habits.

14 Dewitz, H., Zeits. wiss. Zool., Bd. XXV, pp. 174-200, Taf. 11, 12 (1875).
(Tettigonia verrucivora referred to as Locusta viridlissima. )

1919] Walker: Structure of Orthopteroid Insects 281

We may take as an example of the family the common field
cricket (Gryllus assimilis Fabr.)

In this species (Figs. 11-15) the eighth and ninth tergites
are very short in the mid-dorsal line, but the ninth expands
considerably on the sides and extends farther ventrad than
the others. The seventh sternum is very large, but is not
produced caudad as a free flap, while the eighth sternum
forms a small subgenital plate, similar to that of the Tetti-
goniide. The eighth spiracle occupies the pleural membrane
close to the tergal margin. The tenth tergite is laterally very
short, but is dorsally longer and fused with the supra-anal
plate, though the line of fusion is visible. The paraprocts are
larger and more heavily chitinized than is usual in the Tetti-
goniidz. The cerci are long, flexible and tapering, with numerous
hairs and sensilla, being very similar to those of Ceuthophilus,
though larger.

The valvifer is very large and is produced under cover of the
subgenital plate into a strong process, which probably represents
a part of the basivalvula. Its inner surface is without strong
marginal ridges, but bears a stout process or apodeme, directed
ento-caudo-dorsad. Its antero-dorsal angle is connected, as
usual, with the ninth tergal apodeme, which is well developed.
There is also a similar but weaker eighth tergal apodeme.

The ventral valvule are without distinct basivalvule, but
these are represented by a well-defined unchitinized area,
covered by the subgenital plate, and probably also by the
ventral processes of the valvifers.

The dorsal valvule are peculiar in the sudden expansion of
their bases, this part forming part of the sternal surface rather
than the valves themselves. Between these bases and partly
fused with them is the relatively large superior intervalvula.
Like that of Conocephalus, its lower part is deeply depressed
and it is also similarly produced cephalad into a thin median
apodeme. The superior and inferior apophyses are well
developed and the latter are connected by a transverse bar,
which represents the inferior intervalvula.

The shafts of the dorsal and ventral valvule fit closely
together by a tongue-and-groove joint and their enlarged
apices are locked together by a peculiar structure, consisting
of a projection from the dorsal valvula, which fits into a socket
in the ventral valvula.

282 Annals Entomological Society of America [Vol. XII,

The inner valvule are extremely vestigial and quite mem-
branous. They project backward from the lower part of
the superior intervalvula. They were overlooked by Lacaze-
Duthiers (loc. cit.), who mistook the dorsal valvule for them,
owing to their close connection with the inferior intervalvula,
which is suggestive of the connection in many Tettigontide
between this plate and the rami of the inner valvule.

In Oecanthus (Figs. 16-18, Oe. quadripunctatus Beut.) the
genitalia do not differ from those of Gryllus except in com-
paratively unimportant details. The most striking feature is
the exceedingly powerful ninth tergal apodeme, which forms a
complete arch around the base of the tergite; and the otherwise
feebly chitinized dorsal surface of segments 9 and 10. These
features are doubtless related to the habit of the genus of ovi-
positing in woody stems, which would require a greater develop-
ment of the muscles of the ovipositor than is necessary in such
eround-dwelling species as Gryllus assimilis.

The dorsal and ventral apophyses and the ventral pro-
longation of the valvifer are also longer than in Gryllus.

The general proportions of the abdominal segments and
terminal structures are somewhat suggestive of some Tetti-
gontide and of Grylloblatta, but there are no indications of
relationship with the latter form, as has been more than once
suggested by Crampton, who has been apparently influenced
by superficial resemblances of form. As far as the terminal
abdominal structures are concerned, Oecanthus is more special-
ized than Gryllus, and if anything even less like Grylloblatta
than the latter genus is.

Acridoidea (Acridide, Acrydiide).

The appearance of the ovipositor in this superfamily is
singularly different from that of the two preceding groups, but
the fundamental similarity of its structure in all three was
demonstrated long ago by Lacaze-Duthiers (’52, loc. cit.) and
Graber (’70)5. Its peculiarities in the Acridoidea are chiefly
modifications connected with the different method of oviposi-
tion. The dorsal and ventral valvule function as a pair of
forceps for digging the hole, usually in the ground, in which the
eggs are deposited. Each pair of valves acts as one of the

18 Graber, V., Sitzb. K. Akad. Wiss., Bd. LXI, Abtl. 1, pp. 597-616, 1 Pl. (1870).

1919} Walker: Structure of Orthopteroid Insects 283

blades of the forceps, the movements being in the sagittal
plane. The inner valvule take no part in this process, but serve
to roof over the path along which the eggs are passed out.

As a type of the group we may take the common two-
striped locust (Melanoplus bivittatus Say.) (Figs. 22-25). In
this species the eighth tergite is well developed, but the ninth
and tenth are not only greatly shortened, but are fused laterally,
the intersegmental furrow being imperfectly developed. The
supra-anal plate is divided by a transverse suture, the basal
part probably representing the eleventh tergite, while the
apical part is the tergite of the anal segment or telson. There
is also a distinct cercal basipodite, which may represent a
separated lateral portion of the eleventh tergite. The cerci
themselves are very short and unsegmented. The paraprocts
are broad, flat and well chitinized, and are not completely
separated from the tenth tergite at the lateral margin.

The abdominal tergites are in reality pleuro-tergites, since
they bear the spiracles, a short distance from their lateral
margins. This feature is more pronounced in the Acridide
than in other families of Orthoptera. The eighth sternum
forms a large subgenital plate and terminates in a papilliform
process called the ‘“‘egg-guide,’’ another characteristic Acridian
structure.

The ventral valvule are short and very stout at base, but
taper to a slender decurved hook-like apex. They are peculiar
in that the tegument is divided into five distinct sclerites, of
which the lateral and the three ventral ones constitute the
basivalvula, while the long dorsal one represents the shaft.
This is evident by comparison with Acrydium in which there
are but three sclerites. The dorsal valvule are of about the
same length as the ventral, and the apices are likewise hook-
like, but curved upwards. They are narrowly separated at
base, the interval being occupied by the small superior inter-
valvula, which lacks the usual median apodeme, but has two
slight prominences for muscular attachments.

. The superior apophyses are represented only by thin flat
tendons of the powerful elevator muscles of the dorsal valvule.
The inferior apophyses (Fig. 24), on the other hand, are stout
processes which bend inwards from the base of the valvule
to articulate with the inferior intervalvula. This plate has the
form of a curved bar, from the concave side of which a bifurcate

284 Annals Entomological Society of America [Vol. XII,

process arises. The inferior apophyses join the ends of the
bar, while the arms of the bifurcate process are connected with
the rami of the inner valves. These connections are thus quite
similar to those of the Tettigoniide (cf. Ceuthophilus).

The inner valvule are very short, but not vestigial as iu
the Gryllide. They are connected almost to their apices by
the intervalvular membrane, the rami being united distally
by a V-shaped pons valvularum.

Extending cephalad from the basal articulation of the dorsal
and ventral valvulz into the body-cavity as far as the seventh
segment is a somewhat flattened and heavily chitinized rod
(pap), which is entirely concealed in an external view. This
structure appears as though grooved externally, but the groove
is covered over with thin chitin, so that it is really a tubular
invagination. It serves for the origin of the most powerful
muscles of the valvule, i. e., the elevators of the dorsal and
depressors of the ventral valvule. Its morphological nature
has been the subject of differences of opinion. Lacaze-Duthiers
(loc. cit.) interpreted it as the ‘“‘épimerite’’ (valvifer), while
Graber (loc. cit.) thought it represented the superior apophysis
of the dorsal valvule. The latter interpretation is obviously
incorrect; the former is much nearer the truth. The muscles,
which in other Orthoptera arise from the ninth tergal apodeme
and its vicinity and are inserted into the superior apophyses,
take their origin in the Acridide from this rod, the apodeme
being undeveloped in its usual situation. It will be remembered
that this apodeme is continuous with that of the lower edge of
the valvifer in the Tettigoniide and other groups, and that it
tends to be best developed towards its lower end, where it is
sometimes raised into a distinct process (e. g. Diapheromera,
Stagmomantis). As the muscles which usually connect the
valvifer with the superior intervalvula arise in Melanoplus
also from this process, it appears that the latter represents in
part the lower end of the tergal apodeme and in part the
valvifer. That it does not represent the main body of the latter
is clearly shown by an examination of both nymph and adult
of one of the small grouse-locusts (Acrydiide, more commonly
known as Tettigine). Fig. 28 represents the end-segments
of the female nymph of Acrydium ornatum Say. Here we find
a perfectly typical valvifer of considerable size. In Fig. 26,
which is taken from an adult Acrydium, the valvifer is also

1919] Walker: Structure of Orthopteroid Insects 285

present, but is very small and sub-chitinized. The characteristic
rod (pap) is fully developed and is separate from the valvifer,
but its base of attachment is in very close relation to the latter;
in fact, in an internal view of the nymph, they appear to be in
contact with one another.

The first appearance of this rod in the nymph of Melanoplus
is a slight invagination of the soft chitin at the angle between
the valvular bases. This is later continued cephalad as a thin
band-like tube, as seen in the last nymphal stage, assuming its
final form only at the last moult. No external appearance of
the valvifer is present in Melanoplus.

In respect of the presence of a distinct valvifer in the
Acrydiidze we may consider the latter to be a more primitive
family than the Acridide and this view is borne out by other
features of the terminal segments: In the nymph of Acrydium,
e. g., the ventral valvule possess unmistakable basivalvule
(bs), which, however, in the adult became divided into two
sclerites, a lateral and a ventral, the latter representing the
two principal ventral sclerites in Melanoplus. The ninth and
tenth tergites are quite separate and less reduced than in
Melanoplus, especially in the nymph. The tenth tergite
(Fig. 29) is imperfectly divided into median and lateral regions,
of which the former is less heavily chitinized and is produced
caudad into a lobe which divides the eleventh tergite into two
separate lateral plates. The supra-anal plate also shows two
lateral chitinized areas. In the nymph (Fig. 29), the eighth and
ninth tergites are unmodified, but in the adult they also present
a narrow, less densely chitinized median region, which folds
inwards, allowing the lateral parts to come together in a pointed
or Gothic arch.

These peculiar features are very suggestive of the Tri-
dactylide, which also resemble the Acrydiide in some other
respects. The cerci are divided into a stouter basal part,
which is hairy, and a slender smooth apical part. This gives
them the appearance as though composed of two fused seg-
ments, a feature which is also suggestive of some Tridactylide
(dere):

In other respects the female genitalia closely resemble
those of the Acridide, the larger size of the superior intervalvula
in Acrydium as compared with that of Melanoplus being perhaps
the most striking difference noted.

286 Annals Entomological Society of America * [Vol. XII,

The eighth spiracle, as in other Acridioidea, is situated in,
the tergite (pleuro-tergite), but the others retain a more
primitive position in the pleural membrane.

Tridactyloidea (Tridactylide).

This small family has been commonly associated with the
Gryllide, owing to the superficial resemblance of the typical
genus, TJvridactylus, to the mole-crickets (Gryllotalpine), a
resemblance due to the subterranean habits of both. Thus we
find in both Tridactylus and Gryllotalpa a similar form of head
and pronotum, short antenne, front legs adapted for digging
and reduction of the ovipositor, which in Tridactylus 1s vestigial,
while in Gryllotalpa it is entirely wanting. In the Tridactylid
genus, Ripipteryx, however, which is not a distinctly sub-
terranean form, there is a well-developed ovipositor, and it is
not at all like that of a Gryllid, but is remarkably similar to
that of the Acridoidea, as suggested by de Saussure and
Zehntner (’94).16 These authors, however, failed to appreciate
fully this resemblance and made no comparisons between
the two groups.

I have examined the terminal segments and ovipositor of
Ripipteryx forcipata Sauss. (Figs. 19-21), and find the latter to
be closely comparable with that of the Acridoidea, but with
certain important features of its own.

The obliquity of tergites 9-11, which is slightly indicated
in Acrydium and associated with the weakening of the median
region and a breaking up of the dorsum into a number of second-
ary sclerites, occurs in the Tridactylidz in an exaggerated form.
In R. forcipata the eighth tergite is normal in shape and form
except that it is divided along the mid-dorsal line into two
lateral plates, the thin tegument between the plates being folded
inwards. This is also the case in a slighter degree in Acrydium.
The ninth tergite is also completely divided into two lateral
plates, but the obliquity has been carried so far that the original
anterior, ventral and posterior margins are respectively ventral,
posterior and dorsal in position. The antero-dorsal angles of
these plates extend forward underneath the eighth tergite
as two slender processes, which are apparently all there is to
represent the dorsal region. The tenth tergite lies dorsal to

16 Rev. Suisse de Zool., tome II, Fasc. 2, pp. 403-430, Pls. XVI, XVII (1894).

1919] Walker: Structure of Orthopteroid Insects 287

the ninth, and consists of two narrow lateral plates prolonged
forward into a pair of slender, curved bars, which pass between
the similar processes of the ninth tergite, under cover of the
eighth tergite, but meet in the middle line between the two
main parts of the latter and three other small plates which
apparently represent the median elongated portion of this
tergite in Acrydium. ‘The eleventh tergite is possibly rep-
resented by the two small oblique plates at the base of the
supra-anal plate, and two others laterad of these at the bases
of the cerci (cercal basipodites). These relations are also
similar to- those of the Acrydium. The cerci are short,
cylindrical and unsegmented, although in Tridactylus they are
distinctly two-segmented, and in certain other species of
Ripipteryx (R. mexicana Sauss.) they are imperfectly divided
into a number of secondary segments (de Saussure & Zehntner,
lec: cit.):

The supra-anal plate is similar in form to that of Acrydium
but is semi-membranous and undivided. It varies, however,
considerably in form in different species of the family, as do also
the subdivisions of the abdominal tergites, so that it is ques-
tionable to what extent these parts are really homologous to
those of the Acrydiide.

The paraprocts are of very remarkable form, being greatly
elongated, with a terminal segment like a pair of styli or a
second pair of cerci. These, however, have nothing to do with
true styli, for which they were mistaken by Crampton (’18)!’;
for the styli, when present in female insects, are always borne
by the dorsal valvulz or their homologues, and thus belong to
the ninth segment. These peculiar structures, which are present
in both sexes in Tridactylus as well as Ripipteryx, were
correctly interpreted by de Saussure and Zehntner as out-
growths of the paraprocts.

Another feature possessed by Ripipteryx in common with
Acrydium is the position of the eighth spiracle in the eighth
tergite, and the other spiracles in the pleural membranes.

Turning to the sternal region, we find that, except in the
case of the eighth, the sternites overlap the tergites instead of
vice-versa. The eighth sternum forms the subgenital plate,
but is not specially modified for the purpose and lacks an egg-

17 Bull.-Brooklyn Ent. Soc., Vol. XIII, No. 7, Pl. V, Fig. 48, 1918.

288 Annals Entomological Society of America (Vol. XII,

guide. The basivalvule are of enormous size and have the
appearance of a divided subgenital plate, for which they were
mistaken by de Saussure and Zehntner.

The ovipositor as already noted, is essentially Acridian in
type. The dorsal and ventral valvule are sigmoid in form,
the apices of the dorsal valvule curving upwards, the ventral
pair terminating in sharp, decurved hooks, and having also
a shorter external hook (cf. Melanoplus, Figs. 22, 23). In
de Saussure and Zehntner’s Figure (1. c., Pl. XVII, Fig. 27)
what is evidently the same hook is represented as belonging to
the dorsal valvulz, but this is certainly an error, due to the
valvule having been tightly closed in the specimen drawn.
As in most Acridoidea there is no trace of the valvifer, while
the free chitinous rod (pap) is represented by a prominent,
shelf-like apodeme (ap 9), extending from the angle between
the valvule along the ventral edge of the ninth tergite and
projecting a short distance under the eighth. This ridge is
quite similar in relation to the valvule to the free rod of the
Acridoidea, but is a true ninth tergal apodeme, like that of the
Gryllide and Tettigoniide. It is in this feature that the
genitalia of the Tridactylide differ most from those of the
Acridoidea.

The dorsal valvule, as in the Acridoidea, lack distinct
superior apophyses. The superior and inferior intervalvule
have precisely the same positions as in, that group, but are
even smaller and simpler in form. The inferior intervalvula
is a slender transverse bar resembling that of Gryllus in form
and is connected at its outer ends, separately, with both the
small inferior apophysis and the rami of the inner valvule.
The latter are, as in the Acridoidea, very small, but lack the
intervalvular membrane and pons.

There are many other points of resemblance between the
Tridactylide and Acridoidea, and taking all these into con-
sideration, we can hardly doubt that the nearest relations of the
former are the Acridoidea rather than the Gryllide.

1919} Walker: Structure of Orthopteroid Insects 289

Grylloblattoidea (Grylloblattide).

The female genitalia of the Orthoptera are most nearly
approached by those of Grylloblatta, so that we may consider
this form next.

A glance at the end-segments and ovipositor of Grylloblatta
(Figs. 31-85) at once recalls the Orthopterous families Gryllidz
and Tettigoniide, except in the segmented cerci, which are
more like those of certain Plecoptera, or to a less extent, the
Mantide. The form of the eighth, ninth and tenth segments
is very like that of Conocephalus, though more depressed and the
tenth segment more prominent. The eighth sternum is not
flap-lke, but quite generalized in form, although a median pale
or thin area is present, which marks the small cavity into which
the vagina opens.

The ninth tergite (Fig. 31) is somewhat prolonged ventrad
as in Gryllus, the ventro-cephalic margin being oblique, as is
usual in the Tettigoniide and Gryllide. The tenth tergite in
the adult female is fused with the sternite forming a complete
ring, as in many Plecoptera. Its dorsum is shghtly prolonged
behind the bases of the cerci, but does not overlap the supra-
anal plate, which is quite distinct, though small. Both supra-
anal plate and paraprocts, which are subchitinized, are quite
like those of many Tettigoniide. The spiracles, which are
very small, are all situated in the pleural membrane.

In the external form of the valvule and their connections
with neighboring sclerites Grylloblatta is remarkably like a
primitive Tettigoniid, such as Ceuthophilus. The most con-
spicuous, but not the most important difference is in the
basivalvule, which are more clearly defined and more heavily
chitinized, recalling those of the Mantids. The valvifer has
exactly the same relations as in the Tettigonide. Its lower
angle articulates with -the ecto-basal angle of the ventral
valvula, not being prolonged under the subgenital plate as in
Gryllus and Oecanthus. There are well-marked internal ridges.
along its upper and lower margins, the latter being continuous
with the ninth tergal apodeme, all these features recalling
Ceuthophilus or Conocephalus strongly.

The ovipositor is much like that of Ceuthophilus or related
genera, though the valvule are less compressed and not so
closely applied to one another, the ventral ones, particularly,

290 Annals Entomological Society of America [Vol. XII,

being slightly separated from one another along the mid-ventral
line and from the dorsal valvule along their entire length.
The dorsal valvule do not overlap the ventral valvule on the
sides, so that the inner valvule are not concealed as in the
Tettigoniide. The inner valvule are a little shorter than the
ventral, with which they are connected by a tongue-and-groove
joint of the same character as in Ceuthophilus, i. e., there is a
narrow groove with raised edges along the upper margin of the
ventral valves, into which fits a ridge along the lower margin of
the inner valves.

In order to study the inner connections of the valvule the
abdomen of the original paratype specimen was cut across at
the base of segment seven, and the posterior segments treated
with potash.

Fig. 32 shows the ovipositor in dorsal view, with the left
ventral valve forcibly turned outwards; the right valve is
omitted. The dorsal valves are separate from one another
except near the base, where they meet at little more than
a point, the inner margins then diverging again. The superior
apophyses are very like those of Ceuthophilus (Fig. 1) and the
articulations with the valvifer quite similar. The inferior
apophyses are prolonged inwards into a pair of slender curved
bars, which meet one another in the middle line. These evi-
dently represent the inferior intervalvula, but are in no way
distinct from the apophyses.

The inner valvule fit closely into the concavities of the
dorsal valvule, there being no space between the two pairs
such as we usually find (cf. Ceuthophilus). The intervalvular
membrane is well-developed as is also the pons which unites
the rami just beneath the point where the dorsal valvule meet.
The median part of the pons projects below from the upper
surface, as a bilobed prominence (Fig. 35, pp), which recalls
the transverse thickened portion of the superior intervalvula
in the Mantids (Figs. 35, 49). No median apodeme could be
found, however, and it is to this apodeme that the muscles
arising from the valvifers are attached in the Mantide, Tetti-
gonide and Gryllide. This apodeme, when present, is situated
between the valvifers, the muscles running transversely and
if such an apodeme were present in this position in Grylloblatta it
would lie considerably in front of the bilobed prominence.

1919] Walker: Structure of Orthopteroid Insects 291

The fact that the valvifers are well chitinized and bear the
usual ridges for muscular attachment seems to indicate, how-
ever, that the muscles in question are probably present and yet
no sclerite for their inner attachment was found, unless it be
the bilobed prominence. As compared with Ceuthophilus
this prominence would appear to belong to the pons, but in
any case it seems probable that the pons was primitively not
distinct from the superior intervalvula, as is indeed the case
in such forms as the Mantids where no distinct intervalvular
membrane is developed. With the great development of this
membrane in the Tettigoniide, however, the pons became
differentiated, retaining its connection with the superior
intervalvula in some forms (e. g., Tettigonia) while becoming
entirely separated from it in others, such as Ceuthophilus.
Although the matter requires further investigation, it may be
considered probable that the superior intervalvula is at least
partly represented by the bilobed prominence beneath the
pons. If this is the case, Grylloblatta differs from all the other
forms studied in which the superior intervalvula is present in
that this sclerite does not connect the bases of the dorsal
valvule, but lies beneath them.

The rami of the inner valvule are fused with the inferior
apophyses which, as stated above, is not distinct from the
inferior intervalvula. This firm union of the dorsal and inner
valves, together with the manner in which the latter fit closely
within the former, would seem to prohibit any independent
movements of the inner valvule.

A short distance behind the united inferior apophyses is
the opening of the receptaculum seminis, a small, thick-walled
rounded pouch.

The female nymph—(Figs. 36-38). The only known speci-
men of the female nymph was recently described by the writer
(19)!8 and is nearly full grown. The eighth sternum is similar
to that of the adult, except that it is uniformly, though feebly,
chitinized. The ventral valvule arise just behind the sternal
margin and are straight, subcylindrical processes, bluntly
pointed at the apices. The basivalvule are clearly defined by
a transverse groove. The sternal region of segment nine still
has its primitive horizontal position in line with that of segment

18 Walker, E. M., Can. Ent., Vol. LI, 11, pp. 131-189. 1919.

292 Annals Entomological Society of America [Vol. XII,

ten. Across its middle arise the dorsal and inner valves in the
same transverse plane. They are likewise of subcylindrical
form, the inner valvule pointed like the ventral pair, while the
dorsal valvule have well-developed styli upon their apices,
a primitive feature, unlike any of the Ortheptera at so late a
stage. Judging by the comparative lengths of the valvule
the styli evidently go to form the apices of the dorsal valvule.
There is no trace of the intervalvular membrane, this structure
being doubtless formed from the median part of the sternal
area, both before and behind the bases of the inner valvule.
The remaining parts of the sternum go to form the valvifers
(antero-lateral region) and the broad bases of the dorsal valvule
(postero-lateral region).

It will be seen that at this stage the ovipositor of Gryllo-
blatta is distinctly more primitive in character than that of a
Tettigoniid nymph of corresponding stage. This is seen in the
form and position of the valvule and in the presence of styh.
In these respects they approach the Blattide and Mantide,
but the valvule of the ninth segment have not the terminal
position, nor have the dorsal valvule the broad, flattened form
found in these groups. In these respects the Blattids and
Mantids are the more primitive and Grylloblatta occupies a
position between them and the Tettigontde.

Phasmoidea.

In this well circumscribed order the ovipositor is of a some-
what primitive type and is, at the same time, more or less degen-
erate and probably functionless in most cases. It is generally
largely concealed by a hood-like subgenital plate, which as in
the Orthoptera, is the eighth sternum. The end-segments
(8 to 10) show no tendency to become abbreviated and the
sternal surface of segment nine retains its primitive horizontal .
position in the same plane with that of segment ten. The
abdominal spiracles are all situated in the pleural membrane.
The cerci are always relatively short and unsegmented and the
anal valves (supra-anal plate and paraprocts), though promi-
nent, are not usually large nor densely chitinized.

In the remarkable little Phasmid, Timema_ californica
Scudd. (Figs 39, 40), the genitalia appear to be of very primitive
form, but they may perhaps be more correctly described as of

1919] Walker: Structure of Orthopteroid Insects 293

a “larval” type, i. e., they are probably to be looked upon as
degenerate, functionless structures, whose development has
been arrested at an early stage, as is not uncommonly the case
in degenerate structures. The eighth sternum is but little
produced over the bases of the ventral valvule, which are
slender and flexible with broader basivalvule. The ninth
segment is shorter than usual in this order, but the sternum
is horizontal, its lateral parts, the valvifers, being larger and
having a larval position in relation to the dorsal valves of the
ninth tergite. They were erroneously regarded as basal seg-
ments of the dorsal valvule (basivalvula) by Crampton. The
dorsal valvule are flexible, triangular flaps, whose bases meet
the inner edges of the valvifers exactly. The term ‘‘dorsal’’
is here inappropriate, as they retain the larval position external
to the inner valves, with which they are fused to within a short
distance of the apices of the latter. Neither dorsal nor inner
valvule are connected with the ventral pair. Having seen
only one specimen of Timema, which was kindly lent by Prof.
Crampton, I have not studied the internal relations of the
valvule.

The tenth tergite is rather long dorsally, but narrowed
ventro-laterally. Its apex is somewhat produced over the supra-
anal plate, which like the paraprocts, is rather long and pointed,
though but thinly chitinized. The cerci are large and stout.

A less primitive ovipositor, but more typical of the order,
is that of the common stick-insect, Diapheromera femorata Say.
(Figs. 41-43). The valvule of this species are of considerable
size, but quite soft and flexible. They are flattened dorso-
ventrally and arched towards the anus. The inner valvule
lie as in Timema in the primitive position between the dorsal
valvule (although in some genera they lie dorsal to the latter),
with which they are united in a similar manner. This con-
nection seems to be characteristic of the Phasmoids and it was
for this reason that the inner valvule were mistaken by Lacaze-
Duthiers (’52, loc. cit.) for a part of the dorsal valvule. There
is little development of the intervalvular membrane and the
rami are represented only by a short chitinous process on each
side, at the angle of union with the dorsal valvule, these pro-
cesses ending freely, there being no trace of either superior or
inferior intervalvule. The underside of the inner valvule
bears a very distinct though flexible ridge (Fig. 43), which fits

294 Annals Entomological Society of America [Vol. XII,

in a corresponding groove in the ventral valvule (Fig. 41),
but as the parts are soft no union is thus obtained. The dorsal
valvule do not lie so flat as in Timema, but the primitive widely
separated position is retained, their bases not becoming arched
dorsad over the inner valvule. Their only strongly chitinized
parts are the superior apophyses which are well developed and
a small process just above the angle with the ventral valvule,
and apparently representing the inferior apophyses. There is
no trace whatever of the valvifer, but there is a small process:
(pap) which seems to belong to the lower extremity of the ninth
tergal apodeme. Both valvifer and apodeme, however, may be
well developed in the Phasmoidea as shown by Lacaze-Duthiers
in the genus Acrophylla. Their absence in Diapheromera is a
degenerative feature. The basivalvule are also absent in this
genus. The subgenital plate is of large size, but much smaller
than in many other genera. The tenth segment is large, with
an extensive sternal area, but the supra-anal plate and para-
procts, though prominent are much smaller than in Tamema.
The cerci are also relatively smaller.

There is considerable variation in the Phasmids in the size
of the subgenital plate, the relative lengths, form and position
of the valvule, but a firm, coherent structure, such as the
ovipositor of the Orthoptera and Grylloblattoidea, does not,
so far as we are aware, occur in this order. The apparently
primitive form of the Phasmoid ovipositor may be largely due,
as already suggested in the case of Timema, to imperfect or
arrested development, but at the same time, it is probable that
the Phasmoid ovipositor never reached a high degree of develop-
ment.

Mantoidea (Mantide).

Of the single family comprising this order two species were
studied, Stagmomantis carolina L. and Mantis religiosa L.
They are so much alike in the parts with which we are con-
cerned, that we may confine our attention in the main to the
former, this having been the form which was most thoroughly
studied.

In Stagmomantis (Figs. 44-49) there is a marked abbreviation
of the eighth and ninth tergites, which also extend a shorter
distance ventrad on the sides than the seventh and preceding

1919] Walker: Structure of Orthopteroid Insects 295

segments, so that they are separated from their sterna by a
wider extent of pleural membrane. The tenth tergite is also
laterally reduced, but is prolonged backwards over the supra-
anal plate, taking the place of the latter, which is greatly
reduced and feebly chitinized, as are also the larger paraprocts.
The spiracles are all situated on the tergites, near their lateral
margin. ‘The cerci consist of about 15 short segments, which
are but little flattened.

The ovipositor is of considerable size, but almost entirely
concealed by a huge hood-like structure, which is the seventh
sternum and is therefore not the homologue of the subgenital
plate of the groups thus far discussed. Under cover of this hood
and adherent to it is a bifid plate (Fig. 44), which overlaps the
vulva. This is part of the eighth sternum and apparently the
homologue of the subgenital plate of the Orthoptera and Phas-
moidea. The vulva lies between a pair of median shelf-like valves.
Extending forward from between the base of the valves is a
thick fold with a chitinized edge (ar), upon which is a small
opening of the spermatheca. In the natural position of the
parts this orifice lies just above the vulva. The valvule are large,
but of irregular shape and only partly chitinized. . The dorsal
pair overlaps the ventral and both are distally decurved and
peculiarly lobed at their blunt apices. The ventral valvule
are widely separated for some distance from their bases, this
space apparently serving for the exit of the ova and secretions
which form the ootheca. The basivalvule are usually distinct
and heavily chitinized, being the firmest parts of the valvule.
Just above the basivalvula is a small, but well-chitinized valvifer
connecting the dorsal and ventral valvule in the usual way. In
Crampton’s figure of the ovipositor of Stagmomantis® this plate
is incorrectly labeled “‘basivalvula”’ (i. e., of the dorsal valvule).
It bears only one apodeme, parallel to the antero-ventral margin
and continued as a strong ridge across the pleural membrane
to the tergites of segments eight and nine, opposite the inter-
segmental groove, where it is expanded into a short broad
spur (pap), and is thence continuous with the short tergal
apodemes of these two segments. (Fig. 48). The pleural
apodeme is a thickening of the intersegmental furrow, and is
marked externally by a distinct groove.

19 Jour. N. Y. Ent. Soc., Vol. XXV, Pl. XVI, Fig. 7, 1917.

296 Annals Entomological Society of America [Vol. XII,

The dorsal valvule are close together and at their extreme
bases they are joined by a fold of tough cuticle. Just behind
this fold is the large superior intervalvula, which unites the
two bases closely with one another and with the inner valvule.
It consists of a stout, bilobed transverse bar connected below
with a thin horizontal plate (hlm), which projects forward into
the body and bears a median apodeme, which has the same
muscular connections with the valvifers as that of the Gryllidz
and Tettigonide.

The dorsal apophyses are short but stout, the ventral rather
slender, but well chitinized and firmly united with the large,
thin, ventral intervalvula. The inner valvule are of moderate
length, but very close together, not enclosing a passage for the
exit of the eggs. There is a considerable space between their
lateral surfaces and the dorsal valvule. Like the other valvule,
they are lobed and ridged, and they engage the ventral valvule
ineffectively, by a sight tongue-and-groove joint. There is prac-
tically no intervalvular membrane and no pons, but the rami are
strongly chitinized and fused with the transverse bar of the
superior intervalvula. Their anterior extremities, however
(Fig. 49), lie below the level of this sclerite on each side of the
horizontal plate (hlm) which seems to represent an invaginated
intervalvular membrane. There is no connection between the
ends of the rami and the inferior intervalvula, such as occurs
in the Orthoptera.

A pair of small lobes project back from the inferior inter-
valvula between the inner valves. (cf. Blattoidea).

Blattoidea (Blattide).

In the cockroaches we find many of the peculiarities of the
Mantidez in a more pronounced form, as well as special char-
acteristics of their own. The ovipositor is not only degenerate,
but more or less greatly atrophied and completely concealed by
the large ‘“‘subgenital plate,’’ which as in the Mantidze is the
seventh, not the eighth sternum. Crampton (loc. cit., p. 227),
states that in the superorder Pandictyoptera, (later termed
Panisoptera and including Blattide, Mantide and Isoptera),
‘the ventral portion of the terminal abdominal segments 1s
typically overlapped by a backward prolongation of the eighth
segment.’’ He ‘considers Holmgren (Termitenstudien. Anat-

1919] Walker: Structure of Orthopteroid Insects 297

omische Untersuchungen, 1909) as being incorrect in designating
this segment in the Isoptera as the seventh, believing that the
first abdominal segment is not developed ventrally in this
order. Allowance is made by Holmgren, however, for the
missing segments for the subgenital plate is the sixth sternum
by actual count except in some forms where a vestige of the
first persists.

This enlarged seventh sternum is prolonged back to the end
of the abdomen, thus enclosing a very large “genital cavity”
(or ‘‘anal cavity,’ as it is sometimes called), in which the ovi-
positor is entirely hidden. In some genera such as Periplaneta
and Blatta it bears a pair of apical moveable valve-like plates,
which serve to close the opening of the genital cavity.

The eighth and ninth tergites are usually greatly abbreviated
and the tenth tergite, though laterally narrowed and not con-
tinued inwards beneath the cerci to any extent, is often con-
siderably produced backward between the cerci, overhanging
the anus and substituting the supra-anal plate, which in adult
cockroaches is wholly wanting. This tenth tergite is commonly
termed the supra-anal plate by systematists, but it is better to
restrict this term to the structure to which it has been generally
applied, otherwise it loses its morphological significance.

The disappearance of the supra-anal plate in the Blattids
and its substitution by the tenth tergite is the more complete
expression of the same tendency indicated in the Mantids,
where the true supra-anal plate, though present, is reduced and
entirely covered by the tenth tergum. A similar condition is
met with in the Isoptera (q. v.).

The Blattids also resemble the Mantids in having segmented
cerci, though these are shorter and more flattened, and in many
other respects, which will be noted in the following account.

As an example of a typical Blattid with a fairly well
developed ovipositor we may take the common native cock-
roach, Parcoblatia pensylvanica De Geer, better known as
Ischnoptera pensylvanica. Fig. 52, is an oblique or ventro-
lateral view of the terminal segments of this species, the seventh
sternum (‘‘subgenital plate’’) having been removed. Fig. 51
is a ventral view of the same parts, omitting those of the tenth
anal segments. Fig. 51 is a similar view, but with the valves
of the ovipositor forcibly bent forward and the right dorsal
valvula cut away from the base.

298 Annals Entomological Society of America [Vol. XII,

- The ventral valvule, like those of the Mantids, are irregular,
only partly chitinized and spread apart towards their bases.
But here a strange modification is seen. The bases are suddenly
enormously expanded and connected with one another by an
arcuate, chitinous band, which narrows at the sides and,
passing around behind the base, joins a broader plate in this
situation. This arcuate band, or its equivalent, varies greatly
in form in different Blattids and is clearly the homologue of
the transverse sclerite (ar) or chitinous edge of the fold in the
Mantids which bears the opening of the spermatheca. It
appears to be a special characteristic of these two nearly allied
orders. The posterior plates referred to are the basivalvule,
which have reached this position by the spreading apart of the
bases accompanied by an outward rotation. A similar shifting
of the valve bases is noticeable in the Mantidez, but is much less
pronounced. Inthe immature Parcoblatia, up to the last instar,
no such peculiarities are seen, the ventral valvule being close
together with typical basivalvule, and this simple condition is
retained in the adult of Cryptocercus punctulatus Scudd. (Fig. 59),
the ovipositor in this form having been apparently arrested in
development, in a manner closely comparable to that of the
Phasmid Timema californica (see pp. 292-3).

Immediately behind each basivalvula is the large valvifer,
which, as usual, connects the bases of the dorsal and ventral
valvulz and is continuous at its antero-lateral angle with a strong
apodeme, which follows the constriction between segments
eight and nine to the tergal margins of these segments and is
in every way comparable to the similar apodeme in the Mantids.
The valvifer is thus widely separated from the tergal margins
by a membranous area, which is crossed by the intersegmental
apodeme, their relations being essentially the same as in the
Mantide, and but little different from those which are typical
of the Orthoptera, in which the ninth tergal apodeme is really
the homologue of the intersegmental apodeme. The longi-
tudinal groove of the apodeme is plainly seen and marks the
constriction between the two segments.

Between the valvifers are two large plates which are con-
tinuous with the shafts of the dorsal valvule, from which they
are strongly bent outwards and downwards. These are simply
the bases of the dorsal valvule, and are not unlike the expanded
bases of these structures in Gryllus. The dorsal valvule are

1919} Walker: Structure of Orthopteroid Insects 299

decidedly folded and irregular on the dorsal side and bilobed
at the apices, though ventrally smooth. There is a prominent
sub-basal process on the dorsal side (pvd) but it is unchitinized.
It is perhaps represented in the Mantids by a minute lateral
lobe, which is present in this situation. The expanded bases
are united behind by a narrow strip of chitin, in front of which,
closely connected with: the bases, is the peculiarly shaped
superior intervalvula. There is also a pair of short spurs (sap)
which may represent the superior apophyses (not present in all
Blattidz) and a pair of slender rods with thin, anterior expan-
sions, (iap), which surround the bases of the inner valves and
meet in a somewhat heavier median portion (iv), just behind
the ventral valve bases. These rods evidently represent the
inferior apophyses, while the thicker median piece is the inferior
intervalvula. The spermatheca opens a little behind this plate;
in other forms, such as Blatta, directly upon it.

The inner valvule have much the same form as in Stagmo-
mantis and as in this genus they are slender, close together, and
lie in the hollow of the dorsal valvule, close to the latter and
engage the ventral valvule very feebly. In their slightly
expanded, flexible apices, the lack of a true intervalvular
membrane, and the presence of a pair of ventral basal lobes,
they also recall the Mantids. They are not fused with the
superior intervalvula, however, but are connected with the
latter by a peculiar ball and socket joint, the dorsal surface
of the fused valve bases bearing a knob, which fits into a socket
in the transverse part of the superior intervalvula. In front
of the knob the fused valve-bases form a thin plate which is
curved upwards (hlm). This plate is similar in position to
the plate (hlm) in Stagmomantis (Figs. 45, 49) and is doubtless
its homologue, as in Stagmomantis this plate is likewise con-
tinuous with the bases of the inner valvule, as well as the
superior intervalvula, of which it appears to form a part.

It will be seen from the foregoing description that, in spite
of wide differences in the form and proportions of the various
parts they are essentially similar in the Blattids and Mantids,
the differences being of small weight, as compared with the
many points of resemblance.

In addition to the structures described, there is a rather
large, though ill-defined chitinized area behind the bases of the
dorsal valves (ca), and serving for the attachment of muscles.

300 Annals Entomological Society of America [Vol. XII,

This is a secondary deposit of chitin and does not represent a
definite sclerite. It is not present in all Blattide.

A narrow chitinized strip possibly represents the tenth
sternum in Parcoblatta. The supra-anal plate is wholly absent
in the adult though its vestige is present in the young nymph.
The paraprocts, on the other hand, are large, broad and rather
flattened and the dorsal surfaces strongly chitinized. The cerci
are of the usual Blattid type, the basipodite developed on the
inner instead of the outer side.

The immature Blattid (e. g., Parcoblatta, Figs. 55-57) is
remarkable for the very primitive condition of the ovipositor,
which is indicated, (1) by the fact that the valvule all develop
from the hind margins of their corresponding segments; (2) in
the manner in which the dorsal valvule develop (vide infra);
(3) in the broad flat form of their valves, as distinct from the
slender inner and ventral valvule (cf. the Thysanura, Fig. 72),
and (4) in the persistence of styli on the dorsal valvule until
the last moult.

Development of the ovipositor (Parcoblatta). The stages here
outlined were first described by Denny (’94)2° in the case of
Blatta orientalis L. and are well known, but their significance
has not been sufficiently emphasized. Figs 55, 56, 57 and 58
represent ventral views of segments eight and nine, a portion
of seven (the seventh sternum being nearly all removed), of
three immature stages and the adult Parcoblatta pensylvanica.
The youngest nymph (Fig. 55) differs little from the male of
the same stage except in the presence of a pair of small pro-
cesses, on the hind margin of the eighth sternum and a small
apical median fissure of the ninth sternum. Both sterna are
well developed. In the next stage (Fig. 56) the eighth sternum
is much narrower; the two little processes are greatly elongated
and are marked off from the sternum by a shght constriction.
They are now distinguishable as the ventral valvule, and the
basivalvule are also indicated in the median sternal region.
The two stylus-bearing lobes into which the sternum is divided
have become relatively narrower, and the fissure between them
has greatly deepened and widened at base, from which the
rudiment of the inner valvulz have arisen.

20 Denny, A., Rept. 63, Meeting Brit. Assoc. Adv. Science, p. 818 (1894).

1919} Walker: Structure of Orthopteroid Insects 301

In the third stage (Fig. 57) which represents the last nymphal
instar, the eighth sternum is still narrower and has lost its disto-
lateral angles. The ventral and inner valvule have changed
but little, but the two lobes of the ninth sternum have greatly
decreased in width, though the styli are practically unchanged.
They are now clearly recognizable as the dorsal valvule.

The metamorphosis of this primitive condition into the
complex structure of the adult is a transformation worthy of
a higher order of insects, but the most significant feature in
the entire process is the clear indication that the dorsal valvulie
are a part of the ninth sternum, and are homologous with the
flat, stylus-bearing sternal lobes or “‘coxites’’ of the Thysanura.
Their comparatively late development is no objection to their
interpretation as such prinitive structures, in view of the well-
known fact that organs, which are not functional until adult
life is reached, are frequently retarded in development.

The ovipositor of Cryptocercus punctulatus (Fig. 59) presents
an interesting structure intermediate between that of the adult
and late nymphal stages of more typical roaches, such as
Parcoblatta. ‘The ventral and inner valves are quite like those
of an immature roach, while the dorsal valves have something
of the mature form, but are unusually simple. The valvifer
is of remarkably generalised form and together with the well-
marked intersegmental apodeme bears a closer resemblance to
these structures in the Mantids, than is found in any other
Blattid I have examined.

The ovipositor of Cryptocercus is, however, very degenerate,
the valvulz and valvifers being very feebly chitinized and their
inner connections greatly simplified. Thus, in view of the many
details held in common by the more complex types of Blattid
ovipositor and that of the Mantids, the simple structure found
in Cryptocercus must be looked upon as at least partly due to
arrested development, or the persistence of a larviform con-
dition, rather than a truly primitive one. It is closely paralleled
in the Phasmoidea by Timema californica.

It may be added that in certain respects Cryptocercus is
highly specialized, namely in the enormous development of
the seventh tergite, which is prolonged backwards over the
remaining segments, so that the latter are completely concealed
above and below. ji

302 Annals Entomological Society of America [Vol. XII,

Isoptera.

Two species of this order were examined, Termopsis angustt-
collis Hagen and Leucotermes flavipes Kollar. The former is
less degenerate in the structure of the genitalia and is therefore
more favorable for comparison with other groups, besides being
a much larger insect; so that the following remarks, unless other-
wise stated, refer to this species.

The abdomen (Figs. 60, 61), is tolerably broad and Aatienade
not unlike that of a cockroach in appearance, though the tergites
are more uniform in size, the eighth, ninth and tenth being
much abbreviated only towards the lateral margins. As in
the Mantids and Blattids, the tenth tergite is prolonged over
the supra-anal plate, which is obsolete in the adult; the
paraprocts are broad, subtriangular and more distinctly,
though not heavily, chitinized, and the short cylindrical
cerci are composed of five segments, of which the apical
one is much the longest and is probably compound. Termopsis
also agrees with these two families in the concealment of the
genitalia and sternal regions of segments eight and nine in a
‘‘genital cavity,’’ under cover of the backward prolongation of
the seventh sternum or “‘subgenital plate.’’ As mentioned
under the account of the Blattoidea, this plate is regarded by
Crampton as the eighth sternum, but this is certainly erroneous.
If the seventh sternum is cut away, a small chitinized flap is
found, overlapping the genital orifice and also covering the bases
of a pair of flattened lobes. This flap represents the free edge
of the eighth sternum and the two lobes are the ventral valves.
On each side of the lobes is a pair of plates, corresponding in
position to the valvifers of the cockroach and, like the latter,
thickened along the front margin. These plates are evidently
the valvifers, the thickening representing the usual apodeme
which occurs on the margin. It is not, however, continued
laterad of the valvifer, there being nothing here to mark the
line of junction of the eighth and ninth sterna.

Behind the overlapping by the ventral valves is a slender
V-shaped sclerite. This appears to represent the vestiges of
the two slender bars which meet in a similar position in Par-
coblatta and were interpreted as the inferior apophyses of the
dorsal valvule, meeting in the inferior intervalvula. They at
least represent some part of the bases of the dorsal valvulz

1919] Walker: Structure of Orthopteroid Insects 303

which have otherwise disappeared, together with the inner
valvule. The remainder of the ventral surface of segments
nine and ten is covered with a thinly chitinized cuticle.

In Fig. 63 the eighth abdominal .spiracle may be seen in
the conjunction immediately behind the eighth sternum, and
close to the lateral margin, a position similar to that which it
occupies in Parcoblatta.

It is sufficiently evident that the terminal abdominal struc-
tures of Termopsis are essentially like those of a Blattid in which
the ovipositor has nearly disappeared. *° In Leucotermes it
has quite vanished and the cerci are reduced to two segments.

Judging by the more uniform length of the abdominal
tergites, the cylindrical form of the cerci, and many other
characters in other regions of the body it is probable that the
ancestors of the Isoptera were more primitive than any Blat-
toidea of recent age, but the genitalia are distinctly more
suggestive of the Blattids than the Mantids, and the wing
venation of the primitive New Zealand termite Mastotermes, is
decidedly more like that of the Blattoidea than the Mantoidea
or .Protoblattoidea, so that I am inclined to consider the order
Isoptera as an offshoot from primitive Blattoid stock.

The results of these studies of the genitalia of the Mantoidea,
Blattoidea and Isoptera strongly support Crampton’s grouping
of these orders in a superorder “‘ Panisoptera.’”’

Dermaptera.

In this order the genital segments are so highly modified
that they give little information that is of value in determining
the systematic position of the order. While in the majority
of forms the ovipositor is entirely lacking, a small one is present
in some genera of Protodermaptera, notably in the families
Pygidicraniide and Echinosomide (Zacher, ’11),2! so that we
may conclude that the absence of this structure is a secondary
condition. I have not seen any of these ovipositor-bearing
forms, having, in fact, examined critically only two species
_ of the order, viz., Forficula auricularia L. and Anisolabis
maritima Bon. I have reproduced, however, figures of the
ovipositor of Kalocrania and Echinosoma from Zacher (loc. cit.).

"1 Zacher, Friedrich, Zool. Jahrb., Bd. XXX, Syst., pp. 303-400, 80 Figs. (1911).

304 Annals Entomological Society of America [Vol. XII,

In Forficula (Figs. 62-64) the eighth, ninth and tenth
abdominal tergites are fused, but their boundaries are clearly
defined. The eighth and ninth tergites are very short and
concealed by the much larger, overlapping seventh tergite;
while the tenth is also very large, doubtless owing to the great
development of the muscles concerned in the movement of
the large forcipate cerci. The eighth and ninth sterna are
concealed by the very large seventh sternum, and this feature,
together with the reduction of segments eight and nine and the
atrophy of the ovipositor is very suggestive of the ‘‘ Panisoptera”’
(Blattoidea, Mantoidea and Isoptera); but unlike these groups,
the tenth tergite does not replace the supra-anal plate, which
is entirely free, and although small, is heavily chitinized and
divided transversely into two separate sclerites, the distal
of which is situated ventrally between the bases of the cerci,
with which it is articulated. Possibly these two sclerites
represent the eleventh tergite and the true supra-anal plate,
as in the Acridoidea. Where three such sclerites are present
(Pygidicraniide, Allosthetide, teste Zacher) they have been
interpreted as representing the tergites of as many segments,
viz., the eleventh, twelfth and thirteenth, or anal segment
(pygidium, metapygidium and telson, or supra-anal plate),
but the evidence for the existence of an additional segment
between the eleventh and anal segments is quite insufficient.
It is worthy of note, however, that these three sclerites are thus
separately developed only in primitive genera.

Apart from the features mentioned above, there appears to
be no evidence of close relationship between the Dermaptera
and the Panisoptera. —

The eighth and ninth sterna in Forficula are feebly chitinized
and each is divided into two lateral plates, which were con-
sidered by Verhoeff (’03)” to represent coxites. It is probable,
however, that this division of the sterna is related to the former —
presence of an ovipositor, which occupied the median space,
and this view is supported by the presence of an ovipositor
in this situation in such genera as Kalocrania and Echinosoma,
as judged by Zacher’s figures. In these genera the ovipositor
consists of but two pairs of valvula, belonging to the eighth
and ninth segments. The former pair is long and slender

22 Verhoeff, H. W. Nova Acta. Acad. Caes.-Leop., Vol. XX XI, 1903, pp.
277-278.

1919] Walker: Structure of Orthopteroid Insects 305

and obviously represents the ventral valvule. The other pair,
from its lateral position and comparatively broad form is doubt-
less the dorsal valvule. Since the dorsal valvule represent the
coxites of segment nine, the divided sternal plates, which are
also present cannot be correctly termed coxites. They are
more nearly comparable to the valvifers.

In some genera, such as Anzsolabis (Fig. 65) there is also
a small paired tenth sternum, but this is absent in Forficula.
The two large sub-triangular plates (Figs. 66, 67), which are
closely united with the margins of the tenth tergite and form
part of the articulation with the cerci, were regarded by Ver-
hoeff and Zacher as the coxites of segment ten, while Crampton
identified them with the paraprocts. Crampton’s view is the
more probable in my opinion. Coxites are absent from segment
ten in all other primitive insects, so that on a priori grounds
we should not expect to find them in the Dermaptera. Para-
procts, on the other hand, are almost invariably present, and
although these plates appear to belong to segment ten, there is
no reason why, in such a highly modified group as the Der-
maptera, they should not have been developed from the para-
procts, as these are usually closely connected with the margins
of the tenth tergite, as, e. g., in the Blattoidea.

The spiracles all occupy the pleural membrane, the last
pair, as in all the groups discussed, belonging to the eighth
segment.

The unsegmented form of the cerci has probably been
developed within the course of evolution of the order, as in
immature stages of certain primitive forms (Dzplatys, Karschiella
and Bormansia) they are segmented.

Embiidina (Embiide).

This small group, which is undoubtedly of ordinal rank, has
been considered by some writers (Enderlein and others) to be
nearly allied to the Isoptera, while others (Crampton, West-
wood, MacLachlan) find closer relations with the Plecoptera
and Dermaptera. The latter view, is in the present writer’s
opinion, much nearer the truth. The resemblance to the
Isoptera is largely due to the retention in both groups of many
primitive characters and to parallel development along certain
lines, such as the form and venation of the wings, the two-

306 Annals Entomological Society of America (Vol. XII,

jointed cerci, the loss of the ovipositor, etc., but the special
features of each group indicate a different line of descent.

As regards the female genitalia, but little evidence in support
of either view is obtainable from this source alone. In Embia
major, e. g., (Fig. 68) the general appearance of the terminal
segments is much like that of a termite. The carci are two-
jointed as in most termites, the tenth tergite is large and curves
downward, covering the supra-anal plate, to which it is adher-
ent. On the other hand, there is no extension of the seventh
sternum to form a subgenital plate, in fact, no modification
of either seventh or eighth sterna, and no lateral reduction of
tergites eight, nine and ten as in the Isoptera. The paraprocts
are large, but unchitinized and are distinct from the well-
developed cercal basipodites.

Whereas in Termopsis there is evidence that the Isoptera
are descended from ovipositor-bearing ancestors, there is no
indication in Embia or any of the Embiidina that an ovipositor
was ever present, the eighth and ninth sterna being quite
simple. Since an ovipositor of primitive form is present in
some Apterygota (Machilis, Lepisma, etc. of the Thysanura)
and is undoubtedly homologous with that of Pterygote insects,
it must have been present in the earliest representations of the
latter, unless we are to regard the ovipositor-bearing Thysanura
as descended from winged forbears, a view which I believe few
will accept. This being the case, the Embiids, Plecoptera, etc.,
must likewise, (contrary to Crampton’s opinion) be considered
as secondarily without ovipositors. The fact that Campodea
and other Thysanura have also no ovipositor does not affect
the question. There is~evidence that some of the Palaeo-
dictyoptera had no ovipositor, while it is certain that some of
them had one, and it was probably among the former that
the ancestors of the Embiids and Plecoptera existed. The
ovipositor, which was probably never highly differentiated, had
already disappeared before these groups had acquired independ-
ent ordinal rank.

It is scarcely profitable to make a comparison between the
Embiids and Grylloblatta until the structure of the male
genitalia has been considered, but the following characters
held in common between the females of the two orders may be
mentioned: (1) There is no reduction in the length of the eighth
and ninth tergites; (2) the spiracles are all situated in the

| 1919] Walker: Structure of Orthopteroid Insects 307

pleural membrane; (3) neither the seventh nor the eighth
sterna are prolonged into a sub-genital plate; (4) the cerci
are segmented; (5) the paraprocts are feebly chitinized. The
last feature has no special significance; the others indicate the
primitive nature of both groups, but are otherwise negative
in value, when taken alone.

No relationship to the Dermaptera or Orthoptera is even
hinted at in the terminal abdominal segments of the females
of Embiids.

Plecoptera.

As Crampton and others have pointed out, this order is in
some respects the most primitive of existing Pterygote insects,
particularly in the cervical and thoracic sclerites, wing venation
and cerci. In the abdominal segments, of which ten are well
developed, there is a tendency in many forms towards a con-
siderable degree of chitinization of the pleural membrane.
The ninth and tenth segments may be quite ring-like, even
in the adult, while in the nymph all the segments may be
annular. Crampton’s suggestion that this annular form of
segment may be a primitive one seems to me untenable. It is
too exceptional among the Tracheata, and even within. the
Plecoptera there are all grades of chitinization of the pleural
membrane. It is moreover, explained by the non-functional
character of the abdominal spiracles in the nymph, in which
respiration is performed by the tracheal gills, these structures
sometimes (e. g., Pteronarcys) persisting in the adult in a
reduced form.

The abdominal spiracles are all pleural in position, the
eighth sternum is frequently more or less prolonged to form a
subgenital plate, sometimes overlapping the ninth, or even the
tenth, sternum. In some forms (e. g., Megarcys signata
Hagen)” it is bilobed or bifid at apex, these lobes being slightly
suggestive of vestigial ventral valvulea. In most species of
Pteronarcys there is no backward extension of the eighth
sternum as a whole, but it bears a pair of slender processes, at
or near the hind margin, which are probably true representatives
of these valvule, being very similar to these structures as met

23 Klapalek, Fr., Coll. Zool. Selys., Fasc. IV, p. 12, Fig. 6 (1912). Smith, Lucy
Wright, Trans. Am. Ent. Soc., Vol. XLIII, pp. 433-489, Pis. XXIX-XXXIV (1917).

308 Annals Entomological Society of America [Vol. XII,

with in certain Libellulid dragonflies. There is no other trace
of the ovipositor in the Plecoptera, so far as I am aware, the
ninth sternum being quite simple.

The tenth segment is well developed, often little smaller
than the ninth. The tergite is often somewhat prolonged
behind, concealing or partly covering the small supra-anal
plate, with which it may be adherent. The paraprocts are
large, generally well chitinized and usually intimately fused
with the bases of the cerci so that the latter appear to arise
from them. The cercal basipodites are thus not distinct from
the paraprocts.

The cerci are typically multiarticulate, but are very variable
in respect to both the number and the form of the segments.

These characters, taken by themselves, do not throw much
light on the affinities of this group, but point to a very gen-
eralized structure, with secondary loss of the ovipositor. The
primitive multiarticulate cerci are approached by those of the
Grylloblattoidea, more closely than any other order, with the
possible exception of some of the Ephemerida.

Ephemerida.

Although the Ephemerida and Odonata can hardly be called
‘‘Orthopteroid’’ insects, and their lines of descent from the
Paleodictyoptera are undoubtedly quite distinct from any of the
others considered, they deserve a few words, on account of their
having retained certain very primitive characters.

The females of Ephemerida are chiefly remarkable from the
fact that the oviducts open separately, and behind the seventh,
instead of the eighth sternum. In some forms the seventh
sternum is prolonged backwards into a spout-like structure,
which apparently functions as an ovipositor (Morrison, ’19)?4
but this, of course, has no homology with any part of the
Orthopterous ovipositor. The elongate, uniformly segmented
abdomen, multiarticulate cerci and cerciform caudal filament,
borne by the eleventh tergite are all marks of primitive structure.
The anal valves are membranous and very slightly developed.

24 Morrison, Emily Reed, Can. Ent., Vol. LI, No. 6, pp. 189-146 (1919).

1919] Walker: Structure of Orthopteroid Insects 309

Odonata.

The form of the terminal segments of the dragonflies recalls
the Phasmids, as suggested by Crampton, there being no
abbreviation, but on the contrary, an elongation of all of them,
as compared with the usual conditions. A fully developed
ovipositor is present in all of the suborder Zygoptera and some
of the Anisoptera (Aeshnidz:—Aeshnine and Petalurine),
while a more or less reduced and simplified one occurs in the
other groups. This ovipositor is remarkable in several ways.
The dorsal valvule are broad, subtriangular, hood-like at the
apices, and form a pair of flaps or covers for the ventral and
inner valves, these structures serving as the actual instrument
for making the punctures or incisions in which the eggs are
placed. They thus resemble closely the broad sternal processes
of segment nine in Lepisma, Machiliis and other Thysanura,
which cover over the two pairs of valvule (representing the
ventral and inner pairs) in quite a similar manner. They are
still more interesting in the fact that they retain the styli in
adult life as functional sense-organs, the Odonata being the
only Pterygote order 1n which this is the case.

The ventral valvulz also possess well developed basivalvule,
the dorsal valvule superior and inferior apophyses. A superior
intervalvula and intervalvular membrane are also present, but
no inferior valvula, the strong rami of the inner valvule being
articulated with the inferior apophysis.

There is also a valvifer, having typical connections with
the valvule and ninth tergal apodeme.

The occurrence of these features in the Odonata is of interest
in showing that they must be characters of very ancient origin
and are in no way specially characteristic of Orthopteroid
groups.

310 Annals Entomological Society of America [Vol. XII,

SUMMARY OF CHARACTERISTICS OF THE ORDERS.

BASED. ON THE TERMINAL ABDOMINAL STRUCTURES OF THE
FEMALES OF TYPES EXAMINED.

Orthoptera—Eighth sternum more or less modified as a
subgenital plate, but leaving ovipositor exposed; ninth sternum
vertical or nearly so; valvifers (sometimes absent) in contact
with ninth tergum and its marginal apodeme; ovipositor
well developed (rarely vestigial or absent), with three pairs of
valvule; basivalvule rarely well developed; a superior inter-
valvula connecting bases of dorsal valvule, which cover inner
valvulz and have two pairs of apophyses (superior sometimes
indistinct or absent); inner valvule, when not vestigial, with
rami, intervalvular membrane and pons, the rami with longi-
tudinal ridge which fits into groove on ventral valvule; an
inferior intervalvula connected separately with rami and inferior
apophyses; tenth sternal region not defined, unchitinized;
cerci unsegmented (two-segmented in Tridactvlus); supra-
anal plate well developed, sometimes fused with but not con-
cealed by tenth tergite; paraprocts usually well developed.

Nymph: Valvulze all slender, dorsal and ventral pairs
developing from ventral surface of ninth sternum; styli rarely
distinct and usually disappearing at a very early stage.

Grylloblattoidea—Eighth sternum unmodified in form, ovi-
_positor exposed; ninth sternum, valvifers and ninth tergal
apodeme as in the Orthoptera; ovipositor well developed, with
three pairs of valvule; basivalvule exposed, heavily chitinized,
superior intervalvula absent; dorsal valvule covering inner pair;
both pairs of apophyses present, inferior apophyses joining
one another medially, there being no separate inferior inter-
valvula; inner valvulz well developed, with rami, intervalvular
membrane and pons; rami fused with inferior apophyses and
connected with ventral valves as in Orthoptera; tenth segment
annular; cerci, slender eight-segmented; supra-anal plate small,
but not concealed by tenth tergite; paraprocts not large,
unchitinized.

Phasmoidea—Eighth sternum modified, usually forming a
very large flap-lke subgenital plate, largely concealing the
Ovipositor; ninth sternal regional horizontal; valvifers and

1919] Walker: Structure of Orthopteroid Insects ore

ninth tergal apodeme variable in development; ovipositor
generally of considerable size, but the three pairs of valvule not
firmly chitinized; dorsal valvule widely separated at base,
not covering the inner pair and without a superior intervalvula;
superior and inferior apophyses present, the latter widely
separated; inner valvule united dorsal except distally and
having sometimes an ineffective tongue-and-groove connection
with ventral valvule; no intervalvular membrane or inferior
intervalvula; tenth segment with distinct sternal region; cerci
short, unsegmented; supra-anal plate usually small, paraprocts
prominent.

Mantoidea—Tergites of segments eight and nine shortened;
seventh sternum forming a very large subgenital plate concealing
remaining sterna and ovipositor; eighth sternum greatly
reduced; ninth, vertical or subvertical; valvifers separated from
tergites, but connected across intervening membranous area
by the intersegmented apodeme with eighth and ninth tergites;
ovipositor of considerable size with three pairs of valvule,
basivalvule heavily chitinized, widely separated, but con-
nected by a chitinous arch or transverse bar; valvule of irregular
form, and imperfectly chitinized; dorsal valvule covering inner
pair, united at.extreme base; superior intervalvula connecting
both dorsal and inner valvule; dorsal valvule with two pairs
of apophyses, the inferior apophyses joined by a large inferior
intervalvula; inner valvule free from the latter without
intervalvular membrane, but the internal (basal) prolongations
of the rami connected by a thin plate which bears the median
apodeme of the superior intervalvula; ventral valvule engaging
inner by a feeble tongue-and-groove joint. Cerci subcylindrical,
many-segmented; supra-anal plate vestigial, concealed by tenth
tergite; paraprocts little chitinized.

Nymph (Mantis) similar to that of Blattoidea, but styli
in last stage very minute.

Blattoidea—Abdominal segments flattened; tergites of eight
and nine shortened; seventh sternum forming a very large
subgenital plate, concealing remaining sterna and ovipositor;
eighth sternum greatly reduced; ninth obliquely inclined;
valvifers very large, separated from tergal margins, but con-
nected across intervening membranous area with the eighth
and ninth tergites by the intersegmented apodeme; ovipositor
small, the three pairs of valvule of irregular form, more or less

312 Annals Entomological Society of America [Vol. XII,

imperfectly chitinized and not coherent, though an ineffective
engagement between the ventral and inner valves may occur;
basivalvule very widely separated and rotated outwards, con-
nected in front by a chitinous arch or bar; dorsal valvule cover-
ing inner pair, their bases greatly expanded, and medially united;
superior intervalvula connected with dorsal valvule and with
lamella formed by united bases of inner valvule, which are free
from ventral valvule and lack an intervalvular membrane; infer-
ior apophyses represented by slender bars connected with a
vestigial inferior intervalvula; cerci of moderate length, flattened,
segmented; supra-anal plate absent in adult, being substituted
by tenth tergite; paraprocts flattened, generally well chitinized
in part.

Nymph with valvule terminal, dorsal pair broad and flat,
retaining styli until last stage; a vestigial supra-anal plate
present.

Isoptera—Abdominal segments somewhat flattened; seventh
sternum forming a large subgenital plate, covering the remaining
sterna; eighth sternum ill-defined and greatly reduced; ninth,
oblique or almost horizontal; valvifers, when present, widely
separated from tergites, the intervening space not crossed by an
apodeme; ovipositor absent or represented by a very reduced
pair of ventral valvule and traces of the bases of the dorsal
valvule; tergites eight, nine and ten laterally narrowed, the
tenth replacing the supra-anal plate, which is absent in
the adult; paraprocts broad, somewhat chitinized; cerci short,
slender, with 2—5 segments.

Dermaptera—Tergites of segments eight and nine greatly
abbreviated and concealed by the seventh, tenth very large;
seventh sternum forming a subgenital plate and concealing
eighth and ninth sterna, which are medially divided; ovipositor
usually absent, when present, reduced and lacking inner
valvule; ventral valvule long and slender, dorsal valvule
shorter and broader; tenth sternum sometimes represented by
a pair of small sclerites; cerci very large and strong, forcipate,
unsegmented (except in larve of some genera); supra-anal
plate terminal, exposed, heavily chitinized, divided into
pygidium and telson, sometimes with an intervening meta-
pygidium; paraprocts ventral, consisting of flat, well-chitinized
plates.

1919] Walker: Structure of Orthopteroid Insects 313

Embudina—Segments seven to nine unmodified, there being
no specialized subgenital plate; ovipositor wholly absent;
supra-anal plate practically absent, replaced by the large tenth
tergite; paraprocts lobate, unchitinized; cerci two-jointed.

Plecoptera—Eighth sternum generally more or less modified
to form a subgenital plate, sometimes with a pair of lobes or
processes, possibly representing vestigial ventral valvule;
ovipositor otherwise absent; ninth sternum horizontal, unmodi-
fied; ninth and tenth segments tending to be annular; supra-
anal plate variable, sometimes covered by a prolongation of
the tenth tergite; cerci typically long and many-jointed, their
basal segments fused with the large paraprocts.

COMPARISON OF OVIPOSITORS OF PTERYGOTE AND
APTERYGOTE INSECTS.

In the Apterygota the ovipositor is present only in the
families Machilidz and Lepismatide of the order Thysanura.
It consists of two pairs of filiform gonapophyses, arising from
the eighth and ninth sternum respectively. In Machilis sp.,
e. g., the separated eighth sternum has the appearance shown
in Fig. 74. It is deeply bilobed, each lobe bearing a stylus
while the gonapophyses occupy a median position between the
lobes. They are very long and flexible and are divided by faint
constrictions with numerous segments, with regularly arranged
groups of sete. The ninth sternum (Fig. 75) is similar to the
eighth, except that the styli and sternal lobes are much longer,
while the gonapophyses are shorter and. more slender. In the
natural position they reach about the same distance back.
As compared with the eighth and ninth sterna, the preceding
sterna (Fig. 73) differ not only in the absence of gonapophyses,
but in the union of the stylus-bearing lobes and the presence of a
triangular basal plate (ste), these parts being separated only
by sutures. They also differ in the presence of eversible
glands (cgl).

It is now generally recognized that the basal plate is the true
sternite, while the lateral styli-bearing plates are coxttes,
these probably representing flattened coxz of abdominal limbs
which have otherwise disappeared (Haase, ’89).2° The sterna of

25 Haase, Erich, Morph. Jahrb., Vol. XV, pp. 331-435, Pls. XIV, XV (1889);
Verhoeff. Zool. Anz., Vol. XXVI, pp. 60-77 (1903).

314 Annals Entomological Society of America [Vol. XII,

segments eight and nine in Machilis are thus composed chiefly
of the coxites, the sternite having practically disappeared.
In some of the Lepismatide, however, (Nicoletia, Atelura)
(Escherich, ’05) the ninth segment has a distinct sternite
overlapping the bases of the coxites just as in the thoracic
segments. Comparing the eighth and ninth sterna of Machilis
with those of the immature Blattid (Figs. 56-59) we have no
difficulty in recognizing the coxites of segment nine in the flat
stylus-bearing lobes, which become the dorsal valvule, and
the gonapophyses of the same segment in the inner valvule
of the Blattid, while the gonapophyses of segment eight in
Machilis are the homologues of the ventral valvule of the
Blattid. The styli of the latter segment have disappeared and
the coxites are fused with the sternite, to form a ‘‘coxosternum”’
(Verhoeff, loc. cit.). It may be added that in Machilis, Lepisma,
etc., the coxites of segment nine overlap the gonapophyses,
forming a sort of sheath for them, just as they do in such
Odonata as have retained a well-developed ovipositor. In
Aeshna, e. g., (Fig. 71), the ‘genital valves’’ are the coxites
of segment nine or dorsal valvule, and although more complex
in form they have a very similar general position to those of
Machilis, covering the other valvule in the same way, and
bearing styli at their apices. There is no part of the anterior
gonapophyses of Machilis that is distinctly recognizable as the
basivalvule.’”’

The homologies of the terminal abdominal structures of the
‘female with those of the male, and other general questions will
-be discussed in Part II.

26 Escherich, K. Das System der Lepismatiden, Zoologica, Bd. 18, Heft 43,
pp. 1-164, 11 Pls. and 67 text figures (1905).

1919] Walker: Structure of Orthopteroid Insects 315

EXPLANATION OF PLATES.

REFERENCE LETTERING.

ap 8, 9—apodeme of segment 8 or 9. pa—paraprocts.
ar—basal arch. pap—process of intersegmental apodeme.
be—basipodite of cercus. pp—ventral prominence of pons.
bs—basivalvula. pvd—sub-basal process of dorsal valvula.
c—cercus. rm—rami of inner valvulae.
cf—caudal filament. sa—supra-anal plate.
clg—coxal gland sap—superior apophysis.
cx—coxite. sp—spiracle.
eg—egg guide. spth—aperture of spermatheca.
ga—genital aperture or vulva. st—sternum, sternal region.
hlm—internal lamella joining bases of | ste—sternite.

inner valvulae. stl—stylus.
jiap—inferior apophysis. sv—superior intervalvula.
int ap—intersegmental apodeme. tg—tergite.
im—intervalvular membrane. vd—dorsal valvula.
iv—inferior intervalvula. vi—valvifer.
lb—inferior lobes of inner valvulae. vi—inner valvula.
p—pons valvularum. vv—ventral valvula.

PLATE XX.

Ceuthophilus lapidicola, adult; lateral view of end segments and ovipositor,
ventral and inner valvulae bent downwards.

Same, not quite mature; dorsal view of ovipositor, left ventral valvula bent
outwards, right not shown.

Same, adult; ventral view of ovipositor.

Same, inner view of structures at base of ovipositor.

Conocephalus fasciatus, adult; lateral view of end segments and basal part of
ovipositor.

Same; ventro-lateral view.

Same; dorsal view of superior intervalvula and basal connections of inner
valvulae.

Same; anterior view of base of ovipositor.

Same; ventral view of end segments of nymph four mm. long.

Same; same view of nymph five mm. long.

PLATE X XI.

Gryllus assimilis, adult; lateral view of end segments and base of ovipositor.

Same; ventro-lateral view.

Same; dorsal view of ovipositor, left ventral valve bent outwards, right not
shown.

Same; inner view of structures at base of ovipositor, left side.

Same; anterior view of base of ovipositor. ,

Oecanthus quadripunctatus, adult; lateral view of end segments and base of
Ovipositor.

Same; inner view of structures at base of ovipositor, left side.

Same, nymph; ventral view of end segments and ovipositor.

Ripipteryx forcipata Sauss., adult; lateral view of end segments and ovipositor.

Same; dorsal view.

Same; ventral view.

PLATE XXII.

Melanoplus bivittatus, adult; postero-lateral view of end segments and
ovipositor.

Same; postero-dorso-lateral view of cleared preparation.

Same; posterior view of ovipositor, with valvulae forced widely open.

Same; ventral view of ventral valvulae.

Acrydium ornatum, adult; lateral view of end segments and ovipositor.

Same; dorsal view.

Same, nymph; lateral view.

Same, nymph; dorsal view.

Same, adult; ventral view of ventral valvulae.

316 Annals Entomological Society of America [Vol. XII,

PLATE XXIII.

31. Grylloblatta campodeiformis, adult; lateral view of end segments and ovipositor.

32. Same; ventral view.

33. Same; dorsal view of ovipositor, left ventral valvula bent outwards, right
not shown.

34. Same; inner view of structure at base of ovipositor, left side.

35. Same; anterior view of base of ovipositor.

36. Same, nymph; lateral view.

PLATE XGXLV:

37. Grylloblatta campodeiformis, nymph; ventral view of end-segments.

38. Same with ventral valvulae bent forward to expose inner valvulae.

. 39. Timema californica, adult; ventral view of end segments.

40. Same, lateral view.

41. Diapheromera femorata, adult; ventro-lateral view of end segments.

42. Same; lateral view of ovipositor; most of the 9th tergite cut away to show
the superior apophysis. :

43. Same; dorsal view of ovipositor.

PLATE XXV.

44. Stagmomantis carolina, adult; 8th sternum and ventral.

45. Same; dorsal view of ovipositor, with dorsal valvulae spread apart.

46. Same; ventral view of ovipositor with ventral valvulae spread apart.

47. Same; lateral view of terminal segments and ovipositor with 7th sternum
bent downwards to expose the valvulae.

48. Same; inner view of structures at base of ovipositor.

49. Same; anterior view of base of ovipositor.

PLATE X XVI.

50. Parcoblatta pensylvanica, adult; ventral view of ovipositor, the left ventrai
valvula cut off at base.

51. Same; the valvulae bent forward to show their dorsal surfaces and the right
dorsal valvula removed.

52. Same; ventro-lateral view of terminal segments, the 7th sternum removed.

53. Same; ovipositor removed, cleared, and viewed from above. The ventral
valvulae and the dorsal valvulae, except their bases, are omitted.

54. Same, last nymphal stage; ventral view of end segments with the 7th sternum
except a small part, removed.

55 to 58. Same; successive stages in the development of the genitalia, ventral
view, 7th sternum removed, cerci 10th and anal segments omitted. Fig. 58
is from the adult.

PLATE X XVII.

59. Cryptocercus punctulatus, adult; ventral view of end segments, the 7th sternum
removed.

60. Termopsis angusticollis, adult; lateral view of end segments.

‘61. Same; ventral view of end segments, the 7th sternum removed.

62. Forficula auricularia, adult; lateral view of end segments.

‘63. Same, dorsal view.

‘64. Same, ventral view, 7th sternum: removed.

65. Anisolabus maritima, adult; ventral view of end segments, the 7th sternum
removed.

66. Kalocrania marmoricrura, adult; ventral view of ovipositor and neighboring
sclerites. (After Zacher.)

67. Echinosoma occidentale, adult; ventral view of ovipositor and part of segment
10. (After Zacher.)

PLaTE XXVIII.

68. Embia major, adult; lateral view of end segments.

69. Pteronarcys proteus, adult; ventral view of end segments. (After Smith.)
70. Perla lycorias, adult; ventral view of end segments.

71. Aeshna canadensis, adult; ventral view of end segments.

72. Machilis sp., adult; ventral view of end segments.

73. Same; 8th sternum and gonapophyses (ventral valvulae).

74. Same; 9th sternum and gonapophyses (dorsal and inner valvulae.)

ANNALS KE. S. A. VoL. XII, PLATE KX:

E. M. Walker.

ANNALS E.S. A. Vou. XII, PLATE XXI.

|g 10 te9 tes

E. M. Walker.

ANNALS E. S.A, VOL. XII, PLATE XXII.

= 30

Z tg 10 ‘g° tgs

] 5 oe
Yo

te9 tg8

S geiao ia

bs p

vtec 26

E. M. Walker.

ANNALS E. S. A. Vou. XII, PLATE XXIII-

E, M. Walker.

ANNALS E.S. A. VOL. XII, PLATE XXIV.

rm?

sap

pap

E. M. Walker.

ANNALS E. 5S. A. VoL. XII, PLATE XXV.-

E. M. Walker.

ANNALS E.S. A. VoL. XII, PLATE XXVI.

E. M. Walker.

ANNALS E.S. A. VoL. XII, PLATE XXVII.

sr ety,

E. M. Walker.

VOL. XII, PLATE XXVIII.

ANNALS E. S. A,

— re

Sr rns nnamntnirnceenavennraanet een G

ES
\
WAN \ ‘iS
2 SS \\
spt SSSA
oe BBs
OA DDD ue
Lg
“oe
SSK
i Peay 2 if
Vi EG fyi /

y
a2
5 oy Eom

E. M. Walker.

UNDESCRIBED SPECIES OF JAPANESE CRANE-FLIES.
(TIPULIDZ, DIPTERA)

By CHARLES P. ALEXANDER, Ph. D.

The species of crane-flies herein described as new were
included in some extensive and highly interesting collections
made in the vicinity of Tokio by Mr. Ryoichi Takahashi,
during the years 1918 and 1919. The station ‘“‘Saitama,”’
refers to Chichibu, in the province of Saitama, a mountainous
locality about fifty miles from Tokio. The types are preserved
in the collection of the author. I am greatly indebted to Mr.
Takahashi for this very valuable series of Japanese Tipulide.

Dicranomyia Stephens.

Dicranomyia immodestoides sp. n.

Resembles D. immodesta O. S. (Eastern North America); antennz
dark brown throughout; general coloration light gray; wings nearly
hyaline, stigma pale; veins Sc: very long, abdomen dark brown, indis
tinctly ringed with yellow.

Male—Length, 5.8 mm.; wing, 6.6 mm.
Female—Length, 6.8 mm.; wing, 7.4—8.3 mm.

Rostrum reddish brown; palpi dark brown. Antenne with the
scape brownish black, the basal flagellar segments brown, the apical
segments darker. Head light yellowish gray; the front and the vertex
adjoining the inner margin of the eyes bright silvery.

Mesonotum dull gray, the praescutum with an indistinct median
brown stripe; lateral stripes indistinct; humeral areas a little more
reddish; scutum reddish gray, the lobes largely brown; scutellum
reddish gray; postnotum gray. Pleura light gray, indistinctly marked
with darker beneath the wing root. MHalteres rather long, pale, the
knobs dark brown, the base of the stem more yellowish. Legs with
the coxee and trochanters yellow, the former very sparsely gray pruinose,
the fore coxe dark basally; femora dull brownish yellow, somewhat
darkened on the apical half; tibie and tarsi brown. Wings nearly
hyaline, the stigma large, oval, very pale brown; veins slender, brown.
Venation: Sc; ending opposite origin of Rs, Sc, far before the tip of
Sci, the latter being longer than the basal deflection of Cm; r arcuated,
at the tip of Ri; cell 1st M2 open by the atrophy of m; basal deflection of
Cu, at or before the fork of M.

Abdominal tergites dark brown, the segments laterally at the base
and on the caudal margin dull yellow; sternites nearly similar, the
bases of the segments likewise yellowish. In the female the bicolored

327

H
. ‘
‘ 5 : )

328 Annals Entomological Society of America Wools ane

condition of the abdomen is not so evident as in the male. Male
hypopygium with the ninth tergite deeply notched medially, the
adjacent lobes evenly rounded, setigerous; dorsal pleural hook moder-
ately long, at the tip suddenly narrowed into a slender point; ventral
pleural appendage large, fleshy, the proximal basal portion produced
into a fleshy lobe that bears two erect chitinized spines on its caudal
face.

Habitat: Japan.
Holotype, &, Meguro, Tokio, April 1, 1919 (R. Takahashi).
Allotopotype, ?, March 31, 1919.

Paratopotypes, 6 &@ 2’s, March 25-May 26, 1919.

Dicranomyia basifusca sp. n.

Antenne black throughout; thorax gray, the praescutum with an
indistinct median brown stripe; wings pale gray, the stigma and a
rounded spot at the base, brown; Sc ending a little beyond the origin
of Rs, cell 1st M2 closed.

Female—Length, 4.8 mm.; wing, 6.4 mm.

Rostrum and palpi dark brownish black. Antenne black through-
out, the basal flagellar segments short-oval, the terminal segments a
little more elongated. Head gray, the front more tawny.

Mesonotum dull gray, the praescutum more brownish medially
to form an indistinct dorsal stripe; scutellum and postnotum lighter
gray. Pleura dark gray. Halteres yellow, the knobs dark brown.
Legs with the coxee brownish yellow, darker basally; trochanters dull
yellow; femora dull brownish yellow, darkened toward the apices, the
extreme tips pale; tibia yellowish brown, the tips narrowly darker
brown; tarsi dark brown. Wings with a faint grayish tinge, the stigma
short-oval, brown; a conspicuous rounded brown spot at the arculus;
origin of Rs and the cord very indistinctly seamed with darker. Vena-
tion: Sc; ending slightly beyond, Sce exactly opposite, the origin of Rs;
Rs angulated to almost square at origin; cell 1st Mz long, closed, about
equal in length to the veins issuing from it; basal deflection of Cm at
the fork of M. :

Abdomen dark brown. Ovipositor reddish, the tergal valves very
slender.

Habitat: Japan.
Holotype, 2, Meguro, Tokio, April 10, 1919 (R. Takahashi).

Dicranomyia atripleura sp. n.

Head light grey; mesonotum brownish grey, the praescutum with
three pale brown stripes; thoracic pleura with a dark brown longitudinal
stripe; legs pale brown; wings greyish subhyaline, stigma rounded,
brown; Sc short, cell 1s¢ Mz closed, about twice as long as vein_Cm
beyond it.

ies L,
i '
1919] { Alexander: Japanese Crane-Flies 329

Female—Length about 6.3 mm.; wing, 6.5 mm.

Rostrum and palpi brown. Antenne with the scapal segments pale
brown; flagellum broken. Head light grey.

Pronotum brownish grey. Mesonotal praescutum pale brownish
grey, with three pale brown stripes, the median stripe broad, the lateral
stripes indistinct; scutum pale, the lobes darker; scutellum and _ post-
notum heavily light grey pruinose, the latter with a darker median line.
Pleura pale, yellowish grey pruinose; a broad, very conspicuous, dark
brown stripe, extending from the propleura to the base of the abdomen,
passing above the fore coxe and beneath the halteres. Halteres pale
yellow basally, the knobs dark brown. Legs with the coxe and troch-
anters pale yellow; femora and tibiz pale brown; tarsi darker brown.
Wings greyish subhyaline; stigma rounded, brown; veins dark brown,
C, Sc and the veins at the wing-base more yellowish. Venation: Sc
short, Sc, ending a little beyond the origin of Rs, Sc. exactly at the
origin; Rs less than twice the deflection of R145; 7-m about equal to m, a
little shorter than 7; cell 1st M2 long, about twice the section of Cm
beyond it; basal deflection of Cu; immediately before the fork of M.

Abdomen dark brown, the sternites more yellowish brown. Ovi-
positor with the tergal valves slender, strongly upcurved; sternal
valves straight, the tips subacute.

Habitat: Japan.
Holotype, 9, Choshi, Chiba, October 17, 1919, (R. Taka-
hashi).

Dicranomyia mesosternata sp. n.

Antenne dark brown; head dark; mesonotum brownish black, the
pleura gray pruinose; wings pale gray with four brown costal marks and
pale gray clouds along the cord and at the ends of the veins; Sco appar-
ently lacking.

Female—Length, 7.3-8 mm.; wing, 8-8.4 mm.

Rostrum and palpi dark brown. Antenne dark brownish black
throughout, the flagellar segments elongate-oval, with rather long
verticils. Head dark brownish gray. Vertex very narrow between the
large eyes.

Pronotum conspicuous, brown, narrowed anteriorly. Mesonotal
praescutum shiny brownish black, the humeral regions with a paler
brown pollen; remainder of the mesonotum brown, the scutal lobes
brownish black. Pleura shiny dark brownish black, light gray pruinose,
most heavily across the dorsal edge of the mesosternum, which is shiny
blackish between the fore and middle coxe. Halteres with the base of
the stem pale, the knobs dark brown. Legs with the coxe yellowish
brown, darkest on the fore coxz; trochanters dull yellow; femora brown,
paler basally, the tips dark brown; tibiz and tarsi dark brown. Wings
pale gray, with a heavy dark brown and brownish gray pattern, as
follows: four large brown areas along the costal margin, the first at

330 Annals Entomological Society of America [Vol. XII,

the arculus, the third at the tip of Sc; and the origin of Rs, the last being
the stigmal blotch; wing-apex darkened; large, pale brownish gray clouds
along the cord, outer end of cell 1st Mz and at the ends of the longitudinal
veins; veins dark brown. Venation: Sc short, Sc; ending opposite or
slightly beyond the origin of Rs, Scz lacking; a supernumerary cross-vein
at about midlength of cell Sc, located in the second anterior brown
blotch; Rs about two and one-half times as long as the basal deflection of
Riis; cell 1st Mz closed; basal deflection of Cm, before, at or slightly
beyond the fork of M.

Abdominal tergites dark brown, the sternites paler brown. Male
hypopygium with the ninth tergite broad, the caudal margin gently
concave. Ninth pleurite short; ventral pleural appendage pale, the
proximal face produced into a stout beak, set with two widely separated
spines, one subapical, the other basal. Gonapophyses a little longer
than half the length of the penis-guard, slender and gently curved.

Habitat: Japan.

Holotype, o', Meguro, Tokio, October 20, 1919, (R.
Takahashi).

Allotopotype, 9, July 9, 1919.

Paratopotypes, 4c’s, October 20-21, (1919 Snore sex
uncertain, July 12, 1919.

One paratype bears the label, “‘ This insect was flying about
over a stream.”’

Antocha Osten Sacken.

Proantocha, subgen. n.

Antennz short, the flagellar segments oval, the terminal segment
nearly as long as the preceding two taken together. Head very small,
broad, the small eyes protuberant, widely separated both above and
below. Thorax very large, the mesonotum convex. MHalteres short.
Legs very stout, the fore and middle legs covered with conspicuous
long erect hairs, the posterior tibize with numerous tiny black spines;
the coxee, especially the posterior ones, very large and globular, the hind
trochanters being long, narrow, compressed; hind legs very long,
especially the tibia, which is slightly curved before its tip; at the base on
the ventral face a long, slender spine which meets a similar but stouter
tubercle located near the apex of the femur; tarsi remarkably shortened,
especially those of the hind legs where they are less than one-fourth
the length of the tibia; claws long and straight, at about mid-length
with a long straight tooth that is about two-fifths the length of the
apex of the claw alone, an additional tiny basal tooth. Venation as in
the subgenus Antocha. Abdomen short.

Type of the subgenus—Antocha (Proantocha) spinifer sp. n.
(Japan).

1919] Alexander: Japanese Crane-Flies Bie

The genus Antocha has, till now, been one of the most
compact genera in the Tipulidae. The discovery of Antocha
spinifer renders it necessary to subdivide the group. This
latter is a curious fly with long, stout, very hairy legs that
suggests in its general appearance a Trimicra or an Empedo-
morpha rather than an Antocha. The venation alone is normal
for this latter genus. The discovery of the immature stages of
this isolated, generalized fly will possibly result in giving generic
rank to the group here proposed.

Antocha (Proantocha) spinifer sp. n.

Size very large (wing of male over 12 mm.); legs very long and
stout, the fore and middle legs provided with long, dense hairs, the hind
tibiae set with numerous tiny spines and with a large spinous tubercle
on the ventral side near the base.

Male—Length about 9 mm.; wing, 12.3 mm. Fore leg, tibia,
7.8 mm.; tarsus, 4.8 mm.; middle leg, tibia, 7.4 mm., tarsus, 3 mm.;
hind leg, tibia, 13.8 mm., tarsus, 3 mm.

Rostrum short, dull yellow; palpi short, brownish yellow, the
terminal segments more infuscated. Antennze with the scape and basal
three or four segments of the flagellum dull brownish yellow, the
remainder of the antenne brown. Head dull yellow, very sparsely
light gray pruinose.

Mesonotum dull brownish yellow, the praescutum darker brown
medially, the lateral margins -pale. Pleura dull yellow. Halteres
pale, the knobs slightly brownish. Legs with the coxz and trochanters
dull yellow; remainder of the legs dull brownish yellow, only the terminal
tarsal segments brown; the long, dense hairs that cover the fore and
middle legs are dark brown and obscure the ground color of the sclerites
that bear them. The hind legs are covered with numerous blackened
spinous setigerous tubercles. Claws very long, dark brown, longer
than the fourth and nearly as long as the fifth tarsal segment. Wings
milky white, stigma indistinct, faintly yellowish; veins brown, the
costa beyond the point of insertion of Sc tawny and somewhat incras-
sated. Venation: r tending to be obliterated by atrophy; basal deflec-
tion of Cu far before the fork of M.

Abdominal tergites brownish buff, with a distinct dark brown
median stripe that is interrupted at the posterior margins of the seg-
ments; sternites reddish, on the terminal segments darker, brownish,
the caudal margin of the segments broadly pale.

Habitat: Japan.
Holotype, o, Saitama, June 1, 1919 (R. Takahashi).

332 Annals Entomological Society of America [Vol. XII,

Antocha (Antocha) satsuma sp. n.

Head rusty brown; thorax gray, the praescutum with three brown
stripes; wings milky gray, the stigma dark brown; veins Sc, R; and C
beyond the stigma chestnut brown; basal deflection of Cm, before the
fork of M.

Male—Length, 5.3-5.5 mm.; wing, 7.3-7.6 mm.

Female—Length, 6.8 mm.; wing, 8 mm.

Rostrum pale reddish brown; palpi dark brown. Antenne brown,
the flagellar segments a little paler basally; scapal segments large,
tumid. Head pale rusty brown, with a faint bronzy tinge.

Pronotum grayish yellow, broadly dark brown medially. Mesonotal
praescutum grayish yellow with three broad dark brown stripes, the
median stripe broadest, not attaining the anterior margin of the sclerite;
scutum gray, the lobes dark brown medially; scutellum and postnotum
gray. Pleura yellowish gray. Halteres very pale yellow. Legs long
and slender, the coxe pale, sparsely gray pruinose; trochanters dull
yellow; femora pale brown, more yellowish basally; tibize and tarsi pale
brown. Wings milky gray, pale at the base, the stigma dark brown;
slightly darker clouds along veins Rs and 2nd A and at the wing apex;
veins dark brownish black, M and Rs pale basally; Sc, Ri and costa
beyond Sc, light chestnut brown. Venation: Rs long and straight,
the base rather indistinct; r opposite r—m; basal deflection of Cm far
before the fork of M, this distance variable; cell 1st M> small.

Abdomen dark brownish gray, the sternites paler medially, in the
female the posterior half of the intermediate abdominal segments
is slightly paler than the basal half. Male hypopygium yellowish,
the pleurites rather stout; the two pleural appendages are slender,
subequal in length, the dorsal hook chitinized, at the tip narrowed into
a slender point; ventral pleural appendage fleshy with scattered setze
that are larger and stouter at the tip of the organ. Gonapophyses long,
acicular, almost straight, the tips acute.

Habitat: Japan.

Holotype, «#, Meguro, Tokio, April 21, 1919 (R. Takahashi).
Allotopotype, 2, March 25, 1919.

Paratopotype, 2¢’s, March 26—April 28, 1919.

Dicranoptycha Osten Sacken.

Dicranoptycha yamata sp. n.

Antenne black, the scapal segments dull yellow; general coloration
gray, the praescutum with three brownish stripes; legs black, the bases
of the femora narrowly yellowish; abdomen dark brownish black.

Female—Length about 11 mm.; wing, 12.7 mm.

Rostrum and palpi black. Antenne with the scapal segments dull
brownish yellow, the second segment brightest; flagellum black, the
base of the first segment paler. Head light gray.

1919} Alexander: Japanese Crane-Flies 309

Pronotum brownish gray. Mesonotal praescutum dull gray, clearer
anteriorly, with three slightly darker brownish stripes; remainder of the
mesonotum clear light gray, only the scutal lobes a little darker. Pleura
gray. Halteres yellow. Legs with the coxe grayish, the mesocoxe
more yellowish; trochanters yellow; femora dark brownish black, the
bases narrowly yellow; remainder of the legs dark brownish black.
Wings with a strong grayish yellow tinge, somewhat darker distally, the
costal and subcostal cells clearer yellow; veins C and Sc yellow, the
remaining veins dark brown. Venation: Rs strongly arcuated at its
origin, a little longer than cell 1st Me.

Abdomen dark brownish black. Ovipositor with the tergal valves
flattened, the outer faces with numerous erect yellow sete.

Habitat: Japan.

Holotype, ?, Meguro, Tokio, April 23, 1919 (R. Takahashi).
Paratopotypes, 22, April 24—May 5, 1919.

One specimen bears the label “‘in a pine forest.”’

Paratropeza Schiner.

Paratropeza flavitibia sp. n.

General coloration metallic blue; legs yellow, the femora clavate,
with a broad black subterminal ring; tibiz with the tips blackened, the
posterior tibiz with an apical fringe of erect black hairs; metatarsi
yellow, darkened at the tips; wings with three broad dark brown cross-
bands.

Female—Length, 6.8 mm.; wing, 4.7mm. Fore leg, femur, 2.9 mm.;
tibia, 2.6 mm.; hind leg, femur, 4 mm.; tibia, 3.3. mm.; tarsus, 2.4 mm.

Description from an alcoholic specimen.

Rostrum and palpi pale brown, the terminal segments of the latter
about one-half longer than the third segment. Antenne brown, the
second scapal segment and the base of the first flagellar segment more
yellowish; flagellar segments oval. Head dark metallic blue.

Thorax metallic blue, with only the dorso-pleural membranes
whitish. Halteres dark brown, the knobs whitish. Legs with the
coxe metallic blue on their outer faces; trochanters yellowish brown;
fore femora dark brown, more yellowish on the basal half, the extreme
tip and a very indistinct band at about one-third the length dull yel-
lowish; middle and posterior femora yellow with a broad dark brown
subterminal ring; tibiae yellow, more brownish basally, the tips broadly
dark brownish black, broadest on the hind legs; metatarsi yellow, only
the tips brown; remaining tarsal segments dark brown; on the posterior
metatarsi the brown tips occupy one-third of the total length. The
legs are provided with flattened scales; the femora are strongly clubbed,
especially the posterior femora; the posterior tibiae with a broad apical
fringe of outspreading black hairs. Wings hyaline, with three broad .
brown cross bands, the first proximad of the origin of the sector, extend-
ing from the costal to the posterior margin; the second band occupies

334 Annals Entomological Society of America |Vol. XII,

the cord; the third the wing apex; costal and subcostal cells brownish
yellow; a small brownish spot at the origin of. the sector, sometimes
confluent with the basal crossband. Venation: almost exactly as in
P. ornatipennis (de Meij.); Sc short, Sc; extending to about two-fifths
the length of Rs, Sco being proximad of the origin of the latter.

Abdomen dark metallic blue; ovipositor yellowish corn color,
the sternal valves darker; tergal valves strongly curved.

Habitat: Japan:

Holotype, @, Chichibu, Province Saitama, October, 1918
(R. Takahashi).

Edwards: (Ann. Mag, ‘Nat. Hist. «ser. 8, vol. 17; p. 356:
vol. 18, p. 249; 1916) records this new species from Idzu,
Japan (June, 1910), but does not give it a name. P. flavitibia
is closest to P. ornatipennis (de Meij.) but is sufficiently dis-
tinguished by the broad complete basal wing band and the
different leg pattern. The brush of erect or nearly erect hairs
at the tips of the posterior tibia is very conspicuous and sug-
gests the condition found in the recently described P. pennipes
(Brunetti) of India (Records Indian Mus., vol. 15, p. 308, 1918;
as gymnastes). This fly differs from flavitibia in the pattern
of the legs and wings.

Ormosia Rondani.

Ormosia diversipes sp. n.

Belongs to the nigripila group; general coloration black, including
the femora, the tibiz and tarsi abruptly light yellow; wings gray, the
stigma brown, the costal margin with yellowish pubescence; anal veins
divergent.

Male—Length, 4-4.38 mm.; wing, 5.1-5.5 mm.

Rostrum and palpi dark brownish black. Antennze with the basal
four or five segments yellowish brown; remainder of the flagellum dark
brown; flagellar segments (in the male sex) elongate-cylindrical, with
abundant erect whitish pubescence that is little shorter than the length
of the segment that bears it. Head dark.

Pronotal scutellum pale. Mesonotal praescutum blackish with
a heavy reddish brown pollen, the dorso-median area darker, producing
an indistinct stripe. Pleura shiny blackish; a brush of more than a
score of long erect yellowish hairs on the pleura between the bases of
the wings and halteres. Halteres light yellow. Legs with the coxe
dark brownish black, sparsely pruinose; trochanters dull yellowish
brown; femora black, a little lighter basally; tibiae and metatarsi abruptly
light yellow, the tips narrowly darkened; remainder of the tarsi dark
brown. Wings gray, the stigma more brownish, the costal region
more yellowish; base of the wing and an indistinct area near the fork

1919] - Alexander: Japanese Crane-Flies 3935

of M pale; veins brown; wings broad, densely covered with hairs.
Venation: Cell 1s¢ Mz small, closed; basal deflection of Cm slightly
before the fork of M; anal veins strongly divergent.

Abdomen black. Hypopygium of the general structure of the
nigripila group of the genus; pleurites stout, one pleural appendage
much longer than the others, dusky in color, tapering gradually to the
acute tip; the second appendage is shorter, flattened, subhyaline. The
gonapophyses are flattened blades, deeply bifid apically with the
proximal arm about twice the length of the outer arm.

Habitat: Japan.

Holotype, &, Meguro, Tokio, March 31, 1919 (R. Taka-
hashi).

Paratopotypes, 1 o&, March 26; 2.7’s, April 16, 1919.

Ormosia atripes sp. n.

Belongs to the nigripila group; general coloration black, including
the legs; wings grayish brown; anal veins divergent.

Female—Length 5-5.3 mm.; wing, 5.8-5.9 mm.

Rostrum and palpi dark brownish black. Antenne with the first
scapal segment brown, second segment large, broadly pyriform, light

yellow; flagellum dark brownish black, the segments oval (in the female
sex). Head dark.

Sides of the pronotal scutellum yellowish white. Mesothorax
blackish with a sparse dull yellowish brown pollen; scutellum with long
coarse yellow bristles. Pleura black, a sparse group of from six to eight
long erect yellowish hairs between the bases of the wings and halteres.
Halteres light yellow. Legs with the coxe brownish black; trochanters
yellowish brown, brighter basally; femora dark brownish black, paler
basally; remainder of the legs brownish black. Wings with a dark
grayish brown suffusion, brighter basally and in the costal region;
stigma indistinct, brown; a faint whitish spot beyond the stigma and
a larger, somewhat more distinct, one at the fork of M; veins dark brown,
R and Cu yellowish; wings broad with a dense delicate pubescence.
Venation: r just beyond the fork of Roy; on Re; cell 1st Mz small, closed,
the inner end narrowed, the outer deflection of M3 a little longer than m;
basal deflection of Cm at or slightly before the fork of M; anal veins
strongly divergent.

Abdomen black; tergal valves of the ovipositor strongly curved,
black basally, dark horn yellow apically; sternal valves dark, the tips
acute.

Habitat: Japan.

Holotype, 2, Meguro, Tokio, April 10, 1919 (R. Takahashi).
Paratopotypes, 7 2’s, April 9-25, 1919.

306 Annals Entomological Society of America [Vol. XII,

Ormosia tokionis sp. n.

Belongs to the fascipennis group; general coloration gray; femora
yellowish at the base; wings pale gray with a narrow pale brown seam
along the cord.

Male—Length, about 3.8 mm.; wing, 5.2 mm.

Female—Length, about 4.5 mm.; wing, 6.1 mm.

Rostrum dark with a gray pollen. Palpi dark brown. Antenne
dark brown, short in both sexes, the flagellar segments oval in the male,
clothed with a dense white pubescence. Head grayish brown, a little
clearer brown along the margin of the eyes.

Mesonotal praescutum yellowish gray with four indistinct reddish
brown stripes; tuberculate pits a little anterior to the level of the
pseudosutural foveee, separated from one another by a distance equal
to, or a little less than. the diameter of one; scutellum and median area
of the scutum lighter; postnotum gray. Pleura gray. Halteres rather
long, dark brown, the base of the stem yellowish. Legs with the coxz
dark, covered with a yellowish gray pollen; trochanters dull brownish
yellow; femora brown, the bases more yellowish, darker brown at the
tips; tibie and tarsi brownish black. Wings pale gray, the veins
narrowly and indistinctly seamed with brownish, more distinct along
the cord; stigma large, brown; costal and subcostal cells a little more
yellowish; veins dark brown. Venation: 7 on R», about its own length
beyond the fork of R243; deflection of Ri;; longer than r-m; cell 1st Mz
closed; basal deflection of Cu; beyond the fork of M; anal veins divergent.

Abdomen dark brown. Male hypopygium small, the pleurites rather
slender, nearly cylindrical; a single pleural appendage that is narrowed
to the blunt, blackened apex, the surface with abundant setigerous
punctures. Penis-guard a narrow, triangular chitinous plate, which
has a narrow median apical point.

Habitat: Japan.

Holotype, &, Meguro, Tokio, March 23, 1919 (R. Taka-
hashi).

Allotopotype, ¢@, March 23, 1919.

Paratopotype, co’, March 25, 1919.

Ormosia cinctifer sp. n.

Belongs to the fascipennis group; general coloration black, the body
sparsely dusted with gray; wings nearly hyaline, a narrow brown seam
along the cord.

Male—Length, 4.5-4.7 mm.; wing, 6.4-6.5 mm.

Female—Length, 5.6-5.8 mm.; wing, 6.3-7 mm.

Rostrum and palpi black. Antenne black, short in both sexes, in
the male the intermediate segments rounded oval, the apical segments
long-oval. Head dark gray with a sparse brownish yellow pollen.

Mesonotal praescutum black, sparsely dusted with gray, with three
blackish stripes, the median one shiny, the lateral stripes less distinct;

1919] Alexander: Japanese Crane-Flies 337

scutum similar, each lobe with a long-oval blackish area; scutellum and
postnotum black, sparsely gray pruinose. Pleura black, gray pruinose.
Halteres light yellow, the base of the stem dusky. Legs black. Wings
nearly hyaline; stigma dark brown, conspicuous, sending a narrow brown
seam along the cord; outer end of cell /st M2 and vein Cu less distinctly
seamed with brown; veins dark brown; the base of the wing more
yellowish; wings with the pubescence rather short and sparse, almost
lacking in the basal cells, including most of cell WM. Venation: Sc
ending just beyond r; r on Re near its base; cell 1st Mz closed; m shorter
than the deflection of M73; basal deflection of Cm slightly before the fork
of M; 2nd anal vein slightly sinuous, but cell /s¢ A broadest at the wing
margin.

Abdomen black, including the male genitalia. Male hypopygium
with the pleurites stout; pleural appendage oval, tumid, with long
yellowish hairs, on the dorsal face with a chitinized plate, each side
of which is heavily toothed, the outer margin with from five to six teeth
and the inner margin with about three teeth; gonapophyses flattened,
slender, tapering to the subacute tips. Ovipositor with the tergal valves
horn color, darker basally.

Habitat: Japan.

Holotype, «, Meguro, Tokio, April 2, 1919 (R. Takahashi).
Allotopotype, ¢, April 2, 1919.

Paratopotypes, 207’s, 1 9, March 25—April 10, 1919.

Ormosia takahashii sp. n.

Belongs to the varia group; general coloration black, gray pruinose;
wings grayish with the stigma brown; cell /s¢ Ms. open; anal veins con-
vergent; male hypopygium with eight chitinized points surrounding
the penis-guard.

Male—Length, 3.8—4.2 mm.; wing, 5.4-5.5 mm.

Female—Length, 4.6 mm.; wing, 6.2 mm.

Rostrum and palpi dark brown. Antenne brown, the flagellar
segments long-oval, covered with a dense white pubescence, the flagellum
with a single very long verticil before midlength of each segment to
give the organ a secund appearance. Head dull gray.

Mesonotum dull brownish gray, the praescutum with indistinct,
somewhat darker brown stripes, the interspaces with a row of coarse
yellowish hairs; tuberculate pits very close together. Scutum dull gray,
the lobes darker. Pleura black, sparsely gray pruinose; a few very long
yellow hairs ventrad and caudad of the wing-base. Halteres yellow.
Legs with the coxe dark, sparsely gray pruinose; trochanters dull yellow;
femora dark brown, the bases yellow; tibie and tarsi dark brown,
clothed with a golden-yellow pubescence that is especially distinct in
the female. Wings gray, the stigma darker brown, lying beyond r;
veins dark brown, the base of the wing slightly brighter. Venation:
Cell 1st Mz open by the atrophy of the outer deflection of M3; basal

338 Annals Entomological Society of America [Vol. XII,

deflection of Cu; just before the fork of M; 2nd anal vein sinuous, at its
tip bent rather strongly toward the first anal.

Abdomen dark brownish black. Male hypopygium with the pleurites
rather stout, covered with sparse prominent tubercles that bear coarse
yellow sete, which are longest at the tips of the pleurites; pleural
appendages two, the outer appendage fleshy, the outer face with an
abundant short brownish, appressed pubescence, arranged in trans-
verse rows; inner appendage narrowed and chitinized at the tip, the
inner face with about nine or ten stout setae. Penis-guard terminating
in two subparallel slender black spines; two pairs of gonapophyses, the
inner pair simple and ending in an acute blackened point; outer gona-
pophyses bifid, the lateral arm much longer and more slender than the
proximal arm. Eighth sternite with a short, flattened, subspatulate
median lobe.

Habitat: Japan.

Holotype, &, Meguro, Tokio, April 2, 1919 (R. Takahashi).

Allotopotype, 2, April 7, 1919.

Paratopotypes, 19 o& 9, March 25—April 25, 1919; paratype,
@, May 29, 1919; 66 9, October 2-21, 1919:

The autumnal specimens are much smaller than the vernal
material, but show no other differences.

It is with great pleasure that this interesting:fly is dedicated
to its discoverer, Mr. Ryoichi Takahashi.

Erioptera Meigen.

Erioptera bifurcata sp. n.

General coloration gray, the mesonotum with three brown stripes;
legs black with whitish scale-like hairs; wings yellowish, vein Sc yellow;
cell 1st Mz open; anal veins divergent.

Male—Length, 5—5.5 mm.; wing, 6.1—7.1 mm.

Rostrum black, densely gray pruinose; palpi dark brown. Antennz
black, the segments oval, covered with a dense white pubescence, the
verticils short, barely projecting beyond the pubescence. Head-broad,
light gray, the eyes widely separated above, the ventro-caudal angles
almost contiguous beneath.

Pronotum light gray. Mesonotal praescutur gray with three
brown stripes, the median stripe broad, slightly ~tricted by the
conspicuous, elongate pseudosutural foveze, indistinctly bisected by a
pale median line; tuberculate pits small, very widely separated, lying
slightly anterior to the level of the pseudosuturak -.seutum gray,
the lobes indistinctly marked with brown; remainder ot the mesonotum
gray. Pleura clear light gray. Halteres light yellow, the extreme base
dark. Legs with the coxe and trochanters black, gray pruinose;
remainder of the legs black, densely covered with flattened whitish,
scale-like hairs. Wings with a faint yellow tinge, the base of the wing

1919] Alexander: Japanese Crane-Flies 309

more yellowish; stigma very narrow and indistinct, brown, confined to
vein Ri; veins dark brown, Sc yellow. Venation: Sc long, ending
opposite r; r on Ry about its own length beyond the fork of Ro43; cell
1st Mz open by the atrophy of m; basal deflection of Cu; beyond the
fork of M; anal veins divergent.

Abdominal tergites gray, the sixth and seventh segments with a
transverse median brown blotch; the basal segments on either side with
blackish impressed rectangular areas; pleural appendages of the hypo-
pygium yellowish with the tips blackened. Male hypopygium with
the ninth tergite deeply notched medially; pleurites rather slender,
at the apex with two pleural appendages; dorsal appendage long,
slender, the tips blackened; ventral pleural appendage short, deeply
bifid, the cephalic or proximal arm more slender, at its tip with a single
long bristle; the caudal or lateral arm broader, its flattened apex densely
set with roughened tubercles. Gonapophyses very long, almost straight,
tapering to the acute tips.

Habitat: Japan.
Holotype, &, Meguro, Tokio, April 4, 1919 (R. Takahashi).
Paratopotypes, 4 o&’s, April 4-9, 1919.

Limnophila Macquart.

Limnophila (Lasiomastix) pilifer sp. n.

Antenne short in both sexes; wings yellowish at the base, the apical
cells strongly pubescent.

Male—Length 6-7 mm.; wing, 8.6-8.7 mm.

Female—Length, 8.5-9.5 mm.; wing, 8.8 mm.

Rostrum and palpi dark brownish black. Antennz short in both
sexes; first scapal segment reddish brown, sparsely gray pruinose,
second segment reddish; basal segments of the flagellum brown, paler
basally, apical flagellar segments more elongate, dark brown; flagellar
segments clothed with a pale pubescence. Head gray.

Pronotum large, dark colored, light gray pruinose. Mesonotum
shiny jet black with an easily removed, but dense pollen, brownish
yellow on the median area of the praescutum, clearer gray on the
remainder of the thorax, including the pleura. Halteres light yellow.
Legs with the fore coxe black, grayish pruinose; middle cox dull
brownish yellow, dark at the extreme base; hind coxe largely brownish
yellow; trochants:ssdull yellow; femora yellow, the tips broadly dark
brown; tibiz yellowish brown, the tips darkened; tarsi dark brownish
black,. the base of ‘he metatarsi brown. Wings grayish, strongly
yellow at the ho ; color including most of the costal and subcostal
cells; stigiua oval, grayish brown, almost evenly split by 7; narrow gray
seams along the cord, outer end of cell 1st Me, origin of Rs, and the forks
of Rois and Mi42; veins brown, those in the costal and basal regions
yellowish; apical cells of the wings strongly pubescent, this extending
from cell 2nd Ri to cell Cu;. Venation: Sc2 slightly removed from the

340 Annals Entomological Society of America [Vol. XII,

tip of Sc; Rs long, in alignment with R243, this latter shorter than the
basal deflection of Ci; , removed from the tip of Ri; inner ends of cells
R3, Rs, and 1st My» in oblique alignment; petiole of cell M1 about equal
to or shorter than this cell; basal deflection of Cm slightly before mid-
length of cell 7st Mz. Some of the specimens in this series show abnormal
types of venation.

Abdomen dark brownish black, the hypopygium a little reddish.
Male hypopygium with the outer appendage almost straight, split
into two short points at the apex; the inner appendage deeply bifid, the
outer arm flattened, subspatulate, the blade setigerous, the inner arm
much smaller and narrowed to the blunt apex; gonapophyses slender,
acicular, much twisted. Valves of the ovipositor rusty horn color,
very long and slender, the tergal valves almost straight, slightly upcurved
at the tips.

Habitat: Japan.

Holotype, &, Meguro, Tokio, April 8, 1919 (R. Takahashi).
Allotopotype, @, April 19, 1919.

Paratopotypes, 18 o @,. April 7-25, 1919.

Limnophila (Ephelia) subaprilina sp. n.

Mesonotal praescutum yellowish gray, the lateral brown stripes
connected with the intermediate pair by a subtransverse brown lne;
wings with a heavy brown pattern; abdominal segments bicolorous.

Male—Length, 5.6 mm.; wing, 7.38-7.4 mm.

Female—Length, 8.7 mm.; wing, 8.4 mm.

Rostrum and palpi dark brown. Antenne with the scapal segments
dark brown, the basal flagellar segments pale brown, the apical segments
darker. Head brownish gray, more yellowish along the inner margin
of the eyes and on the front.

Mesonotal praescutum yellowish gray with broken dark brown
stripes; the intermediate stripes are separated from one another by a
distance that is nearly equal to the width of one, indistinct in front,
not reaching the suture behind; lateral stripes short, their cephalic
ends with a transverse bar to form a T-shaped mark that connects
more or less completely with the intermediate stripes and with the
lateral margin of the praescutum; scutum yellowish gray; scutellum
and postnotum clearer gray. Pleura yellowish gray with conspicuous
dark brown blotches. Halteres yellow, the knobs dark brown. Legs
with the coxz dark brown; trochanters brown, with a black spot on the
posterior face; femora dull yellow, the tips broadly dark brown; tibiz
brownish yellow, the tips narrowly and indistinctly darkened; tarsi
dark brown, the first and second segment yellowish brown with only
the tips darkened. Wings grayish subhyaline, the costal and subcostal
cells more yellowish; a heavy dark brown pattern along the costal
margin and large grayish brown blotches on the cross veins, deflections
of veins, and along the margin at the ends of the longitudinal veins;
there are seven costal blotches, the basal one broken, subocellate; the

1919] Alexander: Japanese Crane-Flies 341

third at the origin of Rs extending from costa almost to M; the fifth,
largest, at the stigma; the last two at the ends of Ri and Re» respectively ;
a narrow cloud in the anal angle of the wing; the seam along the super-
numerary cross vein sends a very black line along vein Cu to form an
inverted T; veins C and Sc yellow, the other veins brown. Venation:
Rs strongly arcuated at origin; Roy3 and M142 nearly equal in length.
Abdominal segments bicolorous, the basal half of each segment dull
reddish, the apical half blackish. Male hypopygium with the outer
pleural appendage stout, on the outer face before the tip with one
conspicuous spine in addition to the apical, slightly curved, tooth.

Habitat: Japan.

Holotype, &, Meguro, Tokio, April 20, 1919 (R. Takahashi).

Allotopotype, @, April 19, 1919.

Paratopotype, o, April 19, 1919; paratype, o&, Tokio,
May, 1919.

Limnophila (Prionolabis) rufipennis sp. n.

General coloration blackish, femora a little paler basally; wings
brownish yellow with sparse brown seams along the cord; cell 1st M2
subquadrate, basal deflection of Cu; beyond the middle of its length.

Male—Length about 14 mm.; wing, 16.4 mm., its greatest width,
5.0 mm.

Description from an alcoholic specimen.

Rostrum, palpi and antenne black. Head black, very possibly
grayish pruinose in dried specimens.

Thorax black throughout in alcoholic material. Halteres long,
light yellow, the knobs a little darker. Legs with the coxe black;
trochanters dark brown; femora dark brown, only a little paler basally,
especially on the hind femora; tibiz and tarsi dark brownish black.
Wings very broad and ample, deeply suffused with pale brownish yellow,
clearer yellow in the costal and subcostal cells; stigma oval, dusky
brown; very narrow dusky brown seams along the cord, at the outer
end of cell /st Mo, at the origin of Rs and at the fork of Mi42. Venation
generally similar to L. (P.) rufibasis O. S. (Northeastern North
America), but cell /st M2 much shorter, subquadrate; R43 a little
longer than r-m; basal deflection of Cu, beyond mid-length of cell
1st M2; petiole of cell M; a little shorter than this cell.

Abdomen dark brownish black. Male hypopygium with the ventral
pleural appendage irregularly pectinated, one tooth being considerably
larger than the others; dorsal pleural appendage with the apex simple,
blackened.

Habitat: Japan.
Holotype, o, Chichibu, Province Saitama, October, 1918,
(R. Takahashi).

342 Annals Entomological Society of America [Vol. XII,

Polyangezus Doane.

Polyangzus japonicus sp. n.

Thorax yellow, the praescutum with an indistinct darker median
stripe; wings with a heavy brown banded pattern; crossvein r lacking,
m present; supernumerary crossveins in cells R3, Rs, Mi and M; tibie
and tarsi largely white.

Male—Length, 6.6 mm.; wing, 8 mm.

Head small, closely applied to the thorax. Rostrum and palpi
dark brown. Antenne short, the flagellar segments crowded, the
scape dark brown, the flagellum light brownish yellow, the last segment
infuscated. Head dark brownish gray.

Mesonotum light yellow, the praescutum with a broad pale brown
median stripe that is broadest and most distinct in front, becoming
obliterated behind; postnotum brownish. Pleura yellow. Halteres
very long and slender, dark brown, the base of the stem yellow. Legs
with the coxe and trochanters dull yellow; femora dull yellow, the tips
broadly dark brown; tibiz china-white, the tips broadly dark brown;
tarsi white, the apical segments more infuscated. Wings subhyaline
with a heavy banded pattern of dull yellowish, margined with brownish
gray; the costal cell is entirely dark; interrupted crossbands occur at
the origin of the sector, along the cord and very diffusely between the
cord and the wing-tip; dark spots at the ends of all the longitudinal
veins, large and blotch-like at the ends of the anal veins; a row of small
dots in cell Cu, behind the abortive anal vein in that cell; two spots
in the base of cell /st A; the apical pattern consists of yellowish seams
to the crossveins, these markings broadly margined with brownish
gray to produce a somewhat ocellate appearance. Venation: Rs
square and spurred at its origin; r lacking but a strong spur at the angle
of Roz; at the point where r is located in the genotype, P. maculatus;
Ro4s44 longer than r—m; cell 1st Me closed, m being located between Me
and M3; supernumerary crossveins in cells R3 near the tip, Rs at about
two-thirds its length, at about midlength of MM, and in cell M about
in alignment with the angulated base of the sector.

Abdomen brown, the segments margined posteriorly with darker
brown.

Habitat: Japan.

Holotype, o’, Saitama, May 31, 1919 (R. Takahashi).

Polyangaeus japonicus agrees with the genotype, P. macula-
tus Doane (Western United States) in most features of its
organization. It differs in the lack of the radial crossvein,
the closed cell 1st Mz and the presence of an additional super-
numerary crossvein in cell M4.

1919] Alexander: Japanese Crane-Flies 343

Rhaphidolabis Osten Sacken.
Rhaphidolabis flavibasis sp. n.

Antenne with fourteen segments, the last segment enlarged; meso-
notal praescutum light gray, with three dark brown stripes; base of the
wings strongly yellowish; male hypopygium with the outer angle not
produced into a lobe, the appendages borne at the narrow apex.

Male—Length, 7 mm.; wing, 8.7-8.9 mm.

Female—Length, 7.5-8 mm.; wing, 9.4-9.8 mm.

Rostrum and palpi dark brown. Antenne dark brownish black,
the first scapal segment gray pruinose, the second scapal segment a
little brighter apically; there are twelve flagellar segments which grad-
ually decrease in size to the end of the terminal, one much larger than
the penultimate and seemingly produced by the union of two small |
segments. Head light brownish gray, clearest along the inner margin
of the eye; an indistinct dark median line; vertical tubercle low.

The mesonotal praescutum light gray with three dark brown stripes,
the intermediate stripe entire; remainder of the mesonotum plumbeous,
gray pruinose. Pleura light gray. Halteres dark brown, the basal
third of the stem bright yellow. Legs with the coxe gray; trochanters
brown; femora dark brown, brighter on the basal third; tibia and tarsi
dark brownish black. Wings grayish subhyaline, the costal and
subcostal cells a little more yellowish; base of the wings bright yellow,
including the veins; stigma distinct, brown; veins dark brown. Venation:
Rs short, slightly arcuated to feebly angulate near midlength; Re
ublique, its fusion with R, at the wing tip slight; Roi3;4 rather short,
osually a little longer than r-m; cell My present; cell 1st Me open.

Abdomen dark brown, sparsely gray pruinose. Male hypopygium
with the pleurites short and stout, at the apex with the two small
pleural appendages, an outer subcircular fleshy lobe whose inner face
is densely set with short black spinous sete; inner pleural appendage
more chitinized, slightly curved, along the inner margin with about a
dozen spinous setz that are stouter and more crowded near the tip;
lateral gonapophyses slender, bent at a right angle near midlength.
Ovipositor rusty in color.

Habitat: Japan.

Holotype, <#, Meguro, Tokio, March 27, 1919 (R. Taka-
hashi).

Allotopotype, ?, April 4, 1919.

Paratopotypes, 8 o' 9, March 20-April 9, 1919.

One teneral female that may represent still another unde-
scribed species has cell R; sessile, the petiole being entirely
lacking. The praescutal stripes are less distinct than in the
specimens above described.

344 Annals Entomological Society of America [Vol. XII,

Dicranota Zetterstedt.

Dicranota nipponica sp. n.

Antenne with twelve segments, the terminal segment small; meso-
notal praescutum grayish buff, with three dark brown stripes, the
median one of which is split by a capillary line of the ground color;
male hypopygium at the outer angle produced caudad into a lobe.

Male—Length, 7.5 mm.; wing, 8.7 mm.

Rostrum and palpi brown. Antenne dark brownish black, the
flagellar segments with a conspicuous erect white pubescence in addition
to the verticils; but ten flagellar segments are evident, the last much
smaller than the penultimate; first flagellar segment long clavate, much
longer than those following. Head brownish gray, very narrowly
clearer gray along the inner margins of the eyes.

Thorax light grayish buff, the praescutum with thee dark brown
stripes of w hich the intermediate one is distinctly split by a capillary
vitta of the ground color; scutal lobes gray marked with pale brown;
scutellum, postnotum and pleura light gray. Halteres pale brown,
brighter at the base. Legs with the coxe light gray, more yellowish
apically; trochanters dull yellow, the outer margins blackened; femora
brownish yellow, brighter at the base, the tips dark brown; tibia dark
brown, the tips black; tarsi dark brownish black. Wings with a pale
gray tinge; stigma elongate, brown, extending from r to the tip of R14;
veins dark brownish black, the veins at the wing base a little paler.
Venation: Rs very short, strongly angulated to almost square at its
origin, slightly spurred; Ro4344 short, about equal to r-m; apical fusion
of Re with R, slight; cell 1st M2 open, cell Mj, present.

Abdomen dark brownish gray, the tergites very narrowly margined
caudally with paler gray; sternites similarly margined with yellowish
gray and with an indistinct broken median yellowish stripe. Male
hypopygium with the outer angle of the pleurites extended caudad into
a blunt hairy lobe; two pleural appendages, the inner one of which is
curved, flattened and expanded distally, not conspicuously armed with
setae and spines. }

Habitat: Japan.
Holotype, o&, Meguro, Tokio, March 30, 1919 (R.-
Takahashi).

Cylindrotoma Meigen.

Cylindrotoma japonica sp. n.

Antenne rather short, dark brownish black; front and a broad
margin around the eyes yellow; praescutal stripes entirely confluent, the
interspaces with a deep grooved impression.

Male—Length about 13-14 mm.; wing, 10.8-11.5 mm.

Female—Length, 13.5 mm.; wing, 13 mm.

1919] Alexander: Japanese Crane-Flies 345

Rostrum and palpi dark brown. Antenne rather short for this
genus of flies, dark brownish black, the flagellar segments with the
posterior face straight, the anterior or inner face gently convex. Front
and a broad margin around the eyes yellow, remainder of the head
shiny black, microscopically punctured.

Pronotal scutum dark, the scutellum yellow, darker medially.
Mesonotal praescutum with the disk black, subopaque, with deep,
impressed longitudinal grooves in the usual interspaces; scutal lobes
and the posterior half of the postnotum black, remainder of the
mesonotum yellow. Pleura pale whitish yellow, the sternopleura and
mesopleura, as well as an oval spot cephalad of the base of the halteres
shiny black. Halteres pale yellow, the knobs scarcely darker. Legs
with the coxe light brown, the outer face marked with darker brown;
trochanters dull yellow; femora yellow, the tips broadly black; tibize
brownish yellow, tipped with black; tarsi black, the base of the meta-
tarsi a little paler. Wings broad, grayish yellow, the basal and costal
areas brighter; stigma dark brown, oval; veins dark brown. Venation:
cell 1st Mz not greatly elongated; cell M, short-petiolate.

Abdominal tergites reddish brown, the segments margined laterally
and caudally with black; in some specimens these markings completely
cover the dorsum; sternites reddish, indistinctly margined laterally and
- caudally with dark brown; male hypopygium greatly enlarged, black;
ovipositor black.

Habitat: Japan.

Holotype, o&, Saitama, May 29, 1919 (R. Takahashi).
Allotopotype, 2, May 31, 1919.

Paratopotypes, 2's.

Liogma Osten Sacken.

- Liogma serraticornis sp. n.

General coloration deep shiny black; flagellar segments in the male
deeply serrated; wings with the stigma yellow or brownish yellow.

Male—Length about 15 mm.; wing, 12—12.1 mm.

Female—Length about 14 mm.; wing, 12 mm.

Rostrum dark brown; palpi yellowish brown. Antenne with the
scape brownish yellow, the flagellum dark brownish black, the flagellar
serrations in the male sex very long, considerably more so than in L.
kuwanai; there are seventeen segments, the last two of which are slender
and not serrate. Head shiny black.

Thoracic dorsum shiny black, the postnotum sculptured. Pleura
shiny black, the dorso-pleural membranes pale. Halteres yellow.
Legs with the coxe largely black on their outer faces; trochanters dull
yellow; femora pale yellow, the tips broadly black; tibize yellowish brown,
the tips blackened; tarsi black. Wings with a strong yellowish gray
tinge, the costal region, the stigma and the space between Cu and the
abortive anal vein behind it. yellowish; veins dark brown, excepting

346 Annals Entomological Society of America [Vol. XII,

Sc and R, which are yellowish. Venation: r-m sometimes present, but
short, at other times obliterated by the punctiform contact or short
fusion of Ry,; on M42; basal deflection of Cm before or slightly beyond
the fork of M.

Abdomen shiny black throughout.

In the female sex, the antennze are much shorter and less distinctly
serrated; only the second scapal segment is pale; the stigma and costal
region of the wing brownish yellow; the valves of the ovipositor short
and flattened, black, their tips rusty brown.

Habitat: Japan.

Holotype, , Saitama, May 30, 1919 (R. Takahashi).

Allotopotype, 9, May 29, 1919.

Paratopotypes, 2 o’s, 2 9’s, May 29, 1919.

Liogma kuwanai is a gray fly with the stigma dark brown
and the femora brown with only the bases yellow.

Phalacrocera Schiner.

Phalacrocera mikado sp. n. ‘

Antennz simple; mesonotum shiny black, the lateral margins of the
praescutum, the scutellum and the anterior half of the postnotum pale;
abdomen shiny black.

Male—Length about 11 mm.; wing, 12.2 mm.

Female—Length about 12 mm.; wing, 11.5-11.8 mm.

Rostrum and palpi dark brown. Antennz with sixteen segments,
simple in both sexes; in the male elongate-cylindrical, clothed with an
abundant white pubescence; antennee dark brown, the bases of the first
two flagellar segments more yellowish. Head with the vertex between
the eyes pale brown, the remainder shiny brownish black. Head
strongly narrowed behind.

Mesonotal praescutum shiny black, the lateral margins and the
humeral regions pale; scutal lobes black; remainder of the dorsum pale
except the posterior portion of the postnotum, which is darkened.
Pleura pale, the mesopleura and sternopleura shiny black. Halteres
pale brown, the base of the stem paler. Legs with the coxe pale, the
outer faces more or less darkened, especially on the mesocoxe; trochan-
ters pale; femora pale, the tips dark brown; tibize pale brownish white,
the tips dark; tarsi dark brown. Wings pale grayish, the stigma oval,
pale yellowish brown; veins brown. Venation: m and r-m present,
long; in the allotype the tip of vein Re is indistinctly persistent in both
wings, but in the other specimens there is no trace of this vein.

Abdomen shiny black; hypopygium small.

Habitat: Japan.
Holotype, <<, Tokio, April, 1919, (R. Takahashi).

Allotopotype, @.
Paratype, 2, Saitama, May 29, 1919, (R. Takahashi).

1919] Alexander: Japanese Crane-Flies 347

Nesopeza Alexander

Nesopeza tarsalis sp. n.

Mesonotal praescutum with the stripes shiny brown, confluent;
legs black, the. tips of the tarsi white; wings with the stigma dark
brown, with a pale area before and beyond it.

Male—Length about 10 mm.; wing, 12 mm.

’ Female—Length about 138 mm.; wing, 12.3 mm.

Frontal prolongation of the head and the palpi dark brown. Antennz
of the male moderately elongated, if bent backward extending about to
the base of the abdomen, the scape yellow, the flagellum dark brown.
Head yellowish, passing into dark brown on the posterior portions of
the vertex.

Mesonotal praescutum shiny brown, the stripes confluent, the
humeral angles yellowish; scutal lobes dark; remainder of the mesonotum
paler. Pleura yellowish with a shiny black streak or band extending
from the sternum across the mesopleura to the praescutum; a black
spot between the middle and hind legs. Halteres dark brown, the base
of the stem paler. Legs with the coxe and trochanters yellow; femora
dark brown, paler at the base; tibia and tarsi dark brownish black, the
tips of the latter pure white, these about equal in width on all the legs
or a trifle broader on the hind legs where about the apical half is pale.
Wings gray, the stigma dark brown; a whitish subhyaline area before
and beyond the stigma; a narrow seam on the basal deflection of Riis
and the tip of the wing faintly infuscated. . Venation: Rs not so elongate
as in N. geniculata and the medial forks deeper.

Abdomen shiny dark brown, the basal sternites and the sides of the
second and third tergites more yellowish; hypopygium brownish black.
In the female the abdominal tergites are uniformly dark.

In the living specimens the body is probably with decided greenish
tints as traces of these persist in the pinned individuals.

Habitat: Japan.

Holotype, &, Saitama, May 29, 1919 (R. Takahashi).
Allotopotype, @. .

Paratopotype, @.

Tipula Linnzus.

Tipula joana sp. n.

Antenne black, the scape orange-yellow; praescutum gray with three
darker gray stripes, the median stripe split by a capillary black line;
wings strongly petiolate, a seam along the cord and the apex brownish
black; 2nd Anal cell very narrow.

Male—Length, 18-20 mm.; wing, 17-18 mm. Fore leg, femur,
11 mm.; tibia, 13.6 mm.; hind leg, 13 mm.; tibia, 15.8 mm.

Frontal prolongation of the head reddish brown, the dorsal surface
light gray pruinose; nasus distinct. Palpi pale brown, the last segment,

r
!

4 <
As

348 Annals Entomological Society of America [Vol. XII,

with the exception of the base, light yellow. Antennal scape orange-
yellow, the flagellum black, the segments of the latter nearly cylindrical,
with a dense, microscopic white pubescence. Head light gray, the sides
of the vertex. more brownish, set with abundant sete.

Mesonotal praescutum light gray, with three darker gray stripes,
the median stripe broadest, strongly narrowed behind, split by a
capillary velvety-black line; scutum light gray, each lobe with two
darker gray blotches; scutellum and postnotum darker gray. Pleura
dark colored, with a dense blue-gray pollen, the dorso-pleural membrane
conspicuously dull yellow. Halteres brown, the extreme base brownish
yellow. Legs with the coxz blue-gray; trochanters yellow; femora
brownish-yellow, brightest at the base, the tips brownish black; tibize
brownish black, darkest apically; the posterior tibiae with a broad,
indistinct, whitish, sub-basal band; tarsi black. Wings with a faint
yellowish tinge, brightest at the wing-base; costal cell more saturated;
subcostal cell dark; stigma dark brownish black; a broad dark brownish
black seam along the cord; wing-apex blackish, this occupying the end
of cell Re, nearly the apical half of cells Rs; and Rs and part of M4.
Venation: Rs short; Re persistent; 2nd Anal vein running very close
to the anal margin so the 2nd Anal cell is very narrow. The wing is
strongly petiolate.

Abdomen with the basal tergites yellowish, especially laterally, the
fifth to ninth tergites black; sternites paler; abdominal segments mar-
gined laterally and ringed caudally with paler. Male hypopygium not
conspicuous; ninth tergite rather small, the caudal margin almost
straight across or but feebly notched. Outer pleural appendage rather
small, flattened, greenish white in color, broadest at the base, tapering
gradually to the apex. Inner pleural appendage pale greenish, termi-
nating in two subequal blackened spines that are slightly divergent,
claw-like; a broadly spatulate blade projects toward the ninth tergite.
Ninth sternite with a small but deep V-shaped median notch. Eighth
sternite projecting caudally, the margin unarmed.

Habitat: Japan.

Holotype, &, Meguro, Tokio, April 23, 1919 (R. Takahashi).

Paratopotype, o.

‘This exquisite species is dedicated to fine memory of my
mother, Mrs. Jane Parker Alexander. It shows numerous
points of resemblance to T. pedata Wied. and allied forms and
if the group 77pulodina Enderlein is to be recognized as a valid
genus this species will probably have to be placed therein,
although the hypopygium is strikingly different from that of
the genotype of 77ipulodina (pedata). For a discussion of the
status of the group 77pulodina, consult a paper by the writer,
Proceedings of the United States National Museum, vol. 49,
p. 184; 1915.

WING STRUCTURE OF LEPIDOPTERA AND THE PHY-
LOGENETIC AND TAXONOMIC VALUE OF CERTAIN
PERSISTENT TRICHOPTEROUS CHARACTERS.

By ANNETTE F. BRAuN, Cincinnati, O.

I. Introduction.

II. Wing Structure.
(a) Venation.
(b) Mode of insuring synchronous action of fore and hind wings.
(c) Distribution of fixed hairs.

III. Summary and Conclusions.
IV. References.

I. INTRODUCTION.

It is agreed that the insects constituting the two orders
Trichoptera and Lepidoptera respectively are closely related
genetically. All insects included in these orders have the same
general-and characteristic type of venation, which in the more
primitive forms closely approaches the hypothetical primitive
type as figured by Comstock. It is the problem of determining
where the line of division shall be drawn between Lepidoptera
and Trichoptera which presents difficulties. According to the
usually accepted classification, the Micropterygide are regarded
as the most primitive group of Lepidoptera, from which all the
other Lepidoptera have been derived. This group more closely
resembles in venation and mode of uniting the fore and hind
wing in flight the more generalized of the Trichoptera than it
does any other group of insects, evidently approaching the
stem form from which both groups have been derived. On
the basis of certain common and characteristic features in
venation, and the identical structure of the fibula in both,
Comstock (’18) removes the Micropterygide from the Lepi-
doptera, confirming his conclusion by the results of Dr. T. A.
Chapman’s studies of pupze, and makes them one suborder of
the Trichoptera, the Micropterygina, the members of Tri-
choptera as usually recognized constituting the other suborder,
the Phryganeina.

In addition to the Micropterygide, there is another group
of moths, the Nepticulide, the more generalized members
of which possess a fibula identical in structure with that of the

349

ae |
A

3
t

350 Annals Entomological Society of America [ Volk. Xahie

Micropterygide and the generalized Trichoptera,* and a pupa
resembling that of the Microptergyide. The venation of the
Nepticulide (Braun, ’17) shows highly specialized features. ©
It resembles most closely the venation of some of the Hydropti-
lide, but shows extreme anastomosis of main veins and absence
of cross-veins. It differs from that of Hydroptilide chiefly
in the few points in which the Nepticulide approach the usual
type of venation in the frenate Microlepidoptera with reduced
venation. The existence of this group, which shows undoubted
trichopterous affinities, further complicates the question of the
true relationship existing between the Trichoptera, the Microp-
terygide, the Nepticulide, the Hepialide, and the frenate
Lepidoptera.

The discovery of another family, the Prototheoride (Mey-
rick, Annals South African Museum, XVII, 17, 1917), with
characters intermediate between the Micropterygide and the
Hepialide, would seem to make it impossible to place the
Micropterygide and Hepialide in different orders.

In two related groups, any adaptation for a particular
function may have been handed down from the common ancestor
of the two, or it may be a similar modification of an homologous
structure arising independently in each group, because of the
same inherent tendencies in each, or the adaptations serving
the same purpose may be developments of entirely different
structures in the two groups. The two former conditions
indicate community of descent and closeness of relationship;
the third the phylogenetic divergence of the two groups. The
same principles may be applied in the analysis of the entire
structure of the groups whose true relationship is to be deter-
mined. If it can be shown, that the more primitive members of
one group having many characteristics in common with the
primitive members of another group and thus appearing more
closely related to them than to any other group, show the
beginning of divergence in structure from that other group, and
these modifications can be consistently traced in the same
direction through the more specialized members of the first
group, in my opinion the point where the two groups begin
to diverge, that is, the point where the distinctive characteristics
of a group originate, is the logical place to make a taxonomic

* The term Trichoptera is used in this paper in its commonly accepted sense,
not including the Micropterygidae.

f
t fi /

: i

1919] Braun: Wing Structure of Lepid a 351

division, even though an apparently wider gap may exist
somewhere between two divisions of a group, due to incomplete
preservation of connecting forms or insufficient knowledge on
our part.

In the present paper, the relative value of those factors
indicating community of descent and relationship of the insects
constituting the Trichoptera and Lepidoptera and those indi-
cating divergence is considered in its bearing on the question of
classification.

With regard to the taxonomic problem involved, the question
is essentially this: Are certain characteristics possessed by
the Micropterygide in common with the Trichoptera of such
taxonomic importance as to necessitate the conclusion that the
Micropterygide are trichopterous insects or are these char-
acteristics merely retained as a common inheritance, later to
undergo far-reaching modifications which can be traced back
to the Micropterygide.

Certain characters which are possessed in common by the
Micropterygide and Trichoptera and which seem sharply to
separate the Micropterygide from frenate Lepidoptera, will
be shown to undergo far-reaching modification and in this
modified form to persist in many of the comparatively primitive
groups of the frenate Lepidoptera. The fact that the course
of such modification can be traced makes it evident that the
gap between the Micropterygide and the remaining Lepidoptera
is not as wide as is sometimes supposed. This fact, and the
fact that certain acknowledged lepidopterous characters origi-
nate in the Micropterygide, necessitate the conclusion that the
Micropterygide are lepidopterous insects.

II. WING STRUCTURE.

The structure of the wings presents the most obvious and
available characters for the study of the phylogeny and taxo-
nomy of the Lepidoptera and Trichoptera. The following
discussion of the structure of wings of Lepidoptera deals with
the modification and disappearance of trichopterous characters
and with the origin of some distinctively lepidopterous char-
acters. The data are presented under (a) venation, (b) mode
of insuring synchronous action of fore and hind wings, (c)
distribution of fixed hairs.

352 Annals Entomological Society of America [Vol. XII,

(a) Venation.

Certain striking characteristics of venation are held in
common by the more generalized Trichoptera, such as Rhya-
cophila and by the more generalized of the Micropterygide.
These characteristics have been enumerated by Comstock
(The Wings of Insects, 1918, p. 317). In fact, the resemblance
is so close that the only essential difference to be noted is that
in Micropterygide media of both wings has been reduced to
a three-branched condition, apparently by the coalescence
of M; and M,, while in Rhyacophila M3; and ‘M, are separate in
the fore wing, although united in the hind wing. This identity
of characteristics is shared by no insect not belonging to
Micropterygide or Trichoptera. That it is possible to trace
the modification of these trichopterous characters and their
transformation into what are commonly recognized as the usual
lepidopterous characters, such as are found in the frenate
Lepidoptera, will be shown from evidence derived from the
Micropterygide themselves, the Nepticulide, the more primitive
frenate Lepidoptera, and the Hepialide.

The forms for study of venation in the more primitive
Lepidoptera have been chosen because they show, besides the
preservation of media, certain characters—such as the costal
spines on the hind wing, absence of frenulum in the female,
fixed hairs on the wings and folded maxillary palpi—all of
which indicate a generalized condition.

In the Micropterygide veins Cu and 1st A of both fore and
hind wing coalesce at the base of the wing; from their point
of separation Cu extends obliquely across the wing toward
media, then bends and extends longitudinally to the margin.
In the hind wing, the point of separation of Cu and Ist A is
nearer the base than in the fore wing. A short cross-vein, the
posterior arculus, connects media with the longitudinal part
of Cu, forming with the base of media, a serial vein in the
fore wing. The presence of this condition in Mnemonica
and in Rhyacophila is pointed out by Comstock in his book
‘““The Wings of Insects,’’ where wings of both are figured
together with figures of the tracheation in the base of the
wings of Mnemonica, which confirm the homologies of the veins
as determined from a study of adults. In so far as published
figures of other genera of the Micropterygide show the details

1919} Braun: Wing Structure of Lepidoptera 353

of venation at the base of the wings, they indicate that these
conditions are common to all members of the family, although
not without modification which may tend to obscure these
characters. That such modification does take place in the
Micropterygide, is shown by an examination of the wings of
Epimartyria (Fig. 1), which genus, since it precedes the appear-
ance of the tongue, must be regarded as a more primitive
genus than Mnemonica. In the fore wing, the configuration
of these veins is essentially that of AJnemonica. In the hind
wing, Cu extends straight to the margin from its point of
separation from the Ist A, that is, the ‘‘oblique”’ part of Cu
has become so extremely oblique as to be in a line with the
longitudinal part of Cu. As is to be shown presently, this
differs in no way from the course of Cu of the hind wing in
certain undisputed Lepidoptera. In Prodoxus, most of the
tracheae are preserved in the extreme base of the wings. Figure
2 shows the more general features of the venation, with wavy
lines representing tracheze; Figure 3 shows in detail the course
of the tracheze at the extreme base of the hind wing. In the
fore wing the coalesced bases of Cu and ist A contain a single
trachea, which soon divides, one branch following the /st A,
the other branch immediately dividing again, both branches
traversing Cu, but soon shriveling up. That portion of Cu
between its separation from ist A and the posterior arculus
(which is here transverse and contains no trachea) is evidently
homologous with the oblique free portion of Cu in Muicrop-
terygidze and Trichoptera. The tracheze in the base of the
hind wing (Fig. 3) show that the relations of Cu to the 1st A
are apparently the same as in the fore wing, but the separation
takes place extremely close to the base, and the course of the
veins is obscured by the tuberosities at the base of the wing.
It will be observed that the vein containing a branched trachea,
evidently the cubital trachea, is met very obliquely near the
base of the wing by a vestige of what appears to be the posterior
arculus. The evidence afforded by a study of pupal wings of
Prodoxus (Fig. 6) supports the conclusions derived from a
study of adult wings. The coalescence of Cu and Ist A, the
Z-shaped course of Cu, and the formation of the serial vein are
sometimes shown with greater distinctness in the fore wing of
Adela bella (Fig. 5), where these features of the venation are
almost exactly as in Mnemonica; a similar condition is present

354 Annals Entomological Society of America |Vol. XII,

in the hind wing. The course of these veins is also the same
in Cyanauges cyanella, as reference to Figure 4 will show; in
this case the separation of Cu and 1st A occurs almost at the
base of the wing.

The obvious conclusion to be drawn from this series of
examples is that in the course of evolution, the point of separa-
tion of Cu and 1st A moves toward the base of the wing, and
concomitant with this change, the oblique basal part of Cu
becomes more and more oblique until by the time the point of
separation has reached the base of the wing, the formerly
oblique free basal part of Cu is in a line with the longitudinal
part of Cu. There will thus be no evidence in the venation
of this part of the wing in the more specialized Lepidoptera to
indicate that it is derived from a venation like that of
Mnemonica. What appears to be the posterior arculus is
usually obsolescent (as indicated in the Figures by dotted
lines), but vestiges of it sometimes traversed by a persistent
trachea are present in many Lepidoptera’ belonging to widely
separated groups, e. g., in Tineide (e. g., Tzneola), Plutellidee
(e. g., Pliniaca), Eucleide (e. g., Sisyrosea textula), Tortricidae,
and possibly in other groups. The question whether this is
really the posterior arculus or the base of M, is not discussed
here. In the light of what has been shown to have taken place
in more primitive forms, the configuration of veins in the base
of the fore wing of the Noctuid, Renza flavipunctalis (Fig. 7),
suggests the same course of evolution. The coalescence of
the base of the first anal trachea with the cubital trachea,
shown in the pupe of some butterflies, and in some of the
specialized moths (e. g., Samia cecropia), I am inclined to regard
as a secondary connection, following the separation of Cu and
ist A at the base.

The venation of the fore wing of the more generalized
members of the Nepticulide is characterized by the basal
coalescence of media and cubitus, a character unique in Lep-
idoptera, but whose origin can be explained by assuming
that processes similar to what is known to occur in some
Trichoptera have taken place. The cubital and medial trachee
lie within the same vein cavity near the base (Fig. 8), the two
separating at the point where media bends forward to join
radius. This characteristic of the venation of the Nepticulide
can be derived from an ancestral form in which cubitus followed

1919] Braun: Wing Structure of Lepidoptera 355

the course common to Mnemonica and Rhyacophila, if we
assume an anastomosis of Mf and Cu, obliterating the posterior
arculus (such as Comstock shows has taken place in a species
of Rhyacophila) to have proceeded to the base of the wing.
This view is strengthened by the course -of the persistent
cubital trachea in the fore wing of a specimen of Nepticula
platanella (a comparatively generalized species of its genus)
which follows the first anal vein at the base, bending obliquely
toward media which it reaches at the point where media bends
upward toward radius, then extending longitudinally along the
usual course of cubitus. The venation of the hind wing shows
an advance over that of the fore wing, even in the most gen-
eralized Nepticulide, in that media and radius coalesce for
half their length; this condition, which is brought about by the
crossing over of media to radius at the base has been attained |
in part of the genus Nepticula in the fore wing.

In the Hepialide the course of cubitus of the fore wing is
the same as it is in the Micropterygide. In the hind wing Cu
is free from J/st A at its base; there is no evidence from pupal
wings or adult forms to indicate how this condition may have
been derived from one in which Cu follows the same course as
in the fore wing.

The coalescence of the tips of two branches of the third
anal vein with the tip of the second anal vein of the fore wing
is a character common to both the more generalized Trichoptera
and Micropterygide. The second branch of the third anal vein
tends to disappear very early in phylogeny, although the third
anal trachea is often forked in the pupa. Even in the most
primitive Micropterygide, as Sabatinca, this coalescence of
veins 1s shown in one species and not in the other. The second
branch of the third anal vein is but faintly shown in Ep7-
martyria (Fig. 1). In Scoliaula of the Nepticulide, there is a
faint indication of a second branch of the third anal vein,
represented merely by a broad slight thickening of the wing
membrane, but neither it nor the first branch continue far
enough to unite with the second anal vein. In Hepialide, the
second branch of the third anal vein shows no tendency to
unite with the first branch, but runs close to the margin of the
wing. Among the Frenate, the condition of the branches of
the third anal vein in Prionoxystus is similar to that in Hepialide.

356 Annals Entomological Society of America [Vol. XII,

In Prionoxystus robinie, the second branch of the third anal
closely parallels the axillary furrow; the third branch follows
the free margin of the posterior lobe. As shown later, the inner
margin of the fore wing in this species clasps the costa of the
hind wing. In other Frenate, I have found no indication of
the presence of the second branch of the third anal vein. It is-
suggested as a plausible explanation of the atrophy of this vein
that its disappearance is correlated with the loss of the holding
function by this part of the wing.

The cross-vein between the first and second anal veins of
the fore wing, which is present in Micropterygide and Tri-
choptera, is preserved in many of the primitive Frenatz as well
as in the Hepialide. It is shown distinctly in Adela bella
(Fig. 5), and in Prodoxus; in a pupal wing of Pronuba this vein
is as strong as the second anal. It is shown faintly in Cyanauges
cyanella.

In the hind wing, the course of the second anal vein, which
anastomoses with the first anal vein for a distance, is regarded
by Comstock as a distinctly ordinal character, common to
Micropterygidze and Trichoptera (cf. The Wings of Insects,
p. 310). The tracheation in the base of the hind wing of
Prodoxus (Fig. 3), shows the same course of the second anal.
The trachea of the second anal vein bends forward to the first
anal vein, but almost immedtately bends obliquely backward
and after meeting the cross-vein between the second and third
anal, extends in a longitudinal direction to the wing margin.
The cross-vein is a much more distinct and well developed
tubular vein than is the base of the second anal vein itself.
The pupal tracheation of Prodoxus (Fig. 6) shows quite clearly
the anastomosis of the first and second anal veins. In this
instance the longitudinal cross-vein between the second and
third anal veins is preceded by a trachea, which is apparently
the first branch of the third anal itself. A similar condition
‘1s indicated by the course of these veins in Cyanauges and in
Adela; in neither is verification possible through persistence of
trachee. In Adela the base of the 2nd A followed proximad,
becomes indistinct before reaching the Jst A; the cross-vein is
quite distinct.

In Hepialide neither venation nor tracheation in the pupal
wings (MacGillivray, ’12), where there are three free anal
trachez, indicates any such anastomosis.

1919] Braun: Wing Structure of Lepidoptera DOT

Adela bella (Fig. 5) shows a branching of the third anal vein
of the hind wing, similar to that in Micropterygide and
Trichoptera.

It has just been shown that certain essential features of
venation common to Muicropterygide and Trichoptera and
seemingly found in no other insects can be identified in more or
less modified form in most of the more primitive Lepidoptera.
In most cases the steps in this process of modification can be
traced. The evidence is perhaps least satisfactory in the
Hepialide, which would indicate their divergence from the other
lepidopterous groups, a view borne out by other points of
structure. It is apparent that these characteristic features
of venation which the Micropterygide hold in common with the
Trichoptera do not distinguish them from the rest of the
Lepidoptera as sharply as might be inferred by a comparison
of the Micropterygide with more specialized Lepidoptera only.
The manner of specialization in the Nepticulide, with respect
to the course of cubitus, is of especial value in determining
the true phylogenetic relationship of the Micropterygide to
‘the rest of the Lepidoptera. The course of modification in
the more generalized Nepticulide, paralleling that sometimes
occurring in Trichoptera, which is a divergence from the
usual lepidopterous type, together with the possession of the
fibula, undoubtedly indicates a common ancestry with Trichop-
tera, while certain other typically lepidopterous characters,
such as the single spined frenulum of the male, the short tongue
and six-jointed maxillary palpi characteristic of many primitive
Frenatz, gbsolescence of the first anal vein of the fore wing,
the reduction of radius of the hind wing in the same manner as
it has taken place in the Frenatz, place the Nepticulide with
certainty in the Lepidoptera. This peculiar combination of
characters in the Nepticulide, taken in connection with the
undoubted specialization in venation, places them as the end
group of a line of development divergent from that of the other
Lepidoptera. Since these two divergent lines of development
are both lepidopterous, the common ancestor must also be
lepidopterous. It has just been shown, that the course of
modification in venation in both lines of development can be
traced back to Micropterygide, which therefore, on the basis
of evidence derived from a study of venation, must be regarded
as the common ancestor and is hence lepidopterous. To

398 Annals Entomological Society of America |Vol. XII,

remove the Micropterygide from the Lepidoptera, would
necessitate the assumption that the Lepidoptera have originated
twice or have been derived from a hypothetical lepidopterous
ancestor so close to Micropterygide that it can not be separated
from Micropteryide.

(b) Mode of Insuring Synchronous Action of Fore
and Hind Wings.

The methods of holding the fore and hind wings together in
flight show characteristics which are of value in a study of the
phylogeny and relationships of the Trichoptera and Lepidoptera.

In the more generalized Trichoptera and in the most primi-
tive groups of Lepidoptera, the posterior lobe of the fore wing
has been modified to serve as an organ for holding the wings
together, termed fibula or jugum, depending on mode of
functioning. The fibula in the more generalized Trichoptera,
such as Rhyacophila, and in certain of the Micropterygide, as
Mnemonica in the subfamily Eriocraniinez, acts by pressing
downward over the base of the hind wing and clasping the
anterior tuberosity of the hind wing. Tillyard (18, 719) has
described for certain genera of the two remaining subfamilies
of the Micropterygide, the Mnesarchaeine and the Microptery-
gine, a different mode of functioning of the fibula. In these
genera it is described as being bent under the fore wing and
serving as a retinaculum for the series of costal spines of the
hind wing.* My observations on Epimartyria in the sub-
family Micropterygine tend to support this view.

The fibula in the female of the last specialized, genera of
Nepticulide (which find their nearest allies in characteristics of
venation among the Hydroptilide in the Trichoptera) is
identical in structure with the fibula in the more primitive
Trichoptera and in the Micropterygide. In the Trichoptera
the process of modification of the fibula has finally resulted in
such a reduction in size and change in shape that the posterior
lobe of the fore wing no longer bears any resemblance to a
fibula and can not function to aid in holding the fore and hind
wing together. In some forms, while not retaining the char-
acteristic shape shown in the Rhyacophilide, the longitudinal
free margin still shows the downward curve, thus indicating

* That these spines do not constitute a true frenulum and are not homologous
with it, is shown in the pages following.

1919] Braun: Wing Structure of Lepidoptera 359

that it can still act by clasping the anterior tuberosity of the
hind wing. The course of modification of shape in the posterior
lobe of the fore wing in Nepticulida from a fibula in females of
the more generalized genera can be traced through various
changes until 1t becomes merely a narrow lobe whose free margin
is continuous with the inner margin of the wing. Thus we find
that while in the females of earlier genera the fibula has pre-
served its original structure and is apparently functional,
in the males of these genera the posterior lobe is rather promi-
nent but lacks the characteristic shape of the fibula as found in
the Trichoptera and Micropterygide. In Nepticula the posterior
lobe has lost all resemblance to the fibula; it is extremely
narrowed and the axillary furrow is so indistinct as scarcely
to separate it from the rest of the wing. This process, whose
steps can be observed in this lepidopterous family, would seem
to indicate the possibility that a similar process might have
taken place in families of the Frenatz, or in other words, that
the Frenatz have been evolved from ancestral forms in which
a well developed fibula was present. There is some evidence
to support this view. In Prodoxus the posterior lobe of the fore
wing is more than usually prominent and is separated from the
rest of the wing by a very distinct axillary furrow; it shows a
strong tendency to fold under, thus seemingly retaining some
of the function of the fibula, although it has lost its characteristic
shape. In some Tineide, the posterior lobe of the fore wing
bears some resemblance to a fibula, but it is not of a structure
to be functional. In others of the Frenate where it can be
distinguished, it is merely a narrow lobe-whose free margin is
continuous with the inner margin of the wing.

In the Hepialide the jugum, though homologous with the
fibula, differs from it in shape and method of functioning.
The difficulty in the way of deriving the Hepialide from
ancestors with-a fibula disappears if a process similar to that
which is known to take place in the Nepticulide is postulated.
The jugum, in accordance with this view, would be considered
a development in another direction from the posterior lobe of
the fore wing in a more or less reduced condition.

The fact that a fibula of identical structure, though showing
some variation in function, is present in three different groups,
two of which, the Micropterygide and Rhyacophilide, are
primitive and approach one another closely in other char-

360 Annals Entomological Society of America |Vol. XII,

acteristics of the wings, indicates that it is a persistent primitive
character handed down from the common ancestor. The
Nepticulide show marked specialization and are the end of a line
of development, having given rise to no other group. Though
they are not apparently to be easily or directly derived from
any existing group, the possession of this primitive character
undoubtedly allies them more closely to the Micropterygide
than to any other Lepidoptera, and also indicates trichopterous
affinities. However, the possession by Muicropterygide and
Nepticulide of this character of primitive Trichoptera, while
it indicates relationship to Trichoptera, need not be taken as a
basis for regarding them as trichopterous insects, because as
shown for Nepticulide, it has disappeared as a functional
structure in males even of the less specialized genera, and its
function has been taken over by a single-spined frenulum of a
character typical of the males of the more specialized Lepidop-
tera. That is, the Nepticulide retain evidence of descent
in the form of a fibula, but they have progressed far enough in
the lepidopterous direction to have developed a distinctly
lepidopterous structure. The undoubted relationship between
the Nepticulide and the Micropterygide indicates that the
Micropterygide are without doubt also lepidopterous, but being
a more primitive group, have not traveled thus far toward the
usual lepidopterous type in respect to mode of uniting the
wings.

In Rhyacophilide and other groups of the Trichoptera, in
Micropterygide, Nepticulidee and many of the more primitive
Frenate (Figs. 1, 2, 5; 4a, 8), there is a series of slightly curved
stiff spines on the costa of the hind wing near the base, which lie
against one of the anal veins of the fore wing, or catch into a
similar series on the fore wing, or lie in the fold of the fibula, and
aid in holding the wings together. These spines lie beyond the
costal sclerite, not on it, as do the true frenulum spines. They
are proximal to the humeral vein in forms where this vein
is present. This series of spines is without doubt homologous
‘in the various groups in which it occurs; it may be present in
addition to other means of holding the wings together, or it
may be the only method of insuring united action of fore and
hind wings. It may be functional in females, while in the
males of the same species its function has been taken over by
some other structure. Thus in Nepticulide (Braun; ’17),

1919} Braun: Wing Structure of Lepidoptera 361

these costal spines are functional in females, but in the male,
where there is a single spined frenulum, they are rarely preserved
with the same structure and function. In Nepticulide when
functional, they are larger and stronger than in any other
Lepidoptera (except perhaps Opostega); they are decidedly
stronger than in the Micropterygide. The spines are present
in several of the more primitive frenate lepidopterous groups,
often in combination with other primitive characters, and some-
times persist in more specialized groups. In the females of
many groups of Lepidoptera this is the only method of insuring
the united action of the fore and hind wings. These spines
are present in females in Prodoxide, Adelide and Incurvariida,
together with a few weak and useless short spines in the position
of the frenulum. Homologous structures much modified and
without function of holding may be distinguished in the males
in these families, together with a well developed single-spined
frenulum. In many genera of Lyonetiide they persist par-
ticularly in the female where there is also a functional frenulum,
and they may also be distinguished as definite structures in the
male, different from the rest of the scale covering of the wing.
It is questionable whether they are functional except in rare
instances in this family.

The possession of the row of costal spines by many of the
Trichoptera and by more primitive Lepidoptera indicates that
it is a persistent primitive character, and as such indicates
common descent of the two groups. The preservation of these
spines in many frenate Lepidoptera is one of the few connecting
links between them and the Micropterygide.

The series of costal hooks — the hamuli — which have
developed in the more specialized families of Trichoptera as a
means of locking the fore and hind wing together, is a specializa-
tion not found in the Lepidoptera.

The true frenulum spines are situated on the costal sclerite
of the hind wing. Although structures homologous with a
frenulum, consisting of several stiff spines on the costal sclerite
of the hind wing are found in some of the more specialized
Trichoptera, the frenulum in its specialized form, culminating
in the single-spined frenulum with the well developed frenulum
hook of the male, is a distinctly lepidopterous development.
In most Trichoptera, in Micropterygide, in females of Nepti-
culide, in females of Prodoxus, Adela, etc. (Figs. 1, 2, 5, 4a, 8),

362 Annals Entomological Society of America [Vol. XII,

there are merely hairs or weak functionless setz on the frenulum-
bearing area; in these the function of holding the wings together
is performed by some other structure. In the males of some of
these, viz., Nepticulide, Prodoxus, Adela, etc., there is a strong
single-spined frenulum, formed by the fusion of several spines,
in some instances of as highly specialized a character as 1s
found in any Lepidoptera. These forms furnish no evidence
of the phylogenetic process of development of the frenulum or
frenulum hook, but such evidence is furnished by the condition
of the frenulum in some members of several groups which
conserve other primitive characters. In Cosside and Megalo-
pygide the frenulum is in a rudimentary condition, consisting
of a bunch of spines, and in the females of many moths it con-
sists of two or several spines. Of the process of transition
from more primitive modes of holding the wings together to a
functional frenulum, we have very little evidence. In this
connection, the condition found in Pronuba is significant. In
the female Pronuba yuccasella all except the two most proximal
of the row of costal spines have become flattened and scale-lke,
and these two remaining spines are larger than any of the
corresponding series found in Prodoxus, which is an allied,
though more generalized genus. In addition in the female
Pronuba the beginnings of a true frenulum are shown, con-
sisting of a tuft of short, weak spines at the distal end of the
costal sclerite in the same position as the frenulum of the male.
There is also an inwardly projecting row of scales from the
costa of the fore wing, apparently the beginning of a frenulum
hook, which is present in the male. The female frenulum is
not long enough to reach this row of scales, which is, however,
easily reached by the two strong costal spines, which may thus
function as a frenulum before the true frenulum has reached
a sufficient size to be functional. These two spines are not
present in the male Pronuba, which has a single-spined frenu-
lum. In the Nepticulide there is no stage in the development
of the frenulum intermediate between the minute functionless
spines of the female, and the strong single-spined frenulum of
the male. However, the fact that functional costal spines and
frenulum are occasionally present at the same time, indicates
that the costal spines retain their function up to the time that
it is taken over by the frenulum. In the Cossid, Prionoxystus
vobinie, where the frenulum is rudimentary in both sexes, the

1919] Braun: Wing Structure of Lepidoptera 363

underside of the inner margin of the fore wing hooks against
a strong ridge along the costa of the hind wing.

It is apparent then that a functional frenulum has originated
independently in the Lepidoptera, and that it is not a primitive
character derived from the ancestral lepidopterous stalk.

From the facts as above stated, it is equally apparent that
the transition from more primitive modes of holding the wings
together has taken place within the Lepidoptera, although in
most cases the intermediate stages have not been preserved.
That the frenulum originated independently several different
times at least in the Lepidoptera is shown, first, by the fact
that it appears in the Nepticulide, which is an end group not
derived from or related to the frenate Lepidoptera; second, by
the fact that it takes the place of costal spines in Pronuba;
third, by the specialized condition of the frenulum in the males
of certain groups, the females of which have a very rudimentary
frenulum; fourth, by its rudimentary condition in other com-
paratively primitive groups, such as the Cosside, where there
is no evidence of costal spines or similar holding structures,
and the transition stage has been bridged by a very different
means. It may have originated independently in other families,
but we are without direct evidence to this effect; similarly it
may have disappeared independently as is shown for example
by its presence in Euschemon only, in the Hesperide.

The significant phylogenetic feature in the development of
the frenulum is its appearance in the Nepticulide which is the
end of a line of development, and which is related to Microp-
terygide and must be regarded as derived from them, and its
independent appearance in the group usually known as the
Frenatz. No similar course of development is to be witnessed
in the Trichoptera. The evidence for phylogeny derived from
the frenulum in my view points unmistakably to the conclusion
that the branch of the ancestral stem which produced Microp-
terygide and Nepticulide must also have given rise to the
rest of the Lepidoptera and hence all should be included in one
order.

(c) Distribution of Fixed Hairs.

A character, which when present, may without doubt be
regarded as a persistent primitive character is the presence of the
fixed hairs upon the wing surface. These are characteristic

364 Annals Entomological Society of America |Vol. XII,

of the Trichoptera, the Micropterygide and certain other of
the more primitive families of the Lepidoptera, Nepticulide,
Hepialide, Incurvariide, etc., where they are distributed over
the entire wing surface, but even in these most plentiful on
the underside of the fore wing near the dorsal margin. In many
Lyonetiide, e. g., Hvzeroxestis, Oinophila, Coptodisca, besides
the fixed hairs on the underside near the inner margin, there
is a patch in the middle of the fore wing near the base; in
Tischeria and Opostega, there are additional scattered fixed hairs
on the wing surface. In other groups the fixed hairs are confined
to the under side of the fore wing near the base of the dorsal
margin, with the rare presence of such hairs on other parts of
the wing, e. g., in Tineola. The very fact that the fixed hairs
are most numerous in the most primitive groups, tending to
become scattered and later confined to definite areas of the
wing, and finally persisting only in a limited area on the
underside of the fore wing, where they may function to a
sight degree in holding the wings together in flight, shows
that while taken in connection with other characters, their
presence may indicate a comparatively primitive condition of
the forms possessing them, the character is not one upon which
a taxonomic division can be made.

III. SUMMARY AND CONCLUSIONS.

The discussion of wing structure in Lepidoptera has dealt
chiefly with those characters which have been handed down
from the common ancestor of both Lepidoptera and Trichoptera,
and which have been preserved without modification in the
most primitive Lepidoptera, but which have undergone more
or less far-reaching modification in all other groups of Lep-
idoptera. In many instances the steps in the process of modifi-
cation have been traced, and it has been possible to identify
these characters in their modified form in many of the more
primitive groups of frenate Lepidoptera. These changes in
structure have sometimes been correlated with changes in
function of certain parts of the wing or with the taking over of a
particular function by a different organ.

The conclusion reached from a study of certain features of
venation is that the Micropterygide are not as sharply separated
from the rest of the Lepidoptera as might be inferred from a

1919] Braun: Wing Structure of Lepidoptera 365

comparison made only with more specialized groups of Lepi-
doptera. The modifications which are shown in the more
primitive groups of frenate Lepidoptera have their beginnings
in the Micropterygide themselves. In addition we have
the evidence given by the Nepticulidze, which combine in one
group, certain characters found in the Frenatz, with characters
belonging to the Micropterygide and in the manner of
specialization of certain characters of venation diverge from
all other Lepidoptera, paralleling what occurs in some
Trichoptera. The existence of these divergent groups, the
Frenate and the Nepticulide, both of which are derived
from Micropterygide, is conclusive evidence of the lepidop-
terous character of their common ancestor, the Micropterygide,
even if we do not take into consideration such features
. of the Micropterygide as the character of the mouth-parts and
the scale covering of the wings, which unmistakably stamp
them as lepidopterous.

The conclusion drawn from a study of the various modes of
holding the wings together in flight in the Lepidoptera, is that
in the more primitive groups of Lepidoptera, including the
Micropterygide, certain trichopterous structures are retained
and are functional, but in higher groups are modified or dis-
appear and their function is taken over by other wing structures,
chief of which is the frenulum. The frenulum in its specialized
form is shown to have had its origin in the Lepidoptera and to
have developed independently in several widely separated
groups.

Certain general phylogenetic and taxonomic conclusions
follow from these studies. The Micropterygide are close to the
common ancestor of both Lepidoptera and Trichoptera, but
are true Lepidoptera. From them the remaining Lepidoptera
have been derived, not from a single line of descent, but from
several divergent lines, one of which is represented by the
Nepticulide alone; a second line by the Hepialide, with the
Prototheoride apparently forming a link between it and the
Micropterygide, and to which the Cossidze show some degree
of relationship; a third much branched line includes the frenate
Lepidoptera, of which some members such as the Prodoxide,
Incurvariide, etc., conserve some of the trichopterous char-
acters of their ancestry and must therefore be regarded as the
most primitive of the Frenate.

366 Annals Entomological Society of America [Vol. XII,

IV. REFERENCES.

Braun, Annette F. 1917. Nepticulidae of North America. Trans. Am. Ent. Soc.,
Vol. XLITI, pp. 155-209.

Busck, August. 1914. On the Classification of the Microlepidoptera. Proc. Ent.
Soc. Wash., Vol. XVI, pp. 46-54.

Chapman, T. A. 1896. Studies in Pupae. Trans. Ent. Soc. London.

Comstock, J. H. 1918. The Wings of Insects. The Comstock Publishing Co.,
Ithaca, pp. XVIII+430.

Kellogg, V. L. 1894. The Taxonomic Value of the Scales of Lepidoptera. Kansas
Univ. Quart., Vol. III, No. 1, pp. 45-89.

MacGillivray, A. D. 1912. The Pupal Wings of Hepialus Thule. Ann. Ent. Soc.
Am., Vol. V, pp. 239-245.

Meyrick, E. 1912. Lepidoptera Heterocera, Fam. Micropterygidae. Genera
Insectorum, 132 me Fascicule. ‘ z

Meyrick, E. 1917. Descriptions of South African Microlepidoptera, Annals South
African Museum, Vol. XVII, pp. 1-21.

Mosher, Edna. 1916. A Classification of the Lepidoptera based on Characters
of the Pupa. Bull. Ill. State Lab. Nat. Hist., Vol. XII, Art. 2.

Tillyard, R. J. 1918. The Micropterygidae not of the Jugate Type. Ent. News,
Vol. XXIX, p. 90.

Tillyard, R. J. 1919. A Further Note on the Wing-Coupling Apparatus in the
Family Micropterygidae. Ent. News, Vol. XXX, p. 168.

Ulmer, George. 1907. Trichoptera. Genera Insectorum, 60me Fascicule.

EXPLANATION OF PLATE XXIX.

Fig. 1. Wings of Epimartyria auricrinella.

Fig. 2. Wings of Prodoxus quinquepunctellus, female.

Fig. 38. Base of hind wing of Prodoxus quinquepunctellus, female. ~

Fig. 4. Wings of Cyanauges cyanella, male; 4a, base of costa of hind wing of female.
Fig. 5. Wings of Adela bella, female.

Fig. 6. Tracheation of pupal wings of Prodoxus quinquepunctellus.

Fig. 7. Base of fore wing of Renia flavipunctalis.

Fig. 8. Wings of Ectoedemia heinrichi, female.

ANNALS E.S. A. VoL, XII, PLATE XXIX

Raa, a a =e
SSS 3 a

Annetie F. Braun

7

.
‘

+, ; .
‘ as ot
7 + © Se.
oo “ee.
“> (sgh
tL
i pe :
*
. % . : 7 -
: “ve "
2
- com
}
- :
= : re 9
oie
; LJ
. 4 ,
7 i
o
rv :
: ie
¥ y =
2 -

x uo 4
wey | : ,
a : 2. be >?
: 7 ay, z
is i
= 4 ‘ i te,
a 7 = =r
i a =
- a “"
> ™ .— =
te , “
oe ‘ 7
os : :
w a
7 : Mt
: :
- .
we
ee
* A
: we,
a :
; a
: 7) Fr ;
= or ;
ia : 0) € ,
ab
:
: : =
7 * ¥
a a" a oe
P :
— : i ae ke fe

? . % 5
“4 : : dd eo eel voy ae ae a a er ee ee ee eee
a : = te ® : : a t _ 7 i
it ‘4 : >
. 1 a) ‘ Ta
a ’ *
B ws ars 4 r
: r rs : } —
; m x . : fi
‘al - i. iT 7
wt, :
E sah : 7
I : 1 a t 7
7 ‘
- ; 3
‘ - D
_ i 7 = '3os
‘ ae
‘ a+ : ‘ !
: » a8
a's , est 7
~< 7 j 7
x 1 t =
t : n ‘ Le 7 -
i : a
i .
* ow! = §
i a - 7
v = c y
be &.
i = '
- SJ i 7 -
A : -
i i A a ie « fi -
, - ° oa
A i. — . mf r _
‘ :

THE LIFE HISTORY AND EARLY STAGES OF PLATY-
METOPIUS HYALINUS OSB.,* A JAPANESE MAPLE
LEAF-HOPPER IN NEW JERSEY.

By EpGar L. DIcKERSON and Harry B. WEIss,
New Brunswick, N. J.

For the past several years the writers have noted this species .
as occurring more or less abundantly on Japanese maple (Acer
palmatum Thunb.) in various places in New Jersey, principally
South Orange, Plainfield, Springfield, Irvington and Rutherford.
During the summer of 1918 it was very abundant at Freehold,
N. J., on a form of Norway maple (Acer platanoides L. var.
globosum Nich.) and the following notes are the results of
observations made at that locality.

The species overwinters in the egg stage, each egg being
deposited singly just under the bark close to a bud on the
recently made growth of the twig. The egg is firmly embedded
in the tissue which becomes brownish and somewhat hard
adjoining it. The outer bark over the egg is raised and some-
what cracked and a crack is also present where the egg is
inserted. Sometimes the end of an egg protrudes slightly.
The egg is inserted so that its long axis is more or less at right
angles to the tissue and a favorite place for deposition is the
thickened part of the twig around the buds. Sometimes eggs
were found in groups of two to five each egg being separated from
the other, however, by considerable tissue.

At Freehold, N. J., hatching took place about the first
of June and continued for almost a month. As late as early
July it was possible to find first stage nymphs. From eggs
hatching the first few days in June, adults appeared the first
few days of July, showing that about one month is necessary
for the development of the five nymphal stages. This is
probably the maximum time and would be shortened by very
warm weather. On account of the uneven hatching of the eggs,
it is possible to find all nymphal stages and adults present
during the first part of July. By the end of July practically
all of the nymphs have matured and at this time and during
the first of August the adults are most plentiful. Over-

* Identified by E. P. Van Duzee.
369

370 / Annals Entomological Society of America [Vol. XII,

wintering eggs are then ‘deposited. The nymphs feed on the
lower leaf surfaces and seem to prefer the shaded portions of the
tree. The adults are very active, move readily when the
leaves are disturbed and when numerous their activity results
in sounds like rain falling on the leaves.

Ecc—Length 0.9 mm. Greatest width 0.18 mm. Translucent,
subelliptical, somewhat flattened and not quite cylindrical, tapering
slightly to both ends, one end rounded, the other end, at the surface of
the tissue, truncate.

First NympHart Stace—Length, including anal processes, 1.1 mm.
Width of head, including eyes, 0.29 mm. Elongate-elliptical; very
light yellowish white; beginning just behind the eyes there is a reddish-
brown, lateral margin varying in width and extending dorsally on
prothorax, posterior margin of thorax and anterior margin of abdomen
and on some of posterior abdominal segments (interrupted in some
specimens) ; antenne slightly longer than body; head extending forward,
triangular, obtusely angled in front, sides slightly rounded; eyes lateral,
prominent; insect widest across eyes; sides of body slightly rounded and
tapering to posterior end of abdomen; posterior end of abdomen divided
into two minute, spine-like processes; abdominal lateral row of hairs
similar to row on second stage; (dorsal hairs not apparent); rostrum
extending to beyond bases of second pair of legs; indications of spines
on tibia.

SeconpD NyMPHAL STaAGE—Length, including anal processes, 1.85
mm. Width of head, including eyes, 0.36 mm. Shape narrow, more
elongate than that of first stage. . Color varies from sordid white to
light yellow. Lateral reddish-brown margin somewhat similar to that
of first stage, but varying and somewhat broader. Antenne three-
fourths length of body. Head similar to that of first stage; sides of
thoracic segments rounded; metathorax slightly longer than lengths of
pro- and mesothorax combined; broadest across metathorax; abdomen
slightly narrow at base, widening to second abdominal segment and
gradually tapering to posterior part; spine-like processes one and one-
half times as long as last abdominal segment and bearing several hairs;
each abdominal segment bears two dorsal and two subdorsal and in
addition the last three segments two lateral hairs; rostrum extending
to between bases of second pair of legs; two rows of hair-like spines
on outer margins of hind tibia.

Turrp NyMPHAL STaGE—Length, including anal processes, 2.9 mm.
Width of head, including eyes, 0.44 mm. Somewhat similar to pre-
ceding stage; eyes more prominent; lateral reddish-brown markings
more extended, in some specimens much extended on dorsal surface
leaving a dorsal, median band of ground color of varying width (mark-
ings vary considerably in different individuals); anterior margin of head
slightly more angulate; posterior margin of prothorax arcuate, outer
posterior angles margined; mesothorax twice as long at sides as at middle;
metathorax twice as long as pro- and mesothorax combined, posterior

‘, -

1919] Dickerson-Weiss: Platymetopius Hyalinus 371

margin subtruncate, scarcely longer at sides than at middle; abdominal
hairs similar to those of fifth stage nymph; posterior abdominal spine-
like processes equal in length to the two posterior abdominal segments;
rostrum extending to between bases of second pair of legs.

FourtH NyMpHAL StaGe—Length, including anal processes, 4.1 mm.
Width of head, including eyes, 0.58 mm. Shape somewhat similar
to that of fifth stage; dorsal and lateral reddish-brown markings some-
what similar to those of preceding stage, but more diffused, in some
specimens extending further medially on dorsal surface of head and
some of body segments; antennz about one-half length of body; eyes
prominent, lateral, extending slightly posteriorly; anterior margin of
head subacute, sides slightly rounded; prothorax rounded in front to
conform to posterior margin of head, posterior margin slightly curved
and slightly wider than anterior margin; mesothorax with lateral
margins decidedly elongate covering two-thirds of metathoracic lateral
margins; metathorax with lateral margins somewhat elongate, covering
part of first abdominal segment; abdomen broadest across second
segment gradually tapering to end; length of posterior abdominal
spine-like processes more than combined lengths of last two abdominal
segments; spine-like processes bearing long hairs; abdominal hairs
similar to those of fifth stage nymph; rostrum extending almost to bases
of second pair legs; few, minute spines on legs; hind tibial spines similar
to those of fifth stage.

FirtH NyMpPHAL STAGE—Length, including anal processes, 5.5 mm.
Width of head, including eyes, 0.78 mm. Elongate-narrow; vertex
acutely pointed and abdomen gradually tapering from anterior portion
to apex; color sordid white with dorsal and lateral surfaces and wing-
pads mottled with a number of orange-yellow spots often margined with
interrupted reddish-brown lines, spots vary in size and some tend to
form one or two dorsal rows; antennze more than twice the length
of the head, situated on front just below eyes; head acutely pointed,
posterior margin arcuate, eyes prominent, extending backward, forming
outer posterior angles; head as long as width of broadest portion of head
across eyes; prothorax rounded in front to conform to the posterior
margin of head, posterior margin slightly curved, slightly wider than
widest portion of head across eyes; mesothorax about as wide as pro-
thorax; median portion of mesothorax about one-half as long as width,
sides somewhat shorter; wing-pads long, narrow and extending to
posterior margin of first abdominal segment. Metathorax about as
long as mesothorax; wing-pads extending slightly beyond those of
mesothorax; abdomen widest across anterior end and gradually tapering
to last segment which is terminated by two, long, spine-like processes,
which equal in length the combined lengths of the last three segments;
each abdominal segment bears near its posterior margin two dorsal
and two subdorsal and beginning with the third segment, two additional
lateral, fine, long hairs which are equidistant; posterior spine-like
processes also bear a number of fine, long, hairs; ventral surface and legs
sordid white; rostrum extending beyond bases of first pair of legs;
hind tibia bearing a row of hairs on inner margin and a double row of
spine-like hairs on outer margins; few minute spines on legs.

os

ae al Annals Entomological Society of America [Vol. XII,

AbuLTt—Platymetopius hyalinus Osborn. This was described
by Osborn in 1900 (Ent. News, vol. XI, p. 501) from five
females and four males collected in Washington, D. C., June,
1897, by Mr. J. S. Hine, who stated that they were very abun-
dant upon an introduced species of maple. As this description
is available to most entomologists, there 1s no need to repeat
it here. It might be stated, however, that the insect can be
recognized due to the fact that it is quite distinct from any
other species of the genus occurring in this country.: In
Osborn’s description it is characterized as follows: ‘‘Elytra
hyaline with dark points and fuscous bands arranged, one sub-
basal, one median and one subapical. Face bright, sulphur-
yellow, vertex, pronotum and scutellum yellow, with some
infuscations or greenish washes. Length, female, 5.5 mm.;
male,.4.25 mm.”

Van Duzee, in his “Revision of the American Species of
Platymetopius,’’ (Annals Ent. Soc. Amer., vol. 3, 1910, p. 218),
states that “this insect is quite aberrant in its genus by its
uniformly whitish hyaline elytra crossed by three narrow
fuscous bands. Prof. Osborn described it from a series taken
from an imported tree at Washington, D. C., and strongly
suspected that 1t might be an exotic form introduced from
material added to the Botanical Gardens there. In that case
it probably has become acclimated as Mr. C. W. Johnson has
sent me a specimen he took near Philadelphia.”’

While Pennsylvania and the District of Columbia are the
only localities listed in Van Duzee’s Catalogue, it probably
occurs in many other eastern places in view of the wide dis-
tribution of the Japanese maple as an ornamental tree. We
believe that Prof. Osborn’s surmise that this is an introduced
species is correct and that it has been brought into this country
on imported maples.. This is quite possible in view of the
fact that overwintering takes place in the egg stage in the
twigs.

THE PREVALENCE OF PHORMIA’ AZUREA FALLEN
(LARVA PARASITIC ON NESTLING BIRDS) IN THE
PUGET SOUND REGION AND DATA ON TWO

UNDESCRIBED FLIES OF SIMILAR HABIT.

By ©. E. PLatu.

(Together with Descriptions of the Two New Flies by
C. H. T. TownsEenp and J. M. ALprRIcH.)

In two recent papers! the writer pointed out that the larve of
Phormia? azurea Fallen habitually suck the blood of nestling birds,
sometimes with fatal results, and that the adult flies, contrary
to the opinion of most dipterists, are quite abundant, at least
in the San Francisco Bay Region. Last summer (1918), while
visiting Seattle, he had occasion to ascertain whether Phormia
azurea Fallen also occurs in the Puget Sound Region and found
that it is as frequent there as in the San Francisco Bay Region.
During his stay at Seattle (June 15th to August Ist), fifty-four
birds’ nests were examined containing nestlings belonging to ten
different species of birds. Of these fifty-four nests, thirty-three
were infested by the larve of Phormia azurea Fallen,* one
by those of a new species of Phormia, and six by those of a
new fly belonging to the genus Hylemyia of the family Antho-
myide. Drs. J. M.Aldrichand C. H. T. Townsend, of the United
States Bureau of Entomology, have been kind enough to
describe the two new flies, the descriptions being appended to
this paper, and the two flies will hereafter be referred to as

1 Plath, O. E. (a) Parasitism of Nestling Birds by Fly Larvae. The Condor,
Vol. XXI (1919), pp. 80-389. (b) A Muscid Larva of the San Francisco Bay Region
which Sucks the Blood of Nestling Birds. Univ. Calif. Publ. Zool., Vol. XIX

(1919), pp. 191-200.

2 A synonym of Protocalliphora which Dr. C. H. T. Townsend of the United
States Bureau of Entomology considers more appropriate.

3 A large number of specimens of this fly were sent to Dr. C. H. T. Townsend
of the United States Bureau of Entomology who has informed me that the fly was
correctly identified by Mr. C. W. Johnson (Cf. a. The Condor, Vol. XXI, p. 34.
b. Univ. Calif. Publ. Zool., Vol. XIX, p. 194. Dr. Townsend has also called my
attention to the fact that Coutant’s fly (Cf. Coutant, A. F., The Habits, Life
History, and Structure of a Blood-sucking Muscid Larva (Protocalliphora Azurea).
Jour. Parasit., Vol. I, pp. 135-150) is not Protocalliphora (Phormia) azurea Fallen,
as was claimed by Coutant, but Phormia chrysorrhea Meigen. This fact, Dr. Town-
send thinks, may account for the disparity between Coutant’s observations and
mine (Cf. The Condor, Vol. X XI, p. 37. b. Univ. Calif. Publ. Zool., Vol. XIX,
pp. 198-199).

373

%

ro
*

%

374 Annals Entomological Society of America [Vol. XII,

Phormia metallica Townsend! and Hylemyia nidicola Aldrich.
The frequency with which the larve of Phormia azurea Fallen
were encountered in birds’ nests, and the number of larve and

pupe per nest, is shown by the following tables:

TABLE I.
Species of Bird Nests | Infested | Uninfested
examined] nests nests
Nuttall Sparrow (Zonotrichia leuco-
phrys nuttalli Ridgway)...........- 3 3 0
Rusty Song Sparrow (Melospiza
melodia morphna Oberholser)....... 2 2 0
Willow Goldfinch (Astragalinus tris-
tis salicamans (Grinnell)............ 2 2, 0
Oregon Towhee (Pipilo maculatus
OKEZONUS Bell) acer eee eee ee: 1 1 0
Russet-backed Thrush (Hylocichla us-
tulatan (Natta) eee eee ce 5 4 1
Western Robin (Merula migratoria
propinqua (Ridgway).............. 25 18 7
Cedar Waxwing (Ampelis cedrorum
(Vieilot): eaten set gee ue nee ese 3 1 2
Yellow Warbler (Dendroica aestiva
(Gime lina) See oe etre 5 1 +
Cliff Swallow (Petrochelidon albi-
TONSDAY) 2 oes ane eee eee ws 1 6
English Sparrow (Passer domesticus
I Bitabate heyhey yieeney okey eens ees ais ee eae 1 0 1
if Boh orci) (sme pera sen Met ain opsitits o ieee 54 33 21

Infesta-
tion

100%

100%

100%

100%

80%

72%

33%

0%

61%

;Average

4 A species closely related to Phormia chrysorrhea Meigen, the latter, according
to Dr. Townsend, being synonymous with Phormia sordida Zetterstedt and Phormia

splendida Macquart.

1919] Plath: Larve of Phormia 375

TABLE II.
Species of Bird Larvae in Each Totals | Larvae
Nest per Nest
Nuttall Sparrow (Zonotrichia leuco-
phrys nuttalla Ridgway)............ BO MAO Os taney nes Ss: 166 55
Rusty Song Sparrow (Melospiza ie-
lodia morphna Oberholsder).........| 52, OO. ee cee cece eee e ee 107 ey 8 =
Cliff Swallow (Petrochelidon albifrons
SE nee teenie tne SAM ae es ps OTS oes 47 47
Oregon Towhee (Pipilo maculatus
REM ree god eos. See eee eet STR ess hele 45 45
Willow Goldfinch Ce ae tris-
tis salicamans (Grimnell). . oss (ator etree etareceicmn tite 87 43
Yellow Warbler (Dendroica aestiva
(Gnvelin) ages Oe I SONIA So rR eka 39 39
Western Robin (Merula propinqua
CReid camoyay rere. p SO Be Lh letata tte Uy lil, ey, IG is wikSy Pal
. 23, 26, 27, 27, 29, 37,
39, 40, 43, 72, 188...| 607 33
Russet-backed Thrush (Hylocichla us-
PRUETT EAN crs hs cle c ats, ore scce oye DP SOA ais} oy lee oom Be 108 27
Cedar Waxwing (Ampelis cedrorum
(Wino) ast seuiccaes aan Seeee henner Vestal ett "apt Mara so 2 2
Thovzuleeis oo Be ogee Bea Ee Attic oe 1208 36
Average

All of these nests, excepting eight, were located in shrubs
and trees three to twenty feet above ground, while the remaining
eight came from an altitude of about forty feet. Seven of the
latter (the Cliff Swallow nests) were taken from below the eaves
of a two story building on the campus of the University of
Washington, and the remaining one (that of the English
Sparrow) from a box near the top of a high telephone pole.
The comparatively small percentage of infestation of these
eight nests might of course be a coincident, but it is perhaps
more probable that it was due to the location and structure
of the nests.

The thirty-three nests infested by the larve of Phormia
azurea Fallen contained 111 nestlings. Although many of the
larve taken from these nests were gorged with blood, only two

376 Annals Entomological Society of America |Vol. XII,

of the 111 nestlings died. That there were no more fatalities
is perhaps to be explained by the fact that more than half of
the thirty-three infested nests were those of comparatively
large birds, the Western Robin and the Oregon Towhee.

The nest infested by the larve of Phormia metallica Town-
send was that of a Western Robin and contained three nestlings.
When the latter took wing, twenty-six larvae were taken from
the nest. The larve pupated in a few days and both larve
and pup looked so much like those of Phormia azurea Fallen
that the writer was unaware that he was dealing with a new
fly until the adults emerged. From the facts at hand it seems
probable that Phormia metallica Townsend, both as larva and
adult, is similar in habit to that of its near relative, Phormia
chrysorrhea Meigen.

The six nests infested by the larve of Hylemyia nidicola
Aldrich offered an entirely different aspect. When discovered
four of them contained the remains (only bones and feathers) of
nine, nearly full-fledged nestlings. From these four nests, 283
pupz were taken nearly all of which later hatched. The
remaining two nests infested by the larve of Hylemyia nidicola
Aldrich were those of a Cliff Swallow and a Yellow Warbler and
contained one and five dead nestlings respectively. In these
two nests the larve were still at their repast. Fifty-nine
larve were picked from the outer surface of the six dead birds,
though many more could undoubtedly have been obtained from
within the nestlings. None of the six nests infested by the
larve of Hylemyia nidicola Aldrich contained any living nest-
lings when discovered, but judging by the small number of
dead birds in most of the nests, it seems possible that some
of the nestlings may have survived. /

The larva of Hylemyia nidicola Aldrich is much smaller
than that of Phormia azurea Fallen and measures about 8 mm.
in length and 2 mm. in width when mature. It apparently
is not blood-sucking in habit like the larve of Phormia azurea
Fallen, Phormia chrysorrhea Meigen, and other Muscidae. The
pupa® measures about 5 mm. in length and 2.5 mm. in width
and is of a glossy, golden-brown color.

®>It is identical in size and appearance with the seventy-one secured near
hb caer California, the preceding summer (Cf. a. The Condor, Vol. XXI, pp.
36-37. b. Univ. Calif. Publ. Zool., Vol. XIX, pp. 196-197).

1919] j Plath: Larve of Phormia 377

There are only two possible explanations in regard to the
relation existing between the dead nestlings and the presence
of the larve of Hylemyia nidicola Aldrich. The first is that
the young birds died from some cause or other and that the
adult flies were then attracted by the odor of the decomposing
nestlings to deposit their eggs, or maggots, on them. The
second explanation would be that the eggs, or maggots, were
deposited on, or near, the young birds while the latter were still
alive, the larve subsequently causing the death of the nestlings
by penetrating into their body. It has been known for some
time that the larve of certain species of flies belonging to the
genera Hylemyia and Mydaea of the family Anthomyide
attack nestling birds in Central and South America,’ and
the second explanation is therefore not an improbable assump-
tion. Should further investigations prove that the larve of
Hylemyia nidicola Aldrich do attack living nestlings, as now
seems probable, then it will be important to determine the
range and bring this fly under control if we are not to suffer
considerable loss among our wild birds.

As regards subcutaneous parasitism of nestling birds by
fly larve, I should like to relate an interesting experience I had
at Seattle, last summer. During my investigations I came across
a nest containing six Willow Goldfinches which were nearly
old enough to leave the nest. One of these nestlings had a
patch of yellow on its head instead of being of the ordinary
color. I turned this bird over to a lady in Seattle to have it
reared with a brood of canaries. Two or three days later my
attention was called to the fact that some of the feathers on the
bird’s head were standing up and that the young bird was
constantly scratching its head. Upon examining the bird I
noticed three small holes, one on top and two at the side of its
head. Near the opening of each of these holes the anal end
of an active fly larva was visible. The larvze were extracted
with considerable difficulty by means of a pair of scissors and
forceps, and were about 5 mm. in length and 2 mm. in width.

6 Cf. (1) Townsend, C. H. T., (a) The Grass-quit Bot, an Anthomyiid Parasite
of Nestling Birds in Jamaica. Jour. Inst. Jamaica, Vol. II, pp. 173-174. (b) Mydaea
spermophilae n. sp. (Larva Parasitic on Nestling Birds). Trans. Amer. Ent. Soc.,
Vol. XXII, p. 79. (2) Nielsen, J. C., (a) Mydaea anomala Jaenn., a Parasite of
South-American Birds. Vidensk. Meddel. fra Dansk Naturhist. Foren. Bd. 63
(1912), pp. 195-208. (b) On Some South-American Species of the Genus Mydaea,
Parasitic on Birds, zbid. Bd. 65 (1913), pp. 251-256.

\

378 Annals Entomological Society of America [Vol. XII,

They were similar in appearance to the immature larve of
Phormia azgurea Fallen, but showed no blood in their intestines.
I tried to rear these three larve, but all three died before
pupating.. The young goldfinch completely recovered from
the rather severe operation and is still living (March, 1919).

The study of birds’ nests from the viewpoint of the entomolo-
gist and parasitologist seems thus far to have been rather
neglected. While examining Cliff Swallow nests below the
eaves of a two-story building on the campus of the University
of Washington, I found them so thickly infested by bed bugs’
that I preferred to leave the remaining nests alone. Another
blood-sucking insect that seems to use birds’ nests as a favorable
breeding place is the flea. I had encountered numerous flea
larvee in birds’ nests near Berkeley, California, the preceding
summer (1917), but when I saw the almost increditable multi-
tude of flea larve contained in a single sparrow nest which
was taken from a box near the top of a high telephone pole at
Seattle, I was amazed. In addition to the parasites which I
have spoken of as inhabiting birds’ nests, two or three others
were mentioned in one of my recent papers,® and this number
will undoubtedly be increased.. These facts make it evident
that birds’ nests offer a fruitful field of investigation for the
professional parasitologist.

7 Very probably Oecacus (Acanthia) hirundinis Jen., a species closely related to
the common bed bug, Cimex (Acanthia) lectularia Linnaeus, but more slender than
the latter.

8 Cf. The Condor, Vol. X XI, pp. 35-36.

1919} Plath: Larve of Phormia 379

DESCRIPTION OF THE NEw SPECIES OF Phormia.

By C. H. T. TOWNSEND,
United States Bureau of Entomology, Washington, D. C.

Phormia metallica n. sp.

Length, 7 to 8.5 mm. Twenty-eight specimens, all females; eight
pinned from New Hampshire, Virginia, Illinois, Wisconsin; and twenty,
of which ten are alcoholic, from Puget Sound, Washington State,
reared by Mr. O. E. Plath from maggots taken from nests containing
young birds.

Female vertex one-sixth to one-fourth head width. Head brownish,
thinly olive-grayish pollinose, with more or less of a brassy tinge.
Viewed obliquely from below the whole face appears blackish, with a
conspicuous golden spot at upper end of both parafacilia. Antennze
wholly blackish or brown, the third joint one and one-half to two times
as long as second. Palpi fulvous to rufous, the tips fuscous. Thorax
and scutellum dark metallic greenish-black to bright cupreous or brassy-
green, silvery to olive-gray pollinose, leaving three nearly equal and
rather heavy vitte of the ground color. There is a faint narrow dark
vitta barely visible in front close on each side of the median vitta in
some specimens. Abdomen metallic dark bluish-green to bright
cupreous or golden-green, showing thin silvery pollen coat in certain
lights, the anal segment always cupreous to golden-green. Legs
blackish. Wings clear. Tegule white to buff-yellow.

Holotype from Franconia, New Hampshire.

There is a well marked difference in the two forms here
described as one species. The typical form, represented from
all the above localities, is the darker one, with only the anal
segment golden-green, and the tegula white. The vertex is
about or nearly one-fourth head width, and the length is 7 to
8.5mm. Thoracic pollen silvery.

The other form, represented by three New Hampshire and
four Puget Sound specimens, has the abdomen wholly cupreous
to golden-green, the anal segment being concolorous; the thorax
and scutellum cupreous to brassy-green, almost the same shade
as abdomen; and the tegule deep buff-yellow. This form will
perhaps prove to bea good subspecies, or it may even be specifically
distinct. The front is narrower in this form, the vertex running
from one-sixth to little over one-fifth head width, and the
length is 7 to 7.75 mm. Thoracic pollen olive-gray.

The Puget Sound specimens, of both forms, have the third
antennal joint about twice as long as second, the second joint

380 Annals Entomological Society of America [Vol. XII,

being moderately short. The eastern specimens, of both forms,
have the second antennal joint longer, and the third joint only
about one and one-half times as long as second.

The typical form was identified by Coquillett as Calliphora
splendida Macquart, and specimens of the other form were
included in the series so labeled by him. Macquart’s specimen
was from Galveston, Texas, and the size was given as five lines
or 10 mm. The thorax and scutellum were stated to be black,
the abdomen greenish-blue with tip golden. The first two
antennal joints were said to be very short, and testaceous in
color. Also the wings were said to be fuscous on costal border.
On account of these discrepancies, I am unable to identify
the present forms as Macquart’s species, which appears cer-
tainly to be a synonym of chrysorrhea Meigen.

DESCRIPTION OF A NEW SPECIES OF Hylemyia.

By J. M. ALDRICH,
United States Bureau of Entomology, Washington, D. C.

Hylemyia nidicola n. sp.

Male—Deep black, with yellow halteres; calypters infuscated, almost
black, with the fringe dark below.

Front hardly wider than lower ocellus; six pairs of frontal bristles,
beginning at middle; frontal and facial orbits dark, not silvery, narrow;
antenne black, rather large, third joint reaching nearly to edge of mouth,
twice as long as second, arista bare; face flat, edge of mouth very little
protruding; bucca about one-eighth the eye-height, rather bristly;
palpi and proboscis black, ordinary; back of head flat, with coarse black
hair.

Thorax deep black, rather velvety above, with a trace of paler pollen
at inner edge of humerus each side; two to three pairs of large anterior
acrostichals, three posterior dorsocentrals, three sternopleurals; meta-
notum with gray pollen; pteropleura and hypopleura without hairs.

Abdomen black, with a faint, interrupted lighter pollinose cross-
band on each segment and a median dark stripe; fifth sternite with deep
excision and two long black lobes, which do not bear any striking
bristles.

Genitalia rather small; first genital segment shining black, with
about twenty small bristles; second segment subshining, with ordinary
hairs directed backward; inner forceps united, short, beadlike, with
two long hairs on each side near tip; outer forceps dark brown, shining,
twisted, not long, rounded at tip.

Legs wholly black, without striking characters, pulvilli a little
enlarged, brownish; front tibia with one or two bristles on outer hind

!
q

, fy
1919] Plath: Larve of Phormia 381

side; middle tibia with one on outer front, one on outer hind, and two
on inner hind side; hind tibia with two on outer front, six on outer hind,
three on inner hind, and a few cilia on basal half.

Wings uniformly blackish-brown; costal spine almost wanting; last
section of fourth vein one and a half times the preceding.

Female—Color rather uniform dark gray; front almost one-third the
head width; cruciate bristles present; abdomen uniform gray, sub-
shining; chaetotaxy as in male. Calypters yellow, with pale fringe;
wing not infuscated.

Length, 4 mm.

Locality, Seattle, Washington.

Type, male, allotype, female, and one paratype female,
deposited in United States National Museum (Type No. 22131).
Paratype male and female deposited in the California Academy
of Sciences.

The species shows the usual characters of Hylemyia—
sixth vein reaching wing-margin, scutellum with delicate hairs
below, hind calypter not elongated. The bare arista would
place it in Phorbia according to the prevailing system; but I
agree with Malloch (in Trans. Amer. Ent. Soc., xliv, 305, 1918)
that plumosity of the arista is not a generic character in this
case and Phorbia cannot be maintained.

NOTES ON THE BIOLOGY OF THE CARABID GENERA
BRACHYNUS, GALERITA AND CHLAENIUS.*

J. L. Kine, Harrisburg, Pa.

One of the most interesting and unexpected habits recorded
of the Carabide centers about the egg-laying of certain species
of the genus Chlaenius. In a family of beetles so decidedly
terrestrial in their habits we are indeed surprised to find that
certain members deposit their eggs in delicate mud or clay
cells high above the ground on the leaves of trees and shrubs.

Dr. C. V. Riley in the Proceedings of the Entomological
Society of Washington, I, 23, 1884, was the first to record this
interesting method of oviposition through his observations of the
egg cells of Chlaenitus impunctifrons Say. He also stated in
this note he had strong proof that Chlaenius aestivus Say,
Scarites subterraneus Fab. and the genera Dicaelus and Galerita
share with C. impunctifrons its singular mode of oviposition.
However, after the publication of this note Riley does not seem
to have given his proof that the beetles in question do deposit
their eggs in earthen cells as does C. impunctifrons nor, as far
as the writer is aware, do we have further mention of this
anomalous method of oviposition occurring in the Carabidz
until a recent paper by Claassen in 1919.f In this paper
we are given a description and figure of the mud cells of Chlaentus
impunctifrons, the same species reared by Riley in 1884.

The writer first observed these eggs during the summer of
1917, but was not successful in rearing the larvee which issued
beyond the third instar. As all attempts to learn the identity
of the cell builder proved fruitless the study was again taken
up the following year with such success that it was proven
Chlaenius aestivus Say was one of the cell builders, however, a
marked difference in the form and placement of the cells led
to the conclusion there were three species of cell builders
living in the same habitat. Through careful rearing from eggs
to adults this proved to be true as Galerita bicolor Drury and

* A paper on the life histories and biology of the beetles herein mentioned is
in preparation by the author.

7 Claassen, W. P., Life History and Biological Notes on Chlaenius impuncti-
frons Say. Annals Ent. Soc. Amer., XII, 95-99, Pl. VI. 1919.

382

1919] King: Biology of the Carabid 383

Chlaenius impunctifrons Say were reared from the egg cells in
question. After this other habitats were studied and by careful
rearing of all the eggs found and obtaining eggs from beetles
in captivity, it was also proven that Chlaenius cericeus Forst.,
Chlaenius tricolor Dej. and Brachynus cyanipennis Say. also
deposit their eggs in mud or clay cells constructed by the female
beetle.

The form, size and structure of the mud cells of these
beetles seems to be distinct in each species and this, together
with decided variation in the immediate location or placement
of the cells, renders specific identification possible for most of
the species studied with the exception of Brachynus.

THE Mup CELLS OF BRACHYNUS.

The mud cells of Brachynus cyanipennis Say (Plate XXX,
Figs. 1 and 2) are somewhat crescentic or triangular in outline,
rough in surface and measure about 2.25 mm. in length, that is,
the longest axis. The cells of these beetles are commonly
placed on the undersurface of loosely set stones, on dead twigs
and plant stems; they may be placed singly or in groups of
three to ten in number. In captivity the cells were sometimes
constructed massed one upon the other in clusters of eighteen
to twenty-five.

THE MuD CELLS OF GALERITA.

The mud cells of Galerita bicolor (Plate XXX, Fig. 3) are
roughly triangular in outline or purse shaped and finely granular
in surface. They are 5 mm. in length and 3.5 to 4 mm. in
width. The cells of this species are placed singly on the under
surface of smooth leaves, Benzoin and Impatiens being most
frequented by the beetles in the habitat studied.

THE Mupb CELLS OF CHLAENIUS.

With Chlaenius impunctifrons, the species studied by Riley
and Claassen, the mud cells (Plate XXX, Fig. 4) are almost
always smoothly convex and oblong in outline, measuring about
3.9 mm. in length and 2.28 mm. in width. In the habitats
studied it was noted that about ninety per cent of the cells of
this species occur on living plants, the under strrface of smooth
leaves being most often selected by the beetles. The distance

384 Annals Entomological Society of America [Vol. XII,

above the surface of the ground at which the cells are placed
varies from a few inches to seven feet or higher however, one
to two feet is nearer the average.

Unlike the preceding species, Chlaenius aestivus, while
often living in the same habitat as C. impunctifrons, rarely
ever constructs her mud or clay cells on leaves. In the habitats
studied, they were found almost exclusively on dead twigs,
plant stems and the bark of trees and shrubs, the dead thorns
of Gleditsia were much used in one habitat and in another
locality they were occasionally found on the strands of a barbed
wire fence. They are sometimes placed ten feet above ground.

The cells of C. aestivus (Plate XX XI, Figs. 1 and 2) are convex
and oval in form, coarsely granular or rough in surface; length
3.(5 to 4.5 mm., width 2:75 to- 3mm. “The exit of the larva
is effected by the breaking off of the lidlike flap, which covers
one side of the cell.

The earthen egg cells of Chlaenius cericeus (Plate XX XI, Fig 4)
are smooth in surface, 3.75 mm. in length, 2 mm. in width.
In general shape they are somewhat triangular or purse shaped.
The cells of this species have not been observed in nature,
but in captivity the beetles placed their cells on stones and the
lower parts of dead plant stems, all very near the surface of the
moist soil.

The cells of Chlaenius tricolor are small, measuring about
2 mm. in length and 1.25 mm. in width. They are strongly
rounded, oblong in outline, smooth in surface and along one
side there is a distinct fold or flap which marks the point of
closure. This species constructs her cells of fine clay-sand. The
stems of slender grasses and sedges seem to be most frequented
by the beetles. Cells were found in great abundance along
the shore of the Susquehanna River at Harrisburg, Pa., on the
common sedge Eleocharis tenuis.

CONSTRUCTION OF THE Mupb CELL.

As far as the writer is aware, no one has yet noted how these
cells are made. This has been observed by the author, but
will be only mentioned in this preliminary note, it sufficing to
state that the pellet of mud is collected by the beetle at and
around the tip of the abdomen in such a way that the cell
formed is in reality a mold of the caudal abdominal segments;

1919] King: Biology of the Carabid . | ‘88D

the lid of the cell being formed from a thin layer of mud which
covers the dorsal portion of these segments. After the deposi-
tion of the egg the tip of the abdomen is withdrawn and is again
used to bend down the dorsal flap or lid and close the cell, the
lid of which is self sealing, because of the soft texture of the mud
at the time of construction. Thus the mud cells of Chlaenius
may be compared to a pouched envelope in form and mechanism.
The folding flap and the sealing ot the edge when the soft mud
of the flap comes in contact with the opposite lip of the cell
is not different in principle from that of a common envelope.
The clay cells of Chlaentus tricolor (Plate XX XI, Fig. 3a) show this
structure quite distinctly.

As concerns the number of cells made by an individual
beetle, I have noted that C. impunctifrons may construct seven
to twenty cells during one night and eighty-two cells in a
season.

FEEDING HABITS OF THE LARV.

The cell building habit of these Carabids presents several
interesting biological questions. Among these is one relative
to the feeding habits of the issuing larve. As concerns this,
there is but a single note by, Claassen, Annals of the Ent. Soc.
Amer., XII, 96, 1919. I quote the following:

“Various kinds of food were offered to the young larva
(of Chlaenius impunctifrons); larvee and pupe of Lymnaecia
phragmitella Stainton, larve of Nonagria oblonga Grote, Arsi-
lonche albovenosa Goeze, and Arzama obliqua Walk. Flies and
fresh meat were also offered to the larva. At first the young
larva fed somewhat on flies and on larve and pupe of L.
phragmitella, but it showed a decided preference for the little
larvee of A. obliqua.

During the second and third instar all other food was refused
except larvee of A. obliqua.

All the above mentioned Lepidoptera are inhabitants of
Typha. A. albovenosa_is an incidental surface feeder oa the
leaves; L. phragmitella lives in the heads of Typha, while A.
obliqua and N. oblonga both begin their larval activities as leat
miners, later becoming solitary stem borers.

It is possible that the larve of Chlaenius impunctifrons
feed altogether on the larve of A. obliqua. The former hatch

386 Annals Entomological Society of America [Vol. XII,

about the same time that the larve of the latter species leave
the mines of the leaves of Typha to become stem borers. Thus
while they are exposed they might easily fall prey to the Carabid
larvee.”’

From the above quotation it would seem that there is‘a
possible relationship between the occurrence of the mud cells
of C. impunctifrons on Typha and the food of the issuing larvee
as being the Lepidopterous inhabitants of Typha.

The writer believes that so far with all the known species of
Carabide which deposit their eggs in earthen cells on plants,
there is no correlation between the immediate location of the
egg cells and the feeding habits of the issuing larve. He
believes, therefore, that Claassen’s statements are misleading
and for Chlaenius impunctifrons points out the following
reasons:

First, because Chlaenius impunctifrons is often found in
abundance in habitats where Typha and its inhabitants are
entirely wanting. Second, the larve of this species are not
chmbers, it being doubtful if they could climb the smooth leaves
of Typha. As observed by the writer the larve either in
captivity or in nature were never seen to climb and in their
natural state have been observed at night running about
among stones and debris on the muddy shores of the Susque-
hanna. In nature I have never observed the larve feeding,
however, 1n captivity they were general feeders preying upon
all soft bodied insects which were not too active to escape them.
They fed greedily on small Lepidopterous larve (Gelechids and
Geometrids) Collembola, termites, crippled flies and the larve
and pupez of several species of ants. The ant larve and pupez
were easily collected and proved an ideal food for the Carabid
larvee, consequently they were used as the chief food in rearing
all the species mentioned in this paper, with the exception of
Brachynus.

A further lack of such correlation between the place of egg
laying and the feeding habits of the larve is shown in Chlaenius
aestivus which deposits her eggs in cells on dead twigs, tree
trunks and many other objects not supporting any forms of
life at all. Lastly, in Brachynus, which genus also shares this
cell building habit, we have parasitic larve, the hosts of which
are often quite removed from the twigs, stems and stones on

1919] King: Biology of the Carabid 387

which the egg cells are found. The larve of Brachynus cyani-
pennis mentioned in this paper, are parasitic on the pupe of
Dineutes discolor.

PARASITES.

A number of minute Hymenopterous parasites have been
reared from the egg cells of these Carabids. With Brachynus
as high as fifty per cent of the eggs found in nature were
parasitized by tiny winged and wingless Hymenoptera of the
genus Hoplogryon. All parasites of C. impunctifrons, C.
aestivus, and C. tricolor were winged. In one habitat studied
the eggs of C. tricolor were parasitized to the extent of twenty-
five per cent, by a minute Hymenopteron, Prosacantha caraborum
Riley, which is a common parasite of several species of Chlaentus.

ACKNOWLEDGMENTS.

The author is indebted to Mr. H. B. Kirk, entomologist,
Pennsylvania Bureau of Plant Industry, who so kindly con-
tributed his time and patience in taking the photographs herein
used. He also acknowledges the kindness of Mr. A. B. Cham-
plain, Entomologist, Pennsylvania Bureau of Plant Industry, .
for the verification of the species mentioned in this paper, with
the exception of Brachynus cyanipennis, which was determined
by Mr. H. C. Fall. .Thanks are also due to Mr. A. B. Gahan
of Washington, D. C., for the determination of the Hymenop-
terous egg parasites.

388 Annals Entomological Society of America [Vol. XII,

EXPLANATION OF PLATES.

PLATE XXX.
’ (Figures seven diametérs. )

Fig. 1. Masses of mud cells of Brachynus cyanipennis. Cells made by beetles in
~ captivity.
Fig. 2. Mud cells of Brachynus cyanipennis as found in nature on the under surface
of a stone.
Fig. 3. Mud cell of Galerita bicolor on the under surface of Impatiens leaf.
Fig. 4. Mud cell of Chlaenius impunctifrons on under surface of Benzoin leaf.

PLATE XXXI.
(Figures seven diameters. )

Fig. 1. Mud cells of Chlaenius aestivus on dead twig.

Fig. 2. Mud cells of Chlaenius aestivus showing lidlike opening for escape of the
larva.

Fig. 8. Mud cells of Chlaenius tricoler on sedge, Eleocharis tenuis.
a. Lateral aspect showing fold.
b. Lateral aspect showing side opposite fold.
c. Lateral aspect, fold removed showing egg.

Fig. 4. Mud cells of Chlaenius cericeus.
a. Cell which was removed from surface of a stone.
b. Cell on sedge.

ANNALS E. S.A. VoL. XII, Plate XXX.

J. L. King

ANNALS Dp Shee VOL. XII, PLATE XXXI.

INDEX OF VOLUME XII.

Abdominal Structures of Orthopteroid
Insects, 267.
Acridide, 282.
Acridoidea, 282.
Acrydiide, 282.
Achroia grisella, 76.
Acronycta americana, 76.
Adela bella, 353.
Adoretus tenuimaculatus, 171.
Aeromyrma, 175.
Agalena mimoides, 255.
Agrilus walsinghami, 10.
Aldrich, J. M., description by, 380.
Report of Sec., 59. -
Alexander, C. P., articles by, 25, 327.
Allodorus major, 20.
Alsophila pometaria, 76.
Amara erratica, 8.
Amaurobius bennetti, 240.
nevadensis, 239.
nomeus, 240.
pictus, 240.
utahensis, 239.
Anasa tristis, 193.
Anisodactylus, 11.
dilatatus, 11.
Anisolabis maritima, 303.
Anomala, 12.
orientalis, 171.
Anosia plexippus, 193.
Ants of the Genus Metapone, 173.
Antocha, 330.
satsuma, 332.
spinifer, 331.
Apatela americana, 76.
Aquatic Adaptations of Pyrausta
penitalis, 213.
Aranea tusigia, 254.
utahana, 254.
Asida, 9.
Asphondylia websteri, 159, 160, 161,
162, 163, 165, 167, 170.
Athous ferruginosus, 5.
Atteva aurea, 76.
Automeris io, 76.

Basilarchia archippus, 76.

Bathyphantes fillmoranus, 249.
fratrella, 250.

Bellura melanopga, 216.

Bembidium incertum, 8.

Blattide. 296.

Blattoidea, 296, 311.

Bombyx mori, 76, 193, 200.

391

Botis nelumbialis, 214.

Brachynus cyanipennis, 383.

Braun, Annette F., article by, 349. -
Brennus, 2.

Bruchophagus funebris, 161.

Brues, C. T., article by, 13.

Californian species of Malthodes, 31.
Callimone asphondyli, 161.
Callosamia promethia, 193.
Carabid Genera. Brachynus Galerita
and Chlaenius, Biology of, 382.
Carabus taedatus, 4.
Calotermes, 175.
Carabara, 175.
Cecidomyid Galls, Chalcid wasps
from, 159.
Ceratomiz catalpz, 76.
Cerura, 44.
vinula, 47.
Ceuthophilus, 274, 278, 279.
lapidicola, 278.
Chalcid fly parasitic on Australian bull-
dog ant, 13.
Chalcid wasps from Cecidomyid
galls, 159.
Chamberlin, R. V., article by, 239.
Chariesterus and its Neotropical
Relatives, 227.
Chriesterus albiventris, 228.
antennator, 227.
armatus, 228.
balli, 228, 229.
cuspidatus, 228.
gracilicornis, 228.
moestus, 228.
Chlaenius impunctifrons, 95.
Biological Notes on, 95.
Chlaenius aestivus, 95, 382, 384.
cericeus, 383, 384.
impunctifrons, 95, 382, 383, 385.
tricolor, 383, 384.
Chlorotettix fumidus, 237.
productus, 236.
Cibdelis, 11.
Cicadellide, 231.
Cicurina garrina, 255.
idahoana, 258.
utahana, 257.
utahana anderis, 257.
Cirphis unipuncta, 76.
Claasen, P. W., article by, 95.
Clubiona orinoma, 255.
Coelus, 11.

392 Index to Volume XII

Coniotus, 11.
Conocephalus, 280.

fasciatus, 279.
Copris, 12.
Cornicularia monticolens, 251.
Crane Flies of the Hawaiian Islands

(Tipulidae Dipt.), 25.

Craneflies, Japanese, 327.
Cryptocercus punctulatus, 301.
Cryptohypnus bicolor, 8.
Cyanauges cyanella, 354.
Cychrus tuberculatus, 4.
Cylindromyrmex, 173, 175, 176.

meinerti, 175.

striatus, 176.
Cylindrotoma, 344.

japonica, 344.

Datana integerrima, 76.
ministra, 76.
Deltocephalus delector, 233.
latidens, 234.
Deltometopus rufipes, 49.
Immature stages of, 49.
DeLeng, D. M., and Sanders, J. G.
article by, 231.
Dermaptera, 303, 312.

Detweiler, J. D., and Herrick, G. W.,

article by, 44.
Diabrotica soror, 11.
Diapheromera femorata, 293.
Diatraea saccharalis, 76.
Dicaelus, 12, 95.

Dickerson, E. L., article, 369.
Dicranomyia, 25.
foliocuniculator, 28.
grimshawi, 27.
hawaiiensis, 28.
jacobus, 28.
stygipennis, 27.
swezeyi, 29.
Dicranomyia, 327.
atripleura, 328.
basifusca, 328.
immodestoides, 327.
mesosternata, 329.
Dicranoptycha, 332.
yamata, 332.
Dicranota, 344.
nipponica, 344.
Dictyna arundinaceoides, 241.
bellans, 242.
coloradensis, 241.
insolens, 242.
olympiana, 243.
sociella, 242.
sublata, 241.
unitana, 240.
vincens, 243.
Dietz, W. G., article by, 85.
Dioryctria abeitella, 76.

Diplocephalus wamotsus, 251.
Dipterous Genus Tipula, Streptocera
Group, 85.
Dissosteira carolina, 193.
Distribution of Insects in Western
INevAmas, 16
Drassodes gosiutus, 245.
melius, 246. 4
robinsoni, 245.
Dyctinoides, 243.
arizonensis, 244.
Dystaxia, 11.

Edrotes, 9.

Eleodes, 10.

Embide, 305.

Embidiina, 305, 313.

Ephebomyrmex, 175.

Ephelia subaprilina, 340.

Ephemerida, 308.

Epimartyria, 353.

Erax, Robberflies of the Genus, 103.

Erax estuans, 105, 120, 124.
affinis, 105, 121.
anomalus, 105, 119.
apicalis, 105, 125.
aridus, 105, 117.
armatus, 105, 112.
aurimystaceus, 105, 122.
argentifrons, 106, 148.
argyrogaster, 106, 152.
argyrosoma, 106, 144.
auripilus, 105, 145.
aurivestitus, 105, 142.
barbatus, 105, 108, 109.
belfragei, 105, 121.
bicolor, 105, 110.
bimaculatus, 105, 115.
bicaudatus, 105, 138.
californicus, 106, 150.
candidus, 105, 119.
canus, 106, 145.
carinatus, 106, 131.
cingulatus, 105, 111.
comatus, 106.
concinnatus, 106, 131.
coquilletti, 106, 149.
costalis, 106, 134.
cressoni, 106, 134.
dubius, 106, 141.
eximius, 106, 146.
femoratus, 106, 123.
fulvibarbis, 105, 129.
grandis, 105, 111.
harveyi, 105, 115.
haloesus, 105, 130.
inflatus, 106, 146.
interruptus, 107, 153.
jubatus, 106, 138.
kansensis, 105, 122.
latrunculus, 106, 132.

Index of Volume XII 393

Erax (continued)
leucocomus, 105, 114.
loewii, 105, 125.
mexicanus, 105, 123.
nemoralis, 107, 147.
nigrimystaceus, 106, 130.
nigritarsis, 106, 129.
pallidulus, 107, 144.
parvulus, 106, 132.
pernicis, 105, 120.
pilosus, 107, 150.
portoricensis, 106, 128.
plenus, 107, 148.
prattii, 105, 117.
productus, 106, 136.
quadrimaculatus, 105, 114.
tapax, 107, 142.
rufibarbis, 106, 137.
tufitibia, 106, 128.
sagax, 105, 113.
slossone, 106, 121.
snowi, 105, 116.
spiniventris, 106, 135.
splendens, 107, 148.
stramineus, 107, 140.
stylatus, 106, 127.
subcupreus, 106, 133.
subpilosus, 105, 118.
tagax, 105, 113.
tabescens, 106, 126.
texanus, 107, 151.
triton, 107, 151.
truncatus, 107, 148.
tuberculatus, 106, 135.
unicolor, 106, 132.
varipes, 107, 152.
willistoni, 105, 110.
zonalis, 105.
zonatus, 112.

Erioptera, 338.
bifurcata, 338.

Eucharis myrmicie, 13.

Eulabis, 11.

Eupagoderus, 9.

Eurytoma medicaginis, 159.

Euschistus fissilis, 193.
variolarius, 193.

Evarthrus, 12.

Fall, H. C., article by, 31.
Feltia sp., 76.
Forficula auricularia, 303.

Fracker, S. B., article by, 227.

Gahan, A. B., article by, 159.

Galeopsomopsis multisulcatus, 168.

_ transcarinatus, 168.
Galerita, 95.

bicolor, 382, 383.
Glyptoscelimorpha, 11].

Gonomyia, 25, 30.
gracilis, 30.
hawaiiensis, 30.
skusei, 30.

Grammonota orites, 249.
obesior, 250.
simplex, 250.

Gryllide, 280.

Grylloblatta, 289.

Grylloblatta, 270, 271.
campodeiformis, 270.

Grylloblattida, 289, 310.

Grylloblattoidea, 289.

Gryllus assimilis, 281, 282.

Grylloidea, 280.

Harrisina americana, 76.

Hemerocampa leucostigma, 76.

Hepialide, 356.

Herrick, G. W., and Detweiler, J. D.,
article by, 44.

Herpyllus oabus, 246.

Hilaira uta 253. :

Hine, J. S., article by, 103.

Hoplogryon, 387.

Hylemyia nidicola, 374, 380.

Hyphantria cunea, 76.

Immature Stages of Deltometopus
tufipes, 49.
Ipochus, 11.
fasciatus, 11.
Ischnoptera pensylvanica, 297.
Isoptera, 302, 312.

Japanese Craneflies, 327.
Jassids from the Eastern United
States, 231.

King, J. L., article by, 382.

Lachnosterna, 12.
Lagoa crispata, 76.
Lathys trivittata, 244.
Lasiomastix pilifer, 339.
Leiponeura, 30.
Lepidoptera, Male Genital System, 192.
Wing Structure of, 349.
Lepidopterous larve, Olfactory sense
of, 65.
Lepisma, 309.
Leptogenys mucronata, 175.
Lespeyresia molesta, 76.
pomonella, 76.
Leucotermes flavipes, 302.
Libnotes, 25.
perkinsi, 26.
Life History and Biological Notes on
Chlaenius Impunctifrons Say (Cara-
bidze Coleop), 95.

394 Index of Volume XII

Limnophila, 339.
pilifer, 339.
rufipennis, 341.
subaprilina, 340.

Linyphia latescens, 248.
orinoma, 248.
insignis, 248.

Liogma, 345.
serraticornis, 345.

Lucanus, 12.

Machilis, 309.

Macrosiagon ferruginea, 20.
Macrurocampa, 44.
Malacosoma americana, 76.

Malthodes, Californian species, 31.

Malthodes—
appendiculatus, 33, 38.
basalis, 33, 36.
bicurvatus, 33, 39.
complicatus, 33, 40.
fusculus, 33, 41.
laticollis, 33, 34.
mitificus, 33, 37.
magister, 33, 34.
mollis, 33, 37.
var longipennis, 38.
obdustus, 33, 39.
piceolus, 33, 37.
reflexus, 33, 34.
sericeiventris, 33, 40.
tularensis, 33, 41.
vapidus, 33, 41.
vigilans, 33, 36.
visceratus, 33, 42.

Male Genital System of certain

Lepidoptera, 192.

Mantoidea, 294, 311.

MclIndoo, N. E., article by, 65.

Megalopyge opercularis, 76.

Melanoplus bivittatus, 283.

Melanotus, 12.

Metapone, 173.

bakeri, 186.

greeni, 179.

hewitti, 189.

leae, 183.

mjobergi, 181.

sauteri, 182.

tillyardi, 187.
Metrius, 3.
Micropterygide, 349.
Miscodera insignis, 4.
Mnemonica, 352.
Monilema, 9.
Mosher, Edna, article by, 49.
Muir article by, 171,
Myrmecia forficata, 13.
gulosa, 13, 15.

Metapone, Ants of the Genus, 173.

Nebria columbiana, 8.
diversa, 9.
ingens, 9.
kincaidi, 8.
mannerheimi, 4.
Ovipennis, 8.
sahlbergi, 8.
trifaria, 8.
Neoscona utahana, 254.
Nepticula planatella, 354.
Nepticulide, 349.
Nesospeza, 347.
tarsalis, 347.
New Western Spiders, 239.
Nezara viridula, 193.

Notodontid caterpillars, Repugnatorial

glands of, 44.
Nyctoporis, 11.

Odonata, 309.
Oecanthus, 282.
quadripunctatus, 282.
Olfactory sense of Lepidopterous
Larve, 65.
Omus, 2.
Omus dejeani, 4.
Onthophagus, 12.
Ophrastes, 9.
Ormosia, 334.
atripes, 335.
cinctifer, 336.
diversipes, 334.
takahashii, 337.
tokionis, 336.
Orthoptera, 278, 310.
Orthopteroid Insects, Abdominal
Structures of, 267.
Osborn, Herbert, Report of Man-
aging Editor, 61.

Pachysima ethiops, 174.
latifrons, 174.
Paedalgus, 175.
Papilio polyxenes, 76.
Paragaleospomyia gallicola, 167.
Parapsilogaster, 13.
Paratropeza, 333.
flavitibia, 333.
Parcoblatta pennsylvanica, 297.
Pardosa utahensis, 258.
Pasimachus, 12.
Pectinophora gossypiella, 76.
Perilampus, 15.
Phalacrocera, 346.
mikado, 346.
Phasmoidea, 292, 310.
Pheidole fraudulentus, 13,
megacephala, 13.
Philosamia cynthia, 193,
Phlegethontius sexta, 76.

Index of Volume XII

Phlepsius tenuifrons, 235,
tinctorius, 235.

Phloeodes, 11.

Phormia azurea, 373.
chrysorrhea, 376.
metallica, 374, 379.

Phyllophaga, 12.

Pieris brassice, 193.
rape, 193.

Palpigus, 227.

Plath, O. E., article by, 373.

Platymetopius collaris, 232.
rubellus, 231.

Platymetopius hyalinus,

of, 369.

Platynus, 10.
bogemanni, 7.
jejunas, 10.

Plecoptera, 307, 313.

Pliniaca, 354.

Polyangaeus, 342,
japonicus, 342.

Pontia rape, 76.

Prevalence of Phormia azurea, 373.

Prionolabis rufipennis, 341.

Prionoxystus, 355.

Pristoceuthophilus, 279.
cercalis, 279.

Proantocha, 330.
spinifer, 331.

Proceedings Thirteenth Annual

Meeting, 57.

Prodenia ornithogalli, 76.

Prodoxus, 353.

Promecognathus, 3.

Pronuba, 356.
yuccasella 362.

Prosocantha caraborum, 387.

Pseudocatholaccus americanus, 164,
asphondyliz, 164.

Pseudomyrma kunckeli, 175,

Psilochorus utahensis, 247.

Psilogaster fasciiventris, 14.
pallipes, 13.

Pteromalus polyphagus, 164.

Pteronarcys, 307.

Pterostichus, 3, 6.
amethystinus, 5.
brunneus, 9.
caligans, 6.
grandiceps, 6.
herculaneus, 4.
ovicollis, 6.
protractus, 7.
rostratus, 6.
scutellaris, 5.
sphodrinus, 6.
validus, 4.

Pyrausta penitalis,

tions of, 213.

early stages

Aquatie Adapta-

395

Renia flavipunctalis, 354.
Report, Committee on Resolutions, 62.
of Auditing Committee, 62.
of Nomination Committee, 62.
Report of Managing Editor, 61.

Report of Secretary, 58.
Report of Treasurer,
Foundation, 61.

Repugnatorial Glands, 44.
Resolutions on death of S. W.
Williston, 56.
Raphodolabis, 348.
flavibasis, 343.
Rhigopsis, 11.
Rhycophila, 352.
Rhynchites bicolor, 5.
Riley, W. A., Proposed Amendment, 172.
Ripipteryx, 286, 287.
forcipata, 286.
mexicana, 287.

Robberflies of the Genus Erax, 103.
Rosalia funebris, 3.
Ruckes, Herbert, article by, 192.

Thomas Say

Sabatanca, 355.

Samia cecropia, 195, 354.

Sanders, J. G., and DeLong, D. M.,

article by, 231.

Say Foundation, Report of Treas., 61.

Scaphinotus, 3, 4, 6.
angulatus, 4.
angusticollis, 4.
behrensi, 4.
debilis, 6.
imperfectus, 6.
incompletus, 6.

Scarites subterraneus, 95.

Schizopus, 11.

Schizura concinna, 44.

Scolia, 171.

Scoliz manile in Hawaii, 171.

Scoliaula, 355.

Sibine stimulea, 76.

Sima, 173, 175, 176.

Simopone, 173, 175, 176,

Spiders, New Western, 239.

Stagmomantis carolina, 294.

Staluptius marginalis, 227.

Streptocera Group of the Dipterous

genus Tipula Linnaeus, 89.

Styringomyia, 25.
didyma, 25, 30.

Syntomaspis elegantissimus, 163.
medicaginis, 162, 163.
thalassimus, 162.
umbilicata, 162.
warreni, 163.

Sysyrosea textula, 354.

396

Terminal Abdominal Structures of
Orthopteroid Insects, 267.
Termopsis angusticollis, 302.
Tetraponeura, 176.
Tetrastichus sobrinus, 166.
brucophagi, 167.
Tettigonia verrucivora, 280.
Tettigonoidea, 278.
Thelea polyphemus, 193.
Thyridopteryx ephemeriformis, 76.
Timena californica, 292.
Tineola, 354.
Tipula Dipterous Genus Streptocera
group, 85.
Tipula derbyi, 85.
devia, 85, 86.
fragmentata, 85, 87.
joana, 347.
monochroma, 85, 88.
mutica, 85, 91.
opisthocera, 85, 90.
streptocera, 85.
var. pallidocera, 85, 92.
trypetophora, 85, 89.
Tipulide, 327.
of Hawaiian Islands, 25.

Townsend, C. H. T., description by, 379.

Trichoptera, 349.
Tridactylide, 286.
Tridactyloidea, 286.

Index of Volume XII

Tridactylus, 270.
Trigonoscuta, 11.
Trigonurus, 3.
Trimicra, 25, 30.
lateralis, 30.
Trimeromicrus maculatus, 165, 166, 167.
Triorophus, 9.

Uloborus utahensis, 244.

Valentinia glandulella, 76.
Van Dyke, E. C., article by, 1.
Vanessa antiopa, 193.

Walker, E. M., article by, 267.
Weiss, Harry B., article by, 369.
Welch, Paul S., article by, 218.
Wheeler, W. M., article by, 173, 174.
Williston, Dr. S. W., Resolutions on
the death of, 56.

Wing Structure of Lepidoptera, etc.,349.
Wubana, 252.

drassoides, 252, 253.

pacifica, 253.

retrahens, 253.

Zacotas, 3.

Zelotes ater, 247.
tuobus, 247.

Zopherus, 9.

Zorotypus, 270, 271.

NOTICE TO MEMBERS AND CONTRIBUTORS.

The Annals of the Entomological Society of America, pub-
lished by the Society quarterly, includes the Proceedings of the
Annual meetings and such papers as any be selected by the
Editorial Board.

Papers may be submitted to any member of the Editorial
Board and should be as nearly as possible in the form desired as
final, preferably typewritten, and illustrations must be finished
complete ready for reproduction. Plates must not exceed 5x7
inches unless intended to fold. In general, papers to be accepted
must be original, complete and previously unpublished and,
except in connection with the proceedings, it will not be the
policy to publish preliminary announcements or notes. Authors
will be allowed fifty reprints gratis and additional copies at cost
to the Society.

The Managing Editor is provided with the most recent
address of all paid-up members on record in the Secretary’s
office for mailing the numbers of the Annals and members failing
to receive their numbers should present their complaint to the
Managing Editor within four months from the date of the mail-
ing of the issue. After that time the numbers will be furnished
only at the regular published rate.

The regular annual subscription price for the ANNALS is in
the United States, Cuba, Porto Rico, Hawaii and Mexico, $3.00;
Canada, $3.50; other countries, $4.00. Checks, drafts or money
order should be drawn payable to ANNALS ENTOMOLOGICAL
SOCIETY OF AMERICA, and addressed to HERBERT OSBORN,
State University, Columbus, Ohio, U. S. A.

Requests for information as to membership and the, annual
dues of members may be sent to the Secretary-Treasurer,
J. M. Atpricu, National Museum,’ Washington, D. C.

Communications relating to the ANNALS, and all orders for
separate copies or reprints should be addressed to
HERBERT OSBORN, Managing Editor,
ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA,
State University, Columbus, Ohio.

CONTENTS OF THIS NUMBER.

WALKER, E. M.—The Terminal Abdominal Structures
of Orthopteroid Insects: A Phylogenetic Study--- 267
ALEXANDER, C. P.—Undescribed Species of Japanese
Crane-flies..('Tapulides, Diptera) ce Oe ae B27,
BRAUN, ANNETTE F.—Wing Structure of Lepidoptera
and the Phylogenetic and Taxonomic Value of
Certain Peristent Trichopterous Characters-.---- 349
DICKERSON, E. L. anp Weiss, HARRY B.—The Life-
history and Early Stages of Platymetopius hyalinus
Osb. A Japanese Leaf-hopper in New Jersey.--- 369
PLATH, O. H.—The Prevalence of Phormia azurea
Fallen (Larva Parasitic on Nestling Birds) in the
Puget Sound Region and Data on Two Undescribed

Phes of pimilar Habits ae see a Bp:
Kine, J. L.—Notes on the Biology of the Carabid
Genera Brachynus, Galerita and Chlaenius..... - 382

Price List of Publications.
Annals, Vols. I, II, III, IV, V, VI, VII, VIII, LX, X and XI complete, each. $3.00

Annals, Separate Parts except as below, each... .. cc... cece eee c ccc ccceees 1.00
Agnalss Vis. E-anGa lL APart-3,°eache oy 25 oy cigs ecitla hing sisted Mearkcaxbeta's ints aes .50
Annals, Vol IV, Part 4 each (oor es eae Ripe weche's Rade eee ees 1.50

BACK VOLUMES

Of the ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA may
be secured from the office of the Managing Editor and new members
of the Society who may wish to complete a set are advised to secure
the earlier volumes while there is still a supply.on hand and the price.
is kept at the original subscription rate.

Address HERBERT Osporn, Managing Editor,
ANNALS ENTOMOLOGICAL SOCIETY OF AMERICA,
State University, Columbus, Ohio.

hide

a

‘fy
‘S, a)
Y

|
tf

%

e na
4 Ss
£ - xe yy

ty
Tne

t| l ie
Hi Wait Vey

Hu ms

i) ih

i
?
‘

Cie |

|
i

By Loom)
, é iZ
ore!

era

py

aX
‘i Os .
eX,
re

ny
Hig

|

bait

by, | |
< Ne | Cay |

rf
“Ay
S

“
~ 2
~f

| | i,

ys ; | i

<p
“A,

INSTITUTION LIBRARIES

CHI UN

Vill

3 9088 00844 611