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The Entomological Society of America _
_ Founded 1906.
OFFICERS 1919.
President
James G. NEEDHAM. Ste Se . : .
pee . First Vice-President
Justus W. Fotsom .- - 3 - ° ° :
Second Vice-President 3 Bi ate
R. V. CHAMBERLIN . : ‘ rf . Cambridge, Mass.
' Managing Editor Annals ie Siar
Herpert OssoRN - = - = + ~~ se Columbus, Ohio: . %
Secretary- r: reasurer ee
J. M. AtpricH . : :
Executive Committee
THe OFFICERS
AND
F. E. Lutz, ARTHUR GIBSON, £
G. A. Dray, G. C. Crampton. r
f Committee on Nomenclature | «RRC as
E. P. Fett, T. D. A. CocKERELL, NATHAN BANKS.
Thomas Say Foundation RSS
NatHan BANKS, A. D. MacGiirvray, P. P. CALVERT, ae
E. B. WILLIAMSON, . cnt
es J. M. ALDRICH, Editor, E. D. Batt, Treasurer. 9°!

Committee on National Museum
' T. D. A. CocKxEerRELtt, Chairman,

HERBERT OSBORN, Ww. BARNES,
James G. NEEDHAM.

ANNALS
bictogicah
OR= Medical
Suiaja?

The Entomological Society of America

VeLuUme. X1ii;-1919

EDITORIAL BOARD

HERBERT OSBORN, Managing Editor,
COLUMBUS, OHIO.

L. O. HOWARD, WM. A. RILEY,

WASHINGTON, D. C. MINNEAPOLIS, MINN.
VERNON L. KELLOGG, | FRANK E. LUTZ,

STANFORD UNIVERSITY, CALIF.. NEw YorkK 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, EDwIn 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

CAbtyereilinG lays ID WoNtEs G2) Vs ee Be sexcalise sheet aie eT i Sy ee ec ane ET 25
HAH we: — he Californian. species of Malthodes..................:2.-++- 31
HERRICK, GLENN W. and DETWEILER, JOHN I).—Notes on the Repugnatorial

Clandssom ©erhaimsNotodonid Caterpillars: 5.95220... cu eo ee 44
MosHER, EpNA—Notes on the Immature Stages of the Deltometopus rufipes

Wielsen(ColeoptenavlticneniiGae ee, anc. 2. a: ases chy ees ai vale soot seas eee: 49
AypricH, J. M.—Proceedings of the Baltimore Meeting...:.................: 57
Dietz, W. G.—The Streptocera Group of the Dipterous Genus Tipula,

DHOMEBIOSS. 5 ac Goce Sa ne CE ocoslGt.6 OS CER AR INOS Sees Ce eee rae ee 85
CLAASEN, P. W.—Life-History and Biological Notes on Chlaenius Impuncti-

ROWS: GE oget ans co keno Sees 6 pw 6 5 Oo OO ee eee ee ee 95
EINES ASS: —Robpertiies ofthmelGenus rax. ..22.'. 6.566 ge shee ence ee eee ns 103
Gauan, A. B.—Some Chalcid Wasps Reared from Cecidomyid Galls.......... 159
Murr: .— Lhe Progress of Scolia mantlae Ashm... >... 0.0... 0.002.055 0. dece: 171
WHEELER, WILLIAM M.—The Ants of the Genus Metapone.................... 173

RuckeEs, HERBERT—Notes on the Male Genital System in Certain Lepidoptera. .192
WeEtcu, Paut S.—The Aquatic Adaptations of Pyrausta penitalis Grt.

(LIS ONSIOVEPE Vice 208 So, Se Se Ree 8 a 213
FRACKER, S. B.—Chariesterus and its Neotropical Relatives (Coreidae

LEISTRSNLOSONASIH) |. we ane, cic VER ORE Rh oe RNC ae OP resinlet 2 ae os SER. = 227
SANDERS, J. G. and DELonc, D. M.—Eight New Jassids from the Eastern

Ufa BELG IS HIWEC A Se ta SR neo os & 5 Sal Aces Sees Es 3 CCLCRE Mea Panera 231
CHAMBERLIN, RaLpH V.—New Western Spiders....................200002005. 239
WALKER, E. M.—The Terminal Abdominal Structures of Orthopteroid

linsecisrea Abhi ogenetiCnsiliGie serdar es meres en eae ageing cee ae = 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
DIcKerRSON, E. L. and Wetss, Harry B.—The Life-History and Early Stages

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

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

gevrilsyacl MRE Ole Shranbilene [SIZ oni, Sete Rr eGe Roeko On 8 ot ) Olen ReECenn cea ReINS cl ici 373
TownseEnD, C. H. T.—Description of the New Species of Phormia.............379
AtpricH, J. M.—Description of a New Species of Hylemyia................... 380
Kinc, J. L.—Notes on the Biology of the Carabid Genera Brachynus,

feraletiteay ane @ MIACIUS mms s feo sv hae he iors vcdis vis olelslatelacyemeslalein aig etenewy 382

DATES OF ISSUE.

March, 1919, number mailed April 26, 1919.

June, 1919, number mailed July 2, 1919.

September, 1919, number mailed October 21, 1919.
December, 1919, number mailed December 31, 1919.

ANNALS

OF

The Entomological Society of America

Volume XII DECEMBER... [9 19 Number 4

THE TERMINAL ABDOMINAL STRUCTURES OF
ORTHOPTEROID INSECTS: A
PHYLOGENETIC STUDY.

By E. M. WALKER, 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 Needham, 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, 1. e., the Blattide, Mantide, Phasmide, Acrididz
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-

' 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.

1919] Walker: Structure of Orthopteroid Insects 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 Protobiattoidea. 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 Elcanide and Locust-
opside, 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 tibiz.

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. campodetformis). The
former group 1s 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, 16, 17, 18, 719)? 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. 43 (’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 Phasmidz arose, not from Saltatorial Orthoptera
(Locustopsidez), 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, Timema californica.

3. The 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. 343 (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.)

* Crampton, G. C., Ent. News, Vol. XXVI, pp. 337-350, Pl. XIII (1915);
Ent. News, Vol. X XVII, pp. 244-258, 297-307 (1916); Can. Ent., Vol. XLIX, pp.
213-217, Fig. 9 (1917); Ent. News, Vol. XXVIII, pp. 398-413, Pl. XXVII (1917);
Journ. N. Y. Ent. Soc., Vol. X XV, pp. 225-237 (1917); Bull. Brooklyn Ent. Soc.,

oe 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, Embiids 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 8th, 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.

274 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.)"; 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. 2386 (1917). Also termed
“‘Basalstuck’’ (Van der Weele, Tijd. voor Ent., Deel XLIX, pp. 99-198, Pls. 1-3,
ey and ‘‘basal plate’’ (Walker, Univ. of Toronto Studies, Biol. Ser., No. 11,

mM Crampton, op. cit., Pl. XVI, Fig. 7; Pl. XVII, Fig. 12.

12 Crampton, op. cit., Pl. XVI, Figs. 1, 6; Pl. XVII, Fig. 10.

1919] Walker: Structure of Orthopteroid Insects 275

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
17, pp. 207-251, Pls. 10, 11, 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. In the 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 Tettigoniide) 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 Tettigoniide 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 valvule 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 papille 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 Aeshnide), 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 valvulz
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 basivalvule
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 valvulze are much shorter than the other two
pairs and are somewhat widely overlapped by the dorsal
valvule. 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 valvulz, 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
foliowing characters: The distinct basivalvule, the less com-
pressed ovipositor and the less complete cohesion of the valvule
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. f

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 somettmes 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.

‘4 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 sensille, 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 < ~—t+remely 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 Tettigoniide
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
ground-dwelling species as Gryllus assimilts.

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-
gonude 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 Grylloblatia
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)!%. 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 P1. (1870).

1919] Walker: Structure of Orthopteroid Insects 283

blades of the forceps, the mov-~” 3 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 valvulze
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 in
the Gryllidz. 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 valvule 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 (Acrydidz, 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
Acrydiidz 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 Metanoplus. 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-
dactylidz, which also resemble the Acrydiidz 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
(q. v.).

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, Tvridactylus, 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 7ridactylus is 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 See 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 Tridactylide 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 precesses, 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. 4038-480, 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,
loc. 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 valvule or their homologues, and thus belong to
the ninth segment. These peculiar structures, which are present

in both sexes in TJvridactylus 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-

% 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 valvule, 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 Grylloblaita, so that we may consider
this form next.

A glance at the end-segments and ovipositor of Grylloblatta
(Figs. 31-35) 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-like, 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 slightly 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 Tettigoniida. 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 Tettigoniide. 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 valvulz 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, 1. 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 valvulz 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 Mantidz, Tetti-
gonide and Gryllidz. .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. 131-139. 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 Orthcptera 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 styli.
In these respects they approach the Blattide and Mantida,
but the valvule of the ninth segment have not the terminal
position, nor have the dorsal valvulz 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 Tettigoniide.

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 (basivalvule) by Crampton. The
dorsal valvule are flexible, triangular flaps, whose bases meet
the inner edges of the valviters 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 Zimema, 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 femoraia 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 7zmema 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 A merica [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 Timema.
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 hes 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 Gryllide
and Tettigontide.

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 slight 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
Mantidz 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 Mantide 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 is
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 Pertplaneta
and Blatita 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, Parcoblatta 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 Mantide, but is much less
pronounced. In the immature Parcoblatta, 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 7imema californica (see pp. 292-3).

Immediately behind each basivalvula is the large valvifer,
which, as usual, connects the bases of the dorsal and ventral
valvule 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 valvulez, from which they
are strongly bent outwards and downwards. These are simply
the bases of the dorsal valvulz, 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 es [Vol. XIT,

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., Parcoblaita, 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 slight 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 valvule 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 valvulz
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 primitive 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 valvule 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. Itis 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.

302 Annals Entomological Society of America [Vol. XII,

Isoptera.

Two species of this order were examined, Termopsis angusti-
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 flattened,
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
‘“‘senital 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 valyifers, 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 Parcoblatia.

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 [soptera 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, Allosthetidz, 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)*2 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 valvule, belonging to the eighth
and ninth segments: The former pair is long and slender

22 Verhoeff, H. W. Nova Acta. Acad. Caes.-Leop., Vol. XXXI, 1903; pp.
277-278.

1919] Walker: Structure of Orthopteroid Insects 309

and obviously represents the ventral valvule. The other pair,
from its lateral position and comparatively broad form is doubt-
less the dorsal valvula. 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 @ 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 (Diplatys, 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 cerci 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 7hysanura have also no ovipositor does not affect
the question. There is evidence that some of the Palaeo-
dictyopteri 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
tespiration 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 valvula. 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, Pls. 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.

*4 Morrison, Emily Reed, Can. Ent., Vol. LI, No. 6, pp. 139-146 (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 (Aeshnide#:—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, Machiiis 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 in which this is the case.

The ventral valvul& 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
valvule 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: Valvule 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 valvule 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-like subgenital plate, largely concealing the
Ovipositor; ninth sternal regional horizontal; valvifers and

1919] Walker: Structure of Orthopteroid Insects 311

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 valvulz, 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

Embiudina—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 Machilidez 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 covxites,
these probably representing flattened coxz of abdominal limbs
which have otherwise disappeared (Haase, ’89).% The sterna of

28 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 aesions will
be discussed in Part II.

*6 Escherich, K. Das System der Lepismatiden, Zoologica, Bd. 18, Heft 48,
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.
cex—coxite. sp—spiracle.
eg—egg guide. spth—aperture of spermatheca.
ga—genital aperture or vulva. st—sternum, sternal region.
hilm—internal lamella joining bases of | ste—sternite.

inner valvulae. stl—stylus.
jap—inferior apophysis. sv—superior intervalvula.
int ap—intersegmental apodeme. tg—tergite.
im—intervalvular membrane. vd—dorsal valvula.
iv—inferior’intervalvula. vi—valvifer.
1lb—inferior lobes of inner valvulae. vi—inner valvula.
p—pons valvularum. vv—ventral valvula.

PLATE XX.

—_

a ial Ried ks oe alles a a

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 sad 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 XXI.

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.

bt

316 Annals Entomological Society of America [Vol. XII,

Pram XXIII.

Grylloblatta cam podetformis, adult; lateral view of end segments and ovipositor.

Same; ventral view. h

Same; dorsal view of ovipositor, left ventral valvula bent outwards, right
not shown.

Same; inner view of structure at base of ovipositor, left side.

Same; anterior view of base of ovipositor.

Same, nymph; lateral view.

PLATE XUV.

Grylloblatta campodeiformis, nymph; ventral view of end-segments.
Same with ventral valvulae bent forward to expose inner valvulae.
Timema californica, adult; ventral view of end segments.

Same, lateral view.

Diapheromera femorata, adult; ventro-lateral view of end segments.
Same; lateral view of ovipositor; most of the 9th tergite cut away to show ~

the superior apophysis.

Same; dorsal view of ovipositor.

PLATE X XV.

Stagmomantis carolina, adult; 8th sternum and ventral.

Same; dorsal view of ovipositor, with dorsal valvulae spread apart.

Same; ventral view of ovipositor with ventral valvulae spread apart.

Same; lateral view. of terminal segments and ovipositor with 7th sternum
bent downwards to expose the valvulae.

Same; inner view of structures at base of ovipositor.

Same; anterior view of base of ovipositor.

PLATE XXVI.

Parcoblatta pensylvanica, adult; ventral view of ovipositor, the left ventrai
valvula cut off at base.

Same; the valvulae bent forward to show their dorsal surfaces and the right
dorsal valvula removed.

Same; ventro-lateral view of terminal segments, the 7th sternum removed.

Same; ovipositor removed, cleared, and viewed from above. The ventral
valvulae and the dorsal valvulae, except their bases, are omitted.

Same, last nymphal stage; ventral view of end segments with the 7th sternum
except a small part, removed.

5 to 68: Same; successive stages in the development of the genitalia, ventral

9 view, 7th sternum removed, cerci 10th and anal segments omitted. Fig. 58
‘ is from the adult.

PLATE XXVII.

Cryptocercus punctulatus, adult; ventral view of end segments, the 7th sternum
removed.

Termopsis angusticollis, adult; lateral view of end segments.

Same; ventral view of end segments, the 7th sternum removed.

Forficula auricularia, adult; lateral view of end segments. °

Same, dorsal view.

Same, ventral view, 7th sternum removed.

Anisolabus maritima, adult; ventral view of end segments, the 7th sternum
removed.

Kalocrania marmoricrura, adult; ventral view of ovipositor and neighboring
sclerites. (After Zacher.)

Echinosoma occidentale, adult; ventral view of ovipositor and part of segment
10. (After Zacher.)

PLATE XXVIII.

Embia major, adult; lateral view of end segments.
Pteronarcys proteus, adult; ventral view of end segments. (After Smith.)
Perla lycorias, adult; ventral view of end segments.

Aeshna canadensis, adult; ventral view of end segments.

Machilis sp., adult; ventral view of end segments.

Same; 8th sternum and gonapophyses (ventral valvulae).

Same; 9th sternum and gonapophyses (dorsal and inner valvulae.)

ANNALS E. S. A. Vou. XII, PLATE KX:

_E, M. Walker.

ARS EES AC VoL. XII, PLATE XXI.

E. M. Walker.

ANNALS E. S.A. VOL. XTI, PLatTE XXII.

E. M. Walker.

ANNALS E. S. A. VOL. if, PLATE XXIII-

a

—S 5S

E, M. Walker.

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

ne

st 8 bs v

fw A 4]

E. M. Walker.

ANNALS E. S. A. : VoL. XII, PEATE XXV.

E. M. Walker.

ANNALS E.S. A. VOL. <i, PLATE XX Vi-

E. M. Walker.

ANNALS E.S. A.

VoL. XII, PLaTEe XXVII.

E. M. Walker.

Vou. XII, PLATE XXVIII.

ANNALS E. S. As

2. as

SM on
Pee,
r, ee Stee i

one.

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. Halteres rather long, pale, the
knobs dark brown, the base of the stem more yellowish. Legs with
the coxe and trochanters yellow, the former very sparsely gray pruinose,
the fore coxze dark basally; femora dull brownish yellow, somewhat
darkened on the apical half; tibiae and tarsi brown. Wings nearly
hyaline, the stigma large, oval, very pale brown; veins slender, brown.
Venation: Sc, ending opposite origin of Rs, Sco far before the tip of
Sc, the latter being longer than the basal deflection of Cm; r arcuated,
at the tip of R:; 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

328 Annals Entomological Society of America [Vol. XII,

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, 2, March 31, 1919.

Paratopotypes, 6 & 9’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 Me 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 lghter
gray. Pleura dark gray. MHalteres yellow, the knobs dark brown.
Legs with the coxe brownish yellow, darker basally; trochanters dull
yellow; femora dull brownish yellow, darkened toward the apices, the
extreme tips pale; tibiz 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, Scz exactly opposite, the origin of Rs;
Rs angulated to almost square at origin; cell /st Me 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, 9, 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 1st Mz closed, about twice as long as vein Cm
beyond it.

1919] Alexander: Japanese Crane-Flies 329

Female—Length about 6.3 mm.; wing, 6.5 mm.

Rostrum and palpi brown. Antennz 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 coxz 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, Scg exactly at the
origin; Rs less than twice the deflection of R::;; r-m about equal to m, a
little shorter than 7; cell 1st Mz long, about twice the section of Cu
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, 2, 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; Sc. 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, Sc. lacking; a supernumerary cross-vein
at about midiength of cell Sc, located in the second anterior brown
blotch; Rs about two and one-half times as long as the basal deflection of
Ris; 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, 2, July 9, 1919.

Paratopotypes, 4o’s, October 20-21, 1919; 19, 1 sex
uncertain, July 12, 1919.

One paratype bears the label, ‘‘ This insect was flying about
over a stream.”

Antocha Osten Sacken.

Proantocha, subgen. n.

Antenne 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 tibiz 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 331

The genus Antocha has, till now, been one of the most
compact genera in the Tipulide. 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
tibize 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. Antenne 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 Cm, 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, Ri and C
beyond the stigma chestnut brown; basal deflection of Cu 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. Antennz 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. MHalteres 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 M2 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 setz
that are larger and stouter at the tip of the organ. Gonapophyses long,
acicular, almost straight, the tips acute.

Habitat: Japan.

Holotype, #7, Meguro, Tokio, April 21, 1919 (R. Takahashi).
Allotopotype, ?, March 25, 1919.

Paratopotype, 2o’s, March 26—April 23, 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 333

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, 2, 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, 6.3 mm.; wing, 4.7mm. Fore leg, femur, 2.9 mm.;
tibia, 2.6 mm.; hind leg, femur, 4mm.; 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
coxz 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; tibiz 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 tibiz 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, 92, 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 tibie 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 tibie and tarsi abruptly light yellow; wings gray, the
stigma brown, the costal margin with yellowish pubescence; anal veins
divergent.

Male—Length, 4-4.3 mm.; wing, 5.1-5.5 mm.

Rostrum and palpi dark brownish black. Antennz 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 ; tibiz 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 335

of M pale; veins brown; wings broad, densely covered with hairs.
Venation: Cell /st M2 small, closed; basal deflection of Cm slightly
before the fork of MW; 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, o&, Meguro, Tokio, March 31, 1919 (R. Taka-
hashi).

Paratopotypes, 1 #, 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., Antennz 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 wrth the coxz 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 7; 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 M2 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 @’s, April 9-25, 1919.

336 Annals Entomological Society of America [Vol. Dabs

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. Antennz
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 foveze, 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 cox
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: ron Re, about its own length
beyond the fork of Roi3; deflection of Ray; longer than r-m; cell 1st Me
closed; basal deflection of Cu, beyond the fork of WM; 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, 9, March 23, 1919.

Paratopotype, o', 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. Wi ings
nearly hyaline; stigma dark brown, conspicuous, sending a narrow brown
seam.along the cord; outer end of cell 1st Mz 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 7; r on R> near its base; cell 1st M2 closed; m shorter
than the deflection of M3; basal deflection of Cu; slightly before the fork
of M; 2nd anal vein slightly sinuous, but cell /st A broadest at the wing
margin.

Abdomen black, including the male genitalia. Male hypopygium
with the pleurites stout; pleural appendage oval, tumid, with long
yeliowish 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: Japart.

Holotype, «, Meguro, Tokio, April 2, 1919 (R. Takahashi).
Allotopotype, @, April 2, 1919.

Paratopotypes, 27’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 1st M2 open; anal veins con-
vergent; male hypopygium with eight chitinized pomits surrounding
the penis-guard.

Male—Length, 3.84.2 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; tibize 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
vellow 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 sete. 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 # 9, March 25—April 25, 1919; paratype,
Q, May 29, 1919; 64 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 M».open; anal veins divergent.

Male—Length, 5-5.5 mm.; wing, 6.1—7.1 mm.

Rostrum black, densely gray pruinose; palpi dark brown. Antenne
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 praescutum gray with three
brown stripes, the median stripe broad, slightly constricted by the
conspicuous, elongate pseudosutural fovez, indistinctly bisected by a
pale median line; tuberculate pits small, very widely separated, lying
slightly anterior to the level of the pseudosutural fovez; scutum gray,
the lobes indistinctly marked with brown; remainder of 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 339

more yellowish; stigma very narrow and indistinct, brown, confined to
vein Ri; veins dark brown, Sc yellow. Venation: Sc long, ending
opposite 7; r on Re 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. Antenne 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. MHalteres light yellow.
Legs with the fore coxe black, grayish pruinose; middle coxe dull
brownish yellow, dark at the extreme base; hind coxe largely brownish
yellow; trochanters dull yellow; femora yellow, the tips broadly dark
brown; tibize yellowish brown, the tips darkened; tarsi dark brownish
black, the base of the metatarsi brown. Wings grayish, strongly
yellow at the base, this color including most of the costal and subcostal
cells; stigma oval, grayish brown, almost evenly split by 7; narrow gray
seams along the cord, outer end of cell /s¢ M2, origin of Rs, and the forks
of Roi3 and My42; veins brown, those in the costal and basal regions
yellowish; apical cells of the wings strongly pubescent, this extending
from cell 2nd R, to cell Cu. Venation: Sc: slightly removed from the

340 Annals Entomological Society of America [Vol. XII,

tip of Sa; Rs long, in alignment with Rs43, this latter shorter than the
basal defiection of Cm; r removed from the tip of Ri; inner ends of cells
R;, Rs, and 1st Mz in oblique alignment; petiole of cell M; about equal
to or shorter than this cell; basal deflection of Cm slightly before mid-
length of cell 1st 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. ee
Allotopotype, 2, April 19, 1919.

Paratopotypes, 18 @ 9, 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 line;.
wings with a heavy brown Baten abdominal segments bicolorous.

Male—Length, 5.6 mm.; w ing, 7.3-7.4 mm.

Female—Length, 8.7 eae ; 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 cox dark brown; trochanters brown, with a black spot on the
posterior face; femora dull yellow, the tips broadly dark brown; tibize
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 R: and R: 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 Cand Sc yellow, the other veins brown. Venation:
Rs strongly -arcuated at origin; hog and Mj. 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, 2, 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 /st Me
subquadrate, basal deflection of Cu; beyond the middle of its length.

Male—Length about 14 mm.; wing, 16.4 mm., its greatest width,
5.5 mm.

Description from an alcoholic specimen.

Rostrum, palpi and antennze 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: tibia 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 1st Me, at the origin of Rs and at the fork of Miz42. Venation
generally similar to L. (P.) rufibasis O. S. (Northeastern North
America), but cell 1st Mz much shorter, subquadrate; Ro:3 a little
longer than r-m; basal deflection of Cu, beyond mid-length of cell
1st M2; petiole of cell Mj, 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, #, Chichibu, Province Saitama, October, 1918,
(R. Takahashi).

342 Annals Entomological Society of America [Vol. XII,

Polyangeus 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, M; and M; tibiz
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 coxze and trochanters dull yellow; femora dull yellow, the tips
broadly dark brown; tibize 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 Ro,3; at the point where r is located in the genotype, P. maculatus;
Ro.s44 longer than r—m, cell 1st Mz closed, m being located between Mo
and M3; supernumerary crossveins in cells R3; near the tip, R; at about
two-thirds its length, at about midlength of M, 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 M. and the presence of an additional super-
numerary crossvein in cell M..

1919] Alexander: Japanese Crane-Flies 343

Rhaphidolabis Osten Sacken.
Rhaphidolabis flavibasis sp. n.

Antennz 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; tibiz 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; R»
ublique, its fusion with R; at the wing tip slight; Ro4344 rather short,
osually a little longer than r-m; cell M, present; cell 1st Mz 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-setze; inner pleural appendage
more chitinized, slightly curved, along the inner margin with about a
dozen spinous sete 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, 2, April 4, 1919.

Paratopotypes, 8 #@ 2, 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.

Antennz 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 three dark brown
stripes of which 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; tibiz dark
brown, the tips black; tarsi dark brownish black. Wings with a pale
gray tinge; stigma elongate, brown, extending from 7 to the tip of Riz;
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; Roi3;4 short, about equal to r-m; apical fusion
of Re with R; slight; cell 1st M2 open, cell M, 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
setze 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. MHalteres 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; tibiz
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, 9, May 31, 1919.

Paratopotypes, 27's.

Liogma Osten Sacken.

Liogma serraticornis sp. n.

General coloration deep shiny black: flagellar segments in the male
deeply serrated; wings with the stigma y ellow or brownish yellow.

Wie tenoth about 15 mm.; wing, 12—12.1 mm.

Female—Length about 14 mm.; wing, 12 mm.

Rostrum dark brown; palpi yellowish brown. Antennz 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.
kuwanazt; 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. MHalteres yellow.
Legs with the coxz 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 Rus on My42; basal deflection of Cw, before or slightly beyond
the fork of M.

Abdomen shiny black throughout.

In the female sex, the antennz 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 &’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. Antenne with sixteen segments,
simple in both sexes; in the male elongate-cylindrical, clothed with an
abundant white pubescence; antennz 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. MHalteres
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; tibiz 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 R>, 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).

—E——E—————

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 13 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 coxz and trochanters yellow; femora
dark brown, paler at the base; tibiz 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 Ris
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, o, Saitama, May 29, 1919 (R. Takahashi).
Allotopotype, 9.

Paratopotype, @.

Tipula Linnezus.
Tipula joana sp. n.

Antennez 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,

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 setz.

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 coxe blue-gray; trochanters yellow; femora
brownish-yellow, brightest at the base, the tips brownish black; tibiz
brownish black, darkest apically, the posterior tibiz 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 R; and R; and part of Mi.
Venation: Rs short; Ry 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).

Paratapotype, o.

This exquisite species is dedicated to the 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 T7pulodina 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 7ipulodina (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.

Il. 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
tesembles 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 pupe, 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

350 Annals Entomological Society of America [Vol. XII,

Micropterygide and the generalized Trichoptera,* and a pupa
resembling that of the Microptergyide. The venation of the
Nepticulide (Braun, °17) shows highly specialized features.
Tt 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
Hepialida, 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.

1919] Braun: Wing Structure of Lepidoptera 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 patt.

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
Micropterygidz 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 ltepidopterous characters origi-
nate in the Micropterygide, necessitate the conclusion that the
Micropterygidez 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 1s 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 1st 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 Mnemonica. In the hind
wing, Cu extends straight to the margin from its peint of
separation from the /st 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 trachee; Figure 3 shows in detail the course
of the trachez at the extreme base of the hind wing. In the
fore wing the coalesced bases of Cu and 1st 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 Microp-
terygide and Trichoptera. The tracheze in the base of the
hind wing (Fig. 3) show that the relations of Cu to the Ist 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 1st 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 ist 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 Tineidez (e. g., Tineola), Plutellide
(e. g., Pliniaca), Eucleide (e. g., Sisyrosea textula), Tortricide,
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 pupze 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
1st 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 tracheze
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 300

the course common to Munemonica 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 Hepialidz 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 ist 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 Micropterygidz, as Sabatinca, this coalescence of
veins is shown in one species and not in the other. The second
branch of the third anal vein is but faintly shown in Epr-
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 Frenatze, I have found no indication of
the presence of the second branch of the third anal vein. It is
suggested asa plausible explanation of the atrophy of this vein
that its disappearance is correlated with the loss of the holes
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
Micropterygide 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 immediately 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
is 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 1st 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
trachee, indicates any such anastomosis.

1919] Braun: Wing Structure of Lepidoptera 357

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
Hepialidz, 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 typicaliy lepidopterous characters,
such as the single spined frenulum of the male, the short tongue
and six-jointed maxillary palpi characteristic of many primitive
Frenate, obsolescence 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

358 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 Eriocraniing, 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
Nepticulidz (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 Nepticulide from a fibula in females of
the more generalized genera can be traced through various
changes until it 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 Tineidez, 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 Frenatz 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 Hepialidze 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 Micropterygidze
than to any other Lepidoptera, and also indicates trichopterous
affinities. However, the possession by Micropterygide 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, Nepticulide 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 Incurvariide,
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, '4aj'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 is
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-like,
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
robinie, 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 Nepticulidze 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., Hieroxestis, 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
slight 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.

Ill. 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 Nepticulide, 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 pees 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 Cosside show some degree
of relationship; a third much branched line includes the frenate
Lepidoptera, of which some members such as the Prodoxide,
Incurvariidz, 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. XLIII, 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.

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. 3. 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 Renita flavipunctalis.

Fig. 8. Wings of Ectoedemia heinrich, female.

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

Cu risthF ==

~

te.
Neo 277)
veSS

Ree

Annette F. Braun “

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 NyMPHAL STaAGE—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.

SECOND NyMPHAL STAGE—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. Antennz 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.

THIRD 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 H yalinus 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.

Fourta 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 NyMPHAL STacGE—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; antennz 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.

Annals Entomological Society of America [Vol. XII,

Oo
A
bo

ApuLt—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 is 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 1s 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 475 mim. -

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 it 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 O. E. PLaTH.

(Together with Descriptions of the Two New Flies by
C. H. T. TownsEenpd and J. M. Atpricn.)

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. 30-39. (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. XXI, p. 37. b. Univ. Calif. Publ. Zool., Vol. XIX,
pp. 198-199).

373

374 Annals Entomological Society of America |Vol. XII,

Phormia metallica Townsend! and Hylemyia nidicola Aldrich.
The frequency with which the larve of Phornua azurea Fallen
were encountered in birds’ nests, and the number of larve and
pupz per nest, is shown by the following tables:

TABLE I.
Species of Bird Nests | Infested| Uninfested | Infesta-
examined| nests nests tion

Nuttall Sparrow (Zonotrichia leuco-

phrys nuttalli Ridgway)............ 3 3 0 100%
Rusty Song Sparrow (Melospiza
melodia morphna Oberholser)....... 2 2 0 100%
Willow Goldfinch Ce So tris-
tis salicamans (Grinnell)............ 2 ON 0 100%
Oregon Towhee (Pipilo maculatus
orevonus Belle Ai eee eee eee 1 1 0 100%
Russet-backed Thrush (Hylocichla us-
Hilatan (Nira) eet ee eee 5 + 1 80%
Western Robin (Merula migratoria
propinqua (Ridgway).............. 25 18 7 72%
Cedar Waxwing (Ampelis cedrorum
Wirellloi i hak. eee eee 3 1 2 33%
Yellow Warbler (Dendroica aestiva
(Gimelin eae eat ce shied Oe 5 1 4 20%
Cliff Swallow (Petrochelidon albi-
OTUSION Yh Bas tee econ at ets Aeteers if 1 6 14%
English Sparrow (Passer domesticus :
| Grhatnic¥ a0) eer oe iat @ Gen Se ences 1 0 1 0%
Shotalsi, Me dash ce tcrwa adelante 54 33 21 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 nutialla Ridgway)............ DORAOM ST: sole ee ae: 166 55
Rusty Song Sparrow (Melospiza me-
lodva morphing Oberholsder)........%|52, 55. ....5... 0.00006. 107 53
Cliff Swallow (Petrochelidon albifrons
REAM E apie Mera wk ee Bilger te Mee Felt 20 47 47
Oregon Towhee (Pipilo maculatus
Beli). . cao qckedticates Eon et lca eee AN RLS OAT, 45 45
Willow Goldfinch eee tris-
tis salicamans (Grinnell). . BRAS seek. nen cent 87 43
Yellow Warbler (Dendroica aestiva
Ree eR IR Ore S a9 She. witgere et «levee so SOS ou eotets Bon 39 39
Western Robin (Merula propinqua
Muni aycuye) akAee ulead? As kid bo 7, 11, 15, 16, 18, 18, 21,
23, 26, 27, 27, 29, 37,
39, 40, 48, 72; 188... 607 33
Russet-backed Thrush (Hylocichla us-
PANCROM NCAT) octet ccs es se eaves 3 BOM hero Oot tcey fsa a 108 27
Cedar Waxwing (Ampelis cedrorum
MVERMINCHU Ree ee os Sal, oc hdc leuloes Die teretgerinte Pepe Tac 2 2
‘UGHBIS dts S Bao Gooe Bere Oe eee 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 larve were taken from
the nest. The larve pupated in a few days and both larve
and pupez 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
Berkeley, California, the preceding summer (Cf. a. The Condor, Vol. XXI, pp.
36-37. b. Univ. Calif. Publ. Zool., Vol. XIX, pp. 196-197).

——————

1919] Plath: Larve of Phormia ~ avd

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. ‘Fwo 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 larve 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, ibid. 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 azurea 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
larve 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. XXI, pp. 35-36.

1919] Plath: Larve of Phormia _ 379

DESCRIPTION OF THE NEW SPECIES OF Phormia.

By C. H. T. TownseEnp,
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. Antenne
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 tegule 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 [Vohk3erP

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. ALpRicu,
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 middie; 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

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 Chlaenius 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 Chlaenius
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 larvae 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.

+ 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: Buology 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 MupD 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
3.5 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 surface 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. impunctiifrons, 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 X1, Figs. 1 and 2) are convex
and oval in form, coarsely granular or rough in surface; length
3.75 to 4.5 mm., width 2.75 to 3 mm. 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 MupD 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 385

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 celis of Chlaenius
may be compared to a pouched envelope in form and mechanism.
The folding flap and the sealing of 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
beetie, 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.
pbhragmitella, but it showed a decided preference for the little
larve of A. obliqua.

During the second and third instar all other food was refused
except larve 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 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
larve.”’

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
climbers, 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, in 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 pupz of several species of ants. The ant larve and pupe
were easily collected and proved an ideal food for the Carabid
larve, 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 larvez 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 pupz 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, bya minute Hymenopteron, Prosacantha caraborum
Riley, which is a common parasite of several species of Chlaenius.

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 cyantpennis, 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 diameters. )

Fig. 1. Masses of mud cells of Bee cyanipennis. Cells made by beetles in
captivity.

Fig. 2. Mud cells of Brachynus c oe as found in nature on the under surface
of a stone.

Fig. 3. Mud cell of Galerita bicolor on id under surface of Impatiens leaf.
Fig. 4. Mud cell of Chlaentus 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. 3. Mud cells of Chlaenius tricolor 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. - ~ Vou. XII, Plate XXX.

XII, PLATE XXXI.

VOL.

J. L. King

INDEX OF VOLUME XI.

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. O21.
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, Sie
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 catalpe, 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

Coniotus, 11.
Conocephalus, 280.

fasciatus, 279.
Copris, 12.
Cornicularia monticolens, 251.
Crane Flies of the Hawaiian Islands

(Tipulidz 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.

DeLong, 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, 827.
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.

Index to Volume XII —

Diplocephalus wamotsus, 251.

Dipterous Genus Tipula, Streptocera

Group, 85.
Dissosteira carolina, 193.

Distribution of Insects in Western

IN SSA: a
Drassodes gosiutus, 245.
melius, 246.
robinsoni, 245.
Dyctinoides, 248.
arizonensis, 244.
Dystaxia, 11.

Edrotes, 9.

Eleodes, 10.

Embidea, 305.
Embidiina, 305, 313.
Ephebomyrmex, 175.
Ephelia subaprilina, 340.
Ephemerida, 308.
Epimartyria, 353.

Erax, Robberflies of the Genus, 103.

Erax zestuans, 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, 184.
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, 133.
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.
rapax, 107, 142.
rufibarbis, 106, 137.
tufitibia, 106, 128.
sagax, 105, 118.
slossone, 106, 121.
snowi, 105, 116.
-spiniventris, 106, 135.
splendens, 107, 143.
stramineus, 107, 140.
stylatus. 106, 127.
subcupreus, 106, 133.
subpilosus, 105, 118.
tagax, 105, 1138.
tabescens, 106, 126.
texanus, 107, 151.
triton, 107, 151.
truncatus, 107, 143.
tuberculatus, 106, 135.
unicolor, 106, 132.
varipes, 107, 152.
willistoni, 105, 110.
zonalis, 105.
zonatus, 112.

Erioptera, 338.
bifurcata, 338.

Eucharis myrmicia, 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, 1).

Gonomyia, 25, 30.
gracilis, 30.
hawaiiensis, 30.
skusei, 30.

Grammonota orites, 249.
obesior, 250.
simplex, 250.

Grvllide, 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-
bidz Coleop), 95.

*
394 Index of Volume XII ©

Linyphia latescens, 248.
orinoma, 248.
insignis, 248.

Liogma, 345.

serraticomis, 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. it. article by, 65.
Megalopyge 76.
Melanoplus bivittatus, 283.
Melanotus, 12.
Tr eroeti Ants of the ——— 173.

Micropterygidz, 349.

Miscodera insignis, 4.

Mnemonica, 352.

Monilema, 9.

Mosher, Edna, article by, 49.

Muir, F., article by, 171.

Myrmecia forficata, 13.
gulosa, 13, 15.

. 24.

Nepticula planatella, 354.
Nepticulide, 349.
Nesospeza, 347.

tarsalis, 347.
New Western Spiders, 239.
Nezara viridula, 193
Notodontid caterpillars, Repugnatorial

glands of, 44

Nyctoporis, 11.

quadripunctatus, 282.

Orthoptera, 278, 310.
Orthopteroid Insects, Abdominal
Structures of, 267.
Osborn, Herbert, Report of Man-
aging Editor, 61.

Pachysima zthiops, 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.

5.

Phasmoidea, 292, 310.

Index of Volume XII

Phlepsius tenuifrons, 235.
tinctorius, 235.

Phloeodes, 11.

Phormia azurea, 373.
chrysorrhea, 376.
metallica, 374, 379.

Phyllophaga, 12.

Pieris brassice, 193.
tape, 193.

Palpigus, 227.

Plath, O. E., article by, 373.

Platymetopius collaris, 232.
rubellus, 231.

Platymetopius hyalinus, early stages

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, Aquatic Adapta-

fay . tions of, 213.

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, 85.

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, So.
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. ; ba |
Trigonoscuta, 11. ‘Mae
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, a
Vanessa antiopa, 193. :

Walker, E. M., article by, 267.
Weiss, Harry B., article by, 369.
Welch, Paul S., article by, 213.
Wheeler, W. M., article by, 173, 174.
Williston, Dr. S , Resolutions on
the death of, 56.

Wing Structure of Lepidoptera, etc. 349,
Wubana, 252.

drassoides, 252, 253.

pacifica, 253.

retrahens, 253. 3

Zacotas, 3.

Zelotes ater, 247.
tuobus, 247.

Zopherus, 9.

Zorotypus, 270, 271.

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