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UNIVERSITY OF CALIFORNIA PUBLICATIONS
TECHNICAL BULLETINS
COLLEGE OF AGRICULTURE, AGRICULTURAL EXPERIMENT STATION

ENTOMOLOGY

Vol. 1, No. 1, pp. 1-152. September, 1906

CONTRIBUTIONS FROM THE ZOOLOGICAL LABORATORY OF THE
MUSEUM OF COMPARATIVE ZOOLOGY AT HARVARD COLLEGE,
UNDERSTHE, DIRECTION OF E. L. MARK-—No. Ist:

THE WING VEINS OF INSECTS.

BY

C. W. WOODWORTH.

INTRODUCTION.

The wings of insects are more extensively used in classifica-
tion than any other portion of the body. Since wing characters
are peculiarly conspicuous and tangible, it might be supposed
that taxonomists have chosen them for the identification of
groups to a larger extent than their relative importance
warrants, simply because they present easily recognized char-
acters. One has but to make comparison, however, of wing
characters with those based on other parts of the body to be
led irresistibly to the conclusion that they are of exceptionally
high value. It is not too much to say that of all structures
the wings have preserved the most nearly complete record of
the course of the phyletic history of insects. The confidence
with which wing characters are selected for the differentiation
of groups results, in large part, from the recognition of this
high phylogenetic significance, and the conviction that groups
so defined are natural.

Though this fact will doubtless be readily conceded by every
one, it seems strange that so little attention has been given to
the study of the wings by those who have attempted to investi-
gate the classification of the orders of insects from the stand-
point of their evolution. There is, however, a difficulty which
has stood, and still stands, in the way of the utilization of the
data, that may be a sufficient explanation of this failure. It
is the absence of a really comprehensive study of this compli-

2 University of California Publications. |ENtomMoLocy

cated subject. The complexity arises from the plasticity of
the organ upon which its value for systematic and phyletic
purpose depends,—a plasticity that permits the minute differ-
ences between species or varieties to record themselves, thus
making the older history so nearly illegible that one is hardly
to be criticised if he despair of deciphering it.

Aside from the color-pattern,—a matter scarcely ever of more
than generic value, and usually less,—the venation of the wing
includes practically every item that has been found useful in
classification based on that organ. We must turn, therefore,
to the study of venation for the evidence afforded by the wings
upon the questions of their origin and the relationship between
the larger groups of insects.

The important contributions that have been made to the
study of venation may be put into three quite distinct groups,
corresponding to the motive, or point of view, of the investi-
gator. By far the largest number of authorities on venation
are the systematists, or the students of single groups, who
have in most cases ignored similar work by fellow laborers in
other groups, the result being an almost unendurable confusion
of nomenclature. It should be said, however, that we owe to
them a most careful elaboration of the details and in almost
every case a trustworthy establishment of homologies within
the groups upon which they labored. We shall here mention
only some of the most conspicuous authors from the almost
endless list of those who have studied along this line. Among
those most worthy of mention, on account of the quality,
extent, or significance of their contributions, are De Selys
Longchamp (Odonata); Eaton (Ephemeride); Hagen, Wood-
Mason (Embide); Hagen, Kolbe, MacLaughlin, Spangberg
(Psocidee); Brunner von Wattenwyl (Orthoptera); Fieber,
Kolenati (Hemiptera); Heer, Kirby, Kempers (Coleoptera) ;
MacLaughlin (Phryganeeide); Low, Meigen, Schiner ( Diptera) ;
Forster, Schenk, Shuckard, Thompson (Hymenoptera); Com-
stock, Packard, and Spuler (Lepidoptera). The second group
are the paleontologists, three of whom have made very valu-
able and extensive studies on the venation of the wings of
recent insects, as a means of laying a foundation for their
work on fossil forms. They are Brongniart, Heer, and Scudder.

The most important studies of a general nature, besides
those of the authors just noted, are those dealing with the

Nahe!

Vou. 1.] Woodworth.—Wing Veins of Insects.

subject from the standpoint of comparative anatomy, the aim
being to establish a sound nomenclature. Among these are
Adolph, Amans, Brauer, Comstock, Needham, and Redten-
bacher.

The result of all this work is that there has been a more
exhaustive study of the wings than of any other part of the
insect’s anatomy. Thousands of species, representing every
group of any importance, have been illustrated. The details .
of the subject can be said to have been very fully worked out,
but on the theoretical side not much progress has been made.
Most of the theories that have been proposed have been of
limited application, relating generally to small matters of
special adaptations.

What most needs to be done is to develop a general theory
of venation that will serve in the interpretation of the facts
that have been so richly accumulated. The present work is
an attempt in this direction. The studies of the author have
been under way since 1884, and have included the study of
microscopical preparations of the wings of about two thousand
species representing all the principal groups, the examination
of a much larger series of insects with spread wings, and the
inspection of practically all the published figures of insect
wings. Representatives of most of the larger groups have
been studied in their earlier stages, both from simple micro-
scopical preparations of wingpads and from sections. It will
not be useful to attempt to present the mass of material thus
accumulated, as much of it would of necessity be but the
verification of work already well done by others, or the criti-
cism of relatively insignificant details.

The author desires to acknowledge the inspiration and
encouragement of Professors Burrill and Forbes of the Uni-
versity of Illinois, with whom his first work on this subject
was begun, and the invaluable assistance and advice of that
honored and lamented prince of entomologists, Dr. Hagen,
whose invariable kindness during the two years’ work in his
laboratory can never be forgotten; to Dr. Mark, with whom
the concluding work has been done, his indebtedness is fully
as great; to numerous entomologists, who have aided him with
material, and to his students and assistants, especially for the
making of preparations, his thanks are due; he regrets that
he can not mention them all by name.

4 University of California Publications, {ENTomoLocy

PART I.—WINGS.

ORIGIN OF WINGS.

The wings in insects that have simple metamorphosis very
evidently arise as outgrowths of the body wall of the thorax,
as has been well understood by all entomologists, and described
by even the oldest writers. Their development in the higher
orders, however, is much more obscure, though the presence of
the wing rudiments beneath the skin of the larve in the
higher insects has been known since the days of Swammerdam
(1737-38). The earlier authors quite naturally considered
these latter to be entirely different in structure and origin from
the wingpads of the lower insects. Weismann (66) was the
first to discover the connection between these internal wing
rudiments and the hypodermis of the body wall, but even he
believed this to be true only in certain insects (Nematocera).
Later, Kiinckel d’Herculais (75) showed that, in the groups
supposed by Weismann not to have the fundaments of the wings
connected with the hypodermis, there was, nevertheless, evi-
dence of such an origin. All subsequent authors who have
investigated the subject, with the exception of Landois (’74),
Ganin (’76), and Graber (’89), have accepted the idea of the
hypodermal origin; as no one of these three were able to dis-
cover any other source, all the positive evidence is to the effect
that the wings in all the groups, those with complex metamor-
phosis as well as the others, arise as a modification of the cells
of the hypodermis of the thorax.

From this ontogenetic fact the conclusion can be drawn with
much confidence that, in the primitive insects also, the wings
arise as outgrowths of previously undifferentiated portions of
the hypodermis of the thoracic segments. .To some extent,
especially in the lower insects, as will be shown hereafter, the
ontogeny does not exhibit a direct development toward the
imaginal wing. There is a great deal of evidence, as will also
be shown below, to the effect that the wing in its first develop-
ment followed likewise an indirect course; that is, in passing
from the first differentiation of the skin structure to that found
in a perfected wing, the cells assumed functions and developed

Vor. 1.] Woodworth.—Wing Veins of Insects. D

peculiarities quite different from those which they at first
possessed or finally acquired. From whichever point of view
we approach the subject, the evidence points clearly to the
truth of this contention.

On the grounds of the need of continuous utility in an organ.
that is being developed, we must conclude that before it served
as a wing it must have existed for some other purpose and
owed its structural peculiarities to other functional necessities,
since in order to have the slightest value for flight it must have
already attained a very considerable size and developed pecul-
iarities both in its own structure and in that of the segment
from which it arises. The size requirement has been admitted
by all since Gegenbaur (’70) put forth his theory of the origin
of wings, but the other requirements of the function of flight
have been quite overlooked.

Additional evidence in favor of the theory of the indirect
development of the wings may be had from a study of the
structure of existing and fossil wings. The great difference
that is evident between the structure of the wing membrane
and that of the body wall is nowhere bridged over by tran-
sitional conditions, even in degenerate and functionless wings.
It is only in cases where the wing loses in a measure its
flight function by taking upon itself other duties, as in the
case of elytra, halters, etc., that the histological structure
becomes approximately that of the body wall. The invariable
association of strikingly distinct structural peculiarities with
the function of flight and the absence of transitional conditions
indicates that the primitive wing must have arisen from an
organ distinct in function and structure from any of the types
of functional wings known in living insects or preserved to us
in the rocks. Some of the differentiations that characterize
wings arose while the wing exercised this other function.

Finally, the mechanics of the growth of the wing, as far as
we can judge, indicate that there are no influences that are
able to transform directly an unmodified epithelial cell of the
body wall into one capable of producing, for instance, a sec-
tion of a wing membrane, without passing through a certain
course of development by exposure to such conditions as are
presented in the wingpad of a nymph. This matter will be
discussed more fully when treating of the development of the
wing. According to this view the wingpad may be considered

6 University of California Publications, [= NTomoLocy

not only a preparation chamber for the wing, but also as repre-
senting a vestige of that organ within which the character of
the primitive wing is supposed to have been established.

The evidence as to the nature of this hypothetical precursor
of the wing has been but little discussed. The usually accepted
idea, that of Gegenbaur (’70), is that the wings are modified
‘tracheal gills. Plateau (’71) considers the wings as hypertro-
phied spiracles, but since he probably holds the same view as
to tracheal gills, his theory is not essentially different from the
former. Miiller (’75), Pancritius (’84), and Packard (98) are
the only writers who have combated this theory. Miiller,
from his study of the development of the wings in Calotermes,
concluded that they first arose in the same way as they appear
to develop in this young white-ant; that is, as lateral out-
growths of the dorsum. The chief ground for this conclusion
was the absence of the tracheze in the wing fundaments for
some time after these make their appearance. Miiller contended
that this disproved their having been derived from tracheal gills.

Pancritius follows Miiller, and adds the idea that the prim-
itive outgrowth of the body wall may have developed into a
protective body-covering like an elytron, and that this may
have been so modified as to become a wing.

Packard originally adopted the view of Gegenbaur, but in
his later work (Packard, ’98) accepts in its place the theory
of Miiller, and attempts to supplement it by outlining what
he considers to have been the probable course of development
of the wing. According to this modification of the Miillerian
hypothesis the primitive winged insect is supposed to have
possessed the power of leaping and had lateral extensions of
the thoracic segments, which acted as aéroplanes; later, in
some manner not explained, these a€roplanes became articu-
lated, were invaded by trachee, and finally, after the repression
of the organ in the earlier stages of the insect’s development,
gave rise to true imaginal flying organs.

The grounds upon which Gegenbaur’s theory is rejected by
Packard are:

First, that tracheal gills are produced in a variety of situa-
tions, and not in a definite place like the wings;

Secondly, that a gill is always supplied by a single trachea, a
wing by five or six;

Thirdly, that tracheal gills are known only in Neuroptera

Vor. 1.] Woodworth.—Wing Veins of Insects. 7
and Pseudoneuroptera, and are, therefore, supposed by him to
be secondary adaptations, as he seems to doubt that these
orders represent the primitive insects.

In answer to these arguments it may be admitted that the
tracheal gills are of many sorts, and are not necessarily
homologous structures in any strict sense. Gegenbaur did
not suggest that the wings arose from organs extending from
the end of the abdomen, or from those growing on the base of
the legs, but rather that there are ‘tracheal gills on the sides
of the abdominal segments, which by their position and struc-
ture suggest that other similar, perhaps homologous, structures
situated on the corresponding part of the thoracic segments
might have been modified into wings. The second argument
can be easily disposed of by citing the recent work of Com-
stock and Needham (’98-99), where it is shown that the wing
trachee all arise from the side of a single trunk, or at most
from two. The weight of any argument based on the arrange-
ment of the trachee is greatly diminished by the fact that, in
many wings at least, the tracheation is a comparatively late
and: entirely secondary matter. The final argument advanced
by Packard does not seem to have much force. Granting that
none of the groups now existing represents the one in which
wings first arose, and that tracheal gills as they now occur are
always adaptive and secondary, or even that this was true in
the primitive insects, it is hard to see any bearing the facts
would have on the question at issue.

It is not difficult to find weakness in the ideas advanced by
Packard. What, for instance, can be supposed to account for
the first development of broad lateral expansions on the sides
of the body in ancient Paleozoic times? Certainly these were
not developed as a means of protection from the kinds of
enemies that are supposed now to account for such structures.
Again, there is much more difficulty trying to imagine the
process of the conversion of a plate of this nature into a wing
than to imagine one produced from such a structure as a tracheal
gill; and further, there is no evidence that leaping insects
existed in those early times—certainly the remains of jumping
insects from the Paleozoic era are not abundant, and the
family groups possessing this power have never been recog-
nized as ancestral.

Pancritius’s suggestion is open to the same criticism as that

8 University of California Publications. [ENTomoLocy

just made. The paleontological evidence is all to the effect that
no true elytriferous insects occurred till after the Paleozoic
era. The cockroaches of the oldest strata, for instance, still
possess comparatively membranous fore-wings.

Miiller’s theory thus seems not to have been particularly
strengthened by these contributions, and really rests on the
original ontogenetic observations. Unless other facts can be
discovered that will give undoubted support to this theory, it
certainly has no sound foundation, for the well-known tendency
toward short cuts in ontogeny will easily account for the facts
observed by Miiller in regard to the tracheation of the young
wingpads in Calotermes.

Gegenbaur’s theory stands, therefore, not only as the gen-
erally accepted one, but as a theory against which no very
significant objection has been raised, nor has it an important
rival. There are grounds, however, more important than any
that have been raised, for objecting to this theory in its original
and usually accepted form. It may safely be said that the
histological conditions essential to the function of blood aéra-
tion are far different from such as would be necessary in the
construction of an organ of flight. A wing could not have
been directly produced from a tracheal gill. The changes that
must occur in the production of a wing are so great as to
require, as an intermediate stage, the modification of the gill
into an organ with a different function.

When an organ changes from one structure and function
to another it must be possible for the organ to serve for a
while the former function, but there must also be requirements
in the first function that will cause a development of the
organ to a stage where it can serve also in the new function.

In no tracheal gill not specialized somewhat for some other
than the primary function is there a sufficient approach either
in size, form, or structure toward an organ that could assume
in the slightest degree the function of flight. It seems neces-
sary for this reason to modify Gegenbaur’s theory, as has been
done by Lang (’88), so as to recognize the tracheal gill as one
of the two necessary steps before the production of a wing is a
possibility.

No one has yet attempted to consider the detail of wing
specialization, or to inquire into the process or method by which
it is brought about. A complete explanation requires that the

Vor. 1.] Woodworth.— Wing Veins of Insects. 3]

wing should be traced from the first fundaments of an out-
growth of the thoracic integument, through the modifications
of form and structure essential to the performance of its func-
tion in each stage of its evolution, into a fully developed organ
of flight.

The earliest insects of which we have any certain remains
had well-developed wings, and wings have in all probability
arisen but once among insects. All existing insects are, there-
fore, to be considered descendants of flying Paleozoic ancestors
and to have inherited their wings through all these ages. The
meagerness of all ancient geological records and the necessary
absence of all direct evidence in later time make the problem
one of great difficulty. The process would be almost unin-
telligible were it not that organs similar to those from which
the wings have in all probability arisen appear to be easily
acquired by insects; for this reason we have preserved to us
abundant data for our investigation.

The first step in the development of wings, in accordance
with this theory, is brought about as a result of a change of
habitat—an insect, formerly terrestrial, becoming aquatic. If
this occurred in a small insect, which was probably the case,
the only adjustment necessary would be a reduction of the
thickness and firmness of the cuticle. The insect was doubtless
one of those living in damp situations, such as serve at the pres-
ent time as the habitat of Thysanura. The situations in which
these insects may be found vary from subaquatic to almost
terrestrial, and the direct effect of the environment would be
sufficient in the primitive insects to accomplish the differentia-
tion of the older insects into two groups of individuals, not
sharply distinct, but looking toward the specialization that
later occurred: On the one hand, those that remained in the
more moist localities and retained a character of skin that made
possible the free absorption of oxygen from the water; and, on
the other hand, those whose surroundings resulted in the older
individuals quite losing this power, through the drying and
hardening of their cuticle, thus confining the adult individuals
to such situations as permitted the continuous breathing of
free air.

The danger from disease would in time render such a neutral
condition as we have imagined for the subaquatic forms quite
untenable and required a modification that would harden

10 University of California Publications, (ENToMoLocY

the skin in spite of the moisture when the insect retains, as
have the existing Thysanura, the old habitat. Another modi-
fication—one, indeed, that seems to be simpler and easier of
accomplishment-—is the abandonment of the breathing of free
air and dependence entirely on the water for the air needed.
How efficient the skin is as a breathing organ in water in
the case of small insects may be judged from the fact that at
the present day certain small insects, such as mayflies and
dragonflies in their first stages, appear to have no need of
gills, being in some cases quite devoid of them.

As soon as there is an increase in size, there comes to be the
necessity for a hardening of the body wall for muscle attach-
ment, and at the same time the demand for oxygen increases
much more rapidly than does the surface of the body,
requiring that the breathing function be localized and _per-
fected by the special modification of some part of the body
wall. This becomes absolutely essential to the life of the
insect whenever the other demands upon the integument have
reduced the unmodified oxygen-absorbing area to a point where
it is entirely inadequate to the demand. In the case of small
insects, as for instance the young of dragonflies and may-
flies, as stated above, this point has not been reached, but will
be in both these insects as soon as they become larger.

Any part of the body may be specialized for this purpose
either by the adaptation of an organ already existing or the
development of an entirely new structure. A gill in an insect
is simply any outgrowth of the body wall that retains the
soft texture originally possessed by the whole body surface.
When of this simple form they have been called blood gills,
because the distribution of the air absorbed is dependent upon
the general circulation of the blood. This organ becomes
much more efficient as soon as it is invaded by a tracheal twig,
for it then permits the more rapid transfer of the carbonic
acid to the water outside it, and the absorption of the oxygen
from the water through the thin layer of blood and the walls
of the tracheal twig and the gill itself. The exchange of
gases that results tends to approach a condition of equilibrium.
Such tracheal gills appear to be very efficient organs of respi-
ration.

The position of a tracheal gill probably depends upon the
condition of the insect at the time that the gills are specialized.

a ss eo) ee

Vor. 1.] Woodworth.—Wing Veins of Insects. ipl

That region which is most susceptible to the stimulation that
produces growth generally—that where growth last occurred—
is the region where the gills most readily form. We can con-
ceive that the bases of the appendages and the end of the
abdomen where gills most commonly occur are such regions of
easy stimulation. That the region of the thoracic segment at
which the wings are developed belongs to this same category
we can readily believe, on account of the frequency with which
another growth, the so-called margin, occurs in the corre-
sponding parts of adjacent segments, and in these segments in
very young stages.

The simplest form of tracheal gill is a tube-like process con-
taining a simple tracheal twig. The modifications consist of
the development of a brush-like bunch of such simple elements,
a tree-like branching of a simple element, or a leaf-like expan-
sion of the same; all intergradations exist between these types.
There is a practical limit to the length of a gill, chiefly depend-
ent upon the danger of injury in the case of a long organ;
these modifications are the result of an effort to increase the
surface without increasing the length.

Of these three forms of modifications the last is evidently
the most efficient, since the whole gill surface is exposed to
open water and none of it faces an adjacent surface with which
it must compete for the oxygen contained in the intervening
water, as is the case in the other types. . There are compensat-
ing disadvantages in the increased danger of injury, as the
individual portions can not give way as readily when coming
in contact with objects Hable to injure them. The necessity,
therefore, arises of devoting a portion of the surface to strength-
ening ribs or veins, and a greater or less thickening of the
whole upper surface, making it somewhat more resistant and,
therefore, less useful as a gill. The production of the wing
from a gill of this sort would result in the early loss of its
utility as a gill, so we can not look upon it as the precursor
of the wing, even though there may be a striking similarity in
general appearance.

Another specialization, one that affords a better transition
toward a wing, is seen in those insects in which a portion of
the gill is devoted to the function of protecting the remaining
parts, permitting them to retain their most efficient gill struc-
ture even under rather adverse conditions. Just such an organ

12 University of California Publications. [ENToMoLocy

may be seen in the gill cover of a species of mayfly common
in Illinois, Rithogena manifesta Eaton (Baetis debilis Walsh);
as shown in Fig. 1. This is not the most wing-like gill found
in the family, but will
show as well as any the
tendency of its specializa-
tion toward the structure
of a wing.

The shape, while not
exactly like any existing
wing, still in its general
outline suggests very
closely the form of wing
which several authors
have considered the prim-

FIG.1. A portion of the abdomen of a mayfly itive shape.
eam Chicos penta aloud cnawing) | ive Sormamcntag(oinat

the edge by spines and
hairs is so exceedingly wing-like that it might almost be
duplicated in detail by the wings of some of the lower Diptera.

The surface of this gill cover, while not entirely free from
the function of respiration, still has most of its area modified,
and in the apical region possesses an arrangement of surface
hairs not essentially different from that found on the wings
in many groups of insects.

The most convincing evidence of the relation of such an
organ as this gill cover with wings is to be seen in the vein-
like stiffening of the membrane. These structures possess all
the characteristics of veins. They have, moreover, a definite
relation with rows of special spines, such as so often oceur with
veins. The only matter characteristically different is the want
of any relation between these veins and the trachee of the gill
covers. The gill covers are quite well supplied with trachee,
but they do not in any instance correspond with the vein-like
structures in these organs. It is possible that we have here a
stage in the development of the wing preceding that in which
the trachese: become codrdinated with the veins.

Still another matter making these gill covers comparable
with wings is the nature of the articulation. The alar articu-
lation must have been produced from a structure of this sort.

A great many insects living in water have a very curious

VoL. 1.] Woodworth.—Wing Veins of Insects. 13

habit of vibrating the body, apparently to assist in respira-
tion by causing the water bathing the gills to mix with the
surrounding water and so become purer and fresher. When
an insect possesses gill covers, the same change may be effected
by the vibration of these. Any process attached to the body
as the gill covers are, would be caused to vibrate with every
movement of the segmental muscles and particularly by the
contraction of the dorso-ventrals. The gill cover that we have
been considering moves quite freely. The economy of this
means of keeping up the circulation of water is at once evident.
The musculature and the first steps toward the specialization
of the thorax for flight are thus provided for.

A difficulty that has presented itself to the minds of many
students of the subject of imaginal characters is that of account-
ing for the repression of an organ during larval life, or the hold-
ing of its development in abeyance until the final molt. The
precursor of the wing, according to any theory connecting it
with the tracheal gill, must have been functional in the larval
insect. Wings would, therefore, seem to offer one of the most
difficult of these problems; that is, the entire suppression of
all the earlier stages.

A closer study of this subject shows that this is only an
apparent difficulty, for there is no real suppression in the
larval stages in the case of the wings, since they often, and
perhaps generally, appear quite as early as the true gills, and
represent at first nearly the condition of gill covers. The
change into wings (that is, its development beyond the gill-
cover stage) is all that belongs to the penultimate stage.

Another kind of suppression has really occurred in the
case of insect wings, whereby the organs have become limited
to the thorax. Much of the difficulty in this case disappears,
however, when we note that the same problem has been met
in the case of gills that retain their original function. I refer
to the remarkable case of insects of the genus Cwnis, where
one pair of gill covers has become very like an elytron, is
almost or quite gill-less, and protects the naked gills of the
following four segments. On the other segments of the abdo-
men the gills have entirely disappeared. An exactly similar
process, but one resulting in two pairs of gill-less covers and
an entire suppression of all gillsin the last molt, would give us
the condition of the winged insect.

14 University of California Publications. |ENTomoLocy

Résumé.—Thus we see that the wings are hypodermal spe-
cializations of the meso- and metathorax, essentially similar in
all insects, and, according to the most probable theory of
origin, are of indirect development. The theory of Gegenbaur
that the wings came from tracheal gills, when compared with
that of Miiller, either in its original form or as modified by
Pancritius or Packard, is seen to be much nearer the facts, but
must be modified along lines similar to those suggested by Lang.
With this modified theory it is possible to trace the evolution
of a portion of the gill till there is produced an organ approach-
ing so closely to the structure of a wing that the transition
would not seem to present any very serious difficulties.

RELATION TO THE BODY.

There is a considerable difference of opinion as to the place
on the segment occupied by the wing. This is in part due to
apparent differences observable in the place and manner of
origin in different insects. In most of the lower insects the
wingpads appear to be processes of the notum of the meso- and
“metathorax, showing first at the extreme posterior angle as a
slight elongation. In later molts they extend till they com-
monly involve nearly or quite the whole length of the respec-
tive segments and inwardly along the hind edge sometimes a
third of their width. They appear, as just stated, to belong to
the notum, being usually in the same general plane of the
disk of the notum and so little different from the normal
structure that it is not possible, at least at first, to distinguish a
dividing line, while on the other hand the angle they make
with the side pieces sharply separates the wingpads and the
pleure almost from the first.

Exceptions to this statement are found (a) in the saltatorial
Orthoptera, where the wingpads, as described by Riley (76)
and Graber (’77), arise in the ordinary manner, apparently
from the notum, and suddenly at one of the molts reverse
their position and project upward instead of downward, thus
coming to be apparently processes of the pleura, but always
with a distinct line of demarkation separating them; (>) in
the Embiide, where, as pointed out by Wood-Mason (’83), the
wings have no connection with the posterior part of the seg-
ments to which they are attaehed, but come from the extreme
anterior portion; and (c) in the Odonata, where the wingpads

———

Vor. 1.] Woodworth.—Wing Veins of Insects. 15

from the beginning project dorsally, and thus appear to be
outgrowths of the pleure. Comstock and Needham (‘98—99)
remark that “they appear at a time when the tergum and
pleura are very little chitinized and are hardly more identified
with one than with the other.”” The wingpads in Odonata also
differ from those of most of the lower insects in that they
arise from the middle of the segments and only later extend
forward and backward across their whole width.

Among the higher insects the wings seem uniformly to
arise from the middle of their respective segments somewhat
dorsal of the legs, but at a time when there is no evident dif-
ferentiation of dorsum and pleura. They usually appear well

~

2

FIG. 2. Diagram illustrating the relation of the wing to the
dorso-pleural suture.

1. The commonly received view.

2. Comstock and Needham’s view.

3. Calvert’s view.

n, notum; p, pleura; w, wing; s, suture.

up toward the back and le in such a position that the tip is
directed downward and posteriorly, giving one quite the same,
impression as do most of the wingpads in the lower insects,
viz., that they are processes of the dorsum. Packard ('98)
goes still further in the case of the Hymenoptera by regarding
_the wing fundaments as scutal structures. No one else has
attempted to locate them so closely, but their derivation from
the notum has been the conception of nearly all those who
have expressed any opinion in the matter. Two very recent
authors, however, have given expression to other and entirely
distinct views.

Calvert (’93) says, while discussing the Odonata, that the
upper lamina is “tergal and the lower pleural,’ and presum-
ably maintains the same for other insects. No evidence is
cited in favor of this view. Comstock and Needham (’98—99)
incidentally give expression to another idea by describing the

16 University of California Publications. |ENToMoLocy

rudimentary wings as arising ‘“‘at the point where the suture
between the tergum and pleura later develops.” Stated in
other words, these three views are as follows: According to
Calvert’s view both articular membranes are independent of
the dorso-pleural suture; according to the common conception
the lower articular membrane is formed at this suture; while
the idea expressed by Comstock and Needham woud seem to
make the suture the source of both articular membranes.

There is the possibility that these differences in theory and
the apparent differences in fact are the result of actual differ-
ences in origin; that is, wings may not be in all cases strictly
homologous structures. But the degree of uniformity, or
rather the lack of distinctive characteristics for the separation
of wings into groups, indicates very strongly that we have to
do with a single monogenetic organ and the differences must
be considered secondary.

Morphological evidence, as far as I have been able to accu-
mulate it, has failed to yield any very decisive conclusions in
regard to this problem, but does concur with the results
arrived at from somewhat theoretical considerations regarding
the part played by the wing in the evolution of the thoracic
segment. The structure of insects has been quite elaborately
investigated from some standpoints, but the order and method
of specialization of the parts of the thorax have quite escaped
the consideration of investigators.

I will only briefly outline what appear to be the salient
points in the process and those that have a bearing on the
problem in hand. The accepted nomenclature, that of Audouin
(°24), who made the first comprehensive study of the structure
of the thorax, is based on the idea that a segment consists of
a number of separate pieces, called sclerites, grown together
into a more or less solid ring. Audouin recognized one sclerite
belonging to the ventral region, the sternum; two belonging
to each side, the episternum and epimeron; and four dorsal
pieces arranged in a series from before backward, the pre-
scutum, scutum, scutellum, and postscutellum. Besides these
were the legs and wings and their accessory basal pieces. The
tergal pieces, like the pleural, are separated from each other
by transverse lines at right angles to the axis of the body
and to the lines that divide the tergal pieces from the pleural,
and these latter from the sternum.

Vou. 1.] Woodworth.— Wing Veins of Insects. 17

The theory has been suggested by Hagen (’82) and others,
but I believe in most cases only tentatively, that a thoracic
segment, like the head, is a composite region of the body, and
owes its complexity, in part at least, to its possessing traces of
the primitive segments. The wing in this case might even be
considered a modified leg, as it is in birds. It need scarcely
be said that, in this form at least, the theory is certainly unten-
able. Very recently the idea of a double origin of the thoracic
segment has been seriously revived in a somewhat new form
by Walton (00). As the result of a study of the basal portion
of the leg, he conceives that the peculiar structures found there
are the result of the fusion of two successive legs, the posterior
one being rudimentary, but contributing a large portion of the
basal segment. The two pleural pieces, according to this view,
would represent the sides of the two primitive segments. Wal-
ton does not discuss the bearing of this theory on the problems
of the wing and of the notal structures. His idea in the matter,
however, seems to be that the wing belongs to the second prim-
itive segment, and the corresponding organ in the anterior
segment is represented in the mesothorax by the patagia in
Lepidoptera and the tegula in Hymenoptera. The dorsal divi-
sion between the two primitive segments would evidently be
that between scutum and scutellum. All the facts that have
been cited in favor of this theory can be explained fully as
satisfactorily without supposing the segment to have a double
origin. The greater part of what follows regarding the special-
ization of the thoracic somite would not be seriously affected
even if the double origin were demonstrable. We shall assume,
however, that each segment of the thorax is developed from a
single articular somite.

The key to the solution of the problem of the specialization
of the thoracic segment appears to be its skeletal function.
The skin of the insect is the sole place of attachment of the
muscles of the organs of locomotion. The thorax is specialized
for this function and correspondingly relieved of other duties.
The head contains the mouth and most of the sense organs,
and the abdomen, at least in the adult insect, performs most of
the vegetative and reproductive functions, and neither of these
regions shows skin structures that can be definitely identified
with the parts of the thoracic segment, except in a most

general way. Thus, the peculiarities of the thorax both in
2—V

18 University of California Publications. LENToMoLocY

structure and function, and the very evident dependence of the
function upon the region specialized, lend considerable pre-
sumptive evidence in favor of the idea here advanced, that the
specialization is in fact dependent on the function.

There are many facts: bearing on this problem, to which it
will not be necessary to refer in detail, by means of which the
course of the development of the segment may be traced from
the simplest type of structure, such as occurs in the lowest
articulata, or in the apodous larvee

Sel. Peet. of insects, or only slightly more com-
|| | LY plex in the abdomen of many adult
forms to the most complicated.

There are evidently two stages in

the development of the thoracic seg-

ment: first, the one embracing those

changes that follow and are depend-

Es. Em. ent on the development of the leg;
and, secondly, the one embracing

those which are quite as evidently

| to be assigned to the specialization

of the thorax for flight.
Before discussing the nature of
these modifications it will be useful
ee to note the nature of the suture sep-
ee och EEGs ee arating the thoracic sclerites. Con-
os von ternum fm cpimeroni trary to the usual conception, these
ca ee SC aa cuteliums sclerites are not to be considered as
separate pieces in the same sense
that the segments themselves are separate. The division of
the segments is brought about by a specialization of the cells
of the connecting membrane, whereby they do not take part
in the modification that occurs in the other cells of the skin
resulting ina harder and thicker cuticle, but become instead
capable of producing thinner and more flexible cuticle. The
purpose of this thinning is to afford a freer motion between
the harder parts. Another example of the same kind of
specialization is found in the separation of the dorsal from the
ventral portion of the abdominal segment, and it is developed
for a similar purpose, that is, for motion. There is consid-
erable modification for motion, and therefore of the segmental
type, at the base of both the legs and the wings, but it should
be clearly understood that the sutures between the thoracic

Vor. 1.) Woodworth.— Wing Veins of Insects. 19

sclerites have a different nature and have arisen to meet
different needs. The latter consist of linear invagination,
which are characterized by harder rather than softer cuticle
and result in the stiffening of the region in which they are
developed.

The first and most important result of the development and
perfection of the legs is the production of the sterno-pleural
suture. The space occupied by the basal joint of the leg greatly
weakens the segment at a critical point, but it is entirely com-
pensated for by the production of this suture and the stiffening
of the edges of the pedal foramen. This region, instead of
being weakest, thus becomes in fact the strongest and stiffest
part of the segment.

A second change following the development of the leg is the
stiffening of the parts of the segment that bear the strain of
the muscles of the legs. There is some variation in the arrange-
ment and working of the muscles at the base of the legs of
insects, but we shall consider only the more common, and
undoubtedly primitive, arrangement, which consists of three
principal muscles: one extending toward the venter and two
toward the sides, one of which is anterior and the other posterior.
The ventral muscle was necessarily short at first and attached
at an unsatisfactory angle, but gradually became accommodated
by an ingrowing process from the back of the middle line of the
venter, and the difficulty was thus wholly overcome. This
process, which is of various shapes and sizes, forms part of
the so-called internal skeleton; though of much physiological
importance, it affects only very slightly the external appearance.
The side muscles did’ not require such elaborate changes for
their attachment, but did require a stiffening of the body wall
of the side, which was more conspicuous externally. This was
accomplished in the simplest and most effective manner by the
stiffening of the edges and of a line down the middle of the
side between the muscles, thus furnishing on each side of each
muscle the stiff support needed. The sutures thus formed,
showing as ridges internally, conspicuously divide the side into
the two regions, the episternum and the epimeron, which are
distinct below, but not delimited above, where they extend
feebly over the back from side to side. This is the condition
of highest development reached until structures evidently
associated with flight or with some other special requirement
arise.

20 University of California Publications. [ENtomoLocy

The final steps in the perfection of the thoracic segment are
not very dissimilar to those just described. The most striking
part of this specialization is the separation of the notum from
the pleuree to make way for the attachments of the wing, which
results in the completion of the specialization of the pleure.
The thickening which forms the interpleural suture is still
further strengthened to serve at its upper end for the articu-
lation of the wing, and the upper edges of these sclerites
are not only cut off sharply from the parts above, but are also

thickened. This is
Meso-thorax Mela-thorax rf Aud Seg. required by the fact
that the shape of the
base of the wing is
such that the seg-
ment is nearly cut
in two for its attach-
ment. The notum
keeps its connection
with the pleure
solely by the narrow
strips at either end.

Hig: Heer eee een? Oe bere a eet
oe gtermo hetero ee use mau fee guriieiniae Comite
goneollunn Sea ont ore uaa Hue cnues thie Wind Of nenliiays
sue ee eon eerie cements 2 Giond ot pe nnemere

segments necessary
to adapt them to the musculature of the wings, as contrasted
with that for the muscles of the legs.

With the exception of the Odonata and Ephemerida, insects
have a large longitudinal dorsal muscle, the contraction of
which causes an arching up of the notum; since the latter is
attached to the inner end of the wing root, its arching produces
the downward stroke of the wing. To.provide for the attach-
ment of this muscle, the front edges of the meso- and the meta-
thorax and of the first abdominal segment are developed as
deep infoldings, almost making diaphragms across the body
when highly developed. These infoldings are produced by the
chitinization of the connecting membrane as it grows inwardly,
and forms two additional pieces, as seen from the outside,
uniting closely in most cases the backs of the adjacent seg-
ments. The accompanying diagram (Fig. 4) gives the appear-
ance of a segment according to this theory.

Vota] Woodworth.— Wing Veins of Insects. Bil

In the Odonata and Ephemeride there is no deep infolding
for muscle attachment, but the external appearance is not very
different, the additional pieces evidently resulting from the
need of a firmer attachment for the parts that vibrate with
the wings and in the perfection of the anterior and posterior
notal articulation. The prescutum of the mesothorax and
the back of the first abdominal segment are especially devel-
oped to strengthen the thorax, because the remainder of the
mesonotum and the whole of the metanotum are: loose and
beat with the wing.

As a résumé, we may recall that the position of the wing
on the segment, though subject to some variation, always
marks the division between dorsum and pleuron, but there
is difference of opinion as to the exact relation of these parts.
The theories of composite segments in the thorax seem to be
entirely untenable and would not throw light on the question,
if true. The real solution requires a study of the specialization
of the thoracic segment, and must include the consideration of
thoracic functions. Two distinct types of sclerite division may
be distinguished, one for articulation, the other for resistance;
both of these occur in the thorax, but the latter only causes the
separation of the principal sclerites. The first stage in thoracic
specialization follows the development of the legs, and results
in the separation of the sternum from the remainder of the
segment and the beginning of the division which ultimately
separates the episternum from the epimeron and the scutum
from the scutellum. The completion of this division, the
separation of the dorsal from the pleural region, and the
development of the preescutum and of the postscutellum, mark
the final stage, which is brought about by the development of
the wings, and the wings can not, therefore, be considered as
a product of either the pleuron or the notum, but rather as the
means of their differentiation phylogenetically as well as
ontogenetically.

FLIGHT.

The one fact in regard to which all authors who have written
on the subject of flight in insects are in agreement, is, that
insect flight is not comparable with the flight of birds. The
path over which the wing of an insect moves in flight is shaped
like the figure 8, contrasting sharply with the oval path of a
bird’s wing. Furthermore, it is to be noted that while the

22 University of California Publications. [ENTomoLocy

bird’s wing is different on the two sides, that of an insect pre-
sents the same kind of surface to the air in both phases of the
stroke, and finally, as pointed out by Amans (’85), the struc-
ture of the flying apparatus as a whole, as well as in every
detail of structure, is fundamentally different in the two groups.

The path of the wing in flight is evidently correlated with
the nature of the articulation at the base of the wing, but
whether the articular structure requires the wing to pass
through the air in the course indicated, or is simply flexible
enough to permit the air resistance to force the wing out of
what would otherwise be a simple to-and-fro motion, is still a
subject of controversy. This latter theory was first pro-
pounded by Marey (’69), and was the result of very careful
observation and experiment. The opponents of this view
include two of the three who have made a serious study of the
articulation of the wing. The third, Amans (785), does not
take sides with either party, but expresses himself as incom-
petent with only anatomical data to decide the matter.

The observations of both von Lendenfeldt (81) and Lowne
(90-95) were confined to single insects with rather complex
structure, and in the case of von Lendenfeldt not with the one
that was studied for hinge structure. While both of these
authors agree that the course of the wing is determined directly
and entirely by the muscles of flight and the articular struc-
tures at the base of the wing, they differ in a matter of fact of
fundamental importance. The point at issue is the direction
taken by the wing in passing over its characteristic path;
whether, for instance, the wing when passing over the part of
its course farthest above the body as seen from the side is
moving forward, as described by von Lendenfeldt, or backward,
in conformity with the views of Lowne and Marey. The
method employed by von Lendenfeldt, that of photography,
gives with perfect accuracy a series of positions which, by
attention to the effects of air pressure on the different parts of
the wing, should give unmistakable evidence as to the direc-
tion in which the wing is moving. We have no means, how-
ever, of determining, from the figures given, the correctness of
his interpretation of the photographs. In like manner, Marey’s
results, though doubtless clear to him, can not be correctly
interpreted by any one else, because he mentions neither the
direction of the movement of the cylinder nor the position of

Vou. 1.] Woodworth.—Wing Veins of Insects. 93

the head of the insect when making the record. If the insect
were heading toward the direction from which the cylinder
turns, which would be the natural thing to do, then Marey’s
records substantiate his theory. There is the possibility in
all this work that the insect may reverse the action of the
wings—which in some cases
at least it seems to have the
ability to do—and thus
make false records, though
this is not likely to happen
often, for reverse motion is
apparently always of short
duration. Lowne’s work is
based almost entirely on
the action of the wing in a
dead insect, and is open to
the criticism that there is
no good evidence that the
action studied is that which
actually occurs during
flight.

The accompanying fig-
ures (Fig. 5), based on the
curves elaborated by von
Lendenfeldt, illustrate the
motion of the wing of
Agrion in flight. As seen
from above, the path ap-
pears to be practically the
same, whichever course the
wing follows; from the

. FIG. 5. Diagram illustrating the flight
side, however, the route of of Agrion.

stig i ss 1 A. From above, the curved line showing the
the SUSE Us essentially path of the stigma. B. From the side, accord-

different. as is shown in i»gtotheory of von Lendenfeldt. C. From the
? ‘ side, according to theory of Marey.

the diagrams.

This insect, as well as the blowfly, the species studied by
Lowne, moves its wings much more nearly in a direction
parallel to the axis of the body than dorso-ventrally, but the
latter was doubtless the more primitive motion. The common
white butterfly, Pontia rape, moves it wings in a nearly direct
dorso-ventral path. In such an insect the mechanics of flight

24 University of California Publications, |[®NToMoLocy

may be illustrated by the diagramatic projection shown in
Fig. 6. In an insect viewed from the left side let AB repre-
sent the hinge, or axis, about which the wing swings at
each stroke, the direction of progression being from B toward A.
Let the line CD represent the mean inclination of all parts of
the wing during the up stroke, and C’D’ the same during the
down stroke. These lines CD and C’D’ converge behind,
because, as is well known, the more flexible posterior field of
the wing yields more to the resistance of the air than the
anterior margin and, therefore, lags behind in both upward
and downward strokes. Draw from any point O of the line
AB, perpendiculars OE
and OK’ to the lines CD
and C’D’; then the lines
OE and OH’ represent
the direction of the force
exerted by the wing dur-
ing the up and down
strokes respectively.
Perpendiculars to the
line AB from E and EH’
(FE and FE’) represent

c

c the amount of this force
FIG. 6. anes aaa mechanics which is neutralized by

AOR horizontal line; WOR axisofbadyiCD reason of the opposition
pendicular to AB; EG and FE/H, perpendiculars jy direction of the two

strokes, and 20OF the net
resultant force producing forward propulsion.

A more economical motion would be produced by increasing
the obliquity of the lines CD and C’D’ to the base line, AB;
but it would require at the same time a greater rapidity of
stroke to maintain the same rate of progression.

Such motion as that just described requires, in order to
accomplish horizontal flight, that the axis of vibration lie
somewhat obliquely to overcome the effect of gravity. In the
diagram let A’B’ represent the actual line of motion. Now,
drawing the lines EG and E’H perpendicular to A’B’, we have
the lifting power that opposes gravity represented by E’H
minus EG, and the propelling force OG minus OH. At both
the beginning and end of the stroke the two pairs of wings
oppose each other, and thus exert no lifting or depressing

Vor. 1.] Woodworth.—Wing Veins of Insects. 25

action, so that the ratio shown in the diagram between the
vertical and horizontal forces is entirely out of proportion, the
former being much exaggerated.

Besides the to-and-fro motion just discussed, there is also a
motion along the plane of the wing resulting in the figure 8
path already referred to. The width of the loops varies
greatly in different insects, and in the same insect the exact
course is dependent in part on the speed, but more on the
character of the stroke, which the insect seems to have the
power of varying more or less. This power is seen very strik-
ingly in those insects that poise themselves in the air, as do
most of the bees and such flies as Bombilidee and many of the
hawkmoths. '

Now, Marey has shown that a body constructed lke an
insect wing moving to and fro will be forced from its direct
course and caused by the resistance of the air to travel over
just such a path as an insect’s wing is known to follow. The
front part of the wing being the more rigid, the effect of the
air resistance will be such that the whole wing will become
twisted near its base, and if the wing attachment permit, this
resistance acting obliquely on the wing disk will also force the
tip of the wing forward during each stroke. At the end of the
stroke the elasticity of the articulation and of the wing mem-
brane will bring the wing back to its normal position. This
occurring at each stroke, both up and down. the result will be
the characteristic curve. In such an insect as the butterfly, air
pressure, acting as here described, appears to be sufficient to
account for the curve of motion. Marey, indeed, believes the
air pressure alone is sufficient to explain the curved path in
the case of all insects.

There is, however, the possibility that air pressure will pro-
duce just the opposite effect from that described above, provided
the hinge at the base is sufficiently rigid to prevent the forward
motion of the tip of the wing, and provided the membrane is
sufficiently lax to bag and take on the helix shape. This is
the condition which has been insisted on by Pettigrew (71),
and is illustrated in the beetles. The result of the “ bagging,”
in this case, is the pulling backward of the tip of the wing and
its release at the end of each stroke, allowing the wing to
straighten out and the tip to move forward by its own elasticity.
The curve thus produced would resemble that previously

26 Unversity of California Publications. [HNromoLocy

described, but its direction would be reversed. Undoubtedly
this tendency to bagging is a factor in the flight of all insects,
unless the wing is so stiff as not to bend appreciably during a
stroke; but in a beetle it becomes of high importance, if not,
indeed, the controlling factor. As far as I know, the question
of the direction in which the wing moves during the flight of a
beetle has not been investigated.

A second force which modifies the stroke of the wing is the
action of the wing muscles directly attached to the base of the
wing, of which quite a number have been described. By
means of these muscles the wing can be pulled forward and

A backward; moreover,
the angle which its plane
makes with the frontal
plane of the body can be
varied. These muscles
are evidently the means
by which the insect varies
the precise nature of
the stroke of the wing,

FIG. 7. Agrion, showing figure 8 path of wing and also doubtless in
as seen from above and from the side. most insects they are
ciHibD of complete teat. 70s peattion: ofrest with the means of placing the
ee wing into most of the
positions of rest assumed by this organ in different insects and

A

Bs) B

of bringing it again into position for action.

If the action of any one of these muscles should become
synchronized with either set of the muscles producing the
wing strokes, it would doubtless greatly modify the path of the
wing. The muscles which Amans denominates the preaxil-
laries or anteaxillaries pull the wing forward; the postaxillaries
pull it backward. With such a curve as occurs in Pieris a
preaxillary or an anteaxillary would augment the width of a
curve, and a postaxillary diminish or reverse it.

Supposing that such a synchronization occurred, it is prob-
able that in most cases there would not be an equal association
of axillary muscles with both systems of primary muscles, so
that the probable effect would be the exaggeration in the size
of one loop of the curve. An exaggeration of this kind, though
I can not state if for this cause, is exhibited by the posterior
loop in a side view of the curve in Agrion (Fig. 7).

——

Vor. 1.] Woodworth.—Wing Veins of Insects. 27

This same unsymmetrical path could result from air pressure,
if the angle of the wing were different in the two strokes, as
might be produced by peculiarities of the hinge structure, in
which different bearing joints are brought into operation in the
two strokes, the wing lying free between them when at rest.
Another way in which the character of the curve might be
influenced by the hinge structure is by the development of a
catch which should restrain the wing at one or both ends of its
course while the wing is traveling in one direction, and not
affect it on the return stroke. The sudden release of the wing
from this catch would result in an immediate change in the

FIG. 9. Diagram illustrat-
ing flight of Pontiarapx. A

FIG. 8. Diagram of the stroke of the and B, two positions of wing.
wing in the blowfly, according to Lowne. EFGH, a plane intersecting

F, point at which there is a sudden the wings at land J. K, path
change of direction. of motion.

direction of motion. In this way there would be produced just
such an angle as is given by Lowne at point F (see Fig. 8) in
the path of the tip of the wing in a blowfly, at which place,
according to his account, “the hammulus escapes from the
uncinate sclerite.’ This would produce an unsymmetrical
curve, unless an exactly similar structure were developed at
the opposite point. This is probably never accomplished, and
an influence of this kind on wing motion is of only secondary
importance, if indeed it ever occurs.

The three methods of modifying the stroke described above
may all be considered as possibly operating in particular
cases, but only one factor—air pressure—is invariably present.
A wing with a simple up and down stroke modified only by
the influence of air pressure may be considered as representing

28 University of California Publications. [ENTomMoLocy

the simplest and most primitive condition, and is to be seen in
a butterfly like Pontia. Fig. 9 indicates the nature of the wing
motion under these conditions. The angles at which the wings
cut the plane EFGH indicate the oblique position given the
wing by the air pressure, and explain its efficiency for forward
propulsion during both phases of the vibration.

If one watches the flight of this insect when, during its
passage, it is directly opposite him (Fig. 10), or when it is
coming directly toward him, he will notice that the body moves
up as the wing moves down, and vice versa. A still more
striking example of this bobbing motion may be seen in many
Lycenide. In the larger Papilionide and in the higher
Nymphalide the motion is commonly more steady, and in the

SS ie
s 4 2 ELD |

FIG, 10. Flight of Pontia rapx, showing up and down motion of body.

Hesperidew, where the wings move very rapidly, the body
vibration is difficult to distinguish at all, but is presumably
present. <A stroke of this kind is not as efficient as one in
which the wing moves more nearly in the plane of body
motion, because of the loss of power in the process of lifting
the body up and pushing it down again, which must occur
with an up and down stroke.

From this type of stroke to that exhibited by a fly or bee, or
that of a damsel-fly shown in Fig. 5, there are all stages of
transition. _ These may be found in a single family, the
hawkmoths. While the stroke of this type is more efficient it
is of necessity more rapid, for the wings cease to act as aéro-
planes and become paddles. <A single stroke of a broad wing
may carry an insect many times its own length, whereas, if the
stroke is nearly directly backward, the insect can not make much
more than its own length, even counting the movement due to
momentum, which acts, during the recoil of the wing, in keep-
ing up the progress. The backward stroke is the most efficient
propelling force in this type, and while this is being made the
tip of the wing must, to be effective, move backward consid-
erably faster than the body moves forward. The forward stroke

—s es

Vor. 1.] Woodworth.— Wing Veins of Insects. 29

aids the flight chiefly by lifting the body, and toa large extent
the wing simply feathers forward.

The oblique or direct stroke is accomplhshed by shifting the
dorsal articulation of the wing backward, or the ventral
bearing-point forward, or, in the case of the damsel-flies, by
the movement of the whole dorsal region of the meso- and
metathorax into a position almost at right angles to that orig-
inally held. The stroke is from above and behind forward and
downward, or vice versa. ;

In the majority of insects, especially in those in which
change to an oblique direction has not been carried to an
extreme, the course of the wing is probably that described by
Marey. Unfortunately, I can not speak with certainty in
regard to either of the insects (Odonata, blowfly) in which
the contrary motion has been claimed. In regard to them it
may be said that they are both evidently very highly special-
ized; both possess the habit of assuming a peculiar position,
not one of rest, just before beginning to fly. The damsel-
fly shows this particularly well. In the blowfly it is not as
evident as in some of the other Dipterous families, such as
Syrphide, Conopide, and many others. It is not impossible,
therefore, that these insects may pass over the course that
they have been described as following, though it involves a
remarkable change in the nature of the wing motion as com-
pared with that of other insects.

The presence of air sacs in great abundance in the bodies
of all flying insects has often been discussed, and they have
often been compared to the swimming bladder in fishes, the
explanation offered being that they enable the insect to change
its specific gravity. The uselessness of such a contrivance to
aid in flight is so evident that it is strange that text-books
have so long copied the suggestion. The only other explana-
tion of this function that has been offered is that of Packard,
who suggests that they are a means by which a larger supply
of oxygen is provided for the muscles during their very vio-
lent action in flight. This does not appear to be a_ sufficient
explanation, even though the air sacs may aid respiration
materially in the manner suggested. It is especially the dis-
tribution of the air sacs that counts against the idea that this
is the sole cause of their existence.

A possible use of the air sacs that may explain their arrange-

30 University of California Publications. |ENToMoLoGY

ment in some cases is that they may aid in adjusting the
center of gravity; the inflation of air sacs in one region caus-
ing the body fluids to occupy another. This reason, however,
can not account for their distribution.

What appears to be the most important function of the air
sacs is their utility in the recoil after the wing stroke. The
muscles of flight are all so attached that the body contents at
the end of each stroke are greatly compressed and the return
stroke is started as a recoil from this tense condition. The
elasticity of the body wall of the thorax produces this result
without the aid of the body contents, but not as well as with
it. If the body cavity is tense, as by the inflation of the air
sacs, the recoil becomes to a large extent pneumatic.

Résumé.—We have thus’ seen that the flight of insects is a
process not duplicated elsewhere in the animal series; it is
unique not only in the matter of the structure of the wings
and other flying mechanism, but likewise in the character of
the stroke. There is still need of investigations that shall
settle the question of the exact nature of the stroke in the
higher forms. The mechanics of flight in its simplest form is
not very complicated, but in more specialized types of wing
structure several factors contribute toward its complication.
Of the factors modifying the stroke the effect of the resistance of
the air against the surface of the wing is always in evidence,
causing the wing to take an oblique position during each
stroke. The stroke may be further influenced by the action
of the minor muscles at the wing-roots, either temporarily or
continuously by their synchronization with the primary mus-
cles. Peculiarities in the structure of the articulation may
also contribute to the modification of the stroke in many ways.
The slower, vertical stroke contrasts strongly with the more
steady and rapid oblique movement of the wings, and the
latter presents a number of details requiring further investiga-
tion. Finally, a possible relation of the air sacs with the
recoil at the beginning of the stroke is pointed out as the
most reasonable explanation of the development of these
structures.

ARTICULATION.

The articulation of the wings of insects has been carefully
studied by only three investigators: von Lendenfeldt (81),
who described with great detail the structure in one of the

a

Vou. 1.] Woodworth.—Wing Veins of Insects. 31

dragonflies; Amans (’85), who gives us the only comparative
study of the subject and covers all the principal groups; and
Lowne (790-95), who, confining himself to a single insect, the
blowfly, makes a notable contribution by his very complete
and painstaking account of the structure.

The investigation of Amans is the most nearly in line with
the question of articulation now under consideration. As a
result of his investigation he reached the conclusion that all
the parts of the entire articular mechanism are comparable,

FIG. 11. Wing articulation in @sechna. A, from above; B, from within; C, in section.
s, se, scutum; sel, scutellum; pscl, postscutellum; psc, prescutum; ips, inter-
pleural suture; md, muscle disks; msn, mesonotum; mtn, metanotum; w, wing;
wr, Wing-roots; aa, articular apophyses; sm, depressor muscle; em, elevator muscle;
hp, hinge piece; p, pleure. 1-7 veins.
piece by piece, throughout the whole series of insects; he
includes in this category of homologous structures not only
the hinge proper, but also all the veins that reach the base of
the wing. .

Upon this point there is room for a decided difference of
opinion, there being abundant reason for the view, first sug-
gested by Chabrier (’20-22) and adopted by Pettigrew (’71)
and Jousset de Bellesme (’79), that the Odonata present a
different type of structure from that found in the higher
insects.

A reéxamination of the structure of dschna, the insect
studied by both von Lendenfeldt and Amans, is highly desir-
able. Both of these authors recognize five veins as reaching

32 University of California Publications, (HNtomMoLocy

the base of the wing, von Lendenfeldt’s sixth ending, as it does,
on the fifth some distance from the wing-roots. There are
really seven veins (compare Fig. 11), as the first and fifth, on
account of their close proximity to the second and fourth,
were overlooked by both observers. This, however, does not
materially affect the questions regarding the articulation of
the wing.

The bases of the fore and hind wings are easily and strictly
comparable, so that the following description will apply to
either (see Fig. 11):

The first vein, which has rarely been distinguished from the
second, lies immediately on the front margin of the wing.
Except at the base of the wing, the first is distinctly ventrad
to the second, lying in close proximity to it throughout the
whole length and continuing as the marginal vein around the
‘outer and posterior edge of the wing.

The second vein forms at the base the anterior wing-root.
Before expanding into the root it is first narrowed obliquely
and then broadly constricted. The wing-root consists of a
black plate immediately attached to the vein, in front of that
a broader, green-colored expansion, and finally in front of this
a detached, somewhat clavate, dark-colored portion.

The third vein is very far ventrad of the adjacent ones. Its
outer extremity is at the nodus, and its basal termination is a
simple rounded enlargement free from any attachment. * Just
distal to the basal end, this vein is bound to the adjacent
veins by a triangular cross vein, the three veins being attached
to the angles of the triangle. The anterior ventral limb of
this triangular cross vein is somewhat enlarged and bounds
externally the anterior condyloid cavity of the wing-root.

The fourth vein is the strongest one in the wing. At the
base it is somewhat flattened and expanded vertically, form-
ing the front edge of the posterior wing-root.

The fifth vein is usually confused with the fourth, to which
it lies closely appressed. Distally it ends in the arculus. At
the basal end it expands into a large L-shaped portion consti-
tuting the bulk of the posterior wing-root. Between the limbs
of the L projects the green tuberosity of this root.

The sixth vein is deeply ventrad to the adjacent veins,
nearly as much so as the third. The basal end is abrupt and
slightly expanded, apparently for muscle attachment. It is

Vou. 1.] Woodworth.— Wing Veins of Insects. 33

connected by a feeble cross vein with the fifth, and by a
stronger enlarged one with the seventh. This appears to bear
the same relation to the posterior condyloid cavity that the
vein between the second and third does to the anterior. There
may be some connection between the base of this sixth vein
and the green tuberosity, or with the black corrugations
beneath it, but this is certainly not very evident.

The seventh vein ends at the base in a tracheoid structure -
that bends ventrad near the posterior end of the L-shaped
portion of the posterior wing-root, and then proceeds dorsad
and across the body to meet the corresponding structure from
the opposite wing, forming the posterior boundary of the
X-shaped portion of the notum.

_ The parts of the body that come in contact with the wings
are a little more difficult to compare in the two segments, but
in most features associated with the wings there will be no
difficulty in recognizing the homologies.

The closest attachment between the body and wing is where
the tissue about the suture between the side pieces is extended
upward into a process, which is reinforced by a triangular
sclerite bearing articular apophyses (Fig. 11, aa). These scle-
rites are evidently chitinizations of the articular membrane
above the two pleurites. Their processes are strengthened by
a system of rod-like, cuticular thickenings, as shown in Fig.
11, B. The lower lamina of the wing is continuous, as a soft
flexible membrane, with the ends of these pleural processes.

Dorsally the posterior wing-root (Fig. 12) is opposite the
X-shaped portion of the notum, which is evidently composed
of the scutellum and postscutellum, the hard parts of which
meet only at the middle and diverge laterally.with a rather
delicate membrane between. The postscutellum is for the
most part tubular; and it is tracheate and directly continuous,
as already described, with the end of the seventh vein. The
ends of the scutellum project rather deeply into the body and
approach closely to the deeper portion of the end of the hinge
piece lying between the anterior wing-root and the scutum. As
seen from within (Fig. 11, B) the end of the scutellum is found
to be continuous with the longitudinal support of the muscle
disk to which the elevator muscle is attached. The scutum
has a broad, rather smooth surface, which in the vicinity of
the wing dips deeply into the body and there expands into
muscle disk.

Oa)

34 University of California Publications, [ENToMoLocy

The action of the mechanism of the wing, as shown in Fig. 11,
does not resemble in the slightest degree that described and
figured by Graber (’77), but indicates an entirely distinct kind
of motion. The action of the elevators, which are the princi-
pal muscles of flight, results in pulling the whole dorsum of
the meso- and metathorax into the thoracic cavity, causing
the wings to assume a vertical position. When the tension is
_ released, the wing begins its descent by the expansion of the
compressed thoracic contents. Both of the strokes are at the
beginning somewhat in the nature of a recoil. The full depres-
sion of the wing is brought about by the action of the side
muscles of the thorax, which, by contracting the body cavity,
force the dorsum outward toward the back, elevating the inner
ends of the wing-roots.

Although von Lendenfeldt’s description of the structures is
quite full, it is very difficult to identify his parts. Amans
evidently experienced difficulty, and gave a new description,
much more simple and quite easy to follow. His figures,
however, possess more artistic beauty than structural accuracy.
Von Lendenfeldt’s figures are not better, for the parts that can
be identified are grossly inaccurate. It can be safely said that
the many small cuticular wrinkles and thickenings that gave
occasion to von Lendenfeldt’s errors, and the minute muscles
described by him and by Amans as attached at various posi-
tions in the base of the wings, are of only subordinate impor-
tance in flight.

According to the account given above, the articulation in
the case of the wing of a dragonfly is brought about by the
expansion of the bases of the veins into a broad wing-root
corrugated to correspond with the wing itself, and presenting
two broad, flat areas; these become on the under side condyloid
cavities fitting the ends of processes developed above the pleu-
rites in connection with the interpleural suture. The inner
ends of the wing-roots connect by a rather complicated thick-
ening of the membrane with the deeply infolded and internally
expanded edge of the scutum where the elevator muscle is
attached.

Let us now turn to other insects and see to what extent, if
at all, the articulations there found may be reduced to this
same type. One of the insects described and figured by Amans,
the common harvest fly, is quite favorable for study. One

Vor. 1.] Woodworth.— Wing Veins of Insects. 35

species, Cicada tibicen, is not greatly different from the Euro-
pean insect studied by him. In this insect the metathorax
has been very much reduced, the hind wings being scarcely
more than appendages to the anterior pair, and having very
feeble, if any, motion of their own. Only in the front pair do
we find the articular structures in a truly functional condition.
We therefore will not concern ourselves at this place with a
comparison of two segments, but confine our consideration to
the attachments of the anterior wings.

FIG. 12. Mesothorax of Cicada tibicen, showing wing attachment.
A, from the side. B, section.

dm, depressor muscle; em, elevator muscle; wr, wing-root;
awr, anterior wing-root; pwr, posterior wing- root.

At the outset we are confronted with a fact which can not
but have a powerful influence in shaping the structure at the
base of the wing. It is what is known as the flexion of the
wing, a motion by means of which that organ is made to he
along the side of the body when at rest. This flexion, even in
a small insect possessing a wing much constricted at base, can
not be sufficiently provided for by the simple elasticity of the
parts, because the bending must be nearly at right angles to
the path of vibration, and so across the breadth of the wing,
and because the flexed position must be that of rest. A
rearrangement of certain of the basal structures is thus
necessary.

The harvest fly will serve as a type of insects with flexible
wings (see Fig.59). In this insect there is a rather rigid wing-

36 University of California Publications. (ENToMoLocY

root set in a cavity of the body wall, to which the wing is
attached, and upon which it rotates during flexion. Eight
veins may be recognized in the basal attachment of the wing.
The first vein is the strongest one found in the wing. Its basal
end is abrupt and does not articulate with the wing-roots, but
continues toward the anterior root as a soft tubular sheath of
the distal portion of the extensor muscle. This vein is closely
appressed to the second, but is, through much of its course,
always distinct from it.

‘The second vein is about equally prominent on both sides of
the wing, except near the base, where it is entirely ventral.
Just before the base is reached the vein appears to split into
two branches, the anterior of which may be considered as a
cross vein binding this vein to the first. The posterior branch
is doubtless the lower portion of the posterior wing-root.

The third vein resembles closely the fourth vein in dragon-
flies in its shape and distal connections. Proximally it lies
almost directly over the base of the second vein and is contin-
uous with the posterior wing-root above; in the same manner
the second vein connects beneath. These two veins form the
hinge for the flexion of the wing.

The fourth vein is very closely approximated to the fifth,
except at the two extremities. Toward the basal end it
becomes abruptly feeble, but rapidly widens again, and from
some points of view appears double. The extreme base is
formed by a vertical process that articulates above with the
tip of the posterior wing-root.

The fifth vein is quite prominent, being dorsad of the fourth,
and in connection with the sixth vein expands at the base
into a broad, flat, elevated plate, that lies partly over the
posterior wing-root, especially when the wing is flexed.

The sixth vein, besides uniting with the fifth in the forma-
tion of the plate just described, comes into close relation with
the body by fitting into a groove along the side of the scutum
and into the catch formed at the end of the postscutellum; it
also is closely associated with the hind wing, forming the
catch that holds the latter expanded. A portion of the poste-
rior edge of this vein is black and fuses with the seventh.

The seventh vein lies close along the sixth, somewhat ven-
trad to it, and contributes to the formation of the catch for
the hind wing. The whole vein really belongs to the posterior

Vou. 1.] Woodworth.—Wing Veins of Insects. 37

fold of the wing and connects loosely at base with one of the
floating sclerites of that portion of the articular membrane.

The eighth vein is the tracheoid margin of the membrane
arising from about the middle of the seventh and continuing
at the base into the edge of the postscutellum.

The body is highly specialized; only the mesothorax con-
tains important muscles of flight, those of the metathorax
being nearly rudimentary. The hind wings have only the
feeblest motion of their own anel depend upon the front wing,
to which they are attached, for their effective action. The
wing-root is conspicuous, though relatively small and com-
pact. It is the only hard part connecting the dorsum and
pleuron, except at the extreme ends of the segment. The
exact nature of the joint may be seen in Fig. 12.

Contrasting this type of articulation with that seen in the
dragonfly, we notice most conspicuously the much greater
expansion of the notum. This corresponds with its greater
importance in flight in insects with this type of articulation,
and with the more complex and efficient union with the wing-
roots. Almost as striking is the relatively small size of these
roots, already alluded to. The separation of the notum from
the pleuron is nearly as complete as in the dragonfly, but the
Wing-roots occupy scarcely more than a fifth or a sixth of the
width of the articular opening, the remainder being closed by
soft, flexible skin.

Not only in these more conspicuous matters are the struc-
tures unlike, but when one examines the details, a difference
will, be noticeable in every item. The differences are so
numerous and so great that it is difficult to conceive either to
have been produced from the other; they point to a differen-
tiation at a time when the wing-root had not been specialized.
Starting with such an articulation as occurs in the base of a
gill cover, either form of wing-root could have been developed,
but only by growth along different lines.

In the evolution of a wing the wing-root must first arise as
the result of a thinning of the wing membrane, making the
organ everywhere very delicate except at the base, where there
must be greater strength. The further perfection of the wing-
root consists either in the thickening of the bases of all veins,
so as to produce a single solid piece, or in the thickening of
only those veins which come in direct contact with the bear-

38 University of California Publications. |ENTomoLocy

ing points of the primitive wing. The form of the connection
with the notum will be essentially different according to the
method followed. When the wing-root is developed from the
veins immediately adjacent to the bearing points, the structure
will admit of greater movement at right angles to the direction
of the stroke. The perfection of this type of articulation per-
mits the complete flexion of the wing and requires provision
for its expansion.

The attachment of a flexible wing usually shows three
strong veins, as in the insect just described and shown at the

FIG. 18. Diagram illustrating the flexion of a wing. AM, anterior
margin; PM, posterior margin; P, primary vein.

three black areas in Fig. 12. These three consist of the vein
I have called the primary and usually the one immediately in
front of or behind it, which at the base takes a position
directly above or below the primary, the two forming the
hinge for flexion, and in front of these a third vein, which is
broken in one or more places near the base by soft, elastic,
telescopic sections and is connected by a tendon to the exten-
sor muscle, within the thorax. Behind the hinge all the veins
stop short and are separated from the body by a flexible area,
in which there are usually developed a number of floating
articular sclerites very variable in shape, number, and position.
Their weakness and variability prove them to be of very second-
ary importance in flight, but they will doubtless show, when
more fully studied, a distinct and important relation to the

Vou. 1.] Woodworth.— Wing Veins of Insects. 39

character of the basal fold. The accompanying diagram
(Fig. 13) will show the characteristics of this form of articu-
lation.

There is considerable variability in the exact structure of the
wing-roots, probably corresponding with peculiarities in the
shape or stroke of the wing, and the same is true of the acces-
sory sclerites of the articular membrane. We will not attempt
to describe or discuss them, as they do not appear to affect
the venation of the wing. These two types of hinge structure
correspond with essential differences in venation, and, notwith-
standing their variability, exhibit nowhere transitional condi-
tions.

Résumé.—In conclusion, we may remark that there remains
abundant opportunity for study in the structures about the
base of the wing. An examination of the structure of the
articulation of the wing of a dragonfly has shown many points
of difference from the accounts and figures of previous inves-
tigators. It proves also that the former ideas of the mechanism
of the wings must be considerably modified. Likewise, a study
of the articulation of the wing of a harvest fly brings out sufh-
cient differences from the account of Amans to disprove his
contention that there is no essential difference in the hinge
structure of different insects. That there are two distinct
types is further shown to depend upon differences in the prob-
able course of the development of the wing-roots. Finally, it
is pointed out that this corresponds with important differences
in venation.

40 University of California Publications. [ENTOMOLOGY

PART II—VEINS AND VENATION.

VEIN STRUCTURE.

Before it is possible accurately to discuss the problems of
venation, it will be necessary to inquire into the nature of a
vein, in order to determine whether we have to, deal with a
single type of structure or with a number of fundamentally
distinet, though superficially similar, organs. The majority of
authors have accepted one or the other of these views, without,
however, discussing the question. Asa consequence they are
often inconsistent, and in most cases have evidently given the
subject very slight consideration. The points upon which
differences of opinion have been expressed are as follows:

1st. It has been repeatedly maintained that such structures
as occur toward the tip of the wing of a beetle are not veins,
because they possess no lumen.

2d. Most of the authors treating of the wings in Neuroptera,
Orthoptera, and, to a less extent, in other orders, have made a
clear distinction between veins and cross veins, not classing
them in the same category, chiefly because of the variableness
exhibited by the latter.

3d. A difference is often made between cross veins; certain
ones that are very constant in position are supposed to be
somewhat different in kind or origin from the ordinary and
more variable ones.

4th. In a few groups where long veins are numerous, most
of those that are variable are supposed to be different in kind
from those that are more constant.

5th. Very generally veins that arise from the base are con-
sidered as distinctly different from those without evident
basal connections.

6th. Veins appearing as branches of another vein are often
treated as of a different kind from those whose inner end is
free or connected by only a cross vein with principal vein.

7th. The vein that lies along the margin is often distin-
guished from the other veins, sometimes by evident difference in
structure, sometimes because of its position, especially when it

canine ae thee eile

Vor. 1.] Woodworth.— Wing Veins of Insects. 41
makes a complete circuit of the wing and has two basal
attachments, or finally because it often differs from the other
veins in having no trachew in the early stages of its develop-
ment.

8th. This last argument has been carried so far as the main-
taining of an entire difference of nature between all veins
developed about trachez and those without trachee.

9th. An entire difference of nature has been maintained for
veins occupying elevations or depressions of the wing membrane.

10th. A similar distinction has been made between veins
according to the proportion of substance contributed by the
upper and lower layers of the wing membrane.

1ith. Again, a row of hairs or spines, a fold of the wing
membrane, or even a disturbance of the pattern, have been
held to be potential veins, though showing no other evidence of
vein structure. —

12th. Finally, the true vein has been supposed to lie deeper
than the skin and to have no necessary connection with any
of the surface indications.

A typical vein is a tubular structure; the wall of which
differs from the wing membrane by its firmer texture, by more
or less evident peculiarities of pigmentation, both in its own
substance and in the adjacent parts of the membrane, by
peculiarities in the surface ornamentation, especially in the
shape and arrangement of the surface hairs, and by the pres-
ence in its lumen of body fluid, a tracheal trunk, a more or
less evident quantity of connective tissue, and occasionally a
nerve. Ontogenetically the vein is a region of arrested devel-
opment—a region in which cells are proliferated, but not
specialized. The cells of the vein merely remain more like
those of the body wall than do the membrane cells, and for
this:reason retain the power of chitin production unimpaired.
Not only is the chitin production similar to that of the general
body wall, but the pigmentation is also quite comparable. In
position the veins probably in every case represent wrinkles
in the embryonic wing membrane, but it is often difficult to
make this out. The lumen is simply a continuation of the
body cavity into the wingpad, brought about at first mainly
by the smaller size of the vein cells, as compared with the
membrane cells. The position of the vein at the angles corre-
sponding with the folds of the membrane and the crowding,
and consequent bowing out, aid in keeping the cavity open.

42 University of California Publications. [ENtomoLocy

Nearly all the structures that appear as veins in the higher
groups—Diptera, Hymenoptera, and Lepidoptera-—are veins
according to almost any criterion that may be used, and may
be taken as the standard for determining the soundness of the
criteria proposed. We will now consider in detail the signifi-
cance of the characters suggested for the recognition of veins.

The presence of the lumen has been a very common test
ever since the name came into general use. This is partic-
ularly true with English entomologists, who follow Westwood
in the belief that flight is only one of the functions of the
wing, and that another of great importance is the aération of
the blood, the veins being supposed to afford especial oppor-

B C

FIG. 14. Diagramatic section of a wing of Cicada near the base,
showing three principal veins in cross-section.

A, a vein belonging entirely to the lowermembrane. B, one largely
formed by upper membrane; and C, a vein about equally divided
between the two membranes.

tunity for its accomplishment. If this is true, the lumen is of
prime importance. No evidence, however, has ever . been
advanced to prove that the blood could receive any appre-
ciable amount of oxygen from the outside air through the
vein walls, even when the veins have large cavities. The
circulation of the blood through the veins is in most cases an
impossibility, because they are not continuous channels.

The supposition that the vein function is the aération of
the blood is not necessary in accounting for the shape of the
vein. There are two other explanations for the development
and maintenance of this shape. One of these is that the
presence of the blood in the wing prevents the drying of the
wing to such an extent as to make it brittle; a condition
which is quickly reached in dead insects. The other explana-
tion is of greater importance, namely, that a tubular form
combines the greatest strength with the least amount of
material.

Vor. 1.] Woodworth.— Wing Veins of Insects. 43

In this connection it should be recalled that a vein (Fig.
14) is produced in halves, one in each membrane, and that
the lumen is the result of their not fitting together closely,
they being arched in cross-section. Sometimes the curvature
exists in only one membrane, the other contributing merely a
flattened plate, which is sometimes only slightly or not at all
thickened. There is no essential difference between these one-
sided veins and symmetrical ones, as all transitions occur
from a case where a vein is shared equally between the two
lamina to one where, so far as any appreciable structure is
concerned, it belongs wholly to one membrane. This transi-
tion may sometimes occur in the course of a single vein.

The apical end of a vein, especially in Hymenoptera, is often
solid; that is, the lumen ends long before the tip of the vein
is reached, and there is no change in any of the other charac-
teristics of the vein. These peculiarities may be seen in Fig. 15.
The structure in the latter case is unquestionably identical
before and after reaching the point
where the lumen ceases, and for
some distance both lamina con-
tribute to the formation of the solid
vein; imally, whenithe wing appears * ie is portion ofthe Hina
tounvelye but, single layer, there 008 o oe ibeuewiuemeinsbe:
is still no change in structure, only a
gradual diminution in size, which begins long before the lumen
disappears, and is uniform and regular all along its course.
Furthermore, there is a great deal of variation within a species
regarding the point where the lumen ceases. In some species
the lumen may extend to the extreme end of the vein; in
others, it may not exist beyond the point where the vein leaves
the cross vein.

The evidence is thus sufficient to support the proposition
that the presence or absence of the lumen does not indicate
any essential difference in the nature of a vein, but signifies,
rather, different conditions of the same organ.

The presence or absence of a trachea in the vein has ap-
peared a matter of more significance, especially to German and
American entomologists, on account of the association of
the wings with tracheal gills. Not a few writers have sup-
posed the vein to be merely a specialized trachea. The only
basis for this idea is the tracheoid character of many veins.

44 University of California Publications. UZNTomMoLocy

This condition is seen in all wings in the delicate vein that
borders the posterior basal membrane and connects with the
edge of the postscutellum. It comes out sometimes very dis-
tinctly in parts of the vein where folds occur (Fig. 16). In
many cases a whole vein will become tracheoid, and occasion-
ally all the veins of a wing take on this character. Whether
a vein is tracheoid or not is a matter independent of its homol-
ogy, as the character may occur in wings of any group of insects
or in any particular vein.

Tracheoid thickenings occur in both membranes independ-
ently of each other, as may be most clearly seen when halves
of the vein fail to correspond exactly in position, as is not
uncommon in Psocide. In the accompanying figure (Fig. 17),

Qty 14
a

=

FIG. 17. Portion of Psocid
FIG. 16. Bulla at outer, lower cross vein in wing, showing nonconformity
Vespa, showing tracheoid markings. of halves.

where, to prevent confusion, the tracheoid structure of only
one lamina is indicated, the tracheoid ridges of the two do not
correspond. The shape of the two forks also shows that in
this particular case the half veins themselves could not have
been made to superpose.

In their ontogenetic origin the tracheoid markings of veins
are due to excessive chitinization of the outer edges of the
transverse folds of the layer of cells that in the pupa produce
the veins. The markings consist of thickenings of the cuticle
which project on the convex side of the vein. It is these ridges
that, in the wings of crickets, are made use of in the formation
of the stridulating organ. They are developed in the same
position as is the hairy or scaly clothing of the veins, and are
related to the spines which sometimes replace the hairs on the
wings of insects. The wing-clasp of the front wings of the
species of Psocus illustrated in Fig. 18 shows a very interest-

Vou. 1.] Woodworth.—Wing Veins of Insects. 45

ing modification of the tracheoid ridges. Here isa vein that is
conspicuously tracheoid on the lower side of the wing, but free
from these structures on the upper side. The tracheoid struc-
ture is entirely normal along most of the wing, but toward its
tip the ridges begin to project as spine-like processes from the
outer margin of the wing.
These processes are successively
longer and longer, till finally
those nearest the end curve
over and form the clasp that
serves to hold the two wings
together. At the region of the
greatest development of these
spines the vein itself almost
disappears, all the cuticular
substance appearing to be ap-
propriated for the production
of the modified spines.

Tracheoid structures can not, ie is Gino e wae ss
in view of these facts, be con- ee relation to) tracheoid
nected with the trachea, nor are
they criteria for the recognition or classification of veins, but
simply methods of specialization, and might be produced at
any point when the appropriate conditions occur. What
these conditions are can at present only be guessed at, but
we may have a clue to them in the common presence of
tracheoid growths in bulle. These appear to be simply col-
lapsed portions of veins whose cells have failed to produce
sufficient chitin. It is conceivable that such cells would have
a lower osmotic pressure and therefore a greater tendency to
cling together, so that relatively less chitin would be deposited
in the depths of the transverse wrinkles of the developing wing
in this region. The facts that weak veins exhibit a greater
tendency to develop tracheoid characteristics, and that in a
single vein regions which are suddenly narrowed often show
the same tendency, agree well with this hypothesis.

To return to the question of the true tracheation of veins:
we have to deal with a matter that goes back to the early
stages of the development of a wing. The chitinization, which
is the most evident thing about a vein, is produced only just
previous to the last molt; but the vein cavity is evident long

46 University of California Publications. [ENtoMoLocy

before this time and is usually already occupied by a trachea.
In some insects the vein cavities are clearly distinguishable
for a considerable period before the trachew enter them, while
in others the trachee can be recognized before the vein cavities
appear. In these cases it is impossible to deny that the loca-
tion of the veins may have been really marked out, though
unrecognizable to the eye; but if so they were really estab-
lished before the entrance of the trachez, as in other insects.
It is probable that the tracheze enter the cavity between the
lamine and assume their fixed relation to the structure of the
walls, though they later occupy vein cavities.

It is the contention of those who insist upon a close relation-
ship between the veins and trachee that the presence of the
trachez causes the change in the cells close to it which results
in the production of the vein. In the cases where the trachez
are not present, or only later enter these cavities, the supposi-
tion is that the veins, while originally requiring the presence
of a trachea, have reached a condition in which they no
longer thus depend on trachee. . How such adjustment can be
brought about has never been explained, and until it is, the
facts must be considered as throwing a great deal of doubt
upon this explanation.

There is reason to believe that the presence of a trachea pro-
duces a different environment for the neighboring cells; but
the fact is, that there is no appreciable difference in the struc-
ture or appearance between veins which enclose trachez and
others in the same wing which are devoid of them, and
that there is likewise no difference between the regions of the
membrane which are traversed by tracheee and those which
are distant from them; all goes to show that the influence of the
tracheze extends far enough so that.all parts of the wing are
in practically the same condition in this respect.

A trachea always takes a more or less sinuous course through
a vein, never conforms to the short, abrupt turns the veins
often make, and does not exhibit any evidence of closer rela-
tionship to the vein than that which exists between the tubu-
lar antenna or leg and its tracheal trunks.

Evidence has already been cited discrediting the idea that
the wing performs the functions of a respiratory organ; more-
over, the course pointed out as probably that of the phylo-
genetic development of the wing—which gives the vein an

Vou. 1.] Woodworth.— Wing Veins of Insects. 47

origin in the precursor of the wing entirely distinct from the
trachere--suggests the idea that no essential relation exists, or
ever has existed, between these structures.

This view is further supported by the well-known fact that
there is considerable variation in the tracheation within a group,
certain veins being in some cases with or in others without
trachez, though otherwise not different, and particular veins
being served from entirely different trachee. These variations
occur in the lower groups, as well as in the higher; in the
former at least, a definite and constant relationship should be
maintained.

A large amount of very strong evidence would be needed to
explain away the essential identity of structure in tracheated
and non-tracheated veins; the evidence obtainable seems all to
indicate identity rather than difference. We must conclude,
then, that the presence or absence of trachez is an incident
of structure of no special significance in comparing veins.

Other evident differences in the structure of veins and in the
degree of constancy in structure or position within the same
species or small group, where homology is easily traced, have
given occasion for the idea that there are two different kinds
of veins. But the line separating these two sorts is placed dif-
ferently by different authorities, in the ways already pointed
out. The structural differences in addition to those already
discussed in this section are mostly matters of size and pig-
mentation.

In reference to the size of veins, it isonly necessary to point
out that while, in particular cases, veins can be readily grouped
into two classes on the basis of relative size, still one need ‘go
but a short way in the study of related forms to find that this
character is of no significance. Longitudinal and cross veins
are the ones in which differences of this kind most commonly
occur. In most cases no decided difference can be made out,
- and often where cross veins have become parallel with the
costa they are so similar as to be easily confounded with true
longitudinal veins.

The pigmentation of veins is a very characteristic and rather
complicated matter. We must recognize at least three topo-
graphically distinct regions of pigmentation as occurring on
veins. The pigments of these three regions are very uniform,
all being brownish or black oxidization products in the cuticle,

48 University of California Publications. [ENTomoLocy

like those in that of the body wall. A difference of intensity
can generally be recognized, often bringing these three regions
into very sharp contrast one with the other. Pigment may be
absent from any one of these regions, but some pigment is
always present, unless it be in the case of very small wings in
which the veins are presumably degenerating.

The first region of pigmentation is the one usually seen as a
uniform tint over the vein surface. It is often more feeble
than in the other regions, but sometimes persists when that of
the others is not recognizable at all. This may be seen most
evidently in veins without lumina. Sometimes when the other
regions are well pigmented, it is evident only locally, as illus-

NS
FIG. 19. Portion of wing of ; :
Apis, Showing different kinds FIG. 20. Bulla in
of pigmentation at bulla. wing of a wasp.

trated in Fig.19. A differentiation of the surface pigmentation
may be seen in the case of tracheoid veins where the surface
elevations are very much more densely colored than the inter-
vening hollows, thus bringing out the structure of the vein
very plainly. This may be seen in Fig. 16.

The second and third regions mark the border of the vein
and are not always readily distinguishable from each other.
The former lies within the substance of the vein, while the
latter belongs to the wing membrane. Sometimes one is more
prominent, sometimes the other. Differences come out most
conspicuously in the bulla. The second region either stops
abruptly or narrows into a sharp, but pale line marking the
edge of the lumen, while the third shows a tendency to spread
out and follow the fold, as seen in Fig. 19, or narrows into a
line similar to that of the second region, as may be seen in
Fig. 20.

et ete

tet

Vou. 1.] Woodworth.— Wing Veins of Insects. 49

The fact that there are three kinds of pigmentation and
that these may occur in almost any combination as regards
their relative distinctness, affords abundant opportunity for
apparent differences in structure. One can commonly trace
on a single wing, sometimes in a single vein, a large number
of variations in pigmentation. Interesting examples of this
are very common in the wings of the higher flies and of beetles.

A very careful study of the pigmentation of the veins sup-
posed by different authors to represent different types of veins
has failed to reveal a single constant item of difference in any
case when a sufficiently large number of wings were studied.
The pigmentation of the same vein differs in many cases
according to the size of the wing, indicating that perhaps the
size of the vein cavity may have something to do with the
amount or character of pigmentation. However this may be,
it is safe to say that there is no evidence that the pigmenta-
tion indicates in any case differences of structure by which
types of veins may be differentiated.

There still remains for consideration that class of so-called
veins which are represented on the wing either by the arrange-
ment of the surface hairs, by the disturbance of the color pat-
tern, or by folds in the membrane.

Hairs usually occupy a very definite position in relation to
veins. Usually the surface over the vein is either devoid of
hairs, or possesses one or several rows of them regularly
arranged, often of a larger size than those on the membrane,
and sometimes of a peculiar structure. Whenever there are
on the surface of the membrane rows of hairs similar in struc-
ture and arrangement to those on the veins, their position is
always such that if a vein were to replace them, it would not
be considered anomalous in position. The very common pres-
ence of hairs on veins, except such as have been just described,
gives very good grounds for thinking that they could not be
produced except over cavities like the vein cavities, and there-
fore that they represent veins either suppressed or not yet
perfected.

The pattern on a wing, whether due to coloration of the
membrane, or, as in Lepidoptera, to the presence of colored
scales, often shows a very distinct irregularity where a vein
crosses a spot or band. This is conspicuously evident in the
bands along the outer margin of the wing in most Lepidoptera.

1Ny

50 University of California Publications. UENToMoLoGyY

Now, in the area in front of the second posterior vein, although
there is no vein traversing it, the band is interrupted, in many
cases at least, in just such a way as it would be if a vein were
present. In this particular case students are pretty well
agreed that this really represents a lacking vein.

Folds undoubtedly have a close relation to veins. If com-
parisons be made between twa closely allied insects in the
wing of one of which there is one less vein than in the other,
this lacking vein, except when near the margin, is always found
to be represented by a fold that occupies the same position as
the vein. On the other hand, there are many folds in the
membrane of the wings of insects that are never represented
anywhere by veins.

Each of these three modifications of the membrane is thus
seen to indicate relationship with veins. The question arises,
Do they represent veins that have disappeared, or are they
places where veins are in the course of development? The
fact that the lower insects commonly have more veins than
the higher has led many observers to the conclusion that
these modifications represent lost veins. In some instances
this is very evidently the case. A broader view of the matter
would be the conception that there is between every two veins
a possible vein-path, or more than one, and that these may
be indicated by either of the peculiarities we have just been
considering, and that there is no essential difference between
the indications of a forming vein and of a suppressed one.
This path partakes of many of the characteristics of veins, so
that the calling forth of a new vein is not a process that must
result in an entirely new structure, but there is at hand a
region almost ready to take on that kind of development.
Likewise, the suppression of a vein involves merely the lower-
ing a little of the conditions that increase the tendency toward
vein development. These paths of potential veins exist in all
possible conditions from nothing to a complete vein. The
suppression of a vein may be supposed to involve the reduc-
tion of the adjacent vein-paths to the minimum, and their
replacement by a single path that represents the destroyed
vein; the formation of a new vein to involve the development
of two new vein-paths.

Résumé.—The many diverse conceptions of the structure of
veins, which have been based on the assumption that other

—_-7

Vor. 1.] Woodworth.—Wing Veins of Insects. 51

vein-like structures exist in the wing which have nothing to
do with veins, prove, upon analysis, to be untenable. The
typical vein structure is connected by a complete series of
intermediate forms with all the other vein-like structures in
the wing.. The lumen is merely the space left between the
semi-cylindrical halves, and disappears if these two elements
remain flat. Tracheoid structures are simply modifications
of one or both of the layers and are exceedingly variable.
Trachee likewise are found to be variable enough to discredit
their importance as means of defining veins. Neither size nor
pigmentation affords any better method of distinguishing
foreign elements in a venation. On the other hand, the less
tangible influences, such as are expressed in folds, hairs, and
color pattern, do conform to and belong to the venation scheme.

VEIN DEVELOPMENT.

While many investigators have studied the development of
the wings of insects, most of the principal groups having
received attention, practically nothing has been given, even
incidentally, upon vein development. Doubtless the reason
for this is the fact that most of these investigators have been
concerned with the question of the origin of the wing, the
method of its expansion after molting, the clothing scales, or
the tracheation; all matters much more conspicuous than vein
development. While there is nothing peculiar enough to have
attracted students to the study of the formation of veins, still,
for the purpose of the present study, the matter becomes of
more than usual interest.

The development of the wing, in most of the lower insects,
begins by a slight modification of the “margin” of the meso-
and metathorax, though sometimes, as in Odonata, where there
is no “margin,” by a minute tubercle; in the higher groups, by
a much more complicated process involving an invagination of
the tegmentary epithelium and a subsequent evagination of
the organ like that which occurs in other imaginal disk-
producing appendages. In the lower groups of insects the cells
of the wingpad are not distinguishable in any respect from
other hypodermal cells in similar situations, such as those in
the narrow fold at the posterior edge of most segments. In
the invaginated wings the cells are very different from those of
the body wall, but not distinguishable by any histological

52 University of California Publications, (ENToMoLocY

peculiarities from those of other similar invaginations, as for
instance, those of the legs. The wing disks, as a whole, are
from the first different in shape from other invaginations, and
thus distinguishable from the leg disks; later the arrangement
of the different sorts of cells that compose them is characteristic,
and ultimately they are distinguishable by the structure itself
of some of the cells; but this is not true of the wing funda-
ments when first developed.

All wing fundaments are alike to the extent that they are
sac-like organs and that the mouth of the sac opens into the
general body cavity. Being thus open the sacs always contain
blood and are accessible to ingrowths of trachez and connect-
ive tissue; the latter forms in many cases an investment of
the deep ends of the cells which constitute the wing mem-
branes, and is generally conspicuous when the wing is young.
This investment has been noticed by several authors in widely
different groups, and may occur in all cases. Ultimately this
connective tissue may partially or wholly disappear, or may in
other cases increase in amount and contribute not a little to
the stiffening of the wing. An extreme case of this kind may
be seen in the front wing of Belostoma, where it extends as a
strong layer across the whole breadth of the outer third of
the wing and is thickened at the vein so that if the whole
cuticle is stripped off (after loosening it in caustic potash) a
complete venation and membrane still remain. In this case
the cuticle over the vein differs from that elsewhere only
slightly if at all, so that it would be difficult to make out the
venation from the cuticle if it were taken off and examined
alone. The chitinization of the connective tissue here is more
complete than that of the cuticle itself, so that it is harder and
more resistant to the action of alkalies.

The presence of nerves associated with tactile hairs or other
sense organs has often been noted in company with the veins.
They need not be treated of in this place, as they are not uni-
formly present in veins, and certainly are not a necessary or
important component of them.

The problematical “rods of Semper,’ that have been
repeatedly described in Lepidoptera, are only imaginal trachez
in the process of development and not essentially different
from any other trachez in insects’ wings.

In studying the development of veins we need only to take

Vox. 1.] Woodworth.—Wing Veins of Insects. 53

into consideration the constant features of a developing wing,
the hypodermis of the wingpad, and the cuticle that it secretes.

The first trace of the venation in a wingpad is a series of
slight separations between the two layers of hypodermis run-
ning lengthwise of the pad. Exactly when this arises is difh-
cult to make out with certainty; it may exist from the very
first, but the indication is so slight that one can not be sure of
its presence until the cavities have attained considerable size.
By this time the veins can be seen in surface view as well as
in sections. In surface view they appear as slightly paler
stripes with no definite boundaries. They are from their first
appearance in a position approximating that which they occupy
in the adult venation. The paleness of the bands is doubtless
due to the transparency of the blood in the vein cavity. The
vein cavity hardly becomes distinguishable before a difference
begins to appear between the cells of the vein region and those
pertaining to the membrane. The difference is chiefly one of
size, perhaps wholly that.

The early details of the development of the wing vary
greatly and in many particulars in different insects. An im-
portant difference that distinguishes the lower insects from the
higher is the fact that, in the former the wingpad takes part
in each ecdysis, whereas in the latter the wings part from an
old cuticle only at the molt by which the insect becomes adult.
These two processes of growth are not so fundamentally differ-
ent as they at first appear to be, because a complete transition
between the two types is afforded by a single order, the beetles,
indeed in one case by individuals of the same species.

In order to take part in the ecdyses, all the hypodermal
cells must retain the same power of secreting chitin until the
pupal stage is reached. Up to this time the smaller vein cells
are neither more nor less active in this respect than the larger
cells of the membrane. The differentiation of the vein and
membrane cells in the early stages is very incomplete, and
they remain thus similar almost without further change until
after the penultimate molt. In some cases the veins appear
as wrinkles in the cuticle of the wingpad, but in other cases the
wingpad remains smooth in all the stages. When the wrinkles
are present, we can consider the wing to have progressed in its
development to an ontogenetically later condition, but it ap-
pears to be in nowise a gainer by this precociousness.

54 University of California Publications. UENTomMoLocY

The cuticle of the wingpad is quite uniform with that of the
rest of the body, and is simply continuous, at the base of the
pad, with that of the thoracic wall, there being no articular
thickenings. When the wrinkles of the veins appear, one can
also see signs of the developing hinge structure of the wing.
In those insects in which the pupal wingpads are cemented to
the body, the cuticle of the inner (ventral) side is much thin-
ner than that of the outer (dorsal) side, but there are no
differentiations indicating the veins except wrinkles on the
dorsal side.

The increase in size at each molt is provided for by a series
of wrinkles of the layers of the hypodermis, in the same way,
and to much the same extent, as occurs in other parts of the
body.

Thus we see that up to the entrance upon the pupal condi-
tion the development of the wings had not been carried forward
any considerable distance. Within the pupal wingpad the
organ continues its development in a very leisurely way, coming
to its perfection only just before the insect is ready to emerge
as an adult. The wing just before the last molt is a striking
example of close packing. The veins, though composed of
thick cuticle, are stowed away so that they occupy only a small
fraction of their length when extended. This is accomplished
so exactly alike for both the dorsal and ventral faces of the
wing that when expanded the two correspond in every detail.

Since the veins are nearly all produced in two parts it is
necessary for their completion when in position after the final
expansion of the vein that they superpose with great accuracy.
This requires that the cells of each intervenous area shall
develop a cuticle of the same size in the two surfaces. This
coordination of the two membranes I believe to be brought
about by the direct contact of corresponding cells. The fact
that the deep ends of the cells composing the two hypodermal
layers are united has been figured and described for several
groups. In some groups this is supposed to be brought about
by the connective tissue, while in others the cells of the two
hypodermal layers are figured as actually touching each other.
From my own preparations I believe that both methods are to
be met with.

The size of the wing seems to determine to a considerable
extent which of the conditions shall be realized. The best

Vou. 1.] Woodworth.—Wing Veins of Insects. 5D

figures of the condition of the cells in the larger insects are
given by Mayer (96). The section shown in Fig. 21 illus-
trates the cell arrangement in one of the smaller leaf hoppers.
It is always difficult to make out exactly what the nature of
the union of the cells really is,
except in the smaller forms,
where there is no real connec-
tion, but only close contact.

Usually the correspondence of
the two parts of the vein 18 FIG. 21. Cross-section of wingpad
apparently exact,) but one ‘can of nymph of Pypkiocyhe comes, show
easily find abundant examples
of a failure to meet, such as is shown in Fig. 17. A ‘more
common condition is where there is a considerable differ-
ence in the relative size of the parts on the two sides of the
wing; the small part, occupying any position from side to
side of the larger section, though in this case the veins give
one the impression of perfect conformity, except when exam-
ined very carefully. The possible range of variation of this
sort, before the nonconformity would ordinarily be discovered,
is often more than half the diameter of the vein, and cursory
examination might even fail to distinguish a displacement
that is not distinctly more than the diameter of the vein.

The difficulty of the problem arises from the fact that the
wing is not developed flat, but wrinkled to a remarkable degree,
especially in the membrane areas, where, in addition to the
transverse wrinkles, found also on the veins, there is still
greater folding in a longitudinal direction, providing for lateral]
expansion. This difference in the character of the folding,
when it is developed just before chitinization, is the most evi-
dent difference between the vein and membrane regions of the
wingpad. Probably the wrinkling was at first irregular; that
is, with no definite arrangement, but more or less uniform over
the whole wing; with the development of veins, the greater
need of room for the thicker cuticle of the vein regions gradu-
ally led these regions to be folded in one plane only; the
membrane areas, on the contrary, retain the irregular method
of wrinkling, and to compensate for want of lateral wrinkling
in the vein areas, suffered an increased degree of wrinkling in
the lateral direction. With all the changes incident to this
readjustment, it is necessary that the cells of the two sides of

6 University of California Publications. |ENToMoLocy

the wingpad shall everywhere develop alike. This seems
possible only when the corresponding areas lie strictly opposite
each other.

It is not necessary that each individual cell be connected
with a corresponding cell on the other side, but only that
enough points of attachment exist to insure the general ‘cor-
respondence of the areas, and, in larger wings, of the principal
folds. Mayer has pointed out the utility of these connectives
at the time of the ecdysis, but no one has suggested their evi-
dent primary function, that of correlating the position of the
cells in the wing membranes.

The relation of the hairs and scales to these folds is very
evident, those of the membrane areas being at the peak of
somewhat conical elevations produced by the double folding
If the folds of the two sides of the wing exactly correspond, the
hairs or scales in the imago would exactly match. In some
insects this is true to a degree that is marvelous, scarcely a
single hair in the whole wing being without its mate, both
so perfectly placed that in focusing, the transition from one
set to the other may be easily overlooked. In other insects
there is no evidence of any relationship, showing that the
connection was of the most general kind. In Lepidoptera
there is usually a general conformity, but seldom a very strict
one. ;

On the veins there is never the precise opposition of hairs
or seales that is sometimes seen on the membrane. However,
when they occur in rows, there are generally in a given
distance the same number on the two sides. This condition
corresponds with the entire absence of connection of the two
sides in the vein region and to the equality in the size and
number of the folds upon which the straightness of the vein
depends.

The hairs or seales lie much closer together in the mem-
branous region of the wingpad than they do on the veins, for
the reason that the veins do not expand laterally at the time
of emergence, and this probably explains their differences in
structure, usually so evident. One of the functions of the
stronger hairs of the veins is doubtless to aid in the molting
process. Even in the case of Lepidoptera the vein scales,
though appearing much like the others, are very much stronger,
because of their thicker walls.

\

Vor. 1.] Woodworth.— Wing Veins of Insects. atl

es)

Thus far we have designated the corrugations of the surface
of the wingpads as wrinkles or folds. This should be explained
before we begin to consider the production of the cuticle, for
this structure more properly than the hypodermis might be
spoken of in these terms. To avoid confusion, therefore, we
must point out that when speaking of the cell layers reference
is had only to the condition of the outer surface of the cells.
The deep surface may or may not follow the course of the
outer surface, and at best does so only approximately.

The so-called foldings of the cellular layers is rarely or
never a folding in the true sense of the term, since it does not
involve the lower halves of the cell, but is brought about by a
rearrangement of the outer ends of the cells such that the
outer surface presents a series of ridges regularly alternating
with furrows. It occurs immediately before or at the time of
the beginning of chitin production. The chitin is a layer which
is so molded over ridge and groove as to be thrown into folds.
The regularity of the cell grooves would at least suggest that
they passively result from the real folding of a plastic layer of
some kind, its folds being caused by lateral pressure. The
cuticle itself when first formed, or rather the outer portion of it,
might possibly serve as such a layer.

This hypothetical layer might be supposed to meet with too
much resistance along the veins to allow the production of
longitudinal wrinkles in addition to the transverse ones. The
latter may be supposed to arise first, because a short wrinkle
across the wing is more easily formed than one lengthwise.
By this assumption may be explained the manner in which
such a complicated wrinkling as occurs in the membrane could
result in the production of a flat membrane after it is expanded.

The possible history of this hypothetical layer is that it is
produced as a uniform homogeneous layer over the surface of
the wingpad cells before they begin to show any wrinkling
whatever, and that it then swells by the absorption of water
or some other material originating in the cells beneath.
Thus arises the lateral pressure that throws the whole -layer
into folds. The cells separate along lines where the folds press
upon them, thus bringing about the grooving of the cellular
layer. The importance of the connection between the cells of
the two walls of the pad in maintaining uniform conditions has
already been dwelt upon. If the connection is intimate, the

58 University of California Publications, |ENTomMoLocy

two wrinkling membranes would find the points of resistance

practically identical on the two sides, and so might produce

sets of foldings sufficiently the counterpart of each other to

accomplish the codrdination of the two membrane areas (Fig.
22-53):

The amount of expansion necessary in our hypothetical

layer would not be as great as is usually estimated. From a

number of measurements of

wingpads and adult wings, it

appears that an increase of

from three to four diameters

} \\\ c would be sufficient. Even this

is very large as compared

) ) with the power of most sub-

3 hi stances to swell, but certainly

not impossible. Whatever the

cause of wrinkling may be, it

is without doubt the same all

FIG 22. Diagrams illustrating folding oyer the body of the insect.

of wings at time of development of cuti-
cle between two veins of T yphlocyba comes. The folding of the hy poder-

A, longitudinal section of vein; B, of
memDEANe C, tangential sectionof mem- yjjigs in the wingpads is differ-
. ent in degree, but not in
kind, from that which accompanies increase in size elsewhere.
Just before each molt the hypodermis secretes two substances
of peculiar nature. One is watery, or slightly mucilaginous,
and serves first to separate the old cuticle from the cells, pre-
liminary to the production of the new cuticle, and acts as a
lubricant during the process of molting. The other is chiti-
nous, but of a peculiar kind, clearly distinguishable from the
remainder of the cuticle. The nature of neither of these
substances is understood, and one of them may represent the
hypothetical layer.

The hypothesis thus offered as an explanation of the grooving
of the developing veins and wing membrane is, in brief:
(1) That there is secreted by the hypodermal cells shortly
previous to each molt a substance that by swelling throws
itself into folds, and where the hypodermal cells are in the
right condition insinuates itself between these cells; (2) That
there is produced at once a new cuticle, which follows down
into each of the grooves thus formed, but straightens out as soon

e°
k .
ay Rend jeg ri

airs sXd Wen Asin

as the insect emerges.

Vor. 1.] Woodworth.— Wing Veins of Insects. a9

The production of the cuticle on the vein areas presents no
very peculiar features. It proceeds at the same time and in
the same manner as the cuticle production on the membrane,
from which it differs simply in being thicker. There must be
differences in the chemical composition of the cuticle in different
regions, as is shown by the development of color after emergence,
but this is not evident to the eye at first. The cells are not
usually so completely exhausted by cuticle production as are
those of the membrane. They thus often form a considerable
layer around the inside of the expanded vein.

After the final molt all the folds of the cuticle straighten
out, the membranes of the two sides flatten against each
“other, and after drying are usually hardly distinguishable as
two layers. The veins, at first flat, soon become tubular and
harden in that shape.

Résumé.—According to this account the development of the
vein begins with cells not at all differentiated from those of
the body wall, but constituting a sac, opening into the body
cavity, which may or may not be associated with trachee,
connective tissue, etc. The first trace of a differentiation is
the formation of the vein cavities, caused chiefly by a slight
shortening of the cells of these regions. The differences
between the development of veins in higher and lower insects
are not important, as in all cases scarcely any progress is made
in- development until the penultimate stage is reached, in which
the two types closely resemble each other. Even in this stage
the veins are scarcely different from the membrane, except in
the matters already mentioned, and this condition persists until
near the last molt, when the grooving occurs. The veins fold
only transversely, while the membrane wrinkles in a very
complicated manner in both directions. An explanation of
the folding is attempted by assuming the secretion of a layer
that swells, and molds the outer cuticle-producing surface into
an appropriate shape, and thus provides for the ultimate
expansion of the wing into a flat organ. The cuticle produc-
tion and the expansion of the wing then proceed in the man-
ner that is normal for expanding regions in any part of the
body.

60 University of California Publications. [ENromotocy

GENESIS OF VENATION.

There are two chief groups into which the problems of the
origin of venation may be assembled. The first includes the
general questions of the formation of venation of the primitive
wing and the means and methods for the correlation, modifi-
cation, and adaptation of the veins that are the units compos-
ing it. The second group comprises the data bearing upon the
formation of the special types of venation found in the different
orders and families of insects.

As belonging to the category of general questions, we shall
first discuss the theories and conceptions upon which have
rested the schemes of venation that have been current. All
the earliest systems were based upon the very evident, indeed
unmistakable, homologies existing in closely allied forms.
Names were given to the veins according to the needs in
each particular group, in some cases without regard to the
nomenclature employed in other insects. The belief was quite
general that there was an essential unity in the venation of
all insects, and numerous attempts have been made to apply
the terms used in one group to the veins in other groups. The
great difficulty of tracing homologies in insects only distantly
related accounts for the tangle that has resulted. What these
early systems were, we shall not attempt to discuss until we
come to consider the modifications in the separate groups. For
the present purpose it will be sufficient to say that none of them
represents any particular theory further than the names
employed suggest, and that they were simply systems of nomen-
clature made for the sole purpose of rendering the wing char-
acters available for classification.

After the general acceptance of the doctrine of evolution, and
especially after the promulgation of the theory of Gegenbaur
as to the origin of the wing, the venation came to be generally
regarded as depending for its chief features upon the structure
of the organ from which the wing was supposed to have been
derived; the conception being, as has already been explained,
that the veins are produced around trachee, and that these,
and therefore the veins, simply reproduce the tracheal arrange-
ment inherited from the organ that was the precursor of the
wing.

If this view is correct, the surest and best means of study-

dS et

Vor. 1.] Woodworth.— Wing Veins of Insects. 61

ing the homologies exhibited by the venation is to make out
the tracheation of the wingpad. Comstock and Needham
(98-99) are the only ones who have attempted to put into
practice in any extensive manner this plan of studying vena-
tion. They found that in several groups the tracheation could
not in any manner be depended on for this purpose. In other
cases, however, including most of the lower orders, these au-
thors were convinced that the tracheation afforded the sound-
est criterion for the establishment of homologies.

The only other theory to account for venation that has been
elaborated and seriously defended is that of Adolph, as fol-
lows: This author does not attempt to explain the origin of
venation, but bases his explanation of the present condition on
the alternate arrangement of the veins in two series, one occu-
pying the crests, the other the depressions of a series of regu-
lar plications, best seen in the mayflies. He supposes that the
more dorsal veins, which he denominates the convex veins, are
different in origin, structure, and significance from the ventral
(his concave) veins, and that the orderly alternate arrange-
ment of these will aid in the recognition of obscure homologies.
According to this theory the wing represents the fusion of two
semi-venations alternating with each other. Adolph’s theory
does not appear to have gained many adherents, though it has
attracted a good deal of attention and not a little criticism—
first, by Brauer and Redtenbacher (88), and later by many
others. The idea has strongly influenced later students, even”
when they have rejected it as unsound. This is because it
directed their attention to facts that are very evident and, as
] believe, of unquestionable importance in the development of
the venation of many groups, though exactly what they signify
has never been clearly pointed out by any of my predecessors.
I shall presently endeavor to explain their meaning.

Still another theory has been suggested, but not elaborated,
in a recent work by Packard (’98). This latest idea is that
the original venation was determined by the mechanical
necessity of flight. Most authors have supposed that this
factor has been an important element in the specialization of
particular types of structure, but none had previously ascribed
any of the features held in common by all types of wings to
this cause. Packard does not explain any of the details of
the genesis of venation in accordance with this principle, and

62 University of California Publications, |ENtTomoLoey

has not been entirely able to divest himself of his former idea
of an essential connection between the trachesz and veins, even
after becoming a convert to the Miillerian theory of wing origin.

The mechanical theory of the origin of venation really
affords, I believe, an explanation of wing structure much more
reasonable and adequate than either of the other two suggested
solutions of the problem. In an organ adapted for such a
special function as flight, where the accurate adjustment of the
mechanical detail is of such supreme importance, it is very
natural to think that the structure upon which the strength
of the wing depends should be more profoundly affected by its
own functional requirements than by the accidents of the
structure of a temporary adaptive organ occurring in an
ancestor that lived in or before the Silurian epoch, particularly
since the detail of structure of that organ could have been of
no importance in the beginning, and can not have any onto-
genetic necessity at the present day.

The more rational conception is that there existed at the
beginning, and has existed through all time to the present day,
a mechanical necessity, in accordance with which the primitive
venation was produced and all its essential features have been
maintained through all the vicissitudes of the ages. This idea
accords entirely with what has been conjectured to be the
course of the development of the precursor of the wing.

In this ancient organ we may suppose that the general out-
line of the venation of insect wings had already been blocked
out, because that organ was called upon to meet similar needs.
Exactly how far the venation was specialized before the organ
became an organ of flight may never be known, and does not
particularly concern the present discussion. The terms wing
and wingpad will be used, in what follows, in the sense that
they include the precursor of the wing.

Another matter that can not be followed with any assurance
of chronological accuracy is the order of vein specialization,
because there are a number of elements in a venation that
are to a large extent independent of each other. In other
respects the course of the development of venation can be
traced very satisfactorily.

One of the earliest veins to appear is the marginal. Its
ontogeny can be imagined by supposing the wingpad to have
become somewhat more bag-like than usual after one of the

oo ee hes

Vou. 1.] Woodworth.— Wing Veins of Insects. 63

molts. Certain conditions of cell nutrition, probably those
that decreased osmotic pressure, would result in a closer union
of the adjacent surfaces of the pavement-like cells which would
interfere with a sharp bending of the wall in the region which
is to become the edge of the wing. As the wingpad collapsed
during the hardening of the cuticle after the molt, there would
therefore remain all around the edge of the wing an appre-
ciable cavity. As soon as a cavity is produced between the
two cell layers of the bag it acts as a blood sinus, putting the
cells immediately adjacent to it in nearly the same relation to
the food as the cells of the general body wall. This explains
their subsequent secretory activity at the time of the production
of the cuticle. —

The marginal vein is often not of uniform size around the
whole of the wing, being usually strongest on the front mar-
gin, often weak on the hind margin, and quite commonly
entirely absent on the outer (distal) edge of the wing. This
corresponds with the chances these parts have to secure nutri-
tion from the blood. If the channel is not freely open all
around the margin, so that all parts have a nearly equal
chance, then the parts farthest from the body are much handi-
capped, and as between the front and hind margins the former
has the advantage of taking its nutritive supply first.

It is conceivable that a vein arising in the way here described
might in part disappear and afterwards reappear in the history
of the group, corresponding to changes in body-form or order
of development; indeed, that it would be very susceptible to
such changes. This is in accord with the peculiarities of the
occurrence of the vein, for it not only appears in widely
divergent groups, but is subject to variations that are often of
only generic significance.

A vein of much more fundamental importance, one that is
invariably found in functional wings and usually the strongest
vein present, I denominate the primary. Its position is nor-
mally within the anterior third of the wing, but in a few
cases it may be farther back, as in the gill cover shown in
Fig. 1 (page 12), where the anterior margin is strongly devel-
oped and the primary takes a position somewhat behind the
middle for reasons of utility. Among true wings there is a
modification in certain Phasmide, where the primary vein
takes a posterior position, due to the exaggerated development
of the anterior region for another purpose than flight.

64 University of California Publications, [ENtomoLocy

The typical position is dependent on the same mechanical
necessity that determines the position of the bones in the
wings of a bird or bat, or the shaft in a feather. The ontogeny
of this vein is associated with the development of the inter-
pleural suture into an articular process; that is, the position
of the basal end is determined by this association. Probably
the course of the vein was originally determined by the fact
that the longest axis is the weakest line and most liable to
bend. The primary fold would thus be associated with the
primary vein.

The basal attachments of this vein, its primary significance
ontogenetically, and its functional importance in the wing, all
conspire to cause it to take and maintain its preéminence when
a complex venation is built up around it. No vein, unless it
be the marginal, is as easily identified as the primary, and °
there is none in reference to which there is such a perfect una-
nimity of views among students of venation in the matter of its
homology in the different. groups. The many names it has
received are simply due to differences in nomenclature, not to
differences in interpretation of homologies.

If there is any vein that can be homologized throughout the
whole class, it is the primary vein. This vein, then, must be
the starting point in the study of homology, as it is the oldest,
most important, least variable, and most easily recognizable
vein of the wing membrane.

The primary vein is usually conceived of as a branching
vein, though, of course, it must have been at first simple.
Usually there are more branches of the primary vein than of
any other vein in the wing. Sometimes they are given off
from both sides of the vein, more commonly from only one
side, which may be either in front or behind. The largest
number is seen in cockroaches; the simple unbranched con-
dition occurs in many groups. Between these two extremes
there are all intermediate conditions. Perhaps the commonest
number is four, making with the end of the vein itself five
ends, the number supposed by several recent authors to be the
typical condition.

Branches are given off usually at an angle of rather more
than sixty degrees, and proceed outward ordinarily in a more
or less evident curve, the concave side of which is toward the
tip of the vein from which they arise. They are produced at

Vou. 1.] Woodworth.— Wing Veins of Insects. 65

very regular intervals and lie parallel with each other, and
sometimes, when long, come to take a position almost parallel
with the primary vein.

For rather small wings, branches afford what is probably the
most satisfactory mechanical solution of the problem of sup-
porting the wing membrane. When the wing becomes larger,
however, the torsion on the principal vein would begin to be a
serious matter, so that other means of strengthening the wing
become necessary.

The ontogeny of the vein branches is explainable on the
supposition that the tendency toward multiplication instead
of specialization of the cells of the wingpad when carried too
far results in the production of transverse wrinkles, which
open up courses for the food material in the vein to find its
way outward. If this occurs before the specialization of the
membrane cells has gone too far, it may result in the extension
of the vein area in that direction. The course of a branch
produced in this manner would be governed by the same forces
that determined the course of the primary vein; that is, the
vein branch would follow the lines of least resistance. This
would be at first approximately at right angles to the primary
vein, and as soon as it gets beyond the influence of this vein
it would turn forward to reach the margin at approximately
right angles. Sometimes branches occur that leave the pri-
mary in the way indicated, then turn forward and run parallel
to it for a distance, and then turn again to meet the margin
squarely. Such a vein follows both of the lines of mechanical
weakness; that is, parallel to the vein and at right angles to
it. Of course, veins do not bend abruptly, and the tendency
is always to go straight, so there is always a compromise
effected, which reduces the angle of branching considerably
below ninety degrees.

An increase in the size of a wing usually results in an in-
crease in the number of veins. There are many notable
exceptions to the rule that the size and number of veins are
correlated, but these only .emphasize the general rule. In
many of the lower insects the tendency of a wing to develop
a vein wherever there is room is very evident. To this tend-
ency we may look for the explanation of many of the facts
relative to the perfection of the venation system of insect
wings.

o—V

66 University of California Publications, |ENTomoLocyY

Two sets of veins early to be developed constitute the series
which we may denominate the anteriors and the posteriors.
These are independent series, being developed on separate
sides of the primary vein, but they arose in exactly the same
manner. These are the veins that radiate from the base and
occupy the membrane not

arin: served by the branches of
a the primary. They would
be produced in a series one
after the other as the size
of the wing increased, and
may be numbered in the
order of their development,
which is from the primary
outward. The stages of
development are illus-
trated in the accompany-
ing diagrams. (Fig. 23.)
These anterior and _ pos-
terior veins, as they are
produced,will extend them-
selves across the membrane
at its weakest points and
thus divide the area more
or less equaily into two
parts. Probably the first
of these veins to be devel-
oped was the first posterior.
FIG. 23. Diagram illustrating proposed theory The time of the appearance

of the development of venations.

AM, anterior marginal vein; PM, posteriotY of the. next posterior in
marginal vein; P, primary vein; B, branches: .
1A, anterior vein; 1P,2P,3P, 4P, first, second, yeference to the first ante-
third and fourth posterior veins; 1, inde- 3
pendent veins; 12, 14, and fractions 4, }, 3, ete., yjor can not be decided, as

possible ways of designating independents.
the two series have no
necessary relation to each other. The anteriors and posteriors
may become branched, but as they become successively closer
together the probability of branching rapidly diminishes. The
first posterior is often branched, but seldom as much so as the
primary.
All the veins so far considered have arisen directly or indi-
rectly from the base of the wing. While they are hardly to be
designated as outgrowths of the cells of the body wall, they are

AM

Vor. 1.] Woodworth.—Wing Veins of Insects. 67

nevertheless produced from cells that have developed under
similar conditions. These conditions have progressed from the
base of the wing toward its tip, so that these veins may fairly
be regarded as outgrowing veins. They maintain their con-
nection with the body cavity in the adult condition in the
groups which, in this respect at least, we may consider to have
retained the primitive condition; that is, the Odonata and
Ephemerida. In all other existing groups of insects the base
of the wing has been profoundly modified by the development
of the structures that make the flexion of the wing possible.

The vein least affected by this specialization is the primary.
Its relative strength is probably the reason that all the other
veins of the wing gave way while the primary is merely made
flexible, becoming the axis of flexion. The changes in the
articulation of the wing thus brought about have already been
discussed, and it has been shown that the only close attach-
ment of the wing to the wing-roots is by the primary, though
it may be assisted by an adjacent vein. This fact has given
the primary vein still more importance in venation, but has
somewhat comphcated its study, because of the tendency of
other veins to attach themselves to this one. The extent to
which this occurs will be evident from a study of the modifi-
cations in the different groups.

Anterior to the primary vein the effect of flexion has been
most marked in the specialization of the base of the marginal
vein. This vein has been broken into a number of movable, hol-
low segments, separated from each other by areas of soft skin;
a tendon terminating in a muscle passes through these segments
to be attached to the distal elements of the series. Thus the
insect is enabled to pull the distal element toward the base of
the wing, the front edge of which is thereby drawn forward till
the organ is expanded ready for action. Insects usually pos-
sess but a single anterior vein, which loses its basal connection,
attaching itself to the primary and becoming weak except
when it takes part with the primary in the articulation. This
specialization of the marginal vein makes it necessary, if there
is to be any considerable expansion of the wing anteriorly,
that the membrane extend beyond the vein; that is, that the
vein no longer le along the margin. When this occurs and
the vein extends any considerable distance, the membrane so
produced is served by branches on the outside of the anterior

68 University of California Publications, |ENToMoLocy

marginal vein. This condition is not common; it is best seen
in the front wing of the cockroach.

Behind the primary vein all the veins except the marginal
lose their basal connections. The marginal vein is so delicate
at the extreme base that it is generally overlooked, and because
of its weakness it offered no obstacle to flexion, and so escaped
the fate of all posterior veins. Practically the whole difficulty
in homologizing veins arises from the difficulty in identifying
these posterior veins. How great these difficulties are, and
how completely we can follow these vagrant posterior veins,
will be shown as we discuss the different groups.

This brings us to the consideration of a class of veins that
never has had basal connections. These we may denominate
the independent veins. The term “independent” has already
been used in Lepidoptera for these veins, and has never been
used in any other sense. The independents are most uni-
formly present in the area immediately behind the primary
vein, and have here been designated by Comstock and others
as the “media.” Exactly similar veins may occur between
any two veins and develop in the same manner. These veins
may be designated by the vein immediately anterior to them.

The independents arise from the margin of the wing, and
might possibly be considered as ingrowing branches of the
outer portion of the marginal vein; but, since this portion of
the marginal vein is commonly absent, while the independents
are almost always present, this conclusion may seem to be
unwarranted. In insects rich in independent veins, these are
produced wherever there is a sufficient space between two
veins, and they extend up the wing as far as the space remains
sufficiently wide. The order of their development in any
space (compare Fig. 23) is, first, the production of a vein
approximately half way between the two veins bounding the
area; secondly, one on either side of the first independent
vein, and then one in each of the four interspaces thus pro-
duced. This process might go on indefinitely, but generally
three or four ranks of veins are all that can be clearly recog-
nized, except in Ephemeride, where this system is most highly
developed.

The relation of independent veins to branches is such that
it has given rise to confusion in the study of homology. In
the most typical condition the independent veins are strongest

Vor. 1.] Woodworth.— Wing Veins of Insects. 69

at the margin and become weaker as they proceed inwardly
(proximally), the ending being similar to that of the outgrow-
ing veins, but in the opposite direction. In some cases, however,
they have gained an attachment to an outgrowing vein through
a cross vein, becoming so completely united to it that they
have all the characteristics of branches. There is nothing in
the structural peculiarities to determine whether a particular
vein arose as a branch or as an independent vein; the only
criterion being a comparison with the venation of other mem-
bers of the same group, the principle being to class as an inde-
pendent vein anything that anywhere exhibits structures
characteristic of independent veins. It is not certain that
branches in the strictest sense may not, under some circum-
stances, especially that of their relation to a cross vein, take on
some of these characteristics. Such appears to be the belief
of several recent students of venation, who, having adopted a
typical number of branches as belonging to the vein we have
called the primary, include among them in many instances
veins which show very evident signs of their independent
nature. I am convinced that there is no reason why, in any
group, branches might not be decreased or increased in this way
to any extent, and that the application of the typical-number
hypothesis can not be depended on as conclusive evidence of
the nature of a supposed branch. At the same time, while there
is a theoretical typical number of independents in any area—
that is, one, three, seven, fifteen, etc.—a very little inspection
of venations will convince one that the typical number is
rarely attained. The reason is that an independent vein only
approximates the middle of an interspace, so that a vein in
the next series will commonly be quicker to develop on one
side than on the other, and this difference may become more
and more exaggerated as time goes on. In some groups the
evidence may be so plain that there can be but little doubt as
to the source of the vein appearing as a branch; but whether
we can homologize and clearly distinguish between true
branches and independent veins throughout the whole class,
is open to doubt.

A matter which somewhat complicates the question of inde-
pendent veins is that specialization of the wing membrane
which results in what have been called convex and concave
veins. There has been no satisfactory explanation of the cause

70 University of California Publications. |ENToMoLoGY

of these bendings of the membrane, beyond the bare suggestion
that this condition is useful in stiffening the membrane. The
explanation. of Adolph, that they represent fundamentally
distinct elements, has been shown to be without foundation.
The method of development of these convex and concave veins
appears to be as follows: If one examines an insect wing, such
as that of a butterfly, he will notice that the wing membrane
between the veins passes from vein to vein in a gentle curve
the concavity of which is dorsal. The reason for this may be
that, since the hardest stroke is downward, a convexity in this
direction is useful to stiffen the wing. , One can readily see that
such a curvature would have this effect, since the flattening of
each arched area by the resistance of the air would force the
veins apart and thus render the wing more tense. If between
a marginal vein and the primary an intermediate vein were
developed in such a concavity it would be a “concave” vein.
After a vein was produced in such a concavity and became
strong the membrane would no longer make a single curve, but
the portions of the membrane on the two sides of the new vein,
would become independent of each other and respond to the
influences that cause the curving of the membrane in the first
instance, both portions becoming concave above, so that upon
the development of new veins it would in time become a ‘‘ con-
vex” one. According to this theory, then, a concave vein is
merely a young vein, and a convex vein an older one. Thus
if there was but a single independent in an interspace it would
be coneave; and if there were three, the center one, which was
the first one developed, would be convex and the other two, one
on either side, concave.

In cases where the veins are very numerous and their alter-
nate convexity and concavity is very pronounced, this rule is
varied in such a manner that no well-established vein changes
its position either to concave or convex on the preduction of
intermediate veins. The membrane in such a case has the
form of a compound or S-shaped curve (Fig. 24, C7), being in
part concave, in part convex. In this membrane the inde-
pendent vein starts, as elsewhere, with first a single vein and
then a smaller one on either side. The middle vein (Fig. 24
B4 or 5) and the most precocious of the two lateral veins
(C 6 or 8) soon adjust themselves respectively to either the
highest and lowest (C 4 and 6) or to the lowest and highest

Vor. 1.] Woodworth.— Wing Veins of Insects. 71

(C 5 and 8) lines along the membrane and soon greatly out-
strip the third vein (C 7 or 9), which may then become a
middle vein in the next developing set. In groups possessing
this type of vein structure, as the mayflies, the independents are
thus developed in pairs, one convex and one concaye, the latter
being the one next to the convex vein in the interspace.

The cross veins resemble the independents in some particu-
lars, but differ from-all other veins in that they have no
determinable sequence in their first

production. It is probable that they PO A
were developed over all parts of the ee ,
wing at the same time. Like the j 3
independents, they probably corre- ONE pase B
spond to wrinkles in the membrane, A

but not to wrinkles parallel to the A é
. . . ~ . J
principal veins. Cross veins do not Rea: ae
: : = \ “8
always correspond in structure with ¢ *
longitudinal veins, though at times FIG. 24, Diagram illustrating the
5 a ee 5 7 : development of independent
indistinguishable from them. Like veins without disturbing
‘4 mal d 1 ¢ position of older veins.
the inc epen ents, t ACY are often A,B, C, three successive stages;
" . tot] j ‘ . . 1, 2,3, older veins; 4,5, those first
not clearly distinguishable from inde pale ea eieten eae
beamchecy lhe usual: practice: 1s to Hees: of whlel@ and: sremain

feeble.

class a cross vein as a branch, if in

any species of the group the vein in question exhibits a position
resembling that of a branch. This interpretation is probably
in most cases correct, though in considering the relation between
independent veins and branches the opposite method is pur-
sued; that is, if there is any evidence of the vein in question
being unattached, the interpretation is usually that it is an
independent vein.

In most wings with few cross veins one may see the whole
membrane, if held in the right hght, wrinkled in a very regular
manner, suggesting a regular system of cross veins. In order
to transform a simple venation into a netted one, it is only
necessary to assume that the tendency that brought about the
wrinkling corresponding to the longitudinal veins was carried
a step farther, so as to open passageways for the blood from
vein to vein along the course of the transverse wrinkles and
then, in the same way that we have supposed the branches
developed—or for that matter the principal veins them-
selves—the resulting change in the nutrition of the cells

72 University of California Publications, [ENtoMoLocy

bordering these passages would secure a greater chitin-produc-
ing power.

The evidence of such simultaneous production of cross veins
all over the wing is that in all the oldest venations these veins
are most inconstant in both number and position, none seem-
ing to have any preéminence or to show any significant pecul-
iarities. There is, however, a difference between those near
the body and those near the edge of the. wing, the latter being
much closer together, and correspondingly more delicate. In
all these points the cross veins resemble the wrinkles, and
prove that the cause of their production must have been some-
thing that influenced the wing as a whole. If the cross veins
had different origin, or method, or time of development, we
should not expect such a degree of uniformity. Among higher
insects certain of the cross veins exhibit characteristics that
mark them off as distinct from the ordinary cross vein, but in
most of these cases the number of cross veins has certainly
been reduced from a condition in which they were more abun-
dant. These special cross veins, then, are probably later
specializations.

By way of réswmé, we may say that, in place of the earlier
arbitrary schemes of venation, or of those that were based on
the assumption that the veins represent the remains of an
ancient tracheal system, or of the ingenious but unsupported
theory of Adolph, we would substitute a new theory of vena-
tion—a mechanical theory. According to this theory the
utility of the veins in flight is the prime factor determining
their number, position, and character. Several distinct classes
of veins are recognizable, not structurally, but on account of
their place or manner of origin. The marginal vein, which
arose by a modification of the cells at the bend in the wing-
pad, is always present next to the base of the wing on both
margins, but often disappears along the outer edge of the wing,
and occasionally may come to lie at a distance from the edge.
The primary vein is clearly the most important vein in the
wing, because of its relation to the pleural articulation and
later to flexion. In this development the branches, like the
primary itself or any other vein, follow the lines of least
resistance in the membrane, and have very definite relations
to the trunk vein. The anterior and the posterior veins are
developed to meet the increase of the wing surface, and approx-

Vor. 1.] Woodworth.—Wing Veins of Insects. 73

imately bisect the areas in which they develop. The basal
attachment of these veins has been destroyed in wings that are
flexed, and this often gives rise to much uncertainty as regards
homologies in distant groups. Veins of another series arise
independently between the tips of these and grow toward the
base, often becoming attached and thus simulating branches.
Veins are shown to have a very definite relation to the plane
of the membrane, some being above (dorsal), others below
(ventral) that plane. Thus we have the so-called convex and
concave veins; when the alternation of these veins is well estab-
lished the independent veins are produced in pairs between
them. Somewhat allied to the independents in the method of
their formation are the cross veins, which are believed, how-
ever, to have developed simultaneously over the whole wing
surface and later to have undergone specialization, accompa-
nied by a decrease in number.

METHODS OF MODIFICATION.

Having traced the development of these systems of veins
that give character and stability to the wing, there now
remains the necessity of following the organ along the lines
of modification which have resulted in the production of the
various types of venation that exist in the different groups, of
insects.

In these diverse specializations, while each has followed a
course in certain respects peculiar to itself, they have all been
controlled by common limitations. Some of the more important
of these common methods of modification should be considered
before proceeding to the detailed review of the special types of
venation.

A factor constantly influencing venation in an insect wing
is the coordination of the front and hind pairs. Whatever the
solution arrived at in any particular case, the venation can be
confidently expected to offer evidences of the reaction of these
organs upon each other.

The original condition was doubtless one in which the
wings were entirely independent of each other, such as exists
to-day in some of the dragonflies. The only requirement in
these cases is that the wings shall not interfere with each other.
As a consequence these insects have long and slender wings,
and the most conspicuous difference to be observed in their

74 University of California Publications. |ENTomoLocy

venation is the arrested development of the veins of the poste-
rior region of the front wing as compared with the hind wing
when the latter has become somewhat expanded.

The coordination of the two wings permits more variation in
shape than in non-codrdinated wings. We find, therefore, in
these wings, much greater range of variation in venation. Coor-
dinated wings are not usually of the same size. When the front
wing is the larger, it is always longer than the hind wing; but
when the hind wing exceeds in area the front wing, the differ-
ence is wholly in their greater width, except in certain beetles
and cockroaches. In making this generalization we are con-
sidering only cases where both wings are functional as organs
of flight.

The simplest method of coordination is produced by the
development of the adjacent edges so that they overlap each
other, the hind wing always lying beneath the front wing.
The other method consists in the specialization of structures
on both wings, by which these organs are held firmly together.
When wings are bound together they act as a single wing, and
the mechanical needs of the different areas have but little in
common between corresponding parts of the two wings. This
gives ample reason for the differences that have been brought
about.

The rule is that, if there is any difference in the number of
veins in the anterior or posterior areas, the lesser number will
occur in the posterior region of the front wing or in the ante-
rior region of the hind wing. A simple and very striking
illustration of this may be seen in the wings of the honey-bee
(Fig. 92).

There is always a certain amount of disturbance of the vena-
tion in the region where the connecting structures are developed.
Both the character and the position of the veins may be modi-
fied in this region. The attachment may be accomplished by
means of a groove, produced by the folding over ventrad of a
portion of the hind edge of the front wing to the extent of 180
degrees, and the development, on the adjacent part of the hind
wing, of a series of hairs, much enlarged and specially modified
into hooks which fit into the groove above. This is the struc-
ture in the Hymenoptera. Another modification, occurring in
Cicada and related insects, differs in having a portion of the
front edge of the hind wing bent upward into a hook locking

iys iy

ee” ff

ta

Vor. 1.] Woodworth.— Wing Veins of Insects. 75

into a corresponding hook of the front wing. A third method,
seen in such Heteroptera as Belostoma, consists of a pair of pro-
jecting processes on the under side of the front wing near its
hind edge. These are of such shape and position that they
clasp around a thickened portion of the costal vein of the hind
wing. <A similar method obtains in the Psocide, but in some
instances in this group the normal structure is supplemented
or replaced by the clasp shown in Fig. 18.

In the Lepidoptera a rather less efficient structure with the
same purpose is the frenulum. See Comstock (’93), Kellogg
(95). In this group the coordination was probably at first
dependent on the overlapping of the wings. With the primi-
tive forms the wings were probably not particularly wide, so
that their overlapping involved a great loss of effective sur-
face. The development of the frenulum, which prevents the
wings being pulled too far apart, allows the reduction of the
overlap to a minimum.

In any case the effect of coordination is to be seen in the
straightening of the adjacent edges of the wings, and a tend-
ency toward reduction in the number of veins of the adjacent
wing areas. In addition to this, there is the local specialization
of the coordinating organ.

Another entirely distinct form of coordination is that which
exists between the two front wings in the Coleoptera and
Hemiptera. In both orders the codrdination is brought about
to make the organ more efficient as a protection for the hind
wings when at rest, rather than to aid in flight. As the result
of this specialization in the Coleoptera the marginal vein of
the hind edge of the elytron becomes much enlarged and pecul-
iarly modified to form a catch, by means of which the two
elytra are held firmly together when at rest, the edge of the
wing at the same time having been so straightened that it
exactly meets the corresponding edge of the other wing through-
out most of its length.

In the Heteroptera the marginal vein is also strongly de-
veloped, but not brought into such intimate relation to the
opposite wing. The most characteristic matter in this codrdi-
nation is the structure of the apical third of the wing. This
region is so shaped that when at rest it exactly overlaps
in the two wings. As a consequence of this duplication there
is not the need of as much thickening of the membrane in this

76 University of California Publications, |[ENTomMoLocy

region, and the wing remains a more efficient organ for flight.
Also as a consequence the venation in this region does not
become obliterated to the extent that it does in the basal
region. The cross veins along the basal edge of the mem-
branous portion of the wing have become specialized into a
strong transverse vein, from which in many cases all the other
veins seem to arise, because of the obliteration of their basal
connections.

The coordination of the wing with various parts of the body
is, like the last, a development to accommodate the wing when
at rest. Either edge may be involved, or the codrdination
may show itself in changes on the disk of the wing. It
is always the front wing that is modified. The front edge
is sometimes made to conform to the shape of the body so that
it will fit closely. Generally the body is modified at the same
time, the adjustment being mutual. In many beetles and ina
few Hemiptera there is a special provision for holding the wing
against the body: but none of these adjustments greatly affect
the venation, except locally. The hind edge modifications are
of a similar sort, and are equally barren of great results upon
the venation.

The modifications that occur on the disk of the wing consist
of a fold, usually quite abrupt, whereby the wing brings itself
into general conformity with the shape of the body. This
fold usually occurs just in front of the first posterior vein, and
when present divides the wing sharply at that point into two
regions. Commonly the line of fold is bordered on either side
by a strong vein, and is not crossed by cross veins, but only
by the marginal vein at the end. This fold has been usually
identified with a line found in many insects where no actual
fold occurs, and with the first folding line of the hind wings.
All of these lines can not be considered homologous, however,
as will be brought out in the discussion of the venation of the
different groups. Where the fold occurs in the front wing it
certainly influences very profoundly the adjacent venation.

The modification of the hind wing that most nearly cor-
responds to this front-wing fold is the system of folds whereby
the wing is made to occupy as little space as possible when
at rest, so that it will be protected by the front wing. The
fold in these wings is always a full 180 degrees, so that the
folded wing lies flat. All wings, except those of the Odonata

Vou. 1.] Woodworth.— Wing Veins of Insects. an

and Ephemerida, have a flexible area at the base behind,
which, if of considerable size, may contain one or more veins.
Very commonly there is more than one fold, in which case all
possess veins, except it be the basal one. The number of folds
may become very large when there is much disparity between
the size of the two pairs of wings, as in many Orthoptera. In
most beetles, in earwigs, and in some cockroaches the folds of
the hind wings lie in various directions in addition to those
radiating from the base, which we have just been considering.
The effect in all of these cases is the almost complete isolation
of each area bounded by folds. The fold thus produces a pro-
found effect upon venation.

There are two distinct influences produced by folding. One
of these is the severance of the veins, already alluded to, and
the other is the production of new veins to define more sharply
the line of folding, as well as for the stiffening of the folded
area. These will be discussed in more detail when considering
the groups affected.

One of the two methods of specialization recognized by
Comstock and Needham in the production of types of vena-
tion is the addition of veins. Adolph considers the maximum
number the primitive condition, so that all specialization
involves reduction. According to the theory of venation
offered in this paper, all venations may be considered to have
been produced by addition in the first instance, though in
most cases a subsequent reduction takes place. The theory
provides for indefinite increase or decrease along what may be
called normal lines; that is, by the duplication either of
posterior veins, of branches of existing veins, of independent
veins, or of cross veins, or by the suppression of these same
veins in the reverse order. This process can hardly be called
specialization, however, but rather simply normal increase or
decrease. Specialization is an unusual process producing
unusual results. A dragonfly possesses a specialized venation
with many veins, and a Psocid one with few veins; but it
can hardly be said that in the one case the securing of the
many veins, or in the other the loss of them, was the means
whereby their peculiarities were brought about.

A true specialization by addition occurs in insects with
folded wings, where veins are produced in a manner quite dif-
ferent from the normal orderly manner in which new veins are

78 University of California Publications. LENTomoLocy

produced. In other cases where numerous veins occur, the
specialization is one that has fixed the many-veined condition.

Specialization by reduction is a common phenomenon; with-
out doubt much of the reduction that has occurred is produced
by normal decrease, but in most cases part of the reduction
has been brought about through specialization. The speciali-
zation of cross veins is always a reduction process. Whenever
any cross vein becomes strengthened it decreases the chances
for survival of the adjacent cross veins. The cross veins
were produced simultaneously over the whole wing, and the
natural tendency would be for them to disappear in the same
way wheneyer the conditions ceased to be favorable for their
production. Those to survive would be the ones specialized
through some special association with the longitudinal veins.

There are at least four ways in which a cross vein may come
into this unusual relationship: It may so lie as to bring the
end of an independent into close connection with it, and so
become, to some extent, the continuation of the independent
vein; it may lie exactly opposite another cross vein as though
the cross vein were continuous over the longitudinal vein; it
may come to lie in a position parallel with the margin and
be favored more on account of the weakness of the membrane
in this region; or, finally, it may gain its preéminence over
neighboring cross veins by binding the longitudinal veins
together at critical points.

In regard to the first of these methods, it may be remarked
that the connection of the independents to the outgrowing veins,
making them take on all the appearance of branches, is an
extremely common occurrence. An independent vein naturally
follows a furrow, and a cross vein has the same peculiarity.
When the longitudinal furrow of the membrane begins to dis-
appear through the approximation of the adjacent veins, it
would not be unnatural to expect that the vein would have a
tendency to turn aside, if a transverse furrow were at hand.
Thus, the cross vein lying here would be augmented by the tip
of the independent and make such a large opening into the
adjacent vein that the cross vein and independent vein together
would resemble a branch of the stronger vein.

The development of cross veins exactly opposite one another
is a very common occurrence in wings. Their usual place of
development is across a vein that lies mainly on one side of

Vo. 1.] Woodworth.— Wing Veins of Insects. 79

the membrane. If one side of the wing in the vein region
does not take part in the formation of a longitudinal vein, it
is not difficult for a.cross vein to extend its influence across to
the other side. Not uncommonly, where the longitudinal vein
is very much to one side, the cross veins actually establish
a connection across from side to side. The vein thus formed
across the longitudinal vein sometimes becomes quite strong
and important. An extreme case of this sort has been
described in this paper in the triangular cross vein at the base
of the wing of Eschna. (See page 31.)

The advantages a series of cross veins would have, in case
they came to he parallel to the margin, is evidently the explana-
tion of the veins of this nature that are so common in the wings
of certain neuropterous insects. The combined cross veins in
this case come to bear the same relation to the outer margin
that the primary vein originally had to the front margin. It
is doubtless only another expression of the power that tends
to hold all the longitudinal veins parallel with one another.
Sometimes there will be more than one line of cross veins
parallel with the margin. The cross veins on the disk of the
wing that sometimes extend as a series of continuous veins
at right angles to the longitudinal veins, are certainly related
structures, though they have no relationship with the margin.

What gives the cross veins that follow the margin their high
development and permanency is the fact that they connect
veins which reach the margin more or less obliquely. The
influence of these cross veins causes the elements of the longi-
tudinal ones to alternate with each other while the cross veins
become continuous and prominent. The cross veins become
coordinated in spite of the tendency impressed on them by
the longitudinal veins—they become stronger, and then the
logitudinals conform to them in the same way they would to a
marginal vein.

The space between the primary and the first posterior vein
is larger than any other interspace in the wing—usually
nearly or quite a third of the whole wing area. The conse-
quent weakness of this region is not wholly relieved by the
insertion of the independents. Here, then, is a place where a
series of codrdinated cross veins serving as a connective
between these two veins would be of particular utility. Some-
times the veins are not codrdinated, but are equally important

80 University of California Publications. [ENTomMoLocy

as connectives. If the independents are strong, and especially
if they are joined together basally, the necessity of the con-
nective cross veins being covdrdinated is by no means as
evident. On the other hand, if the cross veins are coor-
dinated and strong, there is no need of the independents
extending inwardly from the connecting cross vein, and they
may be entirely suppressed.

There is a similar reason for the strengthening of the cross
veins into connectives to bind the primary to the front margin,
or to connect the first and second posteriors. The connective
between the primary and the front margin is of particular im-
portance, and is found in some form in the majority of insects.
This connective may not be so useful in directly preserving
the stiffness of the wing as it is for bearing much of the strain
consequent on the pull of the muscle that expands the wing.

A matter that has a great influence on venation is the
uneven increase or decrease of wing areas. This subject has
never been adequately discussed, nor have I sufficient data at
hand to treat it in detail, and so can consider only the most
evident facts. The best evidence on this subject is that afforded
by the study of the front and hind wings of the same insect,
because we do not have to consider here the question of differ-
ence in type. Likewise, preference should be given to wings that
are as near alike in general appearance as may be. A very good
type with simple venation for this study is that of the Aphide,
and a more complex one is that of the Hymenoptera. In both of
these cases there is a much simpler venation in the hind wing
than in the front, a difference not wholly explainable on the
theory of the reduced size, but apparently largely a matter of
difference in shape brought about by a change in the apical
part of the wing.

In cases of this kind I think it is safe to assume that increase
in a wing area would do just the same things that a decrease
would undo. Where there is difference in size we need not
inquire whether one or both of these processes were in opera-
tion. In the cases suggested above we may suppose, to judge
by the number of veins in the most nearly related forms, that
the front wings are examples of increase and the hind wings of
decrease.

The most evident fact brought out by an examination of
these wings (see Figs. 63 and 80) is, that the modification is

ee

Vor. 1.] Woodworth.— Wing Veins of Insects. 81

almost, or quite, confined to the area increased or diminished.
Thus, if the two pairs of wings in an aphid were cut across
just before they begin to become narrower toward the tip, they
would have an almost identical venation. In Hymenoptera
the front wing somewhat overhangs the hind wing. If the
front wing is cut across just before the beginning of this
extension and the hind wing also at the corresponding point
as indicated by the end of the first posterior vein, the venation
of the resulting basal portions will agree very closely in their
median area. The difference associated with the codrdination,
since it affects the anterior and posterior areas only, constitutes
about all the difference there is.

In a like manner the Psyllidee, the Psocide, and other groups
show the truth of this general contention. The independence
of the different areas of the wings is further borne out by the
development of the system of plication veins in many of the
Elytroptera, as will be described when treating of those groups.

There are three distinct regions of increase or decrease in
an insect’s wing: the anterior, outer, and posterior margins.
The central area can be but little affected according to this law
of the independence of regions. Each of the three regions is
subject to different methods of vein increase or decrease.

The anterior margin is limited at the base by the marginal
vein, which has become essential to the wing because of its
function of holding the wing spread. There can be, therefore,
little or no addition of veins at the base, though the margin
of the wing does sometimes extend out beyond the vein that
originally bounded it. The amount of increase of this area is
distinctly limited by the flight requirements, which hold the
primary near the front edge of the wing. In the cases where
the wing serves some other purpose than flight, as in certain
Phasmide, this area may have enormous extension. The
added veins of this region always partake of the nature of
branches, though probably originating as independents, or pos-
sibly in some cases as cross veins. Whatever their origin, they
are soon indistinguishable from true branches.

The diminution of added veins in the anterior region appears
to take place mostly from the base outwardly, but this is a
matter very difficult to decide. The subject is one capable of
experimental study by decreasing by breeding methods the size
of insects that are subject to considerable variation in nature.

6—Vv

82 University of California Publications, (ENTOMOLOGY

My own investigations have not gone far enough to show more
than this tentative conclusion. Still further decrease consists
in the suppression of the ends of the longitudinal veins. Either
the marginal vein may disappear, all except the extreme base,
as in many Lepidoptera, where the anterior vein has become
very important by taking part in the articulation; or the
anterior may weaken and fade out, as in Diptera, in which
case the marginal remains strong and only disappears by fus-
ing with the primary, the two melting together more and more
till only the bases are separate.

The outer margin may increase by .elongation, or, by a
broadening, probably in all cases by growth in both of these
directions, though in different relative degrees. There is no
limit to the amount of extension, except the mechanical one
due to the necessity of support by the basal parts. If the
extension is chiefly longitudinal, the primary and the inde-
pendents are simply extended and strengthened and the latter
increased in number. If the extension is very great, they
assume an oblique position. On the other hand, if the exten-
sion is largely across the diameter of the wing, making a tri-
angular wing, the effect is greatly to increase the independents,
as in the Ephemeride, to strikingly change their course, as in
the Psocide and Aphide, or to favor the holding of a consid-
erable number of cross veins, as in the Hymenoptera. The
latter tendency we can consider the normal coursé, as in the
Ephemeride there is as much increase in length as in breadth,
and in the other cases the small size of the wings prevents the
natural tendency from showing itself. The peculiarities of the
reduced wings in Hymenoptera indicate that in general the
diminution is by the suppression first of these cross veins, and
then of the longitudinals.

The hind margin is expanded in the hind wing much more
commonly than in the front wing, though there are some cases
where evidence of this specialization may also be seen in the
front wing. Usually the front wing shows evident reduction.
The method of increase of veins is usually by the production
of additional posterior veins, and if there is a tendency in the
wings toward the production of numerous independent veins,
there may be one or more in each interspace. The character-
istic thing about this area, however, is the duplication of the
posteriors. This area can not be very greatly expanded with-

Vor. 1.] Woodworth.— Wing Veins of Insects. 83

out the developing of folds. The extreme specialization in
this direction is seen in the Orthoptera, where the number of
posteriors has become very large.

The process of decrease appears to follow two lines: The
posteriors may fade out and disappear from the hind angle for-
ward, in the reverse order to their production. The other
method is seen in the cases where the bases of the veins have
been modified into an arch-like structure to hold the base of
the wing flat when lying at rest upon the body. In this case
the tips of the veins fuse together, and this fusion extends
more and more toward the base till the two veins ultimately
entirely coa-
lesce. In the
front wings of
Lepidoptera the
posteriors very
commonly show
evidence of this
fusion. In sev-
eral families of
Diptera the first
stages in the
process -can

FIG. 25. Hout wing of male cricket (Gryllus), showing often be seen all
expansion of posterior region to form a sounding-board for gy >
er readiatine redn. g around the outer

edge of the wing,
and the method of the fusion of the marginal to the primary
is the same process.

In contrast with these evidences of the independence of areas,
we may find abundant examples of modifications of one area
which greatly change the size and venation. We will call atten-
tion to only one of the most striking cases of this kind. In the
accompanying illustration of the front wing of a male cricket
(Fig. 25) we may note the great expansion of the posterior
area for the production of the stridulating organ, which so
impinges upon the space usually occupied by the principal
series of independents that the area has practically disap-
peared, except at the extreme tip. Modifications of this kind
are evidently not for flight, and need not be considered in
detail.

84 University of California Publications. LENTomoLocy

Résumé.—Among the methods of modification of the simple
type of venation, those dependent upon the codrdination of the
two pairs, as we have seen, involve various special modifications
of the immediately adjacent parts, as well as a general adjust-
ment of the whole wingareas. The codrdination to parts of the
body is of another kind, appertaining to the wing at rest rather
than in flight, and so is best seen in wings modified as elytra.
In the hind wing the arrangements for folding are dependent
upon this elytral modification of the front wing. There is a
normal method of increase or decrease of veins, which does not
result in any particular specialization. The reduction in the
number of the cross veins is usually true specialization, because
it is not usually a process involving the whole wing, unless it be
associated with decrease in size. The surviving cross veins may
owe their preservation to their becoming continuous with longi-
tudinal veins, or with each other.

Vo. 1.] Woodworth.— Wing Veins of Insects. 85

PART IIl—TYPES OF VENATION.

In the discussion of the types of venation we shall, for con-
venience, group insects into three series, which may be desig-
nated the Neuroptera, Elytroptera, and Neoptera. These are
not offered as natural groups, but merely convenient assem-
blages of groups from a pterological point of view.

NEUROPTERA.

Neuroptera is used in the Linnean sense, but includes all
the older fossil forms that have been assembled under the
name Palodictyoptera by Scudder, thus representing the
primitive winged insects and all the derived groups excepting
the three with the thickened front wings and the three most
recently evolved orders.

The two problems that present themselves most prominently
to the student in this group are, first, the treatment to be
accorded to ancient venations, and secondly, the criteria for
the recognition of the separation of distinct types.

ANCIENT VENATIONS.

The study of venation is nearly coextensive with that of the
classification of insects. Not only is the venation correlated
with the systematic position of an insect, but all the problems
involved in the grouping of organisms apply with equal force
to the interpretation of the specialization of such an organ as
a wing.

The venation of the most ancient fossil remains of insects
has not thus far yielded as satisfactory evidence on the early
course of the evolution of venation as it has been expected to
give. For this reason it has not figured very conspicuously in
the discussions of venation. Before the close of the Paleozoic
era, insects were evidently already abundant and varied in
form. The remains preserved to us in the rocks are every-
where acknowledged to be extremely fragmentary, and the
relation of these ancient forms to the existing groups is a
matter of controversy.

86 University of California Publications. U&NToMoLocy

One of the many fundamental points upon which there is a
conspicuous absence of agreement is the question of the
division line between groups. In the taxonomy of existing
organisms the question is of less practical significance than in
paleontology, or in the study of phylogeny. The view that in
nature there are really no division lines is a statement of a
fact that, though true, is an evasion of, rather than an answer
to, the question, for as long as there are real differences in
nature, there are real divisions, though neither of these be
absolute.

These divisions are by no means to be interpreted as indi-
cating absolutely isolated, unapproachable structural types, but
do signify diverse lines of evolution. All organisms would
probably be proven to have a single common ancestor, if the
course of the phylogeny of each group could be traced back to
the beginning, and probably one would not have to go back to
the very beginning of organic life to reach the individual from
which all existing organic groups have originated. Likewise, in
any group, as insects, the immediate descendants of the first
representative of that group probably do not represent the sep-
aration of the ancestors of the two oldest orders. When this
first division did occur, it is conceivable, indeed probable, that
these two ancestral individuals did not differ as much from
each other. as they did from other forms then existing. A
study of insects of that time would hardly have suggested the
grouping that the descendants of those insects have revealed
in their subsequent development.

For this very reason a study of Silurian or Devonian in-
sects without a knowledge of the insects of later times, if such
a thing were possible, would quite likely result in an entirely
different system of arrangement from that which we use at the
present day for existing insects.

Scudder’s classification of the Paleozoic insects is the nearest
approach to a system of this sort. His Palsodictyoptera brings
together in one order a number of forms which had already
differentiated to an extent sufficient to make them recogniz-
able as the probable ancestors of different orders. Because of
their evident similarity to each other, and difference from the
existing members of the orders toward which they point, they
are-considered as not yet belonging to those orders.

Brauer (’86), on the other hand, would place the ancient

Vor. 1.] Woodworth.— Wing Veins of Insects. 87

types directly in the groups toward which they are develop-
ing, and deny to them the status of transitional forms not yet
sufficiently differentiated to be separated one from the other.

No one will seriously deny the contention that any really
natural system of classification must be one that is natural at
every stage of phylogenetic development from the earliest time
to the present, and that it is proper to shut one’s eyes to the
later developments when treating the earliest forms. With
equal truth it can be maintained that groups represent the
product of two factors working in their evolution, of which one
consists of those forces or conditions accountable for divergence
in structure, and the other of those natural conditions making
intermediate forms untenable. The greater groups are those
separated by wide, impassable gulfs, and the lesser by nar-
rower but none the less impassable gulfs. As soon as species
are distinct, that is, as soon as interbreeding is impossible,
they become as truly isolated as though the differences were
of a more profound character. Classification therefore repre-
sents not so-much the construction of a genealogical tree as
the expression of the natural barriers which set the bounds to
morphological differentiation.

According to the present practice of nomenclature we have
to recognize in the primitive winged insect, for instance, a
species, genus, family and order. Its first differentiation that
became distinct would be a new species within the same
genus, family and order. A new species would be considered
as established as soon as a form became infertile or incapable
of breeding with other forms of the original species. The
total amount of divergence from the original form has no nec-
essary relation to the question, the whole matter being depend-
ent upon those items of differentiation which influence or affect
the fertility or capacity for interbreeding.

For this reason it is evident that the whole species may
develop away from the original type to any extent and remain
one species, though not necessarily the same species as »the
primitive one. It may also vary to an extreme degree, but as
long as the fertility between the individuals remains there is
but a single species. Such extreme variation may either produce
an inconstant polymorphism, or may be associated with chang-
ing external conditions, the insect becoming sensitive to them,
producing phytophagic forms, seasonal variation, or alterna-

88 University of California Publications, (ENToMoLocy

tion of generation, or may become associated with sex, pro-
ducing the so-called secondary sexual characters, together with
sexual dimorphism and such phenomena. Differences much
less than these, if they are such as to prevent crossing, are
sufficient to differentiate species.

In the formation of higher groups, while there is no similar
criterion of separation such as that which decides the difference
between species, there is the same independence of the amount
of divergence between the members of the group, and the
division is determined solely by the extent of the gap sepa-
rating the group from its neighbors. No matter what may
have been the theory of systematists, it has been the uniform
practice in every department of biology to ignore the question
of phylogeny in the lmitations of groups of all ranks, and
base them wholly upon the degrees of separation recognizable.

In the arrangement of the groups thus obtained the attempt
is usually made, with greater or less consistency by modern
naturalists, to indicate the lines or times of their development.
_ Applying these doctrines to the subject of venation, we must
conceive that the first condition of the wing was one in which
we could recognize but a single type of structure—one subject
to considerable variation. Among these variations would exist
forms suggesting types that later became separated. Every
type of venation now existing is historically connected by
regular gradations with the primitive form. If all of these
intermediate forms existed at the present day we should have
to deal with an organ whose variation was great, but at the
same time of little significance. Only the fact of the isolation
of groups by the suppression of intermediate conditions gives
the variation taxonomic value.

With structures, as with species, there are thus two problems
for study that should not be confused. One is the delimitation
of groups, and the other is the phylogeny ofthese groups. There
appears to be much reason for Scudder’s Paleodictyoptera.
Certainly, as far as the venation is concerned, there was not in
the Paleozoic time sufficient differentiation between the various
types of venation to consider them comparable with the divisions
between orders at the present day. We may consider that at
the close of the Paleozoic era there was only a single order of
winged insects, though the name Neuroptera seems preferable
to Paleodictyoptera. At the same time the phyletic lines of

o 6)

Vor. 1.] Woodworth.— Wing Veins of Insects. 9
several of our existing orders were certainly very definitely
established. These lines of development lead to the present
orders Odonata, Ephemerida, Corrodentia, Orthoptera, and
Hemiptera. Those that prefer to consider these orders already
established will thus recognize six orders of winged insects as
existing at that period.

We will now consider in order the ancient types of venation
that have been preserved to us, and the related modern
groups, where these can be recognized. From the nature of
the case one can not speak with as great confidence in regard
to fossil forms as in regard to living, but even fragmentary
data are of great value in tracing homologies. The classifica-
tions of fossil insects of the Paleozoic era given us by Scudder
and by Brongniart scarcely resemble each other in any par-
ticular, but figures of the fossils classified by these two
authors seem to indicate that, to a great extent, this difference
was justified and that we may almost add the two systems
together without doing violence to either.

MEGASCOPTERID 4.

This is the first of the ancient groups of insects in which
there appeared evidence of a tendency to suppress the cross
veins and still preserve a comparatively small number of lon-
gitudinal veins. The
group is quite sharply
defined from other
known insects, and is
supposed to represent an

extinct branch peculiar

>) S av FIG. 26. Diagram illustrating the venation of
to the Paleozoic era. the Megascopteridze. Dotted lines indicate veins
ry A that may be present or not. Brackets indicate

As will be seen by variable place of attachment of veins.

the accompanying figure

(Fig. 26), the venation is distinctly of a neutral primitive type,
which it might be possible so to adjust as to resemble either one
of a number of venation types. In the shape of the wings, in
the number and arrangement of the veins, and in the peculiar-
ity exhibited by a number of the independents in changing their
attachments, this wing agrees with the Termitide; but the
really characteristic things about the venation of the Termi-
tide are not suggested in these wings, and there is little or

90 University of California Publications. [ENTomoLocy

nothing in the other parts of the body to suggest such a rela-
tionship.

Some members of the family possess a venation that could
be compared rather closely with that of Cicada, but here, like-
wise, there is the entire absence of all the characteristics of a
wing of that group, and the body characters show that here
there is still less ground for comparison.

There is a very evident tendency to specialize one or two
rows of cross veins parallel with the hind margin of the wing.
The same tendency is seen to some extent in the Sialide and
very distinctly in the Hemerobide. The Sialide is a compar-
atively undifferentiated group, so that its venation is not very
distinct from that of the Megascopteride, but there are some
characters which will at once separate them. The Megascop-
teride have a rather more consistent venation than the Sialide,
and in the specializations of the cross veins are also dis-
tinctly in advance of that group. As to the Hemerobide,
it is much easier to homologize their venation with the Sialide
than with these insects. We must therefore concur in the
separation of the Megascopteride from all the existing groups.

Their real relationship I conceive to be with the Odonata,
Protodonata, and Ephemeride. AI] the fossil remains we have
of these insects show them with their wings spread, and the
appearance of the base of the wings would indicate that they
were not provided with means of folding. The structure of
the notum of the thorax, moreover, as far as it can be made
out, lends further support to this idea.

If this supposition is correct, the Megascopteride and Pro-
todonata represent two extremes in the development of this
primitive type, for there is lttle similarity between the two
groups. <A group, doubtless derived from the latter family,
has in later time gone through a process of reduction, in which
the cross veins suffered to quite as great an extent as occurred
in the Megascopteride, and produced the Agrion of the present
day. In all respects Agrion is much in advance of the ancient
Megascopteride.

PROTODONATA.

There can be no doubt of the correct identification of the
insect remains that have been placed in this group as being
allies of the Odonata. The wings possess neither stigma,
nodus, nodal sector, areculus, nor triangle. But this list of

Vor. 1.] Woodworth.— Wing Veins of Insects. 91

characters embraces all of the conspicuous marks of the wings
of Odonata. However, with all of these wanting, the whole
facies of the wing is so distinctly like that of the Odonata, that
one would not hesitate to place it in that group.

The enormous size of the insect requires an immense num-
ber of veins, but most of them are the mere duplication of
independents. The really characteristic veins are given in the
accompanying drawing (Fig. 27). At the base there are five
strong veins, alternately convex and concave. An independent
series interpolated behind the primary reaches nearly to the
base. We will consider rather fully the process of specializa-
tion of this wing, in order to lead to the consideration of that
of the Odonata.

The fact is brought out clearly in this group that there is a
close correlation between the shape of the wing and the char-
acter of the venation. Shape and venation were doubtless
developed together, and each is somewhat dependent on the
other. In the adjustment between the venation and the shape,
the great length of the wing required the stiffening of the
longitudinal axis. This is the one predominating mechanical
necessity. All other matters become of distinctly secondary
importance.

The Protodonatid wing is somewhat specialized in a number
of ways, but in nothing is this carried to the extent seen in the
provision for the longitudinal stiffening. The number and
position of the longitudinal veins can have but minor impor-
tance, because of the regular monotony of their shape, size, and
position. Many of them are scarcely more significant than the
cross veins. All veins are simple, except the independent
systems, and perhaps also the posterior, though all but two, or
perhaps three, of the apparent branches are in reality only
attached independents.

The one important and characteristic specialization of the
Protodonatids, which also gives character to the venation of all
the existing Odonata, consists in the limitation of the basal
veins to five strong ones, arranged two on either side of the
primary. These five veins are situated on alternate elevations
and depressions of the membrane. All these veins are con-
nected by cross veins, and the whole forms a very rigid com-
pound truss structure.

In passing from the base toward the apex the arrangement

92 University of California Publications. |ENTomMoLocy

is disturbed by the interpolation of the independent veins,
which crowd off those behind the primary; but these new veins
fall into the general scheme, so that instead of fiye alternating
veins there are seven. With the branching of the independents
the same process is repeated. All this time the veins have been
coming to he more nearly in one plane, so that the alternation
is not so evident. This wing never exhibits the fixity and
regularity of alternation seen in the Ephemeride, except at
the extreme base.

Among the independents one may see quite distinctly a
feature which later in the Odonata comes to be very charac-
teristic. It is the tendency toward the production at regular
intervals along the outer part of the membrane of a series of

pairs of long, un-

branched veins (Fig.
Hitman AANHNKAMWNWW, GUO $96

27). These can be
clearly distinguished
from the intervening

FIG. 27. Diagram of the venation ge he EOE veins by the fact that
odonata. The short lines represent independents nT
without basal attachments. the surface of the

j
"1,
TP
ny

~\S
> WOOO
WH
Ow

wing membrane be-
tween them is nearly perpendicular to the plane of the wing.
With their connecting cross veins they form a simple truss.
Some Ephemeride exhibit the same tendency, and it is well
developed in most Odonata.

The only other feature worthy of particular note in regard
to this venation is the evidence of a tendency to narrow some-
what the space between the primary and the first posterior
just before they are pushed apart by the independents. This
suggests that at this point arises the arculus, which is the most
prominent structural difference between the wings of the Prot-
odonata and the Odonata.

The structures exhibited by the Protodonata are thus exactly
in accord with the supposed ancestral position of the group,
and clearly indicate, though they do not attain to, the venation
of the Odonata.

ODONATA.

There is no group of insects so nearly isolated from all other
existing forms as the Odonata. The whole thoracic organiza-
‘tion forms a special type. Only the Ephemeride possess ini
common with it the peculiarities of hinge structure and muscle

a es at

Vor. 1.) Woodworth.— Wing Veins of Insects. 93

already described. In the venation we find these groups are
absolutely isolated from others by the basal structure and that
the Odonata are further distinguished by a combination of
characters on the wing disk.

The phrase “combination of characters” is used to express
what is often designated facies. It consists of a general plan
of arrangement, —elastic enough to allow considerable varia-
tion, but only within certain lmits both of kind and amount,
and often bound up by definite correlations,—and in addition
to this general plan a series of details, perhaps never all pres-
ent at one time, but having definite places and characters
when present.

The Odonata are distinguished from the Protodonata by the
specialization of three sets of coordinated cross veins forming
connectives (Fig. 28). The most
conspicuous of these is known
as the arculus. The cross veins
in this connective may extend
entirely across the wing from bor-

Oc . FIG, 28. Diagram of the important fea-
der to border; the minimum dis- tues of the wing of the Odonata.
famechigucom the primary tothe , % Arcos: & wianele; m) nodus,
first posterior. When the arculus
is present, as in all Odonata, the independents never cross this
vein, but end in it.

The next most conspicuous connective is the one at the
nodus, with the production of which it is quite likely associated.
This structure (Fig. 28) is found in all Odonata, and nothing
of exactly this type is found in any of the other orders of
insects. The connective at this point binds the primary to the
veins anterior to it, and it also extends behind the primary
and involves one or two of the independent veins.

The third connective is weaker and nearer to the base; it is
sometimes so disguised as to be difficult to recognize. This is
particularly true in the groups with a well-developed triangle.
The character of the connective is much the same as the one
just described, only that it connects the hindmost independ-
ents with the first posterior. These three connectives are the
only constant characters in the venation of the Odonata dis-
tinguishing it from the Protodonata.

In order to understand the development of these connectives,
we should look, first, for the conditions determining the loca-

94 University of California Publications. [ENToMoLocy

tion of the points where the connectives have arisen; secondly,
for conditions within the wing favorable to the production of
this specialization; and thirdly, for evidence of its utility, by
reason of which natural selection will tend to maintain the
structure when once produced. In regard to the last item little
need be said, as the utility in strengthening the wing is very
evident; the fact that the two connectives whose positions
and structure are the most uniform are the ones where the
utility is the most evident, is distinctly confirmatory of this
view.

There is one fact in the matter of the location of the con-
nectives that is at least very suggestive. When the wingpad
of one of these insects is pulled a little to one side, it will be
seen that the organ does not bend in a straight line, but along

a curve corresponding to that

shown in the accompanying dia-

gram (Fig. 29). The reason for

this is evidently the difference in

the stiffness of the different parts

of the wing, dependent on its

21g, 22, Diagram showtngiing of Shape, Now with the wingpacs

aeanpae placed as they are, a current In

the water, or the movement of

the insect, would result in a slight bending of these organs

back and forth. Along the bending line the cells would have

a slightly different environment, and it is possible that the

cells of this region might become more and more sensitive

to this change until they responded by the production of fully
perfected connectives.

How this might be brought about is explainable in two ways.
It may be that we have here simply a direct response to
mechanical stimulation, which is known in both plants and
animals to produce, under proper conditions, very decided
hypertrophies. Another, and perhaps sounder, explanation is
that the bend resulted in a slight displacement of the principal
veins where the bend occurs, so that cross veins occurring at
these points are accordingly strengthened. A support to this
explanation is the fact, already pointed out, that in the Prot-
odonata the primary and first posterior are somewhat approxi-
mated in what will be the arculus region; further, the very
evident bend of the two veins in front of the primary at the

Vor. 1.] Woodworth.— Wing Veins of Insects. 95

nodus, and the approximation of the first and second posteriors
at the other connective, strengthen this view.

The Odonata, a group rich in problems of venation, are full
of interest, but we shall have to confine ourselves to the con-
sideration of a single one—the development of the structure
known as the triangle. This structure constitutes the sole con-
stant difference between the venations of the two divisions of
the Odonata. There are almost innumerable points of differ-
ence between the venation of typical representatives of these
groups, as for instance between Agrion and Eschna; but within
each group there is such diversity that, if we take up one by
one each point of difference and trace it out through the whole
series, the differences vanish.

The triangle is found among the Odonata only in the
Anisoptera, and is, indeed, wanting in all other insect wings.
All writers have, I believe, uniformly, and probably cor-
rectly, homologized all of the veins in the neighborhood of
the triangle with the corresponding veins in the Zygoptera;
none have, however, attempted to trace the method of the
origin of the triangle. The only discussion of its genesis is
the brief one by Comstock and Needham (’98-99, p. 908). I
can agree with them as to the nature of the bounding veins—
the cubitus and two cross veins—which they were the first to
point out, but am not so ready to agree as to the course of
development which they aim to indicate by their diagrams.
There is no necessity, I think, for conceiving the triangle to
have been, as they assume, originally the whole of a quadri-
lateral cell. Moreover, their figure 65 simply provides
for different positions of the triangle, by variations in the
angle made by the “inner cross vein” with the basal portion
of the cubitus and in that made by the bend of the cubitus;
it does not, nor does the wing of any of the Anisoptera, show
any quadrilateral cell, nor any transition between a quadrilat-
eral and a triangular cell. This view assumes that by the
approximation of the corresponding ends of two cross veins

till they meet, a quadrilateral area is converted into a trian- ‘

gular one. My view is that the triangle is formed by the
division of a quadrilateral cell into two triangular cells, one
of which is the triangle. How, in my opinion, this has been
brought about I will explain directly.

I would particularly object to their interpretation of the

EE Eee ee

a

96 University of California Publications. \ENTomoLocy

relation of the veins they denominate cubital and. anal, which
has evidently influenced their conception of the method of the
development of the triangle. If the ontogenetic evidence
afforded by the trachee at this point is reliable, the trunk of the
first posterior vein (their Cu) should be marked Cu plus A,
since both trachese occupy the cavity of the vein Cu, and the
main trunk of the second posterior designated as the recurrent
fourth of the anal, as the trachea indicates in their figure 61
of the hind wing of Cordulegaster. In an insect like this,
where the apical portion of the second posterior (Cu of Com-
stock and Needham) has come to lie so as to be almost a direct
continuation of the first posterior (their Cu), it is not strange
that the relatively strong trachea of the latter vein should
send a branch to occupy the cavity of the second posterior,
even to the exclusion of the one that it originally contained.
Indeed, a similar shifting of trachee occurs at the nodus, as
was first figured by Brongniart. The interpretation of this sug-
gested by Comstock and Needham (p. 904), that we here have
acase of actual crossing of veins during nymphal stages, is
impossible.

But to return to the posterior veins; it must be borne in
mind that we are dealing with structures in a region that is
constantly hable to suppression, because a little narrowing of
the wing would involve the disappearance of parts of these
veins. Examples of this reduction are to be seen in many
Anisoptera, and an extreme case in Lais, where both posteriors
are involved.

’ The development of the triangle probably occurred in insects
in which this region was of rather more than usual importance
in the process of flight. It will be recalled that the character
of the stroke as to speed and path determines the relative im-
portance of the different parts of the wing. We can suppose
that the triangle first appeared in the wing of a primitive
Odonatid, in which there was an increasing functional signifi-
cance in the anal area; a wing witha venation resembling that
of a Calopterygid, which is not far removed from the Prot-
odonata (Fig. 27). The first step in the process was a more
abrupt bending backward of the posteriors, which occurs below
the arculus, to meet the increasing demands of the anal area.
In Archilestes (Fig. 30) we see, in spite of the narrowing of
the wing, a similar position of the posteriors. Now if the wing

ee

Vou. 1.] Woodworth.— Wing Veins of Insects. oF

were broad, so that there must be a large series of veins radiat-
ing from the second posterior, the strain on this vein would
require the strengthening of the area in front of it, which could
be accomplished by the development of a cross vein along the
already existing ridge, extending from the second posterior to
the hind one of the two independents, or at least along the
outer (distal) half of this ridge. In time this vein would come
to look like the direct continuation of the first posterior, con-
necting it with the hindmost independent, as indicated by the
lower of the two dotted lines in Fig. 30. An area thus pro-
duced agrees in every particular with the condition of the
triangle. The second triangular cell resulting from the divi-
sion of this quadrilateral cell is always recognizable in front
of the triangle, and the cell marked s by Comstock and Need-

Primary vein

Orvove

Ridge
FIG. 30. Portion of the FIG. 31. Venation of Protepheme-
base of the wing of Archi- ride, with cross veins omitted. Dotted
lestes, showing probable lines are veins that are sometimes
origin of triangle. wanting.

ham (e. g., their Fig. 62) is thus shown to be bounded by the
two posteriors and the basal half of the newly formed diagonal
cross vein.

PROTEPHEMERID&.

The Paleozoic insects grouped under this family name pos-
sess four equally developed wings. Aside from this character,
and the structure of the thorax correlated with it, these
insects are not particularly different from the mayflies of the
present day. The unusually abundant development of the
posterior veins resembles the condition in Ephemeride. There
seems to be nothing against the idea that we have in this group
an early equialar stage in the evolution of the modern group.

The character of the venation is well indicated in Fig. 31,
in which all except the cross veins are shown. The latter are
very variable in number, rather regularly placed, and quite
uniform in size, therefore not particularly significant. In one
point, however, these early wings stand in contrast with those
of the modern group. It is the absence of free independents.
The production of free independents prior to connected ones

(Xi

98 University of California Publications, (ENToMoLocY

would seem to be the natural order of evolution, but this evi-
dence certainly does not point that way.

PALEPHEMERIDA.,

This family, founded on two Devonian insects, represents a
very much nearer approach to the Ephemeride of to-day than
the group last considered. The fragments are but fractions of
the wings, and while we can not know the whole venation, that
which is present indicates a shape quite similar to that of the
wing of the modern Ephemeride. This group can be consid-
ered a near ancestral type intermediate between the previous
group and the modern Ephemeride.

EPHEMERID.,

The mayflies have, in recent years, occupied a very promi-
nent place in the discussions of the subject of venation. No
other group with wings that do not fold possesses so many
longitudinal veins. One of the easiest ways to trace homol-
ogies is to assume that the primitive organ contained as many
parts as are found anywhere within the group, and then to
imagine that specialization consists chiefly or wholly in the
cutting out from the supposed primitive form of such members
as is necessary to produce conditions resembling each of the
existing forms. On this principle the Ephemeride have been
chosen by some writers as the nearest living representative
of the ancestral stem form.

There is much reason for this assumption in the Ephemeride,
on account of the rather simple undifferentiated venation
possessed by this group. On the other hand, if the geological
record is significant in this case, we must conclude that the
Ephemerid, instead of being a primitive insect, is really highly
specialized, and has the distinction of leaving fossil traces of
two distinct steps in the process of its evolution. On the
grounds of comparative anatomy there is the objection, that it
is more natural to look to an insect with two pairs of equal
wings (especially for the production of wings like those of the
grasshopper), than to a mayfly, as an ancestor. It is quite as
easy to derive all modern venations from one like a Sialid as
from an Ephemerid.

The alternation in the level of the veins, resulting in the
so-called convex and concave veins of Adolph and his followers,
is to be considered as an acquired character rather than a

Vor. 1.] Woodworth.—Wing Veins of Insects. 99

primitive one, and the explanation of the significance of convex
and concave veins, already given, shows that it is not necessary
to derive all insects from an Ephemerid, in order to explain the
presence of traces of this arrangement in so many different
groups.

The veins are not simply convex and concave, as_ the
accounts given would lead one to infer, but there are many
peculiar changes of level not easily explained. The wing of
Hexagenia, from which Fig. 32 was made, was mounted on a

IP

FIG. 32. Venation of Hexagenia bilineata.

slide, and vertical measurements were made, by means of the
graduated fine adjustment screw of the microscope, at the
points where the veins intersected twelve equidistant lines
crossing the wing at right angles to the costa, to ascertain the
relative heights of these several points. The results are given,
in hundredths of millimeters, in the following table. The first
column is the anterior and the last the posterior marginal
vein; the intermediate columns represent all the veins (except
the minute apical independents) occurring back as far as the
first posterior. The first line is nearest the tip.

Table showing the Variation in the Level of the Veins.

Tan eae (eee ee OURS (OU ws 2 Mise ecc Ah tT US oe ea ee
NOMA eOngco twp G Ol wlO 4 7 ceaty @ See ees ee ee eee 8
Wes @ MB} abot) Oe ee ees BS
eee el Omme eel) ear tela 1 Ded, lS eee

Ome ue eet ae Lee Gh LBS Sieh tO a Oly se) lil

Ceca cee Olea p20 eee el Si toe Oo) wl Smee aos
Ob Gy Raaee ee eee we Sey 2h 27 227 Lb
SOM NIOuee twee = 6 20 Ree VBr ne 125) 2) Py 124) 4S 26
20 ee One oe) eee oe Se Ge earn een 2 2 ee Oger Oe 26
2S OU COPlOM fo Oe meee a as sa es Ol, (Oo) senyiolk 1.3) 12
DOVEIO PE AGn oa ye he oo PE B00" 6 2ON 14 4
Gee Oil Oe eg te Pk os rene se Se a ARON gee OD

100 University of California Publications. \ENtomoLocy

The costa had been crushed in the region of lines 4-6, but
otherwise the wing is normal. The character of some of the
irregularities here indicated is shown much more clearly on
the accompanying figure of a fragment of the same wing (Fig.

TULLE Dati
since WEEE Ti Nj
LUT

LET Tf

FIG. 33. Fragment of the wing of Hexagenia bilineata, showing by
contour lines and shadings the heights of the various portions of the
wing. The unshaded areas are the highest, and the solid black the
lowest.

33). This shows the region between the primary vein and the
first strong convex vein behind it, the one indicated by the
twelfth column of the above table. The levels are shown by
contour lines, the
black and shaded
portions being lower
than the unshaded
part, as the figures
in the table indicate.
These data show
that there is without
question more in this
matter than alter-
nate veins of differ-
ent character. The
mechanical problem
of producing the
adequate stiffness of
/ the wing is the most
important factor.
Aside from the
matters already dis-
cussed and those

FIG. 34. Abnormal yenations in Ephemeride after @F1SINE out of the

Eaton. A, Palingenia. B, Elass wra. COC, Callibetis. ore C
5 one alingenia lassoneura. C, Callibetis reduction of the hind

——

Vou. 1.] Woodworth.— Wing Veins of Insects. 101

wing, the wing of a mayfly exhibits but little that is peculiar.
The venation is essentially of the primitive type. There are
among the mayflies a number of interesting cases of reduced
venation. In Palingenia the arrangement of certain of the
independents in pairs, in the same manner as is common in
Odonata, is carried to a greater degree of perfection than in
any other insect, since it involves here the suppression of the
adjacent independents. In the allied Elassoneura the suppres-
sion of longitudinal veins is carried to the farthest extreme;
there only the marginal, the primary, and a single independent
and posterior vein remain. In Czenis and related genera the
reduction and finally complete suppression of the hind wing
have resulted in a very peculiar wing shape and venation, sug-
gesting somewhat Adolph’s figure of a theoretical dipterous
venation. Among these insects we have the best examples of
the suppression of the cross veins.

The homologizing of the veins in the wings of different
genera of Ephemeride has been very well done by Eaton, but
in the comparison of the veins of the front and hind wings,
Redtenbacher is more successful. He has also indicated the
correct homologies with other insects.

STENODICTYOPTERIDE.

The fossil insects of this group, made known by Brongniart,
are of large size, and show a relatively simple venation of
large veins, with the membrane finely and densely reticulate.
These characters make a very dis-

tinct group. The arrangement of ee

the larger veins is according to the SSVow

simple primitive type, and so does SSS

not closely indicate any relation- ini 135 Diseraniontier yen

Eoipmeparticulacs VAll the fossils “ou -of the, Stenodictyontende,
; P omitting the very dense reticula-

Have the wines spread or broken in “ou Dotted lines show | yetns

2 ice et sometimes wanting.

such a way that it is difficult to

say whether the insects could fold the wings or not. The

character of the venation, as far as regards the larger veins,

is shown in Fig. 35.

HOMOTHETIDA.

The study of fossil insects is beset with so many difficulties
that it is not strange that there should be room for great
difference in opinion. The present group, as limited by

102 University of California Publications. [ENToMoLocy

Scudder, contains insects with very different facies, and has
been somewhat dismembered by later writers. The characters
given by the author of its name may be stated in a few
words. The wing possesses an entirely simple primary vein,
free from both the anterior and the independents. This char-
acter is shared by only the Protodonata and Protephemeride
among the Paleozoic insects. The
condition is not always to be
made out with certainty in a fossil
specimen, and so there may have
been errors in identification in
iG; 3, ,Pingram Mlustating the this group, but it is not likely
abundant andi variable cross vel that all the cases are error) Uke
character is not as significant,
according to our present conceptions of the veins concerned,
as it might be under other theories of venation, but it is about
as good as any to be found in such an undifferentiated type of
venation. The accompanying diagram (Fig. 36) shows as well
as may be the character of the venation in this group. I
agree with Scudder that the group has no very evident affin-
ities with any existing insects.

PALEOPTERINA.

The characters assigned to this group are shown in Fig. 37.
The relatively small size of the first group of independents
distinguishes it from the two following families. I am free to
confess that this char-
acter appears to me to
be of very little value,
and quite insufficient
to differentiate groups
larger than genera.

Certainly, in living FIG. 37. Diagram of the yenation of the Paleop-
era ay eee ‘ _terina, omitting the cross veins and the most vari-
forms with a neurop able of the longitudinal veins.

terous venation more

would be required. I am inclined to believe that venation
does not furnish good grounds for classification among these
insects. If we depended upon the venation, it is probable that
closely allied forms would be separated and more distant ones
united. Such a form as Propteticus shows a body structure
that would at once separate it from the mass of the insects of

Vor. 1.] Woodworth.— Wing Veins of Insects. 103

that time, and there is but little reason to think that all the
other insects grouped under this head have a similar structure.
One should not give too much attention, therefore, to the classi-
fication, if he desires to make a just interpretation of the vena-
tion. Whole series of families whose differentiation was not
associated with flight can, for the purposes of the study of
venation, be considered asa unit. The same dictum will apply
to the higher groups as well as to these lower forms.

XENONEURID.

The insect which is the type and sole member of this family
is one of the smallest of the Paleozoic fossils, and possesses
the simplest venation. It seems to
be distinct enough from everything ca
else to deserve a separate place. /

There is nothing so unusual in the ON Oe

venation, however, as would cause See Ss

. , . 3 ald peel FIG. 38. Remains of the wing
surprise if further discoveries should ok Xenoacuniden aicrsecddee

connect it with the common Neurop-
tera. As will be seen by the accompanying sketch (Fig. 38),
the venation is simply a reduced form of the primitive type.

HEMERISTINA.

This group seems to be another case of a very composite
family brought together by the character of the venation.
The character upon which it is
founded is the relatively greater
size of the first group of inde-
pendents, as compared with
those that follow. This differ-
uiilg, Diagram of tie vountion of C0 WHY De cleanly seen by
and most variable independents. SONI ane ere the ac companys

diagram (Fig. 39) with that of
the Paleopterina (Fig. 37). Members of the latter family, as
well as of the one now under consideration, have been identified
by Brauer (’86) as Sialids.

GERARINA.

The character upon which this group is founded, the direct
attachment of a number of independents to the primary, seems
more likely to prove of value in making a natural classification,

104 University of California Publications. [ENTOMOLOGY

than the venation characters in the groups just considered.
The venations are all comparatively simple, but quite variable
as to number of veins. In
the accompanying diagram
(Fig. 40), the venation shown
at the anal angle does not

fairly represent this region
ee tO. Venton ot ee arereaen aim alll the cwimaeets wplacerigin
See eee: this family; but accuracy
on this point is impossible at present, because of the frag-
mentary character of the specimens.

PLATYPTERID#.

This group of Paleozoic insects, as defined by Brongniart,
includes the genus Lithomantis, placed by Scudder in Hemer-
istina, and a large number of similar many-veined forms which
usually have dense reticulated veins, something after the order
Stenodictyopteridz, but not so dense nor with very numerous
close-lying longitudinal veins. It will not be necessary to
figure the venation, as it is not essentially different from
Paleoperine or Hemeristina. Brongniart compares these
insects with the Sialide.

PROTOPERLA.

The insects placed in this group by Brongniart seem to
belong to Scudder’s Hemeristina. They possess a primitive,
and somewhat reduced, venation, and might easily present an
ancestral form of the Perlide. They show no specific character,
however, that will afford undoubted evidence of this relation-
ship. This is perhaps what should be expected, since the vena-
tion in the Perlide is of such an undifferentiated character.

PERLIDA.

The Perlidee possess two pairs of independent wings lying
over each other and flat on the back when at rest. The front
wings thus in a measure serve as a protection for the rather
more delicate, and usually distinctly broader, folded hind
wings. In one case, Nemura trifasciata, the front wings have
become reduced to a semi-rudimentary condition, giving the
insect somewhat the appearance of certain Cerambycide with
abbreviated elytra.

Vor. 1.] Woodworth.— Wing Veins of Insects. 105

The venation of the two wings never exactly corresponds,
but approaches correspondence most nearly when the wings
are of approximately the same size and shape, as is true in
some species of Perla and Nemoura. The family possesses a
few quite characteristic venations and shows certain very
evident tendencies, but there are intermediate forms so com-
pletely connecting the various types that, with the diversity,
it is difficult to define the Perlid type of venation except by
negations. Even such a wing as that possessed by the Paleo-
zoic Protoperla would not be particularly anomalous in this
family, if it existed on an insect at the present day.

One of the most evident and most constant tendencies notice-
able in these wings is the production of a strong cross vein

FIG. 41. Venation in Nemoura.

beyond the middle, corresponding quite strictly with the
principal connective in the higher groups. This-connective is
not very evident in Pteronarchys, where the venation is of the
primitive type, nor in such forms as Eusthenia, where the veins
have come to lie parallel with the front margin, forming long
rectangular cells arranged as bricks in a wall. It is most
evident in Capnia and in the hind wings of Perla.

The ladder-like arrangement of veins is a very characteristic
thing in a Perlid wing. This consists of a series of numerous,
closely placed cross veins, usually occupying two interspaces,
asin Fig. 41. This ladder serves to locate the first posterior
vein, the sub-median of Pictet, which is the middle vein of the
ladder. There is practically no difference of opinion regarding
the homologies of the veins in this group.

TERMITID#.

The peculiarity of the venation of the Termitidee is in part
dependent on the extremely short period the wings are used—a
structure that will wear well not being required --and in part
on the manner of specialization, whereby the posterior two
thirds practically becomes veinless; ‘all the formative material
is apparently appropriated by the primary and the veins

106 University of California Publications, [ENTomMoLocy

adjacent to it. The region in which the veins are thus weak-
ened, or nearly suppressed, is subject to so much variation
that it is impossible to give a diagram that will clearly show
the condition.

As seen in Fig. 42, the venation is of a primitive and simple
sort, such as might be had by the suppression of the cross veins
of several of the Paleozoic types of venation, and in some cases
the suppression also of a few of the longitudinals. This wing

FIG. 42. Diagram illustrating the venation of the Termitide.
Dotted lines indicate alternative attachment of the independ-
ents. The number of these is somewhat variable. Cross veins
are omitted.

shows clearly the instability of the basal attachment of the
independents, since they may be attached as branches to the
primary, or appear as a continuation of the posterior vein.

Whatever differences of opinion there have been in homol-
ogizing these veins, they have all arisen from the idea that the
independent arises from the base like the other veins. There
are always five veins only—the primary, an anterior, a pos-
terior,and the two marginals. The large size of the wing-roots,
and the feebleness of the attachment of the wing to them, are
characteristic features of this group.

EMBIIDA.

The small family Embiids deserves a more careful study in
regard to the nature of the articulation of the wing. The
very anomalous position of these organs on the segment makes
this particularly desirable. The
venation presents no difficulties
of interpretation, as will be seen
by the accompanying diagram
(Fig. 48). The feebleness of the

FIG 43. Venation of Embiidz. Dot-
ted lines indicate veins that may be veins in the posterior part of

sometimes absent.
the wing allies these insects with

the white ants, with which they are commonly associated.
The general direction of the independent veins, however,
makes their venation very distinct from that of the Termitide.

Vor. 1.] Woodworth.— Wing Veins of Insects. 107

There do not seem to be sufficient grounds for giving the cross
veins the value of branches, as has been done by Comstock
and Needham.

PSOCIDA.

As far as the venation is concerned, the Psocide are very
completely isolated. There is a striking superficial resem-
blance between the venation of this group and that of the
Hymenoptera. Only when one attempts to compare carefully
the two is the illusion dispelled. No other group approaches
the Hymenoptera so closely in the shape of the wings as the
Psocidee. It may be that the similarity results from the veins
adapting themselves to meet an identical mechanical condi-
tion due to the shape of the wings.

The typical venation, such as seen in Psocus, is given in
Fig. 44, and the scheme of venation of both pairs of wings in
Fig. 45. From the diagram it will be seen that, excepting the
Embiide, there is
less diversity than
inanyofthegroups <
of lower insects,
and in this respect
the Psocide are
quite comparable
with the Hymen-
optera or other
higher groups.

The homologiz-
ing of the veins in
Psocide has been a matter of controversy. There have been
two questions: One, the comparison of the veins of the front
wing with those of the hind wing; the other, the comparison
of the venation of Psocid wings with other venations. The
former is often avoided by disregarding the hind wing entirely.
There are two principal theories regarding the homologies of
the front and hind wing. According to one theory the pri-
mary vein is nearly suppressed, as it appears to be in the hind
wing of Psocus (Fig. 44); according to the other, this vein is
the one running nearly to the tip parallel with the margin. A
comparative study of the wings of other genera shows that the
latter is clearly the case.

The other question is more complicated. Recent studies

FIG. 44. Venation of Psocus.

108 University of California Publications. [ENTomoLocy

differ greatly. Thus, Scudder considers the veins I have ealled
independents, to comprise all the branches reaching the margin
in the outer half of the wing. Redtenbacher and Brongniart
supposed that only the first two branches correspond with my
independents, the
others being
branches of the
first posterior;
Comstock and
Needham regard

the first two as

FIG. 45. Diagram of the venation of Psocidz. Con- .
stants of the front Wane are indicated LS apa ee those branches of the
of hind wings by lettere. Variables of front wings are ] > ;
represented by dotted lines; those of the hind wings are ae the next
indicated by the letter v. Brackets indicate alternative threeas independ-
attachments of veins.

ents, and the last
two as the first posterior. I believe that Scudder is right in
regard to this question, and think his conclusion is confirmed
by a comparative study such as is epitomized in Fig. 45. If
these two problems are correctly decided, the rest of the homol-
ogizing becomes a very easy matter. The accompanying
diagram (Fig. 45) gives the plan of the venation of both wings
as made out by a comparative study of all the genera, fossil
and modern.

SIALID®.

The Sialide possess the most primitive venation of all
living insects. The variation within the family is by no means
as great as is found in the earliest fossil insects. The specializa-
tion of the group appears to have been the setting of rather
narrow limits to the varia-
tion, instead of the production
of any special feature that
can be recognized. None of
the known Paleozoic insects
possesses a venation that falls FIG. 46. Diagram of the venation of
within the range:of this fam- Sigllde,, Crossteine andthe mooie
ily, though nearly every fea-
ture in the venation can be duplicated in those ancient wings.
The character of the venation may be seen in the accom-
panying diagram (Fig. 46). There is evidence of a tendency
toward the reduction of cross veins and the development of .a
number of large cells in the disk. The manner in which the

Vor. 1.) Woodworth.— Wang Veins of Insects. 109

second and following posteriors are bound together is also
characteristic of the neighboring families. A very similar
structure seen in the Diptera is not the same in origin.

The homologies of the veins as worked out by Redtenbacher,
Brongniart, and Comstock are essentially alike. This is one
of the very few cases where it can be said that there is general
uniformity of opinion.

HEMEROBID®.

Each of the seven subfamilies constituting this group possesses
a venation that is quite distinct from the others; though the
definition of some subfamilies is difficult, because of the great
diversity of their contents. The venation is not so primitive
as in the Sialide, as in every case there is evidence of some
particular specialization.

A character seen in all members of the family, except in the
Coniopterygine, is a tendency to excessive production of inde-
pendents. These are mostly short veins attached in such a
way as to appear as branches, or rather furcations, of the
longer veins. This tendency is carried to the extreme in cer-
tain Myrmeleonine.

The tendency to coordinate, or approximately coordinate,
cross veins is seen in Mantispine, Hemerobine, and Chryso-
pine, always associated with a more or less evident reduction.
The tendency toward reduction extends to the longitudinal
veins in most cases, though in the Mantispine these remain
very numerous. The smallest number of longitudinal veins
is seen in the Chrysopine; in Coniopterygine there is almost
an entire absence of cross veins. Only in these extreme reduc-
tions are the veins practically constant. Ordinarily, as in
most neuropterous venations, there is much variation, even in
the same species.

There has been scarcely any difference of opinion as to the
homologies of the veins. The very evident relationship with
the preceding family prevents any very serious mistakes. In
smaller matters there is room for some difference of judgment,
but these need not be considered here.

PANORPID.

The wings in this family exhibit a remarkable degree of
uniformity for an insect with numerous veins, contrasting
strongly with the nearest allies. The cross veins show a great

|
|
|

110 University of California Publications, [ENTomMoLocy

deal of diversity, but these have not yet become very signif-
icant in venation.

The character of the venation may be seen in Fig. 47. As
in the preceding two fam-
ilies, there is no room for
controversy regarding the
homology of the veins.

The very close resemblance

FIG. 47. Diageam of the venation of Panor- : :
pide. All cross veins not constantly present between this venation and
ee that of a Tipulid makes it
seem possible that this group is somewhat close to the ancestor
of Diptera.

PHRYGANEID 2.

The Phryganeide are of peculiar interest, because the vena-
tion of some of these insects is almost identical with that of
the lower Lepidoptera. They are probably nearly related to
the ancestor of that group, and possibly to that of the Hymen-

FIG. 48. Diagram illustrating the venation of the

Le TEE eamel ae The numbers are those used by
optera also. These suggestions are made the more probable,
because in the one case there is a series of pointed-winged
Phryganeide with reduced venation, quite comparable with
the Tineina in the Lepidoptera, and in the other case a series
of hooks by means of which the wings are hooked together,
much as in Hymenoptera; in the latter case the venation
resembles that of the Hymenoptera more closely than any

other venation does.

The character of the venation in this group is shown in the
accompanying diagrams (Figs. 48-50), on which for conven-

Vor. 1.] Woodworth.—Wing Veins of Insects. 111

ience have been placed the numbers employed by McLachlin
(74-80). The longitudinal veins compare very closely with
the venation in the Panorpide, but the cross veins have been

FIG. 50. Diagram of the venation of
Phryganeide.

A and B, Cloropsyche; C, hind wing
of Helicopsyche; O, veins that must be

FIG, 49. Diagram of the venation suppressed to produce the venation of
of the pointed-winged Phryganeide. Hymenoptera; St, area in which the
The numbers are those of McLachlin. stigma is supposed to be developed.

reduced to a small number, which are very constant in posi-
tion. The homologies are very evident, and have been uni-
formly interpreted correctly.

HEMIPTEROIDEA.

Under this title Scudder (°85) has assembled a series of
insects belonging to the Paleozoic era which have evident rela-
tionships with the Hemiptera of the present day. It is not at
all certain that some of them should not be placed at once in
the Hemiptera, as is done by most authors. The one possess-
ing the least evident similarity in venation to the Hemiptera,
Eugeron, possesses mouth parts that appear to resemble very
closely those of a true hemipterous insect. The venation of
this insect resembles the ordinary undifferentiated type. The
peculiar development of the anal area precludes the possi-
bility of folding, and resembles the condition in the front
wings of Blattide. It is evident that the wings were capable
of flexion.

The insect nearest like modern Hemiptera is Phthanocorus,
from the Carboniferous, known only by the front wing. This
shows very distinctly all the characteristics of the Heteroptera

112 University of California Publications. [ENToMoLocy

of the simpler type, thus indicating that the heteropterous
wing was very early developed. There is even ground for
supposing that the Homoptera were at one time heteropterous,
though of course this is improbable.

Most of the remains supposed to be of this group are very
unsatisfactory for the study of venation, and it is more than
possible that some of the fossils supposed to be homopterous
do not, in fact, belong here at all. It will be quite impossible
to give a satisfactory diagram illustrating the venation of this
group.

ORTHOPTEROIDEA.

The oldest known fossil insect shows evident relationship
with the Blattide. Numerous similar forms, and others sug-
gesting the Phasmide and Acridide, constitute the fossil group
usually placed in the Orthoptera, but considered by Scudder
as not sufficiently differentiated from other Paleozoic forms to
warrant the assignment.

The Paleoblattide, which have been considered by some
authors as representing the most primitive insects, have the
most characteristic and distinct venation of any of the Paleo-
zoic insects, the other Orthopteroidea being more like the
other insects of that era. If the position is well taken that the
Palewoblattide are the most ancient of known insects, then the
first differentiation was that which separated off the other
Orthoptera, from whose ancestral form the other groups arose
later. A more reasonable hypothesis, however, is, that the
Paleoblattidee represent an early specialization, the extreme
member in a series that specialize in such a way as to pro-
vide for the protection of the hind wing by the front wing, the
cause being the same as that by which the Hemipteroidea were
specialized. The difference between these groups concerns
chiefly the mouth parts, but there are differences in venation
resulting from the adoption of a somewhat different method
of coming to rest.

In the Paleoblattide the thickening of the wing was brought
about by the enlargement and strengthening of the longitu-
dinal veins, causing them to He quite strictly parallel, and.
resulting in a very favorable object for preservation in the
rocks. It thus comes about that there is a large series of these
fossils known, and their venation is very easily made out.

Vor. 1.] Woodworth.— Wing Veins of Insects. 113

The wings are quite free from cross veins. The character of
the venation is shown in Fig 51.

The remaining Paleozoic Orthopteroidea exhibit a general
resemblance to the in-
sects just considered,
but are distinctly more
like other insects of the
time. Cross veins are
always present. The

longitudinal vers ane FIG. 51. Diagram of the venation of the Palo-

1 r rer blattide. The figures indicate the range of
proportionately MC ry: variations in the number of veinsin each area.

much smaller in diam-
eter and not so strictly parallel, but they nevertheless resemble
the Paleoblattide more nearly than do any other insects.

ELYTROPTERA.

This group is unquestionably unnatural from a systematic
point of view, since there are no grounds for supposing that
there is any close relationship between either of the orders
here assembled. Each order has responded in much the same
way to the requirements of a similar environment, and so
they present many characters in common. The name here
proposed (Elytroptera) is therefore simply one of convenience.

The anterior wings show a tendency toward the suppression
of the venation, owing to the membrane becoming throughout
vein-like in texture, so that the whole surface is more nearly
uniform. The posterior wings come to exceed in size the
anterior, and so are folded enough to be covered by them, the
venation becoming involved in the folding mechanism. In
this group, therefore, the study of the folding and its influence
on venation is of paramount importance.

BLATTIDA.

The specialization of the Blattide from the Paloblattide
consists in the thickening of the front wings, the perfection
of the posterior region of the hind wings, and the changes
incident to folding. The effect of the former process is rather
the suppression of venation when carried to the extreme, and
need not be considered in detail.

ov

114

The perfection of the folds is of more significance.
nature of the folding may be seen in Figs. 52 and 53.

University of California Publications. [ENTOMOLOGY

The
Com-
paring these fig-
ures, it will be seen
that the wing is
divided longitudin-

Des

ZY
BN

—

VA
be

HIG 2:
Madagascar.
bling the Hemiptera.
in hind wing.

ent in front of it.

Venation of a species of Blattide
The front wings show a structure resem-
Arrows indicate lines of folding

ally into an ante-
rior unfolded and
a posterior folded
area; that the
folded area is en-
tirely divided into
fields and entirely
concealed under
the anterior area;
and further, that
each field embraces
a longitudinal vein
and an independ-

from

Probably the first fold is the original one,

and the following folds represent wrinkles in the posterior area
separating tracts, each of which, after becoming successively

possessed of a longi-
tudinal vein and the
independent acces-
sory, became a defin-
ite flat field. Accord-
ing to this view, the
whole plicated area is
developed in adapta-
tion to the folds; in-
deed, the veins are
produced in the way
they are, in order to
perfect the folding of
this part of the wing.
In different members

of the family there

~

FIG. 53.
hind wing of a cockroach. Black areas are those

Diagrams illustrating the folding of the

reversed in folding. Central figure shows folded
wing from beneath; lower figure a cross-section of a
folded wing.

may be a greater or less number of folds, and always a cor-
responding augmentation or diminution of the number of veins

in this region.

Vor. 1.] Woodworth.—Wing Veins of Insects. 115

There is another specialization in the hind wings of certain
cockroaches that is interesting, since it shows in an unmistak-
able way the formation of new veins in a folded area, and
because it gives us a hint of the way in which the Forficulid
wing must have been produced. This matter was first worked
out by Saussure (’68) in the manner indicated on the accom-
panying diagram (Fig. 54). In the evolution of Diploptera,
the wing is sup-
posed to have
Succes sively
passed through
a series of stages
similar to the
conditions
shown in the
series of genera
illustrated in FIG. 54. Diennid showing intermediate forms between

1 5 1 the normal hind wing and that of Diploptera, showing the
Fig. 54, In this evolution of the plicated tip.

way the whole

of the folded tip of the wing is shown to have come from an
increase in size of the minute triangular bit of membrane
occupying the angle of the principal fold. In none of these
forms is the plicated area at the tip provided with veins, except
the extreme member of the series (Diploptera), where they
must have been produced simultaneously over the whole area,
just as the reticulations of cross veins are.

The lenticular anal field of the front wings of Blattide,
which was already well developed in the Paloblattide, is
clearly comparable with the anal area of all orthopterous and
hemipterous insects, and is also clearly homologous to the
folded anal area of the hind wings. This anal field is always
very distinctly marked off from the rest of the wing by a line
that is usually very much thinned and often quite transparent.
Another line of the same character is to be seen just behind
the primary vein. Between these two lines the area is occupied
by independent veins, usually clearly distinguishable as forming
three groups. The third of these is often specialized so as to
resemble a reverse of the many-branched primary. Because
of this peculiarity this vein has been identified as vein VII by
Redtenbacher and by Comstock and Needham, but the inter-
pretation of Brauer seems to be more sound and to accord

116 Umversity of California Publications. [ENTomoLocy

better with the paleontological evidence. The posteriors belong
wholly to the anal field, and are the veins modified for the
common anal foldings.
FORFICULID®.
The earwigs possess a venation that is entirely uniform, as
far as known, but remarkably different from anything known

FIG. 55. The venation and folding of Forficulide. The dotted lines
show the lines of folding. The black areas are those reversed in folding.
Letters are to identify areas.

elsewhere. The general features of the venation have often
been figured, but the details necessary for the interpreta-
tion of the venation have never been shown. The whole
venation depends upon the very peculiar method of folding.
The wing is packed away into very small space in the
manner shown in Figs. 55 and 56. The piece A: (Fig. 55)
is attached to the body in the manner in which wings are

Vou. 1.] Woodworth.— Wing Veins of Insects. 17

usually attached, and moves in the same way. Piece B,
shown in black, bends down under A when released, because
of the twist of the vein, which I interpret as the first posterior;
this crosses from A to B at about the middle of their length
(compare Fig. 56). The other veins that are under tension when
the wing is spread are the radiating veins given off at the distal
end of B. These elastic veins constitute the whole mechanism
of folding. The process is as follows: On the relaxation of the
muscle that pulls on the anterior marginal vein to keep the
wing expanded, the piece B folds downward. The pull of the
elastic radiating veins folds downward the piece V, which pre-
viously resisted the strain when the wing was flat, folding the
wing along the line
beginning between V
and A. The apical part
of the wing falls into
folds alternating in
direction and deter-
mined by the position
of the veins, in the
manner indicated in
Pig. 55. While this: FIG, 56. Section across the middle of a folded
folding process goes on, wing of Forficula. Letters indicate areas, and cor-
respond with those on Fig. 55.

the piece B is being

folded along the line between A and B into a position directly
beneath A, and all the rest of the wing takes its position
between the two (see Fig. 56).

When the wing is completely folded, the under sides of A
and A’ are in contact, and then follow in regular order B’, C’,
D’ and so on. Likewise, counting from the body outward, we
find first .B, then C, D, EK, ete., in regular order.

In spreading, the pull of the front margin straightens out
A and B, thus spilling out, as it were, downward and forward,
the rest of the wing, which is finally pulled into position as
the piece V is straightened out at the end of A.

How this very complicated mechanism was brought about
is not an easy problem to solve. The common idea that we
have here simply the fan-like fold of the grasshopper doubled
twice after folding, in order that it could be better stowed away,
is certainly entirely wrong, as the figures and the description
of the process show. The place where the radiating veins at

A

118 University of California Publications. UENTomoLoey

the end of B are attached can not be compared closely with
the basal attachment of the radiating veins in other Orthop-
tera. The real basal structures are found on areas A’ and B’,
The correct interpretation of the parts of this wing is probably
that suggested by Scudder, who compares the folding with
that described by Saussure in the Blattide. This suggestion
has never been followed out, though the subject of the homol-
ogy of the veins has been discussed by both Brongniart and
Redtenbacher.

According to this theory, the added apical areas, which
remain in Blattide much smaller than the rest of the wing,
become in this insect the major part, everything beyond C and
C’ being comparable with these areas. The increase in size
occurs in the posterior of the two areas, the anterior being
represented by the area V. The development of an earwig
wing from the more complicated conditions found in the Blat-
tide is not much more difficult to understand than the first
production of the complex cockroach type. We only need to
imagine that the hind apical area, becoming larger, wrinkles
up fan-like when it is folded, and that the veins which develop
in the membrane adjust themselves so as to facilitate this fold-
ing. The lengthening of the membrane in this region would
result in the bending back of the tips, with which new modifi-
cation the veins would accommodate themselves, and finally
the increase of this area would so encroach upon the anal
region that it would lose its original fold, or perhaps, rather,
merge it with the fold that brings the tip of the wing backward.

This explanation accounts for the difference between the
character of the veins in areas A’ and B’ as compared with the
corresponding areas further on, and seems to present no
particular difficulty of any kind.

There are three kinds of veins in the apical region of the
wing: One having basal attachments; another, alternating
with them, and scarcely more than half as long, without basal
attachments; and a third consisting of cross veins. Thereis a
slightly different union between the cross veins and the two
kinds of longitudinal veins, indicating a real difference between
the longitudinals. The shorter ones have much the appear-
ance of independent veins, and the others probably have grown
out from the vein bordering the fold, in the same way that the
principal veins arose from the base of the wing. It would be

Vou. 1.] Woodworth.— Wing Veins of Insects. 119

hard to determine the order of the development of these veins,
but it is possible that it is from the front backward.

Both Redtenbacher and Brongniart have attempted to
homologize the veins of these insects, and agree in considering
the radiating veins comparable with the radiating veins in
grasshoppers. The posterior would thus be their vein IX as
well as the rays. It is very difficult to conceive the stages in
the migration of the vein IX out beyond the tip of the inde-
pendent, and, besides, the folding of this wing is fundamentally
different from the folding of the anal area of Orthoptera.

ORTHOPTERA.

The Orthoptera, omitting the two families just considered,
are, as far as differentiation is indicated by the venation, much
simpler than the preceding families. The Paleozoic ancestors
bore the same relation to the Paleoblattide that the modern
forms do to the Blattidee. We know that the venation is one
capable of extreme specialization, as is shown by the remark-
able stridulating organs in the males of Gryllide and Locus-
tide (see Fig. 25, showing the wing of the common cricket).

The shape and function of the front wings are not such as to
favor the development of any new or more definite venation,
and the hind wings are taken up with the arrangements for
folding, which requires and maintains a monotonous repeti-
tion, instead of high specialization. There is great similarity
between the venation of the Blattide and that of other Orthop-
tera, except that the Blattid venation is rather more definite
and complete. There are many examples of reduced venation
among the Orthoptera, but in no case is there reduction with
specialization, except it be for some other function than flight.

The Phasmidee are mostly apterous, and when possessing
wings, these have been so reduced that only the more impor-
tant of the longitudinal veins remain. Under these conditions,
unless the wing is too much reduced, the characteristics of the
orthopterous venation are clearly seen, especially in the folded
area of the hind wing.

The Mantide possess the least differentiated wing in the
whole order, and the greatest number of longitudinal veins.
These are extremely variable, as in the Neuroptera, and very
regularly placed.

The Acrididee show something of a reduction in the number

120 University of California Publications. [ENTOMOLOGY

of veins, as contrasted with the Mantide. They also present’
in most cases, a modification of the independent region in the
form of an enlarged cross vein, or an abrupt branching of the
independent veins, which disturbs very strikingly the uni-
formity of the wing. The veins are extremely variable, how-
ever, so that there can be said to be here a tendency toward a
specialization, rather than one already accomplished.

The Locustidee and Gryllide have very simple hind wings,
but the front wings are distinctly changed for the production
of the stridulating organ in the male, and a corresponding
though less pronounced change in this structure in the female.
In the Locustide there has been a great reduction in the num-

FIG. 57. Diagramatic representation of the wing of an Acridid.

ber of the independents without much decrease in the size of
the areas, and there are fewer cross veins, thus making numer-
ous larger cells rather than a dense reticulation. In the
Gryllide the median area is almost entirely suppressed, leaving
only a few independents spreading out fan-like.

The venation throughout these families differs from that in
Blattide, in that the independents of the first group are
attached as branches to the primary. They resemble in this
respect their Paleozoic ancestors. Inthe matter of the homolo-
gies of the veins, there is no difficulty in comparing these
venations with those of Blattide, so the same questions arise as
did in that family.

HETEROPTERA.

*

The Heteroptera stand in a somewhat intermediate position
between the Orthoptera and the Coleoptera. With the former
they agree in the leathery texture of the front wings, and with
the latter in the close application of the wings to the body
when at rest. In neither case is there close similarity, how-
ever—nothing to suggest the probability of a genetic relation-
ship. It is likely that the order of their specialization was
that which the fossil remains thus far known indicate, viz.,

a a et

>

apts

a end On eee as

os

te

Vor. 1.] Woodworth.— Wing Veins of Insects. 121

Orthoptera, Heteroptera, and last of all the Coleoptera. That
they were not derived one from the other is quite evident, but
the exact source of the groups is unknown.

The characteristic of the front wing of Heteroptera is the
division of the area of the wing into three regions, by the sep-
aration of the clavus and membrane from the rest of the wing
known as the corium. This division is very evidently the
result of the action of two distinct influences. The first is the
same as that which separates the anal field in the Orthoptera-
It is probably an adaptation to permit the flexion of the wing
without the excessive narrowing that exists in Coleoptera.
The transverse line, that marks off the membrane, is asso-
ciated with the overlapping of the tips of the wings. In the
wing of a cockroach (Fig. 51) we see an exactly similar modi-
fication of the front wing, such as
must have occurred in the first special-
ization of the heteropterous wing.
The separation of the embolium is
evidently produced by the line that
is seen in Orthoptera, just behind the
primary, and the cuneus may pos-
sibly represent a stigma.

Mhewenation oftheiront.wing (Fig. 4 «5 pinganade soos

x. OO. lagramatic repre-

58) is very much obseured. The pri- sentation of an Heteropterous
venation.

mary is usually at some distance from

the front margin, and the posterior is on the edge of the clavus.
Between these is a single independent, which forks at about
the middle of the wing; each of these forks is connected to the
adjacent vein by means of a cross vein. Usually these veins
can scarcely be made out, and it may be that in other cases,
where the venation is obscure, other independents may also
extend to the corium.

At the edge of the membrane there is much variation. My
interpretation of the structures here presupposes two trans-
verse veins bordering the division line between the membrane
and the corium, one belonging to each. These may both be
considered as accessory veins. The inner one comes into con-
nection with both the primary and the independent, and in
some cases many become quite prominent. The membrane
vein runs parallel with the inner one, and usually turns out-
ward at both ends, but in some instances it crosses the division

122 University of California Publications, [ENTomoLocy

line, becoming attached at both ends to the veins of the
corium. From this accessory membrane-vein the discontinu-
ous independents, having reéstablished themselves, proceed to
the margin.

The hind wings are unlike anything in other orders. The
venation is simple and not particularly difficult to interpret-
The primary is strong and on or near the margin. A little
beyond the middle it makes a bend and is thickened or gives
off a spur-like branch toward the tip. Just behind this line
there is a wrinkle corresponding to the anterior longitudinal line
of the front wing. There are two independents fused together
at the middle, and connected to the primary by a cross vein
in most cases. The posterior of these two independents unites
at the tip with an accessory vein along the first fold. Between
this fold and the next there are two of these accessory veins,
which unite with each other toward the base, and are some-
times connected by a cross vein. Still behind this is a forked
first posterior and, finally, a single second posterior.

Redtenbacher is the only one who has attempted to trace
the homologies of the veins in this group, and I differ from
him here, as in Orthoptera, and for the same reason. He
believes that the vein VII should lie before the anal field in
the front wings. The basal attachments of the veins show
that his interpretation is not correct.

HOMOPTERA.

If one excludes the Phytophthires, this group becomes fairly
uniform in the character of its venation. The front wings
vary from a condition in which both pairs of wings are equally
transparent, as in Cicada, to one in which they are leathery
and nearly opaque: but never to a stage where the course of
the veins can not be made out with certainty. There is much
difference between the venations of the two pairs of wings,
making it difficult at times to homologize the veins.

The family Cicadide forms the most distinct group of the
Homoptera. The venation is shown in Fig. 59. The most
curious feature of this venation is a mark (dotted line, Fig. 59)
extending across the wing, which can be seen only in certain
lights upon the membrane: but wherever this line crosses a
vein, it is very evident, because the vein is here entirely inter-
rupted. There seems to be no other explanation of this line

Vor. 1.] Woodworth.— Wing Veins of Insects. 123

than that it is a vestige of the line which, in the Heteroptera
separates the corium from the membrane. In this connection
a similar structure in the fossil Fulgorina is, as already pointed
out, of interest. It is possible that these insects, instead of
representing the most primitive condition among the Rhyu-

oN
~ Se
SNe ee”
FIG. 60. Diagram of the venation

of Fulgora_lanternaria, The reticula-
. 59. Wings of Cicada tibicen. tions and finer branches are omitted,

chota, are in fact derived from an ancient ancestor resembling
the Heteroptera in the structure of the wings. If Hugerion
represents the primitive shape of head in this group of insects,
it is certain that the primitive shape is more nearly realized in
the Heteroptera than in the Homoptera, the latter standing
intermediate between Heteroptera and the Phytophthires.

The front wings of the Homoptera always possess a very dis-
tinct clavus and are adapted
to the body in exactly the
same way asin the Heterop-
tera. In the Cicadide, where
the wings are not elytral
organs, there is still to be
seen this adaptation in the
long wing-groove in the
notum, already described and
Shown in Fig. 12. This is
certainly confirmatory of the
idea of the Heteropteral origin
of all Rhynchota. The Hom-
optera may, nevertheless, still retain the venation most like
the primitive ancestral form; for the more completely a wing
remains membranous, the better may one expect the venation
to be preserved.

FIG. 61. Venation of Typhlocyba comes.

124 University of California Publications. |ENTomoLocy

The hind wings are never as peculiar as those of the Heter-
optera, but resemble quite closely the front wings. Neverthe-
less, one can readily trace in the hind wing of Fulgoride,
which clearly represent the most primitive group among the
Homoptera, all the veins of a heteropterous wing. We may

conclude, therefore, that the hind wing of most of the families |

of Homoptera have more nearly kept pace with the front wings
in their specialization than have those of the Heteroptera.

In the matter of the homology between the front and hind.

wing, Comstock and Needham have clearly made a mistake.
There are three less cells in the hind wing of Cicada than in
the front wing, and the missing three are the apical ones.
These authors and Redtenbacher place their vein VII one vein

too far forward, as they have consistently done throughout:

this order and in the Orthoptera.

PHYTOPHTHIRES.

The four families constituting this group possess very simple
venations, which are quite different in the several families. In
only the Psyllide is the venation extensive enough to show
clearly their affinity to the
higher Homoptera. The small
size of all these insects is
probably accountable for the
simplicity of their venation.

The venation in the Psyl-

lidee is shown in Fig. 62. The
We largest part of the area of the
as wing is occupied by the inde-
Algo Diagram of the venation of the pendents, which appear as ¢
fold; the other dotted Tues tepiesenyyciy= juWiCe-lOrked, Dr anchimuconoauie
primary. The marginal ex-

tends all around the wing and there is but one posterior.

The Aphids differ from the Psyllide by the suppression of
the posteriors, and by some differences in the attachments of

the independents, as shown in the accompanying diagram
(Fig.638). The hind wing is more reduced than in the Psyllide,
and its maximum venation is nearly the same as the minimum
of the front wing. The additional independent and the cross
vein at the tip are very rarely present.

The Aleurodide exhibit a still further reduction. The front

—

Vo. 1.] Woodworth.— Wing Veins of Insects. 125

wing is the same in its maximum condition of venation as the
maximum of the hind wing of Aphide, and the hind wing the
same as its minimum. (See Fig. 64.)

The Coccide have the hind wing almost suppressed, and the

FIG. 64. Diagram of the vena-

FIG. 63. Diagram of the venation of the tion of the Aleurodide. Dotted

Aphid. Dotted lines represent veins that lines indicate veins that are
are sometimes absent. sometimes absent.

venation of the front wing the same as the minimum of the
front wing of Aleurodide. (See Fig. 65.)

The venation of the Phytophthires has a nomenclature of its
own, not relating it to any other groups, the only investigator
who has attempted to homologize the veins being Redten-
bacher. This author admits
only two branches as belonging ee a ee
to vein V (corresponding to my Te Gat
independents) in the Psyllide,
and sees none at all in the
Aphide. The posterior he con-
siders to be vein IX, apparently on the ground that the fold
represented by the fine dotted line is uniformly considered as
vein VIII. The unsoundness of this view has already been
shown.

FIG. 65. Venation of the Coccide.

PHYSOPODA.

The thrips are uniformly small insects with very simple
venation. Both pairs of wings are narrow and fringed with
long hairs, as is not uncommon in very small insects. The
function of these hairs is to extend the amount of available
wing surface, since they are close enough together not to comb
the air, but to press upon it as though forming a flat mem-
brane. Such an arrangement would be an impossibility in a
larger wing. Hairs beyond a certain length would not be stiff
enough. The fringe that exists on the wings of some of the
higher insects may be nearly, or quite, as wide as in the thrips,

126 University of California Publications, [ENToMoLocy

but they are so small as compared with the width of the wing
that they are a very unimportant addition to the effective
width.

The front wing is generally distinctly wider than the hind
wing, and when this is the case, the venation of the latter is
very much simpler. The general
character of the venation is shown
in Fig. 66. No one has _ before
attempted to homologize the veins,
tee of the theeopoda’ “Doited because the venation was so simple
lines indicate velns that maybe. thaws, could enor bemnrde toga

into the complex schemes of vena-
tion of previous investigators. According to the scheme here
proposed, there is no difficulty in the matter. We see the
marginal, the primary, a posterior, and an independent. The
latter, with the cross veins being often wanting.

COLEOPTERA.

The front wings in Coleoptera are so completely transformed
into protective coverings for the hind pair, that their venation
is almost indistinguishable. The hind wings possess veins
of two types, usually quite distinct. Those near the base are
cylindrical, and resemble the veins of more typical wings,
while those toward the tip are usually solid for the most part,
and tend to fade out around their edges so as to become very
indefinite in shape. These are usually not treated as veins,
but we have previously shown that they are not essentially dif-
ferent from undoubted veins. In this case they are certainly
not to be homologized with any veins in other orders, in spite
of their essentially identical structure.

The most characteristic thing about the hind wing in this
order is the manner of folding. These wings exhibit a good
deal of variation in this respect, but there is one point in which
they all agree, if the wing folds transversely at all—abortive
wings, or those not fully covered by the elytra, lose the charac-
teristic fold that occurs in all normal wings. This common
character is the dividing of the area between the two strong
divergent veins—the primary and the first posterior —into four
triangular areas by the lines of folding. These triangles may
be seen in Figs. 67-70. Besides these, there is always one, and
sometimes two, basal folds, and there are also extremely

Vo. 1.] Woodworth.— Wing Veins of Insects. 2%

variable apical folds. In Fig. 67 the veins show very distinctly
the lines of folding for a double basal fold, but in this case the
wing is folded only once. The breaks in the veins in this case
are doubtless vestigial.

The method of folding is as follows: The largest white area,
the third coming from the front margin near the base, is the
only one that remains uncovered. All the adjacent areas
bend under it. The area in which the primary vein lies is
thus brought partly beneath this area. The two small triangles

FIG. 67. Venation and folding of Harpalus caliginosis. Dotted

lines indicate lines of folding; black areas those that are reversed

in folding.
also fold so as to lie partly over the primary vein. The tip of
the wing beyond the triangles folds first longitudinally along
a slight curve, which causes the extreme tip to fold back upon
the more basal portion. The folding is brought about by the
approximation of the tips of the primary and the first posterior
by their own elasticity, and the extension of the wing by a pull
on the anterior marginal by the extensor muscle.

In the form represented in Fig. 68 the same method of folding
occurs for all but the anal area, where there is one more com-
plete fold. In this wing, as in the one previously considered,
the area (white) containing the first posterior vein is upper-
most when the wing is folded, all the adjacent areas being
doubled beneath it. The most striking feature of this wing is
the arrangement of the veins whereby this third area is carried

128 University of California Publications, ENTomoLocy

forward, as contrasted with the wing previously studied. An-
other type of folding, as seen in Dermestes and other beetles
with rather short wings, is shown in Fig. 69. This can be
compared, area for area,
with the wing last con-
sidered, but the result is
a single rather than a
double folding of the
apical region. These
three types of folding
may be considered as
representing three stages
in the development of
a complicated folding,
such as is found in the
long-winged forms, the
fundamental, or short-
winged, type being represented by Dermestes, and the culmi-
nating stage being seen in Staphylinus.

The Staphylinid type is illustrated in Fig. 70. Here there
are three transverse folds and several added longitudinal folds
in the apical region of the wing. The method of inserting a
longitudinal fold is clearly
indicated in the diagram.
The carrying of the basal

FIG. 68. Diagram of the venation and folding
of a beetle with double basal fold. Dotted lines
are lines of folding; black areas those reversed
in folding.

transverse fold across the
anal region 1s another fea-
ture in which this insect is
peculiar.

The venation of beetles,
as will be seen in these
iUlustrations, consists of two

distinct types, as was first

FIG. 69. Diagram of venation and folding
pointed out by the author in Dermestes lardarius. Dotted lines indicate

folds; black areas those reversed in folding.

(Woodworth, 89). One is
seen in Fig. 67, the other in Figs. 68-70. In the former, there
is a cross vein beyond the first transverse, and the whole
venation about the fold is unique.

The most conspicuous elements of a beetle’s venation are
the primary and the first posterior; between them there are
two independents. Basal to the fold, one or both of these may

Vou. 1.] Woodworth.— Wing Veins of Insects. 129

become connected with the principal veins, and they appear
often as recurrent branches. Beyond the fold the veins are
broad, flat, and solid. There are also two independents
between the first and second posteriors. These are often more
or less fused and are generally connected to the adjacent veins
by long cross veins. The second posterior is generally double
and branching, with one or two cross veins. Behind the sec-
ond there is at least one other posterior vein.

Redtenbacher, and Comstock and Needham (’98—99, p. 561 ff)
have overlooked the independents in front of the first poste-
rior, and have locat-
ed vein VII (=Cu)
on the independ-
ents behind this
vein. In this they
are certainly wrong.
None of them have
advanced any con-
vincing argument in
support of their iden-
tification. Atracto-
cerus is an aberrant
rather than a primi-

: 3 FIG. 70. Venation and folding of Staphylinus cin-
tive member of this ”optarous. Dotted lines indicate folds; black areas
~ are those reversed in folding.

order, but there is

nothing in its venation that would lead to the conclusion that
the veins have the homologies assigned to them by these
authors, and the data in reference to tracheation are, to say
the least, conflicting. The evident character and position of
the veins certainly conform to the other interpretation.

NEOPTERA.

The Diptera, Hymenoptera, and Lepidoptera, which consti-
tute this group, have been supposed by many investigators to
form a single natural super-order, but the transition stages
connecting these orders are difficult to conceive of without
going back to their neuropterous ancestors. They are the only
large groups, unless it be the Coleoptera, that were entirely
absent in the Paleozoic era. In all other cases there were
species present that possessed enough of the characteristics
of existing orders to make their recognition quite easy.

9—Vv

|

130 University of California Publications. [ENToMoLocy

There is but little similarity between the venation of the
wings of these three groups. The venation in each case is repre-
sented by only a few veins; all the cross veins present are
specialized, and almost as constant and important as the lon-
gitudinal veins. This cross-vein specialization is the keynote
of the venations of the Neoptera.

DIPTERA.

The higher flies possess a venation which is very character-
istic and uniform. In no group is there as large a number of
species with so little variation in their vein arrangement as is
true in the higher families of flies. The lower flies present
abundance of variation in venation, so that at times it is
rather difficult to carry out homologies with them.

The character which at once sharply separates the flies from
all other insects is the special modification of the hind wing
into an organ only remotely associated with flight. This
requires a somewhat different development of the front wing
to suit it for independent action. In other orders, where the
hind wings are greatly reduced, there is still some connection
between the two wings. In the mayflies, for example, either
the reduced hind wing overlaps the base of the front wing, or,
in case it is so narrow that it would too readily slip off, there
has been developed a hook-like structure that holds the wings
together in much the same way as occurs in all Hymenoptera.
In the males of scale insects the hind wings are minute and

yod-like, but here, again, they are hooked to the front wings.

In conformity with the peculiarities of Diptera just pointed
out, the venation exhibits a character in the hind areas which
contrasts strongly with that found in any front wing, except
in those Neuroptera where the wings remain somewhat inde-
pendent; but in these cases the resemblance is evidently not a
true homology. In the Sialide and other related neuropterous
insects, in the posterior region of the wing are three veins that
arise from a common center and are bound together by cross
veins, producing basal cells in the manner so common in
Diptera, but every other indication is to the effect that the
veins inclosing the basal cells in Diptera lie in front of those
surrounding similar cells in the Neuroptera. We doubtless
have here analogous structures produced by similar mechanical
needs.

Vor. 1.] Woodworth.— Wing Veins of Insects. 131

The families in which the largest number of veins occur are
the Tipulide and the Psychodide; the smallest number is
found in the Cecidomyide. These three families are all closely
related, and the group to which they belong is, by general
consent, placed at the lower end of the series.

The Cecidomyide are very closely related to the Myceto-
philide. The venation of both
families is shown in Fig. 71.
With the exception of the vein
marked CM, which may disap-
pear in both families, the max-
imum venation in Cecidomyide

is exactly the same as the mini-
mum venation in Mycetophilide.
As far as the venation is con-
cerned, the two families may be
considered as representing de-
grees of development along

FIG. 71. Diagram representing the
venation in Cecidomyide and Myce-
tophilide. Solid lines represent con-
stants in Cecidomyide; line CM is
variable in both families; lines M are
variable in Mycetophilide, and en-
tirely absent in Cecidomyide; the
other dotted lines are constant in
Mycetophilidez and variable in Ceci-
domyide.

exactly the same line, the Mycetophilide furnishing the transi-
tion from the simplest condition, that of the Cecidomyide, to
one complex enough to make easy comparison with the mini-
mum condition of the Tipulide.

The Tipulide are a large and quite varied group. The veins
are not particularly difficult to homologize, but their homology
is often disguised by the
relative position of the
forkings, which results in
a series of patterns so

different in general ap-

FIG. 72. Diagram of the venation of Tipu- pearance that the vena-

lide. Solid lines represent constants, dotted 1 ‘ a4
lines variables. Bracketsshow varying attach- tion seems at first sight

ment at veins. B, indicates concave veins, ‘
according to Adolph; R, convex veins: RB, not to be comparable.

veins considered by that author as concave in 1 ‘ 7 rQYr19-
some species and convex in others. Fig. 72 shows the varia

tion within the group.
The venation of Tipulide is easily comparable with most of
the types of fly venation, and may be considered as represent-
ing, if not the primitive, at least the normal venation of Diptera
and the most available type for comparison and the determina-
tion of homologies.
The Dexide constitute a very small family most closely
allied to the Tipulide. The venation is given in Fig. 73, but
shows nothing particularly significant.

132 University of California Publications, [ENTomotocy

The Psychodidee are minute flies rather isolated from other
families. The venation is shown in Fig. 74.

The Culicidee, Chironomid, and Blepharoceridz constitute
a group with very little difference in venation; they are the

FIG. 74. Venation of the Psycho-
; ; ; didz. Dotted line shows vein which is
FIG. 73. Venation of the Dexide. otten absent.

nearest allies to the Psychodidw, their wings being somewhat
intermediate between the venation of that family and the nor-
mal type. (See Fig. 75.)

—<

FIG. 75. Diagram of venation of FIG. 76. Diagram of venation of
Culicide, Chironomide, and Blephar- higher Nematocera. Maximum is
oceride. Minimum venation is that found in Rhyphide; minimum in
of Chironomid; adding veins Orphnephilide; Bibionide and Simul-
marked B gives venation of Blephar- ide are intermediate. Letters at end
oceride. The maximum venation is of dotted lines indicate families in
that of Culicide. which these veins are variable.

The higher Nematocera exhibit but lttle variation in actual
venation, but vary greatly in the arrangement and relative
strength of the veins. The Bibionide vary most in the former

aN

hae

J' FIG. 77. Diagram of venation of Bra-

chycera. Vein marked DC is wanting _ FIG. 78. Wing of Midas, show-
- in Dolichopodidze and Lonchopteride. ing extreme of fusion and shifting.
Brackets indicate tendency of union of In this case two anterior pair of
tips, and arrows direction of shifting. veins have become entirely fused.

particular, and in the Simulide the independents and the
veins in front of them are strong, while the rest are reduced to
scarcely more than wrinkles of the membrane. (See Fig. 76.)

The Brachycera (Fig.77) have a very uniform venation, but
several groups exhibit an evident tendency toward the

Vou. 1.] Woodworth.— Wing Veins of Insects. 133

fusion of the tips of certain veins and the shifting of the
united tips forward. These tendencies are responsible for
whatever difficulty there may be in the interpretation of the
venations. The figure of Midas (Fig. 78), which is an extreme
ease, shows the effect of such modification on the general
appearance of the venation.

The Cyclorapha are the most uniform of all the groups in
the matter of venation. The accompanying figure of the vena-
tion of the house
fly (Fig. 79) indi-
cates the character
of this venation.
The only variation
of much. signifi-
cance is the weak-
ening, or entire Te
suppression, of one
or more of the posterior veins, or the posterior cross veins, in
the smaller species. This reduction process is carried to the
extreme in the Phoride. In no case is there any difficulty
in recognizing the character of the remaining veins.

The Pupiparia possess a venation which is evidently of the
same type as that of the last group. It lacks the cross veins,
but does not differ greatly from the wings of the Orthorapha
that have undergone this reduction. As far as the wings indi-
cate, there is little ground for making the Pupiparia a group
of as high rank as is usually done.

There are three principal schemes of nomenclature for the
dipterous wing. One of these is that of the systematists, in
which the evident strong veins of an Orthoraphid wing are
numbered, from in front backward, first longitudinal, second,
etc. In the lower groups this is quite consistently followed,
the additional veins being usually treated as branches or as
added veins. In the reduced venation of the Cecidomyide a
mistake was early made in the attempt at homologizing the
veins, and this error, though recognized, was held to for sake
of uniformity within the family. The terms, therefore, do not
have the same significance in that group as elsewhere.

The scheme proposed by Adolph, and followed by Redten-
bacher, is the most elaborate attempt yet made to bring the
Diptera into line with other orders in vein nomenclature.

Wing of a house fly.

134 University of California Publications. [ENToMoLocy

Adolph’s scheme of the dipterous wing includes not less than
twenty-two alternating convex and concave veins, which, with
the exception of a number of veins at the two ends of the series,
includes a concave and convex vein for every vein actually
existing in the vein of one of these insects. In Fig. 72 it is
shown that in a single family every one of the independents
has been treated in one species as convex and in another as
coneave. According to this author, as has already been
explained, there is a fundamental difference between a convex
and a concave vein; consequently, veins that have been sup-
posed to be the same in two insects are actually different, if in
one case they are a little above the average level of the mem-
branes and in the other below it. If this were true, the whole
nomenclature of the systematist would be wrong.

The third system was worked out by Comstock and Com-
stock (95) and agrees in nearly every particular with that
adopted here, except in the terms used as names for the veins,
and in other slight differences dependent upon our different
conceptions of insect venation. This system does no violence
to the ideas of systematists, whose knowledge of the compara-
tive anatomy of this organ is certainly not to be despised.

HYMENOPTERA.

The venation of the Hymenoptera exhibits a remarkable
degree of uniformity. From the sawflies at one extreme to the
bees at the other, we find only
a single type of arrangement.
Such variation as occurs is all
in the nature of reductions
from the typical venation.
It is usually found associated
with small size, and is most
prevalent among the parasitic
wasps.

FIG. 80. Diagram illustrating the vena- While within the order the
pou og he Teor Dotted lines problems of homology are rel-
atively easy, this is not true

when we attempt to compare this venation with those of other
groups. As none of the other higher orders approximates
closely in venation that of the Hymenoptera, there is ample
ground for difference of opinion. I am compelled to take a

VoL. 1.] Woodworth.— Wing Veins of Insects. 135

positive position very distinctly opposed to the latest study
on this subject, that of Comstock.

The uniformity of structure makes it quite immaterial where
we begin the study of this venation. The order contains no
families with anything that may be called an ancestral vena-
tion connecting it in any peculiar manner with another group,
or showing how the peculiar venation of this order has arisen.
The only unusual forms are the reduced venations found in
several families, and the slightly increased venation of the
Tenthredinide (Fig. 80). This group has been considered the
lowest in the series. Here, therefore, rather than in the groups
with reduced venation, may we hope to find suggestions on
the phylogeny of this venation type.

The Tenthredinide and Siricide (Fig. 81) differ from all
other members of the order in the possession of an additional

Cisse ; aie
—— SS

—
=S a
\ Saiiaweee=

‘

ie

es

FIG. 81. Diagram of a Siricid wing. DU Hea Tes wee See each
Dotted line shows variable cross vein. suppressed.
posterior vein in the front wing, there never being more than
the merest rudiment of it in other families. ‘They also some-
times possess additional cross veins, one in the neighborhood
of the stigma in the front wings, and a couple near the tip in
the hind wings; very rarely there is an additional independent
at the tip of the front wing.

The parasitic forms show the extreme in the reduction of the
venation, where the only vein remaining may be the primary.
From this all conditions may be found up to the almost
complete venation of the larger Ichneumonide. The Cynipide
(Fig. 82) are peculiar in the great distance between the primary
and the margin, which is even more than is shown in Fig. 82;
when it alone is present, the appearance is very misleading.
The Proctotrupide (Fig. 83) show best the range of venation
among this series of families. The Ichneumonide (Fig. 84)
show the suppression of the basal end of the anterior inde-
pendent. The Pelicinide (Fig. 85) show a remarkable approach

136 University of California Publications. [ENToMoLocY

toward the venation found in the wings of ants. The sup-
pression of veins in both the Pelicinide and Formicide is
associated with large size and is, therefore, to be considered of

Se ws

FIG. 84. Diagram of the

venation of Ichneumonide.

FIG. 83. Venation of Proctotrupidee. Dot Dotted cross vein sometimes
ted lines show veins liable to suppression. wanting.

a different kind from that in the smaller parasites. The
Evaniide (Fig. 86) and Brachonide (Fig. 87) both have
reduced venations, the latter family to an extreme degree.

er

FIG. 86. Diagram of the vena-

FIG. 85. Diagram of the venation of tion of Evaniide. Veins some-
Pelicinide. Dotted lines indicate times absent indicated by dotted
veins sometimes absent. lines.

The ants (Fig. 88) possess as distinct a venation as any
group of Hymenoptera, having no close neighbors, unless the
resemblance in Pelicinide proves to be more than superficial.

bore

FIG. 87. Diagram of venation FIG. 88. Diagram of venation of Formi-
of Brachonide. Dotted tip of cide. Dotted lines indicate veins that
vein often ‘suppressed, may be wanting.

Among the wasps, the most significant venations are seen in
Pemphredonide (Fig. 89), Crabronide (Fig. 90), and Scoliidee
(Fig. 91). In the higher wasps and bees the variation is only
slight, and consists in the suppression of one or two cross
veins and the ends of the longitudinals.

%¢

Vor. 1.] Woodworth.— Wing Veins of Insects. 137

It will thus be seen that there is not a great deal of diversity
in the Hymenoptera, and that what there is consists of the
simplest type of reduction.

The comparison of the front and hind wings is not very
difficult, on account of the many reduced front wings that

FIG. 89. Diagram of venation FIG. 90. Diagram of venation
of Pemphredonide. Dotted tips of Crabronide. Dotted veins
of veins often wanting. often absent.

approach the normal venation of the hind wings. The com-
parison with other orders isa much more difficult matter. The
nomenclature in common use among systematists is peculiar,
and is not claimed to indicate
necessarily relationships with
other orders. Redtenbacher
considers the vein V of his sys-
tem to be represented by the
spur shown only in the figure
MIG oe Dinerae bet eenatton’ ot of Tenthredinide. Comstock
Scoliide. Dotted tips of veins and cross applies the corresponding term
veins may be absent. r ‘
media to the outer portion of
Redtenbacher’s vein VII, while I would apply the term to all
of Redtenbacher’s vein VII and to Comstock’s veins V and
VII. Redtenbacher does not attempt to name cross veins, but
Comstock identifies most
of the veins that have the
appearance of cross veins
as being in fact longitud-
inal veins,..or their
branches, that in this
venation assume positions
distinguishing their na-
ture. The only ground for FIG. 92. Venation of Apis.
this interpretation ap-
pears to have been the author’s unbounded faith in a yena-
tion scheme that has a definite number of branches to each
vein. I believe he has done violence to the facts in attempt-

138 University of California Publications. |HNromoLocy

ing to overthrow the conceptions in regard to the nature of
the elements of this venation which have stood the test of
practical use in the system of nomenclature adopted by the
systematists. )

If the suggestion made, when considering the venation of
the Phryganeide, in reference to the ancestry of this group,
is to be given any weight; if the homologies I have indicated,
which were determined on purely anatomical grounds, are to
be given any weight, they will entirely support the system of
nomenclature here proposed. In most schemes of classifica-
tion the Hymenoptera are placed nearer the Diptera than to
the Lepidoptera. The structure, however, really supports the
suggestion of relationship shown by the resemblance of the
larvee of sawflies and those of moths. These two orders are
the youngest groups, since the Diptera are shown by the
geological record to be distinctly older. The suggestion of
Phryganeid origin is a very attractive one, for if the figure
of the fore wing of Chloropsyche is compared with that of
the Hymenoptera, the veins supposed to be suppressed will,
in each case, correspond with irregularities in the course of
veins not otherwise easily accounted for. Thus there is a
vestigial structure shown in many Hymenoptera, indicating
the suppression of a longitudinal vein in the median and first
two discoidal cells, and the lanceolate cell in sawflies suggests
the lacking posterior veins. If all these are added and the
third transverse cubital and second recurrent suppressed and
each longitudinal vein forked, we have exactly the venation of
the front wing of Chloropsyche. The hind wing of Helico-
psyche corresponds very closely with the venation of the hind
wing of a sawfly.

LEPIDOPTERA.

Throughout the greater part of the Lepidoptera the venation
is remarkably uniform. The wings are notably free from cross
veins, and the number of longitudinal veins is small, even
though some of the insects in this order are very large, the
largest of any existing insects. Increase in size in this order
evidently does not have any necessary influence on the vena-
tion. Extremely small members of the order, however, often
exhibit decided decrease in the number of veins, giving us the
only types that cause any trouble in determining the homologies.

Vor. 1.] Woodworth.— Wing Veins of Insects. 139

To Comstock we are indebted for the fullest study of the
venation of the Macro-lepidoptera, and to Spuler for the only
extensive comparative study of the Micro-lepidoptera. These
two authors are in essential agreement as to the homology of
the veins within the order, but they differ in regard to nomen-
clature and in the comparisons they institute with other insects.
There is a decided difference between the opinion of these later
writers and the older views held by the systematists as to the
relation between veins, though not in the determination of
homologies.

Spuler considers Micropteryx as representing the most primi-
tive existing condition of venation, but Comstock supposes it
to be Hepialis. These two venations are really very similar,

FIG. 93. Diagram illustrating the vena-
tion of the Jugate. Solid lines show con-
stants in Macropterygide, dotted lines the

variables. The venation in Hepialide is FIG. 94. Diagram of the vena-
like the minimum venation of Macroptery- tion of Tineina. Solid lines are
gide with the veins marked H in addition. constants, and dotted lines varia-
Vein marked HX occurs only in Hepialide. bles.

as will be seen by examining the accompanying diagram (Fig.
93). In Hepialis the front and hind wings are a little more
nearly alike and the resemblance to the venation of the Phry-
ganeide a little more complete, so that we must agree with
Comstock in regarding this the more primitive form.

The resemblance of the venation of these insects to that of
the Phryganzide is so perfect that there seems to be but little
doubt that there is a very close relationship between the two
groups. This idea is strengthened by the fact that in reduced
venations, such as occur in the Tineina, we see the same
method of reduction exhibited as that seen in the Phryganei-
dz. (Compare Figs. 49 and 94.) If our determination of the
veins is right in the latter group, there will be no difficulty in
establishing the homologies in this case.

One vein at the end of the discal cell has long been recognized

i
1

)

140 University of California Publications. [ENTomoLocy

as being different from a simple branch, and early received the
name “ discoidal vein,” or the more suggestive one, ‘“ independ-
ent,” which we have chosen as a common name for all similar
veins. Redtenbacher called this vein and the one in front of
it vein V. Spuler added
the vein behind, making
three, which he denom-
inated vein III. Com-
‘ stock agrees in this
point, but uses the Red-
- tenbacher number V,
~ and also calls the vein
media; with this idea I
also coneur, but sub-
stitute, however, the name independent for these veins.

Just behind the first posterior vein (vein IV of Spuler, VII
of Redtenbacher and Comstock, the one usually called media)
is vein V of Spuler, or VIII of Redtenbacher, which is usually
represented by only a fold. I consider this an independent
vein. These two are the only important points on which there
is a difference of opinion.

The accompanying diagram (Fig. 94) shows the extent of
the reductions that occur in the smallest moths, the Tineina.
Only one independent in both wings is constant, and in the
hind wings, when very
greatly narrowed, all
the veins and branches
niay be suppressed. The
variation presents no
unexpected features.

Other small moths
are represented by the
diagrams of the Sesiide
(Fig. 95), Pyralidina
(Fig. 96), and the Tor-
tricide (Fig. 97), all of

FIG. 95. Venation of Sesiidee.

: FIG. 96. Venation of the Pyralidina. Dotted
which are remarkably line is a variable vein.

uniform in contrast to

the Tineina. The suppression of the basal portion of the inde-
pendents indicates that they are really quite differentiated.
The groups that come closest to Hepialis and Micropteryx,

Vor. 1.] Woodworth.— Wing Veins of Insects. 141

aside from the Tineina, are the Cosside, Psychide, and related
Bombycoid moths. These are the ones separated by Comstock

FIG. 98. Diagram of the venation of
the Cossidze and other Bombycoid
FIG.97. Venation of the Tortri- moths, in which the independent

cide. crosses the discal cell.

asthe “ Generalized Frenate.” The venation of these is shown

in Fig. 98.
The variation in the higher moths and in the butterflies, as

; FIG. 100. Venation of butter-
FIG. 99. Diagram of the venationof the higher flies. Dotted lines indicate
moths. Dotted lines are variable veins. variable veins.
shown in Figs. 99 and 100, is very slight. A great many
families and minor groups can be distinguished in their vena-

142 University of California Publications. |ENToMoLocy

tion by such characters as the relative length of parts, position
of forks, ete. Such questions, however, are beyond the scope
of the present paper.

Table of Comparative Nomenclature.

SEMPER ET AL. BEDE SPULER. Comstock. Tue AUTHOR.
ii 1-Costa Anterior Margin

Costal II I II Subcosta Anterior
Subcostal Til II IIL Radius Primary
Branches 1 1 1 1 Branch 1

2 2 2 2 2

3 3 3 3 3

4 4 4 4 4

5 5 5 5 5

6 Veni MOE. a V,1 Media Anterior Independent
Discoidal V5.2 2, 2 Middle Independent
Median Vil
Branches 1 1 3 3 Posterior Independent

p 2 IOWA al VII, 1 Cubitus First Posterior

3 3 2 2 Branch of Posterior

VIII V VIII Anal Secondary Independent

Submedian IX A x Second Posterior
Anal XI B XI Third Posterior

Vor. 1.] Woodworth.— Wing Veins of Insects. 143

RESUME AND CONCLUDING REMARKS.

The purpose of the foregoing study has been to establish a
rational theory of venation. The development of this theory
has involved numerous radical departures from the views
usually held, not only in regard to the venation itself, but also
to the origin of the wing and its effect on the evolution of the
thorax. I believe there have been ample grounds for the
changes that have been proposed, and all that fit together to
make a comprehensive general scheme whereby the nature of
venation as a whole, and a host of peculiarities in individual
venations before unexplainable, are made clear.

The wings are conceived of as organs whose function is so
definite, and the requirements of which are so exacting, that
the mechanical necessities are the dominant factors in their
first production and in all subsequent specializations. Thus,
in the origin of the wing the most important matter was its
utility for flight. It was this that was responsible for the
location of the wings on a particular region of the insect’s
body, even for the determination of the part of the segment
occupied. A wing could not have been produced in another
situation, since flight is dependent upon equilibrium. The
shape and structure of the organ are determined by its func-
tion, to a predominating extent. There must be a wide,
expanded area, coupled with lightness and strength, and the
latter must be distributed so as to meet the particular strains
to which the wing is to be subjected. The relation to the body
must be such as to permit the proper motion of the wing, even
though it require the readjustment of the whole thoracic
structure—indeed, it is clearly responsible for the most profound
change in the structure of the body wall that the segments
have experienced. It could also be shown that the internal
structures were subjected to a similar readjustment.

In the wing itself, the one specialization of importance for
aérial locomotion is the production of the veins. These struc-
tures are developed primarily forstrength. Any other function
is certainly secondary. Their structure when best developed

144 University of California Publications. [ENtomoLocy
Yy

is that which is mechanically the strongest for the amount of
material used, a hollow cylinder, though the veins may become
rod-like, or flat, or even disappear in situations where they
have become useless. There is no essential difference in either

the structure or origin of the different sorts of veins; this

indicates that their functional utility is the important reason
for their existence. Insects have veins in their wings, pri-
marily because they are serviceable.

The arrangement of veins is in like manner the expression
of the mechanical needs of the wing. The venation is con-
ceived of as receiving nothing from the precursor: of the wing
except veins that were developed in the same way and to meet
identical needs with those of the organ after it became adapted
to flight. A system of trachee that were developed for the
purpose of respiration can not, according to this view, have
any relation with the veins subsequently developing in order
to strengthen the organ, since the functions are incompatible,
and since the production of a vein in a tracheal gill would, by
just that much, reduce the breathing surface.

The detailed study of the venation of the various groups of
insects has only strengthened the idea of the predominating
influence of the functional requirements, and confirmed the
writer in the view that in this we may find a basis for a true
theory of venation. The theory, stated very briefly, is, (1) that
»ach vein is produced for mechanical reasons: the marginals
to stiffen the edges of the wing, the primary to serve as the
dominant vein, the anterior and posteriors to supplement
the primary at the points of greatest need on either side, the
systems of independents and cross veins to occupy the areas
of the wing remaining poorly provided with longitudinal
veins, and, finally, plication veins in the Elytroptera to meet
the special requirements at the points of folding; (2) that the
production of different types of venation has proceeded along
comparatively narrow lines bounded by inflexible physiological
requirements, by which the distinctness of the groups has been
maintained,

There will be found, therefore, in all venations certain fac-
tors in common, and in each group certain features that can
not be so strictly compared, because they were produced under
conditions where the mechanical requirements were different.
To the extent to which these requirements are uniform, and

Vor. 1.] Woodworth.— Wing Veins of Insects. . 145

only to that extent, shall we find the venations comparable.
The nomenclature of venation in each group may be to that
extent independent. Thus, in the Odonata we may be able to
use with entire propriety the term primary vein at the same
time that we use it in the Diptera, but it will not do to apply
the term triangle, or basal cell, to structures in the two groups
interchangeably. This principle has never been clearly recog-
nized by those who would establish a uniform nomenclature.
There must be a distinct nomenclature for each group, with
only such terms in common as are clearly homologous. How
far homologies can be carried will doubtless be for a long

' Neoptera

Orth at
Orthopterar<]

Ephememeridee
Falephewremeridze

Protephemerig ra Subulicornes

FIG. 101. Diagram illustrating the phylogeny of insects. Extinct
Be abs plese: Comnitely, ae to existing groups; those
time a disputed point.’ My own conviction is that strict
homology (that is, the use of terms for veins completely com-
parable) is not possible throughout the whole series of insects,
in the case of any vein, not even the primary, because cross
veins and independents become, to all intents and purposes,
branches of the other veins, and it is not unlikely that true
branches often become disconnected, and would then be indis-
tinguishable from independents. It is possible, however, to
use terms in a more elastic sense and to speak of the primary
with the knowledge that in one case it has branches not
strictly comparable with those in the other. The marginal,
the primary, the anterior, and the posteriors can, I think, in
this sense be used in all the orders of insects, as has been done
in the preceding pages. The independents and most of the

1 Vv

146 University of California Publications. UENToMoLocy

cross veins are, as a rule, so diverse that only in allied groups
can comparisons be made with any confidence.

The relationship between the groups is more clearly shown
by the wings than by any other set of characters. There are
a number of cases of parallel development, producing groups
that are convenient, though not natural. The Subulicornes
have two entirely distinct venations, showing the two groups
not to be really closely allied, but should have been brought
closer together than in the accompanying diagram (Fig. 101).
The groups of the Corrodentia are probably not as distinct as
indicated on the diagram and possibly should be derived from
the same line as the Orthoptera, notwithstanding the absence
of ancient fossil remains. The Elytroptera certainly consist
of independent groups. The Neoptera are all closely allied to
each other, but here, also, the group is probably not mono-
genetic. While the diagram here given is based on the
venation, it is nowhere contrary to evidence obtainable from
the study of other characters, and, modified in the manner
just indicated, represents the author’s views of the phylogeny
of the groups of insects. The sequence of the orders should be
as follows:

. Aptera.

. Neuroptera.
Odonata.

. Ephemerida.
. Corrodentia.
Orthoptera.
Hemiptera.

. Coleoptera.
Diptera.

. Hymenoptera.
. Lepidoptera.

1
FPODDNAARWNH

pe

Vou. 1.] Woodworth.— Wing Veins of Insects. 147 a

LITERATURE.

The following works are cited in the text:

Aarons, Frank.
1886. On somenew Psocide. Proc. Acad. Nat. Sci. Phila., Vol. XX XVIII,
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Adolph, Ernst. ;

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1879a. Ueberabnorme Zellenbildungen einiger Hymenopterenfliigel. Ibid.,
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1883. Zur Morphologie der Hymenopterenfliigel. Jbid., Bd. XLVI, pp.
41-132, tab. 6.

1885. Die Dipterenfliigel, ihr:Schema und ihre Ableitung. Jbid., Bd.
XLVII, pp. 269-314, tab. 24-27.

Amans, P. C.
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Audouin, Victor.
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Bellesme, J. de.
1879. Sur une fonction de direction dans le vol des insectes. C. R., Tome
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Brauer, Friedrich.

1869. Betrachtungen iiber die Verwandlung der Insekten im Sinne der
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1878. Part II, ibid. Bd. XX XVIII, pp. 151-166.

1885. Systematisch-Zoologische Studien. Sitzungsb. d. k. Akad. d. Wis-
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1 taf.

1886. Ansichten iiber die paleozoischen Insecten und deren Deutung.
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Brauer, Friedrich, und Redtenbacher, Josef.
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Brongniart, Charles.
1893. Recherches pour servir & l’histoire des Insectes Fossiles des temps
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148 University of California Publications, [ENToMoLocyY

Brunner von Wattenwyl., C.
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Burmeister, Hermann.
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Calvert, Philip P. |
1893. Catalogue of the Odonata (Dragonflies) of the vicinity of Philadel-
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Chabrier, J.
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Comstock, John Henry.
1893. Evolution and Taxonomy. Wilder Quarter Century Book. 114
pp., pl. 1-2. ,

Comstock, John Henry, and Comstock, Anna Botsford.
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Comstock, John Henry, and Needham, James G. |
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Eaton, A. E.
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Fischer, Leopoldi Henrico.
1853. Orthoptera Europea. Leipzig, 1853. xx 454 pp., 18 pl.

Ganin, M.
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Gegenbaur, Carl.
1870. Grundziige der Vergleichenden Anatomie. Leipzig, 1870. 2te Auflage.
xii+ 890 pp.
Gonin, J.

1892. Recherches sur la métamorphose des Lepidopteren. Bull. Soc.
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Graber, Vitus.
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Vou. 1.] Woodworth.—Wing Veins of Insects. 149

Griffiths, George Charles. are On
1898. On the Frenulum of the Lepidoptera. Trans. Ent. Soc. Lond., 1898,
pp. 121-182, pl. 4.

Haase, Erich. :
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Jahrg. XIV, pp. 116-117.

Hagen, Hermann August,

1870. Ueber rationelle Benennung des Geaders in den Fliigeln der Insekten.
Stett. Ent. Zeitung, Jahrg. XX XI, pp. 316-320. °

1881. The Devonian Insects of New Bronewiok: Bull, Mus. Comp. Zool.,
Vol. VILI,.pp. 275-284.

1881. Some Psocina of the United States. Psyche, Vol. III, pp. 195-196,
207-210, 219-223. ;

1882. [On the Tri-regional Division of the Thoracic Segment.] In Proc.
Camb. Ent.-Club, Dec. 18, 1882. Psyche, Vol. III, p. 417.

Hampson, George F.
1896. On the Classification of Three Subfamilies of Moths of the Family
Pyralide: the Epipeschiine, Endotrichine, and Pyraline.
Trans. Ent. Soc. Lond., 1896, pp. 451-550. .

Kellogg, Vernon L.
1895. The Affinities of the Lepidopterous Wing. Am. Nat., Vol. X XIX,
pp. 709-717.

Kempers, K. J. W.
1899-1900. Het adersysteem der Kevervleugels.. Tidschrift Entomol-
ogie, 42 Deel, pp. 180-199.

Kolbe, Hermann Julius.
1889-93. Einfuhrung in die Kenntnis der Insekten. Berlin, 1889-93,
pp. xii + 709.
Lang, A.
1888. Lehrbuch der Vergleichende Anatomie. Jena, pp. iv + 566.

Lendenfeldt, R. von. |
1881. Der Fltig der Libellen. Sitzungsb. Akad. Wissensch. Wien.
Bd. LX XXIII, Abt. 1, pp. 289-376, Taf. 1-7.

Lowne, B. Thompson.
1890-95. The Anatomy, Physiology, Morphology, and Development of
the Blowfly. London, 1890-95. 2vols. pp. x + viii + 778.

MeLachlan, Robert.
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Marey, C. J.
‘1869. Mémoire sur le vol des.Insectes et des Oiseaux. Ann. Sci. Nat.
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Mayer, Alfred Goldsborough.
1896. The Development of the Wing Scales and their Pigment in Butter-
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ee

4 ih!

150 University of California Publications. [ENTomoLocy

Mayr, Gustav L.
1855. Formicina Austriaca. Verh. Zool.-Bot. Gesellsch. Wien, Bd. V,
pp. 273-478, Taf. 1.

Meinert, F.
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UNIVERSITY OF CALIFORNIA PUBLICATIONS
TECHNICAL BULLETINS

COLLEGE OF AGRICULTURE, AGRICULTURAL EXPERIMENT STATION

ENTOMOLOGY
Vol. 1, No. 2, pp. 153-198 October, 1906

CATALOGUE OF THE EPHYDRIDAE, WITH BIBLI
OGRAPHY AND DESCRIPTION OF
NEW SPECIES.

By BURLE J. JONES.

The present paper represents part of the scientific results
of the study of the mosquito problem near Burlingame. One
of the flies described herein was the most conspicuous inhabitant
of many of the mosquito-infested pools. Three species of this
family have been hitherto credited to California; ten others are
now identified in the University collection, of which four are
new.

NEW SPECIES OF CALIFORNIA EPHYDRIDAE.

Notophila varia, n. sp., ¢ and p.—tThe lighter speci-
mens of this species are similar to N. bellula Williston, from
which they can be distinguished by the brown vitta of the
pronotum, the smaller size, and usually the lighter color of
the antenne. The brown spots on the upper part of the
mesopleurz seem also to be lacking here.

Varying from black to dull brown; front yellowish brown,
with sometimes a trace of a broad, darker-brown stripe from
the orbits on either side at the vertex to the lower edge of
the front. The narrow orbits gray. Antenne yellow, rarely
reddish yellow, the upper part of the second and third joints
and sometimes the tip of the third joint brownish or blackish.

Es

ABE.

i

{

oo

154 University of California Publications. [E#NTomMoLocy

Face and cheeks yellowish, often grayish directly beneath the
eyes. Palpi light yellow; all of the femora except the knees,
the lower part of the pleure, and the abdomen with close
orayish pubescence. In most specimens a black or dark-brown
cloud extends from the lower part of the pleure on each side
upward across the shoulders, thence backward diagonally across
the thorax, meeting at the center above, thus leaving a triangu-
lar light-brown spot on the pronotum. A reddish-brown vitta
crosses this triangular spot from the front backwards, dis-
appearing in the darker cloud behind. ‘The dark color is
usually slightly dispersed or entirely wanting on the scutellum ;
oceasionally it crosses the center of the thorax diagonally
or is otherwise irregular. Abdomen hoary like the femora,
with brownish spots irregularly arranged on the anterior part
of each segment, generally leaving the lateral border and a
median line above grayish. Tips of femora, upper part of
front tibie, middle of hind tibie and the tarsi, except their
tips, yellow; lower part of front tibiz and their tarsi blackish.
Tips of all the tarsi slightly blackish. Wings uniformly a
trifle brownish; halteres pale yellow. Face falling off con-
siderably beneath the antenne for this genus, with a slight
median ridge above. Face narrow; entire body slender. Pat-
tern of thoracic macrochete and venation of wings normal.
Length, 3 to 3.25 mm. Middle and Southern California.

Parydra aurata, n. sp., ¢ and p.—Very similar to
Parydra bituberculata, but differs fundamentally in the absence
of the conical warts of the scutellum, also in the absence of
brownish dusting on the face, in the lighter color of the
antenne, and the possession of uniform golden pubescence.
Entire insect clothed with golden-yellow pubescence, under
which it is black, more or less shining, and sometimes with a
steel-blue reflection on the abdomen. Epistoma more densely
pubescence toward the oral margin; antenne brown or brown-
ish yellow; tibim reddish yellow, sometimes distinctly brownish
in the center; tarsi reddish-yellow, last joint brown or black.
The characteristic bristle on each side of the face long and
slender; a distinct impression above and toward the center of
the face from this. Orbits and cheeks broad; clypeus very
prominent; epistoma with two broad, very flat transverse
ridges crossing the center above the clypeus (these are some-

E~ 5

Vor. 1.] Jones.—Catalogue of the Ephydridae. 155

times almost indistinguishable). Antennal arista long and
slender, base pubescent, bare toward the tip. Vitte of the
thorax almost obsolete; the four characteristic rows of tho-
racic hairs slender, black, strongly reclinate. Wings brown,
veins brownish black; a hyaline spot at each side of the broad
brownish band of the cross veins, a fifth spot at the base of
the marginal cell; second segment of the costa nearly twice
as long as the third. Second longitudinal vein without ap-
pendage; third and fourth longitudinals with a slight diverg-
ence at the tips; fifth longitudinal scarcely attaining the
margin of the wing. Length, 3.5 to 4 mm. San Francisco,
California.

Ephydra millbrae, n. sp., d and » .—Front bronze bluish
green with sparse grayish pubescence, epistoma shading from
a dark brown above to a light chestnut brown at the oral
margin, sides of the face beneath brownish pruinose; antenne
dark brown, first segment lighter; proboscis black above,
yellow at apex; palpi ight brown; orbits purplish blue imme-
diately beneath and behind the eyes. Thorax from the lateral
margins above marked with alternating vitte, five of blue and
four of green, iridescent and sometimes indistinct; body
beneath light green, with very fine grayish pubescence;
abdomen green, slightly bronzed above; general color of the
legs brownish green, trochanters lighter, second pair with a
decided yellowish pubescencé; upper half of the tibie light,
with a row of orange yellow hairs on the upper half behind;
knees yellowish; wings hyaline, halteres lemon yellow. Front °
sparsely clothed with short bristles, regularly arranged ; ocellar
protuberance considerably raised, a pair of stout macrochete
midway between the lower and two upper ocelli, a lateral row
of 2 to 3 fine hairs in the center of the ocellar triangle; a slight
depression below the lowest ocellus. Epistoma densely
pruinose, a row of bristles on the upper half in front, con-
verging with the oral margins laterally; the entire epistoma
clothed sparsely with short hairs, a row of. slight bristles
fringing the oral margin. Second joint of antenne with
numerous fine hairs, third joint destitute of a lateral pile.
Sides of the thorax with only two stout macrochete. Veins
of wings yellowish at base, darker toward the apices. Pattern
of the thoracic macrochete above showing distinctly the

156 Unwersity of Califorma Publications. [EZ NTomMoLocy

generic arrangement as given by Professor Becker in his
monograph of this family, Berliner entomologische Zeitschrift,
1896, Tafel vu, Fig. 19 (reproduced here in Fig. 105, 6). The
two hairs at the center of the field posterior are so small, how-
ever, that they might be overlooked in a casual examination.
(Fig. 105.) Length, 3.75 to 4 mm.

Egg.—-White obovate, somewhat irregular in outline, with-
out definite markings; clothed with hairs about the smaller
end, usually sparsely; attached by the larger ends to floating
bits of vegetation or puparia. (Fig. 102.)

FIG. 102. Eggs of Ephydra miilbra.

Larva.—Length, 10 to 12 mm. with the anal siphon and its
two tubes extended. Length of siphon, 3.5 mm. Densely and
uniformly pubescent, excepting a number of small, very dense
clusters of black hairs irregularly arranged on the last six
segments, dorsad. Abdominal segments with eight pairs of
false legs, non-articulated, ventral exterior end of each with
two rows of transverse, curved hooks; all but last pair with
first row containing four hooks, second five; behind these are
three or four irregularly arranged. Each leg of the last pair
bears thirteen claws in three irregular rows, the first two with
three claws each and the third with seven smaller ones. Mouth
parts composed chiefly of a pair of large median or foot hooks,
provided with smaller, hook-like processes on the ventral sur-

2 tal

Vor. 1.] Jones.—Catalogue of the Ephydridae. 157

face. Antenne rudimentary, with first lobe comparatively
long, second and third short. First segment back of the head
with a pair of fan-like spiracles or gills; the number of branches
varies from three in the young larva to seven in the adult,
directly connected with the anal siphon by the tracheal sys-

FIG. 103. Larva of Ephydra millbrx.

tem; siphon tapering, semi-transparent. When feeding, the
larva draws the food in by a process of invagination in which
the mouth parts are folded back into the head. The larva
resembles in some respects that of E. californica Pack., but
differs in the number of hooks on the abdominal tubercles and

FIG. 104. Pupa of Ephydra millbrx.

in the form and size of the anal siphon and length of its acces-
sory branches.

The puparium differs from that of EZ. halophila most notice-
ably in size, and from E. gracilis in the length of the anal
siphon, which is much longer in the latter species; it is also
much larger than #. gracilis. (Fig. 103.)

Pupa.—Puparium brownish black; length, about 12 mm.
Seventh pair of legs small. Attached by last abdominal seg-

158 University of California Publications, U2Ntomo.ocy

ment, anterior and extends into water, anal siphon protrudes.
Pupa (young) white, naked; mouth parts pressed to the breast,
indistinguishable; legs folded along the abdomen, head high
between the eyes; wings reach to tarsi of second pair of legs.
(Fig. 104.)

Note.—These flies have become very abundant along the
southwest shore of San Francisco Bay between the small towns
of San Mateo and San Bruno, the center of the colony seem-
ing to be about Millbrae, where the floating puparia and adults

FIG. 105. Ephydra milibre.

a. Head from aboye. b. Thorax from above, showing position of spines. c. End of
abdomen of female. d. End of abdomen of male. 4

often cover the entire surface of the small salt-water ponds.
Like mosquitoes, they seem unable to breed in water affected
by the tide, but prefer the smaller pools that are practically
without motion. The salinity of these marshes, owing to
eradual evaporation during the summer and autumn months,
often becomes much greater than that of the bay itself. I
have found the flies living in ponds where the salinity was as
high as 4.2 per cent, being almost one per cent higher than
that of the average sea water. The migratory propensities do
not seem to be very great and the immense colonies move about
from pond to pond only as compelled by the absolute drying
up of their habitat. The length of the life cycle is about the

Vor. 1.] Jones.—Catalogue of the Ephydridae. | 159

same as that of the salt-marsh mosquitoes, which are often
found in the same ponds. The adult flies abstract their
nourishment from the surface of the water in which their
larve live. They are especially fond of decaying animal
matter, and will collect in swarms on water containing dead
erabs or other animal bodies. The puparia are fastened in
clusters to floating bits of vegetation and some even to the anal
siphons of others. The adults crawl freely about over these
floating puparia and lay their eggs upon them. These clusters
of flies and puparia are shown in Plate I.

The puparia are very susceptible to the attacks of Chalcid
parasites, and furnish an excellent breeding ground for them.
From an aquarium in which I have bred out about seventy
Ephydrids, seven of these Chaleid flies have emerged from the

pupe.

Ephydra cinerea, n. sp., ¢ and ¢.-- Related to FL. hians
Say, but differs in the vitte of the thorax, the green of the
front and the very light color of the lower part of the legs.
Entire insect densely cinereous pruinose, giving it a gray ap-
pearance seldom seen in members of this genus. Front brassy
ereen, only slightly shining and densely pruinose; ocelli light
orange yellow, ocellar triangle with dense fulvous pruinosity ;
third antennal joint also fulvous, almost umber; eyes spotted
with black and deep orange yellow in varying proportions.

Thorax above with three broad vitte, varying from olivaceous —

at the margins to brassy green in the center; these stripes
sometimes merge into an olivaceous patch with silky luster on
the back of the mesothorax, usually obsolete on the scutellum.
Abdomen usually concolorous, sometimes becoming yellowish
toward the tip, with purplish bronze reflection, which is in-
visible except when the dense grayish pruinosity is rubbed off.
Joints of trochanters, knees, tibia, and tarsi, except last joint,
pale honey yellow densely clothed with grayish white pubes-
cence; last joint of tarsi concolorous with body or slightly
darker. <A pair of strong macrochete, as strong as the four
outwardly directed above each eye, just outside the ocellar
triangle at the center of its sides; a very small, erect bristle
just above the lowest ocellus; another, slightly larger, directly
above this at the center of the triangle; a slightly longer pair
barely inside the upper ocelli, and two more pairs directly

160 University of California Publications, [ENTOMOLOGY

beyond these back of the vertex. Front very sparsely clothed
with short erect hairs. Second antennal joint beset with short
bristles; third joint without lateral pile; arista densely pubes-
cent at the base with longer hairs, mostly above, toward the
tip and last one fourth bare. Epistoma considerably protrud-
ing, strongly arched above, a row of five to six strong lateral
bristles bordering it above on each side, usually with a smaller
bristle between each two of the larger ones; another row of
strong bristles about the oral margin. Thoracic pattern of
macrochete normal above; pleure with a row of five to six
strong bristles at the base of the wings, anterior portion usually
beset with short hairs, a strong macrocheta directly above the
intermediate coxe. All of the macrochetze and hairs black.
Claws unusually long and straight. Wings grayish hyaline;
veins light yellow at the base, darkening toward the apices;
costa beset with short, stout hairs; unusually short, stout spines
occurring at regular intervals to between the third and fourth
longitudinals. Halteres pale yellow, almost stramineous.
Length, 4.5 to 5 mm. Southern California.

EPHYDRIDAE OF THE UNIVERSITY COLLECTION.

Besides the species described above as new, I find the follow-
ing in the collection of the University of California, not here-
tofore known to occur in this region:

Notiphila virgata Coquillett. The general color is much
lighter than the type, and the vitte of the thorax are brown
rather than black. The pattern of these vittee and the corre-
spondence of the markings to the type leave no doubt as to its
identity. Bakersfield, California.

Notiphila sealaris Low. Though averaging about one
fourth mm. larger than the type described by Professor Low
from the Middle States, the specimens here correspond very
closely in color and markings. Southern California.

Hydrellia seapularis Low. This is very similar to the
type in many eases, specimens varying somewhat. Some have
a whitish dot above the antennae, and in others the mark-
ings of the legs vary slightly. A few specimens, which I regard

PLATE I. Breeding places of Ephydra miilbre.

Vor. 1.] Jones.—Catalogue of the Ephydridae. 161

as a variety of H. scapularis, have the face decidedly whitish
instead of dark ochraceous as in the type; they are, however,
identical in other respects. San Francisco and Rivera, Cali-
fornia.

Parydra appendicuiata Low. Apparently identical with
the type. Berkeley.

Seatophila hamifera Becker. Although representatives
of this genus have not before been found in this country,
there seems to be no doubt but that the specimens before me
belong here. The macrochetal pattern of the thorax, the
larger opening of the mouth, the sparse hairing of the face,
and the fact that the costal vein ends at the third longitudi-
nal point unmistakably toward Scatophila. The coloration
of the various parts, the peculiar color pattern of the thorax
above, as illustrated by Professor Becker (Berl. Ent. Zeit.,
Vol. XLI, Taf. V, Fig. 11) and the number and position of the
wing spots, mark our specimens as identical with the Norwegian
Scatophila hamifera. Alameda, California.

Caenia bisetosa Coquillett. Redlands, California.

5 LITERATURE.

The following bibliography of the family Ephydride is
primarily concerned with systematic works, but includes also
all available references to insects in recognized genera. The
method of reference carried out in the catalogue of species
which follows consists of placing the date first, then the initial
of the author, and finally the page reference. There are several
cases in which similar initials oceur at the same date, but in
every such case the page reference will enable one to determine
which author is referred to.

1776. Degeer, Carl. Mémoires pour servir a l’Histoire des Insects. VI.
Stockholm.

1782. Degeer, Carl. German translation of the above, with noces by
J. A. BH. Gotze. VI.

1792. Fabricius, Johann C. Entomologia systematica emenda et aucta,
secundum classes, ordines, genera, species, adjectis synonymis,
locis, observationibus, descriptionibus. Hafnize.

1798. Fabricius, Johann C. Supplementum Entomologie Systematics.
Hafniz.

162

1805.

1809.

1815.

1820.
1823.
1830.

1838.

1845.

University of California Publications, |! NTomoLocy

Latreille, P. A. Histoire Naturelle, générale et particuliére des
Crustaces et des Insects. .Ouvrage faisant suite aux ceuvres
de Leclerc de Buffon et partie du cours complet l’Histoire
Naturelle redigé p. C. S. Sounini. Paris. XIV.

Fabricius, Johann C. Systema Antliatorum. Brunsvige.

Latreille, P. A. Genera Crustaceorum et Insectorum secundum
ordinem naturalem in familias disposita, inconibus exem-
plisque plurimus explicata. IV.

Fallen, Carl F. Beskrifning ifver nagra i Sverige fume yvatten-
flugor. (Hydromizides. ) Handlingar JKonliga Svenska
Vetenskaps Akademien. XXXIV, 240-257.

Fallen, Carl F. Oscinides Suecie. Dissertat. Lund, Berling.

Fallen, Carl F. Hydromyzides Suecie. Dissertat. Lund, Berling.

Meigen, Johann W. Systematische Beschreibung der bekannten
europaischen zweiflugligen Insecten. VI.

Robineau-Desvoidy, Andre J. B. Essai sur les Myodaires.
Mémoires de le Savants étrang. Académien. Paris. II.

Say, Thomas. Descriptions of North American Dipterous Insects.
Proceedings, Academy of Natural Sciences, Philadelphia, VI,
149-178.

Wiedemann, Christian R. W. Aussereuropaische zweiflugliche
Insecten, als Fortsetzung des Meigenschen Werkes. II.
Curtis, John. British Entomology. being illustrations and descrip-
tions of the genera of insects found in Great Britain and

Treland. IX. London.

Haliday, Alexis H. Catalogue of Diptera occurring about Holy-
wood in Downshire. Entomological Magazine, I, 147-180.
Bouche, Peter F. Naturgeschichte der insecten ebsonders in
Hinsicht ihrer ersten Zustande als Larven und Puppen. I.

Berlin.

Brulle, Auguste. Coup d’eil sur l’Entomologie de la Morée.
Annales Science Naturelle, IIT.

Macquart, Jean. Histoire Naturelle des Insectes Dipteres.
(Suite 4 Buffon éd. Poret.) II. Paris.

Haliday, Alexis H. Descriptions of New British Insects indi-
cated in Mr. Curtis’s Guide. Annals of Natural History,
series 1, IT.

Macquart, Jean. Diptéres Exotique nouveaux ou peu connu. I.

Meigen, Johann W. Systematische Beschreibung der bekannten
europaischen zweiflugligen Insecten. VII.

Zetterstedt, Johann W. Insecta Lapponica descripta. ( Diptera.)
Lipse Revue Zodlogie, 1838, I.

Haliday, Alexis H. Remarks on the Generic Distribution of the
British Hydromyzide. Annals of Natural History, series 1,
III, 217-224.

von Heyden, Carl H. G. Ueber Insecten die an den Salien leben.
Stetteiner Entomologische Zeitung, IV, 227-229.

Stenhammar, C. Forsok til Gruppering och Revision af de
Svenske [Ephydrine. WHandlingar Kongliga Svenska Veten-
skaps Akademien, 75-273.

von Heyden, Carl H. G. Ueber Insecten die an den Salinen
leben. Stetteiner Entomologische Zeitung, V, 200-205.

Staeger, C. Groenland’s Antliater. In Kroyer’s Naturhist, Tid-
skrift, new ser., I, 369.

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1846.

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1857.

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1862.

Zetterstedt, Johann W. Diptera Scandinaviz deposita et
deseripta, Lund. V, 1739-2162.

Loew, Hermann. Dipterologisches. Stetteiner Hntomologische
Zeitung, VIII, 368-376.

Perris, Edouardo. lettre sur une Excursion dans le grande
Landes. Mémoires Académie de Science Lyon, II, 433-506.

Rondani, Camillo. Nova Species generis Ochthera. Annals de la
Société Entomologique de France, ser. 2, V, 29-31.

Loew, Hermann. Ueber die Arten der Gattung Gymnopa.
Stetteiner Entomologische Zeitung, IX, 13-15.

Walker, Francis. Catalog of the Specimens of Dipterous Insects
in the Collection of the British Museum. London. IV, 1098
et seq.

Boheman, Carl H. HEntomologiska Antekningar unter en resai
Sédra Sverige Vetensk. Acad. Hand]. 53-210.

Haliday, Alexis H. Insecta Britannica, LI.

Walker, Francis. Insecta Britannica, Diptera. London. II,
247, tabs. 11-20.

Haliday, Alexis H. Descriptions of Insects figured and references
to plates illustrating the Notes on Kerry Insects. Natural
History Review, II. Proc. 59-64.

Zetterstedt, Johann W. Diptera Scandinayis deposita et
descripta. Lund. NII.

Haliday, Alexis H. Insecta Britannica, Diptera. London. III.

Loew, Hermann. Neue Beitrage zur kentniss der Dipteren. IV,
50-57.

Rondani, Camillo. Dipterologie Italice. Prodromus. Parma.

Walker, Francis. Insecta Saundersiana; or, Characters of Unde-
scribed Insects in the Collection of W. W. Saunders. London.
V, 406.

Walker, Francis. Insecta Britannica Diptera. London. III, 352.

Walker, Francis. Characters of Undescribed Diptera in the
Collection of W. W. Saunders. Transactions of the Mntomo-
logical Society of London, series 2, LV.

Loew, Hermann. Beitrage zur kentniss der Dipteren. Berlin.
WADI:

Walker, Francis. Catalog of the Dipterous Insects collected in
Makassar (Celebes) by Mr. A. R. Wallace, with Descriptions
of New Species. Journal of the Proceedings of the Linnean
Society of London. Supplement to vol. IV, 163-169.

Loew, Hermann. Diptera Americe septentrionalis indigena.
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von Frauenfeld, George R. Jeitrage zur kentniss der Insecten-
Metamorphose. (Diptera, etc.) Verhandlungen Zodlogische-
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Egger, Johann. Beschreibung neuer Zweiflugler. Verhandlungen
in Wien der Zoblogische-Botanische Verein, X.

Loew, Hermann. Diptera Americzs septentrionalis indigena.
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tute. I, 127-172. (Smithsonian Miscellaneous Collections,
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1862. Loew, Hermann. Bidrag til) kannedom om Africas Diptera.
Ofvers WKongliga Vetenskaps Akademien Forhandling, XVII.

Schiner, J. R. Vorlaufiger Commentar zum _ dipterologischen
Theile der Faunica Austriaca mit einer nahern Begrundung
der in derselben aufgenommenen Dipterengattung. Weiner
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1863. Loew, Hermann. Diptera Americ septentrionalis indigena.
(Centuria quarta.) Berliner Entomologische Zeitschrift, VII.

1864. Bergenstamm, Julius von. Ueber die Metamorphose von Dis-
comyza incurva. Verhandlungen der Zodlogische-Botanische
Gesellschaft in Wien, XIV, 715-716.

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Loew, Hermann. Ueber die zu Durenburg beobachteten
halophilen Dipteren. Zeitschrift fiir die gesammten Natur-
wissenschaft, XXII, 346.

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Loew, Hermann. J)iptera common to Europe and America.
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Schiner, J. R. Fauna Austriaca. Die Fliegen, nach der
analytischen Methode bearbeiten, II. Vienna.

1865. Dallas, W. S. Zodblogical Record, IT, 663-664.

Loew, Hermann. Diptera Americe septentrionalis indigena.
(Centuria sexta.) Berliner Hntomologische Zeitschrift, IX.

Walker, Francis. Descriptions of the Dipterous Insects of New
Guinea. Proceedings of ‘the Linnzean Society of London,
VIII. (Jan. 13 and Dec. -5, 1865.)

1866. Dallas, W. S. Zoblogical Record, III, 512.

Frauenfeld, G. F. Zobdlogische Miscellen, VII, 2. Der Metamor-
phoses von Hydrellia albilabris. Verhandlungen Zodiogische-
Botanische Gesellschaft in Wien, XVI, 973-974.

Loew, Hermann. Diptera Americe septentrionalis indigena.
(Centuria septima.) Berliner Entomologische Zeitschrift.
2G, HD).

1868. Dallas, W. S. Zoblogical Record, V.
Packard. A. S. On Insects Inhabiting Salt Water. American
Naturalist, II, 278-829. (Also, Proceedings and Comm. of
the Essex Institute, VI, 41-54. March, 1869.)
Rondani, Camillo. Diptera aliqua in America meridionali lecta
a Prof. S. Strobel annis 1866 et 1867. Annuario della
Society dei Naturalisti in Modena, III, 32.
Shiner, J. R. Reise der osterreichischen Fregatte Novara um die
Erde. YZodbdlogischer Theil. Diptera. 242.
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(Dipteren.) Haft 12, 590-595.
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(Centuria octava.) Berliner Entomologische Zeitschrift,
XIII. :
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Berichtung naturh. Verein in Augsburg, NN, 39-59.
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lungar, no. 6, 97-105.

Loew, Hermann. Diptera Americe septentrionalis indigena.
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Rye, E. C. Zodlogical Record, IX.

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10 of Meigen’s Systematische Beschreibung der bekannten
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Loew, Hermann. Ueber die Gattung Canace. Berliner Ento-
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schrift fiir die gesammten Naturwissenschaften Neue Folge.
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Osten Sacken, C. R. Catalog of the Described Diptera or North
America. Smithsonian Miscellaneous Collections, XVI,
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California. Proceedings of the Boston Society of Natural
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Entomologische Zeitung, XIV, 137-138.

Giglio-Tos, E. Dittere del Messico. Porte Quarta. Memorie
della Reale Accademia della Scienze di Torino, XLY, 1-74.

Johnson, Chas. W. Diptera of Florida. Proceedings of the
Academy of Natural Sciences of Philadelphia, 303-540.

Becker, Th. Dipterologische Studien IV, Ephydride. Berliner
Entomologische Zeitschrift, NLI, 91-27

Coquillett, D. W. A New Subfamily of Ephydride. Entomological
News, VII, 220.

Sharp, D. Zodlogical Record, XXXIII, 290-292.

Slosson, Annie. List of Diptera from the White Mountains,
N. H. Entomological News, VII, 264.

Wheeler, W. M. The Genus Ochthera. Entomological News,
Philadelphia, VII, 121-123.

Williston, S. W. On the Diptera of St. Vincent (West Indies).
Transactions of the Entomological Society of London, 389-404,
pls. VILI-XIV.

Williston, S. W. Manual of the Families and Genera of North
American Diptera, 110.

Kuntze, A. Hine Neue Ceenia. Deutsche Hntomologische Zeit-
sehrift heraus gegeben von der Deutschen entomologischen
Gesellschaft. Berlin. 159.

Sharp, D. Zodélogical Record, XXXIV, 265-266.

Williston, S. W. Diptera Brasiliana, Part IV. The Kansas
University Quarterly, VI, 1-12 (Jan., 1897).

Townsend, C. H. T. Entomological News, IX, 168.

Coquillett, D. W. Description of a New Psilopa. Canadian
Entomologist, XX XI, 8.

Coquillett and Howard. Larva of Psilopa petrolei in Crude
Petroleum. Entomologist, 235.

Gunther, R. T. Contributions to the Natural History of Lake
Urmi, N. W. Persia, and its Neighborhood. Journal of the
Linnzan Society, Zoology. London. NNVI, 415.

Howard, L. O. Scientific American (Feb. 4th), LXNXNX, 75

Johnson, Chas. W. Catalog of the Diptera of New Jersey.
Supplement to the 27th Annual Report of New Jersey State
Board of Agriculture, 615-699.

Sharp, D. Zodlogical Record, XXXVI, 259-240.

Coquillett, D. W. Report on a Collection of Dipterous Insects
from Porto Rico. Proceedings of the United States National
Museum, No. 1198, XXII, 259-261.

Coquillett, D. W. New Genera and Species of Hphydride.
Canadian Entomologist, NN NIT, 33-36.

Coquillett, D. W. Papers from the Harriman Alaska Expedition ;
Entomological Results (3). Proceedings of the Washington
Academy of Sciences, IT,, 462.

Czereny, L. Neue esterreichische Dipteren. Wiener Entomolo-
gische Zeitung, XIX, 205.

Howard, L. O. A Contribution to the Study of the Fauna of
Human Pxcrement. Proceedings of the Washington Academy
of Sciences, II, 541-604.

168 University of California Publications, [ENTomoLocy

1900. Lundbeck, Will. Diptera Groenlandica, IJ. Nidenskaf elige
Meddelelser Naturhistoriske Forening i Kjébenhayn, 302-303.

Sharp, D. Zodlogical Record, XXXVII, 308-310.

Strobl, G. Spanische Dipteren, Part 8. Wiener HWntomologische
Zeitung, XIX, 1-4.

1901. Coquillett, D. W. Papers from the Hopkins-Stanford Galapagos
Expedition; Hntomological Results (2), Diptera. Proceedings
of the Washington Academy of Sciences, III, 377-378.

Grimshaw, P. H. Diptera Fauna Hawaiiensis, III, 49.

Hutton, F. W. Synopsis of the Diptera Brachycera of New
Zealand. ‘Transactions and Proceedings of the New Zealand
Institute, XX XIII.

Kertesz, C. Zwei Neue Wphygrobia-Arten von Singapore.
Termeszetrajzi Fuzetek kiadja a Magyar nemzeti Museum
(Budapest), XXIV, 81.

Sharp, D. Zodlogical Record, XX XVIII, 272-274.

1902. Cockerell, T. D. A. Some Insects of the Hudsonian Zone in New
Mexico. Psyche, 346-347.

Coquillett, D. W. New Acalyptrate Diptera from North America.
Journal of the New York Entomological Society, X, 182-184.

Slosson, Annie T. List of Diptera from the White Mountains,
N. H. Entomological News, XIII, $8 and’*320.

1903. Coquillett, D. W. A New Ephydrid from Australia. Entomologi-
eal News, Philadelphia, XIV, 324.

Speiser, P. Hine Neue Dipterengattung mit rudimentaren Flugeln.
Berliner Entomologische Zeitschrift, XLVIII, 65-67.

1905. Aldrich, J. M. A Catalog of North American Diptera. Smith-
sonian Miscellaneous Collections, XLVI, 623-631.

CATALOGUE AND SYNOPSES.

In giving a synopsis of this family whose members have such
a wide geographical distribution, it is often a considerable
task to bring the genera and species of the various authors into
their proper relation to each other. The pattern of the thoracic
macrocheetz has been most constantly and consistently used
by the German systematists as a distinguishing character and
has reached its perfection in the slightly conventionalized
drawings used by Professor Becker in his monograph of the
family in the Berliner Entomologische Zeitschrift for 1896,
in which many of the genera are very briefly and concisely
separated by this method. The fact that it has been largely
overlooked by many English and American writers makes it
quite difficult to define the generic limits of this family which
is so well represented in both Europe and America, and also
renders uncertain to those not familiar with the specimens the
exact relation of a new genus to those already established, not

Vor. 1.] Jones.—Catalogue of the Ephydridae. 169

to mention the difficulty of effecting a compromise between
the German and American synopses.

A genus not included here was described by Marquart
(38M411) under the name Blepharitarsis with a type species
ornatus, Which was figured in Dipteres (Planches Vol.), taf.
34, fig. 5. There was a decided discrepancy, however, between
the description and the figure, and the genus was left open to
question in the later works of Low, von Roder, and Becker,
and will probably never be settled without reference to the
original type which was found in Africa and was in the
cabinet of M. Viard when seen by Maecquart. If there is a
distinctive mark of this genus at present available it is prob-
ably the long bristles of the hind tarsi.

Another genus, Pegophila, was deseribed by Rondani (06R129)
with a type species meridionalis, but it differs from Notiphila
only in the possession of a somewhat shorter second longitudi-
nal vein, and Professor Becker does not regard this as of
generic value.

A subfamily, Lipochaetinae, was established by Coquillett
(96C220) with a single genus and species, Lipochaeta slossonae,
to accommodate some specimens taken in Florida. ‘The follow-
ing year this insect was relegated to the family Agromyzidae,
subfamily Ochthiphilinac, by Professor Williston (97W7).
Mr. Townsend, in deseribine his species fexensis, states that it
belongs to the genus Lipuchaeta, but differs from slossonae
in that ‘‘the head is even broader than the thorax, clypeus
projects fully the length of the face, front almost as wide but
usually narrower than the oral opening, scutellum one third to
one half as long as thorax, claws slender, a little elongate but
not large or stout. The third and fourth veins converge toward
their tip, but not strongly so.’’ Every one of these characters,
except the last, are given by Coquillett as generic and not
specific, and every one differs from Mr. Townsend’s species.
Suppose this insect belongs to the genus Lipochaeta, even
though the description makes this impossible, then the only
specific difference we have is the following:

“The third and fourth veins strongly, converging toward
PIMOS AM APs. ch eka Cs see ee slossonae Coq.

“The third and fourth veins converge toward their tips, but
MMA OLED LV; ASO). de ys chee Wea oo e-4 mle tya cade eo udd neem out texensis Towns.”’

2—E

170 University of California Publications. [HNtomoLoey

In a family where so many discrepancies already exist I
think we may be pardoned for not accepting one so marked as
this.

The exact limits of this family and of several others that
rank under the general terms of Calyptrate and Acalyptrate
Muscide, are certainly becoming very indefinite as new genera
and species are added, and it is to be hoped by students of
Dipterology that some authority will revise these at an early
date and that when certainly defined descriptions of both new
and old forms be published, always under the name of the
family to which they belong rather than the more general
term, under which many of them may be overlooked in giving
the synopsis of a single family.

KEY TO SUBFAMILIES.

1. Diseoidal and hind basal cells united, anal cell lacking: auxiliary
vein uniting at its extrémity with the first longitudinal ;
third antennal joint more or less flat, not rounded.......

Discoidal and hind basal cells distinct, anal cell present;
auxiliary vein separate from the first longitudinal through-

be

out; third antennal segment rounded........... Canacenae

2. Second joint of the antennze unguiculated, or at least outer side

of middle tibize with a few strong bristles....Notiphilinae

Second antennal joint not unguiculated, outer side of middle
tibis ‘without’s bristles 5. crac, ss a okhetecusieoche se eicee ee ee 43

3. Oral opening small, eyes with distinct hairing, seldom with

Mab MEM INRE SK Gk oooh oedoodduus Sabo egos Hydrellinae

Oral opening large, eyes without visible hairing....Ephydrinae

KEYS TO GENERA.
Subfamily NOTIPHILIN AE.

1. Second joint of antennze with short, spinous bristle at the

antenior Upper. | CMMs: Saicseriele rs ove lc cise eels she elehel dene le iatetetoes 2

Second joint of antennz without spinous bristles ............. a1

2. Costal vein reaches to tip of third longitudinal................ 3

Costal vein reaches to tip of fourth longitudinal..-............. 4
Costal vein reaches beyond fourth longitudinal................

Se NE eee EM GINS OOo. I ROO Ptilomyia Coquillett

3. Tip of male abdomen with elongated bristles. .Dichaeta Meigen

Tip of male abdomen without elongated bristles. .Notiphila Fallen

4. Head with deep concavity at base of antenne................

5 fo ceo a Bintan At oee i eeteele ieee nto eae Parephydra Coquillett
Head without deep concavity at base of antenne............. 5

. ‘Abdomen broad): flats jc.) Se acs ace eins smear Discomyza Meigen
Abdomen’ snot! @broad)s 35s lecwetts 5 oregano een ata me el uses reretete te 6

oO

Vou. 1.] Jones.—Catalogue of the Ephydridae.

-]

10.

ike

12.

18.

14.

. Third joint of antenns lenticular........ Discocerina Macquart

Third joint of antenne not noticeably lenticular...............
Upper side of intermediate tibize with a few long bristles......
Upper side of intermediate tibiz without long bristles.........

. Wings with a costal spine, posterior transverse vein perpendic-

ular, legs not prolonged, clypeus prominent. .Paralimna Liw
Wings without costal spine, posterior transverse vein oblique,
legs prolonged, clypeus hardly showing. .Corythophora Liw

SrAntennal saristay Naked s cc. aes se sc es cues oF Gymnopa Fallen

Abdomen sharply edged, first and fifth segments greatly short-
enedainebothySexesis sacciacec cise Trimerina Macquart
Abdomen not sharply edged, fifth segment not noticeably short-
ened, orsshortened only mathe males: .seue- see ee eee ceee
Upper part of face flat, not carinate............ Psilopa [allen
Upper part of face more or less carinate..............ee-00-
First and fifth abdominal segments shortened in the males....
All abdominal segments distinct in both sexes................
Preescutellar bristle wanting, only one sternopleural bristle... .
5 OPED eee iat EDICT ets aed Sere Hecamede Haliday
Two prescutellar bristles, two sternopleural bristles........
56 BB Ce SOG CRA RO RGAE RIPEN ew hc APP rea Allotrichoma Becker
Clypeus prominent beneath the oral margin. ..Athyroglossa Liw
Clypeus hidden or projecting but little beyond the oral margin. .
Epistoma arched; with only one lateral bristle..............
sedoS 6 Sista mote Date SE DiS OCIS eee Ephygrobia Schiner
Epistoma either slightly concave or kieled above, more than one
Let Te AMES OTIS Gl Cates semmetetestete oh chs on onsticcnkeyerty suc eli oomal ae RsEN ober a fate Gohie

. Thorax with one supraalar bristle........ Clasiopa Stenhammar

Thorax with three supraalar bristles...... Paratissa Coquillett
WS Seem Ornadltaneenpen ct werariceymers ns crcions tists, ns tals d Sree oat Mean Sar sth
Wino swsinrall lore dimentarvisens terra). cere lecietieineccis e eile ciate
Hind femora thickened and first joint of tarsi very long........
ene TMT SRS s apenas pen sts ee a eras w faveiaval sree: aveteues Amalopteryx Eaton
Hind femora not greatly thickened and first joint of tarsi normal
Abdomen decidedly arched, pointed, first longitudinal vein long,
archedmateextremityarii ae see eiinicia ciaise > Cnestrum Becker
Abdomen not decidedly arched or pointed, first longitudinal vein
ATO NIN Aes eprbarehatee sce, eas Cire cian teeawedere crehevedai'e: s6) SS/avsaeree a ort Events
Eyes haired; costal vein with two bristles at junction of first
JON Sibi ys eens ceoasheLs teesee cin Sue stereo ee Atissa Haliday
Wyves; nakeds costal without SpINeS =<) neceaie ce deloee « emee

. Epistoma with a nose-like ridge above........ Ilythea Haliday

Epistoma arched above, or without nose-like protuberance......
Hpistoma decidedly arched................. Philotelma Becker
Mpistomaenot adecidediy ssarched: ya. cea eee oie ciccicis clcissiaieic

. One small fronto-orbital bristle on each side, tibise without

bristles, oral opening small........ Beckeriella Williston
Two fronto-orbitals on each side, all of the tibise with a row
of bristle-like hairs on the outer side, oral opening of
IMOGERALETISIZE ears Sore cle cess e Seis aie eisustetie Gastrops Williston

. Wings short, scaly, halteres small............ Apetaenus Faton

Wings rudimentary, clothed with hairs, halteres of usual size... .
Uy GT OC DE DIS ORIEL oe aE aor Chamaebosca Speiser

iL:

wl

10

15

16

18
24

19

20

21

22

23

8.

We)

10.

ial

University of California Publications. (ENTOMOLOGY

Subfamily HYDRELLINAE.

Hy.es) thickly) pubescent z cpeaversskecsyeuet clad step etton et owe eeaien = pilersts
Hiyes' sparsely (pubescent. or) Dare mci) oreo cis esenerates sited -etat et = elena te
Back of thorax without evident machrochiete, antennal arista
bare or: feebly pubescent... .:........- Glenanthe Haliday
Back of thorax with distinct dorso-central machrochete........
MARCO « CONVER 2542 ba iepe sorta, ce rie ere tence Hydrellia Rob.-Desy.
Hitces slichithyacomesivierecicr cris steric eerie: Nostima Coquillett
Costal vein extending to third longitudinal...... Axysta Haliday
Costal vein extending to fourth longitudinal. 225... 2 cece
Thorax with three pairs of dorso-central and two noto-pleural
machrocheets, sides of face without bristles............
TCH ne OI IN ES OR oe Philygria Stenhammar
Thorax with only one pair of dorso-central machrocheetse and
always one noto-pleural machrocheta, sides of face with
bristles.( 9.2 sie ewes telemee mein eps oiaena te Hyadina Haliday

Subfamily EPHYDRINAHE.

Clypeus! sproyjechim eer sitter iielstnetatceteetleveromtrer eter tenene nei 56
Clypeus: hidden’. oi este oe cioc issues cious alee ovis ition snes re roere ee rereenete
Oral’ opening: “smalls cic cyse wes ete: ons aha vevcreeanieu sls sre au steus ysie e elec eneneases
Oral: -openinioy Taree nt, fas scat « cacccdeteels aw 5 Gh cane ees iesens, sv ouceeverenel otek
Front femora much thickened, first posterior cell narrowed at the

JXoredvene Cope Heavy ANI Soo aan oooocnodece b Ochthera Latreille
Front femora moderately thickened, third and fourth veins
parallel. 23ers. cksie were ise seco toe le Ochtheroidea \Villiston

ron Lemoray WO tbh CKeM Cd clea srtmerietterc late sit cp ereien siento tere nears
Hind legs long, extremity of proboscis bent back to form a hook

ws Se Boa ancl cauertoue Siebeoneles eileen e ate kare iran bene eke op sacags Ectropa NSchiner
Hind legs not lengthened, extremity of proboscis not hook-shaped
Anterior part of mesonotum without bristles....Pelina Haliday
Anterior park, Of mMeESONoOLmM! with brIStLeSia + 44m aera

Third joint of antenns with an obtuse carina above..........
a eS Or eG eanie cit oIon tho, oe.ae i Lytogaster Decker
Third joint of antenns nearly round, large...... Domina Hutton

Costal vein reaches to third longitudinal. ..Brachydeutera Low
Costal vein) reaches to fourth longitudinal. 2. =~. 25. eer oe
Oral margin with several erect bristles laterally, auxiliary vein
distinct from the first longitudinal throughout, thorax with
five distinct dorso-central machrochete..Halmopota Haliday
Oral margin without bristles laterally, auxiliary vein coalescing
with the first longitudinal except at its proximal end, thorax

with only two to three dorso-central machrochete.......
miieudhens Se segetonre sa, Slesisice ela pele aROMeL The Paahieds det siete coh R Ray Te Parydra Haliday
Claws almost straight, pulvilli indistinect...... Ephydra Fallen

Claws eunved-eniulivalln@istin Gtr peeeee sisi nek naira ren een
Costal vein reaches to third longitudinal). .3.6. 2 «sce. eae ve
@ostal vein reaches to fourth longitudinale .-.55-...- 0-4-2. e8
Thorax with only a pair of humeral machrochets in front....

Te Se TT TA RIC eG ONT CLO OE o ,---...Scatophila Becker
Thorax with four rows of bristles extending to the anterior part

ausoy\os So os vats Ts uscoae TNETs sat leiwtelis aishe Myst amen meno NsReR meets Pelomyia Williston

me bk

10 Oo Ww

6

10
11

12

Vou. 1.] Jones.—Catalogue of the Ephydridae.

12. Antennal arista bare, first antennal joint forming an angle with

vs SCrexonael Hal selovidls GaSe an aneocoe< Teichomyza Macquart
Antennal arista not bare, segments not forming an angle.......

15. Antennal arista pubescent............- Scatella Rob.-Desvoidy
Nomigraanll, iene yoycummenle a Gowanonnocues cn0 0000 sno oun OR
ieeeeHemora Wot icrassated......4-- <> «1... Caenia Rob.-Desvoidy

Femora slightly incrassated, metathorax elliptical, enormously
developed, covering whole abdomen, legs short, stout, tibise
curved, wings concealed beneath metathorax..Nomba Walker

Subfamily CANACENAE.

1. Hind basal and anal cells distinct, auxiliary vein separate from
aiesye kovaveat byahbavatllic ces oucidlicloct ClnicaroRCienons Cech oir Canace Haliday

NOTIPHILINAE.
PTILOMYIA Coquillett. Type enigma.
enigma Cog. O0C262, OOS309, O5AG24. Porto Rico.
DICHAETA Meigen. Type caudata.
Norra AMERICAN SPECIES.

1. Last abdominal segment comparatively short and blunt....

ao eS tao ASD DIO Oe EG eae brevicauda Low.
Last abdominal segment prolonged into a conical point or
HELD CREM OM apee ted aa Nat eriees cats sy Stes oases ccehe yee ehecer er shcnsie Breen auc

2. Mesonotum with three indistinct brown vittse............
ERSTE Cheese PLm at ene mice ante ne bate wh ahisepanal eels furcata Coquillett.
Mesonotum with one brownish and four whitish gray viltie
Sieg hb: COSTER SEE ASTOR OLS IDI ane a anor eee erie caudata Wallen.
brevicauda Lw. 621133, TSOS8200, 951388, 9615268,

05A623. Middle States, New Jersey, Florida.
caudata Fall. 620133, T8OS200, 96B268, O2S8, 05AG623.
Middle States.
furcata Coq. 02C182, 02S2638. Florida.
ISUKOPEAN SPECIES.

2. brevicauda Inv. G60L5, 648236, 96B108. Silesia.

2. caudata Tall. 30M62, 448194, 538W251, 56R129, 6OL5,
648236, 9613103, Notiphila 23F8, 38Z717, 46Z1853. Whole

of Hurope.
tibialis Brule. 52M318, 96B104. Europe.

NOTIPHILA Fallen.

Norri AMERICAN SPECIES.
eC eaterapancsor armbkemmecs yellows csc % su <ists oceehcrve ie suctel shane oiefltors
Third jont of antennee yellowish at base. ..decorata Williston.
Third joint of antennee reddish or reddish yellow at base... .
Greater parc Ot antennsesneddish=a.-- eee cekes aim ella ceenelersne
Antenne black throughout, or at least with third joint
BO watts Ey arege one crs citecncdone oxecsse (oct cecnertine Sie bee eiekelene terete cack ae os

174

University of California Publications, U2NToMOLocY

NortTH AMERICAN SPECIES—Continued.

Sp

9.

10.

Pronotum with a median brownish vitta......... varia nN. sp.
Pronotum without such) vittarye-c esse). virgata Coquillett.
Palpi ‘blackish’s ics: fi) stealer cre exes oes fo Geet pulchrifrons Low.
Palpi.- yellowish: isiscie che etercitteauete sisne sole yo shonsiiseclia fe Clekelomtenel holopeens
Upper side of thorax with broad brown lateral stripe........

Dinca 5 aoe STONE HERRON OS ioatar ay snes elekeroustersaatee vittata Low.
Wpper side ‘of thorax without such Striper .crrcre eet tts) eert =
Abdomen with irregular brownish semifasciz. each formed

DY EW! SDOUSis storey elses ste cieiereeieteuciete ot ekea,akegiotane carinata Low.
Abdomen with two brown spots on each of the intermediate

SCCIMENUS) eeu enrocr stone rete ne hemor etek or ten ie unicolor Low.
Abdomen dull, but without brown spots.......... avia Low.
Palpi- blackistr. =n te veten- ko otel oy heyepore tenets oeuay =e solita Walker.
Palpiereddishisy.ell owsereencnctretersictieth-ia- tone erythrocera Low.
Pallpiy liohitsy ellowi terete ete cies -feleteieierenerarerear bellula Williston.
Palpimyellowishis soc esha ct oelerenesy eles «kere tslieonelone cee bella Low.
Palpis blackishy peperrnleserete shale ctersse ects cusconener se suerexcne neuen tense
Front with two black stripes separated by the ocellar triangle

ee ity Sort Oho RO acre Ola Gore decoris Williston.
Hront without: such Stripes. «ace octal e sty a seen caeisielene
Hind metatarsi with a strong black bristle on the under side

Teen Sasigs asahe Cetera oo orto we dM ve to notaeu cP oes lene ees macrochaeta Low.

Hind metatarsi without such bristles.......
Wings slightly notched at the end of the first oneccadinane

Shee IE Wehave ce, ole Sees DLO e ee eekeeimen se scsune producta Walker.
Wines: swithout suche moOtehesrrvreie eieietet oe etelei= ere) = ctenen-teit terete
Chest with: two gray Stripes)... ..cs 6s. transversa Walker.
Chest avithoutesuChie Strip cStereiesteveerrevercteneteicr ath teenie neers

Upper side of thorax with five fine brown lines. ..scalaris Low.
Upper side of thorax with fewer or no vitte. F
PICO rca rR cic omrco MO CIO COO quadrisctosu homison:
argentata. See Brachydeutera.
avia Lw. 781193, 7T80S200, 7T8K244, 96B268, 05:A623.
Hudson Bay Territory.
bella Lw. 621133, T8OS200, 96B269, 05A623. Middle
States.
bellula Will. 96W390, 968291, 054623. -—St. Vincent.
brevicornis. See Hyadina nitida.
carinata Lw. 621137, 7808200, 96B269, 05A628. Middle
States, New Jersey.
decorata Will. 96W389, 968291, 054623. St. Vincent.

decoris Will. 93W258, 938231, 054623. Panamint Valley’

(California).

erythrocera Lw. 781194, 78K244, T8OS201, 96B269,
05A623. Cuba.

gquttata. See Hyadina.

guttata var. brevicornis. See Hyadina nitida.

macrochaeta Lw. 781192, 7T8O0S200, 7S8IX244, 96B269,
05A623. Texas.

producta Walk. 49W1099, 7808201, 96B266, 05A623.
Hudson Bay.

pulchrifrons Lw. 721102, 72L84, 72R389, 7808200,
96B269, 9TW5d2,. ODAG23. Texas.

e0

10

11

Vor. 1.] Jones.—Catalogue of the Ephydridae.

NortH AMERICAN SpEeciIES—Continued.

quadrisetosa Thoms. 681594, TOV442, 7T8OS8200, 96B271,

05A623. California.
repleta. See Scutella.

scalaris Lw. 621134, 7T80S200, 96B268, 05A623.

States, New Jersey.

solita Walk. 56W406, 7808201, 96B266, 05A623.

States.

Middle

United

transversa Walk. 56W407, 7808201, 96B266, O05A625.

United States.

unicolor Lw. 621137, 64D559, 7808200, 96B268, 054623.

Middle States.
Varia nN. sp.

virgata Cog. 00C259, 008309, O5AG625. Porto Rico.
vittata Lw. 621136, 651134, 7808200, 96B268, 0546238.

Middle States, New Jersey.

ISUROPEAN SPECIES.
albicans. See Hecameda.
albifrons. See Hydrellia nigricans.
albilabris. See Hydrellia.

annulipes Stenh. 148203, 60L7, 6482388, 96B113.

dinavia, Germany, Silesia.

Scan-

aquatica Beck. 9618114, 968291. Silesia, Poland.
australis Lw. 6017, 648239, 96B110. Germany, Greece,

Asia Minor, Italy.
caesia. See Hydrellia.
caudata, See Hydrellia.
chalceata. See Clasiopa.
chamaeleon Deck. 96B114, 968291. Pavia.

cinerea Fall. 13F250, 23F8, 30M64, 35M521, 382717,
448200, 4671855, 60LT, 648259, 96B112, Keratocera

palustris BORD788. Europe.
compta. See Ephygrobia.
concolor. See Hydrellia.
discolor. See Hydrellia.

dorsata Stenh. 448198, GOL7, 648239, 96B113.

navia, Germany, Silesia.
erythrostoma. See Hydrellia nigripes.
flaviceps. See Hydrellia discolor.
flavicornis. See Hydrellia.
fulviceps. See Hydrellia.
fusca. See Hydrellia.
genicula. See Hydrellia.
‘glabrata. See Ephygrobia.
glaucella. See Clasiopa.
grisea. See Hydrellia.
griseola. See Hydrellia.

guttiventris Stenh. 448206, 4621860, 60L7,

96B111. Scandinavia, Germany.
incana. See Hydrellia ranunculi.
incurva. See Discomyza.
interstincta. See Philygria.
laticeps. See Hydrellia.

Seandi-

6482389,

175

176 University of California Publications, U2NTOMOLoGY

[EUROPEAN SPECIES—Continued.

maculata Stenh. 448201, 46Z1862, 6OL7, 648239, 96B111.
Scandinavia, Germany, Silesia.

madizans. See Trimerina nigella.

major Stenh. 448196, 46Z1857, 96B111. Sweden.

mutata. See Hydrellia.

nigrella. See Psilopa.

nigricauda. See Philygria.

nigricornis Stenh. 448202, GOLG, 6482387, 96B109. Scan-
dinavia, Germany, Austria.

nigripes. See Hydrellia.

nymphaeae. See Hydrellia.

obscurella. See Clasiopa.

picta. See Philydra.

pilitarsis. See Hydrellia.

plumosa. See Clasiopa.

punctinervosus. See Philygria.

punctipennis Wied. S0W590, 96B266. Hurope.

riparia Meig. 380M65, 35M522, 448204, GOLT, 648255,
96B112. Scandinavia, France, Hngland, Germany,
Silesia, Asia Minor.

stagnicola Stenh. 385M522, 448197, 4621854, GOLG6, 648259,
96B109, Keratocera 830RD789. Scandinavia, Germany,
Silesia.

taursata. See uliginosa.

uliginosa Halid. 89H922, GOLG, 6482387, 96B109, tarsala
448207. England, Scandinavia, Austria.

venusa Lw. 56L55, 60L7, 648238, 96B112. Scandinavia,
Germany, Austria, Silesia.

vittipennis. See Philygria.

OTHER COUNTRIES.

albiventris Wied. 30W589, 96B266. East Indies.

alboclavata Bigot. 88B41, S8S295, 96B272. Cape Horn.

bipunctata Lw. 62112, 96B268. Swakop.

brasiliensis Walk. 56W408, 96B266. Brazil.

carbonaria Walk. GOW169, 65D663, 96B268. New
Guinea. :

chinensis Wied. 30W592, 96B266. China.

ciliata v. d. Wulp. 81W55, 81K253, 96B271. Sumatra.

costalis Walk. 56W408, 96B266. Brazil.

difficilis Wied. 30W591, 96B266. South America.

dorsopunctata Wied. 30W591, 91W215, 928201, 96 B266.
Hast Indies.

exotica Wied. 30W590, 96B266. Montevideo.

fasciata Wied. 30W589, 96B266. East Indies.

flavilinea Walk. 60W170, 96B268. Celebes.

granifera Thoms. 6817594, 96B271. Insula Rossi.

ignobilis Lw. 62112, 96B268. Cape of Good Ilope,
Swakop.

immaculata Wied. 30W592, 96B266. China.

indica Wied. 30W591, 968266. Hast Indies.

insularis Grims. 01G49, 0182738. Oahu (Sandwich
Islands).

Vor. 1.] Jones.—Catalogue of the Ephydridae.

OTHER CoUNTRIES—Continued.
lineosa Walk. 6O0W170, 96B267. Celebes.
obscuricornis Ly. 62112, 96B268. Swakop.
ortaloides Walk. 60W169, 641D559, 65D663, 96B268.
Mysol.
peregrina Wied. 80W592, 96B266. China.
2. pulchrifrons Will. 97W5, 9782638. Brazil.
quadrifasciata Walk. GOW170, 96B267. Celehes.
radiatula Thoms. 687595, TOV442, 96B271. China.
sinensis Schin. 688241, 68D374, 83K251, 961B273.
Hongkong.
smaragdi Walk. 49W1098, 96B266. Sierra Leone.
sternalis Thoms. 6817593, 7TO0V442, 96B271. Manila.
striata Will. 97W5, 978265. Brazil.
triangulifera Schin. 6858241, 68D374, 96B273. South
America.
unicolor Walk. 60W169, 96B268. Mysol.
unilineata Walk. 60W169, 65D663, 96B268. New Guinea.

~

bo

PAREPHYDRA Coquillett. Type humilis.
humilis Cog. O2C183, 054626.

DISCOMYZA Meigen. Type incurwva.

Norrim AMERICAN SPECIES.

ieeeAmtenn se) VellOwiSMy PEWsre 4 ccc) « wietee s:2 <2 eicic oe balioptera Low.

PNG GTI 2 el eC Ky taeevenemnvstel cio Reis /atecltot se fotieyeyenisv'el'e) <6 dubia Williston.

balioptera Lw. 621140, 7TS8OS201, 96B268, O5A624.
Cuba.

dubia Will. 96W392, 96S290, 05A624. St. Vincent.

ISUROPEAN SPECIES.
cimiciformis Halid. 3S8H124, 88G99, 888295, 96129.
Germany, Ireland, Hast coast of Hurope.
incurva Fall. 3OM76, 35529, 448265, 60L8, 64167135,
64D559, 82B14, 821K 252, 96B129, Psilopa 28F6, 4671941,
Notiphila 648241. Germany, North and Middle Europe,
Britain.

OTruHEeR COUNTRIES.
amabilis Kertez. O1IK421, OIS272. Singapore.
obscurata Walk. 60W169, 96B267. Philippines.
pelagica Frnfld. 619451, 968275. Nikobara.
punctipennis yv. d. Wulp. 81W56, 81K253, 96B271.
Sumatra.
tenebrosa Walk. 60W169, 65D663, 96B267. New Guinea.

DISCOCERINA Macquart. Type lewcoprocta.

NortH AMERICAN SPECIES.
HeOnbit OL eyes ihe) SHUMINS. 6 «cece deci o eiclers «ie orbitalis Liw.
Oxrbit.of eyessnot shining, white... .2.....5-220-.- 25+ emer ne
Pee Hdomen plackwatileast abt DAS. cusc oc cls oleic 2 cleicle sles: eele 6
PNG OMeRM eT OGab Ss Cheer cis chia cd ccs c-ctaceseichctsns: sisters) eyes sleet evelchareiers

os |

Dw iv

178

Unwersity of California Publications, |ENtomMoLocy

NortH AMERICAN SPECIES—Continued.

»
o-.

4.

fxd) |

-]

Last segment of abdomen white or silvery gray..............
last segment of abdomen nogewhite. ..-) hci see ceeeeeeeene
Last segment and front angles of the two preceding segments of

abdomen whitish pruinose.:............. incisa Coquillett.
White confined entirely to last segment of abdomen........

sighs Bibs knw teein, stolen sects ae twa eee SOLE soaet ete RRnT ae leucoprocta Low.
Face silvery gray with a medium black stripe. .nana Williston.

Face vaulted, oral opening very large...... facialis Williston.
Oral opening of moderatersizesee ce cee eee eee ie ieee
Thorax black, shining through brownish dust..............
SU ROT IE, ea cave pohetene te oe RCRO ROE Rene oes tete i eee obscura Williston.
Thorax ashi seray. Opaques. vas 44 acieeiee ie siete eee parva Low.
Antenne reddish yellow, third joint a little infuscated on the
apical “maroing pee ei eo ereoee lacteipennis Low.
Antenne black, a whitish pollinose dot on the upper edge of
the:'second joint. oS eicisle os cue bivtelee oem eieterenrs simpler Low.

calceata. See Clasiopa.

compta. See Ephygrobia.

facialis Will. 96W396, 968290, 05A626. St. Vincent.

incisa Cog. 02C182, 028263, 05A626. Porto Rico.

lacteipennis Lw. 621145, 7T8OS8201, 96B268, 054626.
Washington (1). C.), New Jersey.

leucoprocta Lw. 611355, 621.148, 7808201, 966269, -

96W395, 00C261, 054626. Maryland.

nigritella. See Ephygrobia.

nana Will. 96W396, 968290, 05.4626. St. Vincent.

obscura Will. 96W397, 968290, 054626. St. Vincent.

obscurella. See Clasiopa.

orbitalis Lw. 611354, 65L91, 621147, 7808201, 968369,
05A626. Washington (D. C.).

parva Lw. 621146, 78OS201, 96B268, 000261, 00H592,
05A626. Washington (D. C.), Florida.

pulicaria. See Clasiopa.

simplex Lw. 611354, 621147, TSOS201, 96B569, 054626.
Maryland.

PARALIMNA Liw. Type appendiculata.

NortH AMERICAN SPECIES.

ie

“

Hace brownisheeray seein eae cients obscura Williston.
Mace*erayish “whites s1,. cs nce oa lo cis oe ow one lettacoe cis one eneee pe each otene
Face. ‘yellowish white. 2 Geran acre cee cone ence eral emia temo net none
First joint of front tarsi at base and first joint of hind tarsi

LOA, cays osig oils Aw atstanstanencititecr ened tiewenete telenors appendiculata Low.

First joint of front and hind tarsi concolorous or yellowish
aicares Ero Ma eo Mtetomeel saree ae eR RTS Ra ere multipunetata Williston.
Thorax black, antenne black throughout....... decipiens Low.
Thorax with grayish pruinosity, third antennal joint yellow
PRR Te a IC Nae foes) ndiricco Oke Seno to. nuda Coquillett.

appendiculata Lw. 621188, T8SOS201, 95.358, 96B268,
05A624. Middle States, New Jersey.

G

oO

)

-1

bo

ae

~

wy

Vor. 1.] Jones.—Catalogue of the Ephydridae. 179

NorTH AMERICAN SPECIES—Continued.

decipiens Lw. 78L195, 7808201, 78K 244, 96B270, 000259,
05A624. Texas, Georgia, Florida.

multipunctata Will. 96W390, 968291, 05A624. St. Vin-
cent.

nuda Coq. O2C182, 028264, 05A624. Mexico.

obscura Will. 96W391, 968291, 054624. St. Vincent,
Porto Rico.

EUROPEAN SPECIES.
2. appendiculata Lw. 6211388, 96B115. Germany.

OTHER COUNTRIES.

albonotata Lw. 62118, 96B268. Caffraria.
confluens Lw. 62118, 96B268. Caffraria.

javana v. d. Wulp. 91W215, 928302, 96B271. Java.
limbata Lw. 621138, 96B268. Caffraria.

picta Kert. 01K423, 018278. New Guinea.

CORYTHOPHORA Liw. Type longipes.
longipes Lw. 62113, 96B116, 968290. Caffraria.

GYMNOPA Fallen. Type subsultans.

aenea. See subsultans.

albipennis Lw. 48014, 96B127. Germany, Messina.

nigra. See subsultans.

subsultans Fabr. 48L13, 56R128, 648254, 96B127.
nigra 30M137, aenea 23F10, 30M187, Ulida 4622679,
arcuata 35M505, Syrphus 1798F304, Mosillus arcuatus
1809L389, Glabrinus mororum 56R132. Hurope.

TRIMERINA Macquart. Type Psilopa madizans Meig.
nigella Meig. 60L8, 648240, 94B117, madizans 35M529,
Notiphila 30M72, Psilopa 23F7, nigella 448263. Ger-
many, North and Middle Hurope. ;
tibialis Macq. 35M528, 96B117. Europe.

PSILOPA Fallen.

NortH AMERICAN SPECIES.

1. Entire legs, including the cox, yellow. ...mellipes Coquillett.
Wessenotentinely syellowi acre(s 2s eles clele etleke t=icerey ile) ois)+le)l'-) o)ie)- 2,
». Thorax black anteriorly, posterior part scoriaceous........

SP SF Ne oor ret Ne ao oes is) Se aliases ahaa Woe ab et aligt’ lat'elis scoriacea Low.
Mesonotum of thorax deep brown......... desmata Williston.
Mesonotum and scutellum subopaque, slightly scabrous, thinly

Famiy (ews. onaoacogsuceos poospenduoaG similis Coquillett.

Mesonotum and scutellum thinly yellowish dusted.........

Ave SBS CD AO OTE SOD DCO SET ECR Une a ae nigrimana Williston.
Mesonotum and scutellum yellowish, usually polished......

ee Ieee ttn dhs, Salis ay cn gr ew anitanans, @ 6 flavida Coquillett.
Mesonotum and scutellum gray or dusted with gray.........
Mesonotum and scutellum green or tinged with green.......
Mesonotum and scutellum black..............2-0s--eeeeeee

cS es) ae)

180 University of California Publications, |ENtomMoLocy

NortH AMERICAN SPECIES—Continued.

3. Abdomen’shining steel blue.............. cocruleiwentris Low.
Albdonien. blackinc sas ccs eecere ete: chore a tere Oe cae i otenete {
{eAntenns Teddi Shueyellowepeeiieienetsie i is eee umbrosa Low.
‘Anitennse’ Vblack sc alstevs ureters tote mie ousitie me <lousiahars nigra Williston.
ey AB Aoryen geal ryWoligrorn gro rt cacoaA sco Moat 5 cola c nobilis Low.
Elypopyeium mot winter 225 fee a sai a) sone caso e eed oaetsweme ais) echoes G
6. Hirst joint of anterior tarsi white........... pulchripes Low.
First joint of anterior tarsi not white...... caripes Coquillett.
iT. “Rhorax cinelyacicul ates: ccm camer ecitenacicherere aciculata Low.
Thorax not tnely -aciewlate.,. «miss acc cece erect 5
S. A minute blackish spot at the tip of the third vein..........
Renee ern eB Monn AaN eR ary Seat heme ale Ati oe nigropuncta NVilliston.
No black spotratutipmon chum dimeimns -rteccrte ec scetetenc cri acetenenens 9
9). Halteres’ white; petiole blackish =. «1 lee reais atra Low.
Halteres white, petiole yellowish.......... petrolei Coquillett.
Petiole’ of halteres concolovoussie. = 22 <1eleleist ey = ee eeeeenen one 10
10: Hntire abdomen shinine jbhlackzs sao sce erertesn nitidula Wallen.
Abdomen with a greenish reflection........... atrimana Low.

Abdomen black, dull toward the tip.........aenea-nigra Low.

aciculata Lw. 621142, TSOS201, 96W394, 96B268, 9TW4,
00C260, O5AG24. Cuba.

aenea-nigra Lw. 7TS8L196, 78K244, 7TSOS201, O5A624.
Texas.

atra Lw. 621143, 7T8OS201, »v6GB268, O5AG24. Middle
States.

atrimana Ly. T8L197, TS8IK244, TS8OS201, V05A625.
Texas, New Jersey.

coeruleiventris Iw. 620144, 98201, 96B268, OF5AG25.
Cuba.
desmata Will. 96W295. 968291, O5\625. St. Vincent.

flavida Cog. 00C33, 008310, 05.4625. Massachusetts.

mellipes Coq. O00C260, 05A625. Porto Rico.

nigra Will. 96W393, 968291, 05A625. St. Vincent.

nigrimana Will. 96W398, 968296, 97W4, 000260,
ODAGZS. St. Vincent, Porto Rico.

nigropuncta Will. 96W393, 968291, O5A625. St. Vin-
cent.

nitidula Fall. 181252, 054625, Notiphila 49W1089, Lphy-
grobia 96B266. Martin ralls (Canada).

nobilis Lw. 621229, 72L92, 7508201, 05A625. District of
Columbia.

petrolei Coq. 99C8, 99C&H235, 99H7T5, 998240, 05A625.
California.

pulchripes Lw. 7SL197, TSK244, T8OS201, O5A625.
Texas.

scoriacea Lw. 620,142, T8OS201, 96B268, 054625. New
York, New Jersey. :

similis Coq. 00C33. 008310, 054625. Florida, Louisiana.

umbrosa Lw. 621145, T8OS201, 96B268, 054625. Cuba.

varipes Coq. 90033, 008310, 054625. British Coiumbia.

HUROPEAN SPECIES.
apicalis. See Ephygrobia.
compta. See Ephygrobia.
girschneri. See Ephygrobia.

Vor. 1.] Jones.—Catalogue of the Ephydridae. 181

EXUROPEAN SPECIES—Continued.

lewcostoma. See Ephygrobia.

nigrotaeniata Bezzi (Diasemocera new subgenus). 95B137,
958344. Italy.

madzgans. See Trimerina nigrella.

marginella. See Ephygrobia.

maritima. See Ephygrobia.

nigrella. See Trimerina.

nitidula Fall. 188252, 28F7, 448261, 4621932, Hydrellia
Notiphila 49W1098, 60L10, 89156, Hphygrobia 648242,
96B143. Silesia, South Russia, Sarepta. See also
Ephygrobia compta.

obscuripes. See Ephygrobia.

polita. See Ephygrobia.

rodent. See. Ephygrobia.

tarsata. See Ephygrobia.

OTHER COUNTRIES.
metallica Schin. 688242, 97W4. Brazil.

HECAMEDE Haliday. Type albicans.

abdominalis. See Allotrichoma.

albicans Meig. 39H224, 53W254, 56W344, 60113, 96B121,
Notiphila 30M65, 3874760, Clasiopa globifera 52B204.
Coast of Europe.

aurella Strohl. 938256, 938321. Styria.

glaucella. See Clasiopa.

vanthocera. See Clasiopa.

ALLOTRICHOMA Becker. Type lateralis.
Norrie AMERICAN SPECIES.
abdominalis Will. 97W4, OOC260, 05A624, Hecamede
9GW398. St. Vincent.
HUROPEAN SPECIES.
bezzi Beck. 96B123, 968290. Italy.
filiformis Beck. 96B1238, 968290. Sarepta. Italy.
lateralis Lw. 60113, 96B122. Italy, Sicily.
trispinum Beck. 96B124, 968290. Silesia, Malta.
OrTnER COUNTRIES.
2. abdominalis Will. OO0C260. Brazil.

ATHYROGLOSSA Loéw. Type glabra.

Norrim AMERICAN SPECIES.

1. Tarsi white, except the last segments, which are brown......

ORCS cre Vser ORS ee ions vacioie ans speyeisliel. als glaphyropus Low.
INST S cI O bee WLLL praerevres ctrtierreyl a atle a, wrotraeveiiel syayoucpsehporet ait, «tos fore orcerebotay ere) © 2

PPE VEIITES! iver Chis cee ettarensc asta. eecs.einy aver sl evafersle cies glabrata Meigen.

Wate ieieryetdl syatdil Jato ooo oecuosucooa > nitida Williston.

glabrata Meig. 56W254, 60112, 648254, 968134, 05A625,
Notiphila 30M69. Washington.

glaphyropus Lw. 78L198, 78OS202, 78K244, 96B270,
05A626. Texas.

nitida Will. 96W397, 968290, 00C260, 054626. St. Vin-
cent, Porto Rico.

182 Unwersity of Califorma Publications, [ENTOMOLOGY

EUROPEAN SPECIES.
glabra Meig. 30M69, 53W254, 601.12, 648245, 96B134.
Germany, Silesia.
nudiuscula Lw. 73L307, 738R426, 73L50, 96B134.
Hungary, Kasan.
ordinata Beck. 96B135, 968290. Orsova.

EPHYGROBIA Schiner. Type nitidula.

EUROPEAN SPECIES.

apicalis Perris. 648242, 96B1388, Hydrellia 47P494, Psilopa
60L9, 89R56. Germany, France.

compta Meig. 30M68, 648243, 96B143, Discomyza var.
B. nitidula 448261, Psilopa 46Z1932, Hydrellia compta
35M524, Psilopa 60L56, 89R56. Europe, Silesia.

girschneri v. Rod. 96B141, Psilopa 89R55, 89S287.
Fundort, Saxony, Poland.

leucostoma Meig. 648243, 96B142, Notiphila 30M68,
Hydrellia 35M524, Psilopa 448261, 46Z1936, 60L9, 89R56.
Sweden, England, France, Germany, Hungary, Russia.

marginella Fall. 96B1389, Psilopa 23F7, 46Z1939, 55Z4762,
Discomyza 448266. Europe.

maritima Perris. 96B139, Hydrellia 47P494, Psilopa
73L306, 89R56. France.

nana Lw. 96B142, Psilopa 60L9, 89R56. Constantinople,
Sylt Island.

nigritella Stenh. 648242, 96B144, Discomyza 448262,
Psilopa 46Z1935, 60L10, 8S9R56. Scandinavia, Germany,
Austria, Silesia.

nitidula. See Psilopa.

obscuripes Lw. 96B144, Psitlopa 60L10, S9R56, 948124.
Greece, Asia Minor, Hungary.

plumosa. See Clasiopa.

polita Macq. 648243, 96B144, Hydrellia 37M524, Psilopa
60L10, 89R56, tarsata 4621934. North and Middle
Europe, Silesia.

roderi Girsch. 96B140, Psilopa 89G873, 91T244, 918275,
Diasemocera nigrotaeniata 95B137. Fundort, France,
Italy. :

OTHER COUNTRIES.
metallica Schin. 688242, 69D374, 96B273. South
America.
nigricauda Bigot. 91B278, 918275. Canary Islands.
pollinosa Kert. O1K81, 018273. Singapore.
singaporensis Kert. 01K81, 018273. Singapore.

CLASIOPA Stenhammar.

HKUROPEAN SPECIES.
aurifacies Strobl. 9858255, 93883820, 96B157. Styria,
Silesia.
aurivillii Beck. 96B158, 968290. Sweden.
bohemanni Beck. 96B159, 968290. Sweden.
brevipectinata Beck. 96B149, 968290. Norway.

Vor. 1.] Jones.—Catalogue of the Ephydridae. 183

FuROPEAN SPEcIES—Continued.

calceata Meig. 64S244, 96B157, nigrina 448254, 46Z87,
Notiphila 30M69, Discocerina 35M524, 60L11, var.
flavoantennata OOS1, OOSS08. North and Middle
HKurope, Silesia.

cinerella Stenh. 448251, 96B158. Germany, Sweden.

costata Lw. 60114, 96B160. Turkey, Asia Minor.

coxalis Strobl. 938253, 938320, 96B157. Styria.

dimidiatipennis Strobl. 938255, 988320, 96B150. Styria.

duplosetosa Beck. 96B162, 968290. Malta, Orsova.

fulgida Beck. 968156, 968290. Orsova.

glabricula Fall. 448256, 648244, 96B152, WNoliphila
138251, 23F10, 46Z1898. Silesia.

glaucella Stenh. 448253, 96B160, Notiphila 46Z1885,
Hecamede 60L14, 648245, 948126. Hungary, Sweden,
Styria, Silesia.

globifera. See Hecamede albicans.

nigerrimana Strobl. 938254, 938320, 96B151. Styria.

nivea Beck. 96B151, 96S290. Malta. Silesia.

niveipennis Beck. 96B162, 96S290. Silesia.

obscurella Fall. 448254, 648244, 96B148, MNotiphila
18F251, 23F10, 30M73, Discocerina 60L11. North and
Middle Europe.

olivacea Beck. 96B155, 968290. Herkulesbad.

palliditarsis Beck. 96B155, 968290. Silesia.

pallidula Stenh. 448257, 96B150. Scandinavia.

plumosa Fall. 60110, Hphygrobia 648242, 945125,
96B150, Psilopa 23F9, 55Z27, longula 448259, Notiphila
30M73. North and Middle Hurope, Silesia.

pulicaria Halid. 53W254, 96B155, fuscella 448256,
46Z1893, Discomyza 39H224. Hurope.

xanthocera’ Lw. 96B161, Hecamede 69L58, awrella
938250. Germany, The Alps, Silesia.

OTHER COUNTRIES. : :
albitarsis v. d. Wulp. 81W56, 811K 253, 96B271. Sumatra.

PARATISSA Coquillett. Type pollinosa.
pollinosa Will. 00036, 05A626, Drosophila pollinosa
96W404. Florida, St. Vincent.

AMALOPTERYX Eaton. Type maritima.
maritima Haton. 75E58, 96B272. Kerguelen Islands.

CNESTRUM Becker. Type lepidopes.
lepidopes Beck. 96B118, 968290. Germany, Silesia.

ATISSA Haliday. Type pygmaea.
durrenbergensis Lw. 641346, 64D512, 968131. Thur-
ingen.
limosina Beck. 96B132, 968290. Norway, Christiania.
pygmaea Halid. 33H174, 53W258, 648251, 89G223,
898287, 96B131, ripicola 6(0L24. Germany, Ireland, Italy.
ripicola. See pygmaea.

184 University of California Publications, [ENtTomoLocy

ILYTHEA Haliday. Type spilota.

NortH AMERICAN SPECIES.
1. Legs brownish black with yellow rings at the knees..........

AHO nis Sous oo So one Oro iS acc spilota Curtis.
Legs yellow, the tips of the tarsi brownish. . flavipes Williston.
flavipes Will. 96W4038, 97W4, 00C260, 05A625. St.
Vincent, Porto Rico.

?oscitans Walk. S7W233, TSOS262, 96B268, O0C2G60,

O5AG625. United States.

See Ephydra and Scatella.

spilota Curt. 60L37, T8OS204, 05A625. North America. —

EUROPEAN SPECIES.

2. spilota Curt. 3820418, 53W264, 648263, 96B135, notata
448186. North and Middle Hurope, Silesia.

PHILOTELMA Becker. Type anomala.
anomala_ Beck. 96B164, 9GS291. Kkohlfurter Moor
(Silesia).

BECKERIELLA Williston. Type bispinosa.
bispinosa Thoms. 97W2, 978265, Hphydra_ bispinosa
681593, 70V442, 96B271. Rio de Janeiro.

GASTROPS Williston. Type niger.
1. Antenne red, third joint at tip and on upper part black... .

SSO Cra MOC pOnc adh coon SuiaTo enor niger Williston.
Antenne, except upper edge and sometimes broad apex of third
Joint yellow. akrne s oe sae ieee Giawie a ee nebulosus Coquillett.

nebulosus Coq. OO0C34, O5A624. North Carolina, Georgia.

niger Will. 97W3, 97S265,, O5A624. Grenada (W. I.),
Rio Janeiro.

APETAENUS Haton. Type litoralis.
litoralis Eaton. 75E58, 96B272. IWKerguelen Islands.

CHAMAEBOSCA Speiser. Type microptera. «
microptera Speis. O3S67. Chile.

Notre.—Mr. Coquillett seems to have recognized that the species oscitans
(Walker) belongs in this genus (00C260). In Smith’s Catalog of New
Jersey species it is placed in Scatella. The original Walker description,
which places it in Ephydra, does not make the generic characters suffi-
ciently certain. It appears that Coquillett or Johnson (Smith’s Catalog)
should give us a description of this species by which an agreement might
be reached in regard to its generic relations. If an Ilythea, oscitans
differs from both spilota and flavipes in that the antennal avista is bare.
while in the former there are six to eight rays and in the latter eight.

Vor. 1.] Jones.—Catalogue of the Ephydridae.

HYDRELLINAE.
GLENANTHE Haliday. Type ripicola.
fuscinervis Beck. 96B165, 968291. Norway.
ripicola Halid. 39H404, 53 W258, 60L16, 648246, 96B165.
EKurope.

HYDRELLIA Robineau-Desvoidy. Type griseola.

NortH AMERICAN SPECIES.

ee TMIne MESS ey eM OW! s atcictstos Gite. cls wre ara aiee os gilvipes Coquillett.
WESSHNOEMENBIFE]Y My CllOWE ae.ce hts ns secs Sale deere eb 5 oe adele ss

DEAN oi teu ecoxse yellows ciaciesie ccs ee elicit se pulehra Williston.
Only sthe-anterior, cox yellow... J. 05.2256 00 ischiaca Low.
INOM ee OREEN eco xe VellOWineiss sem a6 chs Stoned eeaneiee Bale oa ee
LN Come lnmimocnwrlnitelanie, « ciefehsi sts cie cd che ete evclansteus formosa Loiw.
HIACERSNOW VsAWIILC) teens as ele ctatelne wc ee cie ow otto hypoleuca Low.
Hacenpal rye cana c tere ater ogee oleae Siclasereloro evel £ Sractieué-e ateser6 6
Face dark yellow, narrow, much widened below............

LEM ene sE ONES ol chielik evalicns, Soetere elas micron stece dose scapularis Low
Hacesbrownish=black*opaguies. «..o.css cae tench veesie nc “peabeet

4. Face rather broad, but little widened below...... valida Low.
Face narrow, much dilated below as in scapularis..........
REN NSaS get here) clive, erates: Shas Dios nie he aveorstae le aero conformis Low.

5). Abdomen brownish metallic green, somewhat glossy........

Bee Pree ee a ce tennis iekcto. cfeieie Sr cio wien elanere An es obscuripes Low.
Abdomen black but little shining, thinly grayish dusted......

FB nis Gide BreRe EN EASA er EDEL ANG AEE ea | parva Williston.

apicalis. See Ephygrobia.

compta. See Ephygrobia.

conformis Lw. 69141, 72L75, 69D444, T80S202, 96B270,
05A626. Newport (R. I.).

formosa Lw. 611355, 621154, 65L94, 7T8OS202, 96B270,
00H593, 054626. Pennsylvania.

gilvipes Cog. O00C261, 008309, 054627. Porto Rico.

hypoleuca Lw. 621151, 7808202, 96B269, 05A627.
Middle States.

ischiaca Lw. 621150, TSOS202, 96B269, 05A627. Middle
States.

maritima. See Ephygrobia.

nitidula. See Psilopa.

obscuripes Lw. 621152, T8OS202, 96B269, 02S8, 05.4627.
Middle States, White Mountains (N. H.).

parva Will. 96W399, 96S291, 05A627. St. Vincent.

pulchra Will. 96W400, 968291, 05A627. St. Vincent.

scapularis Lw. 621153, 7T80S202, 96B269, 00C461, 028320,
05A627. New Jersey, [llinois west to California.

valida Lw. 621153, 7T8OS202, 96B269, O5A627. Middle
States.

EUROPEAN SPECTES.

albiceps. See mutata.

albifrons. See nigricans.

albilabris Meig. 53H258, 56H345, 60L18, 648248, 66F973,
66D512, 96B183, argyria 830RD793, argyrostoma 448236,
Notiphila 46Z1895, albilabris 30M71. North and Middle
Burope, Silesia.

oe

186 University of California Publications. |ENTomoLocy

BDUROPEAN SPEcIES—Continued.

argyrostoma. See albilabris.

arygrogenis Beck. 96B185, 968290. Milan.

caesia Stenh. 448214, 60118, 648249, 96B176, Notiphila
46Z1880. Sweden, Germany.

cardamines. See flavilabris and laticeps.

concolor Stenh. 448216, 601820, 648247, 96B178,
cinerascens 35M526, Notiphila concolor 46Z1877.
Sweden, Germany, Austria.

discolor Stenh. 448230, 60L22, 648250, 96B179, flaviceps
39H345, Notiphila 30M72, discolor 46Z1900. England,
Sweden, Germany, Hungary, Silesia.

erythrostoma. See nigripes.

flavicornis Fall. 96B177, Notiphila 2310, 46Z1873.
Germany, Sweden, Hungary, Silesia. See also nigripes.

flavilabris Stenh. 448235, 9458126, 96B173, cardamines
39H402. Styria.

frontalis Lw. 60119, 648249, 96B1838. Silesia.

fulviceps Stenh. 448231, 60121, 648248, 96B178,
Notiphila 4621901, chrysostoma 30M67. Germany,
Sweden, Silesia.

fusca Stenh. 448225, 60L24, 648250, 96B178, Notiphila
46Z1896. Sweden, Germany.

genicula Stenh. 448224, 60L18, 648248, 96B174, Notiphila
46Z1878. Germany, Sweden.

grisea. Stenh. 448227, 60L21, 648249, 96B179, Notiphila
46Z1898. Sweden, Germany, Russia.

griseola Fall. 448220, 60L22, 648247, 96B180, Notiphila
138F254, 28F9, 30M66, 38Z717, 46Z1869. Europe,
Silesia.

hispanica. See nigricans.

incana. See ranunculi.

lamina Beck. 96B184, 968291. Leignitz.

lapponica Stenh. 448287, 96B174. Pavia.

laticeps Stenh. 448229, 60L20, 648248, 96B172,
cardamines 39H402, Notiphila laticeps 46Z1899. North
and Middle Europe, Silesia.

maculiventris Beck. 96B175, 968291. Silesia.

modesta Lw. 60128. 648250, 96R181. Europe.

mutata Zett. 60L19, 648247, 948126, 96B176, plumosa
448218, albiceps 56W345. Notiphila mutata 46Z1876.
Sweden, Germany, Silesia.

nigricans Stenh. 448234, 648250, 96B183, nigrina 60124,
var. hispanica 0081, 00S309, albifrons 448223, Notiphila
23F10. Sweden. Germany, Austria.

nigrina. See nigricans.

nigripes Zett. 60122, 648248. 96R181. flavicornis 448232,
56W345, erythrostoma S8W257. Notiphila 30M69,
nigripes 38Z717. North and Middle Purope, Silesia.

nymphaeae Stenh. 448227. 601.28. 96B182. Notiphila
46Z1897. Sweden.

pilitarsis Stenh. 448219, 60120. 648249. 96R173,
Notiphila 4671881. Sweden, Germany.

plumosa. See mutata.

Vor. 1.] Jones.—Catalogue of the Ephydridae.

EUROPEAN SPEcIES—Continued.
ranunculi Halid. 39H402, 538W256, 60L23, 648247,
96B182, incana 448222, Notiphila 46Z1871, griseola
23F9. Europe, Silesia.
thoracica Halid. 39H402, S3W256, 60L18, 648249,
96B183. England, Germany, Silesia.
transsylvana Beck. 96B184, 968291. Transylvania.

OTHER COUNTRIES.
tritica Cog. 08C324. Australia.

NOSTIMA Coquillett. Type slossonae.
slossonae Cog. OO0C35, 008310, 05A627. Florida.

' AXYSTA Haliday. Type cesta.
cesta Halid. 33H177, 53W262, 60L28, 648255, 96B167,
Philygria punctulata 448241, T'rimerina coeruleiwentris
35M529. North and Middle Europe.

PHILYGRIA Stenhammar.

NortH AMERICAN SPECIES.
1. Posterior portion of abdomen differing in color, or at least in
MUStregirommancerior POLLWONeaeeciacie sscleiaelce aie nm aereciioe a. «
AHdOMEensCOncOlorous) LhrOuUgn OU wn. os 2 -ecursiei) cies c djeleeisioms
2. Prevailing color blackish cinereous, antenne entirely black... .
SGC. O CIEL OU Oe OO DEI IO IRE TE Roar eater ae debilis Low.
Prevailing color brownish gray, third antennal joint reddish,
vellowabeneathinn sires cassis, setselors.ciens vittipennis Zetterstedt.
35 IPneyeuilbine @alkore [signin canabodss cob ous uoeeaasueoooodnoebos
Be valin SCO] OTM La CKeiegsens eas stetetss Sle iss eee c ehee sts ene mlceaoiste ans

4. Transverse veins margined with blackish brown............
fuscicornis Low.

wi w)je le Me)relefisiie}\6)\e) (4 felis «eee ¢| ef@ 6: ele 4110.0) 6,2! 6 « 0) ee 2 ee .=

Transverse veins broadly clouded with black... .opposita Low.
Oe @oxcay lnehityryell Overseas se) acess Ao oe fern 3 yar nitida Williston.
(Glipsis NIETO 5 6 sais Aerie Cie nara caer nitifrons Williston.

debilis Lw. 611357, 621157, 65L96, 7T8OS202, 96B269,
0288, 05A627. Pennsylvania, White Mountains (N.H.).
fuscicornis Lw. 62L155, T8OS202, 96B269, 0288,
O05A627. Middle States, White Mountains (N. H.).
nitida Will. 96W400, 968291, 05A627. St. Vincent.
nitifrons Will. 96W401, 968291, 054627. St. Vincent.
opposita Lw. 611356, 621.156, 65L95, 7808202, 96B269.

Pennsylvania.
vittipennis Zett. 780S202, 96B270. Greenland.

EUROPEAN SPECIES.

abdominalis. See Lytogaster.

femorata Stenh. 448245, 648253, 96B190, Notiphila
23F12. Scandinavia, Germany, Silesia.

flavipes Fall. 35M525, 448244, 60L26, 648252, 96B191,
Notiphila 23F12, 46Z1916, Ephydra 30M123. LBurope,
Silesia.

interrupta Halid. 53W261, 60126, 648253, 96B189,
Hydrina 34H176. Middle Europe, Silesia.

18

(

i)

a

188 University of California Publications, | NTomMoLocy

EUROPEAN SPECIES—Continued.

interstincta Fall. 448246, 648253, 96B190, Notiphila
138F254, 23F12, 46Z1919. Europe.

nigricauda Stenh. 448248, 96B192, Notiphila 46Z1923.
Scandinavia, Germany, Silesia, Transylvanian Alps.

obtecta Beck. 96B122, 968291. Silesia.

picta Fall. 448248, 60125, 6482538, 96B192, Notiphila
13F 254, 23F11, 46Z1913, Hphydra 30M125. North and
Middle Europe, Silesia.

punctatonervosa Fall. 448247, 583W260, 60L25, 648252,
96B188, Notiphila 13F254, 23F12, 4621921, Ephydra
30M123. North and Middle Europe, Silesia.

punctulata. See Axysta cesta.

sexmaculata Beck. 96B191, 968291, Hphydra interstincta
30M122, 35M539, Hydrina maculipennis 20RDT95.
North and Middle Europe.

stictica Meig. 60125, 6482538, 96B189, Ephydra 30M121,
Germany, Silesia.

2. vittipennis Zett. 448250, 4621924, 60126, 648253,

96B193, Notiphila 38Z718. FBurope.

HYADINA Haliday.

NortH AMERICAN SPECIES.

1. Third antennal joint reddish, infuscated toward the tip......
Re eer on ARE TS ran eit Gens REA SiG IER ONEECe NOE Bao -o8e gravida Low.
Third antennal joint black above, yellow beneath............
ithe tol hs tcuaee diaper eRe De at ereee lean eRe pawegereaa esac albovenosa Coquillett.
albovenosa Coq. 00C34, OOS309, O5A627. Georgia,

Louisiana. :
gravida Lw. 631325, 72L98, 78OS202, 96B270, 054627.

Sitka.

EUROPEAN SPECIES.

guttata Fall. 30RD795, 448239, 53W261, 60127, 648254,
96B194, O0S2, O0S809, Hphydra 380M125, Notiphila
18F 2538, 23F11, 382718, 4671914. North and Middle
Europe.

interrupta. See Philygria.

humeralis Beck. 96B195. 968291. Germany.

nitida Macq. 60128, 648254, 96B194, Hphydra 35M539,
Notiphila brevicornis 46Z1912, guttata var. brevicornis
448240. North and Middle Europe, Silesia. —

scutellata Hlalid. 39H406, 53W262, 96B195. [reland,
Silesia.

seamaculata. See Philygria.

EPHYDRINAE.
OCHTHERA Latreille. Type mantis.

Nortu AMERICAN SPECIES.
1. Face with impressed black lines radiating from an_ orbital
ETOOVE cts ee cash susie lees gees ate ine oteneae eben lauta Wheeler.
Eaceswithout Such! dines. samunmieciciocmie tr ecer rien rreneyieenenen re 2

Vor. 1.] Jones.—Catalogue of the Ephydridae.

NortTH AMERICAN SPECIES—Continued.

2. Face with deep black furrows and dots........ exsculpta Liw.

Haceawithourblack turrows and dots)...0..0..4.0oeeeues... -
3. Mesonotum with three dark purple and coppery stripes

Dac CYCLO alot OER CRUG G EROIC RC ICICIO CERO ICT eee cuprilineata Williston
Mesonotumewithouwtesuch) Stripes... <5 22)s 2 feels nieiieiere eae «

4, Wirst joint of hind tarsi but little swollen. ...mantis De Geer.
Hirst yOmMbtor Hindetars) much swollemea.).-% cel easel se. -
Dears black e1a Ce w DLO ccc s)c.c ee aco cose ele sl ete slolele rapax Low.
Narsicered race NALLO WE cise /oe-s es as csc iieie ee tuberculata Low.

cuprilineata Will. 96W402, 968291. St. Vincent.

exsculpta Lw. 62L160, 7T80S202, 86W307, 953338, 96B269,
ODd5A628. Cuba, Florida.

lauta Wheel. 96W121, .968291, 05A628. Milwaukee
(Wis. ).

mantis Deg. 621161, T8OS202, 95GT66, 96W123, 96B269,
968264, O5A628. Middle States, White Mountains
(N. H.), Connecticut to California, Mexico.

rapax Lw. 621162, 7808202, 96B269, 05A628. Carolina.

tuberculata Lw. 621161, TSOS202, 953338, 96B269,
96W123, 05A628. Illinois, Wisconsin, Florida.

EUROPEAN SPECIES.

2. mantis Deg. 1809L848, 35M519, 38Z715, 448166, 4621804,
60L30, 621161, 648256, 96B204, I/usca 1782D61,
manicata 17T98EK334, Macrochira mantis 30MT8, Tephritis
manicata 05F323, 283F2. Whole of Europe.

mantispa Lw. 4712738, 60L30, 96B205, Schembri 47R29.
Italy, Greece, Asia Minor.
schembri. See mantispa.
OTHER COUNTRIES.
chalybescens Lw. 62114, 96B268. Cape of Good Hope.
humilis Will. 97W6, 978265. Brazil.
innotata Walk. 6O0W171, 96B267. Celebes.
praedatoria Lw. 62L14, 96B268. Caffraria.
regalis Will. 97W6, 978265. Brazil.
rotunda Schin. 688243, 69D374, 96B273. Nikobara.

OCHTHEROIDEA Williston. Type «atra.
atra Will. 96W401, 968291. St. Vincent.

ECTROPA Schiner. Type viduata.
viduata Schin. 688243, 68D374, 96B201. Sydney.

PELINA Haliday. Type aenea.

NortH AMERICAN SPECIES.
truncatula Lw. 78L198, 780S202, 78K244, 96B270,
05A628. Texas.
EUROPEAN SPECIES.
aenea Fall. 188253, 238F11, 46271926, 60L30, 648255,
96B197, glabricula 30M124, Telmatobia 448209.
Europe. é

ivy)

190 Unversity of California Publications, [ENTomoLocy

EUROPEAN SPECIES—Continucd.

aenescens Stenh. 448201, 46Z1928, 60L30, 96B198, aenea
382718, Mikvi 9388280. Scandinavia, England, Austria,
Silesia.

guttipennis Stenh. 96B199, Telmatobia 448212. Europe.

nitens Lw. 73L309, 73R427, 96B199. Calabria.

subpunctata Beck. 96B198, 968291. Dalmatia.

ventruosa. See Lytogaster abdominalis.

LYTOGASTER Becker. Type abdominalis.
abdominalis Stenh. 96B208, Philygria 4458238, Pelina
ventruosa 73L3810. Silesia.

DOMINA Hutton. Type metallica.
metallica Hutt. 0O1H90, 018272. New Zealand.

BRACHYDEUTERA Loéw. ‘Type argentata.

NortH AMERICAN SPECIES.
argentata Lw. 05A628, dimidiata 621163, T7T8OS203,
95J338, 96B269, Notiphila 56W406. District of
Columbia, Florida.
dimidiata. See argentata.
EUROPEAN SPECIES.
argentata Walk. 96B201, dimidiata 62L163, Hphydra
06W407. Europe.
dimidiata. See argentata.

HALMOPOTA Haliday. Type salinaria.
mediterranea Lw. 60L384, 96B206. Asia Minor.
salinaria Bouche. 56W346, 60134, 648260, 961B205,
Hphydra 34B99. England, Germany, Silesia.

PARYDRA Stenhammar.

NortH AMERICAN SPECIES.
es Scutellum’ watheconicalliiwatsterercir ieee nae nenencne ane

Scutellums without) conical es wartsaeeeeeer eee eter ecieeiene
2. Scutellum with one wart, apex not bearing a spine.........
Scutellum with two warts, apices with spines..............
Scutellum with four warts........... quadrituberculata Low.

3. Abdomen subopaque, tips of femora, base and tips of tibie.
and metatsrsi «dark reddishis «asc steers imitans Low.

Abdomen black, legs without reddish coloring..............

5 ata Mignacel Sei Stes real oer orate Fata etemeie be aaah star sian ae unituberculata Low.

4. Tubercles exceedingly small, hairing of face snow white......
sib: wee baigenh ol Soot rious ate Shel errai ahonetenabanewewen ie caneaete ts foesued SETS pinguis Walker.
Tubereles not unusually small, hairing of face not white......
bituberculata Low.

cles aie (eleje sete ei ee \e ele 6 0.\6 [ele \siniie) wilel ele ale (mele) owas)

oO. Hace nearly perpendicular) erectile eieisiiar breviceps Liw.
Face more or less oblique......... ee Ae ar iirpa kr ra OTS
6. Clypeus and cheeks excessively narrow......... paullula Low.

Clypeus and cheeks not excessively marrow..................
Second longitudinal vein with a branch near its end.........
Second longitudinal vein without such branch.............

me Co OURS

Wort. gl

Jones.—Catalogue of the Ephydridae.

NortH AMERICAN SPECIES—Continued.
8. Fourth longitudinal vein with a small but very distinct gray

fringe near its apex, third with a similar spot less distinctly
WASIDIC iaper st cpetois siti sra ei aicroue oi Aiea a ie doniclers oak appendiculata Low.

Third and fourth longitudinal veins without such markings... .
9. Wings with seven rather large hyaline spots. ..abbreviata Liw.
Wing siwathetiverhyalinesspous. «1s. ce canes. ade no sl varia Low.
10. Wings brownish with five hyaline spots, cross veins brown....

2 BD CO CODE Piadol 5 CODERS 6 COIR SID ORO ae aurata DD. sp.

Wings hyaline, cross veins blackish........limpidipennis Liw.

abbreviata Lw. 611357, 62L168, 65L97, 7808203,
96B269, 05A629. Pennsylvania.

appendiculata Lw. 78L202, 7808203, 78244, 96B270,
OSA629. Texas, California.

aurata n. sp. Southern California.

bituberculata Lw. 62L165, T8OS203, 96B269, 05A629.
Middle States, New Jersey.

breviceps Lw. 62L167, T8OS208, 96B269, 05A629.
Middle States.

imitans Lw. 78L201, 7808208, 78Kk244, 96B270, 054629.
Massachusetts.

limpidipennis Lw. 78201, 7808203, 78244, 96B270,
05A629. District of Columbia.

paullula Lw. 621167, 78082038, 96B269, 000462, 054629.
Middle States, Alaska.

pinguis Walk. 78L199, 7T8OS203, 78K243, 05A629, Ephy-
dra 56W409, 96B266. District of Columbia, Texas,
New Jersey.

quadrituberculata Lw. 621165, 648258, 7808203,
953338, 9613269, 054629. Middle States, Florida.

unituberculata Lw. 78L200, 78O0S203, 78IK244, 96B270,
05A629. District of Columbia.

varia Law. 631326, 651100, 7808203, 96B270, 054629.
Sitka.

IUROPEAN SPECIES.

affinis. See fossarum.

aquila Fall. 448187, 4621819, 60L32, 648259, 64D559,
96B211, Hphydra 23F4, 30M117, 85Md537, 382716,
North and Middle Europe, Silesia.

bicuspidata Kars. S81K15, 81K253. Porto Allegre.

coarctata Fall. 448189, 46Z1821, 60L33, 648258, 96B214,
Ephydra 2384, rufitarsis 30M126, 35M356, stagnicola
30RD799, hecate? 53H263. Whole of Europe, Silesia.

cognata Lw. 601382, 96B212. Prussia, Sicily.

fossarum Halid. 53H175, 60L32, 648260, 96B211, afinis
448192, 46Z1824. North and Middle Europe, Silesia.

furcata. See quadripunctata.

littoralis Meig. 380M116, 60133, 648259, 96B214. Prussia,
Germany, Silesia.

nigritarsis Strobl. 938S280, 988321, 96B215. Styria.

nubecula Beck. 96B212, 968291. Prussia, Silesia.

obliqua Beck. 96B215, 968291. Crete, Italy.

pubera Lw. 60132, 96B210. Sicily, Calabria.

193i

192

Unwersity of California Publications. |! NToMoLocy

EUROPEAN SPECIES—Continued.

pusilla Meig. 30M126, 60L382, 648259, 96B211, Hphydra

infecta 33H175, nasuta 448192, 46Z1825. North and

Middle Europe, Silesia.

quadripunctata Meig. 80M117, 382716, 46Z1822, GOL33,
648258, 96B214, furcata 448190. North and Middle
Europe, Silesia.

undulata Beck. 9618213, 968291. Russia, Berlin.

OTHER COUNTRIES.

bucculenta Lw. 62114, 96B268. Caffraria.
humilis Will. 9T7W7, 978266. Brazil.

EPHYDRA Fallen. Type riparia.

NortH AMERICAN SPECIES.

ff eBhoraxy Jemoney.clloweere ee ote cece eine lutea Wiedmann.
Thorax ashy, with slight greenish reflection........:........
Thorax’ black sor ‘browaishy black. <i. «ee cr ere itcl sere ctareiioreeen
Thorax: dark, Seen sis otitis cee scicevera cine erste weiter aio anemones
Phorax Metallic Orn COpPeLryvaUSeMai-n= ert clehene/-kelcls tenskameyeiene eters

2. Thorax with three broad vittx, front brassy green. .cinerea nN. sp.
Thorax without vitte, front dark bluish.......... hians Say.

3. Wings except along veins blackish or smoky...............
Wings eray FOr WiyaiMeny ie sos.) apelelee) lies <1 ee levers suelo ney enn erereicnetelstene

4. Front except the lateral margins, shining, slightly greenish....
Re SER eI a ALS Ira eeL RES iekG 6 GOTO californica Packard.
Front opaque velvety black............... pugmaea Williston.
Front grayish white, dusted with brownish. .thomae Wiedmann.

yf Morar lly ceils cco. stan ios setae evetel oxetenceliotench tiene = opal Seonterenenene totale
“Nii ob ancl iboibiynh aereomc oocOne Coes banc eor cin De do bcd C606

6: Abdomen shimine. 5 cc..cserer spss le cis eiretete sceucketiets brevis Walker.
Abdomen! note SHIMITIO sie cerns keris cteretenel tens tarsata Williston.

fee LUSITANS posopadcugaconc ror oonanouncdocaaoes lata Walker.
Megs paler yellowie caesar S steadier) here. nana Walker.

Sa Thirdsantennalejoimtewithyaeiacera liaise sae ite tt leeteietene items
Third antenna] joint without a lateral hair.................

9. Wings clouded with blackish gray........... atrovirens Low.
Wings grayish hyaline, not clouded...... pilicornis Coquillett.

10). LhoraxOpadies am sara setarecrchasie re ciccrierneerers obscuripes Low.
Mhioraxsshining.\~ Hose eae eae ole iol ie elere austrina Coquillett.

11. First joint of front tarsi incrassated........ crassimana Low.
Hirst joimt of fronts tarsi mot IMCrasSaved errs a1. leveae leven) etetene

12. Third antennal joint minute, sunken in the head..........
BP end Or Oe OAC HUG OOIEE CO all" halophila Packard.
Third antennal joint not minute, not sunken in the head....

13. Antenne dark brown, arista with short pectinations........
Ben te TNO SI crore Oat MLO DIE eroTOa-00 1 OG millbrae n. sp.

Antenne black, arista with long pectinations...suwbopaca Low.

Norr.—E. gracilis described in larval stages only.
Norr.—oscitans (Walker) certainly differs from all others of this
genus in the markings of the wings, and if in this genus at all probably
belongs near pygmaea or thomae, from which it differs in that the front is
dark brown.

ad] |

Vor, 1.] Jones.—Catalogue of the Ephydridae. 193

NortH AMERICAN SPECIES—Continued,

atrovirens Lw. 621169, 78082038, 96B269, 05A629.
Middle States.

austrina Cog. 00C36, 05AG629. Georgia, Florida.

brevis Walk. 57W233, 78OS203, 96B268, 05A629. United
States.

californica Pack. 71P103, 7T1R897, 84K266, 84W90,
05A629. California, Nevada.

cinerea n. Sp. Southern California.

crassimana Lw. 651182, 65LS8, 65D664, 7808200,
96B270, 05A629. Mexico.

gracilis Pack. 71P105, 71R3897, 918235, 05A629. Great
Salt Lake (Utah).

halophila Pack. 68P46, 69D444, 7808203, 966270,
05A629. Illinois.

hians Say. 308188, 888871, 831X250, 883P976, T8O0S200,
96B270, 05A630. Mexico.

lata Walk. 57W233, 96B268, 054630. United States.

lutea Wied. 30W593, 96B266, 0O5A630. West Indies.

millbrae n. sp. Millbrae (California).

nana Walk. 57W234, 953339, 96B268, 05A630. United
States.

obscuripes Lw. 66150, 66D512, 7T2L92, T830S200, 96B270,
968290, 05A630. Massachusetts.

octonotata. See Scatella.

oscitans Walk. See Ilythea and Scatella.

pentastigma. See Scatella.

picea. See Scatella.

pilicornis Coq. 20184, 028264, 05AG630. Florida.

pygmaea Will. 96W402, 968290, O5AG630. St. Vincent.

striata. See Scatella.

subopaca Lw. 64198, 64D559, 65L99, T8O0S8200, 95J359,
96B270, 05A630. Connecticut, New Jersey, Florida.

tarsata Will. 98W257, 938320, 05A630. Owens Valley
(California ).

thomae Weid. 30W593, 96B266, 05A630. St. Thomas
(West Indies).

HUROPEAN SPECIES.
aquilla. See Parydra.
argentata. See Brachydeutera.
argyrostoma. See Scatella sorbillans.
attica Beck. 96B222, 968290. Greece.
aurata. See micans.
beckeri (new name) obscuripes. 96B222. Sarepta.
bivitatta Lw. 60135, 96B219. Prussia, Sicily.
breviventris Lw. 60L37; 648261, 96B219. Southern ,
Europe.
cribrata. See Scatella.
fenestrata. See Scatophila despecta.
flavescens. See Scatella lutosa.
flaviceps. See Philygria.
flavipennis. See Scatella aestuans.
gramium. See Scatella quadrata.
guttata. See Hyadina.
halophila. See riparia.

194 Unversity of California Publications, [ENtomotoey

HUROPEAN SPECIES—Continued.

hecate. See Parydra coarctata.

infecta. See Parydra pusilla.

longipennis. See Teichomyza.

macellaria Egg. 62E779, 648262, 96B220. Europe,
Southern Russia.

micans Halid. 383H175, 60L36, 648261, 96B218, aurata
448167, 46Z1810, riparia 23F4. Whole of Europe,
Silesia.

nasuta. See Parydra pusilla.

nubilipennis. See Scatella sibilans.

obscuripes. Preoceupied, see beckeri.

ochrostoma Brul. 385B689, 96B224. Hurope.

opaca Lw. 56L55, 96B222. Egypt, Southern Hurope.

pinguis. See Parydra.

punctinervosus. See Philygria.

quadrata. See Scatella.

riparia Fall. 138F246, 28F3, 30M117, 38Z715, 448169,
46Z1807, 53W268, 60L35, 648262, 96B221, salina
43H228, halophila 44H203. North coast of Europe. See
also micans and Caenia fumosa.

rufitarsus. See Parydra coarctata.

salina. See riparia.

salinae Zett. 46Z1912, GOL36, 96B219. North and Middle
Europe.

salinaria. See Halmopota.

sexmaculata. See Philygria.

Scholtzi Beck. 96B220. Silesia, Poland.

stagnalis. See Scatella and Scatophila despecta.

stagnicola. See Parydra coarctata.

stenhammari. See Scatella.

stictica. See Philygria.

OTHER COUNTRIES.

aquaria Hutt. 01H90, 018273. New Zealand.

australis Walk. 56W409, 96B266. Van Dieman’s Land.

bispinosa. See Beckeriella.

borboroides Walk. 60W171, 96B267. Celebes.

caesia v. d. Wulp. 83W58, 838K251, 96B271. Argentina.

ciligena Rond. 68R82, 68D374. Buenos Ayres.

gilvipes Cog. O1C377, 018272. Galapagos.

maculicornis Walk. 60W171, 9618267. Celebes.

margaritata Wied. 30W593, 96B266. Egypt.

ochropus Thoms. 681592, 70V442, 96B271. Montevideo.

pictipennis Wied. 30W593, 64D559, 96B266. Cape of

Good Hope.
x pleuralis Thoms. 681591, 70V442, 96B271. Manila.
prionoptera Thoms. 681590, 70V442,- 96B271. Pata-
gonia.

taciturna Walk. 60W169, 96B267. Amboyna.
urmiana Gunth. 99G415, 998239. Northwest Persia.

SCATOPHILA Becker. Type caviceps.
NorrH AMERICAN SPECIES.
hamifera Beck. (See “‘University Collection.”) Alameda
(California).

Vor. 1.] Jones.—Catalogue of the Ephydridae.

EUROPEAN SPECIES.

caviceps. Sten. 96B240, 968292, Scatella 448269,
46Z1837, 60142, 648265. Scandinavia, Germany,
Silesia.

contaminata Stenh. 96B240, Scatella 448185. Sweden,
Silesia.

cribrata. See Scatella.

despecta Halid. 96B241, Scatella 39H409, 60L48, 648267,
Ephydra stagnalis 23F5, fenestrata 448181, 46Z1831.
North and Middle Europe, Silesia.

halterata Beck. 968245, 968292. Kohlfurt, Silesia.

2. hamifera Beck. 96B242, 968292. Norway.

laevigata Lw. 96B243, Scatella 60L44. Silesia.

pumilio Lw. 96B244, Scatella 60L44, 648267. Silesia.

quadrilineata Strobl. OOS4, 008810. Spain.

signata Lw. 96B241, Scatella 60L48. Sicily, Hungary.

tetra Beck. 96B244, 968292. Léw’s Collection (prob-
ably Silesia).

unicornis Czer. O00C205, 0OS310. Austria.

variegata Lw. 96B243, Scatella 60L48, 648267. Silesia,
Dorpat.

PELOMYIA Williston. Type occidentalis.
occidentalis Will. 93W258, 938321, 96B274, V00C461,
05A628. Monterey (California), Sordova (Alaska).

TEICHOMYZA Macquart. Type fusca.
fusca Macq. 305M585, 60145, 648268, 86G160, 868312,
90H239, 908290, 96B206, NScatella urinaria 30RD6,
Ephydra longipennis 38M382. England, France, Ger-
many, Italy, Silesia.

SCATELLA Robineau-Desvoidy. Type stagnalis.

NortH AMERICAN SPECIES.

Ibe

to

Face in profile almost perpendicular, decidedly haired and
AGA a tks Bic ioe Agee OG tee eae stagnalis Fallen.
Face not unusually perpendicular, hairing not pronounced... .
Thorax, coxwe, femora and tibix whitish cinereous, wings
irregularly streaked with whitish hyaline. .mesogramma Low.
Little or none of the body whitish cinereous, wings usually with
GEMTILCH ESD OLS yep ocho aloes ae Sion Ate ala Gt tatlape a. ohisraitalinl oper te eerie

SPS POLS OtEthemwAnesm Myaline: One WHIGIS cetera cle seel shel eileen
SOUS Ole HOO AMADA lomo itoemron es one pened Cem pe oe conor ooboc

dbo MVE Alt, TRO \wdINlie KNOWS moms Gaceaeaedon acco ao gadU6
WWamesse waltham invel Wal GO ESOC: sre ajcycuereseras elas) suansys: sJellspelerstaleneversiel
Wings with seven or eight white spots.....................
Wines witheteninwiite SpPOUS.: «+e. ave scene’ striata Walker.

5. Two large brown spots between the first cross vein and the
point of the wing beyond the third longitudinal........

MEU PRL re atatey 0 fy sfstece cobs, cats, oid suatso stereee 4 8% quadrata Fallen.
Three or more brown spots beyond the third longitudinal and
hehindimbher teste CrossivielMicncisn eis seis s ciecreis cists cic cis ciel
Gapbascwotethemtarsin blacks cs. ste sissies «ere ere cts stenhammari Zett.

Base of the tarsi more or less reddish. ..........5 sejuncta Low.

~

fon oes MS) io ne

=

196

University of California Publications, | NToMoLocy

NortH AMERICAN SPECIES—Continued.

Ue

10.

Wing with a small pitchy spot at the tip of the first longi-

GUGM] oo cep svelesvalslate ovale a etecetete elersis o elerereniere picea Walker.
Wings! without such Spotectiacicee icicle eioitaretene ratte

2 Body: ash: 28a acic< 3 tis ost s ols chee sss wis + scveretiets favillacea Liéw.
Body, olive green.c}sicvaaiveccieta ser ereiete ere ausheiens triseta Coquillett.
Body, DroWwilicmiectave. acres orceioclereiterasin ewe hele octonotata Walker.

. Head and chest blackish opaque, densely bluish-gray pruinose
EE Er Or PRY SAO AITIOG Oo Sloe odo AEB Oo URE setosa Coquillett.
Head and chest blackish with olivaceous pruinosity........
ateifaueiayyarea eve eras lo detoneteus[anetehoNonerensiene fitch erate pentastigma Thomson.
Head and chest yellowish cinereous............. obsoleta Low.
Headvand (chest) ferrnusineusicereirreeietens vepleta Walker.
Abdomen flecked with white laterally...cribrata Stenhammar.
Abdomen not flecked with white................ lugens Low.

cribrata Stenh. OOL303, 05A630. Greenland.

favillacea Lw. 621170, 78OS208, 96B269, 05A630.
Middle States, New Jersey.

lugens Lw. 621171, T8OS203, 953339, 96B289, 05A630.
Middle States, Florida.

mesogramma Lw. 69142, 69D444, 72L74, T8OS2038,
96B270, 05A680. Newport (R. I.).

obsoleta Lw. 621172, 65L98, T8OS204, 96B269, 054630.
Washington (D.C.).

octonotata Walk. O5A630, Hphydra 49W1106, 96B267.
Martin Falls (Canada).

oscitans Walk. See Ilythea and Ephydra.

pentastigma Thoms. O5A630, Hphydra 68T591, TOV442,
7808204, 96B271. California.

picea Walk. 0O5A630, Ephydra 49W1105, T8OS204,
96B266. Martin Falls (Canada).

quadrata Fall. 2355, 641317, 648265, 7T5O0S204, 96B270,
05A630. North America.

repleta Walk. O5A630, Notiphila 49W1099, 96B266.
Martin Falls (Canada).

sejuncta Lw. 631326, 65L99, T80S204, 96B271, 05A631.
Sitka (Alaska).

setosa Coq. 00C462, 008310, 05A631. Alaska.

stagnalis Fall. 458169, 72H97, 7808204, 86G162, 868312,
96B271, 968264, 9813803, 00C462, 00H593, 05A631,
obscura 96WA403. Greenland, Alaska, New Jersey,
Georgia, Arizona, St. Vincent, Montreal, White Moun-
tains (N: H.).

stenhammari Zett. TSOS204, 96B271. North America.
North America.

striata Walk. 05A631, LHphydra 49W1107, 96B267.
Martin Falls (Canada).

triseta Coq. 020184, 02S265, 054631. Arizona.

Nore.—oscitans, if a Scatella, is probably most closely related Lo
S. striata, from which it differs in the absence of five hoary stripes on

the chest.

Vor. 1.] Jones.—Catalogue of the Ephydridae.

EUROPEAN SPECIES.

aestuans Halid. 33H176, 648266, 96B283, Ephydra flavi-
pennis 448173, gilva 60L41. Asia Minor, Southern
Hungary.

callosicosta Bezzi. 95B70, 958545, 96B232. Calabria,
Orsova, Silesia.

contaminata. See Scatophila.

crassicosta Beck. 96B234, 96S291. Sylt Island.

2. cribrata Stenh. 60142, NScatophila 648265, 96B240,
Ephydra 448269, 46Z1835. North and Middle Europe.
defecta Halid. 33H174, 46Z1813, 60L38, 96B236, Caenia
53W265. Triest.
despecta. See Scatophila.
dichaeta Lw. 60L40, 96B228. Harz, Sylt Island, Sweden.
indistincta Beck. 96B2351, 968291. Hungary.
laevigata. See Scatophila.
lutosa Halid. 33H176, 60142, 648266, 96B2386, Ephydra
flavescens 448175, 46Z1830. North and Middle Europe,
Silesia.
pilosigenis Beck. 96R229, 968291. Hast coast of Hurope.
pumila. See Scatophila.
2. quadrata Fall. 30M119, 39H410, 448182, 4621840, 60L40,

648265, 96B230, Hphydra 23F5, graminum 330176.
Whole of Europe, Silesia. See also stenhammari.

quadrisetosa Beck. 96B229, 968291. Norway.

sibilans Halid. 33H175, GOL40, 648265, 96B228, Hphydra
nubilipennis 448180. North and Middle Europe, Silesia.

signata. See Scatophila.

silacea Lw. 60141, 648266, 96B233. Silesia.

sorbillans Halid. 33H176, 60141, 648265, 968231,
Ephydra argyrostoma 448176, 46Z1831. Whole of
Hurope, Silesia.

2. stagnalis Fall. 448178, 4621827, 60L42, 648266, 96B235,
Ephydra 13F248, 23F5. North and Middle Furope,
Silesia.

2. stenhammari Zett. 60140, 648266, 96B230, quadrata

448183, Ephydra stenhammari 46271842. North and
Middle Europe, Silesia.

urinaria. See Teichomyza.

variegata. See Scatophila.

OTHER COUNTRIES.

hawaiiensis Grim. 01G49, 018274. Oahu (Sandwich
Islands).
3. stagnalis Fall. 97W5, 978266. Brazil.

CAENIA Robineau-Desvoidy. Type palustris.
NortH AMERICAN SPECIES. -
1. Front black, clothed with long bristles..........spinosa Low.
Middle of front bronze green, partially covered with short
hairs, but without long bristles......... bisetosa Coquillett.
bisetosa Coq. 020183, 028263, 054631. Utah, California.
spinosa Lw. 64199, 651100. 64D559, TSOS204, 955339,
96B271, 054631. New York, Florida, New Jersey.

198 University of California Publications. UENToMoLocy

EUROPEAN SPECIES.

beckeri Kuntze. 9718154, 9758265. Rome.

carricola. See palustris.

fumosa Stenh. 448171, 4621813, 60138, 648264, 965207,
Ephydra riparia 234. North and Middle Europe,
Silesia.

obscura Meig. 30M115, 53W265, 60L38, 96B208. Ger-
many, England.

palustris Fall. 23F4, 30M115, 35M530, 382716, 448172,
46Z1815, 60L38, 648264, 96B207, carricola 30RDS800.
Whole of Europe, Silesia.

NOMBA Walker. ‘ype tecta.
tecta Walk. 60W169, 96B267. Celebes.

CANACENAE.

CANACE Haliday. Type nasica.
nasica Halid. 39H411, 53W269, 60L29, 648269, 74180,
96B247. Ireland. See also ranula.
ranula Lw. 74L81, 74R449, 878297, S7G1, 96B247,
nasica 55H64. North coast of England, Germany.
salonitana Strobl. 0O0S63, OOS308. Salona.
snodgrassi Coq. 01C3878, 018272. Galapagos.

GENERA APPEARING ONLY IN THE SYNONYMY.

Diasemocara. ,
nigrotaeniata. See Psilopa and Ephygrobia roderi.

Drosophila.

pollinosa. See Paratissa.
Glabrinus.

mororum. See Gymnopa subsultans.
Hydrina.

guttata. See Hyadina.
Keratocera.

palustris. See Notiphila cinerea.

tarsata. See Notiphila.

Macrochira.

mantis. See Ochthera.
Mosillus.

arcuatus. See Gymnopa subsultans.
Musca.

manicata. See Ochthera mantis.
T elmatobia. ,

aenea. See Pelina.
Tephritis.

manicata. See Ochthera mantis.
Ulida.

arcuata, See Gymnopa subsultans.

‘

4 i eet

ES Tay UNIVERSITY OF CALIFORNIA PUBLICATIONS

TECHNICAL BULLETINS
' COLLEGE OF AGRICULTURE, AGRICULTURAL EXPERIMENT STATION

a : ENTOMOLOGY

“Vol. 1, No. 3, pp. 199-216 February 2, 1916

ia Does

SYNOPTICAL KEYS TO THE GENERA OF
THE NORTH AMERICAN MIRIDAE

eae BY

EDWARD P. VAN DUZEE

UNIVERSITY OF CALIFORNIA PRESS
BERKELEY

UNIVERSITY OF CALIFORNIA PUBLICATIONS

TECHNICAL BULLETINS
COLLEGE OF AGRICULTURE, AGRICULTURAL EXPERIMENT STATION

ENTOMOLOGY

Vol. 1, No. 3, pp. 199-216 February 2, 1916

SYNOPTICAL KEYS TO THE GENERA OF
THE NORTH AMERICAN MIRIDAE

BY
EDWARD P. VAN DUZEE

The following keys cover all but eight of the genera of the
Miridae thus far recorded from America north of Mexico. These
elght genera were omitted on account of the want of material
for study or because their occurrence in this country is a matter
of much uncertainty. Dr. Reuter’s great work on the Capsidae
of Europe (Hemiptera Gymnocerata Europae, 5 vols., 1878-1896)
and his later studies in the North American fauna have formed
the foundation for the present paper, although the keys given
here are for the most part original. I have found it impossible
to work out his subfamilies of 1910 in a satisfactory analytical
form, and, while accepting them in my catalogue of our Hemip-
tera, I have ignored them in the preparation of these keys,
using only his tribes, or divisions as he terms them. In addition
to these tribes I have found it both practicable and useful to
establish groups of a lower category in two of the larger tribes
which have been denominated divisions with the termination -aria.
All synonymy has been omitted here, but it will be given in the
catalogue.

One fact comes out plainly in these studies: that certain
characters that are useful for diagnosis in one group may fail in
another. This arises from the well-known fact that a character
once discarded in the evolution of a group is never revived. Thus
we find that the hamus, or vestigial vein, found in the wing-cell

200 University of California Publications. [ENTomoLocy

in most of the Phylaria, is apparently always absent in the
Orthotylini where it seems to have been discarded, but in the
Oncotylaria, which is intermediate between these groups, it may
be either present or absent in the same genus, possibly in the same
species or individual.

The characters of the aroha present a similar case. Their
form seems to be constant for each tribe but in any, at least of
the larger ones, it may be entirely absent in certain genera. I
would not, however, consider the arolia a vestigial character as
is the hamus in the wing-cell.

In these keys I have attempted to arrange the tribes and
genera in what seems to me to be the correct descending order,
but here there certainly is a large field for investigation and
many changes will probably have to be made. The claspers, or
genital hooks, of the male form excellent specific characters in
many eases, but there are groups of species here and there in
which these hooks exhibit scarcely any appreciable differences
between what are undoubtedly good species.

The following are the eight genera omitted from the keys:
Neocapsus Dist., Pallacocoris Reut., Neoborops Uhler, Eccrito-
tarsus Stal, Teleorhinus Uhler, Cyllocoris Hahn, Orthocephalus
Fieb., Microsynamma Fieb.

As a matter of convenience the following terms are explained
here.

Arolia.—The pulvillae between the base of the tarsal claws, sometimes
free, sometimes united with the claws beneath.

Bucculae.—A narrow plate lying either side of the base of the rostrum;
rarely used in the Capsidae.

Callosities.—A more or less elevated area on either side of the anterior
lobe of the pronotum, usually distinguished by an impressed bound-
ing line, at least posteriorly.

Cheeks or gena.—The two sclerites below the eyes and between the
clypeus and gula. Between them is frequently a narrow segment
called the lora. The inner or upper cheeks may be nearly flat or at
times considerably elevated or tumid.

Clavus.—The inner area of the elytra next to the scutellum and separ
ated from the corium by the claval suture. It is usually long-
triangular in form, with its apex near the base of the membrane.

Clypeus or tylus.—The median lobe of the head below the front and
reaching to the base of the rostrum,

Collar or collum.—The narrow anterior margin of the pronotum., Gen-
erally separated from the disk of the pronotum by an impressed
line which may or may not be continued over the side.

Vou. 1.] Van Duzee-—Synoptical Keys to Genera of Miridae. 201

Corium.—The main portion of the elytra lying exterior to the clavus;
its outer margin being formed by the costa.

Cuneus.—A triangular piece, joined by a suture to the apex of the
corium.

Facial angle.——The angle between the line of the buceulae and that of
the clypeus when viewed from the side.

Fracture.—The notch between the apex of the corium and the base of
the cuneus on the costal margin of the elytra. .

Front.—The front of the head between the eves, below the vertex and
above the clypeus.

Gula.—The throat, or lower surface of the base of the head lying
beneath the rostrum.

Lorae.—The narrow segment lying between the upper and lower cheeks
at the base of the rostrum.

Membrane.—The membranous apical portion of the elytra. It carries
a looped nervure at base forming one large areole and usually a
second smaller one next the apex of the cuneus.

Scutellum.—The basal lobe is usually convex and separated from the
apical by a suture. This basal lobe is often more or less covered
by the base of the pronotum and in using this character allowance
must be made for the depression of the pronotum.

Tylus.—Same as clypeus.

Vertex.—The basal portion of the superior surface of the head between
the eyes. It merges insensibly into the base of the front.

Vestiture——The covering of hairs on the surface of the body. These
hairs may be soft or stiff, or they may be flattened and seale-like,
and are often deciduous and very easily rubbed off.

Xyphus (prosternal).—The triangular piece on the prosternum between
the bases of the anterior coxae.

In the Phylaria three new genera have been established for
which there are as yet no described species, and they are there-
fore invalid here, but they will soon be validated by the publica-
tion of species. These genera are: Leplotylus, Oligotylus, and
Strophopoda. One hundred and twenty-five genera are treated
of here which, with the eight omitted genera, make a total of
one hundred and thirty-three genera recorded from America
north of Mexico.

The following is a fairly close translation of Reuter’s key to
his subfamilies of 1910:

1 (16). Membrane biareolate, or with one areole distinctly dilated at
apex, very rarely without an areole but with several irregular
longitudinal veins more or less distinct. Elytra with a distinet
cuneus which very rarely becomes confluent with the corium.

2(8). Arolia large, free, approximate at base between the claws,

toward their apex very distinctly divaricate and frequently
dilated. 9. Mirinae

202 University of California Publications. | ENTomoLoey

(2). Arolia differently formed or wanting.

(5). Membrane distinctly pilose. Claws destitute of arolia.

8. Bothynotinae

5 (4). Membrane glabrous.

(7). Pronotum without a collar, but with its apical area gibbous-
convex, anteriorly frequently more or less produced above the
vertex; always roundedly produced posteriorly but not surpassing
the sides, the lateral margins attaining the apical. Arolia short,
united to the claws or wanting. First tarsal joint longer than the
second. 6. Ambraciinae

7. (6). Pronotum with or without apical collar; destitute of a gib-
bous posteriorly rounded apical area.

8 (9). Arolia wanting. First joint of hind tarsi long or very long,
rarely not longer than the second. Tibiae frequently mutic and
very distinctly more slender toward its apex. Wing-cell with the
hamus wanting or very rudimentary. 7. Cylapinae

9 (8). Arolia present, rarely wanting, in this case with the first joint
of the tarsi short, or the wing-cell with a distinct hamus, or the
body constricted at the middle. First joint of the tarsi very
rarely long, in this case the cell of the wing with a hamus or the
arolia present. Tibia very attenuated toward its apex.

10 (11). Apical joint of the tarsi more or less distinctly incrassate,
rarely sublinear. Arolia laminate, rarely short, frequently large,
always approximate to or connate with the claws. Tibiae always
destitute of spines. Lorae confluent with the cheeks. Cell of the
wings without a hamus. 5. Bryocorinae

11 (10). Apical joint of the tarsi linear, rarely a little thicker toward
its apex, in this case the arolia free and connivent at apex.
Tibiae frequently distinctly spinose.

12 (13). Prothorax with an annular collar at apex, in brachypterous
females sometimes obsolete above in the middle. Arolia none, or
with the arolia varying in length and closely approximated to
the claws with which they are connate, at least at base, fre-
quently for their whole length. Lorae linear, well distinguished
on either side. 4. Macrolophinae

13 (12). Prothorax without an apical collar, sometimes with the apical
margin slenderly depressed, in this case with the arolia free and
connivent at apex.

14 (15). Arolia free, slender, parallel or connivent at apex, very rarely
none, in this case the wing-cell destitute of a hamus, or the body
constricted at the middle, or the last two joints of the antennae
thicker than the others. 3. Heterotominae

15 (14). Arolia connate with the claws, very rarely free, in this case
closely approximated to them, sometimes expanded at apex with
the claws minute, faleiform; frequently narrowly laminate,
rarely none, in this case the wing-cell furnished with a hamus.

2. Phylinae

Vout. 1.] Van Duzee.—Synoptical Keys to Genera of Miridae. 203

16 (1). Membrane with but one areole, the vein mostly parallel with
the suture. Elytra destitute of an embolium and cuneus. Pro-
thorax without an apical stricture. First tarsal joint long.
Arolia none. 1. Lygaeoscytinae

Of these, the subfamily Lygaeoscytinae is Australian; the sub-
family Bothynotinae is confined to the Old World; the subfamily
Phylinae is equivalent in our fauna to my Phylini, Bryocorinae
to my Bryocorini, and Cylapinae to my Cylapini. Reuter’s sub-
family Heterotominae is the same as my Orthotylini, but his
typical division Heterotomaria was first founded as Litosomidae
by Douglas and Scott in 1865; but their genus Litosoma being
a straight synonym of Orthotylus Fieb., the tribe, or division of
Reuter, must be called Orthotylini and the subfamily Ortho-
tylinae. Reuter’s Macrolophinae embrace my Dicyphini (Macro-
lopharia Kirk., 1906 is antedated by Idolocoridae Dougl. and
Seott, 1865, the typical genus Idolocoris Dougl. and Scott, 1865,
being a synonym of Dicyphus Stal, 1858), and my Hallodapini
which is equivalent to Cremnocephalaria Reut. (first established
as Eroticoridae Dougl. and Scott, 1865, the typical genus Eroti-
coris Dougl. and Seott being a synonym of Hallodapus Fieb.,
1858). Lastly Reuter’s Mirinae include my Myrini, Capsini and
Horistini; the latter, termed Restheniaria by Reuter, was first
distinguished as Lopidae by Douglas and Scott in 1865, their
Lopus being equivalent to Horistus Fieb., 1861. It will be noticed
that Reuter has entirely ignored the work of Douglas and Scott,
who were the first to break up the great family Capsidae into
smaller divisions. That their divisions were sometimes made too
limited in scope and were termed families is no reason for ignor-
ing them entirely. Reuter uses the termination -ina for his sub-
families, which I have changed to -inae to make them conform to
modern usage.

KEY TO THE TRIBES

Apical margin of pronotum without a collar, swollen or elevated in a hood
NOONE) WHEY LOGIT) OIE WHNYE) NENA eee cee cerse eee oreo eee Clivinemini
Apical margin of pronotum not swollen or elevated in a hood above the
[SVE NSKE) OWE TELE). EGE See ee ee rere 1

1. Third tarsal joint thickened toward its apex; membrane in our genera
LENO OLE G G meee e eee aN eee Net Ole ee Rote Sa Rohe ee Bryocorini

Oo

—I

University of California Publications. | ENTomMoLocy

. Pronotum with a distinct apical collar, or with a flattened anterior

paneenefenu ay fspuanyolleeranalesl hy COVES Coa eee ee ee eo eee eee nee Pee CE 3

Px ono ban awalb ho wt eal (Coll cary eee eee eee cena ea ese a eee 9

. Pronotal collar convex, separated from anterior disk by a distinct
TIN CISO: CMM Ao Scn sof eats eee cee eee eee cee cas see ee cere cee ee oe eee 4
Pronotal collar flat, without an incised line behind it, or wanting;
body elongated) totem jim eer Sesser tee eeee cece ace cee rene aera eee rmeeereeaseees 8

. Head viewed from above short, vertical, produced below the eye for

nearly twice the length of the eye; antennae long and slender, much
longer than the entire body, inserted the length of the clypeus above
its base; basal joint of the tarsi as long as the following two
together; arolia wanting; vertex deeply suleate; pronotal collar very
slender sive sch eee SE Bees ae ee A ee eee Cylapini
Head not greatly produced below the eye; antennae rarely longer than
the entire body, inserted about on the line of the base of the
clypeus; basal joint of the tarsi shorter than the following two
taken together) ose ces cee aoc cle case e race cece acters ae oe nae 5

. Rostrum long, passing the middle of the venter; head produced, hori-

zontal or nearly so; tibiae smooth, or with minute pubescence only;
tarsi slender, basal point but little shorter than the following two

GO Me ther alr OLN elegy a TUG LI geen eee cee cece ese en eee Fulvini
Rostrum shorter, scarcely surpassing the hind coxae; tibiae armed with
bristles or clothed with longer hair, rarely smooth .....................-.--- 6

. Pronotal collar broad, convex, about as broad as the callosities; tibiae

thickly clothed with soft hairs but without rows of stouter bristles;
body opaque, black, marked with red or fulvous.................... Horistini
Pronotal collar narrow, convex, often linear, rarely broad; then flat
with the body elongated, tibiae smooth and pronotum broadest
before: te. maid cil eis ee ee eee 7

. Body elongated, often linear; base of scutellum usually exposed; tibiae

smooth or nearly so; arolia minute and united with base of the
claws, or as long as the claws and lying close to them........ Dicyphini
Body rarely elongated, with the base of scutellum exposed and tibiae
smooth or nearly so, arolia in this case free and divergent at
APO esi sse ee 8 eee ea ee Ee ce ee Capsini

. Tarsi long, first joint longer than the following two together; pronotal

collar a mere flattening of the anterior margin, or sometimes want-
ing; vertex often sulcate; arolia large, free, often clavate......Mirini
Tarsi shorter, first joint not longer than the third, usually shorter,
* pronotal collar often wanting; arolia united with the claws or want-
TN Peete ee eesees Re Soe 2 ee ee eee Hallodapini

. Arolia free, parallel, or converging toward their tips; wing-cell with-

COL 6 ee: el «W008 C= ne ae ee eee ry Be ae Orthotylini
Arolia wanting, or parallel with and usually united to the claws at
base, wing-cell normally with a hamus......---.--2...222-c:--cc---enorecee Phylini

Vou 1.] Van Duzee.—Synoptical Keys to Genera of Miridae. 205

KEYS TO THE GENERA
TRIBE MIRINI Douglas and Scott

Head exserted, distinctly narrowed behind the eyes, which are not con-
tiguous to the anterior angles of pronotum; pronotum with a dis-

YHOU OXON on I oe a Bey i a ee ee PE oor pe eee eae 1
Head not obviously exserted, eyes contiguous to the anterior angles of
FOG OU. G UTM OTseING Ais yar S Opec .ce seer ws seen rest sea sees e scene oe cee oe eee cu eae eee 2
1. Head strongly exserted, eyes being located at about the middle;
median suleus of the vertex short but distinet.......... 1. Collaria Prov.

— Head little exserted, eyes being located near to the hind margin of the
head but distinctly separated from the anterior angles of pronotum;
vertex transversely depressed between the eyes, median sulcus

TAME, Ors GPPUNES OLSON ENS) ornare eee 2. Miris Fabr.

2. Base of pronotum truneated or a little emarginate at the middle........ 3
— Base of pronotum coneavely arcuated, leaving the base of seutellum
[OvsRORE KOU Dyye eb se oN OPES NG es ere ee ee ee er ee ee ee eee 5

3. Basal lobe of scutellum covered by pronotum............ 3. Stenodema Lap.
== IBe geil Moos) one Sco uefa og OSC | a eee 4
4. Body narrow, elongated; head one-third longer than broad, nearly or
quite as long as’ the pronotum’ --........2...2.2-2.....-- 4. Megaloceraea Fieb.

— Body more ovate; head short, not longer than broad, much shorter than
Te Way ONTO CUITIN ae eae eee earn reek a aa te oe canteen 5. Mesomiris Reut.

5. Large areole of the membrane entirely hyaline; head long and pointed,
Withethelmediansulleus) deep) se sce--ee-eee eee! 6. Trigonotylus Fieb.

— Large areole of the membrane, or at least its outer half, opaque pune-
tate; head short, transversely flattened at base, median sulcus
Obsolete mor ne aulivg SO} eeescccce sss cnc csete esses ess eeezes tee 7. Teratocoris Fieb.

TRIBE HorRISTINI n.n.

Head short, vertical, when viewed from the side nearly square at apex,
gula almost obliterated; elytra parallel, or the costa regularly and
feebly areuated; second joint of hind tarsi not more than half the
Teanga ea OLE ELAS TAN TH esc cee ee ee ee eee eee it

Head a little oblique, when viewed from the side distinetly produced and
narrowed toward the apex, gula quite long, oblique; elytra con-
siderably expanded beyond the middle; second joint of hind tarsi
nearly or quite as long as the first -............-...---.----- 2. Opisthuria Reut.

1. Vertex and front more or less convex, front not at all tumidly pro-
jecting before the clypeus; sides of pronotum anteriorly carinate
only across the incisure separating the colluin....1. Platytylellus Reut.

— Front tumid, projecting prominently before the base of the elypeus;
pronotal margins anteriorly carinate to behind the callosities -.........
i he a Oe A 3. Oncerometopus Reut.

TRIBE CAPSINI Reuter

KEY TO THE DIVISIONS

Body linear, constricted at the middle; pronotum swollen at its middle
and as wide there, or wider, than on hind margin....1. Myrmecoraria

206 University of California Publications. |ENToMoLoGy

Body rarely linear and constricted at the middle, pronotum in this case
widest: Webi a. 2.cccccce sc ste saeco casas woos psec ce cemee cee eccsstenesuepacestonbonteccesepees 1

1. Arolia free, divergent, usually more or less curved and clavate .......... 2
— Arolia absent, their place taken by two parallel setae; membrane often
uniareolate; body robust, polished ..................-.---.----- 5. Deraeocoraria

2. Body above impunctate, or with fine aciculate punctures only............ 3
— Body above, or at least the pronotum, coarsely distinctly punctate,
polished VeallosubieskpomOmin Mb ecesecennecesecesese eens eeecnereenene renee 4. Capsaria

3. Form more elongate, parallel or subparallel, cuneus at most but slightly
depressed and the fracture smal] ..-.....-2-.-....c-ccesse-e-core 2. Phytocoraria

— Form more ovate, elytra more distinctly convex, the cuneus strongly
deflexed and the Hractumes deep ---scsceceesss-eeeeene === 3. Dichrooscytaria

DIVISION 1. MYRMECORARIA Reut,

Head constricted into a short neck behind the eyes; clypeus prominent,
convex, its base distinet from the front; pronotal collar with a dis-
tinct) amieis ede lime moe hand eseeeweseesenee ane sen ease eee 1. Mimoceps Uhl.

Head not constricted behind the eyes; clypeus depressed, merged with the
front; stricture of pronotal collar evenescent at its middle ~............
Sach euscbastesyeatecetcs tet sceseaebemsnscatcetee Pacts Sacadacdt cassuesseasasbedeeeenen 2. Pithanus Fieb.

DIVISION 2. PHYTOCORARIA Reut.

Bod yaa owerop acre eum dls (Url Cait Cee nseeeeeneeenneenceceeseneeeeeeceneeressmeeemeaneeneeneaee 1
Body above more or less distinctly polished, sometimes shagreened or
aciculate-punctate: and allmost ope Cie gece = seeeee areas mene eee tarec ns aceneneee 4

1. Form linear, constrieted at the middle; pronotum produced, almost
cylindrical before; vertex suleate ................-.-.---- 1. Paraxenetus Reut.

— Body not constricted at the middle, pronotum trapezoidal; vertex not
Olwlousliy, Sullic ait © yee sees ec e cee c cree ne ne eee ee a ae ase ee 2

2. First antennal joint thickened and clothed with flattened hairs ............
Seek cee ee ee es Se ca eae Ae we ee Sea rere et 2. Neurocolpus Reut.

— First antennal joint without flattened hairs...............2.22..222:2:-2seees-e-- 3
3. Hind femora linear, terete or nearly so ................------ 4, Ecertebia Reut.

— Hind femora ligulate, flattened, broadest near the base and tapering
fromm) mind dil Gib On ap Cesena setae cee ct cee Secs eee 3. Phytocoris Fall.

4. Head not or searcely vertical, when viewed from the side distinctly

narrowed below antennae: cula Oblique: eee eee cee seeecenees 5

— Head vertical, thick and cylindrical below antennae; gula nearly or
quite parallel” swaths ‘ty lus) Soe aaa aces eee nears tana ese en tree seee een mentee 10

5. Second antennal joint strongly clavate, fusiform, more tapering toward

DEUS CYS seoeey oe aro cn eee Se ee ee 10. Garganus Stal.

— Second antennal joint sometimes moderately thickened but not strongly
Clavate:jci=ta8.2 BAe Se Se ee ee 6

6.. Vertex not suleate atibase: 22.2 se cece eccecee cerca neces eee 7

— Wertex sulcate wat: base cccccccciccs eile ccctecctecsssces cect eetest seca career ance 8
7. First joint of hind tarsi shorter than the second.....9. Ganocapsus Van D.

— First joint of hind tarsi much longer than the second....8. Stenotus Reut.

Vor. 1.] Van Duzee.—Synoptical Keys to Genera of Miridae. 207

8. Face opaque, distinctly obliquely striate; suleus conspicuous; clypeus
not at all polished; callosities inconspicuous........ 5, Creontiades Dist.
— Face polished, without distinct striae; sulcus inconspicuous; clypeus
polishedss Callosities: CONSPICUOUS! 22-.2.en.cececcceccce crete wceseeceseceeecerereseecesnes=s 9
9, Sides of pronotum carinate; collar broad; head subhorizontal; eyes
Sra et ero Wo I Cyl © pee eee nes eae cote ce cuca cereue-sbcacesveesvies 6. Allorhinocoris Reut.
— Sides of pronotum ecarinate, rounded; collar very slender; head nearly
vertical and thick at apex, extending but little below the large
YU GURLEORN CEN GIES Sere cee eee e 7. Adelphocoris Reut.
10. Second antennal joint clavate, clavate portion occupying the apical
third and strongly flattened and suleate above; vertex without a

suleus; surface polished mude ................-......-.. 11. Ectopiocerus Uhler
— Second antennal joint linear, not at all clavate; surface above clothed
AVAIL) OV G2, CTIGYSVE) TORU OLESEN LE) nee ee ee all

11. Whole upper surface closely, minutely shagreened, giving the insect an
opaque aspect; vertex sulcate at base; antennae inserted much
Bel Owasthe CviCS tcc voces crea cece fede stacovce cn tacdssuseeeesteveaes 13. Thyrillus Uhler
— Upper surface more polished; base of vertex with a transverse groove
but searcely prolonged at the middle in a sulcus; antennae inserted
close against the lower angle of the eyes ................-- 12. Irbisia Reut.

DIVISION 38. DICHROOSCYTARIA Douglas and Scott

Upper surface opaque or nearly so, clothed with short pubescence -.......... i
Upper surface highly polished, nude; form broad-ovate; cuneus strongly
le El esse clearness ce eet ewe ar ae ee ee NS eat a Aa ee Seg Sees ed Aes ae 6
1. Pronotum marked with a pair of round black points, occasionally
Wanihinomnesp ecimens) nob) tulllivs Colon Ul eee se cece weenceneeetesencenee ae 2
— Pronotum without the pair of round black points 4
Pebasalejoimbsotantennae terete. Or mMearly:, SO) 22ccesc-c--seseceeee nen sneceeenneeser ee 3
— Basal joint of antennae strongly compressed, nearly as wide as hind
CQ TYN ©) 1,51 meee Se wat ee sere ae ES Pee ee 2. Lampethusa Dist.
3. Second antennal joint linear; first joint with but few minute hairs,
not, or scarcely, longer than the head ...................... 4, Calocoris Fieb.

— Second antennal joint thickened toward the apex; first joint densely
pubescent, hairs nearly or quite as long as the thickness of the

STOO eo Sea 3. Paracalocoris Dist.

4, Antennae stout, second joint strongly clavate, third and fourth abruptly
slender and together scarcely longer than the first; body broad oval,
densely sericeous-pubescent; scutellum tumid.... 1. Pyenocoris Van D.

— Antennae slender, second joint linear, third and fourth joints together
MenrlysoLrquLteras ons; asthe) Sec om dl Sass ses enc teee sean eee 5

5. Head exserted, the small rounded eyes not overlapping pronotal angles;
base of elypeus when viewed from above much anterior to the inser-

GTO OtepeDMNG OTN AIC pastes eee eee oo ee wee es 5. Poeciloscytus Fieb.

— Head broad and short, closely set against pronotum, the large eyes
overlapping its anterior angles; base of clypeus when viewed from
above in a line with base of antennae ............ 6. Dichrooscytus Fieb.

208 University of California Publications. | ENtTomoLoey

6. Rostrum long, reaching at least to intermediate coxae; pronotum with
obscure scattering impressed points, but not at all punctate —......
wos ccbaebedecsssasiaccesutean ie ceestee Seva scdetucescnscesssdeececcuas sock “Woceetnctecearess 7. Horcias Dist.

— Rostrum short, not surpassing anterior coxae; pronotum obscurely
PUN HUE ds ake eo sk aecccnccs cnsed se este sccebaceutuscust-basee 8. Poecilocapsus Reut.

DIVISION 4. CAPSARIA Reut.

Vertex suleate and transversely striate; second joint of hind tarsi much
ShortersGhamerins Gear det bin caeesesensesees eres enema 4, Platylygus Van D.
Vertex more or less polished, scarcely striate or suleate; first and second

Geir ali] OT NGS eS Uy e Cyr elle eee see a ee 1
1. Elytra nearly flat, cuneus at most but moderately deflexed __..........-.... 2
— Elytra more convex, cuneus much deflexed and the fracture deep...... 6

2. First and second antennal joints rather thick, the second linear and
searcely thinner than the first, or in the female slightly attenuated
Git DASE ;AMG AY CR eae ace casa sere ca scene sac cece eee ec seran net ae aees Seer ae ae see enmeeeeees 3
— Second antennal joint more or less distinctly thickened toward its
apex; sometimes sublinear but then distinctly thinner than joint

OMG: 6 22.2ho5cdeg8k setae cose cin etch sense ec Sac oe See ns Sates Ss acs en cas Bae eae ee emcees +
3. Base of vertex flattened, hind margin carinate; base of scutellum but
little exposed; elytra oblong, parallel ........................ 3. Lygidea Reut.

— Vertex convex, polished, its base ecarinate; base of scutellum broadly
exposed; body oval, distinctly broader behind the middle; color

TG Ch cee eee ee ere NE ae cs StL oS BR Bee oe) a AE 2. Coccobaphes Uhl.

4. Third and fourth antennal joints abruptly thinner and _ together
scarcely more than half the length of the second joint, which is
linear and moderately thickened but thinner than first ........-........-..

ER Tne Atk NAT OE en MAAR RIES GRE oA AEE ee Tak | 6. Tropidosteptes Uhl.

— Antennae slender, third and fourth joints setaceous and together at
least: two-thindssthenlenethvotav be 7S ec or Ce ssseseerse ese eeeenet near rarenenass 5

. Form more ovate; sides of the pronotum carinate......7. Neoborus Dist.
— Form more elongated and subparallel; sides of the pronotum ecarinate
ore UN NS ioe Coed Banas este etn sans aes earet vege aoe 8. Xenoborus Reut.

6. Inner cheeks tumidly convex, forming almost a tubercle beyond base
of antennae; second antennal joint clavate; pronotum coarsely
punctured; head broad behind and concentric with the anterior
Margin), OL promot uy Lele. see sce sees ences eae scenes 1. Capsus Linn.

— Inner cheeks convex but not prominently tumid; second antennal joint
but little thicker at apex; pronotum more finely punctured; eyes
rounded behind, head not concentric with the anterior margin of

PON OT OGM eins aoa cess t cece cetew nescence vane rsewaceeoecoroestencaesnanesenee 5. Lygus Hahn.

ON

DivISION 5. DErRAEOCORARIA Douglas and Scott

Vertex transversely striate and longitudinally suleate; second joint of
hind! tarsi much shorter) thant first em tian cle eee il
Vertex more or less polished, searcely striate or sulecate; first and second
tarsal! joints sulbeqtall) <-22.ssccco--. coc 2 cesta cessececueessnecscsescevecesaet oer aeeeeeees 2:

Vou 1.] Van Duzee.—Synoptical Keys to Genera of Miridae. 209

1. Second antennal joiut clavate; third and fourth short and thick, fusi-
form; prosternal xyphus convex, but slenderly margined ....................
ee ..-.2 Diplozona Van D.

— Antennae linear, of nearly equal thickness throughout; second joint
searcely enlarged at apex, third and fourth linear....1, Cimatlan Dist.

2. Elytra punctate, not bullate behind; basal joint of hind tarsi not pro-
duce daib.el owambey,Om date SCC OTC eee s sens anne eee ene eee 3}

— Elytra impunctate, bullate behind, cuneus almost vertical; basal joint
of hind tarsi thickened, oblique at apex and attaining the apex of
second joint; head nearly vertical, but little produced before the
QOS et tence nee a ns ces es ce cssctst sua ss cesvasecvasneseselsseceestcee oes 4, Klopicoris Van D.

3. Head strongly produced and nearly horizontal, surpassing apex of
short first antennal joint; second antennal joint thick and very long,
nearly linear, longer than the remaining three joints taken together;
sides of pronotum ecarinate ..............-....-.. 3. Eurychilopterella Reut.

— Head less produced, not surpassing middle of basal antennal joint...... 4

4. Sides of pronotum carinate, antennae rather short and slender, apex of
the second joint distinctly thickened; membrane often uniareolate;
xyphus sometimes convex on the middle ........ 5. Camptobrochis Fieb.

— Sides of the pronotum ecarinate; antennae longer, basal two joints
stout, the first surpassing the apex of the head by two-thirds its
length, second a little thicker apicallv............ 6. Deraeocoris Kirsehb.

TRIBE BRyocoRINI Douglas and Scott

Form oblong, more or less elongated, elytra parallel or subparallel............ 1
TONE SHAG MELETES OYENLEEY (ONE HEU OO 12S ee ee a 4
1. Eyes on a suberect stylus which is at least as long as the width of the
ORO) eoteceencn cc Sees eee 9. Hesperolabops Kirk.
SPV CSMSCSSI Or eatmmMOSt mo UbeSUbs tila te) sess ce see cee crews seten ness eeeeee ence 2
2. Eyes large, exserted or substylate; callosities convex, oblique, con-

tiguous at middle of pronotum, leaving a transverse triangular

JORUHACEURSYOL EWE, LONER EOS) ees crease cee ee 8. Caulatops Berger.

— KHyes smaller, not at all stylate; callosities more transverse, not con-
OWES GE Waa KTMNG) ONE [ORONO UNM ene ese ee rere eco ee

3. Body opaque, pubescent; second joint of antennae long, about equal-

imew basally waldithyons mom ob Uae snes ae cee cece eee 7. Dacota Uhl.

— Body smooth, more or less polished; second joint of antennae short,
MONG Utorayerere qlee rar aya lino Cope alk ENG | eye ee cee ee eee 6. Sysinas Dist.

4, Pronotum with a prominent linear collar .............. 5. Monolocoris Dahlb.

== IPAROMOHEDIAM yaleloyoyy BY Chis wrbaK Oy COCO) UE a eee eee eer ee eer ee 5
5. Seutellum without a triangular discal impression .........222..220..2.---e 6

— Scutellum with a triangular discal impression ~..........222----.2-:-:--ee--0---=- 7
6. Embolium broadly expanded, about as wide as hind femora; pronotum

strongly convex and bullate behind, with three longitudinal impres-
ST OTIS eee ecw ea ed PRES st eis ae ere 4, Pycnoderes Guer.
— Embolium linear; pronotum convex but not at all bullate or longi-
tudinally impressed behind ..................-2.------0---0-+- 3. Sixeonotus Reut.

210 University of California Publications. [ENtomMoLocy

7. Antennae inserted close to apex of the eye, basal joint very short, one-
third shorter than width of front .....................- 2. Halticotoma Reut.
— Antennae inserted some distance above apex of the eye, last joint
considerably longer than width of vertex and much surpassing
CON 14 0 SS ae eee oe Ee Se eae er EEE 1. Cyrtocapsus Stal.

TRIBE CLIVINEMI Reuter

Second antennal joint linear; sides of pronotum without a distinct carina
wotevs Sidaccccceicecckse-cs thas senecatecassteees cee sessessbeseccencessncnsemeneceeueeese 1. Clivinema Reut.
Second antennal joint stout, clavate, apical two short and abruptly slender;
sides of pronotum distinctly carinated ................ 2. Largidea Van D.

TRIBE CYLAPINI Reuter

Form oval; head short, vertical; vertex with a deep longitudinal impres-
sion; antennae very long and slender, much surpassing tip of mem-
brane; basal joint thickened, fusiform .....................----- 1. Cylapus Say

TRIBE FULVINI Uhler

Body elongated; costa but feebly arcuate; sides of pronotum concavely

arcuate, humeral angles prominent ._.......---2.---22c-secceece--- 1. Fulvius Stal.
Body broad-oval; costa strongly arcuate; sides of pronotum not at all con-
cavely arcuate, humeri not prominent .................. 2. Peritropis Uhler

TRIBE HALLODAPINI n.n.

Marsaliclawsiwathyvaroliay mime) Oxy wien oy estes scree cee ee semanas eee mee 1
Tarsal claws with long parallel arolia; females sometimes wingless, for-
MIUCIFOTM 222.5. 401... ee eRe ee ee ee ee 5

1. Hind margin of pronotum with a median spine, behind which the edge
IS MO CCH) x1 s8 wee oD ee ee ee 1. Dacerla Sign.

— Hind margin of pronotum without spine or noteh .........22....22:::2200--------+ 2
2. Head short, vertical, but little produced below the eyes .....................--- 3
— Head long, oblique, produced below the eyes for a distance nearly as
preat as themlenp thy ofthe eye) eecce essere ee eee 4

3. Posterior lobe of scutellum tumidly elevated....2. Cyrtopeltocoris Reut.
— Posterior lobe of scutellum transversely moderately convex, hori-
ZO tally, Ae See NO eae se ee ae ee Ween ee ee 3. Sericophanes Reut.
4. Posterior lobe of scutellum moderately convex, subearinate; sides of
pronotum almost rectilinear, a little curved outward at the humeri
seantalen shells tec Maczech Mth EIS eae oe toons es 4, Closterocoris Uhler
— Posterior lobe of secutellum tumidly elevated, subconical, as high as

DEUS © Osta TUG Ta ee eae eee ere 5. Cyphopelta Van D.
5. Hecond antennal’ jounitr clawateses.cs coe eeeeeeeeeeeeeeee 6. Orectoderus Uhler
——) econd. antennal joint limes) meses seen erence 7. Coguillettia Uhler

TRIBE DICYPHINI Reuter
Hind margin of pronotum rectilinear, or slightly concavely arcuate; head
vertical before, when viewed from the side not projecting before
eyes; hind margin of callosities located considerably before the
uaa BKGKG UKE) COME, LANs) YOWON AVON BEN OL Seo nee cpeccan RoR EeE aN SOSESC ECCS 1. Hyalicdes Reut.

VoL. 1.] Van Duzee.—Synoptical Keys to Genera of Miridae. 211

Hind margin of pronotum concavely arcuate, leaving base of scutellum
exposed; head sometimes nearly vertical but then narrowly but
distinctly surpassing front line of eyes; hind margin of callosities
nearly or quite attaining middle of pronotum ................2...22::.-200--0-- il

1. Head when viewed from the side distinctly produced and oblique;
angle of face (angle of tylus and bucculae) subacute; eyes small

and oblique, placed at middle of the head .......... 4, Macrolophus Fieb.
— Head vertical or nearly so; apex of head truncated, facial angle a
reel og, Byavedlas yyy} le wekets) CUAL ASCE ere ee eee eee 2

2. Head produced in a distinct neck behind eyes, space behind the eye
when viewed from the side about as long as width of the eye
accep SSSR aE cone eS SER Se aS ae ea 2. Dicyphus Fieb.

— Head searcely produced behind the eye, forming but a very slender
margin there of about the width of pronotal collum ......................----
a ee eee ete Leh PI a Nee ere Ese eee 3. Engytatus Reut.

TRIBE ORTHOTYLINI n.n.
KEY TO THE DIVISIONS

Head broad with eyes stylate, their inner margins being beyond the pro-

SA OSLECEUL Ce fe KS S eiene er e e e ne 1. Laboparia Reut.

Head sometimes broad but eyes not at all stylate —....22..22eeeeeeeeeeeee eee 1
1. Head broad, hind margin sharp, concentric with or overlapping an-

(CMON TMMENEANTA, Ole TOWN AOL AUIN Sees re eee ee 2

— Head not unusually broad, not concentric with or overlapping anterior
margin of pronotum; eyes rounded behind .............0...--..---...--eecee-eee-- 3

2. Form broad-oval; hind femora often broad, saltatorial; clypeus distinet

TEYECOW, “Cela E) SONA se ee a 2. Halticaria Kirk.

— Form elongated, body often constricted at the middle; hind femora
normal; clypeus depressed and fused with the front ......-....000...2...--
ec Reh el te AE nk 3. Pilophoraria Reut.

3. Apical two joints of antennae not thinner than the second ......................
aN er A a eer es TE ie cae ach ig ee AG 4. Ceratocapsaria n.n.
— Apical two joints of antennae abruptly thinner than the second, seta-
EXD A TASER 2 IO a aa ee ee 4
4, Base of vertex and sides of pronotum sharply carinate; anterior mar-
gin of pronotum sometimes distinctly elevated in a ridge by a trans-
verse depression before callosities; body opaque....5. Lopidearia n.n.

— Base of vertex and sides of pronotum sometimes more or less carinate;
anterior margin of pronotum in this case not at all elevated ............
spec COCO CSSSEELEEEEE EES re ep ae ee Se a aS 6. Orthotylaria n.n.

DIVISION 1 LABOPARIA Reut.

Simtmonem NOmbhe Am eric am PONUS) c....2.6-<cs-cx-c-2eccceecesee--eceeceecsteceeseles Labops Burm.

DIVISION 2. HALTICARIA Reut.

Antennae long, setaceous, as long as entire body; hind femora greatly
HA TeMe dems allibait OVO eer s--ereese 2. cteee sie. ese eeeee 8 eee 1. Halticus Hahn.
Antennae seareely longer than elytra; hind femora not greatly widened
ORAS Oi eNO Tel etl lyesse ee ao nes ese, we eee es NE 2. Strongylocoris Blanch.

212 University of California Publications. | #NtomoLoey

DIVISION 3. PILOPHORARIA Reut.

Head with eyes little wider than anterior margin of pronotum; sides of
pronotum not arcuated; body not constricted at the middle; upper
surface ornamented with dots of silvery hairs ..........-......--:-t-s--s-e0--+°
eI ee Ae ee ee a Res Re Se Sh yet ....-.......-L. Heterocordylus Fieb.

Head much wider than anterior margin of pronotum; sides of pronotum
arcuate; body constricted at the middle; elytra sometimes with
LLANMS VETS el Mine Smo feasillivie Tayam lca S)eeeseenee eine eee ee il

1. Second antennal joint distinctly clavate; body constricted at the
middle; elytra usually ornamented with transverse lines of pale
scaleslike: hairs) ee ee ue A Ao See 2. Pilophorus Hahn.

— Second antennal joint almost linear; body scarcely constricted at the
middle; elytra without transverse lines of scale-like hairs -.................
St atROL OEE A Te Tt ENON Ce en eed ree Meee a 3. Alepidia Reut.

DIVISION 4. CERATOCAPSARIA n.n.

Pronotum anterior to the middle nearly cylindrical, then rather abruptly
flaring to the humeri; elytra sparsely clothed with long hairs ............
Sl Sates AE AN EE IS ee Nn ne ee 1. Pamilia Uhl.
Pronotum regularly narrowing anteriorly, its sides not constricted at the
TUN CL LIM socks Sacco ee ee oe ae eee nee 1
iP eElytraypanalllele mh ealdkaye rite ai eeeene. see sees ene ee ese eeseeee eee 2. Tiryas Kirk
— Elytra with costa more or less arcuate; head obviously oblique -.............
SEA A MAREE Dee we PRS PALEME DED Re hee Re Ee OTE BPE ey OF fh 3. Ceratocapsus Reut.

Division 5. LOpIDEARIA n.n.

Vertex prominent, convex, its base strongly carinate across its whole
width; anterior edge of pronotum not elevated....3. Hadronema Uhl.
Base of vertex carinate only at the middle, carina not reaching the eye,

anterior Maron OF pronotumiyel eyjalt ed ee eeeeee ee cece aeegn eee ene 1

1. Basal two joints of antennae incrassate, the second narrowed toward
its apex and sometimes flattened in males........ 1. Lomatopleura Reut.

— Second antennal joint linear or nearly so...................------- 2. Lopidea Uhl.

DIvISION 6. ORTHOTYLARIA Douglas and Scott

Pronotum constricted behind eallosities, constriction continued over the

sides; body: elongated. sek.ci 3 oes coses obo ekece csc cece cee cee i
Pronotal stricture if present not continued over the sides ...............--.-------- 2
1. Posterior coxae distant; basal joint of antennae a little longer than

head; pronotum with a collar-like constriction -................-..--.0------00----=-

ksi cdscasveial uaeks Sea tiha ba 2 ere eee eee oan ae ee 1. Pseudoxenetus Dist.
— Posterior coxae contiguous; basal joint of antennae short, searcely
SUTPASSLNG: Clip CUS: xia le. cec sce c sees cece ee cee cco eemeeee a eeeaer sees 2. Globiceps Fieb.
2. Eyes placed near or before the middle of sides of head .......................- 3
— Eyes placed on hind margin of head, vertex not continuing around
behind, the Cys. a2 2.2..cs<.sccks. 2500 sae sss 22 sacs densa sessnuccesssusSheee eeu tee see eee ene eeeeeeee 4

3. Head a little triangularly produced before the eyes; vertex and front
together convex, base of the former rounded over and not at all
earinate; elytra parallel; membrane biareolate....3. Paraproba Dist.

Vou. 1.] Van Duzee.—Synoptical Keys to Genera of Miridae. 213

1

GO

10,

ILM

Head truneate before, not produced before eyes; vertex broadly ex-
cavated, base arecuated and carimate; elytra broad, the embolium
well developed; areoles of membrane merged into one, dividing
nervure nearly or quite obsolete, their surface similar in structure
to the diaphanous corium and cuneus ................ 4. Hyalochloria Reut.

Eyes rounded behind, in conformity with the curve of the base of the
vertex, thus bringing eyes away from pronotal angles; base of
vertex ecarinate in the male, very obtusely carinate in the female;
elytrasampleytlart ssi iiyalln me) sess. scsecsre ese cesee teases 5. Diaphnidia Uhl.

Hind margin of vertex and eyes forming nearly or quite a straight
line, thus bringing eyes into proximity to pronotal angles .............. 5

Tender whitish insects with elytra a little wider than humeri; basal
joint of antennae lineate with black exteriorly; vertex ecarinate.
NN A ee ede Sees Seasan teeters JccuddWoak state cuazvesecsdes gecacvécecettsseent« 6. Reuteria Put.

Sometimes tender and whitish, then with vertex carinate at base and
HAS MEST HOUT, Ayial AORN G eh [OMEN TEAS) eee een es ese 6

Vertex with a distinct carina at base behind a transverse impression,
this impressed area sometimes with a foveate dot on either side.... 7

Vertex without a distinct basal carina; sometimes tumid at base,
simulating an obtuse carina, with surface before it broadly tri-
amo malar Ce presse Ce testes teen coos eos cscatas ce oe = cceuacce¢-cesnsa len scoucteceesesteer=eees 8

. Head viewed from the side short, apex never produced the length of

the eye below its lower angle; pronotum without a round black
SPO lowlewinGl Colo ys eC) Sessa eee 7, Orthotylus Fieb.
Head viewed from the side longer, apex produced at least the length
of the eye below its lower angle; pronotum with an impressed
round spot behind outer angle of callosities.............. 8. Ilnacora Reut.

. Head vertical, with an oblique impression either side on vertex;

clypeus prominent, convex, well distinguished at base; body above
smooth, clothed with deciduous black hairs and minute silvery scale-
like hairs (type Macrotylus angularis Uhl)....11. Pseudopsallus n. gen.
Vertex without an oblique impression at base ................--.----------0-000-0- 9
Pronotum with its base elevated above base of scutellum, strongly
declinate anteriorly, its vertical height about equal to that of head;
basal two joints of antennae flattened; body opaque, hirsute, elytra

marke diwabhne blachesam dl wil ite =... - cece seeeseecenceesesesceesans 9. Semium Reut.
Pronotum not greatly raised above base of vertex and little above
the level of scutellum; antennae not flattened ...........-.2..--...-:----e-0-- 10
Hamdgrenlonamnonmial, moteseultat Orval sess sce nceres cceese nse. esses ssaceseeceae 11

Hind femora very broad (nearly one-third its length), saltatorial; head
short, vertical; vertex ecarinate; basal antennal joint short. scarcely
SUT MASSIMO Hay OX Oi WU VINIS) eecesnccececcee eco ceeesctessnsecee= 15. Parthenicus Reut.

Head transverse, vertical, viewed from above not projecting before
eyes; pronotum campanulate, humeri prominent; males with their
second antennal! joints Clawate....---.2c--2.2---2-2c2--ee--ee=-- 10. Mecomma Fieb.

Head viewed from above angularly produced before the line of the
eyes; pronotum trapezoidal, humeri not abruptly prominent; an-
(E@IBTNENE: TENSE: CVE EE Se csc or ee RRR eee 12

214 University of California Publications. | EN tToMoLoGy

12. Head vertical; dimorphic, females ovate, with eyes strongly arcuated
and membrane abbreviated, males with elytra parallel; green or
ereensandmblackw in Se Citisy esses cesersaes eae eee 12. Labopidea Uhl.

— Head oblique, produced before eyes for about length of eye -............... 13

13. Large green ovate species, with the body narrowed before and behind,
tylus moderately produced; rostrum reaching to middle of venter
(type Macrotylus vestitus Uhl.).............-..------ 13. Macrotyloides n. gen.

— Smaller and more slender species, with the body linear; tylus com-
pressed and semicircularly prominent, occupying about half the
length of head beyond apex of eyes .............. 14. Argyrocoris Van D.

TRIBE PHYLINI Douglas and Scott

Prosternal xyphus depressed on its disk, its margin more or less elevated;
arolia rather long and flattened, usually united with the claws ........
SD Ae eb Mirae ahs SRS MPR Te Division 1. Oncotylaria Reut.
Prosternal xyphus convex, immarginate; arolia short and united with the
Claws: Osawa: bin oes: ee Division 2. Phylaria D. and S.

DIVISION 1. ONCOTYLARIA Reut.

Tarsal claws short and strongly ineurved; arolia free, laminate, as long
as the claws; clypeus prominent, viewed from the side strongly
CUT VIER! (222. ee TA Sie Correo eee cee eee 1. Macrotylus Fieb.
Tarsal claws longer, nearly straight, or somewhat curved toward their
POX seal ee ee ee 1
1. Head long, rostrate-produced, when viewed from the side projecting
before the eye for about twice the length of the eve; prosternal xyphus
sometimes moderately convex, but a distinct marginal carina can
be seen when vestiture is removed; body broad, cvate, much nar-
rowed before; surface clothed with a fine whitish vestiture inter-
mixed with stiff black hairs ........................--- 2. Haplomachidea Reut.
— Head not produced before the eye for more than length of the eye;
body oblong or elongated, clothed with a minute or uniform vesti-

2. Clypeus prominent, its base but poorly distinguished from the front,
placed much above the line of antennae; basal lobe of scutellum
much exposed; arolia long, exceeding apex of claws, with which
they are united for their whole length............ 3. Onychumenus Reut.

— Clypeus moderately prominent, its base well distinguished from the
front and placed a little above the line of antennae; base of pro-
notum rectilinear, leaving basal lobe of seutellum but narrowly
exposed; arolia slender, becoming free toward their apex and sur-
passing the middle of the claws.....................:c200---++ 4, Oncotylus Fieb.

Division 2. Puy arta Douglas and Scott
Head more or less produced; when viewed from the side having facial
angle (between bucculae and line of elypeus) less than a right
ON Ge: s...5.cceteeceee ese e Seceets Se ee eee ee 1

Vou. 1.] Van Duzee.—Synoptical Keys to Genera of Miridae. 215

Head not or scarcely produced, when viewed from the side having the

facialeangle nearly or quite a right) angle: <2 cee ceccccceccccecccccecceeceeeeee-- 10

. Clypeus broad, depressed; first antennal joint not or searcely sur-
FOES SUT O MC liv} @ US eeece-kecce: Sec eeeaets (tcc ssa. ciesectscuastecssstsenaceds 3. Sthenarus Fieb.

Olli DeuSm oN. MIN ET IT pC OTC Xap oe coc ae 2 nee cs saeeen sc cen Saaene se cunu tern scesvaseeheeetecsen sees 2

. Femora black or pale and dotted in longitudinal series; rarely pale

without dots, then with tibiae dotted and marked with a darker

SE VOPR Sy OKO CEN FOLENS Ss ee eo 3
Femora pale and irregularly dotted with darker or black; often with-
out dots, then with tibiae pale without black points ........................ a

. Head rostrate-produced, projecting before the eye for a distance much

greater than length of eye; second antennal joint broadly flattened
nneGie mma Lee sheml Ora) ila Cksessceeccesesseeseccceeseccereseeceeceees 5. Criocoris Fieb.
Head not produced before the eye for a space greater than width of
the eye; second antennal joint linear in both sexes ............22.-2..------ 4

. Femora black; body above conspicuously clothed with white hairs.

ceesetect “bcos SSS ECS EEE EEO ee a ee eee ee ee 10. Apocremnus Fieb.
Femora pale, dotted in longitudinal series; rarely black, then the body
abowemwabhka tines pale) pubescence onlliv, sees secre neerce ates eeeee cence 5

. Tibiae dotted; antennae with second joint uniformly colored, or if

panty iolored paler at middle or apex: <.cc-s:2-cce--ce-e-neeeeesenceseseeeeeeeseseee 6
Tibiae pale without dots, bristles only dark; femoral dots sometimes
obsolete above; antennae with second joint black on apical half.
See cee ASAECEL ESSE a oe eS 7. Rhinocapsus Uhl.

6. General color of whole body red or reddish-brown ..............--..---.0-----------
2 8. Gerhardiella Popp.

— General color pale, whitish or tinged with yellow, marked more or less
with fuscous or black areas, sometimes entirely black ......................--

osc esa SEER a 9. Plagiognathus Fieb.

7. Femora pale and irregularly dotted, at least below........ 2. Psallus Fieb.
See UNO LAM Alem wall NO Wig OLS eset cee teres ctece cee te sees onc a ua. gecthec este cceeesd vos-besceetecceacteee 8
8. Color uniformly black or nearly black above, legs pale ......................---- 9
— Coler pale, more or less marked with fuscous or black above ............-..-
a ee se ea ante Roa cecoecadsccesse 1. Reuteroscopus Kirk.

9. First antennal joint surpassing clypeus by more than half its length;
body polished and nude above ............-.-.---------- 6. Microphylellus Reut.

— First antennal joint scarcely surpassing apex of elypeus; body above
clothed) with pale pubescence 222-2... --o2------oen ee ae 4, Leptotylus n. gen.

10. Vertex distinctly carinate at base; rostrum short, but little surpassing
TPIS) EVO ENED CKO 21.2 ae 15. Myochroocoris Reut.
VCTLeS: without a basal carina; rostrum longer -.....-...0..--2c::.-c-sc.cce-oes 11
11. Second antennal joint linear, neither clavate nor flattened ................ 12
— Second antennal joint elavate or flattened and broader than first
5] CUT ame eee EE we A el CO ee eR Neigh hee eee ee 21

12. Clypeus well distinguished from the front by an incised suture ........ 13

Clypeus confused with the front or with suture very obscure; size
SOME CASH aT) ee a eee ee eee Tf

216

13.

University of California Publications. [| ENTomoLocy

Hind femora not at all saltatorial, not wider than the eye viewed from
the side, pale, dotted in longitudinal series; tibial bristles black and
inserted in black points; first joint of rostrum scarcely attaining

base (of Nea ste n5. os sec seeccccccsccetecctess iotes sees egeees eee 14
Hind femora saltatorial, much thickened or flattened, distinetly wider
than lateral: widthot: theveryie ps5. ee ee 15

. Head broad, its width three-fourths that of hind margin of pronotum;

viewed from the side produced below the eye for a distance as great
asi lenis the Of CVC) ee meee eres eer eee ART See 12. Bolteria Uhl.
Head narrower, hardly more than half as broad as basal width of the
pronotum; viewed from the side shorter, produced below the eye for
hardly more than half length of the eye; body densely clothed
Walt hevniGeRde cr dito sisi ain sseeese ees ene aes neers 11. Oligotylus n. gen.

. Femora black, tarsi annulated with white; body small, black ..................

ees eee eae _...---- -20, Strophopoda n. gen.
Femora pale, dotted with fuscous or black points ..........-.....22-.21:-0e---- 16

. Base of clypeus on a line with the insertion of antennae ................--------

seg NEE ot a tie cee en ee gone eee Oe 16. Atomoscelis Reut.
Base of clypeus above line connecting base of antennae ...............-.---------
SaGh oon Ee Sed RE RR AMS CSE cee nn LC 17. Huropiella Reut.

. Third joint of hind tarsi as long as first and second together; arolia

short, linear; attached to base of claws...............----- 18. Tuponia Reut.
Third joint of hind tarsi subequal to second, or shorter -............-....------ 18

. Head broad, about one-fifth narrower than hind margin of pronotum;
body black, hind femora black ....................-... 19. Chlamydatus Curt.
Headenarrowers) body. ameludin cw les steele yemeseseseeeee ane eee 19

. Head viewed from the side forming a squarish projection below the

eyes; legs and antennae pale, immaculate; tibial spines black; body
pale, clothed with short, thick black hair....23. Maurodactylus Reut.
Head forming a very short angular projection below the eyes............ 20

. Legs pale, femora immaculate; second antennal joint annulate with

TRG ig od ot ed a aoe oT eR EE 22, Cylloceps Uhl
Legs pale, hind femora dotted with black; first antennal joint annulate
With, lackey: ts-f. es eS Se oe 21. Campylomma Fieb.

. Second antennal joint terete, but little thickened at apex, longer than

Phin drande rout hist 0 ge thier essen ease eee eee 24, Rhinacloa Reut.
Second antennal joint of the male broadly flattened ..............2.--.2---------- 22

2. Body black, elytra pale brown, bifasciate with white .........-.....---..--::---+--<

EES GR Ai PE Ree SREY AAD By SOR ON ne? eh Eee es OY 14. Leucopoecila Reut.
Body entirely blue-black, or only coxae pale........ 13, Atractotomus Fieb.

so |

MOLOGY

(HEMIPTERA

yo ms

PRESS

INIVERSITY OF CALIFORNIA
=. "<== BERKEEEY, «

2
ox

yeep Ve

UNIVERSITY OF CALIFORNIA PUBLICATIONS

TECHNICAL BULLETINS
COLLEGE OF AGRICULTURE, AGRICULTURAL EXPERIMENT STATION

ENTOMOLOGY

Vol. 1, No. 4, pp. 217-227 March 3, 1916

NEW OR LITTLE KNOWN GENERA AND
SPECIES OF ORTHOTYLINI
(HEMIPTERA)

BY

EDWARD P. VAN DUZEE

Hyalochloria bella n. sp.

Larger than Orthotylus compsus Reut. from Jamaica; fulvo-
testaceous, elytra white, beautifully varied with clear green
blotches. Length 3.5mm. to tip of membrane.

Head small, vertical. Vertex broad and with base of front
much flattened and depressed between the prominent eyes; apex
of front convex; clypeus narrow and prominent. Eyes rather
large; viewed from the side ovate, reaching two-thirds of the way
to gula. Antennae inserted near to and a little above inner angle
of the eye; first joint not so long as head, second more than three
times longer. Rostrum long, reaching well over base of venter.
Elytra ample, costa expanded for the first one-fourth of its
length, beyond that subparallel. Bristles of hind tibiae pale and
very minute.

Color fulvo-testaceous. Pronotum in fully matured examples
yellowish, with about six green blotches on posterior lobe. Scutel-
lum sometimes varied with green. Elytra whitish hyaline, clavus,
corium, cuneus and membranal areoles quite regularly blotched
with irregular squarish pale green spots. In compsus the green
is more irregularly distributed and on the membrane is confined
to the small areole and the apical half of the larger one. In
bella there is a series of green points alone the costa which is
wanting in compsus, and, as mentioned above, the size is con-
siderably larger.

218 University of Califorma Publications. [ENtTomoLocy

Both of these species are intermediate between Hyalochloria
and Diaphnidia, but the short head and excavated vertex would
seem to place them nearer to the former, although the elytra are
less expanded, with the embolium much less complete.

Described from one male and four female examples taken by
me at Alpine, San Diego County, California, in June and July,
1913. It is worth noting here that Mrs. Slosson has taken
Hyalochloria caviceps Reut. at Biscayne Bay, Florida, thus bring-
ing the typical form of the genus within our territory.

Genus Labopidea Uhler

Form oblong in the male, ovate with abbreviated elytra in the
female. Color green, sometimes marked with black, and rather
densely clothed with pale appressed hairs. Head broad, vertical,
with eyes wider than anterior margin of pronotum. Vertex broad,
at least three times as wide as eyes, triangularly depressed, this
depression really formed by two oblique, impressed vittae, behind
which the surface is lower but leaving the hind margin convex
and almost carinated across the middle. Front broad and convex
in both diameters. Clypeus broad and prominent, its base almost
attaining the line of the eyes, sharply distinguished from the
front. Antennae rather long, inserted near lower angle of eyes;
first joint short, second at least three times longer, third and
fourth becoming thinner. Eyes small, prominent, surpassing by
nearly their whole width the pronotal angles; viewed from the
side short-oval, scarcely reaching to the middle of the side of the
head. Pronotum transverse, rather convex, its length about half
the basal width, sides strongly oblique, straight, anterior angles
well rounded; hind margin depressed, nearly covering basal lobe
of seutellum. Elytra well developed in the male; short, with
costa strongly arcuated in the female, in which the membrane is
frequently reduced to a mere margin. Membrane when developed
biareolate. Rostrum scarcely attaining the hind coxae. Arolia
free and converging. Wing-cell without a hamus.

Dr. Uhler describes the eyes as almost pedunculate, but that
appearance is caused by their rounded, bead-like form when
viewed from above, and perhaps in part by the rounded anterior
angles of the pronotum. The genus was founded upon a female
very like the female of sericatus, but somewhat marked with
black. The large thick green bodies clothed with a close hoary
pubescence, the thick vertical head and the short rounded elytra

Vou. 1] Van Duzee.—New Orthotylini. 219

of the females will distinguish this genus from Orthotylus and
its allies. Our species may be distinguished as follows:

PAWTIIG TINTS 2G) AG Kegs nee eee ts SR ce ge ewes cee ate eas ea cure tcnnapscees te cancenecesatencrceee il
ANTIIBETOTDES) TOG aces ace sccr oe See he ee Se 2
1. Legs pale, the body beneath mostly eae teeter Ss pets 2. simplex Uhl.
— Legs black, the body beneath mostly black __........... 1. nigripes Reut.
A, “Wniijomeeyl ssyowbaness leven leres [oye Me) a ere 3. sericata Uhl.
== IMlonenl Gyonbnes) Chowarene, LIE Wel See ee 4, atriseta n. sp.

1. Labopidea nigripes (Reut.)

A large, stout, bluish-green species with minute pale pubes-
cence, and antennae, legs and all beneath black. Male nearly
the form of Ilnacora malina UhL., elytra about parallel, with apex
of corium almost attaining tip of abdomen; female broad-ovate,
widest at base of membrane, tip of membrane scarcely if at all
surpassing abdomen. Length 5.5mm. to tip of membrane.

Head large; vertex broad, nearly four times as wide as the
small eyes, hind margin arcuate and somewhat thickened or
subearinate, transverse depression appearing foveate. Front
prominent, convex, searcely polished. Antennae rather long;
first joint scarcely as long as head; third two-thirds length of
second ; fourth short, yee one- Alersh length of third. eT eT
short, transverse ; callosities small, prominent and distant. Hairs
on antennae and legs pale and inconspicuous.

Dextral hook of male genitalia ovate at base, incurved apex
narrower but obtuse; sinistral hook broad, convex basally, pro-
duced distally, with an acute hook at either angle.

Color uniform bluish-green above, becoming yellowish on
vertex and front of pronotum; marked with black as follows:
eyes, antennae, a large round spot on front reaching to antennal
sockets, tylus, legs, sternum, membrane and abdomen, or at least
a broad, median vitta on tergum and venter. Callosities some-
times black.

The type was from Ormsby County, Nevada, and I took a
small series at Fallen Leaf Lake, California, in July, at an alti-
tude of 6300 feet. Dr. J. C. Bradley has sent me two females
that he took at Roger’s Pass in the Selkirk Mountains, British
Columbia, in July, 1908. These differ from the type form in
having a darker vestiture and in having a black mark covering
the callosities and most of the front of the pronotum and a black
dot on either side of the base of the vertex.

220 University of California Publications. | ENtTomoLoey

2. Labopidea simplex (Uhler)

Similar to preceding in size and form, but the elytral mem-
brane is more developed in all the females I have seen. Like
nigripes, 1t has a bluish-green color above, with pale pubescence
and black antennae, but here the legs and lower surface are
mostly pale. Length 5 to 5.5mm. to tip of membrane.

Basal depression of vertex large but sometimes obscure, with
hind margin quite distinctly carinate. Callosities small, promi-
nent.

Dextral genital hook of male enlarged into a nearly circular
hirsute plate; sinistral broad as in nigripes, but obtusely pro-
duced exteriorly and acutely hooked at its inner angle.

Color dull bluish-green, becoming yellowish on pronotum
anteriorly, on head, legs, and beneath. Frontal black spot
obsolete, or indicated by a broad brown are on either side; elypeus
black, at least at base. Antennae black, middle of second joint
more or less distinctly brown; ecallosities generally with black are
bounding them behind. Basal suture of scutellum_ blackish.
Elytra with costal border and cuneus paler, membrane quite
uniformly smoky. Tarsi fuscous or black. Pale pubescence
longer and denser than in nigripes.

The types were from Colorado. I took both sexes at Boulder
in that state, July, 1903, and the Academy of Natural Sciences
of Philadelphia possesses a series from Alamogordo, New Mexico,
taken in Apml and May, 1902. As nearly as I can judge from
Reuter’s description, his Hyoidea grisea seems to be a straight
synonym of this species.

3. Labopidea sericata Uhler

Size and form nearly of nigripes, male with very long elytra.
Color whitish-green, with disk of pronotum and elytra strongly
tinged with bluish-green ; upper surface densely clothed with long
white pubescence intermixed with deciduous silvery scale-like
hairs. Length, female 4.5mm.; male 7 mm., to tip of elytra.

Head proportionately longer and more pointed than in simplex.
Base of vertex distinctly carinate in male, scarcely so in female,
in which the transverse depression is much reduced. First
antennal joint short, scarcely as long as dorsal aspect of the head ;
second about as long as hind margin of pronotum; third two-
thirds length of second; fourth about half length of third. Pro-
notum short, in male strongly narrowed anteriorly; callosities
large, oval, prominent. Elytra in male very long, apex of corium
a little surpassing tip of abdomen; membrane long, whitish-
hyaline, scarcely infusecated; nervures green; in female shorter

Vou. 1] Van Duzee.—New Orthotylini. 221

than abdomen, with membrane reduced to a mere border to apex
of corium. Legs short, tibiae and some minute dots on apex of
femora green.

Dextral hook of male genitalia small, nearly circular at apex;
sinistral subtriangular, its inner angle produced in a short tooth,
the outer prolonged in an incurved subacute horn.

Color pale bluish-green, becoming yellowish on head, pronotum
before, and on lower surface. Antennae pale, scarcely darker
at apex. Sides of pectus and abdomen more or less greenish,
perhaps wholly green in life. Tibiae pale, with minute pale hairs
and a few longer brownish bristles. Elytral costa paler; mem-
brane whitish, with a very faint apical clouding, nervures green.

The types of this species were from Colorado, and I found it
common at Denver, Boulder, Ward, and Sunset, Colorado, in
July, 1903. The female has been determined for me as Labopidea
chloriza Uhler, but it wants the black markings mentioned in
Uhler’s description, and for the present I prefer to consider
chloriza as a species unknown to me.

Labopidea atriseta n. sp.

Closely allied to sericata Uhler, but distinguishable by having
infuseated antennae, and tibiae armed with conspicuous black
bristles set on black dots, while in sericata these bristles are
weaker and fewer and are not inserted in black points. Color as
in the preceding form. Length 5mm. to tip of membrane.

Head as in sericata, vertex feebly carinate at base; front con-
vex, clypeus prominent. Antennae a little longer than in allied
species; second joint about three times as long as basal; third,
three-fourths the length of second, the fourth short. Rostrum
a little shorter than in sericata, scarcely attaining apex of inter-
mediate coxae. Elytra with membrane developed in all my speci-
mens, apex of cuneus attaining tip of abdomen.

Color a soiled whitish, more or less tinged with dull green
below ; disk of pronotum and scutellum and the elytra dull bluish-
green, costal margin broadly pale; cuneus pale except its inner
angle. Membrane obviously infuscated, with a deeper shade out-
wardly beyond the areoles. Antennae infuscated, basal joint pale.
Hind femora distinctly dotted with greenish-brown exteriorly,
tibiae armed with stout black bristles much longer than the width
of the joint, each springing from a black point. Tarsi and tip
of rostrum black.

Described from four females taken by me at Alpine, Mussey’s,
and Sweetwater Valley, San Diego County, California, in April
and June.

222 University of California Publications. [ENToMoLoey

Genus Macrotyloides, n. gen.

Elongate, ovate, widest near apex of clavus. Head strongly
produced, subhorizontal, with eyes as wide as pronotum at trans-
verse incisure and a little more than half its basal width, its
length about equal to the median length of pronotum. Vertex
sometimes flattened or impressed before the base, leaving the
edge obtusely prominent, but scarcely carinate as in Orthotylus.
Vertex and front together when viewed from above scarcely
longer than wide, but slightly convex. Clypeus prominent,
arcuated before, its base well distinguished from the front and
on a line with the antennal sockets. Inner cheeks prominent, the
outer narrow. Eyes rather small, their inner margins feebly
divergent; viewed from the side oval, a little oblique, reaching
over half-way to gula; gula nearly horizontal. Antennae long,
inserted close to lower angle of eyes, first joint with two setae
within near their apex. Rostrum long, attaining middle of
venter. Pronotum trapezoidal, its anterior margin about half the
posterior; sides rectilinear, obviously carinated ; callosities large,
little elevated. Basal lobe of seutellum exposed. Prosternal
xyphus with margins carinate. Elytra rather short, with costa
quite strongly arcuated. Wing-cell without a hamus. Legs rather
long; tibiae with short, feeble bristles. Hind tarsi linear, joints
but slightly distinguished, basal shorter than the other two,
which are subequal; claws short, arolia rather large, free and
connivent, not equalling the claws. Vestiture not very dense, of
soft white hairs sometimes intermixed with stiffer fuscous ones.

Type of the genus Macrotylus vestitus Uhler.

This genus includes rather large green Capsids suggesting
Labopidea Uhbler, but with smaller eyes and a produced oblique
head as in Macrotylus; the tylus less prominent and the claws
large, with their arolia free and approaching at apex. When
the elytra are closed the body is conspicuously narrowed both
before and behind. Our species may be distinguished as follows:

Membrane about equally and lightly infuseated..................- vestitus Uhl.

Membrane with the apex abruptly fuscous ........................-- apicalis n. sp.

Macrotyloides vestitus (Uhler)

Long ovate, narrower before; pale grass-green, becoming
yellowish when dry; membrane very faintly and uniformly en-
fumed. Length 5 to 6mm.

Head nearly horizontal. Vertex hardly flattened, more than
twice as wide as the eyes in the male, its basal margin appearing
a little thickened in some individuals; front, feebly convex

VoL. 1] Van Duzee.—New Orthotylini. 223

obscurely transversely striate. Basal joint of antennae surpas-
sing tip of elypeus by nearly half its length; second slender,
about four times the length of first; third and fourth together
nearly as long as second, fourth about equal to first. Rostrum
reaching on the third ventral segment in the female, to the fifth
in the male, first joint surpassing base of head. Pronotum nearly
flat and horizontal, the surface before callosities scarcely de-
pressed, anterior edge with a slight median sinus; sides straight,
subearinate. Scutellum equilateral. Elytra elliptical, nearly flat,
cuneus scarcely longer than broad; abdomen reaching to middle
of cuneus in the male, nearly to apex of membrane in the female.
Membrane surpassing cuneus by about one-third its length.

Male genital pieces large; dextral hook very large and curved
in conformity with the margin of anal opening, in form ligulate,
its subacute tip reaching over onto apex of tergum; sinistral
almost equally long and similar in form, produced either way
and lying parallel with and just below the dextral, its inner end
obtuse, the outer acute. Female pygofers short, beginning behind
middle of venter, hind edge of fourth segment almost straight
across.

Color pale green, becoming yellowish on head, pronotum, legs,
antennae, and lower surface in dried specimens. Antennae in-
fuseated toward apex. Tibial bristles fuscous; apex of tarsi
black. Vestiture composed of a fine soft pubescence and short
stiff fuscous hairs intermixed. Membrane nearly hyaline,
nervures green.

Redeseribed from five male and three female examples taken
in San Diego County, California, at Alpine, Mussey’s, and Fos-
ter’s, from April to June.

Macrotyloides apicalis n. sp.

Closely allied to the preceding; a little shorter behind, with
apex of membrane deep smoky brown. Length 5.5 to 6mm. to
tip of the membrane.

This species is very close to the preceding in most of its
characters, but the elytra are shorter, making the insect appear
broader; tibial bristles shorter and more slender, and the basal
joint of the rostrum scarcely passes the base of the head. Oviduct
of the female longer, beginning distinctly before middle of venter,
with fourth ventral segment strongly oblique, not practically
transverse as in vestitus. Color light green, becoming yellowish
on head, pronotum, and below, as in vestitus. Antennae infus-
cated at apex. Membrane whitish-hyaline with its apex beyond
the tip of cuneus abruptly blackish-fuscous.

224 University of California Publications. [ENtTomMoLocy

Described from six females from southern California. I took
it at Alpine, San Diego County, in June, and Mr. Fordyce
Grinnell has sent me specimens taken at Pasadena, June, Los
Angeles, September, and from the Santa Rosa Mountains.

Pseudopsallus n. gen.

Aspect of Plagiognathus nearly, but having cell of wing with-
out a hamus and aroha free and connivent. Body clothed with
long black hairs and short, appressed, silvery scale-like hairs or
tomentum, which soon wear off, leaving the insect smooth.

Head broad and short, vertical. Vertex with an oblique
impression either side, leaving the base prominent but hardly
earinate. Front moderately convex, perpendicular or nearly so.
Clypeus broad, somewhat prominent, but little compressed ; base
well distinguished from the front; apex of head blunt, produced
below the eye for less than length of the eye. Eves large, promi-
nent, viewed from the side ovate, reaching below middle of side
of head. Antennae inserted near lower angle of eye, short, stout ;
basal joint not longer than head viewed from above, second
scarcely thinner at base than first. Pronotum transverse, smooth,
anterior margin about half the length of the posterior, sides
ecarinate, nearly straight, anterior angles broadly rounded. Cal-
losities large, little elevated. Basal lobe of sceutellum covered.
Elytra broad, costal margin considerably arcuated in the female,
nearly parallel in the male. Rostrum reaching onto base of
venter; oviduct of the female long, beginning before middle of
venter. Wing-cell without a hamus. Legs thick, irregularly
dotted, tibial spies stout, black. Tarsal claws small but longer
than in Macrotylus; arolia free and connivent.

Type of the genus Macrotylus angularis Uhler.

The type species has much the aspect of a large Plagiognathus,
but the absence of a hamus in the wing-cell and the free connivent
aroha places it in the Orthotylin, where there is no established
genus that will receive it.

Pseudopsallus angularis (Uhler)

This species and the next are broad forms, and have httle in
common with the other genera of the Orthotylin, but they must
be placed in that tribe. Angularis is a slaty-brown species with
the claval suture, costa, the two principal nervures of the corium,
and the membranal nervures whitish, sometimes tinged with
yellow on the costa. The region of the callosities is black varied
with white, and the head is white, with the oblique impressions

Vou. 1] Van Duzee—New Orthotylini. 225
of the vertex, a large are on either side of the front, some marks
on. the base and sides of the clypeus and the lorae black. The
antennae are black, the second joint as long as the base of the
pronotum and one-fourth longer than the third and fourth taken
together. The legs are white, with the tibiae dotted with black.
The white markings, especially on the head and pronotum may
become more or less orange.

Dextral hook of the male genitalia apparently a flattened
truncated seale scarcely longer than broad; sinistral very large,
expanded on the right to base of dextral hook and exteriorly
narrowed and curved upward and inward to superior aspect of
anal opening.

I took three examples of this insect from the white sage grow-
ing on the mesa at East San Diego, California, during April and
May, 1913.

Pseudopsallus verticalis (Uhler)

This is a smaller, pale brown or testaceous form, with the
head and anterior lobe of the pronotum fuscous, the base of the
vertex marked with pale, and the legs pale dotted with fuscous.
The types were from California. I possess one female taken by
Mr. Fordyce Grinnell at Pasadena, May 25, 1909, and another
taken in Colorado was sent to me by Professor Baker labelled
Macrotylus verticalis Uhl. MS. It probably was by an oversight
that Dr. Uhler failed to mention the Colorado locality in con-
nection with his description.

Argyrocoris femoratus n. sp.

Smailer and darker-colored than scurrilis ; slender ; testaceous-
grey, more or less infuscated, cuneus dotted with red. Length
4mm. to tip of membrane.

Head strongly oblique; vertex convex, ecarinate ; front moder-
ately convex. Eyes large oval, viewed from the side oblique and
attaining gula, coarsely granulated. Antennae long; basal joint
surpassing head by one-half its length; second over three times
the length of first; third and fourth together about equal to
second. Pronotum transverse, its length about three-fifths its
basal width, anterior margin about two-thirds of posterior; hind
edge depressed, covering basal lobe of scutellum. Rostrum long,
reaching middle of venter. Elytra long and narrow; tip of
abdomen not attaining apex of corium, cuneus slender, its length
nearly three times its basal width in the male. In the female
the abdomen reaches to about the middle of the cuneus, which is

226 University of California Publications. [ENTOMOLOGY

broader and shorter than in the male. Hind femora long and
much flattened.

Male genitalia small; dextral hook about twice longer than its
basal width and tapering to a point; sinistral oval and lying
along the sinistral notch.

Color greyish-testaceous; callosities narrowly black; disk of
pronotum with two broad fuscous rays more or less distinct.
Seutellum infuseated or almost black. Elytra dotted with san-
guinous on cuneus and sometimes more sparsely on corium and
clavus. Membrane deeply infuscated, nervures pale dotted with
red. Wings somewhat infuseated, with blackish nervures. Body
beneath infuseated. Antennae and legs pale, femora dusky, hind
pair more or less dotted with red points; tarsi black at apex.
Antennae slightly infuseated toward their apex. Upper surface
clothed with minute silvery pubescence, obscurely arranged in
lines which are more distinct on the head, where they form three
longitudinal vittae.

Described from numerous examples beaten from chaparral in
San Diego County, California, at Coronado, National City,
Alpine, ete., in May and June.

At first sight this species seems very different from scurrilis,
but a closer inspection shows that it possesses similar generic
characters and there can be no doubt that it should be placed in
the same genus.

Parthenicus picicollis n. sp.

Allied to psalliodes, but with pronotum, scutellum and com-
missure piceous or almost black; beneath more or less infuscated,
above pale, more or less closely dotted with sanguinous points,
elytra appearing dark red; membrane black. Length 3mm. to
tip of membrane.

Head vertical; vertex and front conjointly flattened above, a
very little convex below, base of vertex ecarinate. Clypeus
prominent, but somewhat less so than in Macrotylus, its base
well distinguished and distinctly above line of antennae. Eyes
large, prominent, projecting for nearly their whole width beyond
the pronotal angles; viewed from the side broad-oval, vertical,
reaching almost to gula, face but ttle prominent before the eyes,
almost half of this being projection of clypeus. Antennae rather
short, first joint but little surpassing clypeus; second longer than
basal margin of pronotum. Pronotum moderately convex, eal-
losities small and little elevated, distinguished behind by a deeply
incised line. Basal lobe of secutellum covered. Costal margin of
elytra feebly areuated. Cuneus about as long as wide at base in

VoL. 1] Van Duzee.—New Orthotylini. 227

the female, longer in the male. Hind femora about one-third as
wide as long in the female, hardly one-fourth in the male.

Dextral hook of the male genitalia slender toward its apex,
bent at a right angle dorsally; sinistral somewhat crescentic, its
inner (dextral) angle produced.

Ground color pale yellowish-testaceous. Head, legs, and
elytra dotted with dark sanguinous, the latter closely, almost con-
fluently so, appearing as if washed with red. Antennae pale,
basal joint red, apex slightly infuseated. Pronotum and seutel-
lum deep piceous or almost black; elytral commissure evenly and
indefinitely clouded with fuscous, this clouding scarcely extending
onto inner angle of corium. Membrane blackish, with a pale line
around apex of cuneus, nervures red. Face sometimes dusky,
elypeus and cheeks clear red. Lower surface and base of hind
femora more or less infuseated; tibial bristles fuscous, springing
from sanguinous dots. Upper surface clothed with a fine pale
pubescence.

Described from fifteen examples beaten from Adenostoma in
San Diego County, California, from July to October. Although
closely allied to psalliodes, this form seems sufficiently distinet by
its dark pronotum, scutellum and lower surface, and the con-
colorous inner angle of the corium

Transmitted January 3, 1976.

2 4 is ea a ess ee

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EGE OF AGRICULTURE, AGRICULTURAL EXP
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HEMIPTERA

A

ate ~

_ NEAR LAKE TAH

ESN
ee } og ag

Soe)

EDWARD P. VAN DUZEE

_ UNIVERSITY OF CALIFORNIA
BERKELEY

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Vol. 1, No. 5, pp. 229-249 May 23, 1916

NOTES ON SOME HEMIPTERA TAKEN NEAR
LAKE TAHOE, CALIFORNIA

BY
EDWARD P. VAN DUZEE

In the summer of 1915 it was my privilege to spend about
five weeks in the Sierra Nevada with the entomological field class
of the University of California Summer School at Fallen Leaf
Lake, Eldorado County, California. We reached Fallen Leaf
Lodge, our headquarters, on June 21 and left on July 31, a
period which covered the season of greatest abundance of insects
in certain of the orders, notably the Hymenoptera, Diptera and
Lepidoptera, and the Coleoptera were perhaps at their best before
we left; the Hemiptera and Orthoptera were, however, just
coming into season, so the material obtained in these orders was
but fragmentary. Circumstances were such that I was able to
devote much of my time to the collection of insects for the
University Museum, and Professor Woodworth added a number
of interesting forms during two brief visits to the Lodge, while
other members of the party turned over to me some good things
taken by them. Altogether over six thousand mounted insects
were brought home, a study of which will undoubtedly make
valuable additions to our knowledge of the Sierran insect fauna.

The present paper deals with the Hemiptera taken, but.
owing to the earliness of the season, must be but an imperfect
representation of the hemipterous fauna of that portion of the
Sierra. Of the one hundred and forty species enumerated, per-
haps one-third are known to inhabit the coast region of Cali-
fornia and about one-fourth are common to the eastern and

230 University of California Publications. [ENTOMOLOGY

northern portions of the United States and Canada. A number
of forms characteristic of the Rocky Mountain region were taken.
as well as several found in the hilly back-country of San Diego
County, California. However, before any really useful studies
on the distribution of the Hemiptera of this portion of the Sierra
Nevada can be made, it will be necessary to do systematic collect-
ing there during August and early September. It is interesting
to note that twenty-one of the species taken, or approximately
one-seventh, are new to science, which shows how little is now
known of the Hemiptera of the Lake Tahoe region.

The following notes on localities will indicate the conditions
under which most of the material was taken.

Lower end of Fallen Leaf Lake.—On the west side of the lake
from Cathedral Park and the sawmill to the outlet and some dis-
tance north was an interesting locality with low, rich woods and
open, marshy fields. Mosquitoes were a serious hindrance here
but the Hemiptera were more numerous than elsewhere, perhaps
because the season was earler at this altitude, 6300 feet.

Lateral Moraine.—A sharp ridge along the eastern side of the
upper end of Fallen Leaf Lake, with an elevation of about 800
feet above the lake. This ridge was well covered with chaparral
and until about the tenth of July made an excellent collecting
ground for the Hymenoptera, Diptera and Lepidoptera. After
that date it became too dry for good collecting.

Angora Lakes.—Situated under the eastern escarpment of the
Angora Ridge at an altitude of about 7500 feet. Collecting here
was poor.

Glen Alpine Springs and Creek are situated in a valley run-
ning west from the upper end of Fallen Leaf Lake. Below Glen
Alpine Springs the valley is well wooded and afforded excellent
collecting places. Toward the last of July the northern side of
the valley formed by the southern slope of Mount Tallac proved
to be one of our best collecting grounds.

Cathedral Lake, in a valley on the eastern slope of Mount
Tallac at an altitude of about 7500 feet, was a moderately: pro-
ductive place, especially lower down near Floating Island Lake.

Mount Tallac.—The sloping alpine meadow on the western
aspect of the mountain was a wonderfully interesting place from
about the twentieth of July. The altitude here varied from about
8000 to over 9000 feet.

Vou. 1] Van Duzee.—Lake Tahoe Hemiptera 231

Angora Ridge.—The western slope of this ridge was an alpine
meadow of equal altitude and hardly less interesting than that
on Mount Tallae.

Half Moon Lake under Dick’s Peak at an altitude of about
8000 feet proved to be an interesting place in late July. Several
eastern forms not found elsewhere were taken here.

Grass Lake in the valley above Glen Alpine Springs at an
altitude of about 7500 feet did not yield much of interest, but
might have been better if visited later in the season.

HETEROPTERA

Thyreocoris anthracinus Uhler. July. Taken in numbers
from a low plant growing in grassy places along the roadside
from Tallae to the sawmill at the lower end of Fallen Leaf Lake.

Homoemus bijugis Uhler. Taken with the preceding.

Eurygaster alternatus Say. Abundant with the foregoing
species,

Trichopepla atricornis Stal. Found occasionally on rank
weeds about the lower end of Fallen Leaf Lake, July 17.

Thyanta custator Fabr. Not uncommon on trees and bushes
everywhere below 8000 feet.

Banasa sordida Uhler. <A single specimen swept from weeds
at Half Moon Lake in July.

Tollius curtulus Stal. Low ground near Tallae, July.

Alydus pluto Uhler. One example taken on weeds growing on
a dry sandy spot near Tallae in July.

Harmostes reflerulus Say. Not uncommon with the preceding.

Corizus scutatus Stal. Common on the low lands between
Tallae and Fallen Leaf Lake in July.

Corizus indentatus Hambl. Taken with the preceding.

Corizus hyalinus Fabr. Generally distributed, but not com-
mon, up to 8500 feet.

Corizus crassicormis Linn. Half Moon Lake, July 23.

Aradus debilis Uhler. This, our largest western aradid, is
paler than most of our species, has the abdomen of the female
much produced and attenuated at apex and the third antennal
joint white with its extreme base and the large apical joint black.
Mr. Ralph Hopping took a series on fungi under the bark of a
dead pine tree near the margin of Fallen Leaf Lake, July 2, Dr.

232 University of California Publications. [ENTomMoLocy

Van Dyke found one in a similar situation, and I took a male
near Glen Alpine Springs, June 30.

Aradus hubbardi Heid. A fine series of this species was taken
from fungus growing on a fallen pine tree along the trail to
Angora Lakes at an altitude of about 7000 feet, July 11. In all
the specimens taken the tip of the second antennal joint is scarcely
paler.

Aradus persimilis, new species

Very close to hubbardi, differing principally in having the seutellum much
broader at apex, the expanded portion of the costa longer and the antennae
a little thinner, with their second segment perhaps a little longer and
the third without a pale apex. Length, male 6.5 mm., female 7.5 mm,

Head as in hubbardi; anterior process thick, cylindrical, compressed
toward the apex, reaching nearly to basal third of second antennal joint.
Spine at base of antennae almost attaining the tip of first joint and armed
exteriorly with a very short tooth. Occiput with a tubercle before the
eye and another near its hind angle. Antennae about as in hubbardi,
perhaps a shade thinner; first segment about one-half longer than wide,
second about as long as the head, a little thickened at apex; third hardly
one-half the length of second; fourth still shorter, narrowed to its base
with a short conical tip. Pronotum about as in hubbardi, slightly longer
and more broadly expanded about the humeri; sides irregularly dentate;
four discal carinae nearly parallel, exterior percurrent, not becoming
obsolete before as in the allied species. Abdomen as in hubbardi; genital
segment of the female a little shorter and less expanded; genital lobes of
the male shorter and more transverse. Rostrum nearly attaining hind
margin of mesosternum.

Color fuscous-brown, becoming more ferruginous on the cephalic pro-
cess, basal joints of antennae, pronotal carinae, principal elytral nervures,
extreme tip of scutellum, and in places on abdomen. Antennae becoming
black on apical two segments, conical tip of fourth sericeous pubescent.
Expanded basal portion of costa and broad humeral areas and marginal
serrations whitish-testaceous; elytral reticulations sometimes of the same
pale color. Legs and beneath usually paler. Abdomen becoming more
castaneous toward its margin, where it is often minutely pointed with
green, the hind edge of the connexival segments pale; membranal veins
distinctly pale.

Described from three male and two female specimens taken
near Glen Alpine Creek, June 25 to July 3. While very near
hubbardi this form seems to have good specific characters and is
at least a valid subspecies. I possess one female, taken by Dr.
J.C. Bradley in the Santa Cruz Mountains, California, in May,
1907, that differs from the types only in having the humeral
expansion of the pronotum concolorous.

Vou. 1] Van Duzee.—Lake Tahoe Hemiptera 233

Aradus borealis Heid. Professor Woodworth took one ex-
ample of this species on the eastern slope of Mount Tallac along
the trail from Cathedral Lake to Floating Island Lake, on July 6,
at an altitude of 7000 feet.

Aradus behrensi Bergr. A few examples were taken from
trees near the upper end of Fallen Leaf Lake, June 25. The next
day I took near the lower end of the lake two specimens of what
I believed to be brachypterous or imperfectly developed speci-
mens of the same species. In these the elytra reach only to the
base of the fifth abdominal segment.

Aradus insolitus, new species

A small, black species marked with pale granules; humeral angles sub-
acute, latero-anterior margins a little concavely arcuated, second antennal
joint narrowly white at apex. Length 4mm.

Form narrow ovate, about as in tuberculifer Kirby. Head with a short
oblique impressed area on either side interior to the eye. Antenniferous
tubercles with a large acute spine which surpasses the middle of first
antennal segment. Antennae regularly but moderately thickened to apex
of third joint; basal scarcely longer than broad, second about as long as
the distance between the eyes, uniformly thickened toward its apex, white
apical portion about as long as the thickness of the joint at base; third
and fourth about equal in length, the latter fusiform. Pronotum rather
small; humeri prominent, subacute, the margins behind them feebly
rounded, the latero-anterior margin a little concavely arcuated with the
anterior angles prominent and subacute; entire lateral margins quite
regularly and minutely crenulate; disk a little depressed across the
middle, marked posteriorly with four parallel carinae which become con-
fused anteriorly. Scutellum narrow, about as long as pronotum, obtuse
at apex, margins but little elevated. Elytra reaching nearly to tip of
abdomen, but little narrowed apically; costa regularly but moderately
dilated at base. Nervures of corium and membrane prominent. Rostrum
reaching but little beyond the base of the head, encroaching upon base of
the prosternum about the width of the sternal suleus at its middle; this
suleus expanded at base and apex. Genital lobes of the male about as in
similis, the oblique apical margins a little more rounded.

Color almost black; narrow apex of second antennal segment, tips of
the humeri, broad but indefinite apex of scutellum, knees, tips of the
tibiae and apical angles of the abdominal segments pale or whitish.
Whole surface irregularly dotted with minute greenish-white granules,
most numerous on the head, disk of seutellum and elevated areas of
pronotum and elytra; granulations of connexivum becoming a clear sea-
green in places.

234 University of California Publications. [ENtomoLoey

Described from two males taken about the upper end of Fallen
Leaf Lake, July 12. This species is very distinct from any other
known to me by its subacute humeral angles and the concave
latero-anterior margins of the pronotum. The short rostrum
allies it with Quilnus and in some respects it is intermediate
between that subgenus and the more typical aradids.

Aradus fallent Stal. One specimen taken from a fallen pine
tree near Cathedral Lake on the eastern slopes of Mount Tallac,
July 5.

Mezira moesta Stal. A single individual of this common form
was taken near the upper end of Fallen Leaf Lake on July 2.

Lygaeus reclivatus Say. Found in numbers on a patch of
milkweed near the sawmill at the lower end of Fallen Leaf Lake,
July 265.

Lygaeus truculentus Stal. A few taken on flowers near the
lower end of Fallen Leaf Lake in July.

Lygaeus bicrucis Say. Taken at Grass Lake and near Fallen
Leaf Lake in late June and early July.

Nysius californicus Stal. Common at lower levels but also
found at Half Moon Lake and up to 8500 feet on Mount Tallae.

Nysius ericae Sehill. Half Moon Lake, July 23.

Nysius ericae minutus Uhler. A few taken near Tallae in
July.

Tschnorrhynchus resedae Panz. One specimen was _ beaten
from chaparral along the trail on the west shore of Fallen Leaf
Lake, July 5.

Geocoris pallens decoratus Uhler. One example taken on the
low lands near Tallae on July 25.

Ligyrocoris diffusus Uhler. With the preceding.

Sphragisticus nebulosus Fall. Swept from grass near the
lower end of Fallen Leaf Lake, July 17.

Rhyparochromus angustatus Van D. This form may be
roughly distinguished from sodalicus Uhl. by its having the third
antennal joint entirely black, the lateral edges of the pronotum
concolorous and the clavus ferruginous at base. It is a long
narrow form and northern in its distribution. Here it was found
near Tallae in July.

Scolopostethus thomsoni Reut. Three examples were taken at
Tallae and about the upper end of Fallen Leaf Lake in July.

Ploiariodes sp. One example taken at the Lodge in July.

ese

Vou. 1] Van Duzee.—Lake Tahoe Hemiptera 235

Nabis ferus Linn. Found everywhere. Here the species has
every appearance of being a native insect and possibly it is in-
digenous to all the northern portions of this continent.

Anthocoris bakeri Popp. Taken near the Lodge, June 21.
Poppius’ paper describing this species never reached me and I
was unaware of its existence until after the publication of my
species ornatus, which is undoubtedly identical with bakeri.

Anthocoris antevolens White. Taken on the low lands about
Fallen Leaf Lake in July.

Anthocoris melanocerus Reut. Two examples were beaten
from the chaparral along the western shore of Fallen Leaf Lake,
July 5.

Saldula interstitialis Say. One taken at Tallae July 25.

Saldula sp. Taken on the low lands about the lake and up to
8500 feet on the western slope of Mount Tallac.

Gerris orba Stal. Found on Fallen Leaf Lake, Cathedral
Lake, Angora Lakes and elsewhere, in July.

Gerris gillettei L. & S. One example taken at the sawmill at
the lower end of Fallen Leaf Lake, July 25.

Stenodema vicina Prov. Taken on grassy openings along Glen
Alpine Creek, at Half Moon Lake, and on the western slopes of
Angora Peaks, in July. No males were found but the females
were not uncommon.

Platytylellus intercidenda Dist. Occasional on rank vegeta-
tion about the lower levels after July 16.

Phytocoris eximius Reut. Glen Alpine Springs, July 30.
This species and the next were only just reaching maturity when
I left, the last of July.

Phytocoris inops Uhler. With the preceding. This is a
smaller species having the membrane varied with whitish and
fuscous in coarser blotches, the antennae longer and more slender,
with the basal joint more obviously banded and the pale incisures
broader.

Ectopiocerus anthracinus Uhler. Oceasional in July at the
lower levels.

Poeciloscytus uhleri Van D. Taken at the level of Fallen
Leaf Lake and up to 8500 on the west slope of Angora Peak,
toward the last of July.

Poeciloscytus venaticus Uhler. Not uncommon on rank weeds
about the lower end of Fallen Leaf Lake in July.

236 University of California Publications. [ENtoMoLoey

Dichrooscytus speciosus, new species

Allied to suspectus but more polished. Black; disk of head, pronotum
and elytra deep rich wine-red; scutellum, base of cuneus and legs white;
antennae soiled white. Length nearly 6 mm.

Head broader and shorter than in suspectus, vertex strongly impressed
before the basal margin and more broadly on either side; middle of
vertex quite strongly swollen and polished, without the transverse striae
seen in the allied species; basal carinae prominent across the whole width.
Pronotum narrower anteriorly, meeting the vertex near the inner angle
of the eye; surface more strongly punctured, callosities smaller and
highly polished; median line obscure but evident. Secutellum prominent,
narrower than in suspectus, polished, a little transversely shagreened
toward the base. Elytra polished, very obscurely punctured and
shagreened, almost parallel, costa straight basally.

Color black, highly polished; base of vertex obscurely reddish. Pro-
notum deep wine-red with anterior and posterior margins black. Secutellum
ivory-white, becoming black on anterior lobe beneath base of pronotum.
Elytra dark wine-red; broad margins of clavus about scutellum and the
apex of corium black; euneus black, the basal half ivory-white omitting the
slender costal margin. Membrane deep fuscous, iridescent, the nervures
concolorous. Legs including coxae white, coxae and trochanters some-
times tinged with greenish; apex of the tibiae and the tarsi slightly
infuscated, last tarsal joint becoming black at apex. Antennae soiled
white, a little more infuscated toward the apex. Tergum and apex of
abdomen more or less sanguinous.

Described from numerous examples beaten from juniper trees
during July. mostly on the south slope of Mount Tallac. The
first mature examples were taken near the Lodge on July 5 but
the adults did not become abundant until after the middle of the
month. This is one of the most beautifully colored capsids known
tome. The immature have paler colors but the pattern is distinct
and characteristic.

Dichrooscytus suspectus Reut. Common on juniper trees
during July. These individuals are more uniformly colored with
rufous than those from Colorado determined for me by Dr.
Reuter, but they can hardly be distinct.

Dichrooscytus irroratus Van D. Common on juniper and
cedar trees everywhere between 6000 and 8000 feet. In most of
these examples the elytra are of an almost uniform pinkish color
with a deeper tint on the cuneus, and marked with a fuscous vitta
on the apex of the corium; an oblique vitta on their middle and
the apex of the membranal areoles are also fuscous.

Vo. 1] Van Duzee.—Lake Tahoe Hemiptera 237

Dichrooscytus elegans Uhler. Taken with the preceding forms
but less abundantly and a little earlier.

Lygidea rebucula obscura Reut. Three examples were taken
about the Lodge July 20 and 21. These differ in no respect from
specimens taken in the state of New York.

Platylygus luridus Reut. Not uncommon on Jeffrey pines
from July 8. These are larger than eastern specimens but do not
otherwise differ.

Lygus pratensis Linn. The pale form with yellowish scutel-
lum which is characteristic of the Californian fauna was common
at all the lower levels and up to 8000 feet. Three examples of
the dark eastern form were taken on July 23 at Half Moon Lake.

Lygus plagiatus Uhler. This form was common above 7500
feet but rarely was taken at lower levels.

Lygus campestris Linn. Common about Fallen Leaf Lake and
up to 7800 feet at Half Moon Lake.

Xenoborus canadensis Van D.? One example which may rep-
resent a distinct species differs in being smaller and in having
the basal joint of the antennae and the tylus concolorous.

Camptobrochis validus Reut. Four examples were taken
about Fallen Leaf Lake and Grass Lake from June 23 to July 5.

Camptobrochis fulvescens Reut. Beaten from pine on the
south slope of Mount Tallae on July 30. These specimens are
glabrous and have much the aspect of nitens but the male genital
characters are those of fulvescens and I prefer to so place them
for the present.

Deraeocoris ingens, new species

Large, deep black, polished; second antennal joint greatly thickened.
Length 8 mm.

Aspect of Capsus ater, but much larger and more elongated, with
antennae more clavate. Head as in Camptobrochis, prominent; hind
margin of vertex and eyes nearly rectilinear. Vertex almost flat, basal
margin subearinate behind a shallow depression. Antennae stout, inserted
before the eyes at a distance about equal to the thickness of first segment;
this segment as long as the median width of vertex; second (3.5 mm.)
longer than basal width of pronotum, at base more slender than first but
regularly thickened to near its apex, where it is as thick as the antero-
posterior diameter of the eye, its surface closely pubescent and armed
with a few longer hairs; third and fourth segments short and slender
but not setaceous, together equal to length of the anterior tarsi, third
searcely longer than fourth. Pronotum shaped much as in Camptobrochis

238 University of California Publications. [ENtToMoLoey

nigrita, much narrowed anteriorly, where it is a little wider than the
base of the vertex; surface convex in both diameters, strongly punctured;
callosities large, highly polished, impunctate, scarcely elevated and poorly
defined; sides nearly straight or slightly coneave. Scutellum more finely
punetured. Elytra coarsely punctured, long, narrowed from the middle.
Membrane long and narrow, nervure dividing the areoles obsolete or
nearly so. Prosternal xyphus depressed with a carinate margin. Basal
joint of hind tarsi stouter, as long as second and third together. Arolia
as in Camptobrochis, wanting or very minute and connate with the claws,
between which are two slender parallel setae. Male uncus formed about
as in Camptobrochis nitens with a sharp eurved hook at distal end which
almost attains the line of the proximal extremity.

Color a uniform deep coal-black, brilliant; membrane a dead blackish-
fuscous, scarcely lighter next the point of the cuneus; base of vertex and
knees touched with fulvous. Second antennal joint nearly to its apex
and the tarsi piceous, verging toward castaneous. Osteolar margin white.

Described from nine examples representing both sexes, all
beaten from Jeffrey pines along the south slope of Mount Tallae
above Glen Alpine Creek during late July. This is one of our
largest capsids, and it seems strange that it should have remained
unknown for so long unless it be restricted to the Jeffrey pine,
which has a range in the Sierras and northwardly where little
work on Hemiptera has been done. Dr. Poppius has recently
sunk Camptobrochis as a synonym of Deraeocoris, but I think
it better to restrict the latter genus to those species having a
clavate second antennal joint.

Deraeocoris fraternus, new species

This form seems to differ from ingens principally in being smaller
(6mm.) and in having the basal one-half of the second antennal joint
and the legs pale or even bright rufus, with the apex of the tibiae darker
and the tarsi black. Generally there is a pale annulus beyond the middle
of the tibiae and the margins of the pleural pieces are whitish, usually
broadly so.

Described from ten examples representing both sexes taken
with the preceding. I would consider this but a color variety of
ingens were it not for its much smaller size and the constaney
of its characters.

Largidea grossa, new species

Form nearly that of marginata, but larger; dull rufus-brown, whole
upper surface closely and coarsely punctured. Length to tip of membrane
6 mm.

Vou. 1] Van Duzee.—Lake Tahoe Hemiptera 239

Head impunetate, polished, clothed with sparse, short appressed pale
pubescence; viewed from above short, transverse, gently swollen before;
width about twice its greatest length; when viewed from before five-
angled, portion above the antennae almost quadrangular with sides
straight and parallel, apex obtuse; base of elypeus but feebly dis-
tinguished from the front. Rostrum attaining the base of the inter-
mediate coxae. Antennae as in marginata, inserted near lower angle of
eye; basal joint searcely longer than the eye; second joint is as long as
vertex and pronotum taken together, flattened horizontally and broadly
lanceolate, its width equal to length of first joint, its lower surface
feebly convex, the upper deeply suleate nearly to the apex; third and
fourth thin, fusiform, each subequal to the first in length. Pronotum
broader and less narrowed anteriorly than in marginata, sides straight,
distinctly earinate, attaining the middle of the eyes; posterior lobe
deeply closely punctured; anterior lobe nearly smooth, gray-pubescent,
transverse impressed line deep, black, bent back in a barb at either
side of the median line. Scutellum strongly convex, closely but not
deeply punctured. Elytra closely punctured, the costa bent beyond the
middle. Cuneus long, but moderately depressed. Membrane long, sur-
passing the abdomen by one-half its length. Basal joint of the tarsi
broad and flattened, scooped out below, wider than the apex of the tibiae
and as long as slender third joint; second short, slender like third, deeply
inserted in upper surface of broad first joint. Oviduct beginning before
middle of venter.

Color dull rufus-brown inclining to wine-red but not as deep a red as
in the allied marginata. Whole surface closely but not conspicuously
pale-pubescent; disk of head and much of lower surface blackish; costa
paler; membrane black, nervures black at base, paler beyond. Antennae
reddish. Tarsi and venter blackish.

Described from a single female example taken on Jeffrey pine
on the south slope of Mount Tallae near Glen Alpine Springs,
July 30, at an altitude of about 7000 feet. It is possible that
the expanded basal joint of the tarsi and the fact that the second
antennal joint is scooped out above instead of below would entitle
this form to generie distinction, but it does not seem advisable
to separate it until the male has been examined. In the type
specimen the thickened apex of the second antennal joint is
somewhat bent, making the antennae appear to be five-jointed
when viewed from the side.

Dacerla inflata Uhler. This singular looking ant-mimie was
quite abundant after the middle of July at most all locations but
especially along the southern slope of Mount Tallaec. A few
were seen above 8000 feet on Angora Ridge. It was most abun-
dant on the lodge-pole pine, but it occurred on other trees and

240 University of California Publications. [ENToMoLoGy

a few were swept from weeds near the lake. The males scarcely
differ from the females and in both sexes some individuals are
paler or reddish. The pronotum and parts of the elytra that
are gray in the black form are ferruginous in the pale, but most
of the material taken answers in every respect to the excellent
deseription given by Dr. Uhler.

Coquillettia insignis Uhler. Three males of this pretty species
were swept from weeds near the sawmill on Fallen Leaf Lake,
July 25, and one was taken next day from the western slope of
Angora Peak above 8000 feet.

Dicyphus californicus Stal. A few were taken about Fallen
Leaf Lake late in July.

Dicyphus agilis Uhler. With the preceding but more abun-
dant. I distinguish this from californicus by its larger size, paler
colors, the almost entire want of red on the elytra, the more
tumid and exposed base of the scutellum and especially by the
shorter basal joint of the antennae.

Dicyphus vestitus Uhler. One example was taken July 17
near the lower end of Fallen Leaf Lake. ‘

Dicyphus crudus, new species

Form and size of agilis; pale greenish-white. Length 5mm., width
across the elytra little more than 1 mm.

Vertex less swollen than in agilis, eyes a little more prominent, with
antennae longer and pronotal collar more elongated. Elytra very long,
parallel, membrane surpassing abdomen by two-thirds its Jength.

Color pale greenish-white, deeper green on anterior lobe of pronotum,
pleural pieces and base of scutellum. Eyes and tips of the tarsi black.
Neck with a fuseous cloud behind each eye. Elytra almost pellucid,
sparsely clothed with short apressed white hairs. Membrane with a faint
longitudinal dusky cloud behind the areoles. Arolia long, lying near the
small claws. Apical joint of the antennae, oviduct and genital segment
of the male slightly embrowned.

Described from one pair taken on the flats near the lower end
of Fallen Leaf Lake. This insect might be taken for the im-
mature form of agilis were it not for the longer elytra, broader
collar and flattened vertex. The fuscous cloud behind the eyes
is also distinctive.

Labops hesperius Uhler. One example was taken July 31 on
the shore of Lake Tahoe near the Hotel Tallac.

Vou. 1] Van Duzee.—Lake Tahoe Hemiptera 241

Lopidea nigridea Uhler. Several specimens of what I take to
be a smaller form of this species were taken about Fallen Leaf
Lake during July. This small form is but 4mm. to the tip of
the abdomen, the surface is more slaty-black with the stiff hairy
vestiture more pronounced, and the markings are paler, being
but rarely tinted with red. The male genital characters are
nearly identical and it does not seem best to try to distinguish
them at present.

Hadronema militaris Uhler. One example was taken on the
shore of Half Moon Lake, July 23.

Hadronema robusta Uhler. A single specimen was taken with
the preceding.

Paraproba hamata Van D. Upper end of Fallen Leaf Lake,
June 21. A single specimen.

Orthotylus tibialis Van D. <A good series of this very distinct
species was taken in July from a low hirsute weed growing along
the road between the sawmill on Fallen Feaf Lake and the Hotel
Tallac. The descriptions of this and other new species in this
genus will appear about simultaneously in the Proceedings of the
California Academy of Science.

Orthotylus insignis Van D. A very pretty species, of which
three examples were taken July 24 on the ‘‘short trail’’ to Glen
Alpine Springs, where a tiny streamlet caused a rank growth of
vegetation.

Orthotylus viridicatus Uhl. Not uncommon on the rank weeds
growing near the lower end of Fallen Leaf Lake, at Half Moon
Lake and near Glen: Alpine Springs. The black membrane with
conspicuous white veins will distinguish this pretty species.

Orthotylus uniformis Van D. <A plain green species of which
numbers were taken near Fallen Leaf Lake in July, mostly on
the chaparral.

Orthotylus ovatus Van D. Common on junipers on the south
slope of Mount Tallae late in July.

Orthotylus formosus Van D. Two examples of this very dis-
tinct form were beaten from alder bushes in a damp spot on the
south slope of Mount Tallae about 200 feet above Glen Alpine
Creek on July 30. I was unable to find any on the lower levels
along the creek, but it is possible that the warmer location on the
sunny hillside brought them out earlier than in the shaded valley.

Orthotylus affinis Van D. Abundant everywhere on willows

242 University of California Publications. [ENTOMOLOGY

at the lower levels, but I also took them near Half Moon Lake
at an altitude of about 8000 feet.

Orthotylus cuneatus Van D. Several specimens of this form
were taken on willows about Fallen Leaf Lake in company of the
preceding.

Labopidea nigripes Reut. A small series of this very distinct
species were taken near the lower end of Fallen Leaf Lake in
July.

Macrotylus multipunctatus Van D. The unique type of this
interesting insect was taken by me near the lower end of Fallen
Leaf Lake July 17. Its pale surface, closely dotted with round
black points, will at once distinguish it from all the allied forms.

Macrotylus lineolatus Uhler. A few examples of this form
were beaten from bushes about the lower end of Fallen Leaf Lake
late in July.

Macrotylus infuscatus Van D. Taken with the preceding but
in greater abundance. Unfortunately I neglected to note the
food-plant of these species, but it may have been the willow which
grew in several localities about there.

Microphylellus alpinus, new species

Aspect of Microphylellus rubricans Prov. but larger and darker; piceoas
black with castaneous legs. Length 4.5mm, to tip of membrane.

Head nearly as in modestus, a little shorter when viewed from the
side; projecting below the eye for scarcely more than the width of the
eye. Vertex without a basal carina. First joint of antennae a little
shorter than in the type species, surpassing the elypeus by about half its
length; second joint linear, distinctly incrassate, as thick as basal joint;
third and fourth together about as long as second, setaceous. Pronotum
proportionately shorter than in modestus; humeral angles slightly promi-
nent, sides feebly sinuated and anterior angles well rounded; eallosities
large, distinct, the impressed line strongly arched either side; the disk
minutely transversely wrinkled, sides subacute but hardly carinate.
Rostrum attaining apex of hind coxae, basal joint surpassing base of
the head.

Color piceous-black, deeper on the head, pronotum and scutellum;
antennae uniformly concolorous; rostrum a little paler. Legs castaneous,
becoming piceous on the coxae, on the base of the femora, at least the
posterior, and on the tarsi.

Described from one male example taken on the alpine meadows
on the western slope of Angora Ridge at an altitude of 8500 feet,
on July 26. This species is not typical of genus Microphylellus,

VoL. 1] Van Duzee—Lake Tahoe Hemiptera 243

differing in the shorter pronotum, more prominent callosities,
larger basal joint of the rostrum and the stouter second segment
of the antennae.

Microphylellus bicinctus Van D. Taken on the chaparral
along the western shore of Fallen Leaf Lake early in July.
Apparently rare.

Gerhardiella delicata Uhler. Two examples were beaten from
the chinquapin oaks along the western shore of Fallen Leaf Lake
late in July. There seem to be no good structural characters by
which to distinguish between this genus and certain related
genera such as Psallus, Apocremnus and Plagiognathus. By
separating out the forms having the femora dotted in longi-
tudinal series and the tibiae dotted with black we have a fairly
homogeneous group including the types of the genera Plagiog-
nathus and Gerhardiella, and I can see no harm in using the
latter name for the red forms, as they have quite a different
aspect from the more typical forms of Plagiognathus. If we do
this, it would seem advisable to divide genus Psallus, placing
those forms with ovate blackish bodies and dotted tibiae in genus
Apocremnus. It is possible that further study may disclose good
structural characters for distinguishing these genera, but I have
not vet detected any that seem at all constant.

Apocremnus politus Uhler. Taken about the upper end of
Fallen Leaf Lake and at Half Moon Lake in July. This species
is smaller and more slender than the more typical anchorifer
Fieb., but for the present I prefer to place it here rather than
in Plagiognathus, as it has the black femora and the white seale-
like vestiture of Apocremnus. In this species the second antennal
joint is entirely black.

Apocremnus nigerrimus, new species

Larger than politus, deep black, polished, clothed with coarse deciduous
white hairs; apical joint of antennae concolorous, tibiae obscurely dotted.
Length 3.5mm. to tip of membrane.

Head about as in anchorifer, viewed from the side shorter, with the
facial angle more acute; projecting below the eye for a distance equal
to the greatest width of the eye. Vertex ecarinate, a little flattened.
Antennae hardly shorter than in anchorifer; first joint surpassing elypeus
by about one-third its length; second as long as pronotum and one-half
the head; third and fourth together as long as second. Pronotum rather

244 University of California Publications. [ENtTomMoLocy

long; its length a little more than half its basal width; the sides straight,
with the anterior angles scarcely rounded. Rostrum attaining the pos-
terior coxae, first joint reaching base of the head. Oviduct of the female
long.

Color deep black, polished above, clothed with conspicuous white hairs;
base of vertex with a fulvous sinuate line which (as in politus) simulates
a carina; third and fourth antennal joints obscure castaneous, their bases
narrowly blackish. Tibiae above obscure whitish, bristles and dots at
their base black. Membrane almost as deeply colored as the corium,
marked with a small pale spot next the tip of the cuneus, invading more
or less the apex of the nervures. Osteolar orifice soiled white.

Described from twelve examples representing both sexes.
Eight of these were taken by Mr. Fordyce Grinnell at Pasadena,
California, June 5, 1909, and four I took about the upper end
of Fallen Leaf Lake in July.

Bolteria picta hirta new subspecies

Smaller and darker-colored than picta, clothed with stiff black hairs;
dull yellowish-green, most of the head, two transverse bands on the
pronotum, scutellum in part, inner margins of the clavus, a broad distal
vitta on the corium, an oval spot on the cuneus and most of the lower
surface black; femora dull fulvous, marked with large fuscous dots in
longitudinal series. Length 4mm.

Head shorter and less prominent than in picta. Vertex a little de-
pressed before the base, which is thickened but hardly carinate. Front
convex, polished. Antennae shorter than in picta; first segment but
little surpassing the clypeus; third and fourth together subequal to
second. Pronotum, scutellum and elytra as in picta, impunctate, polished,
clothed with stiff black hairs which are somewhat longer and more dense
on the head. Rostrum attaining the intermediate coxae.

Color a soiled greenish-yellow. Head polished black; a point next the
inner angle of the eye and base of the vertex fulvous, median line of the
front sometimes pale. Antennae and rostrum black, extreme apex of first
antennal joint pale. Pronotum marked with a broad transverse band on
the callosities and a narrower one on the hind margin, black. Incised
line on seutellum black. Narrow scutellar and commissural margin of
clavus, a broad longitudinal vitta on corium, a short mark within the
costa placed near the apex and a large oval spot covering the disk of
the cuneus, black. Membrane infuseated, a cloud in the apical areole and
a curved ray beyond deeper black. Beneath black, polished, osteolar
region and the lower margin of propleura whitish. Legs obscure fulvous,
femora infuscated at base and apex and dotted with blackish in longi-
tudinal series; tibial bristles black springing from small black points.

Described from eight female examples beaten from sagebrush
(Artemesia) on the shore of Fallen Leaf Lake near the Lodge

Vou. 1] Van Duzee.—Lake Tahoe Hemiptera 245

on July 5. Typical examples of Bolteria picta Uhler received
from Mr. Heidemann show but few black hairs and they seem to
be entirely wanting from specimens taken by me on the plains
about Denver, Colorado.

HOMOPTERA

Okanagana rimosa Say. Common at all levels from Fallen
Leaf Lake up to 8500 feet on Angora Ridge, the first individuals
appearing on July 3. This species has the habit of resting in
trees fifteen or more feet above the ground, and being wary are
very difficult to capture.

Okanagana ornata Van D. On July 11 I took from low oak
bushes growing on the top of the lateral moraine east of Fallen
Leaf Lake three male examples of a form agreeing entirely with
ornata except that the orange colors are replaced with pale
testaceous. Possibly the local conditions might account for this
difference in color or these specimens may not have been fully
colored.

Aphrophora permutata Uhler. Common at lower levels and
first reaching maturity about July 15.

Stictocephala pacifica Van D. Not uncommon about Fallen
Leaf Lake during July.

Telamona barbata Van D. On July 30 I took numbers of
what may prove to be this species from a willow having pale
green foliage, growing on the southern slope of Mount Tallae.
These have the crest more erect and elevated than the types from
New York State.

Telamona obsoleta Ball. One example of this form was beaten
from a willow near Glen Alpine Springs, July 22.

Orgerius rhyparus Stal. Two fully grown examples were
taken near the lower end of Fallen Leaf Lake July 17.

Elidiptera woodworthi, new species

Closely allied to septentrionalis; broader, more mottled with white;
front whitish with its base and the clypeus fulvo-testaceous. Length
7mm.

Vertex a little shorter than in septentrionalis, scarcely as long as
broad, the impressed median line almost obsolete in the female. Front
a little narrower than in the allied species, sides almost rectilinear, at
apex curving inward to base of elypeus. Pronotum as in septentrionalis,
narrowly produced to about the middle of the eyes and truneate at apex.

246 University of California Publications. [ENTOMOLOGY

Elytra broad as in variegata and pallida. Rostrum reaching to apex of
penultimate ventral segment. Hind edge of genital segment feebly
arcuated in the female, quite deeply notched in the male, leaving a con-
spicuous lobe or tooth at the lower angle next the ventral insicure; dorsal
plate of the male long and triangular at apex. In the male of septen-
trionalis the genital segment is truncated, without a ventral notch, and
the dorsal plate is short and truncated, not exceeding the plates.

Color fulvo-testaceous, becoming more gray on vertex and pronotum;
elytra milky-white with pale nervures, varied with testaceous-brown and
dotted with fuscous, the brown forming a longitudinal series of three
squarish patches on each elytron, the anterior two sometimes obscurely
united by a brownish cloud along the claval suture; inner apical cells
mostly brown crossed by white nervures and veinlets; the fuscous dots
arranged along nervures and on costal margin, where they form two
irregular marks before the apical areoles. Wings infusecated at apex.
Pronotum irrorate with brown, the fulvous mesonotum more obscurely so.
Beneath and legs almost uniformly fulvo-testaceous with a dark lateral
vitta covering the eye and shading below to black with the extreme edge
of the inferior aspect of the pronotum clear white. Front whitish, with
the base to a little below the upper angle of the eye darker and irrorate
with brown. Edges of the pleural pieces and ventral segments whitish.

Described from numerous examples taken on the Jeffrey pine,
especially along the southern slope of Mount Tallac, after July
19. They were also beaten from cypress bushes growing on this
same slope and here the young were taken with the adults. This
is of a clearer gray than any of our other species of Elidiptera.
It has much the aspect of the eastern variegata but that is a larger
and darker form with a shorter vertex. In my key of 1907 it runs
directly to septentrionalis but may be distinguished from that
form by its more whitish aspect, broader form, the mostly pale
front and the different genital characters.

Catonia nemoralis, new species

Size and aspect of grisea, to which it is closely related; ashy gray
with the elytra irrorated and mottled with fuscous and dotted with white
veinlets, the front uniformly whitish-testaceous. Length 5 mm.

Vertex short, truncate, broader than in grisea, not as long as width
of each compartment. Pronotum at the middle scarcely as long as vertex.
Mesonotum shorter than in grisea, about twice length ot vertex and pro-
notum together (in grisea three times as long). Front broader than in
grisea, but little wider toward eclypeus. Last ventral segment of the
male truncated, with a subacute triangular median tooth about one-third
the length of the valves, while in grisea it is nearly half their length.
Plates oblong, a little narrowed at apex.

Vor. 1] Van Duzee.—Lake Tahoe Hemiptera 247

Color of head, pronotum and mesonotum fulvo-testaceous, becoming
more brown on mesonotum, where the dark irrorations are more con-
spicuous. Pronotum darkened either side of the median carina, with
about four brown points behind each eye; vertex sometimes showing
dark irrorations. Elytra testaceous-gray with a tinge of brown and
marked with two obscure transverse brown bands, one, more distinet, at
the basal third, the other at the apex of the clavus, the costa with a
narrow fuscous line; all the nervures, including the costal, pale, slenderly
edged with fuscous and broken by numerous white transverse veinlets.
Wings fuseous at apex with darker nervures. Front and elypeus whitish-
testaceous, immaculate or nearly so. Pectus and legs brown, inferior
aspect of pronotum testaceous, knees, tips of the tibiae and tarsi pale.
Abdomen black, white-pruinose, the slender edges of segments and genital
pieces pale. Antennae brown, the setae black.

Described from numerous examples taken June 20 to the last
of July from the level of Lake Tahoe up to 8000 feet. This
species seems to live entirely on the lodge-pole pine, although it
was occasionally captured on other trees whence it had evidently
flown from the pines.

Catonia succinea, new species

Form and aspect of majusculus; a little smaller and nearly immacu-
late but closely allied to it in most of its characters; pale amber-brown
above, almost white beneath. Length 6 mm.

Vertex distinctly longer than in majusculus, passing the eye by about
one-fourth its length, anterior edge parabolic and obviously carinate;
median carina prominent, abbreviated before; hind edge almost truncate.
Pronotum shorter than vertex, carinae prominent. Mesonotal carinae
sharp, feebly arcuate at apex. The closed elytra almost parallel, a very
little wider toward their apex. Front narrower than in majusculus, not
obviously wider at apex, carinae prominent. Last ventral segment of the
male truneate at apex, without a median tooth, plates contiguous at base,
oblong, oblique at apex.

Color a uniform obscure amber-brown, quite strongly tinged with
fulvous on mesonotum; surface of the elytral areoles subhyaline and
beautifully green-iridescent when closed over the wings. Beneath whitish,
tinged with fulvous or amber on the front, clypeus, mesopleura and apex
of the tarsi; lateral carinae of the front pale.

Deseribed from twelve examples, representing both sexes,
taken about the upper end of Fallen Leaf Lake and along Glen
Alpine Creek during July. Most of these I beat from cypress
bushes and that probably is the native food-plant of the species.

Oliarus fidus Van D. One pair taken near Glen Alpine Creek
in July. These are a little larger and darker than those from
San Diego County but do not seem to differ otherwise.

245 University of California Publications. [ENTOMOLOGY

Cirius basalis Van D. Not uncommon during July about
Fallen Leaf Lake and at Half Moon Lake. This species has been
taken in the Selkirk Mountains in British Columbia and in
Colorado, and is common in the northeastern states and Canada.

Laccocera obesa Van D. Several females were found on the
flats at the lower end of Fallen Leaf Lake, July 17.

Liburma magnifrons Crawf. Taken with the preceding.

Iiburnia sp. A species near osborni was very abundant on a
marshy meadow near the Angora Lakes at an altitude of 7200
feet.

Koebelia californca Baker. Abundant on lodge-pole pine.
They began reaching maturity about July 12 and by the last of
the month few nymphs were seen.

Koebelia irrorata Ball. One pair was taken with the preced-
ing on July 22. It is larger, with a longer vertex, and is paler
in color and less mottled, and the genital characters are distinct.

Oncopsis variabilis Fitch. On alders about the upper end of
Fallen Leaf Lake, July 5 to 19. These are all pale yellow or
testaceous and are without the oblique elytral vitta. One male
does not differ from the female in color.

Oncopsis prunt Prov. Two males taken at Half Moon Lake
and one near Cathedral Lake, July 5.

Tdiocerus amoena Van D. Taken at the lower levels during
July.

Agallia californica Baker. Angora Lake, June 22, and Fallen
Leaf Lake, July 16.

Helochara communis Fitch. Lower end of Fallen Leaf Lake,
not uncommon.

NXerophloea viridis Fabr. One large rather gray specimen
was taken with the preceding.

Gypona angulata Spangb. A single specimen taken near Glen
Alpine Springs, July 12.

Errhomenellus maculatus G. & B. Two young and two adults
of what I believe to be this species were taken along the Tallae
short trail, July 5.

Parabolocratus viridis Uhler. Several were found near the
sawmill at the lower end of Fallen Leaf Lake, July 17.

Aligia jucunda Uhler. Beaten from chinquapin oaks on the
western short of Fallen Leaf Lake, July 5.

Vou. 1] Van Duzee.—Lake Tahoe Hemiptera 249

Aligia inscripta Van D. Taken with the preceding and on the
southern slope of Mount Tallac.

Deltocephalus affins G. & B. Flats at lower end of Fallen
Leaf Lake, July.

Deltocephalus vanduzer G. & B. Not uncommon on the low
lands and on the meadow near the Angora Lakes.

Deltocephalus punctatus O. & B. Two taken near Fallen
Leaf Lake. This seems to be scarcely more than a pale form of
fuscinervosus.

Phlepsius ovatus Van D. Taken at the lower end of Lake
Tahoe, July 17.

Phlepsius occidentalis Baker. Lower end of Fallen Leaf
Lake, July 25.

Euscelis exitiosus Uhler. Common up to 7500 feet.

Euscelis striolus Fall. One example was taken on the mead-
ows near the Hotel Tallac, July 31.

Thamnotettic geminata Van D. Common about the lower end
of Fallen Leaf Lake in July.

Thamnotettix longiseta Van D. Two examples taken with the
preceding.

Cicadula laeta Uhler. One pair taken with the foregoing,
July 17.

Cicadula sexnotata Fall. Abundant about the Angora Lakes,
Half Moon Lake and Grass Lake. Here as elsewhere it lives on
fine grasses growing near water or in damp situations.

Balclutha impicta Van D. Two females were taken in a
swampy meadow at the foot of Angora trail in July.

Eugnathodus abdominalis Van D. Not uncommon in fields
about Hotel Tallae during July.

Dicraneura carneola Stal. Abundant in a swampy meadow
near the Angora Lakes and more rarely in grassy places about
Fallen Leaf Lake and Grass Lake.

Empoa commissuralis Stal. Near Glen Alpine Creek, three
examples.

Aphalara calthae Linn. Beaten from chaparral along the
western shore of Fallen Leaf Lake in July.

Euphyllira arbuticola Crawt. Three examples taken with
the preceding.

Euphyllura nevadensis Crawt.
the preceding.

Common on Ceanothus with

Transmitted March 20, 1916.

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. Synoptical Keys to the Genera of the North American Miridae, by
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Vol. 1, No. 6, pp. 251-299 June 17, 1916

LIFE-HISTORY AND FEEDING RECORDS OF
A SERIES OF CALIFORNIA
COCCINELLIDAE*

BY
CURTIS P. CLAUSEN

The studies herein outlined of a series of California Coccinel-
lidae were undertaken by the writer at Sacramento, California,
during the spring and summer of 1913, and coneluded at Berke-
ley and Riverside during the season of 1914. Due to the change
of locality, a certain unavoidable lack of continuity was made
necessary, and the variable contributing factors of temperature,
humidity, aphis supply, ete., were thus more strongly accentu-
ated than would have been the case had the experiment with the
different species been conducted simultaneously under definite
and regular conditions. A further discussion of this matter will
be presented in connection with the particular cases as they
arise.

The coccinellid species used for the tests are all of fairly
general distribution over the state, and in general may be said to
be the eight most important aphis-eating species in California.
They are: Hippodamia convergens Guer., Hippodamia ambiqua
Lee., Coccinella californica Man., Coccinella trifasciata Linn.,

*Paper No. 14, Citrus Experiment Station, College of Agriculture,
University of California, Riverside, California.

The investigation of this problem was begun and largely completed by
the writer while field assistant in the Bureau of Entomology under the
immediate direction of Mr. W. B. Parker, and concluded while in the
service of the University of California. Permission for publication has

been granted by Dr. L. O. Howard, Chief of the Bureau of Entomology,
United States Department of Agriculture.

252 University of California Publications. [ENtToMoLoGy

Olla oculata Fab., Olla abdominalis Say, Cycloneda sanguinea
Linn., and Adalia bipunctata Linn.

The writer makes no assertions with respect to the validity of
any of the above-named species. The belief is quite general that
Hippodamia ambigua is a varietal form of Hippodamia conver-
gens, and the same claim has been advanced regarding Olla ab-
dominalis with respect to Olla oculata. Investigations are now
under way to determine the exact status of these forms.

LABORATORY METHODS

At the time the investigations were undertaken an extensive
series of tests was made to determine the most satisfactory method
of caging the individuals during the period covered by the ex-
periment, in order to secure as nearly as possible ideal conditions.
Potted plants infested with aphis and covered with chimney
glasses were used for a time, but did not fulfill the requirements,

due to the fact that moisture often collected on the inside of the |

glass, and also because the adult beetles showed a strong ten-
dency to ascend to the top of the glass and there remain inactive,
rather than to feed normally on the infested plant as desired.
Later plaster of paris containers, jelly glasses, ete., were used
but with indifferent suecess. By far the best results were finally
secured by confining the individuals under observation in ordi-
nary three-inch vials with cotton stoppers. Experiments upon
larvae necessitated the covering of the stoppers with thin tissue-
paper to prevent injury through entanglement in the cottony
fibers. The stoppers used were light enough to permit a reason-
ably free passage of air, thus maintaining approximately outdoor
conditions as regards temperature and humidity.

In the feeding tests, both with larvae and adults, a certain
definite number of aphids of as nearly a standard size as pos-
sible were placed in each vial early in the morning and the count
made at the corresponding time the following day. The number
of live aphids present was determined, and this number sub-
tracted from the original number gave the total for the day.
Cheek vials containing one hundred aphids each were also main-
tained to determine the approximate mortality from natural
causes, and the proportion determined subtracted from the total
of all reeords for the current day.

Vou. 1] Clausen.—Life-Histories of California Coccinellidae. 253

In the experiments conducted at Sacramento the hop aphis
(Phorodon humuli) was used for all feeding purposes, unless the
contrary is stated. At Berkeley and Riverside the rose aphis
(Macrosiphum rosae) was used. More difficulty was here experi-
enced in securing a standard size than was the ease with the hop
aphis. As nearly as possible individuals approximately two-
thirds the maximum size of the adults of the species were used, in
order to obviate the possibility of the production of young during
the day.

In the lfe-history experiments, observations were ordinarily
made once each day and the cast larval skins removed. The moult
was credited in each case to the day upon which the exuvium was
removed, as it was impracticable to make observations corre-
sponding to fractional days. The date of pupation was recorded
upon the day during which the median dorsal split appeared in
the fourth larval skin, even though this may have taken place
several days after the larva, having ceased feeding, attached
itself to some solid base, and assumed the general appearance and
form of the pupa.

In order to secure complete egg records it was found neces-
sary to make the count and remove all eggs three times each day,
because of the strong tendency of the females at times to devour
their own eggs, even though an adequate supply of food was
present at all times.

In the tabulations as given in this paper the following sym-
bols have been used to designate certain things. They are as

follows: L = Eggs deposited

H = Hatched
* — Moulted, or death of male
P = Pupated
EK = Emerged
M = Mated
iD =—Dead
C = From cold storage

Coccinella californica Mann.

The individuals of this species are the largest in size of any
of the common California coccinellids, the fully developed larvae
measuring 10.2 mm. and the adult beetles approximately 8.0 mm.
The elytra of the adult normally exhibit no markings of any
kind, yet specimens have been found which exhibited spots of

254 University of California Publications. |ENtToMoLocy

various sizes and in no definite position. Upon rearing the pro-
geny of these individuals it was found that to a considerable
extent the offspring bore the characteristic markings of the
parents.

Oviposition—Upon the basis of observing ten individuals, a
period of time varying from two to three days was found to in-
tervene between emergence and mating, while from mating to
egg-laying the time elapsing varied from ten to fourteen days,
with an average of 11.9 days. As regards the period of oviposi-
tion, a range of from 20 to 51 days was found, averaging 31 days.
During these periods the proportion of days upon which eggs
were laid varied from 60.0 to 74.1 per cent, with an average of
69.5 per cent. The maximum number of eggs laid in one day
was 24, the maximum number of eggs for the period was 360,
and the minimum 127, thus giving a daily average of 8.0 eggs
and a period average of 247 eggs.

EeG-LAyIne Recorps—Coccinella californica Mann.
Berkeley, 1914

Vou.1] Clausen.—Life-Histories of California Coccinellidae.

E@G-Layinc REcorpsS—Coccinella californica Mann.—(Continued)

Date
May 1

16

eS eS
© OO -]

+ ©

UH ow bo

a

bo & po tf po & to

~_

Total

Average

1

335

sll

Berkeley, 1914

4

5

6 7
12 ef
a es
9 16
AY og
ae 12
PR eee
8 8
Ap, +e
a. 14
15 o
if 6
11 15
eas 1
12 Ie
ok 8
JQP ee
4 21
nae 6
3 14
Dy eee
— 10
—. 9
esses D
205 8245
7.3 7.2

259

NT
mcaranpbwwHNo.

10
11
12

13

ic
z e

a a a zn a ax a Pian

&h = Be = 3a + Se x os f6 os oa
5 Ss «0 2 «8 po a eo 6S Seas
an g Sg Bg Oe Se ee) se oe
Cem ty} os A os co os co os ee Oe S| Os
AS ica) =, a Hn a peal ors ony Hh SS Sr cal Ha
5/1 ay 6) 5/5 385/20) 5 5/728 3 OEY) oo Of 4
YA yfT/ 6 5/17 10) 5/2 4/24 3 os 7 (GD) ane
6/3 6/8 5 6/13 5 6/17 4 £46720 3) § 6728 8 7/2 4
6/3 6/8 5 6/18 5) 6/l6o= 3 6/20 4 6/26 6 7/1 5
6/3 6/8 5 6/3 5) 6/17 4 16720) 73) 6/27 tt 7/2 5
6/3 6/8 ys GAR fb GAG 8 GRO 2 GAO @ Zl 5
6/3 6/8 Gye sy Alyy 2h VAI By 7/2 5
6/3 6/8 by yale) Bs yale ZB Ga 27/ 7 7/2 5
6/3 6/8 5) 36/18) 5) 6/16) 38)" 6/20" 4 6/726 Oi 2 6
6/5" 16/1 5k 6/6) 6/19" 2) 16722) se 6729 if 7/4 5
6/5 6/11 6 G/16" 5 6/18 2)" 6/22" 46/729) 7 w/e: 5
6/5 6/11 6 6/16- 5- 6/18 2 6/22 4 6/29 a 7/4 5
6/5 6/11 6 6/16 5 6/19 3 6/22 4 6/29 -7% 7/4 5
Average 5.4 5.7 3.3 3.4 6.8 ah

256 University of California Publications. |ENTomMoLocy

Life-History.—The records of thirteen individuals give the
average hatching period during normal summer conditions as 5.4
days, the second period from two to five days, the third period
from three to four days, the fourth period from five to eight
days, and the pupal period from four to six days. The averages
for these periods were 5.7, 3.3, 3.4, 6.8, and 4.5 days, respectively,
which, in connection with the egg stage of 5.4 days, gave an
average life-cycle of 29.1 days. A rather peculiar variation is
noted in Nos. 1 and 2, in which ease the first larval period was
lengthened to eight to ten days, respectively, which is an increase
of three to five days over the normal of five days as represented
by the ten other individuals. No noticeable variations appeared
in the later stages and the life-history of the first individual was
complete in 31 days, or slightly less than two days over the
normal. No appreciable difference in size was noted. Individual
No. 2, referred to above, died shortly after pupation. In the case
of nearly all individuals, a period of from one to three days
elapsed from the time feeding ceased until the last larval skin
was cast. This is the only species in which this period of in-
activity was so uniform and pronounced.

Lire-History—Coccinella californica Maun.

Sacramento, 1913

25 Total days
— of stages

16

4

[=

ponmnnwnnwnnwnwwhds Ww
Nonrwnwvdr © O

co

June 1
9

a

Total

Average

Vou. 1] Clausen.—Life-Histories of California Coccinellidae. 257

Feeding Habits—The larvae of this species, as would be ex-
pected, consumed by far the largest number of aphids as com-
pared with those of other species during its period of life. The
records of thirteen individuals give a daily feeding average of
24.9 aphids, while for the entire period the maximum reached
600 and the minimum 394, the average for all individuals being
475 aphids during the larval period of approximately 24 days.

The unusual number recorded for May 22-23 was due to the
excessive heat, all the aphids in the check vials being dead upon
these dates. It was not possible therefore to make a propor-
tionate reduction in the individual totals.

LARVAL FEEDING RECORDS—Coccinella californica Mann.
Sacramento, 1913

1 2 Date 3 4 5 6 uf 8 9 10 ak 12 i133
H H June 8 H H H H H H H EDR eg baler ae ee.
2 iL Oe Se es rie a oy Cees Bg oes, ae ee ee
5 5 10 it 2 2 1 2 3 ey ee eth en ene
3 4 ial 3 4 2 3 2 4 5 3 H H H
4 4 12 4 4 4 7 6 4 8 6 0 1 0
7 6 13 Gea ee ee ee One edn Be Gc et 3 il
9 9 14 9 5 6 6 8 8 6 8 2 4 4
on 4 i aki 9 12 8 7 @) alal 7 3 3 5
8 8 16 Oeee he ae ae ee Oke 9 Ce TD ei ae
13 9* Wey NB hay aS ale oer Iz al} 9 8 8
14 10 IS 24 Si 2358 Se 21 22) 235" 24 9 Bur (Ge
6 11 ORS Oke OM 2D 2S os eo Oreo lem ee llGime Omni
14* 9 210) BX By Bile Be Bie Gye Ber Bie BAL aly? al
ie Ue 21 «6400 490—CO GB 4RC CC OST
1) als 22, 97% 92 88 100 92 96 87 #92 G66 “69 72%
2OR 25 2B al; ey aS ey aie ey aly aly 7) rh tS}
46 26* 2A 82 54 74. Vol 26 41 63) 47 O69 59
79 43 Sy Pat BIL | es ee ene AL) ee tae Meee 83 74 76
127 109 PAD nee Pgh. 2 IPSS oe cad eee ee P 86 66 49
3 63 DH eit seen te 8 10 Jy) eee 12 12 eae 54293) b3
ses 28 28 2 de ee Og WS ey ee eee ree ee 49 45 43
12S anes 2 OF eae Pe mime rience, pts GRRE Se oe ah fee 12 1 12
5 SS Se eee veeea tay gn ee ee Se vials WAS a AL 5 TR ew Sng eee Cee
are P Sully al ee Rie ee Y Re ee eee eee Bett ae Seated ee perm Cee
Beer) Yt 2 1D) ieee NY Ween peels E E Rik oS sson eee, aes
aes D Soot Beate ea Pa aS ONO ee a ES OR he EO «ee
Be) 222s A ge OR te BGs heres ee Ex | WR ws E E E
435 394 525 441 472 450 423 444 471 456 569 600 528
22.9 19.7 39.8 29.4 31.4 30.0 28.2 29:6 31.4 30.4 35.5 35.3 33.0

258 University of California Publications. |ENToMoLoey

Ten adults during a fifteen-day period consumed an indi-
vidual maximum of 661 aphids and a minimum of 414, with an
average for the series of 500 aphids. Upon a daily basis this
would be 34.0 aphids per day. During the period the number
eaten daily ranged from 16 to 62, which is considerably lower
than the record of Hippodamia convergens, a condition not to be
expected from the relative sizes of the individuals of the two
species. This numerical relationship is not evident in the case
of the larvae, and the difference may in all probability be at-
tributed to the variation in climatic conditions existing at Sacra-
mento and at Riverside, California. In this experiment the rose
aphis (Macrosiphum rosae) was used exclusively.

ADULT FEEDING RECORDS—Coccinella californica Mann.
Riverside, 1914

Date 1 2 3 4 5 6 7 8 9 10
Sept. 26 31 41 26 37 34 42 18 37 ais) 24
27 26 33 29 30 25 26 29 26 29 30

28 47 28 41 21 41 35 23 18 45 19

29 49 51 37 29 49 17 41 36 28

30 62 29 35 3 3 24 36 24 2 39

Oxetis. Al D1 43 23 20 40 29 20 18 23
2 29 47 41 28 32 32 iL%/ 29 34 27

3 58 54 32 26 47 27 30 27 43 32

4 47 35 60 42 36 41 21 42 29 17

5 39 44 28 31 25 36 26 18 38 45

6 42 26 33 37 16 21 18 33 52 42

i 26 38 41 25 34 28 35 15 26 36

8 51 47 39 34 29 35 32 41 44 24

9 57 2 25 38 26 19 46 28 39 30

10 46 51 39 24 37 24 33 22 37 52

Total 661 609 529 465 524 436 428 414 574 468

Average 44.0 40.6 35.2 31. 34.9 29.0 28.5 27.6 38.2 31.2

Coccinella trifasciata Linn.

This species is not of any great importance in California be-
cause it occurs only in limited numbers in most sections, though
in the San Francisco Bay Region it may be found in very con-
siderable numbers, at times to an extent even greater than Hip-
podamia convergens. The adults ordinarily pass the winter
singly, hidden away in bunches of grass, leaves, and in other
protected places. This is one of the first species to appear in the

VoL. 1] Clausen.—Life-Historics of California Coccinellidae. 259

spring, and in several cases observed has done good work in the
control of the rose aphis.

Oviposition—No records were secured of the time interven-
ing between emergence, mating, and egg-laying. The period of
oviposition extended in one case over 42 days, while the average
for ten specimens was 29.2 days. During the period the propor-
tion of days upon which oviposition occurred ranged from 66.6
to 78.1 per cent, with a general average of 75.0 per cent, which
indicates a quite general uniformity in this respect. The maxi-
mum number of eggs deposited by an adult was 369, the minimum
109, and the average 249 eggs; the maximum number deposited in
one day was 31, while the average for all individuals was 8.4, with
a range of from 4.7 eggs to 10.4 per day. It will be observed
from the following table that the individuals depositing the larg-
est number of eggs, as a general rule also gave the highest daily
record.

Ea@e-Layine REcorpsS—Coccinella trifasciata Linn.
Berkeley, 1914

Date 1 2 3 4 5 6 iti 8 9 10
N70 keel ieee ee a eee 1M eer oa SS SN eS
1 SS ee oa Oye gescsetel> Sey eee) Seasee | eeers
COMERS Gen kal) hiae OS 22am Reine) Sen, ete Boek

21 (0% seme ye ees Tiaccesr ee EIR On ae cere A aetiee

22 (3) Sy ee Le eee eyes LS) aoe eee peer see

PAY eee I> ees all epee Py cae Oe See ee
24 12 15 11 a) 4 ie ene aes 2 12a

25 9 9 8 7 0 Se Se Sip ue

26 13 22 6 4 13 2 1 eee Se eee

1 le Nese a Greg se 3) eae als) Om Saat 3

28 11 8 3 11 5 ial 3 23 16 8

PA) ee rest 2 a 9 aly 20 16 16 hy es
30 irs Veeces Teli ieee’ Smeets Pea Fee 19 13
May 1 Ys ago ea eee aye 1S pee 13 12 9
2 15 13 14 2 1 6 18 8 Cees

3 6 ieee 14s Be 5 20 6 32 19

4 ay A py ee 6 Dees | ce ret Vile eee

5 2 VG Same 16 WP Meee 14D eee 30

Gp weed eee ts 5 8 Ae ete 4 ee ee 5

7 25 2 ers Ib), howe Ly 8 17 12 M7

8 22 13 Gee: 5 16 19 6 Sai Pes

9 14 Zope Medea’ F cu2tes! 11 23 apt 2 23 16

10 Oe gee 6 Us Se 8 ee 1 21
1 Clog wrens Meee NG yet ee D> ests 6

12 4 15 5 13 2% 15 (ieee 9 #)

260 University of California Publications. [ENToMoLoey

Eee-LAyIne Recorps—Coccinella trifasciata Linn.—(Continued)
Berkeley, 1914

Date 1 2 3 4 5 6 7 8 9 10
ie. eee 2 2 21 21 13 ut as eee ee 2

15 Oe D 12 Gi Sc Ste eee Ge S32

16 14 3 e me 6 D 9 2039 =) ee D
any, if De eee iL See ee the GS gia

18 15 6G. (ES ee 1 a a LQ y Gee

Gigs ae BF ee i 5 VG_7 Se Oy

20 O30 eee ee a eee ee 2G. yah Sipe

2a le ees 2 ee NPA ezeyse 19 Qi” eS (ee

22 22 IDS te PAV) ae 4 Singh ae 122

23 So la ee Geeereg eo 8 Die Meee 8x ope
24 oo ames sat eee ee i Gis ge =: “eee es ie ets 13) ote

25 NGi2 ii 4) Se Uf Fes : GS 4

26 Da Stee eee TIS 4 sere Des ae ee ree j= “eee

Di i" geRsay Ree We a 14 D 6. Se

28) o fit TEN © - eg eee eS Dis pea <2 Re a

29 Dy hme geo a Moe Ar AY ce

30 DUS ey nee (he ae i eer one eae Poe 5° jae

Billy, “es Rete) A Se oe Se Dit) 2)» ES eee
iC ale en ee ee | Ce ag 2 Ss eee Soe
DP Ae A E22 oe eee Tig Ores ee ae en ee 16°

Sask Noe Seer, pS ne =< > ie. Sees 3) ee

Aa tom ee ee Be) oes ei, Oe eae LS 9

ES rE eee eee eee ae ee ee Di =
Total 349 251 1109) 303) 209) 2845 353 sso seelos

Average 8.9 8.6 4.5 (ott 7.4 8.1 10.0 8.1 8.4 8.3

Iife-History.—The entire series of individuals used in this
test was secured from a cluster of eggs deposited by one female
on April 20. These hatched six days later and the adult beetles
emerged in from 23 to 29 days. The first larval period, with
three exceptions, required five days, the second three to five days,
the third from three to four days, the fourth six to eight days,
and the pupal period from five to seven days. The average for
twelve individuals for the entire cycle from egg to adult was 31.8
days. .
Feeding Habits—With complete feeding records of twelve
larvae as a basis of comparison, it was found that the total num-
ber of aphids eaten varied from 217 to 375, and averaged 294.
This, when reduced to a daily basis, represented 15.8 aphids per
day during the stage of approximately 26 days.

Vou. 1] Clausen.—Life-Histories of California Coccinellidac. 26

Date eggs
laid

bo No
ee
SoS
dS bo
SS

tS
™~
bo
oO

4/20

Average

oman SD OP &
i
~
bo
(=)

Lire-History—Coccinella trifasciata Linn.
Berkeley, 1914

Length of
stage, days

4
4/26
4/26
4/26
4/26
4/26
4/26
4/26
4/26
4/26
4/26
4/26

SS Eggs hatched
rors

DAARVWAAAAIAGAAHGAS

a |

a 1st moult
=

Length of
stage, days

aAaNMtAaoaoa»#»aa&a»n#nhaaa4an»§

5.25

d moult

5/4
0/4
5/5
5/5
5/9
5/95
5/7
5/6
5/6
5/6
5/6
5/6

a

we RS

On x

qd 3

=e )

ee 8

on ro

Ha oD
5/7

oo
SS SS
coon

SS a
on
S<
Ya)

~_

ie a
32 a6 32 Zs S
ais) a8 «at So a
o 8 Sa 08 = ®
Hh ae cs) 4
3 5/14 7 5/720
3 5/14 7 5/14
3 5/16 8 5/21
3 8/15 T 5/21
3. 5/14 6 5/19
4 65/16 7 5/22
3 5/18 8 5/25
4 <d/17 7 5/23
3 S/17 8 5/22
3. 5/17 8 5/22
4 5/18 8 5/24
3 S/1T 8 5/23
3.20 7.4

i

ew bo op Total days
© © of stages

AANMAANA AH 1 oO stage, days
(Se)
=

5.7 31.8

The adult records are given for ten individuals for a fifteen-
day period; the number of aphids eaten per day ranged from 6
to 44 per day. The period totals varied from 383 to 470, with an

average of 435.

Date
April

26
27

a or WwW DY FH

The feeding, with the exception of an occasional
day, was very uniform, due no doubt to the evenness of the tem-
perature conditions during the time covered by the experiment.
Medium-sized rose aphids (Macrosiphum rosae) were used
for feeding purposes.

LARVAL FEEDING REcoRDS—Coccinella trifasciata Linn.
Berkeley, 1914

1 2

H H
0 2

3 2
4 6

7 8
8* 5*
7 8
10 9
Oye AG
12 14
il if

Gnea on jens

*

—

e
wOonnNIoaur- Oo wFwr

> dt be

*

*

*

qe) Ses ES TN

12

14
O*

iy

*

YNoupoaaw tt?

10 ital
H H
0 1
2 i
4 3
6 5
7 OF
10* 9
13 9
18 11
12 16
12* 24*

Qnanorauw

*

pay ey pe
oon ee
*

262 University of California Publications. [ENTomMoLocy

LARVAL FEEDING RECORDS—Coccinella trifasciata Linn.—(Continued)
Berkeley, 1914

Date 1 2 3 4 5 6 i 8 9 10 ial 12
May 7 145 18 22 LG 20 16 250 ele 18 15 23 14
8 12 16 NG) yale 8) 22 18 23 dl M7/ 19
9 21 24 20 23 31 Pia ke) 16 Pa as NL 23*

10 2 22 31 18 26 25 23" 2il* ail 26 30* 27

ial 23 31 25 NG, 29 9 27 30 19 33 26 22

12 20 24 15 26 25 16 24 18 26 31 29 26

13 29 27 22 34 30 23 34 26 29 28 17 33

14 le 12) 24 7 P 32 27 32 18 19 32 19

WO fercse5 ee See U I ene 19 19 28 25 dl 37 22

GS Oe eee Meg eee) ae 124 8 It/ 36 23 25 16

WNT whey A eee co eee 12 12 12 12 19 Ie

WS. ees ee ED EE ee PO ees

OP Wess by ee By hs (Se: Si ee eee

20 El (Alle Sky VE cel ee ee

2 ee yy ee E My, es. et eet!) GOED eg Ge

YE ee ae SAN | tier ig (ern es. E De ceces:

23 ees) Yor 2 ee pee peasant meer Ei oto een eee E

DA nskcie Eencc, eA Pees eects, ee ee ee BE

25 ea See Settee Dees en eee I A229 be eee
Total 217 249 269 254 266 279 315 304 329 350 365 313

Average 13.5 146 14.1 14.12 15.6 15.5 15.7 15.2 164 184 17.3 15.6

ADULT FEEDING REcorRDS—Coccinella trifasciata Linn.
Berkeley, 1914

Date 1 2 3 4 5 6 U 8 9 10
May 2 27 58 32 21 34 27 31 18 36 29
3 32 25 35 19 32 29 37 23 34 23

4 29 23 12 32 Uf) 22 7, 15 39 18

5 ig) 31 18 26 7 30 12 36 27 26

6 3 33 19 i 22 26 18 31 22

7 34 29 30 29 18 37 14 22 23 27

8 30 37 25 38 31 34 18 29 30 32

9 36 36 37 22 24 42 36 25 25 37

10 35 ug 32 2 30 26 29 33 43

11 31 28 44 24 28 32 30 30 26 22

12 20 33 26 36 27 39 24 28 21 30

13 37 39 37 41 1g 28 33 29 29 36

14 34 28 31 33 37 27 42 37 31 18

15 28 31 38 35 28 33 28 26 33 24

16 39 35 23 30 if 38 3 38 27 31

Total 468 465 439 444 383 470 388 407 449 418

Average 31.2 31.0 29.2 29:6 25.5 313 25.8 27.1 29:95 ais

Vou.1] Clausen.—Life-Histories of California Coccinellidae. 263

Hippodamia convergens Guer.

The twelve-spotted coceinellid (H. convergens) is very gen-
erally distributed throughout the state and is by far the most
important member of the coccinellid group as regards aphis con-
trol by natural means. This species is shipped by the millions
into Imperial Valley each spring for the purpose of combating
the melon aphis (Aphis gossypii). During the late summer and
fall months the adult beetles gather in great colonies in moun-
tain canons and other cool and protected places, and may be
found in great quantities among the dead leaves and brush or
clustered upon rocky surfaces.

The aphids upon which this coeeinellid feeds are extensive in
number, but the main species infesting cultivated plants are the
hop aphis (Phorodon humuli), the rose aphis (Macrosiphum
rosae), the beet louse (Pemphigus betae), the oleander aphis
(Aphis nerit), the bean aphis (Aphis rwmicis), the plum aphis
(Hyalopterus arundinis), and the melon aphis (Aphis gos-
sypu). Feeding has been observed to a comparatively limited
extent upon the walnut aphis (Chromaphis juglandicola), the
woolly apple aphis (Schizoneura lanigera), and the cabbage
aphis (Aphis brassicae). The elm aphis, though very numerous,
was very seldom attacked.

The Egg—The eggs of H. convergens are 1.4 mm. long,
spindle-shaped, and bright yellow in color, and are deposited ver-
tically in groups of ten to thirty on the under sides of leaves and
in other sheltered and protected places. A short time previous
to the time of hatching, the egg loses its bright yellow color and
assumes a grayish tint. The emergence of the young larva ordi-
narily takes place in the course of the day during which this
change occurs. In the great majority of eases all the eggs of a
given cluster will hatch within the period of one hour, but oceas-
ionally the intervening period may be considerably longer, in
which case the larva first emerging may devour the contents of
the remainder of the cluster.

In order to determine the approximate percentage of fertile
eggs among the total deposited four hundred were collected from
the foliage of a prune tree and an accurate count made of the
emerging larvae. <A total of 332, or 83 per cent, was secured.
Of those failing to hatch, a few were broken by young larvae,

264 University of California Publications. [ENtToMoLocy

but the great majority were sterile, as was evidenced by their
shrinkage in size and the loss of the bright color characteristic
of fertile eggs.

The Larva—tThe larva emerges from the egg through an ap-
perture of irregular outline across the upper end. For a period
of from fifteen minutes to half an hour or more it remains with
the body protruding vertically about two-thirds of the way out
of the shell, during which time the body becomes hardened suf-
ficiently to permit locomotion. At this time the larva is bright
yellow in color but gradually assumes the markings characteristic
of the species. Twelve to twenty-four hours usually elapse before
feeding commences, the larvae in the meantime remaining
grouped upon or in the immediate vicinity of the cluster of
broken shells.

The Pupa.—At the time of discarding the fourth larval skin
the pupa is bright yellow in color, but in the course of one to
three days, depending upon the conditions of temperature and
light, the dark reddish brown base color with the black spots is
developed. At the time of emergence the pupal skin breaks at
the median line of the thorax and the adult beetle pushes its
way out.

The Adult—At the time of emergence from the pupal skin
the adult is a uniform yellow over the entire surface of the body
with the exception of the head, which is black. The natural color
appears within the course of a few hours, becoming considerably
darker, however, after one or two days. Temperature and lght
apparently are very strong contributing factors as regards the
degree of color development.

Oviposition—F ull records of oviposition were secured of six
females taken from cold storage at Sacramento, California, May
2, 1913, and of five females reared from pupae collected at Berke-
ley, California, during the latter part of March, 1914. Mating
took place between beetles from storage from one to ten days
later, the average intervening period being 4.5 days. The time
elapsing between mating and oviposition varied from eight to
thirteen days, with an average for the six specimens of 10.5 days.

The length of the period of oviposition as determined from
eleven individuals varied from 11 to 50 days, averaging 33.3 days.
Within these periods the proportion of days upon which eggs
were actually deposited extended from 28 to 72.7 per cent, with
an average of 63.8 per cent.

Vou. 1] Clausen.—Life-Histories of California Coccinellidae. 265

The maximum number of eggs deposited by a single individual
was 609, and the minimum 94, with an average for the eleven
specimens of 299. The daily averages ranged from 5.9 to 15, with
a general average of 8.9 eggs per day. The maximum number
deposited by a single individual upon any one day was 43 eggs.

Eec-LAvYInG REcoRDS—Hippodamia convergens Guer.

Sacramento, 1913 Berkeley, 1914

Date 1 2 3 4 5 6 Date Of 8 ) 10 11
May 2 C C C C C C Mirehy 24nihe sy ge ietal tee Mee Brees Sal
Me 0 esc Bisse seek Sagas Dy dies eas cesta SMe ori eee Ne
Ae eee) stecee eS teks aleve 2 Oe OM ese ee bute nsdee:
5 le gS eed cee en eee 2i( i eRe eee Metta DS oie 12
CE) ighek gs Anna Re eke 2S et Ds SOAs soe (crane ee
emer) eek LS tae Lalas 29 7 oe eae 9
Ce eer ee Sg OS seats sees BAO) cet Gs ieee ee 14
4) eed oe 31 Sie wits: OP 2se2t Nese
aI RReeneneeen ce. Po een etic s eS) pie te OX 0) ert) Wal Ca es eee heres eee Meee
TL coseel ERR eee eee eeene s Cree oe IZ eS aeeesees igh
12 gees. ge Se ee Die eo See See ee
1G} See Pins cored Wieceane Utter A BW RES * esc! eee 10
14 Sr es Sie OP eS 16 0 yalogee
Eee Ne ecetes ceccct) nce (Sacsath, ° dunt Gig Oe eee: 16 Se
ILS), Se eer LOR ames aera 7 che my AUD teas 12 18
> es MOP steed nt re tuaee ne. bebe Sih as gee Digeer 22
1c): eee lees Peer 16 amie ea ie ee 14
1) eee ee A Oe fees eee. hee 10 26 23 20 8 6
AD) pee Way IG) Gee 21 Tat Se ona 17 + =20
PAW erecta 5 5 UI sae eae eee 42 eee (oO 22 eo
23H la Ce See ZO ee ee 3 30 34 O) ase 31
23 dk PA SIPS Na WY ere eres I a ee 12) 2,
comely. 20) 2 SO) 52.23 16 ey 2 Oe ee 21 5
come ta 16) 2... NZ etcilie eatees 16 S29" lS 10h als
Omeeeltsn 43) 2222 Phebe eer amy eee ee ete Gt wide ineccees
re neces gl Aen as) vee 18 Seep np eee LZ ali2
223) ee LD SYS ee eee MOeF ly ech ints Oe ce 6
OMe Wye A ees! “elec 25 22 PAD) eee Sig eee 23 «618
sO M3 24 22... Ae ee 23 2h 12 O elit~ Nel6e (ec2
set | 30 sce Sy nice 7 OVA) Bare 19% SLO Se ao
June 1 22 See: Dee es 23s 28a tee Aipok dusts
ies: 329) 22. 2s. See 2A i 29%) 22 O 9
S) Zhe) MUG gee een es eres 2Ow 2G 24 eo 5 23
it) SP. ae 8 14 26 .e eae 8 4 Sea a ae
i ee Te recast) Gezaces, Sees 15 7 (eG ae ee UD eee
6 20 O) ENaereNeer Dy see: ee 5 aera) 2 Seeker ibe
ih Pail IG Sees eee 14 PAST ncaa ae eRe eee
ae ch? Gees) aN G10) ies 10s Web eee 4

University of California Publications, [ENTomMoLocy

WN
op)
lor)

EeG-LAyIne REcorpS—H ippodamia convergens Guer.— (Continued)

Sacramento, 1913 Berkeley, 1914
Date 1 2 3 4 5 6 Date 7 8 9 10 11
Ain YY Ae ees ae cee Pail eee May 1 Ou) eta ee eee D
11 O eel: pia ill Go ster 5 eae eee Pip eee EEE Gas 22)
OE er ai CBS ere eet Berar, ae ai eee 23)
12 22 (Pee, Serres Ceri ores A ees NG Acca
tS Pa Ong 61 eee ge ee 5 D eee D 2
Ane WH tab 22 eS eee es ees 6. ee 26h «22h oe
15 u Sy ae a ee tere 2 so. Se
16 Ops Sha D Bigset) es
1 oe Zl), 22 ee 14 oa Vee
18 DY? AG a. oe See ee NO: ) 2 eee ee
1 = D9 sec ee D 20 ae ee id 2
7A) eee Gig iv C ake eee 22 Ne ere 2.
Bil eee: Dot tek) (ieee ee ee U3) ae 23) 3 eee
Vp Roe ee Eerie, ees yal eee 21 4 Ose hy Oe
DBs 2 ceeees ly Soe e®, gee Re ee ee alisyy eto DD!
ee ee [D9 FA 7 ees ee 9 WG) ei ee ee
QO. ( Sstee) Gti Coeds. A aes D WG 2222 Pee) oe eee
Total 425 466 94 141 115 234 371 609 176 280 326
Average 11.4 13.7 5.8 11.7 4.6 6.1 97 12.17 5:8 Ooo

Life-History.— Under normal summer conditions in the Sacra-
mento Valley the life-cycle of this species from egg to adult is
accomplished within a period of approximately four weeks. The
variation in this respect of eight individuals from the same parent
extended from 27 to 34 days, averaging 28.75 days. This period
was divided as follows: egg stage, 5 days; first larval, 3.9 days;
second larval, 3.6 days; third larval, 2.8 days; fourth larval, 6.5
days; and the pupal stage, 7.5 days. The period of incubation
of the eggs varied only a few hours among the eight individuals,
while the variation in the first larval stage was from three to
eight days, the second larval from two to four days, the third
larval from two to three days, the fourth larval from six to eight
days, and the pupal stage from seven to eight days. It will be
seen that with few exceptions the period intervening between
moults was quite constant and distinct for each stage.

Vou. 1] Clausen.—Life-Histories of California Coccinellidae. 267
Lire-History—Hippodamia convergens Guer.
Sacramento, 1913
=
Z Ey)
2 5
ag n a a ra ey a 2
& Scares of = Se = Se 58 Se a SG
ey See emer ee ee 5 a Peas Pee ay Vee
2 fea ee SL ey a Seer sie ee Se
ios) 5/23 5 5/26 3 5/30 34 6/1 2 6/7 6 6/14 of
Peo sis. 5/23 5 5/26 3 5/30 4 46/1 21 6/8 a 6/16 8
eos 5/23 5 5/26 3 445/30 4 &«& 6/1 PA (aye (3) yl) fs)
Pc o/23 09 0/26 3 5/30) 4 £4£6/1 DOV 6 6/14 7
memos 5/23 5 5/al 8 6/2 2 @66/5 3 6/13 8 6/21 8
Oemosis 5/23. 5 5/26 3 5/7380 4 6/1 2 6/7 6 6/14 7
fmemoyls. 5/23 5 5/26 - 3 5/30 4 6/1 2 6/8 7 6/15 8
SOAS 5/23 © 8/28 5 989/31 3 £42673 3 6/9 6 6/16 ff
Average 5 3.87 3.63 2.25 6.5 7.5

Feeding Habits.—The larvae immediately after hatching con-
sume one or two aphids per day, but this number rapidly in-
creases until near the end of the larval stage, when fifty or more
may be eaten each day. There is no appreciable diminution in
the number eaten upon days when moulting takes place, inas-
much as this requires only a short time, and the larvae immedi-
ately afterward resume feeding very actively. Throughout the
experiment the hop aphis (Phorodon humuli) was used, with the
exception of May 16-17, when the small rose aphis (Macro-
siphum rosae) was substituted. The total number of aphids eaten
during the stage ranged from 232 to 487, with an average of 349
for twelve specimens. The daily individual averages varied from
17.2 to 24.2 aphids per day, with a general average of 20.7 aphids.
These data were compiled upon the basis of the number of days
intervening between hatching and pupation, even though in
nearly every case the larvae remained inactive and without food
for one or more days preceding the casting of the fourth larval
skin. All individuals were full size at the time of emergence with
the exception of No. 6, which, strangely enough, consumed the
largest number of aphids during the larval period.

Unwersity of California Publications.

LARVAL FEEDING RECORDS—Hippodamia convergens Guer.

268

Date it
Maye ds ee
24 2

25 3

26 4*

27 9

28 5

29 3

30 10*

Bill 24

June 1 20*
2 45

3 44

4 33

5 41

6 > x

7 P
Situs
Oe:

NOU ee

lil ere

ee are
Sie

14 E

Say pee

NG es

1 (pees
Sees
Total 243
Average 18.6

*

ow on pp bv

bo bo
a"

~S)
ese
PS

Wf

23.0

Sacramento, 1913

3 4 5 6
1 3 2 0

2 4 6 2
(hei 8* oye 4

8 9 8 a

5 2 10 3

5 2 4 2
(ee 8* 8 3
21 25 eet aes
mabe UN aay 16
33 41 35 24*
40 50 26* 35
46 52 58 46
33 49 55 37*
foe eee 51 31
P Pp 46 57
pescvet a eaakts 39 61
ig Jhao, § @ pce ape 54
Bere eee 1 32
Bee Sees sees 29
ed Gee, teeth 32
AER ee RD ee Ve
eee | ree eee
| Digna | re ae grees
oak apenas 1 ON Bere
se OES) ES E
232 272 386 487
17.8 20.9 24.1 25.6

ai

8 9 10
3 i 0
4 3 3
8 5* 4*
5* 6 a
2 5 a
il 4 4
Ue 6* (He
23 21 24
21 225 a
36 43 26
37 42 29
28 24 47
36 53 52
39 42 41
42 BY peer
48 37 iP
57 A eee
17 OM fh Meech
WSs cites «ee
39 Di eee
2A) Tc eee
Des
sedis BP ees E
490 4388 264
23.3 24.3 20.3

| ENTOMOLOGY

The adult feeding records were taken upon the basis of pairs
from storage May 2, 1914, and were continued till the death of
the individuals. In every ease the death of the male, the date of
which is indicated in the table by an asterisk, took place within
two weeks after the beginning of the test, while the females
lived considerably longer.
eight-day period from May 15 to May 23 will be taken, using
specimens Nos. 1, 2, 4, 5, 6, and 7. The maximum number for

the period was 515, the minimum 380, and the average 449 aphids.
The daily averages ranged from 46.6 to 64.2, giving a general
group average of 56.1 aphids per day.

For use as a basis of comparison the

Vou. 1] Clausen.—Life-Histories of California Coccinellidae.

ADULT FEEDING RECoRDS—Hippodamia convergens Guer.

Sacramento, 1913

Pairs of Adults from Cold Storage, May 2, 1913

Date
May 8
9
10
ial

June 1

Total

9°

50
65
31
32

1033

3

50
56
44

972

5
50

5d
40

2583

8
50
28
45
38
50
48
58
65*
42
48
43
50

824

9
50
32
51
33
39
50
60
Com
65
67
46
58
36

2783

10
50
30

ial
50
21

»)

a

69

270 University of California Publications. [ENToMoLocy

Hippodamia ambigua Lec.

The same general considerations may be applied for this
species as were given for H. convergens. The elytra, being red
and devoid of marking, easily serve to distinguish this species.
So far as known no detailed breeding has been carried on to de-
termine the exact relationship of these two forms.

Oviposition.—Egeg-laying records were secured for eight in-
dividuals, with results corresponding quite closely to those
secured with H. convergens. The maximum number secured in
one day was 24 and for the entire period 421, with a minimum of
199 and a general average of 312 eggs. The daily individual
averages ranged from 6.5 to 8.4, with a general average of 6.3
eggs per day. The total length of the egg-laying period ranged
from 25 to 59 days, averaging 48.1. The proportion of days upon
which eggs were deposited was 61.4 per cent, the variation being
from 48.0 to 25.0 per cent.

Ea@a-LayIng REcorDsS—Hippodamia ambigua Lee.
Berkeley, 1914

Date 1 2 3 4 5 6 7 8

Mareh.25) 322; 10 een ye Ae eeneenan YESS me Gs ho
26) yee 6" BR ida ee, > ea ee ee

27 BL a eee Bee eee TOM poke eee

PR = tea end eee Wil Ae) oan § eee +

29 a HAY cess. PAN Sener 140) aay 385

3) nee 5 ON ese Tyee eR ce 5

Sit eee Nolte Oige cee- ee 7 tal
April 1 6 3 13 5 14 Cire ee 18
2 On Mkcoee (0) iy eee 8 18 1B? ee
ope A) = eee ial Oh tee eee 21

4 ital 5 4 Beets 4 Sy eres

5 ile 2A” 2288. 8 15 16 9 15

GY es ae 12 PA 18 lt? yee 20

U 5 16 8 Gi qaee2 19 11 18

8 1 foe oe Oe 3 22 Orr VE

9 12 18 IG pw Seectas 16 8 21 i

10 16 Aas ees 1 4 qal 16 14

11 8 14 Bi Qe: Be 4 aig) eee

12 22 Ope 24 12 NG tee 19

38 Eee 1) ee a eS p22 ee 14 ee

14 5 3 Ou ee eee 13 6.4 Yaa 26

5 TE ee Seas Di os 0 Bee

6 A eae 16 U2 eee 9 Sie Pee

Vou. 1] Clausen.—Life-Histories of California Coccinellidac.

Eec-LAaying REcorps—Hippodamia ambigua Lee.—(Continued)
Berkeley, 1914

Date
April

May

18
19

1 2
4 13
ee 6
aS} eee
ees 5)
Sig kee
6 16
int eee
14 8
DA Wn yes ie2
Gy eee
OS Pease
aes D
Sy sees
oa
Sie ese
Bh ees
A ey eee
Does
311 236
6.7 6.3

atl

3

z e

a ea ae a2 eee oa

Za Ars ca] =h7) a =e eS) Sn or =e +> Ha A Ht
1 4/2 4/8 6 4/11 A VIGs be 4/19) 3) 4726 (- weAE
2 4/2 4/8 Gay 247213 § 4/17 4 4/20 3 4/28 8 4/7 9
3 4/2 4/8 6 4/11 3 AVAIG) 55 4/20 4 4/27 7 5/5 8
4 4/2 4/8 6 4/12 4 4/18 6 4/22 4 4/29 7 5/6 U
5 4/2 4/8 6 4/11 5) 4/15 4 4/18 3 4/26 8 5/3 7
6 4/2 4/8 6 4/120 45 4/7 5 472 4 4/2 6 5/5 8
@ 4/2 4/8 6 4/11 3 4/15 4 4/19 | 4/25 6 BY/3} 8
8 4/2 4/8 6 4/13 Dye ASI on ee yall 3 4/29 8 5/%f 8
9 4/2 4/8 G42 AS A G4 4720) a 429 Oo 8
10 4/2 4/8 Gr 4/2 4 4/17 5 4/21 4 4/28 gaye 9
Average 6 3.8 4.7 3.6 7.3 8

212 University of California Publications. [ENToMoLocy

Iife-History.—The life-cycle of Hippodamia ambigua from
egg to adult at Berkeley, California, during April and May was
33.2 days. The various stages required the following periods of
time: egg stage, 6 days; first larval, 3.8 days; second larval, 4.7
days; third larval, 3.6 days; fourth larval, 7.8 days; and the
pupal stage, 8.0 days. The variations within these periods were:
first larval, three to five days; second larval, four to five days;
third larval, three to four days; fourth larval, six to nine days,
and in the pupal state, seven to nine days.

LirE-Htstory—Hippodamia ambigua Lee.
Berkeley, 1913

Feeding Habits——The eight individuals used in the larval
feeding experiment were the same as those from which the life-
history records were secured. During the period extending over
approximately 28 days a maximum of 396 and a minimum of
269 aphids were eaten, giving an average of 312 for the period;
the daily average extended from 10.3 to 14.8, with a general
average of 11.4 aphids per day. Of particular interest is the
comparatively low number consumed during the first eight or
nine days, and even after this the number was very low, 37
aphids being the largest number eaten by an individual in one
day. The temperature during the early part of the test was

ew w ow co Total days
Hm Co Ol LO of stages

31

Vou. 1] Clausen.—Life-Histories of California Coccinellidae. 273

rather low, the daily maximum being from 65° to 75° Fahrenheit.
For feeding purposes rather large rose aphids were used through-
out, these being somewhat larger than the hop aphid used in the
feeding tests at Sacramento, California. In the adult feeding
records for ten individuals the period total varied from 370 to
429, with an average of 397 aphids. This represents a daily con-
sumption of 24 to 28 aphids, with a general average of 26.5 per
day. The largest number eaten by an individual in one day
was 42.

LARVAL FEEDING RECORDS—Hippodamia ambigua Lee.
Berkeley, 1913

Date 1 2 3 4 5 6 7 8

April 8 H H H H H H H H
9 1 2 0 0 1 3 2 1
10 2 1 1 3 3 2 il 0
11 2 2 0 2 2 3 2 2
12 3 0 2 3 4 3 5 6
115} 4 3 4 6 2 6 2 5
14 5 5 6 3 3 5 + 9
15 7 7 9 5 6 9 ) 8
16 6 8 12 9g 8 6 13 10
17 5 11 10 8 15 6 8 6
18 12 10 14 10 10 9 10 of
19 8 12 16 8 9 13 11 13
20 13 101 9 13 13 15 15 10
21 10 14 7 16 12 14 8 18
22 16 21 15 12 Iliff 12 16 5
23 15 15 22 10 15 19 14 8
24 12 12 12 15 8 15 13 12
25 5 2 10 18 11 10 19 16
26 ii U7 13 24 16 8 12 10
27 13 15 16 27 15 14 7 13
28 18 23 if) 12 26 ue 15 9
29 15 20 15 17 2 23 18 15

30 26 30 27 15 18 26 23 19

May 1 32 16 15 2§ 22 18 16 24
2 30 12 18 22 27 24 27 21
Beeps 37 24 15 12 16 le 17
4 12 28 16 Se ae: OI eee 19
Syn ee 24 12 BY haere Pige sss 2
OGi ee OR lee, pe Gp es 10
oo ee eee ence) ettensa) We seed d tees 12

Total 271 392 312 324 283 317 270 320

Average TOL cL alee alee alntasy ale Naleey alale!

274 University of California Publications, |ENTOMOLOWY

ADULT FEEDING RECORDS—Hippodamia ambigua Lee.
Berkeley, 1914

Date 1 2 3 4 5 6 1 8 9 10
Aug. 25 KS) 26 23 28 16 22 30 ig) 21 25
26 28 29 27 23 3 27 29 22 26 27

27 24 20 18 29 24 31 33 8 23 20

28 35 27 Bs) 26 29 20 37 26 19 23

29 27 29 20 37 21 26 24 29 24 18

30 Ue dl 22 33 26 29 20 24 30 27

3 2: 26 19 29 34 24 26 18 21 24
Sept. 1 2 38 33 42 3 18 23 20 27 33
2 21 25 oT 28 2: 30 19 36 29 17

3 39 27 2 16 29 27 27 23 26 28

4 30 17 26 3 18 26 35 15 30 29

5 22 32 22 24 20 3 29 27 16 30

6 28 37 3 28 3 18 27 BS) 28 39

7 dl 25 31 22 26 18) 32 32 21 27

8 19 28 24 26 ) 2: 38 26 29 34

Total 391 417 393 424 399 371 429 370 370 404

Average 26.1 27.8 26.2 28.3 26.6 24.7 28.6 24.7 24.7 26.9

Olla abdominalis Say

This species is one of the most important aphid eaters in
California, and is especially abundant in the southern part of
the state. Feeding has been observed upon the hop, rose, melon,
and eabbage aphids, but a very decided preference is usually
shown for the walnut aphis (Chromaphis juglandicola). The
adults hibernate singly under fragments of bark, dead leaves,
ete., and emerge in the spring somewhat later than Hippodamia
convergens.

Oviposition.—The time intervening between emergence and
mating was found to average 1.7 days, with a range of one to
three days. From mating to egg-laying, the time varied from
seven to ten days, with an average of 8.6 days for the nine indi-
viduals under observation.

The period of oviposition ranged from 19 to 47 days, with an
average of 34.7 days. The proportion of days upon which ovi-
position occurred was 73.8 per cent as a maximum and 66 per
cent as a minimum, and averaged 70.3 per cent. The maximum
number of eggs deposited in one day was 25 and for the full
period of oviposition 298, with averages of 6.3 and 294 respect-

Vou. 1] Clausen.—Life-Histories of California Coccinellidae. 275

ively for the nine individuals. The minimum number deposited
by one female was 130 eggs.

EGe-LAyineé REcorpS—Olla abdominalis Say
Berkeley, 1914

Date 1 2 3 4 5 6 7 8 9
Mia Chip2 A tmeere Meese ee Sh 6 ee. 1 OG epee peer ieee
cess 1D), | ee SR ee E A Cpa ahs oar
260) Pas M 1D eee Mig gee’ ges at ee
27 ee eee esse M M M q
28 IMIS eet M Vi ree he ee ng ee M
ee ee ore ee.) Oe Oo
3 re a ed, Poe Roy tt) Me le ee ee oe
BL eee, een BU RIE eee ee ee ge Pe
PASO Teil] tReet MP ye) efecewsl cesses iC rceeseh, © adie) —(heease A coteesl) sede
ene ne se ee) pee eek bee
See eee Re ee eet eens 0 ees
A Re en reg et, Soe ee ee eee
Bh reece Pow «Bre Oy pee Die ane Bee eee
6 4 Ae: 7 5 ee B 10
7 5 11 4 9 Oe 12 5 5
8 GIS eee 2 re Rees eee Sis
9 16 16 id eee 2 eee 1G waa lal
EO Riga ee 2s 5 9 Hal ii 6 10 17 4
il 14 13 a) lo © sete 10 4 10 16
12 12 10 14 14 11) = 9 10
13 See eee 12 7 OP) eee 6
14 Gyre: 6 & 8 16 5 iO Sepaces:
15 10 ANA csi 12 i eee 4 5
Gy Azeetee 13 TD y aiceete se 9 14 8 6 9
17 8 6 14 16 5 12 Gy eco eee
18 3 Dey ot: NO) cease 7 8 14
19 5 12 7 i Saee Ee 25 TEI wi ess eee
20 2 ee 15 7 6 18 14 11 5
21 6 5) ieee ee, [Aes 15 Ot pees 12 16
pay Noakes Oe = 5 17 ial 9 12
23 11 7 8 a [0 ) gers 20e 8 i eg eI) re
ny = Se 5 11 16 21 2 3 14 §
Sy ieee Btu ie eee 8 4) ee at ie aan
26 2 4 4 See See 3 Thaiposs. 15
27 Tp. geeks i} 4 iS aaa D 6 7
OSE pee: 9 NO a ee 12 Gig) aes 18 6
29 Sig =a 6 7 5 PAD ae 15 9
30 12 lal ily Qos NGS Peet See Tees
May 1 9 See Se 13 8 Dae 12 8
2 14 12 AN Ne et 16 O)hj Peep ce aa

276 University of California Publications, [ENTomMoLocy
Eee-LAyING REcorpDS—Olla abdominalis Say—(Continued)
Berkeley, 1914
Date 1 2 3 4 5 6 a 8 9
4 6 8 5 19 4 a (ee See 10
Oy ee ee 3 iD ae 12) ee 3 18
Gi bes A ee 4 D SP cee, ee ee
7 D 14 1 Gm 22s) af aes ate u 9
Sy er a ee eee GO ae 9 6
aah: Sec D ON rea Ae 11 7,
DK 0) ee ee A ee ee eee Se ot
LL gO ae ee eee Siueeee=: i 10 8
2a OD Ae ee ee i ee
Oa ee cae eee IB ee eee dg ee 2 4
A ae ee See 1D) eee Di Reet eee
W559 eee Sy Gee peel an, ETERS ook eee D 2
GS Leen Seared geese me oe Tee Gee eee
Ih eee 1D ac ae era) Beer 16. Le eee
eee Rene et Meer re ee ee D
Qs Oe ee eetee Eee: Ae ene OAS eee
PA | Sr ee srens, Maen Reet ee See MS Teor pear hes
Dil tere hee ie Cio gee ate O aeee ig eee ee
Doe ely, eae aie Apres a ee | ee pet!
28 eer Se a Se ee eee ee St ath
fe ee Ree reo | D a Re ar ee

Total 218 288 188 264 225 298 130 236 256
Average Ud) GS Bs: 6.7 7.5 6:0 §6:5 G0) (6:0

Iife-History.—Eight of the twelve eggs used in the experi-
ment hatched within four days, while the remaining four re-
quired five days, even though the temperature conditions were
practically identical throughout the ten days during which the
three clusters of eggs were incubating. The first larval period
required 3.3 days, the second 2.3 days; the third 2.8 days; the
fourth 4.7 days; and the pupal stage 3.5 days; giving a total of
21 days from egg to adult. The variations in time within the
periods were, first larval, three to four days; second larval, two
to four days; third, two to three days; fourth, four to six days;
and in the pupal period, three to four days. The range in the
total life-cycle was from twenty to twenty-three days, showing
a rather unusual uniformity.

A 2
E a:
ie 6/18
pe 6/18
3 ©6718
4 6/18
pe O/ 18
6 6/18
7 6/25
8 6/25
9 6/25
10 6/25
1 6/24
12 6/24
Average
days.
Date 1
June 22 H
23 2
24 1
25 fe
26 4
oe. 12*
Aye wales
29 14
30 23*

Vou. 1] Clausen.—Life-Histories of California Coccinellidae.

a

w a Eggs hatched

for)

Length of
stage, days

LEAMA E RPE PDP

Lirre-History—Olla abdominalis Say

1st moult

Sacramento, 1913

Length of
stage, days

> OD
2d moult
on

er)
Ko)

>) oO
IS SS a
bo bw bd bw bo
© eo}

ND
Ww DS
bo
oo

oO

Length of
stage, days

pHonwmonwmwnw Ww Ww Ww Ww Pk bh bo

|
|

bo
ao

PAO
a

n => n Aa

Ee = SP og sone
scee ao) & se

ge cee aes 80

ss s2 82 8 gs

3 7/6 4 7/10 4

50 Were oe W983

3° 7/6 95 7/9 3

Sa) 7 /5e 7 / Se 08

3 97/6 5 7/10) 4

7/6 - 16) 7/9) 3

2 T/12) 45 TG 4

S07.) 4 eT /1G 04

2 7/12 4. 7/6 4

2 7/12 5 7/16 4

7/6 89 W/il 85 Wie 3

7/6) is n/N)
2.8 4.7 3.5

Feeding Records.—For the larval records three distinct series
of experiments comprising thirty individuals were carried on,

and of these fourteen finally reached maturity.

The range in

total number of aphids eaten extended from 196 to 266, with an
average of 240, which, when reduced to a daily basis, represents
19.8 aphids per day for the larval period of approximately 17

ately after the second moult.

A marked increase in the feeding was observed immedi-

Larval FEEDING ReEcorpsS—Olla abdominalis Say

w

owt e

*

14
ial
Ne
18

Sacramento, 1913

u

8 9 10 11 12 13
SPIRE Dew cee fie ge Nese eee
See EEN eae yeeewe
Be oe SRO ie ee, eR ee
A I Ae 2 aS es A Lage
het Se ae ee ey ener Sree
ALG Be yee ese eect ces ae eal ee
13 it aE Serer: ee ree
20* 0 1S ees eee, ate
29 3 2 H H H

bo bo bo bo BS bo bo be bo Po bo ty Total days

bo
KF |SSSSASESLSRSSA Sotstages

Total

Average

278 University of California Publications.

LARVAL FEEDING REcorDS—Olla abdominalis Say—(Continued)
Sacramento, 1913

| ENTOMOLOGY

if 2 3 4 5 6 7 8 9 10 fal 12 13
27 24" 32-3 Pallis 32) XO) 36* 4* 6* 3 2 3
19 36 28 33* 37 28 17 19 9 8 + 5 +
41 31 39 47 26 37 ol 25 yr 1a (he (hs 5*
46 43 47 43 39 41 46 39 14 20 12 9 1
37 0 21 29 le 30 39 42 27 Passel | rails Als) 21*
12 le 12 Pee Je IE 12 36* 18 29 23* 16
ers, Web.gae ieee 2 Rhee, reek a Sei eeomeeees Se ee 29 31 NG 36 29
ecg ee Pea Querees BOUL eel ges aes 42 37 38" 928" om
aa E 1 er eer ees JO eee 49 40 47 39 38
IH lee feeacn Sysco 1 ioe ae E 30 32 3 46 41
Sg a ere ee) eens, | (oeeeaa seve acre Te’ IE 41 33 49
ee aN we yeh ira eA te AS oe Pee e hliied © eee le IE Ie
Picts, ogee SSW plo. [Geese Oe eee. pene E De 2
Dyret by Nhe AAD ass eae, Reet eeeeeay, Ris ages ae E E E
250 196 240 263 205 259 238 256 254 231 258 246 240
19:2 15.0 18:6 20.2) 17.0 19:9 18:3 19:6 21 192! 23:4 22:3) 2s

In the adult feeding records, complete data were secured for

nine individuals, six of which were females. For

the entire

fifteen-day period the number of aphids consumed daily ranged

from 16 to 47, with an average of 30.4 aphids per
period totals varied from 407 to 534, averaging 457
the entire series.

Medium-sized rose aphids (Macrostphum rosae)
for feeding purposes throughout this experiment.

ADULT FEEDING RECORDS—Olla abdominalis Say

Uplands, 1914

Date 1 2 3 4 5 6 7
June 8 26 37 22 40 28 19 30
9 18 25 28 Bo) 23 Atl 21
10 35 27 25 32 29 36 27
11 PAL 40 36 18 Bie 28 34
12 36 36 45 20 26 22 19
13 32 29 32 26 33 29 15
14 41 oi Pl 32 46 18 26
15 28 16 26 Bf 19 37 29

day. The
aphids for

were used

25 32
28 26
33 4]
31 2

26 19
42 22
37 40
53 ays)

Vou. 1] Clausen.—Life-Histories of California Coccinellidae. 279

ADULT FEEDING RECORDS—Olla abdominalis Say— (Continued)
Uplands, 1914
Date 1 2 3 4 5 6 7 8 9
June 16 32 25 40 24 17 26 40 60 43
iy 21 38 29 29 28 33 28 27 3
18 37 47 32 38 26 38 26 19 22

19 16 28 21 32 29 17 34 38 29
20 35 33 37 41 37 20 37 35 43
21 39 3 39 2 34 29 23 41 Ue

22 41 34 28 35 22 33 18 39 28

Total 464 472 461 466 434 412 407 534 460
Average 30.9 31.4 30.7 31.0 28.9 27.4 27.0 35.6 30.6

Olla oculata Fabr.

This species is found generally distributed throughout the
state and is particularly abundant in the southern section. Its
constant association with O. abdominalis gives considerable
strength to the belief that it is not a true species, but merely a
varietal form of 0. abdominalis Say. Other notes given in refer-
ence to O. abdominalis may be considered as applying to this
form as well. A rather curious circumstance is that in a number
of individuals of this species, at the time of emergence from the
pupal skins, the elytra, which have a black ground color with an
irregular dark red spot near the center, show faintly the mark-
ings of O. abdominalis, the ground-color of which is gray, with
from six to nine small black spots on each wing-cover. These,
however, are soon obscured by the rapidly developing black
ground-color as it eventually exists.

Oviposition—Complete records for the entire period of egg-
laying were secured from nine individuals. The time from mat-
ing to oviposition varied from eight to thirteen days and aver-
aged 10.7 days. The number of eggs deposited averaged 347, the
greatest number secured from any species. The maximum num-
ber was 489 and the minimum 171 eggs. The highest daily record
was 22, and the average for all individuals for the entire period
was 9.8 eggs per day.

The period of oviposition in the nine females under observa-
tion ranged from 17 to 46 days, with an average of 35. During
these periods the proportion of days upon which eggs were de-

280 University of California Publications. |ENToMoLocy

posited varied from 84.3 to 95.5 per cent, averaging 89.3 per
cent for all individuals, which may be said to be a decidedly
unusual condition.

Ecec-LAYING REcorRDS—Olla oculata Fabr.
Riverside, 1914

Date 1 2 3 4 5 6 vf 8 9
May 26 Mey 8 M WC? eee MEPs M M
PA fae Wi ence Ms ES 2 ene IM = (2.ces eee
280 «sacar Mie Git, eS ee eee
290" wach Ye ae Rly we eee Sse a kee Jl one ee
SO ee eles) Dees ghee 5 Rites By «ete ae ee
BU <ehiz— EE” De Gu scce aete eR eee
JUN): Who’ « geese kee SE a Tees a
2 Wile OR ee Ed eee ee
Be ee Recon ee eee eR Ss Se ce
ee Bieta eee Pe
5 Du) heutige TY, neem ee gms ec ne 10
6 2g eek eee Dit cress 11 ew preeee 8
of i= Gone aaa 13 2 7 14
8 6. ee 2 12 5 5 Of 2 5
9 Dg. Se se aa 6 3 12 5 0 11
10 21 13 16 8 7 9 14 10 16
11 vs 8 14 14 6 10 9 18 7
12 8 lf 10 5 4 WG} ae 5 12
13 14 16 13 13 12 sl) eee 10 6
ne oes fers 5 9 21 Gaweeees 6 9 17
15 19 20 {haar 21 3) 5 8 9
16 12 6 12 19 14 9 14 nef 21
iy 11 14 18 13 ial 20 12 5 13
If} Anes 8 11 8 ii 20 afl ile
19 20 5 22 PA ek 16 i7/ 9 10
20 U 2 19 15 19 13 14 16
21 6 12 10 6 9 12 11 8 12
222 7 18 5 13 8 14 16 5
23 12 12 16 9 17 5 6 20h) ee
24 13 21 15 20 6 11 8 D 14
25 5 iY 9 12 8 14 oe eS 22
26 15 14 3 14 12 7 (EN aeeeses 6
OT” ete” “lect eR eee
28 8 Diy aceeees 8 16 10 Se
29 12 3 PA Geeeee 19 14 1G pee eee 13
30 13 21 ili 4 1183 12 SY eae 17
July 1 10 iil 14 9 5 jul LG? ees 12
2 aes 18 3 20 yy Silas 14
3 5 8 10 1 14 19 adage 22
4 11 20 a 1: eee 8 13 143oe oe 16

© of stages

Vou. 1] Clausen.—Life-Histories of California Coccinellidae. 281
Eaec-LaYING Recorps—Olla oculata Fabr.— (Continued)
Riverside, 1914
Date 1 2 3 4 5 6 7 8 9

5 C 18 20 De see: 8 NZ beret 2

6 8 15 Dc eee 13 22 GS ie er 12

7 2 16 Bh oe eeeere 19 12 LO 6

Sh geese 9 i 7 19 Gy ees 8

9 D 4 ify eee al 3 eee 5

TOT corse ere Wey gare 12 UG Si) ee 3

lk Pee 3 D4 Mere 9) 9 Pee Te 9

ep ee ee See 3 18 hig See Res

3s es D NOH « yee reese fer Peete on cas ae 1

Nees ae Roe PAD), Sees D 12 Dy 3

iby* see) Wee a tems ay eae Se RE ie Se ees 2

Gen Ue, “ees eres ob Pagar sg Baten oe i

ILCs ae oe aces Weer GMP esa, Byes Se protons Ae ee D

Tcl = cae a es {Shi al ex sheen De see Salen Seg err gee

Uo eo earn A re) Gas iS ene re. Melee may ZAP a

PAD) 1 eee eaneneee 1D yteeees eee mee 2 Oe eee emer es rs

i eee ee a ee ee Ee Ne petty 2 Vege) Beas

0 ESE PEE OR a 1 Benge corey Wo, Remon

Total 335 38438 460 276 342 489 313 171 = 406

Average 98 10.3 10.9 8.9 9.5 10.6 8.2 10.0 9.6
Lire-History—Olla oculata Fabr.
Riverside, 1914
E
n E oD ou a Sao cena peaiee Ba
eee 2 = ae 2 G2 4 Ge 22 32 a Ss
o D tod g Eo a boo I be 8a, too A bo
C} ee et ag +« ee a Bes os os 48 6s 8 Sa
a a) isa) = = Hu a Ha rons =P +o 0h ca] Ha
jm5/26 6/1 5 6/5 4 6/9 4 G/13 4 6/20° 7 6/25 5
2 C/A nG/ tO 6/10 8 SG /1e Ane 1672) 936/262 5
BmovZom6/l 95 6/6 5 6/10 4 G6/l4 4 46/21 7 6/26 5
4 C/A G/b) 4) 1679 A GAOT 6/20) ee O/ Zon ro
5 GA Bh GAS 2b GAL Ge GAl “ay Gye (y/ 6
6 2 GAL Gypsy 48 — XyAlO) Say GyAIS © eb aye aneGy Ziaur
ino/209 6/1 95 6/6 5 6/11 5 6/14 3 46/21 ff WR | &
Bo /26 6/1 5 6/5 + 6/9 + 6/13 4 6/21 8 6/26 5
9 : CAR oO; Oe (6/11 416/16) 5) 6/23 7 6/28 5
10 C/o O/ Ga ero 6/10") 45> 16/14 45 16/20) 6 6/2 5
6/1 5 6/5 4 46/9 4 6/14 5 6/22 8 6/27 5
5.0 4.6 4.3 4.2 7.0 5.1

oS

9 ww ow po Total days

282 University of California Publications. |ENtomMoLocy

Life-History.—All individuals used in the life-history experi-
ments were secured from a single cluster of eggs deposited May
26. These hatched five days later, and reached the adult stage in
approximately twenty-six days, the larval and pupal stages re-
quiring twenty-one and five days, respectively. The first larval
stage required from four to six days, the second three to six days,
the third three to five days, the fourth six to eight days, and
the pupal period a constant five days with a single exception,
which required six days. No striking deviations from the normal
can be noted here, and the range of from twenty-nine to thirty-
two days in the total period represents only a comparatively
slight variation for a life-eyele of that length.

Feeding Records.—The larval feeding records were secured
from ten individuals fed throughout the period upon medium-
sized rose aphids. The number eaten varied from 299 to 349 for
the period of approximately 25 days, with an average of 326
aphids. The daily average for all specimens was 17.2, repre-
senting a range of from 15.3 to 19.3 aphids per day. Feeding
was continuous up to the day of pupation.

The adult individual records for the fifteen-day period varied
from 589 to 672, and averaged 624, which is very uniform as
compared with the variation secured in experiments with other
species. The daily records as derived from this ranged from
39.2 to 44.8, with an average of 41.6 aphids per day. The largest
number eaten in one day by a single adult was 77 aphids.

Medium-sized rose aphids (Macrosiphum rosae) were used
exclusively for feeding purposes in this experiment.

LARVAL FEEDING REcoRDS—Olla oculata Fabr.
Riverside, 1914

Date 1 2 3 4 5 6 0 8 9 10
June 1 jal lal H H H lal H. H H H
2 2 1 1 1 1 3 1 2 1 0

3 Yr 2 1 1 2 1 i 1 2 2

4 4 5 3 6 5 4 4 3 iy 6

5 4 6 5 Uf 6 6 5 5 4 7

6 9 7 5 13 10 8 11 7 6 8

7 13 10 12 14 12 10 8 1s 9 12

8 17 16 8 12 10 14 ital 19 15s 5

9 19 ii 14 20 15 9 16 23 18 13

10 16 15 uly 14 21 19 17 15 22 18

Vout. 1] Clausen.—Life-Histories of California Coccinellidae. 283

Date
June 11
12
i153
14
15
16
We
18
19
20
21
22
23
24
25
27

28
Total

Average

bo po
an

op 0 bo
oo

CSoaonN QD 8 Pp

Total

Average

LARVAL FEEDING RECORDS—Olla oculata Fabr.— (Continued)
Riverside, 1914

1 2 4 5 6 7 8 9 10
23 18 22 16 24 14 25 21 19 22
27 25 19 28 25 18 23 27 26 20
21 20 23 18 22 23 3 22 19 25
Bil 24 26 19 23 27 18 26 22 30
ais} 24 29 26 30 16 20 22 12 19
21 28 19 22 35 2 25 23 18 24
7 24 28 24 40 29 18 oil 29 23
29 28 32 27 31 37 25 33 19 29
7 ae 41 30 S37 19 2 30 22 36
12 35 32 12 12 26 Bil 18 12 12
weed 1D IP IPT Nieto 30 P 12 29 le ee
ES ey ars ee ee 12 fsa! ee 14 ase
ep er ND 3 ge AEN). 0S de ts legen ee
[Dis AR eee E gies cg i Sey Uae ey Mie ee ean kone
oe E DD ey Sa a aren Mae kek ge DH eee ae ear
tele, git ri iece sii tice vane fe ne E HPS pete BE E
ys U5) 338 338 310 349 333 315 341 323 299

(fs alae aller ales} aI) as NaN 3a

ADULT FEEDING RECoRDS—Olla oculata Fabr.
Riverside, 1914

1 2 3 4 5 6 7 8 9 10
3D 41 28 33 26 44 27 31 38 47
29 24 36 28 29 23 36 29 32 39
42 SY 45 26 37 32 41 40 35 22
63 54 52 46 61 49 53 Sil 48 39
27 31 33 40 28 22 29 30 34 27
41 32 39 36 43 27 36 29 28 37
52 63 70 55 61 48 59 43 60 2
5Sf/ 48 51 50 39 44 52 37 42 38
43 37 21 39 28 36 35 43 27 29
Bil 23 29 26 35 29 40 24 32 41
22 27 26 53} 21 32 37 31 26 22,
49 52 37 41 39 40 32 46 41 37
62 50 46 61 47 54 56 49 60 48
65 ae 59 81 77 84 66 59 63 69
54 4 40 52 36 57 49 61 44 53
672 635 612 648 607 621 648 589 #4=610 ~=#&£600
44.8 423 40.8 43 40.4 414 43.2 39. 40.6 40

284 University of California Publications, [ENTOMOLOGY

Adalia bipunctata Linn.

This species is very abundant in the San Francisco Bay
Region, and is found in comparatively small numbers in other
parts of the state, though none were found by the writer in the
Sacramento Valley. During the winter the adults hibernate in
sheltered and protected places, as many as fifty having been
found in one colony under a loose strip of eucalyptus bark, this
being a location much favored by this particular species. Al] of
the common aphis species are preyed upon by this beetle and, so
far as observed, no particular one can be termed its favorite
food.

Oviposition.—The period intervening between emergence and
mating was found in ten individuals to vary from one to two
days, with an average of 1.6 days, while the time from mating to
egg-laying ranged from eight to eleven days, averaging 9.6 days.
The complete period of oviposition required from 20 to 39 days,
and during this period the proportion of days upon which eggs
were actually deposited varied from 60.0 to 76.3 per cent. The
averages were 28.2 days and 69.4 per cent, respectively. The
maximum number of eggs deposited in one day was 23, while the
average daily production by the ten females was 6.7 eggs per day.
For the entire adult period, the variation in number extended
from 94 to 269, with an average of 190 eggs for each individual.

Ecac-LAyIng Recorps—Adalia bipunctata Linn.
Berkeley, 1914

Date 1 2 3 4 5 6 fi 8 9 10

April 1 Here we Oe ad te erent E E Hee 22
Zim cee E E Mere Bi! Cece See eee See

3 M My ge eee E M M M M Y
Apes a peers I =o gi ag M

By 9 pasecd. Madre toe, Pacey eaeeees IME ee de

ee AE Seve Oe mere. en ceeces, | ste

Tid Vip seh cgi eat 1 ee Se ee ee
renin meee Mes eee ew PRO REE eh eer | Sorte Seabee

ie ee a ee ae ee nya ern WAR eed

VON eis ee eee te Beate = feces, eee ee

Ao ve Or eta eee ee rere ccc

12 7 ia he AO Pe ang Mec Ceres ge porte he gain eee

13 Se eee BS ts. My a ie a ee ia

Vou. 1] Clausen.—Life-Histories of California Coccinelliidae. 28

Eae-Layine Recorps—dAdalia bipunctata Linn.— (Continued)
Berkeley, 1914

Date 1 2 3 4 5 6 7 8 9 10
Aprill4 — ...... (ees 5 Oy) oe IPN Sees Gene eres
15 6 9 cia eres al le ree 4 Bae eres
16 1) eae 11 4 9 3 8 6 5 9
/ 8 GES * Bese 13 eee 5 elt aaa eas f
VS p sece 12 LAE Gh) #2 oe of BI
19 io} eer uN 9 8 12 OMe ies 12 4
20 14 18 ol eee uf ROM gsxcte 13 8 of
21 (ae 21 Oe Sects: 6 14 Op ees )
22 12 a Sipe esse: it Dt © oo ote 13 10
23 Be Wie 6 thle eae 21 8 4 (Ga ReAreee
24 es. 9 Wey seers 15 8 16 20S. 18
25 18 14 8 23 Y) VO) ees 7 12 15
26 NG. fetes 6 il” 14 dal Gis
27 4 16 3 15 4 Ile 2 eee 3 9 12
2c 8 NAS eects OL eke 17 Di ress: Le Ss
29 9 20 8 3 6 9 120 Aaa 13
30 Ly 7e8! (re 3 eat wee 5
May 1 7 12 3 NG gegen ee es a= 18 2 10
2 Gy Ake: Did ett 5 A Gy 4c * sees
Be ee | Reds SEE ey ee ee acer eee 13
4 12 22 D 8 1D eae 10 19 2 8
DUM rey tekaty y cake An Batis of en eS tig pees
6 WG; «te Dae 3 4 8 Oe
a 3 ene ea ee 8 1 2 = 22
8 1 es ae ee ee 5 IDR > Ose 10
eee ESS = Sea ee OP re gee PAY Sees
10 b> Deh east oe Oe Pe BAS heer Df Peso, USS
11 8 Oe | eee Reese ee AS Bese ~ Les 12 3
dee eee Gm e Res ee ic ee es D 6 1
13 a Tees ree meee a errr Ones ee 14 D
14 9 Bi) Goes PSE! cles Sie eee Dee
15 ES arora etme ee ere ee ele, SS ee ae
ICG) eee MOS, evseN9, gttinn QR ee Ale) Boe ye es Go Op
lg Gi ces Weert ase DOS feces eae ore
iS or Dn eee eee ee ORS NS eas Db ® ets
19 TG) A cera Pee ee ta heh Ul ese) phe meee pesien, Be oir
PAN, - acc Ee ne teen REE 4a ete a bees 2 | Fes
21 SI eS See Sig tein es Dye) ees. Scenes, git Lobe. ees
22 TD) RR, RS eee Cee Nie me eee eT Ow Peres. Bea 2a
DB rectal |) pee aaa eco are mee ls eee) a See see
PN ee ee et eee TDG Tee eee we Seep rescore

Totals 269 244 153 169 94 267 134 174 209 183

Averages 6.7 fell 7.2 7.6 4.7 6.5 5.1 6.4 5.9 6.

286 University of California Publications. |ENToMoLoGY

Life-History.—On the basis of seven individuals, the entire
period from egg to adult was found to be 26.7 days, with a range
of from 25 to 29 days. The different stages required approxi-
mately the following periods of time: egg stage, 5 days; first
larval, 4.6 days; second larval, 2.9 days; third larval, 3.0 days;
fourth larval, 5.6 days, and the pupal stage, 6.0 days. It will
be observed that, while the total period required by the four
specimens reared in May and three in August is practically
identical, the early brood required approximately 5.75 days for
the first larval stage and 2.25 days for the second, as compared
with 3.0 and 3.7 days, respectively, for the later brood. All indi-
viduals were full-sized at time of emergence.

Lire-History—Adalia bipunctata Linn.

Sacramento, 1913

3
i 5

= ca ea ca ra a w ae

to = Se = 6a Ss ce = oa 36 ~ oad oa

e ES nee ie ea ae, Bison eg als

g a tg es On ee eee ee ee

S Be eb aS : Sa =I oS $3 5, Bs E os

A AS i) Ha 4 wa QA AR © Ha WH Ae H He
15/20) 5/25) 5) 674 7 §6/3° —25 (6/6593) 56/10 50. 1G /N7 eG
9, 5/20 5/255 5/30) “5 6/1 2) 6/55 4 6/2 Oe 67k ee
3 5/20 5/25 5 5/30 5 6/2 3 6/4. 2 6/8 4 6/14 6
4 5/20 5/25 5 5/aill Glyn 6/25 2) Oo 3 GVAE ay GAS
He 8/25) 8/3) 5) 9/72 3 9/6 45 978) 2) OA OF 9/20 as
6 8/25 + 8/73l 5 972 2 ey 8 My YAW @ MAI) 6
88/25) 3873 San 3972 3.0/6. 4° (9/8 2. O/NSe (7 RO/2lTo

Average 5.0 4.6 2.9 3.0 5.6 6.0

Feeding Records.—The results from ten individuals through-
out the entire larval period of approximately 22 days gave the
maximum number of aphids eaten during the period as 308, the
minimum 220, and the average 252. This givessan average of
14.1 aphids per day for the entire period. It will be observed in
the table that a very sudden increase in the feeding took place
about eight days after hatching, and this was found to corre-
spond very closely to the time of the second moult, the number
of aphids eaten was approximately doubled at this time.

vo po Total days
© © of stages

bo
On

Vou. 1] Clausen.—Life-Histories of California Coccinellidae. 287

The consumption of aphids by the ten adult beetles under
observation ranged from 215 to 305, with an average of 251
aphids for the fifteen-day period. The largest number eaten in
one day was 30, while the general daily average was 16.7 aphids.

Medium-sized rose aphids (Macrosiphum rosae) were used
exclusively.

LARVAL FEEDING REcOoRDS—Adalia bipunctata Linn.
Riverside, 1914

Date 1 2 3 4 5 6 of 8 9 10
June 4 H Jal H H H H H H H H
5 1 0 0 0 i 1 0 il 11 0

6 1 1 2 0 3 2 2 1 2 1

df 3 5 3 2 + 3 4 3 5 2

8 3 4 6 4 3 5 2 6 6 4

9 5 5 G 3 6 7 4 6 7 3

10 U 6 9 6 8 4 6 7 5 5

11 6 8 3) 9 7 5 Uf 8 6 8

12 8 11 9 10 3 8 10 12 i) 11

13 19 12 13 12 6 10 14 9 11 a

14 12 15 19 13 10 17 21 16 12 15

15 18 14 19 17 12 21 20 ley 15 12

16 23 19 26 19 20 28 25 19 20 22

17 21 26 24 19 25 27 32 35 29 26

18 27 29 23 25 22 32 30 27 24 31

19 30 31 26 18 25 29 31 23 29 27

2 28 36 29 26 29 33 26 18 31 19

21 21 22 18 23 35 28 P 29 16 27

22 12 18 30 18 12 ite eeee 3: 12 qe’

PN Veneers 12 12 PAU Bae dees 26 Py seo

2 ee Pee ial tere oe Jee ees gee! gene D2 Srey ee

PAS) Sackecgl S Gecsseh | Micon Oy cece Serer cee Bose 22s
Total 233 262 268 253 225 277 234 308 240 220

Average U3 145 148 1353 82 15.3) 14.6 «(16.4 13.3" 12.9

ADULT FEEDING REcorpS—Adalia bipunctata Linn.
Berkeley, 1914

Date iL 2 3 4 5 6 7 8 9 10
April 1 16 12 18 10 6 14 18 15 Wil 16
2 12 9 14 12 15 ial 16 10 13 8

3 24 20 iTS 21 18 22 20 ef 16 ny)

4 18 12 16 15 7 11 10 14 13 7

288 University of Califorma Publications. |[ENToMoLocy

ADULT FEEDING RECoRDS—<Adalia bipunctata Linn.—(Continued)
Berkeley, 1914

Date 1 2 3 4 5 6 7 8 9 10
April 5 13 17 6 19 18 22 21 15 U7 16
6 22 18 24 23 25 21 17 12 15 14
7 27 26 21 30 27 22 8 29 25 9
8 21 18 19 16 23 20 17 26 19 Wy
9 19 14 16 12 17 8 15 11 U7 12

10 23 20 18 19 15 ‘

13 16 18 20 16 19 14 21 ) 12
14 20 7 16 12 13 10 Wi, 16 14 ug)
15 19 12 13 9 Hal 14 12 15 8 13
Total 305 246 258 256 266 247 254 240 227 215

Average 2053) LGA ie TOF ie 16:4 1629 aliG: Oa el eelae

Cycloneda sanguinea Linn.

This species, while not particularly abundant in any part of
the state, may be found quite generally distributed over the Sae-
ramento and San Joaquin valleys and in certain coastal regions.
The adults pass the winter singly in sheltered places upon trees
and shrubs. The larvae have been observed feeding quite ex-
tensively upon the woolly apple aphis (Schizoneura lanigera),
as well as the more common aphid species.

Oviposition.—The interval of time between emergence and
mating in the ten specimens under observation ranged from one
to three days, with an average of 1.9 days, while the period
elapsing between mating and oviposition varied from seven to
twelve days, averaging 10.1 days.

The period of oviposition extended from 19 to 42 days, aver-
aging 28.8 days. The proportion of days upon which oviposition
of the ten individuals actually oceurred was 73.6 per cent, with a
range of from 64.1 to 78.2 per cent of the total number of days
of the adult life following the deposition of the first eggs. Dur-
ing this period the egg-production totals varied from 121 to 318,
with a period average of 201 and a daily average of 7.0 eggs.

Vou. 1] Clausen.—Life-Histories of California Coccinellidae. 289

EeG-LAviIne Recorps—Cycloneda sanguinea Linn,
Berkeley, 1913

Date 1 2 3 4 5 6 7 8 9 10
Aug. 20 SB al ige-cees Wy eke 1) 2 E Diet eee Het 226
21 M 1D Wea eee 1D Cer sree) eee E
Ome eto dae gee: E M Mee pec M E M M
v8) epetelt Ene als 6 eee me a eee IVI ple pager ee Se eas
AD ae M Nee, Ocak, © tee f Tse NIMS ce Sect
2 ee me Ms ee ec Wr s | Bias 8s ih Bs ieee Oe. Bet,
AS)! cece. gage pie Be. 0 a et eee ne ee EO CRS
21 ee nS ee Me eo oeee. Wt Aes) is Ss Oa pos ree sce Oia
2G eee NORM ate | BENE ice, Re Mgt My Sone
PB) Ee ed eee nate ee en en I ee Ce re ee ar
SX), eetestr gt atta! vases Mere, OL ett Den eteag ce ige © bee ea re
31 5 Bs Seeabade ete Ae ae ENT nai Frets Ee ee as ee
Sept. 1 8 OMe es ete: oe terry ert ls ee hed ere Ag ee
2 13 Ota 4 Ul ene aS SM eee 5
ot ee 5 7 Se aie 3 5 7 7 3
4 1 es ee 12 14 5 9 Gv ae 14
Be ee 6 Tal 6 9 2 LS ee GbE * cake e
6 12 4 1/4 20 9 1 8 10 12
7 14 ew ee? 14 6 10 9 5
cle ae eee 6 10 11 3 14 11 18
9 20 eeele 14 3 VON gsete# gee ta Meese
10 7 23 12 7 IBY Sate e! Nero aes ii
11 26 9 2A ae sae 24 3 8 ] Sieh cee
12 3 De eee ORs S yy 2 2 5 ae 14
13 11 16 tS ae 8 5) 20 eee t 11
14 Owe pa 16 1K?) eeee 6 6 12 5
i) a eee 5 9 4 113) al oe SS eee 3 19
16 7 10 ial 8 7 I ee 19 BU peeierers
iY 1 toe ee ee 3 TOMI te OF, tonite 9 8
18 9 8 US}, oT eae 16 5 11 14 i Sea:
9 eee (he Wemeese 7 9 Is) ees 2k estes 16
20 12 15 9 6 12 NY ie eee ses 11 3
Pile soe nee 4 Dee heme oe) ae 2 AUST Vice sa
22 19 2 6 D ghee 8 3 14 5
23 6 SLA Ty eee Oe cess 21 3 12 D Sits:
DA pee Diageo Sa (aes. 2 AS yest F SSe De poe
25 5 7 Bs) uses: Same. (one pace 13 D
26 1 14 1) eee. ees 7 Se ees alee
A oaks eee | DE tae 11 6 Gas (ome eee
28 D Did eee, ee aes 14 Ae. Wee Sy Phe
29 gaa See eee ISP beer 14, Me! ye
3 ee eee en Sg fie 5 9 LOM Ses DNF aieeet
Oct ee ee ones ae ep erate 113) 5 ae ets rete 1G Aa

290 University of California Publications. [ENtTomMoLocy

Eee-Layine Recorps—Cycloneda sanguinea Linn.—(Continued)
Berkeley, 1913

Date 1 2 3 4 5 6 it 8 9 10

Otis ate 22, (Se ee 10 2 © ain ae aes ee Bt. © eke

ED ey Ia AE 9 ee nen DD gee Be eee

5 eapiee mmm, Aap eS baie. Les ieee Ore itt 119) eee

Gl een Pees ule ee aes 6 Die shee a ne

(eee ety mr eves BR ae eee Es (Pare

Sitee cS Sete sate ey Ue oo Gu Gee

Oh Re el ie renee Die (etetes oR cts: A ee eee

A Oy oe Re eee ees clip suse S.) lect inde LP ages

A eee en eee OnE Bs is ere etsy miele tad Dee

MOR F285 atcha ae eh ae Deans | Veni,

Total 220 207 173 121 318 180 198 137 308 155
Average 7.8 6.4 7.2 6.0 Urge 5.2 6.4 6.5 8.1 6.7

Iife-History—In the following table is given the results of
two series of life-history experiments, the results of which are
quite noticeable in their difference. The eggs for the first set,
of which three survived, were secured May 1 and hatched six
days later. The first larval period varied in length from five to
six days, the second from four to nine days, the third from three
to five days, the fourth from five to eight days, and the pupal
period from five to seven days. It will be noted that the varia-
tion in the second and fourth stages was very great, but the in-
verse ratio seemed to exist in the remaining stages, so that the
complete life eyeles for the three individuals varied only two
days. In the remaining eases the egg stage required only five
days, the first larval period a constant four days, the second
period two to three days, the third period two to three days, the
fourth period three to five days, and the pupal period three to
four days, giving a total of from 20 to 23 days, as compared with
33 to 35 days with those reared seven weeks previously. The
difference in mean temperatures at the time of the two series of
experiments was approximately 10° F.

Vou. 1] Clausen.—Life-Histories of California Coccinellidae.

Lire-History—Cycloneda sanguinea Linn.
Sacramento, 1913

=
Bo Ge eg poe Sek eae ic) aac ae
= a nS =a tied a) a", 6s 85
o n = bo S = wo = mo sa mo 8
SR » &s # 82 «s« 88 s« 88 £8 &s &
a QS co) Ha = Hn na a7 oO AR FW Ht cS)
iL yal 5/7 Geo /2ze oe 5/21 9) 5/26" 5) 673 8 6/10
2 5/1 5/7 @ ByAlBy (a) aye toy YEE a bay/PAY) tay C/o}
3 5/1 5/7 Ge o/137 6 bi, 2 19/2271 5/29) i 673
4 6/24 6/29 5 £47/3 4 7/5 2 7/8 3) C/A AS 7s
5 6/24 6/29 5 7/3 4 7/5 2 7/8 3 7/12 4 7/16
6 6/24 6/29 5 7/3 4 7/5 nS Se he Se Ale:
mu 6/24 6/729 5 7/3 4 7/6 3 7/9 3/3) ATA
8 6/24 6/29 5 7/8 4 7/5 ST, Pa Ola eb 7h fats)
9 6/24 6/29 5 7/3 4 7/6 3 67/8 2s) 205 0/6
10 6/24 6/29 5 7/3 4 7/5 20/8 3) YVAls} Gy Ales
Average 5.3 4.5 3.7 3.2 4.9

291

Length of pupal

stage, days

wounuwn

Pow Pp PO

Feeding Habits—Complete feeding records were secured of
ten individuals for the entire larval period, and the range in the
number of aphids eaten was from 147 to 427, with an average
of 216 aphids for the period. The daily average, computed upon

the basis of approximately twenty days, was 14.5 aphids.

The feeding tests of ten adult beetles for a period of fifteen
days gave a variation of from 205 to 260 aphids eaten, a rather
unusual uniformity. The period average was 234, and, on a
daily basis, 15.6 aphids. The daily individual average ranged

from 8 to 30, with a general average of 15.6 aphids per day.

LARVAL FEEDING RECoRDS—Cycloneda sanguinea Linn.
Sacramento, 1913

Date 1 2 3 Date a 5 6 cl 8 @)

Wie? OU se isi eis! Junie ORs Eh Ee Er ae
oat Bey tp Meper te Reese 30 2 il 1 0 2 1

) 3 3 2 July 1 2 4 3 1 3 1

10 6 8 5 2 6 i) 8 ae Pali 7
lst i il 3 3 AS 2 ee ca aelOR Oe ales

2 8 4 a 4 13 17 9 6p eel

13 7 on” NO* 5 Se aise ae IY

14 i 8 5 6 7 3 6 ne (eames

15 3 4 9 Mey zal "aks} IGS 23%) 49

16 SiS) a Bye eh RES ily PAG alls}

Wy / yf Mal oF S26 as 32 428* 34 28

18 9 6 8 10° 32)" 36°«22 3%) 3' 826

19 8 10 8 Hil ales 8 P 42 33

20 8 fold 12 Poaceae PAILS eee 27

21 9 0% ake Sea | SUR eases | a ere P

co v Total days

WwNrR WOOD FH Ww No stages

bo Ww

bo po bo po LW

a)

bo
oF

U

Tniversity of California Publications.

| ENTOMOLOGY

LARVAL FEEDING RECORDS—Cycloneda sanguinea Linn.—(Continued)

Date
May 22
23
24
25
26
27

Total

Average

Date
May 16
7,
18

b> Do ro
o ©

Cw pp

bo bo
HS

Total

Average

1 2
is; AS)
We AG)
23* 29
41 39
27* 46
23 8621
44 12
5.0 aeons
{ees
BA eecene
Sy eee
P E
ID) eee
418 258

Sacramento, 1913

3 Date 4 5 6 ui
18* 1 eee oe iy ee
155 15 THe ie eee
He 129 Geers Ee eee eee
36 gL fe eke ee) Sat EK
35 A foyer A-One Bees See
16 NO) Se, ee eee
12
E
245 149 200 147 199
2.2 2A GG) 1323) Woes:

156

14,

ADULT FEEDING RECORD—Cycloneda sanguinea Linn.

Riverside, 1914

1 2 3 4 5 6 7 8
12 UY) 15 8 14 ly wy 7
16 12 6 14 16 23 12 16
14 18 13 17 Hal 16 14 12
22 12 10 16 22 19 22 19
19 17 18 14 15 18 15 24
16 26 If/ 19 17 12 17 26

3 14 15 11 12 15 10 16
18 18 20 9 14 Hal 18 U7
26 15 11 8 10 14 24 18
15 12 16 13 19 30 16 12
19 14 19 15 15 9 21 20
1 ial 15 13 17 3 12 17
Ne 18 14 ily ial 16 13 19
14 20 12 16 14 21 Uy 14
19 16 8 15 20 12 19 23

251 242 209 205 225 246 249 =. 2.60
Gyre ileal ale)

206

1 15

8

194

14.9

Vou. 1] Clausen.—Life-Histories of California Coccinellidae. 293

COMPARATIVE RECORDS

Time from Emergence to Mating and Egg-laying—Complete
data in regard to these periods were secured for only five of the
species under investigation. In the case of Hippodamia con-
vergens and Olla oculata, records were secured only in regard
to the time from mating to egg-laying, and none whatever in
the case of Coccinella trifasciata. The periods varied only to a
comparatively slight extent between the different species. From
emergence to mating the usual intervening period was one to
three days, the averages ranging from 1.6 days in the case of
Adalia bipunctata to 2.7 days for Coccinella californica. From
mating to oviposition a corresponding variation occurred, the
shortest time being seven days for Cycloneda sanguinea and Olla
abdominalis. The general averages for the different species
varied from 8.6 for Olla oculata to 11.9 days for Coccinella
californica.

EMERGENCE TO MATING AND E@G-LAYING

Emergence to mating days Mating to egg-laying days

Number AW —, —,

specimens Max. Min. Aver. Max. Min. Aver.

Coecinella californieca...... 10 3.0 2.0 2.7 14.0 10.0 11.9
Hippodamia convergens.. 6 ae a = 13.0 8.0 10.5
lel, etal oy Mea bie eee ee 9 3.0 1.0 Ee ARO 8.0 9.6
Olla abdominalis —.......... 9 3.0 1.0 Wee 10.0 7.0 8.6
Ooculata 22 9 aes a mere 13.0 8.0 10.7
Adaha bipunetata .......... 10 2.0 1.0 1.6 11.0 8.0 9.6
Cycloneda sanguinea...... 10 3.0 1.0 ES 12.0 7.0 10.1

Periods of Oviposition—The period over which egg-laying
will extend is largely dependent upon the conditions under which
the individuals exist. Egg-laying normally takes place daily,
with an oceasional exception, approximately two weeks after
emergence until death. Inasmuch as all the experiments upon
this point were not conducted under similar conditions the re-
sults will naturally vary to a greater or less degree.

The maximum number of days included in the oviposition
period was found to be 59 in the case of Hippodamia ambigua,
Coccinella californica was second with 51, and Adalia bipunctata
last with 39 days. The average number of days for the indi-
viduals of each species was 48.1 for H. ambigua, 35.4 for Olla
oculata and last, A. bipunctata with 28.2 days to its eredit.

294 University of California Publications. [ENTomoLocy

The proportion of days upon which eggs were actually deposited
was found not to have any direct bearing upon the comparative
totals, the individuals of Olla oculata averaging 89.3 per cent
as compared with Hippodamia convergens at 63.8 per cent, even
though the final total laid was in favor of the latter by a margin
of 48 eggs. The maximum proportion of days in which eggs
were deposited by a single individual was 95.5, this also being
to the credit of O. oculata. The minimum was 28.0 per cent in
the case of H. convergens.

PERIODS OF OVIPOSITION:

Length of period, Proportion of days eggs
days deposited, per cent

Number A —,
specimens Max. Min. Aver. Max. Min. Aver.
aC UhORMIC Ape ee ee 10 51 20 31.0 74.1 60.0 69.5
trikasciatay 10 42 14 29.2 78.1 66.6 75.0
COD CLO CUS eee 11 50 iI Saee V2.7 28.0 63.8
5 NOMA eee see 8 59 25 48.1 75.0 48.2 61.4
abdominals) = 9 47 19 34.7 73.8 66.0 70.3
PEO Clilataeseee eee 9 46 117/ 35.4 95.5 84.3 89.3
bipunetatayes a 10 3 20 28.2 76.3 60.0 69.4
) SAN OUI C ayes 10 42 19 28.8 78.2 64.1 (O40

Rate of Oviposition—A very considerable difference was
found to exist in the number of eggs deposited by the various
species. As would be expected from field observations, several
individuals of Hippodamia convergens deposited a considerably
larger number of eggs than those of any other species, the maxi-
mum number secured from one female being 609 as compared
with 489 from Olla oculata, the next highest. This difference,
however, was not found to hold in regard to the general averages
of all individuals of each species, inasmuch as O. oculata ranked
first with 347, Hippodamia ambigua second with 312, and H. con-
vergens third, with an average of 299 eggs for the full period.
As regards the maximum number of eggs deposited in one day, it
will be seen that H. convergens leads with 43, Coccinella trifas-
ciata second with 31, and O. oculata last with 22 in one day.

Olla oculata ranks first in respect to the daily average for the
entire period of oviposition, with 9.8 eggs per day, while the
lowest average is 6.3 eggs per day in the ease of O. abdominalis.
General observations lead one to believe that H. convergens under
field conditions is far more prolific than any of the other species.

Vou. 1] Clausen.—Life-Histories of California Coccinellidae. 295
The record of 1550 eggs deposited within a period of slightly over

two months reported by E. K. Carnes' tends strongly to bear out
this assumption.

RATE OF OVIPOSITION

Daily records Period records

Number — ne

specimens Max. Aver. Max. Min. Aver.
(Cy GANONG) sete 10 24.0 8.0 360 171 207
CHtiritasciata 10 31.0 8.4 353 109 249
EI conver sens) ss ila] 43.0 8.9 609 94 299
Vals GON) OMS U EE, cece nes 8 24.0 7.3 421 199 312
OMabdominalis 222. 9 25.0 6.3 298 130 234
@Oeeocul attains eee 9 22.0 9.8 489 Ne fall 347
ING |Gij WUC ERIE, ceases 10 23.0 6.7 269 94 190
(Co ise nayeqvanaleyy, Soa ieee 10 26.0 7.0 318 12] 201

lafe-History.—In a comparative study of life-histories it is
essential that due allowance be made for variations in conditions
under which the experiments are carried on in ease they are not
made simultaneously. The records of Hippodanua convergens,
Coccinella californica, Olla oculata and Adalia bipunctata were
secured at Sacramento during the period from April 25 to August
20, 1913. Those of Coccinella trifasciata and Hippodamia am-
bigua were secured at Berkeley during the spring of 1914, while
the life-history of Olla oculata was determined at Uplands during
May and June, 1914. As regards climatic conditions in the three
localities, Sacramento was considerably warmer than either of
the other places, the temperature often reaching 105° F. and
oceasionally higher. At Uplands the conditions more nearly ap-
proached those best suited to the maximum development of the
beetles, the average daily maximum temperature being approxi-
mately 85° to 90° F. At Berkeley the temperature was some-
what lower and the nights very cool.

In computing the period lengths in the table given below, the
averages were taken of all individuals of each species and no
particular account made of individual variations. No very great
range was found in the egg stage, the longest period being 6.0
days for Coccinella trifasciata and the shortest 4.3 days for Olla
oculata. The first larval stage required from 3.8 to 5.7 days, the
members of the genus Coccinella apparently requiring longer

1K. K. Carnes, Monthly Bulletin, Calif. Hort. Com., Sept., 1912, p. 820.

296 University of California Publications. (ENtomoLocy

than any other. In the second larval stage the variation was
from 2.3 to 4.7 days, and no special differences are noticeable be-
tween the various genera. The third stage required a minimum of
2.3 days for Hippodamia convergens and a maximum of 4.2 days
in the case of Olla oculata. Coccinella and Hippodamia required
from 6.5 to 7.4 days for the fourth larval stage, as compared with
4.7 to 5.6 for the other genera, except for Olla oculata, which
covered 7.0 days. The pupal stage required the longest time
with Hippodamia, the average being 7.5 and 8.0 days, respect-
ively, for the two species. The variation for the remaining species
was from 3.5 to 6.0 days in the eases of O. oculata and Adalia
bipunctata, respectively.

The complete life-cycle from egg to adult varied from 21.0
days in the ease of Olla abdominalis to 33.2 days for Hippodamia
ambigua. Strangely enough, the period lengths up to and in-
eluding the third larval stage were approximately equal in all
the species, but a very considerable range was presented in the
fourth larval stage and pupal stage, resulting in the consider-
able difference between species as noted above. As stated previ-
ously, however, the possibility is very strong that this was, in
part at least, due to the varying conditions under which the in-
vestigations were conducted.

Lirr&-HISTORIES

Egg Larval stage, days Pupal

Number stage, A -—- stage, Total

specimens days First Second Third Fourth days days

| calvfornica =.= NS 5.4 Da. 3.3 3.4 6.8 4.5 29.1
tritasciatay = 12 6.0 5.3 4.2 3.0 7.4 Soll 31.8

2 COMVELS ens) s-- 8 5.0 3.9 3.6 2.3 6.5 fe) 28.8
5 TNO, eee 10 5.9 3.8 4.7 3.6 7.3 8.0 33.2
. abdominalis —.... 14 4.2 3.3 2.3 2.8 4.7 oD 21.0
seOcUl ata serene 11 5.0 4.6 4.3 4,2 7.0 Bill 30.2
= bipUMC tata 7 5.0 4.6 2.9 3.0 5.6 6.0 26.7
sanguinea ........ 10 5.3 4.5 Soff 3.2 4.9 4.2 25.3

Larval Feeding Records.—Individual feeding records were
secured for the entire larval period for the different species in
numbers ranging from eight for Hippodamia ambigua to four-
teen for Olla abdominalis. In the ease of each species, enough
tests were started to secure approximately ten complete records
making due allowance for the possible mortality during the
larval period. In the following tabulation the length of the

Vou. 1] Clausen.—Life-Histories of California Coccinellidae. 297

larval period is given to facilitate a comparison of the daily
records. The period averages ranged from 475 aphids for
Coccinella californica to 216 as a minimum for Cycloneda san-
guinea. The maximum individual record was 580 and the mini-
mum 147 for the two above-named species, respectively. Feed-
ing in the ease of Olla oculata was very regular, the difference
between the limits of the ten individuals being only 50 aphids
for the period, while a difference of 280 was noted in the case
of Cycloneda sanguinea. In regard to Coccinella californica, it
must be remembered that for a period of one to three days
previous to pupation no aphids were eaten, a condition which
considerably reduced the daily average as compared with other
species which fed normally up to the day of pupation. In spite
of this, C. californica heads the list with 24.9 aphids per day for
the entire period; H. convergens 1s second with 20.7; and JH.
ambigua last with 11.4 per day.

LARVAL FEEDING RECORDS

Period totals Averages
Length ot —"—— a #{ FF
Number period, Max., Min., Period, Daily,

specimens days aphids aphids aphids aphids
Cacalitornie@a 13 23.7 600 294 475 24.9
(Gh, (erent ISYOneN Re), Sees eee 12 25.8 365 217 294 15.8
ie comvergens) 22..2....-- 12 23.8 490 232 349 ° 20.7
Tal,, Glvtm ob) eases ee ee 8 Bel 392 269 312 11.4
OFabdominalis == 14 16.7 256 196 240 19.8
©, oat) 2a 10 25.2 349 299 326 17.2
AS bipunctata, <-.------.-- 10 21.7 308 220 252 14.1
Ch sanouimea, 2----.<--- 10 20.0 427 147 216 14.5

Adult Feeding Records——The feeding records for the adults
of the various species as given in the following table are for a
period of fifteen days with the exception of Hippodamia con-
vergens, an eight-day record of which was taken from the com-
plete adult feeding data of individuals from storage, as previ-
ously explained in the diseussion of that species. The number
of beetles of the different species ranged from seven for H. con-
vergens, and nine for Olla abdominalis to ten in the case of the
other species. The maximum period average was 624 aphids for
Olla oculata, while the minimum was 234 to the credit of Cyclo-
neda sanguinea. The individual maximum was 672 and the mini-
mum 205 aphids, respectively, for the same two species above

298 University of California Publications. |ENToMoLocy

named. On a daily basis the variation extended from 56.1 aphids
for H. convergens to 15.6 for C. sanguinea. In general it may be
said that the number of aphids eaten varied practically in direct
proportion to the size of the individuals, though a conspicuous ex-
ception to this generalization may be noted in the ease of Coc-
cinella californica, with a record of only 34.0 aphids per day.

ADULT FEEDING RECORDS

Period totals Averages
‘ Length of

Number period, Max., Min.. Period, Daily,

specimens days aphids aphids aphids aphids
Cy calitornicay 10 15 661 414 500 34.0
CARGEMEAS Claim eeeenes 10 15 470 383 435 28.9
Heiconviercens) ss 6 8 515 380 449 56.1
Vel, GYR ONSAD EY acne 10 15 429 370 397 26.5
©; abdomimnalis: =222-— 9 15 534 407 457 30.4
©. oculatay ee 10 15 672 589 624 41.6
JA bipunictatay es 10 15 355 215 251 16.7
CSisanguin en esa 10 15 260 205 23 15.6

Measurements.—In the following table is given the measure-
ments of the egg, full-grown larva, and adult of each species. In
every ease the figure given represents the average of ten indi-
viduals, and particular care was taken to secure larvae of
normal development. The variation in length of eggs was com-
paratively shght, those of Adalia bipunctata and Cycloneda
sanguinea measuring 1.3 mm., of Coccinella californica 1.5 mm.,
and of the remaining species 14mm. The larvae ranged in
size from 6.0mm. for Cycloneda sanguinea to 10.2 mm. in the
ease of Coccinella californica. This ratio also held for the
adults, C. sanguinea measuring 4.5mm. and C. californica
8.0 mm.

COMPARATIVE MEASUREMENTS

Eg Larva Adult
Ce Calitornic@aye ee 1.5 mm. 10.2 mm. 8.0 mm.
COG HOB RACONEN IED eee teeta Sects 1.4 8.0 6.5
I COMVCT.O:CTS grees eee eens 1.4 8.2 7.0
EL Oo Uae eee 1.4 8.1 7.0
@Fzabdominalises ee 1.4 fel 6.0
© 0culatawe 1.4 7.0 6.5
INS, | Ot) OWEN arene sees serececnee 1.3 600 5.0
CUP SEU OUIM C eeeceeneeereeceenesecese 1.3 6.3 4.5

VoL. 1] Clausen.—Life-Histories of California Coccinellidac. 299

SUMMARY

Some of the main points determined in the investigations as
herein outlined may be briefly stated as follows.

1. The lfe-history of an aphid-feeding coceinellid under
normal summer conditions in California, based upon the results
secured from eight species, may be given as twenty-seven days,
divided as follows: egg stage, five days; first larval stage, five
days; second, three days; third, three days; fourth, six days,
and the pupal stage five days.

2. The number of aphids eaten by the larvae of the different
species is In proportion to the size of the individuals.

3. The above, to a limited extent, may be said to be true in
the ease of the adults also.

4. Temperature and humidity are very strong contributing
factors in the development and behavior of the various species.

5. The number of eggs to be expected under normal field
conditions will vary from 200 to 500, or occasionally more, and
extending over a period of from four to eight weeks where the
female has lived the full adult hfe under optimum conditions.

6. The period intervening between emergence and mating is
one to three days, and from mating to oviposition, eight to
eleven days, thus giving a period of from ten to fifteen days after
emergence before oviposition may be expected.

7. Oviposition normally takes place daily, with occasional
exceptions.

8. Only one fertilization is necessary during the life of the
female, fertile eggs having been produced in one instanee fifty-
five days after mating.

Transmitted September 25, 1915.

‘Vol. 1, ‘No. Th PP. . 301- 346 S July 20, 1917 :

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dae, by Curtis P. Clausen. Pp. .251-299.. June, 1916 — 2.2.2
7. Aphididae of California: New Species of Aphididae and Notes from Dh
Various parts of the State, but Chiefly from the Campus of the Uni-
Te of California, Berkeley, California, by = O. Essig. Pp. 501-.

Bice Editors. Price per volume, ‘$3, 50.

Vol. 1, 1. The Distribution and. Activities of Bacteria in Soils of the Arid _ > =
: Region, by Charles B. Lipman. Pp. 1-21. October, 1912 _-.... 1... -)

2. Studies in the Phenoldisulphonic Acid Method for Determining Nitrates
in Soils, by C. B. Lipman and L.-T. Sharp. Pp. 23-37. October, 1912.

3. The Effects of Calcium and Magnesium Carbonates on Some Biological
Transformations of Nitrogen in Soils, by W. P. ee Pp. 39-49.
BECOMBSYs SOTO aN ee ar eee as, ate Gnateaattentees 3

The Aluminum Reduction Method.as Applied to the Determination of
Nitrates in ‘*Alkali’’ Soil, by Paul S. Burgess. Fp. 51-62. May,
oA es nee Ube area Oath Ny hea a eg UNE ON oA RSA RE Qiao eacerapeONCont en Ge. 1 ashy biaeae al

5. Studies Upon Infiuences Affecting the Protein Content of Wheat, by G. iz
WShaw,. Pp. 63-1262; October; 19133: a ae a ance com |

6. The Effect of Copper, Zinc, Iron and Lead Salts on Ammonification ahd é te
Nitrification in Soils, by ©. B: Lipman and P. S. Burgess. Pp. 127-
gS Papen eS We} 1 eee 8 BB hpe Yast eet asouceh ae it YANNI seca nnd SAN NCP PONS Deer, V aROES A te

7. Studies on Ammonification in Soils by Pure Cultures, by C. B. Lipman
and P. S. Burgess. Pp. 141-172. April, 1914). cece

8. Humus and Humus-Nitrogen in California Soil Columns, by R. H.
Loughridge. . Pp. 173-274. August, 1914-22... occ eecccce eect eee cence

9. New Experiments on Alkali Soil Treatment (Preliminary Report), by .
Charles B. Lipman and Leslie T. Sharp. | Pp. 275-290, plates 1-4.
DUNST AG er re a ct lp woh cre eons Lot tne apse laren

10. Fundamental Interrelationships between Certain Soluble Salts and Soil
Colloids, by L. £. Sharp. Pp. 291-339. . April, 1916 -.2....2.20... cet ;
11. The Influence of the Composition and Concentration. of the Nutrient oy
Solution on Plants Grown in Sand Cultures, by Arthur Hugo Ayres. ~~
Pp, 341-394, plates5-15; January 19L7 «2.05 ee
12. Certain Effects under Irrigation of Copper Compounds upon Crops, by
R. H. Forbes. Pp. 395-494, plates 6-9. April, 1917 -..22.20-22- ee
13. Experiments on the Effects of Constituents of Solid Smelter Wastes on
Barley Growth in Pot Cultures, by C. B. Lipman and W. F. Gericke. Ry
BD; 495-DS 7. EAL CR LOU ak eS EE OMS A ect aes nee ae res

>

UNIVERSITY OF CALIFORNIA PUBLICATIONS

TECHNICAL BULLETINS
COLLEGE OF AGRICULTURE, AGRICULTURAL EXPERIMENT STATION

ENTOMOLOGY
Vol. 1, No. 7, pp. 301-346 July 20, 1917

APHIDIDAE OF CALIFORNIA’

NEW SPECIES OF APHIDIDAE AND NOTES FROM VARIOUS
PARTS OF THE STATE, BUT CHIEFLY FROM THE
CAMPUS OF THE UNIVERSITY OF CALI-
FORNIA, BERKELEY, CALIFORNIA

BY
E. O. ESSIG

CONTENTS Sie

BE EOCHIG ETO were see oe thee ee Chau Pol ee rN eet I bee ee 302
MescripiionwotemewaSpeCles: sist... Sacer Re iy ee 302
Mivzo calliswarancimamigdes sl SY es 25.2 cen cscee 2 codec sandeacocceensnceulevesctsesneses 302
Winged viviparous female—virgogenia —............2-.-...:ceceeseeeeeeeeees 303
HOSts mlocalitivesmdatestanducolle@tong ssa ee 305

COLORES] OES) iach ts Rs nk i aR a aah te Re NO Ne ee 305

Mey 7oeallisvanundicolens (Olarke inte ee A eo 305
Winged vipiparous female—virgogenia ................2c..-2ceeeeeeeeeeeeees 305

ES NGL calls cl enna EAS eee ke oo ate ek nn 306

BS GEST EU sy permeates ere eee ee eee Re Le ide a skh navel a ay 307

DE CSSOaED I oa Oe 2 asa nm ne 307

DYES) reese A SR AO POSE NY FO RE 307

Vai Gigenme enemies nensniser reere Raia a eee Oa MCE tute oe dee OF) ei Soe 308

Hosts ~localitvestadatesmanducolllectors, ses te see ee 310

STEM HCG) OND SIO Hoa grt RROUITE: = yap ce <0 aye eat ks ae a 311
Apterous viviparous female—virgogenia —......0...220.0.2eeceeeeeeee ene 311
Winged viviparous female—virgogenia —.....20.....2......cceceeseceeeeeneees 312°

Host plant, localities, dates and collectors ..........-..0..--c20cecee0-2 os Bil}

Mi ziiseea mile gta Gr MsiSps onset en Oe te Ne Ge coe ee aE 314
Apterous viviparous female—virgogenia _..0220 0 eee 314
Winged viviparous female—virgogenia ................2...-.-sc1eeeeeee eee 315

Host plants, localities, dates and collectors ..........2-..:.--0.-0--000c0-0---- SLT
JIMS CRIBS aris) ORS anaes Seaburn as OAD PSION aoeiee GMa a en ee eae Taf 317
Apterous viviparous female—virgogenia W000. 318
Winged viviparous female—virgogenia —....0.......2..2022.2:eeeceeeeeeeeee 319

Host plants, localities, dates and collectors ....00.002.2020ceeceeeeeeee-- 320
TOUS Mee A Se, A ne SO eee eee OR ae ee ee 321

_1This paper is No. XI in the author’s series ‘‘ Aphididae of Califor-
nia,’’ published in the Pomona Journal of Entomology and Zoology.

CS

m AUG 2+ 1917 #
WV, |

Sonal Mus

302 University of California Publications [ENToMoLoey

INTRODUCTION

California offers a specially inviting field for the study of
the Aphididae not only because of the comparatively large num-
ber of new species vet undescribed, but also because of the great
and varied adaptations constantly going on to the everchanging
vegetation at the hand of man. The campus of the University
of California with its many native plants growing under per-
feetly natural conditions and the hundreds of introduced species,
together with numerous parks such as Golden Gate Park, San
Francisco, furnish large and excellent fields for study and in-
vestigation. In such environment there are not only many new
species of insects to be had, but there are the constant changes
of food plants and distribution of old and well known species.
The material at hand was taken under such conditions and rep-
resents but a small part of a season’s work. Unless otherwise
specified the data was taken in the field by the writer.

DESCRIPTION OF NEW SPECIES

Myzocallis arundinariae, n. sp.

1. Davidson, W. M., Jour. Econ. Ent., vol. 2, p. 301, 1909, Calltp-
terus arundicolens (Clarke).

2. Essig, E. O., Pom. Coll. Jour. Ent., vol. 4, no. 3, p. 762, 1912,
Myzocallis arundicolens (Clarke).

3. Essig, EH. O:, Inj: & Ben. Ims. Cal} ed: 1; p: 835, 1913s figs 65;
Myzocallis arundicolens (Clarke).

4. Davidson, W. M., Jour. Econ. Ent., vol. 7, pp. 129-130, 1914,
fig. 2, Hucallipterus arundicolens (Clarke).

5. Essig, E. O., Inj. & Ben. Ins. Cal., ed. 2, p. 84, 1915, fig. 67,
Myzocallis arundicolens (Clarke).

During the year 1911 the writer received from Mr. C. W.
Beers, Horticultural Commissioner of Santa Barbara County, a
few leaves of bamboo taken at Carpinteria, California, and in-
fested with a yellow and black plant louse which agreed so well
with the deseription of Callipterus arundicolens (Clarke)? that
it was unquestionably listed as that species. The same insect was
taken and received by the writer from a number of localities
throughout the state since that time, but it was not until the past
year, when a close study of the species described by Professor

2Can. Ent., vol. 35, p. 249, 1903.

Vow. 1] Essig.—Aphididae of California 303

Clarke was made in the type locality at Berkeley, that it was
discovered that the former insect had some marked and constant
characters not common to the latter and that the differences were
sufficient to separate the two. The former has therefore been
described as a new species, Myzocallis arundinariae from the
generic name of the bamboo most commonly infested. Only the
winged viviparous females—virgogeniae—could be secured,
though the writer made a special trip to Sacramento where he
had previously taken the species, and enlisted aid from all pos-

Fie. 1. Myzocallis arindinariae, n.sp. Winged viviparous female. 4,
adult; B, lateral aspect of head showing protuberance; C, cornicle; D,
ecauda and anal plate; H, lateral view of the dorsum of the abdomen
showing dark tubercles and cauda; F, and G, antenna. (Original.)

sible sources in the attempt. The apparent absence of sexuales
is also in contrast to Myzocallis arundicolens (Clarke) which had
an abundance at this time.

WINGED VIVIPAROUS FEMALE—VIRGOGENIA
The color varies from whitish to pale yellow with noticeable
black markings and bright red compound eyes. The average
length is 1.4 mm. and the width near the base of the abdomen
0.4mm. The head is often slightly dusky and has two lateral
and a dorsal longitudinal black vittae. The antennae (fig. 1,

304 University of California Publications [ENToMoLocy

F, G@) are yellow and black and often covered with a white fluffy
material giving a bluish cast to the black beneath. Articles I,
II and VI are dusky, while all of III excepting the middle is
black and IV and V are black at the extreme bases and the
apices. The length of the antennae is more than twice that of
the body; that of the respective articles being: I 0.1 mm., II
0.08 mm., III 0.98 mm., IV 0.65mm., V 0.58 mm., VI 0.64 mm.
(base 0.29 mm., filament 0.35 mm.), total 3.03 mm. The sensoria
are transversely oval and variable in size, but the majority are
about equal in length to half the width of the article. On
Article III there are from four to nine confined to the basal third.
The following tabulation will give an idea of the number on the
pairs of antennae of forty-four individuals, the colon separating

the individuals and the comma the articles, 7, 7:6, 5:5, 5:6, 6: 7,
D1, 06102 OO DOD, Oe (102 SCO ie Or OO ino i aieene:
8:6, 736.724, 5- 6: 6:6, DOr eiOnle Oe Ol Oneal OM aoe
024,625, 0% 0, 02D. 625,602 O12 On OF/ON Te O).0 20; OF Ot OME

Sia:

The usual number occur on articles V and VI. There are but
few very inconspicuous hairs on the antennae. <A very large
protuberance is located on the underside of the head just in front
of the rostrum (fig. 1, B) which it equals in length. The rostrum
is very short, reaching only to the front coxae. The thorax is
yellow with amber or shghtly dusky muscle lobes and a dark
vitta on each side in front of the bases of the wings and two
indistinct dusky lines on the dorsum. The venation of the wings
is shown in the accompanying drawing (fig. 1, 4). The legs
are yellow with the apical portions of the tarsi dusky. The
abdomen has two longitudinal rows of dorsal tubercles (fig. 1,
A, E,); there being eight separate pairs and one large patch
made by the fusing of the ninth pair. A small hair or spire
arises from each of the dark areas and two from the largest
posterior one. The cornicles (fig. 1, C) are yellow, short and
much wider at the base than at the mouth. <A very long hair
arises from a tubercle near the middle of the hind margin. The
length is 0.1mm. The cauda (fig. 1, D) is knobbed and con-
spicuously dusky or black. The anal plate (fig. 1, D) is dis-
tinctly bilobed and pale or dusky yellow.

The nymphs are pale yellow or very bright amber in color
and covered with numerous long knobbed spines or hairs.

———— ee

Vow. 1] Essig. —Aphididae of California 305

HOSTS, LOCALITIES, DATES AND COLLECTORS

1. Bamboo, Arundo, sp., Stanford University, Cal., October, 1908?.
W. M. Davidson.

. Bamboo, Arundinaria, sp.?, Carpinteria, Cal., July 20, 1911. C.
W. Beers.

3. Bamboo, Arundo, sp., San José, Cal., Oct. 15, 1911. W. M.
Davidson.

4. Bamboo, Arundinaria japonica 8. & Z., Capitol Park, Sacramento,
Cal., April 30, 1912. The writer.

5. Bamboo, Arundo, sp.?, Exposition Park, San Diego, Cal., April
27,1916. A. F. Swain.

6. Bamboo, Arundinaria, sp.?, Montecito, Cal., June 27, 1916. C.
W. Beers. ;

. Bamboo, Arundinaria japonica 8. & Z., Capitol Park, Sacramento,
Cal., Dee. 14, 1916. The writer.

ine)

“I

The writer has also had reports of its occurrence elsewhere in
Southern California but has not been able to verify them.

In all eases the insects work on the leaves, usually the under-
sides, and produce sufficient honey-dew to cause considerable
smutting.

COTYPES

All of the descriptions made by the author have been from a
series of individuals which of course become cotypes. These have
been placed in the collections of the U. S. National Museum,
Washington, D. C., and the California Academy of Sciences, San
Francisco, Cal., while a good series in each case has remained in
the author’s collection. In ease of the species herein described
they have been deposited as directed above.

Myzocallis arundicolens (Clarke)
WINGED VIVIPAROUS FEMALE—VIRGOGENIA
1. Clarke, W. T. (orig. dese.), Can. Ent., vol. 35, p. 249, 1903,
Callipterus.
2. Essig, E. O. (host index), Pom. Coll. Jour. Ent., vol. 3, no. 2,
p. 458, 1911, Callipterus.

3. Essig, E. O. (list), Pom. Coll. Jour. Ent., vol. 4, no. 3, p. 263,
OU

The color of the winged viviparous female is from whitish
to pale lemon-yellow with bright red compound eyes, a pitch

3 The writer received a mounted slide of specimens taken by Mr.
Davidson at San José, California, which proved to be this species. These
specimens are probably the same as those which he also took at Palo Alto
as recorded above.

306 University of California Publications [ENtTomMoLocy

black cauda and black or dusky antennal markings. It very
much resembles the corresponding form of the preceding species
in size, color and shape, but lacks the black tubercles on the
dorsum and has the sensoria on article III of the antennae
grouped about one-third the distance from the base instead of
being confined to the basal third (fig. 2, D). The number of
sensoria is also smaller as will be seen from the following tabu-
lation of thirty-one individuals 6, 6:7, 7:5, 5:6, 8:4, 5:4, 4:5,

Fie. 2. Myzocallis arundicolens (Clarke). Winged viviparous female.
A, adult; B, cornicle; C, cauda, and anal plate; D and E, antenna; F,
article III of the antenna of a sexupara.

6:5, 62 56: 6: 6: Db. 52 6.1615. O16) 62 On O 2 fol Onn OmbaGr
62 4.°6:.6,67216, 6.25, 62.9,,600. 070, 625.670) Oc 65.6, O- eee le
stigma of the wings usually lack the darker spots at the base
which is so constant in Myzocallis arundinariae, and the cauda
is usually blacker.

SEXUPARA
Fig. 2, F
The sexupara is very much lke the virgogenia, but is usually
a little larger. There are also more sensoria on article III of the
antennae (fig. 2, /’) as shown by the following tabulation of
eighteen individuals: 7, 8:10, 10:7, 7:8, 9:6, 9:8, 9:6, 1027,

elmo Ie fede /Groh waren Wie Wi TOE fe le tobe ioe 7 IMG).

Vou. 1] Essig —Aphididae of California 307

SEXUALES

FEMALE

The sexual female is large, robust and apterous, varying from
whitish to very pale yellow and immaculate excepting the dark
patches on the antennae and tarsi as well as the pitch black cauda.
The length averages 2mm. and the width 1.3 mm. The dorsum
has four longitudinal rows of hair-like spines which arise from
quite large tubercles. The antennae (fig. 3, C, D) are longer
than the body, the individual articles being as follows: I 0.11
mm., II 0.07 mm., III 0.9 mm., IV 0.57 mm., V 0.5 mm., VI 0.66

Fic. 3. Myzocallis arundicolens (Clarke). Sexual female. 4, side
view; 6, hind tibia showing sensoria; C and D, antenna; ££, cornicle.
(Original. )

mm. (base 0.31 mm., filament 0.35 mm.), total 2.81 mm. The
sensoria are transversely oval and similarly located as in the
other forms, but are fewer in number, varying from none at all
to three or four. The hind tibiae (fig. 3, B) are swollen and
covered with a great many small circular sensoria.

Eaas
The eggs are pale yellow or whitish when first laid, but soon
become shining black. They are oval with a conspicuous blunt
stipe or pedicel at one end. The length averages 0.07 mm. They
are deposited singly, in uneven masses or more often in rows
on the undersides of the leaves and are at once conspicuous to
the naked eye.

308 University of California Publications [ENTomovoey

MALE
The male is winged and much darker in color than any of the
other forms. The average length is 1.5 mm. The head and °
articles I, If and VI of the antennae are dusky, while article IIT
is almost black throughout, the apical half of IV is also black

Fic. 4. Myzocallis arundicolens (Clarke). Eggs. The pale ones are
freshly laid and become black soon afterwards. Three are greatly en-
larged to show the pedicel at one end. (Original. Photo by Div. Sei.
Illust., Univ. Calif.)

and V is a little darker than VI. The antennae (fig. 5, B-E’) are
much longer than the body, the lengths of the respective articles
being: I 0.07 mm., II, 0.07 mm., IIT 0.93 mm., [V 0.58 mm., V
0.48 mm., VI 0.68 mm. (base 0.31 mm., filament 0.37 mm.), total
2.81 mm. The sensoria are transversely oval and numerous on

Vou. 1 | Essig.—Aphididae of California 309

article III, numbering from 21 to 30 and covering the entire
length excepting the extreme base. On article IV there are from
3 to 8 which are usually confined to the apical two-thirds of the
joint. On article V there are from 2 to 9 distributed indefinitely
along the entire length. There are from 1 to 6, not including
those in the process, on article VI located usually near the middle
of the base. The following tabulation, which does not include the
sensoria in the process of VI, will serve to give an idea of the
variation in numbers on the respective articles :

Fic. 5. Myzocallis arundicolens (Clarke). Male. A, dorsal aspect;
B E, antennae. (Original.)

Individuals IEEE IV V VI
1 27 3 8 2
26 4 8 2

2 29 5 9 3
26 6 5 3

3 25 7 6 2
25 4 6 6

4 21 8 5 2
25 4 6 5

5 26 3 5 3
¥

26 Uf 3

oo

OO
em DO OS
bo bo

310 University of California Publications [ENToMoLoGy

Individuals III TEV: Vv VI
7 26 3 9 3
28 6 7 3
8 28 6 8 article missing
30 3 7 2
9 24 4 fa article missing
29 7 8 article missing
10 24 4 5 article missing
26 5 8 article missing
fal 25 4 6 il
26 5 7 2

The legs are quite long, dusky yellow with almost black femora
which have rather prominent oval or circular pale areas showing
specially well in the mounted specimens. The thorax is yellow
or dusky with brown or black muscle lobes. The abdomen is
yellow with black spots along the margins and numerous black
patches on the dorsum usually arranged in transverse rows. The
cornicles are short and dusky yellow. The cauda and anal plates
are from dusky to pitch black. Short spines arise from the dusky
areas on the dorsum, being arranged in longitudinal rows.

HOST PLANTS, LOCALITIES, DATES AND COLLECTORS

1. Bamboo, Arundo, sp.,4 Berkeley, Cal., 1903. W. T. Clarke.
2. Bamboo, Arundinaria japonica 8. & Z., U. C. Campus, Berkeley,
Cal., 1916. Different forms taken by the writer as follows:

Virgogeniae—Abundant on the undersides of the leaves
throughout the summer and fall. Those described were col-
lected on June 1 and July 17.

Sexuparae—Quite numerous in the late fall. Collected Nov.
28 and 29 and Dee. 28.

Sexuales—The females abundant from the last of October to
the first of December and a few stragglers remaining until
after January 1. Collected Nov. 28 and 29 and Dec. 28. The
males not abundant at any season, but most numerous about
the middle of November. Collected Nov. 28, and 29. None
were to be found as late as Dee. 28.

Eggs were laid the last of October and a few still being
deposited in late December, but on Dee. 28 very few freshly
laid eggs could be found, practically all having already
assumed the shiny black color.

4 This plant is undoubtedly Arundinaria japonica 8. & Z., which is the

common species infested on the campus where the type specimens were
also taken.

Vou. 1] Essig.—Aphididae of California 311

Symydobius® agrifoliae, n. sp.

This species was first collected by S. H. Essig and the writer
in 1911, but as only apterous forms were secured at that time no
attempt was made to describe it. During the spring of 1916 a
number of winged and apterous viviparous females were taken
by S. H. Essig and forwarded to the writer. As previously
believed it proved to be a new species, which I am naming after
the species of the host plant. It is with gratitude here that I
acknowledge the great help constantly given by my brother men-
tioned above.

Fig. 6. Symydobius agrifoliae, n.sp. Apterous viviparous female. 4,
dorsal aspect; B, cauda and anal plate; C, cornicle; D, antenna.
(Original. )

APTEROUS VIVIPAROUS FEMALE—VIRGOGENIA

The mature apterous female is rather robust and of a dark
reddish brown color with hghter amber or yellowish markings
describing roughly the letter Y on the back. The dorsum and
sides are also obscurely marked with black blotches. The length
averages about 1.6 mm., and the greatest width is 1.1 mm. The
head is rather narrow and pale amber at the bases of the
antennae. The eyes are dark red. The antennae (fig. 6, D) are

5 The genus Symydobius has been wrongly spelled Symdobius in the
author’s former articles.

oie University of California Publications [ENToMoLocy

dusky with the greater portion of article III and the bases of
articles IV and V pale. The length is nearly as great as that
of the body, the measurements of the respective articles being:
T 0.08. mm., II 0.07 mm., ITT 0.41 mm., IV 0.3 mm., V 0.26 mm.,
VI 0.16 mm. (base 0.13 mm., spur 0.03 mm.), total 1.28 mm.
There is considerable variation in the measurements of different
individuals, but the relative lengths are fairly constant. The
usual sensoria are present on articles V and VI, but very few
hairs are present on any of the articles. The rostrum is light
amber in color and reaches slightly beyond the third coxae. The
legs are rather short and dark brown with lighter areas in the
middle of the tibiae of most specimens. The cornicles (fig. 6, C)
are dark, very short, wide at the base and narrow at the mouth,
which has a very wide lip or flange and small opening. The
cauda (fig. 6, B) is dark, broad at the base with a rounded tip
and normally hairy. The anal plate (fig. 6, B) is dark, rounded
and has a small but distinet incision or notch in the middle.
The nymphs are shghtly paler in color than the mature forms.

WINGED VIVIPAROUS FEMALE—VIRGOGENIA

The winged forms are dark reddish brown with black mark-
ings. The length averages 1.9 mm., and the greatest width 1 mm.
The head is light or very dark reddish brown. The antennae
(fig. 7, C) are dusky or black with the greater portion of article
IIT and the basal halves of TV and V pale yellow or amber. They
are about three-fourths the length of the body, the measurements
of the respective articles being: I 0.1 mm., IT 0.07 mm., IIT 0.44
mm., [V 0.8 mm., V 0.28 mm., VI 0.18 mm. (base 0.13 mm.,
spur 0.05 mm.), total 1.37 mm. There are from four to eight
large and small circular sensoria on article III. Of fourteen
antennae examined to ascertain the number of sensoria on article
IIT it was found that but one had 4 sensoria, three had 5, six
had 6, two had 7, and two had 8. Articles V and VI have the
usual sensoria. The rostrum is amber in color and reaches to the
third coxae. The thorax is reddish brown with very dark brown
or black muscle lobes. The wings are subhyaline with distinct
dusky bordered veins. The venation is shown in the acecompany-
ing drawing (fig. 7, 4). The legs are dark with lighter areas in
the middle of the tibiae. The abdomen is reddish brown with one
or two rows of dark or black markings near the margins from

Vou. 1] Essig.—Aphididae of California 313

which arise short spines. The dark areas increase in size towards
the posterior end. The cornicles (fig. 7, B), the cauda and the
anal plate are much the same as in the apterous form already
described.

HOST PLANTS, LOCALITIES, DATES AND COLLECTORS

This species feeds in compact colonies on the bark and very
rarely on the leaves of the coast live oak, Quercus agrifolia Nee.
As already stated apterous forms were first taken by 8. H. Essig

Fic. 7. Symydobius agrifoliae, n.sp. Winged viviparous female. 4,
dorsal aspect; B, cornicles; C, antenna. (Original.)

and the writer in Santa Paula Canon, near Santa Paula, Cali-
fornia, on July 21,1911. Apterous forms as well as winged were
also taken by the former collector in the same locality on May
27, 1916, as well as along the Ventura River near Ventura, Calli-
fornia, on June 21, 1916.

Symydobius agrifoliae, n.sp. is closely related to Symydobius
albasiphus Davis, but is at once separated by having a much
shorter spur on the sixth antennal article as well as other differ-
ences.

314 University of California Publications |ENtomoLocy

Myzus aquilegiae, n. sp.
APTEROUS VIVIPAROUS FEMALE—VIRGOGENIA

The apterous female is very pretty, being of a pinkish or
reddish color with a very large dark brown or nearly black
irregular blotch on the middle of the back. The body is of
medium size, beset with short knobbed hairs and averaging 1.7
mm. in length and 0.7 mm. in width. The antennae (fig. 8, F,
G) arise from definite but short frontal tubercles (fig. 8, B),

Fic. 8. Myzus aquilegiae, n.sp. Apterous viviparous female. 4,
dorsal aspect; B, head and frontal tubercles; C, cornicle; D, cauda; L,
tarsus; F-G, antenna. (Original.)

are shghtly longer than the body, dusky throughout excepting
the base of article III, imbricated and supporting a number of
knobbed hairs. The lengths of the respective articles are: I
0:09: mm., IT, 0:07 mm., If! 0.5 mm. TV 0:4 mm. V 0:33 ams
VI 0.63 mm. (base 0.1 mm., filament 0.53 mm.), total 2.02 mm.
The sensoria are circular and of various sizes. On article III
they are distributed throughout the length, but are much more
numerous near the middle. On IV there are usually none, but
one or two may sometimes occur. The usual number may be
found on V and VI. The following tabulation gives the variation
found on articles III and IV from eighteen individual females:

Vou. 1] Essig.—Aphididae of California 315

Number Article III Article 1V Number Article III Article 1V

il 26 3 10 18 0
30 4 15 0

2 26 it Il 20 0
2 0 24 0

3 19 0 12 i; 0
22 i 14 0

4 17 0 3} 16 0
16 0 11 0

3) 18 0 14 14 0
14 0 12 0

6 11 0 1155 26 2
13 0 25 0

fl 17 0 16 iS 0
21 il 12 0

8 22, 0 WZ 10 0
0 15 0

9 16 0 18 19 0
14 0 23 2

The rostrum is pale with dusky tip and reaches slightly beyond
the third coxae. The legs are pale yellow with the apices of the
tibiae and all of the tarsi dusky. The tarsi (fig. 8, EF’) are very
small. The abdomen is yellowish, pinkish or reddish in color
with a large brown or black patch on the dorsum and a marginal
row of small dark spots on the sides. The large dorsal dark
patch may in some eases cover all of the dorsum excepting the
extreme base and the area behind the cornicles. The cornicles
(fig. 8, C) are whitish or yellow, faintly imbricated, nearly
cylindrical, with slightly wider base and small flare at the mouth.
The length averages 0.47 mm., or about three times the length of
the hind tarsi. The cauda (fig. 8, D) varies from yellowish to
pale pink or reddish, is gradually pointed and about two-thirds
as long as the cornicles.

WINGED VIVIPAROUS FEMALE—VIRGOGENTA

The winged viviparous female is yellow or reddish with a
number of dark brown or black markings over the body. The
length averages 1.4 mm., and the width 0.45 mm. The body is
sparsely covered with simple and knobbed curved spines. The
head is black and the compound eyes red. The antennae (fig. 9,

316 University of California Publications [ENTomoLocy

F, G) arise from small, but distinct frontal tubercles (fig. 9, B),
are black throughout with few knobbed hairs. The length is
greater than that of the body, the respective articles measuring:
I 0.07 mm., II 0.06 mm., III 0.52 mm., IV 0.4 mm., V 0.36 mm.,
VI 0.8 mm. (base 0.12 mm., filament 0.68 mm.), total 2.21 mm.
The sensoria are circular, of various sizes and numerous on
articles III, IV and V, with the usual ones in the process of VI.
The number varices considerably on the different articles, as the
following tabulation will show:

Article Article Article Article Article Article

Number III IV Vv Number III IV Vv
1 24 0 0 8 34 17 8
26 J 0 31 18 6

2 30 21 4 9 34 21 5
36 16 9 37 16 8

3 31 18 5) 10 42 23 3
32 17 9 38 18 4

4 44 23 3 Hal 34 14 5
42 23 2 32 16 i)

5 38 19 6 12 24 15 5
35 23 8 27 14 3

6 35 19 5) 13 26 11 6
29 15 3 23 16 6

7 33 16 3 14 38 14 9
34 19 5) 34 18 5

The rostrum is pale with the apical half dusky and reaches
slightly beyond the third coxae. The thorax is black with reddish
areas at the sides and between the segments. The wings are
normal in length with brownish veins and stigma. The venation
is shown in the accompanying drawing (fig. 9, A). The legs are
nearly all black, excepting the middle of the tibiae and the
bases of the femora. As in the apterous form the tarsi (fig. 9,
FE) are exceptionally small. The abdomen is reddish with small
irregular dark markings on the sides and dorsum. The cornicles
(fig. 9, C) vary from pale dusky to black, slightly imbricated,
eylindrical with a shghtly flaring mouth. The length is 0.32 mm.,
or about three times the length of the hind tarsi. The cauda
(fig. 9, D) is yellow or reddish and about two-thirds the length
of the cornicles.

Vou. 1] Essig.—Aphididae of California 317

HOST PLANTS, LOCALITY, DATES AND COLLECTOR

A native columbine, Aquilegia truncata F. & M., cultivated
in the botanical garden on the campus, appears to be the pre-
ferred food plant, though a few specimens were found through-
out the season on a nearby species, A. chrysantha Gray. The
small tender shoots and buds are invariably infested and often
the plants are considerably injured by the attacks. The species
passes the entire year on the columbine, going as far down

Fic. 9. Myzus aquilegiae, n. sp. Winged viviparous female. 4, dorsal
aspect; 6, head showing frontal tubercles; C, cornicle; D, cauda; EH,
tarsus; # G, antenna. (Original.)

around the crown as possible during the winter. The specimens
described were collected by the writer on June 27 and July 3,
1916. It was first noticed by the writer in 1914.

G. O. Shinji, a student, tells me that he has taken this
species on a wild columbine in Marin County, but I have not seen
his material to verify the same.

Aphis cari, n. sp.

A small plant louse occurring in compact colonies on the
stems of the common sweet or wild anise, Carum kelloggi Gray,

318 University of California Publications [ENTomMoLocy

was of considerable interest to me when first taken because of
the characteristic and beautiful coloring. A few specimens were
also taken on Angelica tomentosa Wats, which led me to believe
that the species might be Aphis angelicae Koch, reported as
occurring in this state by H. F. Wilson,® but a microscopic
examination at once proved it to be widely different from the
species described by Wilson. Not being able to place it in any
published description I have decided to describe it as new and
to name it after the genus of what appears to be the favorite
host plant.

Fic. 10. Aphis cari, n.sp. Apterous viviparous female. A, dorsal
aspect; B, tarsus; C, eauda; D, cornicle; H, antenna. (Original.)

APTEROUS VIVIPAROUS FEMALE—VIRGOGENIA

The typical apterous viviparous female is transparently
white, yellow or pale yellowish green with very noticeable darker
ereen markings on the dorsum and particularly a distinct trans-
verse patch between the bases of the cornicles. The form is
robust, the length averaging 1.7 mm. and the width 1.2 mm. The
head is usually pale whitish, yellow or pale yellowish green with-
out darker markings. The antennae (fig. 10, E) are whitish or
yellow throughout the basal half, the remainder being dusky or
black. Most of the articles are distinctly imbricated. The
length is about three-fourths that of the body, the lengths of the
respective articles being: I 0.06 mm., II 0.04 mm., III 0.32 mm.,

* Jour. Econ. Ent., vol. 2, pp. 348-349, Oct., 1909.

Vou. 1] Essig. —Aphididae of California 319

IV 0.21 mm., V 0.16 mm., VI 0.36 mm. (base 0.09 mm., filament
0.27 mm.), total 1.15 mm. The usual sensoria occur on articles
V and VI with none on III. The rostrum is pale, transparently
white and reaches to the third coxae. The prothorax has a large
lateral tubercle at the base and on each side of the body a tubercle
is located at the unions of the prothorax and mesothorax and the
metathorax and abdomen. The legs are normal in length, trans-
parently white with the tips of the tibiae and the tarsi dusky.
The cornicles (fig. 10, D) are dusky or black, somewhat wider at
the base with a medium-sized lip at the mouth, imbricated and

Wie. 11. Aphis cari, n.sp. Winged viviparous female. A, dorsal aspect ;
B, tarsus; C, eauda; D, cornicle; H, antenna. (Original.)

nearly twice as long as the hind tarsi (fig. 10, B), the length
being 0.33 mm. and the width at the base 0.09 mm. The cauda
(fig. 10, C) is dusky, nearly as long as the cornicles, rather
slender, with a distinctly wider base for about two-thirds the
length when viewed dorsally or ventrally.

WINGED VIVIPAROUS FEMALE—VIRGOGENIA
The winged viviparous female is pale yellowish or greenish
with dark green or black head, antennae, thorax, cornicles, apices
of the femora, and tibiae and all of the tarsi. The length aver-
ages 1.7 mm. and the width 1 mm. The antennae (fig. 11, 2)
are relatively short, reaching only slightly beyond the middle of
the abdomen. They are dusky or black throughout and imbri-

320 University of California Publications {|ENToMoLocY

cated excepting the basal fourth. The lengths of the respective
articles are: I 0.07 mm., IT 0.06 mm., IIT 0.43 mm., IV 0.21 mm.,
V 0.18 mm., VI 0.41 mm. (base 0.09 mm., filament 0.32 mm.),
total 1.36 mm. Article III is covered the entire length with
numerous circular sensoria of different sizes. The number varies
considerably, from fifteen to fifty, the average being about thirty-
five. On article IV of a great many individuals there are no
sensoria, while on a few the number varies from one to four.
There are the normal number on V and VI. The venation is
shown in the accompanying drawing (fig. 11, 4). The rostrum
is dusky and reaches slightly beyond the third coxae. The pro-
thorax has a distinct lateral tubercle on each side of the base.
The legs are transparently white or yellow with the apices of the
femora and tibiae and all of the tarsi dark. The abdomen is
yellow or pale yellowish green with a few darker green mark-
ings on the dorsum, and a distinct narrow transverse patch
between the cornicles, already referred to. The cornicles (fig.
11, D) are nearly cylindrical or with the base somewhat wider
than the mouth. They are black or very dark in color, imbri-
cated and about twice the length of the hind tarsi (fig. 11, B),
the length being 0.25 mm. The cauda (fig. 11, C) is much the
same as that of the apterous female.

The nymphs vary from pale yellow to faintly yellowish-green
with few or no darker markings.

HOST PLANTS, LOCALITIES, DATES AND COLLECTORS

As previously stated this species was first taken on the native
wild or sweet anise, Carwm kelloggu Gray, which appears to be
the normal food plant. The insects feed in compact colonies on
the stems, usually near a fork and sometimes entirely out of
sight beneath the sheath of a leaf petiole. The above infested
plants were apparently growing in a wild state along a creek bed
in Napa County near the town of Rutherford. Collected by the
writer on June 21, 1916.

The second lot, consisting of but few specimens, was taken on
the leaves of Angelica tomentosa Wats, another native plant
growing in the botanical garden on the campus in close prox-
imity to a clump of the wild or sweet anise, which at the par-
ticular time showed no infestation. The specimens were collected
by the writer on June 27, 1916.

Vow. 1] Essig —Aphididae of California 321

On June 27, 1916, Professor C. W. Woodworth took at St.
Helena a plant louse from Carum kelloggu Gray which appears
to be the same species. The apterous viviparous females, which
were the only forms collected, agree in size, shape and coloration,
but the cauda is wider and the antennal joints somewhat shorter.

NOTES ON OTHER APHIDIDAE
CHIEFLY FROM THE CAMPUS OF THE UNIVERSITY OF CALIFORNIA,
BERKELEY, CALIFORNIA

Phyllaphis, sp. (fig. 12.) A pale greenish or yellow plant louse
which secretes over the body a quantity of white woolly material
and which has commonly been ealled Phyllaphis querci Fitch.
It is not that species, as has been pointed out by Baker’ and is
probably not P. quercifoliae Gillette.s It occurs in considerable
numbers on the undersides of the leaves of the coast live oak,
Quercus agrifolia Nee., on the campus, but only apterous forms
have been secured here thus far. Specimens were received from
Altadena, California, August 14, 1916, taken on the same host
plant. What also appears to be the same thing was collected at
Rutherford, California, June 21, 1916, on valley oak, Quercus
lobata Nee. The lot included one winged male which agrees
very well with Gillette’s description of the same sex of P. querci-
foliae.

Phyllaphis fagi (inn.). A small yellow and black species
densely covered with white woolly material was taken on the
undersides of the leaves of Fagus tricolor in a lath-house at
Oakland, California, July 25, 1916. All forms appear somewhat
smaller than specimens which I have under that name in my
collection.

Drepanosiphum platanoides (Schrank). <A large dark or pale
green species infesting the leaves of Norway maple, Acer plata-
noides Linn., and silver maple, A. saccharinum Linn., in Berkeley,
and neighboring cities. The dark sexual forms are abundant in
September and October. This is the commonest species attacking
the maple here.

Drepanaphis acerifolii (Thomas). This very pretty insect
is gray and black in the winged form and pale yellow in the

7 Baker, A. C., Ent. News, vol. 27, p. 361, 1916.
* Gillette, C. P., Ent. News, vol. 25, pp. 274-275, 1914.

322 University of California Publications |ENToMoLocy

apterous, and is at once separated from other closely related
species by the large dark tubercles on the back of the winged
forms. It has been taken during several successive years and
becomes abundant in a few localities. The sexual forms were
taken September 16, 1916. The Norway maple, Acer plata-
noides Linn. and the silver maple, A. saccharinum Linn. are
attacked.

Fig. 12. Phyllaphis, sp. Colonies on the undersides of the leaves of
the valley oak, Quercus lobata Nee. The bodies are entirely covered with
the white woolly material. Rutherford, Cal., June 21, 1916. (Original.)

Calaphis betulaecolens (Fitch). A large green species com-
mon in certain restricted areas on the European white birch,
3etula alba Linn. Collected on the campus June 5, 1916.

Euceraphis betulae (inn.). A large green and black plant
louse which is commonest on the birches in this section. It occurs
on the leaves of the European white birch, Betula alba Linn., the
eut-leaved birch, B. populifolia laciniata Hort., and the paper or

Vou. 1] Essig.—Aphididae of California 323

eanoe birch, B. papyrifera Marsh. The species is abundant
throughout the year and causes much smutting of the trees.
Eucallipterus flavus Davidson. A pale green species of large
size taken in few numbers on the old leaves and in abundance on
the tender shoots of the white alder, Alnus rhombifolia Nutt.,
in Wildeat Canon, near Berkeley, California, May 10, 1916.
Eucallipterus tiliae (linn.). <A beautiful yellow and black

Fic. 13. Myzocallis castaneae (Fitch). On leaf of a chestnut tree,
Campus, Oct. 10, 1916. (Original.)

species with black clouded wings which often becomes very
abundant on linden trees, producing quantities of honey-dew
and giving the trees a black and offensive appearance. Speci-
mens were taken on the American linden, Tilia americana Linn.,
in Golden Gate Park, San Francisco, California, May 25, 1916,
and on white or silver linden, 7’. tomentosa Moench, Berkeley,
California, June 17, 1916.

Myzocallis castaneae (Fitch) (fig. 13). A small yellow aphid
which has been called this species occurs on hickory and chest-
nut on the campus and is often in large enough numbers to

324 University of California Publications |[ENtTomoLocy

cause a general smutting of the trees. Viviparous females occur
throughout the spring and summer. Sexual forms were taken
on October 10, 1914.

Myzocallis coryli (Goetze). This very small pale yellow
species occurs in great numbers on the undersides of the leaves
of an introduced hazel or filbert, Corylus maxima Mill, growing
in the botanical gardens on the campus. The insects secrete
quantities of honey-dew and cause severe smutting of the fohage.
Collected in great numbers on June 28, 1916, when apparently
at its maximum development.

Myzocallis pasaniae Davidson. <A yellow species of median
size infesting the undersides of the leaves of the tanbark oak,
Pasania densiflora Oerst., growing on the Campus, June 27, 1916.

Myzocallis quercus (Kalt). This small species varies in color
from pale yellow to light green and has four rather conspicuous
dark tubercles on the dorsal base of the abdomen. It generally
infests a number of deciduous oaks on the campus, including the
English oak, Quercus robur Linn. Collected October 10, 1914,
and June 20, 1916. It was also taken on what appears to be the
same oak in Golden Gate Park, San Francisco, California, May
25, 1916. The species was questioned for some years but was
recently verified for W. M. Davidson by P. van der Goot and
for the author by A. C. Baker. (The recently published paper in
The Entomological News, vol. 28, no. 2, Feb., 1917, pp. 62-63,
plate 7, by G. O. Shinji, a student here, has added to the con-
fusion. He described two species, Wyzocallis essigi and M. wood-
worthi from M. quercus (Kalt.). M. essigi is the common forms of
M. quercus and M. woodworthi is the male of the same thing. All
of the specimens were taken from the tree from which my speci-
mens of M. quercus were taken. Had Shinji submitted his paper
to some member of the entomological department as should have
been done such an error could have been avoided. Mr. A. C.
Baker examined the type slides also and agrees with my deter-
minations. It might be stated here that there are a number of
different species on the type slides and it was necessary to have
Mr. Shinji designate the type deseribed in each case.—March 1,
OAT)

Myzocallis ulmifolii (Monell). A bright yellow species occur-
ring on the leaves of English elm, Ulmus campestris Smith, and
other elms at Berkeley, California, September 20, 1916, and at
Sacramento, California, July, 1914.

Vou. 1] Essig.—Aphididae of California 325

Chromaphis juglandicola (Kalt.). The common small yellow
aphid occurring in great numbers on the undersides of the leaves
of the cultivated English walnut, Juglans regia Linn. It is
abundant throughout the state where this tree is grown and often

Fig. 14. Pterocomma flocculosa (Weed). Three apterous viviparous
females almost hid by the white flocculence. On arroyo willow, Salix
lasiolepis Benth. Berkeley, Cal., June 15, 1916. (Original.)

causes much smutting of the foliage. So far I have never taken
it from any other species of the walnut’.
Thomasia salicicola Essig. A very dark species with a median
dorsal longitudinal line on the backs of the apterous females
9 A common yellow species oceurring on the leaves of various black
walnuts in this region does not seem to agree with the descriptions of

Callipterus caryae (Monell) or Monellia caryella (Fiteh), however it has
been determined as the latter by A. C. Baker.

326 University of California Publications [ENTomoLocy

occurring in very dense colonies on the terminal shoots of wil-
lows. Taken on the campus, June 13, 1916, infesting the arroyo
willow, Salix lasiolepis Benth.

Pterocomma flocculosa (Weed) (figs. 14 and 15). Taken at

Fig. 15. Pterocomma flocculosa (Weed). Eggs laid on arroyo willow,
Salix lasiolepis Benth., Sacramento, Cal., Dec. 14, 1916. (Original. Photo
by Div. Sei. Illust., Univ. Calif.)

Berkeley on arroyo willow, Salix lasiolepis Benth., June 15, 1916,
at Rutherford, California, June 21, 1916, on the same willow,
and at Sacramento, California, December 14, 1916, on the same
plant. In the latter case the oviparous females were depositing
quantities of eggs which were partially concealed by white

Vou. 1] Essig.—Aphididae of California 327

woolly material. This bark feeder is at once distinguished by
the partial covering of thin white wool and the bright orange
cornicles.

Pterocomma populifoliae (Fitch). This dark gray and brown
species occurs plentifully on Salix, spp., on the campus, March
29,1916. Lacks the cottony covering of the preceding species.

Tuberolachnus viminalis (Fonse.) (fig. 16). Our largest
species occurring on willow and easily recognized by the gray
color and the large black tubercle on the back between the
cornicles. Abundant on the trunks and branches of various
willows and feeding in large compact colonies. Taken on yellow
willow, Salix lasiandra Benth., red willow, S. laevigata Bebb.,

Fic. 16. Tuberolachnus viminalis (Fonse.). Apterous viviparous females
on longleaf willow, Salix longifolia Muhl. Berkeley, Cal., June 10, 1916.
(Original. )

longleaf willow, S. longifolia Muhl., and arroyo willow, S. lasio-
lepis Benth., on the campus and at Berkeley and on the arroyo
willow at Rutherford, California, June 21, 1916.

Lachnus pini-radiatae Davidson. This very dark green
species is partially covered with a thin whitish woolly secretion
and usually feeds in colonies in rows on the pine needles. Taken
on the campus in small numbers on yellow pine, Pinus ponderosa
Dougl., January 1 and March 11, 1915.

Lachnus tujafilinus Del Guereio. A robust brown and grayish
species occurring in dense colonies on the twigs of common arbor-
vitae, Thuya occidentalis Linn. Received from Chas. B. Weeks,
Red Bluff, California, February 8, 1916.

Nectarosiphum rubicola (Oestlund). A large normally green
species occurring in colonies on the undersides of the leaves of

328 University of California Publications |ENtTomoiocy

the thimble berry, Rubus parviflorus Nutt., on the campus, March
29, 1916. At this date there were also a number of young pink
forms not observed during the summer. The insects were especi-
ally abundant on the thimble berry in Wildcat Canon near
Berkeley, May 24, 1916. The species is easily recognized by the
dusty tip of the primary wings.

Macrosiphum albifrons Essig. A large green species covered
with a white pulverulence, which feeds upon various lupines, in-
eluding Lupinus latifolius Agarth. and tree lupine, LZ. arboreus
Sims., on the campus, March 29, 1916.

Macrosiphum californicum (Clarke) (syn. MW. laevigatae
Essig). A bright green, large very slender species infesting the
terminal shoots of the young growth of various willows, especially
the arroyo willow, Salix lasiolepis Benth., and the yellow willow,
S. lasiandra Benth. Oceurs on the campus throughout the sum-
mer and collected June 1, 13, and July 26, 1916. Also taken on
the arroyo willow at Rutherford, California, June 21, 1916.

Macrosiphum granarium (Kirby). A large green and black
aphid common on various grasses and taken as follows: oats,
campus, March 10, 1916, and at Martinez, California, June 12,
1916; on meadow fox-tail, Alopecurus pratensis Linn., campus,
April 25, 1916; on Ampelodesma tenax Link. and wild rye,
Elymus, sp., campus, June 27, 1916. In case of the last two
plants the flower heads were entirely covered with the insects.

Macrosiphum lactucae (Schrank). <A large bright green and
black species common on the tender terminal shoots of chicory,
Chicorium intybus Linn., growing in an apparent wild state at
Rutherford, California, June 21, 1916.

Macrosiphum ludovicianae (Oestlund). A large green species
with the bodies of many covered with a thin white pulverulence.
Collected on the campus March 10, 1916, and at Rutherford,
California, June 21, 1916. In both cases the terminal tips of
the common wormwood or California mugwort, Artemisia hetero-
phylla Nutt., were infested..

Macrosiphum rhamni (Clarke). This large species is green
with a noticeable large reddish, nearly cireular blotch in the
middle of the dorsum at the union of the thorax and abdomen.
This coloration is internal and disappears with mounting.
Apterous and winged forms were specially abundant on the
leaves of coffee berry, Rhamus californicus Esch., and less com-
mon on caseara sagrada, R. purshiana DC., on the campus during

'

Vou. 1] Essig —Aphididae of California 329

1916. Taken on the former, July 5, when it appeared to have
reached its maximum development, and on the latter, June 29,
1916.

Macrosiphum rosae (Linn.). This is the common large green
and reddish louse so common in the rose gardens of the state.
Besides being found on all varieties of roses, this species has
also been taken on the stems of the heads of the fuller’s teasel,
Dipsacus fullonum Linn., June 7, 1916, the red valerian or
Jupiter’s beard, Centranthus ruber DC., June 23, 1916, at
Berkeley, California, and on wild rose. Rosa californica C. & 8.,
at Rutherford, California, June 21, 1916.

Macrosiphum solanifolii (Ashmead). A large pale yellow or
green aphid attacking a large number of plants. On the campus
it was collected on yarrow, Achillea millefolium Linn., March 29,
1916, and on tobira or Japanese pittosporum, Pittosporum tobira
Ait., July 5, 1916. Also taken on the tender shoots of the
American, white or water elm, Ulmus americanus Linn., in Golden
Gate Park, San Francisco, California, May 25, 1916.

Macrosiphum stanleyi Wilson. A beautiful large yellow and
black species abundant on the undersides of the leaves of the red
elderberry, Sambucus racemosa callicarpa Jepson, on the campus
throughout the spring, summer and fall months. It was taken
in fewer numbers on the tender growth of the blue elderberry,
S. glauca Nutt., growing along the creek in shady places on the
campus, May 15, 1916.

Idiopterus nephrelepidis Davis. A small pitch black species
with whitish legs and black clouded areas in the wings. It is
becoming of considerable economic importance as a pest to green-
house and household ferns. During the past year it has been
observed to be quite injurious to the common and improved
Boston ferns, Nephrolepis, spp., as well as to the holly fern,
Cyrtomium falcatum J. Smith.

Amphorophora latysiphon Davidson. <A large green and black
species with black conspicuously swollen cornicles. Large num-
bers of apterous and a single winged individual were taken on
the crowns at and under the surface of the ground of Shasta
daisies, Chrysanthemum, sp., Berkeley, California, August 15,
1916, and on the tender opening buds of the common garden
cowslip, Primula, sp., December 17, 1916, at the same locality.

Amphorophora rubi (Kalt.). A large green aphid attacking
the terminal shoots of the cultivated blackberry, Rubus, sp., and

330 University of California Publications [ENTomMoLocy

wild blackberry, R. vitifolius C. & S., Berkeley, California,
March 25, and June 6, 1916.

Toxoptera aurantiae Fonsc. A small black species attacking
the leaves and terminal shoots of lemon, Martinez, California,
June 12, 1916, and camellia, Camellia japonica Linn., in a lath-
house at Oakland, California, July 25, 1916.

Phorodon humuli (Sehrank)
(fig. 17). A pale yellowish green
aphid which during the past season
was first observed on the tender
leaves of plum on the campus,
April 7, 1916. At that time only
apterous females oceurred, but on
April 14 a great many winged
migrants were present and by
April 25 practically all were
winged. The species was also
abundant on plum in Golden Gate
Park, San Francisco, California,
May 25,1916. By the middle of the
summer all had become winged and
left the plum trees. Later in Sep-
tember specimens were taken on
hops.

Rhopalosiphum arbuti David-
son. This is a fairly large green

and black species occurring com-
eer Ree monly on madrone, Arbutus men-
plum leaf, Berkeley, Cal., April zeisit Pursh. on the campus. Other
ie eae ABR trees infested are: the strawberry
tree Arbutus unedo Linn., campus,
May 18, 1916; the tender tips of the mountain holly or Christmas
berry, Photinma arbutifolia Lindl., campus, May 15, 1916. It
was also collected on the strawberry tree in Golden Gate Park,
San Francisco, California, June 5, 1916. E. P. Van Duzee of
the California Academy of Sciences collected it on the common
manzanita, Arctostaphylos manzanita Parry, at Hoberg’s Resort,
Lake County, California, August 2, 1916. This is undoubtedly
Rhophalosiphum nervatum Gillette.
Rhopalosiphum howardi Wilson. A medium-sized, shiny
dark red or almost black species associated with Macrosiphum

Vow. 1] Essig.—Aphididae of California 3a

granarium (Kirby) on the leaves, stems and in the heads of oats
at Martinez, California, June 12, 1916. It was later taken in
the heads of barley at Rutherford, California, June 21, 1916.
This is very close if not synonymous with Rhopalosiphum rhois
Monell.

Rhopalosiphum lactucae (Kalt.). A normal-sized green and
black aphid first taken by the writer on the common sow thistle,
Sonchus oleraceus Linn., at Santa Paula, Califormia, April 3,
1916, and erroneously labeled Rhopalosiphum persicae (Sulzer).
Last year it was collected on the same host plant at Berkeley,
California, June 28, 1916, and at San José, California, July 14,
1916

Rhopalosiphum nervatum Gillette. A green species smaller
than the common rose aphis, Macrosiphum rosae (linn.), and
distinguished from it by the darker wing veins and swollen
ecornicles. Occurs throughout the year and collected, February
20, 1916. (See R. arbuti Davidson).

Rhopalosiphum nymphaeae (Linn.) (fig. 18). A large dark-
red plant louse covered with a fine white pulverulence or often
a shiny dark green form as appears in the late summer. Winged
and apterous viviparous females, all pink or dark-red, were first
taken on tender twigs of a plum tree on the campus, April 7,
1916, where they occurred in large, compact colonies. The
species, the same color phase, was also taken on plum in
Golden Gate Park, San Franciseo, California, May 25, 1916. By
the middle of June none could be found on the plum in either
locality, but on June 21, 1916, the dark green and shining phase
was very abundant on water plantain, Alisma plantago Linn.,
and pondweed, Potamogeton natans Linn., at Rutherford, Cal.

Rhopalosiphum persicae (Sulzer). This is a very common
green species with a conspicuous black patch in the middle of the
abdomen of the winged form. It is an omnivorous feeder, attack-
ing a very large number of plants. Among those observed during
the past year were: Lavatera assurgentiflora Kellogg, eampus,
June 27, 1916; Clarkia elegans Dougl., campus, March 29, 1916;
Pentstemon spectabilis Thurb., campus, Mareh 29, 1916; Cali-
fornia laurel or pepperwood, Umbellularia californica Nutt.,
campus, April 16, 1916; nasturtium, Hayward, California, April
19, 1916 (coll. by Roy E. Campbell) ; tulip tree, Liriodendron
tulipifera (Linn.), campus, April 24, 1916; tobira or Japanese
pittosporum, Pittosporum tobira Ait., campus, July 5, 1916;

332 CO University of California Publications [ENToMoLoGy

tarata, Pittosporum eugenioides A. Cunn., campus, July 5, 1916;
Victorian box, Pittosporum undulatum Vent., campus, July 5,
1916; Baccharis douglas DC., Berkeley, California, June 17,
1916, red valerian or Jupiter’s beard, Centranthus ruber DC.,

Fig. 18. Rhopalosiphum nymphaeae (inn.). Colony on twig of plum.
One mature apterous viviparous female and the remainder nymphs.
Berkeley, Cal., April 18, 1916. (Original. Photo by Div. Sci. Ilust.,
Univ. Calif.)

Berkeley, California, June 23, 1916; the fullers’ teasel, Dipsacus
fullonum Linn., campus, June 27, 1916, Escallonia pulverulenta
Pers., campus, June 27, 1916; Melianthus major Linn., campus,
June 27, 1916, and sunflower, Helianthus annuus Linn., Berkeley,

California, August 15, 1916.

|
|
|

Vou. 1]

Fic. 19.

Essig.—Aphididae of California 330

Myzus cerasi (Fab.) on the underside of a cherry leaf and

curled leaves caused by the attacks. Berkeley, Cal. April 25, 1916.

(Original.

Photo by Div. Sci. Ilust., Univ. Calif.)

334 University of Califorma Publications [ENtomoLocy

Rhopalosiphum rhois Monell. A dark green or black shiny
aphid occurring abundantly on the tender shoots of the poison
oak, Rhus diversiloba T. & G., during the spring and early sum-
mer, but entirely disappearing by the middle of summer. Often

Fig. 20. Myzus varians Davidson. Curled leaves of tender peach shoot
caused by this insect. Berkeley, Cal., June 7, 1916.

causes severe curling of the leaves. Campus, July 3, 1916—only
a very few were to be found at this date. (See R. howardiu
Wilson).

Myzus cerasi (Fab.) (fig. 19). A shiny black aphis with long
cornicles, occurring in large dense colonies on the shoots and

Vow. 1] Essig.—Aphididae of California 335

young fruit of cherries and forming large curled pseudogalls on
the former. Very common in this district and often the cause of
much worry to orchardists. Collected at Berkeley, California,
April 25 and August 21, 1916. Disappears soon after the last
date.

Myzus circumflecum (Buckton). This is a very pretty yellow
or pale green and black species which is a common greenhouse
pest infesting maidenhair, asparagus and holly ferns and a great
many other plants already listed by other writers. During the
past year the species was taken on the tender shoots of the Cali-
fornia laurel or pepperwood, Umbellularia californica Nutt.,
campus, April 16, 1916, on the flower heads of Pentstemon specta-
bilis Thurb., campus, March 29, 1916, and on the lower leaves of
both of the show lihes, Liliwm elegans grandiflorum and L.
speciosum rubrum growing in a lath-house at Oakland, Califor-
nia, July 25, 1916.

Myzus fragaefolii Cockerell. A small pale yellow aphid in-
festing the undersides of the leaves of the cultivated strawberry
and at times causing considerable smutting of the foliage. Taken
at Niles, California, June 16, 1916.

Myzus varians Davidson (fig. 20). A pale yellow or green
and black species somewhat resembling Rhopalosiphum persicae
(Sulzer) which occurs in large colonies on the host plants and
often causes much smutting of the foliage. Taken throughout
the spring and summer on clematis, Clematis brachiata Thurb.,
on the campus and appear to reach their maximum development
when collected, June 9th to the 29th, 1916. It was also taken
July 7, 1916, at Berkeley infesting the terminal shoots of peach,
causing the leaves to curl and become reddened much as if in-
fected with peach curl. Within the pseudogalls thus formed were
great numbers of the aphids; one or two trees were severely
injured by the attacks.

Pentalonia nigronervosa Coquerel. A very interesting small
black species greatly resembling [diopterus nephrelepidis Davis.
Specimens were received from Mr. E. M. Ehrhorn, who collected
it on ferns at Honolulu, H. T., in Mareh, 1916. This insect does
not oceur in California, but might be introduced or may have
been, and could be easily mistaken for the other species which it
resembles. It has previously only been reported on banana.

Hyalopterus arundinis (Fab.) (fig. 21). An average sized,
pale green and black species with the bodies usually covered with

336 University of California Publications [ENTomMoLocy

quite thick whitish pulverulence and occurring in dense colonies
on the leaves of the young growth of prune and plum trees. It
was also taken in considerable numbers on apricot trees at Niles,
California, July 11, 1916. The Japanese varieties of plums
appear to be immune to their attacks as was exemplified last year
at Martinez, California, where a few prune trees growing in a
thrifty young orchard of Burbank plums were very seriously
attacked, while not a single infestation could be found on any
of the plum trees. It is also of interest to know that the cracking,
usually a cross, at the ends of prunes in the Santa Clara Valley

Fic. 21. Hyaiopterus arundinis (Fab.) Prunes showing the character-
istic splitting at the ends as a result of the presence of this insect on
the foliage. Morgan Hill, Cal., July 14, 1916. (Original.)

has been definitely found to be due to the attacks of this insect,
according to Horticultural Commissioner Earl Morris, and the
annual loss occasioned is sometimes considerable.

Aphis albipes Oestlund (fig. 22). The apterous viviparous
females are dark with whitish pulverulence arranged in definite
patterns on the back as shown in the illustration. The apterous
forms (fundatrix) were taken on the shoots of the snow berry,
Symphoricarpos racemosus Michx., on the campus, March 25,
1916. Great numbers of both winged and apterous viviparous
females were taken from the terminal shoots of the same plant
on May 15, 1916. The leaves are often curled in dense irregular
masses and form splendid protection for the insects. The species
was determined by John J. Davis.

Vou. 1] Essig —Aphididae of California 330

Aphis avenae Fab. A small dark green species abundant on
corn, grain and grasses on the campus. Collected March, 1916.

Aphis bakert Cowen. <A small pale yellow or pinkish and
black species taken sparingly on the flower heads and at the
bases of the leaves of red clover, Trifolium pratense Linn.,

Fig. 22. Aphis albipes Oestlund. Apterous females on the leaf of snow
berry, Symphoricarpos racemosus Michx. Campus, March 29, 1916.
(Original. Photo by Div. Sci. Hlust., Univ. Calif.)

campus, June 29, and July 25, 1916. Not abundant here at any
season.

Aphis brassicae Linn. (fig. 23). A pale green species covered
with whitish pulverulence and feeding in compact colonies on the
common cruciferous plants of the state. Especially injurious to
cabbage, cauliflower, brussels sprouts, kale, mustard, radish,
turnips, ete. Present throughout the year.

338 University of California Publications [ENToMoLocy

Aphis cardui Linn. (fig. 24). <A fairly large green and black,
shiny aphid. The apterous fundatrix occurred in small colonies
on the tips of tender plum suckers on the campus, April 10, 15,
24, 25,1916. It is very common throughout the summer in great
colonies entirely covering all parts of the bull thistle, Cirsiwm
lanceolatum Linn.

Aphis ceanotht Clark (syn. A. ceanothi-hirsutt Essig). A
small dull reddish and black species which becomes very abundant

Fic. 23. Aphis brassicae Linn. A winter colony on the leaf of brussels
sprouts. Berkeley, Cal., Nov. 6, 1916. (Original. Photo by Div. Sci.
Illust., Univ. Calif.)

on various species of Ceanothus throughout the state. Taken at
Berkeley and on the campus infesting Ceanothus thrysiflorus
Esch. and soapbush, Noltea africana Reichb., June 2, 1916. Also
very common on soapbush in Golden Gate Park, San Francisco,
California, May 25, 1916.

Aphis gossypii Glover. A small dark green or black aphid
usually feeding in dense colonies. Abundant on the creek milk-
weed, Asclepias speciosa Torr., and a few on cascara sagrada,
Rhamnus purshiana DC., on the campus, July 3 and 5, 1916.
Also taken on show lily, Lilium speciosum rubrum, Oakland,

ore eRe

Vou. 1] Essig.—Aphididae of California 339

California, July 5, 1916; on woolly milkweed, Asclepias vestita
H. & A., Visalia, California, July 19, 1916, by W. P. Baird, and
on honeysuckle, Lonicera, sp., and catalpa, Catalpa, sp., at St.
Helena, California, June 18, 1916, by C. W. Woodworth.

Aphis herderae Kalt.? A small dull black species occurring
in compact colonies on English ivy, Hedera helix Linn., and a
number of other plants is certainly not Aphis rumicis Linn., so
is provisionally listed as this species. Abundant at Berkeley,
California, and collected June 2, 1916.

Aphis heliantht Monell is a pale or dark green species with
black markings on the winged form and occurs abundantly on
the undersides of the leaves of the common and wild sunflower,

{

Fic. 24. Aphis cardui Linn. Apterous viviparous females taken from
plum tree, Campus, April 10, 1916. (Original. Photo by Div. Sci. Ilust.,
Univ. Calif.)

Helianthus annuus Linn. Unreeorded specimens first taken by
the writer at Pomona, California, in 1909.

Aphis heraclet Cowen.’® A medium dark species with greenish
or reddish mottlings on the dorsum. Occurs abundantly on the
old yellow leaves of Heracleum mantegazzianum Sommier &
Levier in the botanical garden on the campus, June 9, 1916.

Aphis malifoliae Fitch (syn. Aphis sorbi Kalt. for here). A
common rosy and black species also slightly covered with a
whitish pulverulence. Occurs on the young fruit and curls the
leaves of apple. Often a serious orchard pest.- Collected at
Berkeley, California, April 28, 1916, by E. L. Prizer.

10T am indebted to Mr. John J. Davis for the determination of this

species and for the information that the specific name is also preoccupied
by Aphis heraclei Koch, thus necessitating a new name for the above.

340 University of California Publications [ENTomoLocy

Aphis maidis Fitch. A small dark green aphid received from
Bend, California, July 6, 1916, seriously infesting milo maize.
The common species on corn throughout the state, however, is
Aphis avenae Fab.

Aphis medicaginis Koch (fig. 25). The adults are shiny black
with conspicuous whitish legs. Specimens were taken by the
writer at Moorpark, California, August 11, 1911, where they were
doing great damage to the foliage and pods of the blackeye bean,

Fie. 25. Aphis medicaginis Koch. Nymphs and adults on lemon leaf.
Sacramento, Cal., 1913. (Original.)

Vigna catjang Walpers. It was also taken on the leaves of
lemon, Capitol Park, Sacramento, California, July, 1913.

Aphis middleton Thos. A pale green and pulverulent species
first taken on the roots of rough pigweed, Amaranthus retroflexrus
Linn., and the roots of asters, Santa Paula, California, October
20, Onde

Aphis neru Fonse. A pretty dark yellow and black species
common on the oleander during most of the year and also on
milkweed during the summer. On the campus it attacks oleander
and is specially abundant during the summer months on the

Vou. 1] Essig. —Aphididae of California 341

ereek milkweed, Asclepias speciosa Torr. At Niles an oleander
tree was seriously infested July 28, 1916. It was also taken on
woolly milkweed, A. vestita H. & A., at Visaha, California, July
27, 1916, by W. P. Baird. This is without doubt the common
milkweed species deseribed by Montell as Aphis lutescens and
occurs generally throughout the state.

Aphis oenotherae Oestlund. A small dark green, somewhat
pulverulent species occurring in large numbers among the buds
and on the flower heads of the evening primrose. Taken on
Oenothera biennis Linn. on the campus, June 7, 1916, and on
the common evening primrose, O. grandiflora Ait., and on E'pi-
lobium adenocaulon occidentale Trelease in Golden Gate Park,
San Francisco, California, May 25, 1916.

Aphis persicae-niger Smith. A small shiny black species often
abundant on the tender shoots of peach trees. During the past
few years it has been received from a number of localities as
follows: Peach, Berkeley, California, April 29, 1915; peach,
Dinuba, California, April 30, 1915; peach, Yuba City, California,
March 25, 1916 (H. P. Stabler) ; peach, Fresno, California, May
10, 1916; peach, Ventura, California, July 17, 1916 (S. H.
Essig). In the latter case the infestation was severe and much
of the fruit was also infested. This insect, though distributed
throughout much of the state, has never become a serious orchard
pest and only abundant on rare occasions.

Aphis pomi DeGeer. This small green and black aphid is a
serious pest to a number of valuable ornamental trees in this
section, notably the English hawthorn, Crataegus oxryacantha
Linn. At Berkeley it has also been taken on the following:
Crataegus spp., laurestinus, Viburnum tinus Linn., tarata, Pittos-
porum eugenioides A. Cunn., tawhiwhi, P. tenuifolium Gaertn.,
Cotoneaster franchetw Bois., apple and pear, as well as a number
of other undetermined plants. The foliage is covered with
honey-dew and soon collects dust or becomes covered with the
black smut fungus so as to assume a most undesirable appearance.

Aphis rumicis Linn. <A dull black species occurring in dense
colonies and generally confused with another species which I
am calling Aphis hederae Kalt. What, according to Davis,
is true A. rumicis has been taken at Berkeley, California, on the
Western dock, Rumex occidentalis Wats, April 13, 1916, and on
other species of Rumer and on the common garden. poppy,
Papaver, sp., June 15, 1916.

342 University of California Publications [ENToMoLocy

Aphis salicicola (Thomas). <A pale green species with bright
orange-colored males. Very abundant on the tender tips of the
arroyo willow, Salix lasiolepis Benth., Rutherford, California,
June 21, 1916. The species occurred on the same plant through-
out the summer and fall at Berkeley and on September 20, 1916,
the sexual males and females were also abundant and mating
freely.

Aphis sambucifoliae Fitch. The winged and apterous vivi-
parous females are dull black while the sexuparae are amber or
dull brownish. The former occur throughout the summer on
the blue elderberry, Sambucus glauca Nutt. The sexuparae were
taken in few numbers on the leaves of the red elderberry, S.
racemosa callicarpa Jepson, on the campus, September 21, 1916.

Myzaphis abietina (Walker). A very small dark green aphid
common on the smaller branches of the Norway spruce, Picea
excelsa Link, in Golden Gate Park, San Francisco, California,
May 25, 1916.

Myzaphis rosarum (Kalt.). A small dark green species often
very abundant on the leaves of roses and usually the most serious
rose pest, causing severe smutting of the foliage. Taken at
Berkeley, California, June 16, 1916, and throughout the summer
and fall.

Siphocoryne capreae (Fab.). The apterous forms are pale
green and the winged forms pale green and black. Common here
on willows, including the arroyo willow, Salix lasiolepis Benth.,
campus, June 13,1916. Also abundant on the wild or sweet anise,
Carum kelloggu Gray, at St. Helena, California, June 18, 1916
(C. W. Woodworth) and at Berkeley, California, June 20, 1916.
It is the commonest aphid attacking the latter.

Siphocoryne xylostet Schrank. <A dull green species often
with thin coating of whitish pulverulence. Common on the tender
shoots of the honeysuckle, Lonicera, spp. Taken at Berkeley,
California, July 20, 1916, by Miss A. Williamson.

Tiosomaphis berberidis (Kalt.) A small yellow or pale green
aphid often very abundant on various species of the barberry
and causing considerable smutting. Taken on Berberis vulgaris
Linn., Golden Gate Park, San Francisco, California, June 5,
1916; on Berberis, sp., campus, June 7, 1916, and on B. vulgaris
Linn., Niles, California, September 30, 1916.

Cerataphis lataniae (Boisduval) (fig. 26). This is a very
interesting plant louse and more often taken for a scale insect or

Vou. 1] Essig —Aphididae of California 343

the immature form of a white fly than for an aphid. The young
apterous viviparous females are dark or pale brown and when
mature acquire a white fringe around the margin as shown in
the illustration (fig. 26). The winged forms are dull brownish
or blackish in color. The species occurs in greenhouses through-

Fie. 26. Cerataphis lataniae (Boisduval). Apterous viviparous females
on leaf of orchid. Oakland, Cal., July 25, 1916. (Original.)

out this district and sometimes becomes a pest to orchids. Taken
on orchids, Oakland, California, July 25, 1916.

Pemphigus populitransversus Riley (figs. 27 and 28). The
fundatrix is pale amber or yellowish with tufts of white cottony
material arranged in rows on the dorsum. The virgogeniae are
black with more or less cottony material at the posterior end of

344 University of California Publications [ENtomoLocy

Fic. 27. Pemphigus populitransversus Riley. Galls formed on the leaf
petioles of poplar. Berkeley, Cal., Aug. 17, 1916. (Original.)

Fig. 28. Pemphigus populitransversus Riley. Stem-mother or funda-
trix, and virgogeniae. Campus, Aug. 17, 1916. (Original.)

Vou. 1] Essig.—Aphididae of California 345

the abdomen. Forms large galls on the leaf petioles, near the
leaves, of poplar. Campus, August 15, 1916.

Eriosoma, sp.? (fig. 29). An undetermined species designated
by the common term Wilson’s bonnet gall-louse, by Patch.

Fig. 29. Eriosoma, sp. Large pseudogall produced on the leaf of
English elm, Ulmus campestris Smith. Berkeley, Cal. July 8, 1916.
(Original. )

Fig. 30. Eriosoma lanigera (Hausm.) Colony of the woolly apple aphis
on apple. (Author’s illust., Ben. & Inj. Ins. Cal.)

346 University of California Publications [ENtTomoLocy

The winged forms are dark and often very numerous in large
pseudogalls on the leaves of the English elm, Ulmus campestris
Smith, at Berkeley, California, July 8, and Hayward, California,
July 14, 1916.

Eriosoma lanigera (Hausm.) (fig. 30). A dull reddish species
with the bodies entirely covered with long white woolly material
giving rise to the common name woolly aphis. A very common
species throughout the state on apple and rarely on pears.
Sexuparae begin to appear the last of September on the campus,
but so far I have not succeeded in taking the sexuales.

Chermes cooleyi Gillette. A very small pinkish or dark,
nearly globular plant louse occurring on the needles of various
conifers and usually covered with white cottony material and
appearing as small white specks on the trees. Sometimes very
abundant and injurious. Taken on the campus on Monterey
pine and other conifers, March 29, 1916.

Transmitted January 8, 1917.

es SRE a % pate Sank P rex v palleres CN a age
a 3 ; BA :
. "UNIVERSITY OF CALIFORNIA PRESS TUE Go Bs

ae BERKELEY ’ Cie Rui ly dy Pee

> 3. { er

Ps i Me A

Si

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ENTOMOLOGY.—C. W. Woodworth, Editor. ie 14
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Pp. 541-394; plates: 5-15. January, VOUT ios cic ee ee OU
12. Certain Effects under Irrigation of Copper Compounds upon Crops, by 00

R. H. Forbes. Pp. 395-494, plates 6-9. April, 1917 2220 v
_18, Experiments on the Effects of Constituents of Solid Smelter Wastes on

Barley Growth in Pot Cultures, by C. B. Zipman and W. F. Gericke,

PP. AOG-BS Te: WaT CD Sy LOL Gis il ccty ket agacadihntite hin basSfibhonsanbr stv enkehalabt nas Midateath '

UNIVERSITY OF CALIFORNIA PUBLICATIONS

TECHNICAL BULLETINS
COLLEGE OF AGRICULTURE, AGRICULTURAL EXPERIMENT STATION

ENTOMOLOGY

Vol. 1, No. 8, pp. 347-367 March 28, 1918

NEW GENERA AND SPECIES OF ENCYR-
TINAE FROM CALIFORNIA PARASITIC .
IN MEALYBUGS (HYMENOPTERA)

BY

P. H. TIMBERLAKE

Hawaiian Sugar Planters’ Experiment Station

The material which forms the basis of this paper has been
assembled during the last three or four years, mainly through
the efforts of Mr. C. P. Clausen while engaged in a study of
Pseudococcus and related genera in California. In all, three new
genera and six new species are described, the types of which will
be deposited ultimately in the United States National Museum.
Notes on two foreign or introduced species under Tanaomastixr
are included to complete the generic conception.

Tribe MIRINI
Acerophagus' Emily Smith

Of the three species included in the material two are new and
all belong to the A. texanus group, the five species of which now
known may be separated as follows:

FEMALES

ip Wines wibhOUL a CONSPLCMOUS: CTOSS DANG 2... --21 osccc ac 2tceceeen ee secneeceee see 2

Wings with a smoky band across the disk ~......... A. fasciipennis, n. sp.

2. Frontovertex about one-third to three-fourths longer than wide ~........3
Frontovertex about twice as long as wide, the ocelli in an angle of

| FIGUT ESE eteed te eet ois SN Soll ct A. notativentris (Girault)
1 For a recent revision of this genus see Proc. U. 8. Nat. Mus., vol. 50,
pp. 574-79. The antennal club of A. tewanus (Howard), represented as

solid in figure 52, plate 31 (loc. cit.), in reality is composed of three closely
fused joints except in the male.

a

348 University of California Publications. |ENtTomMoLoey

Go

3. Ocelli small, the posterior pair their own diameter from eye-margin,
coloration! pale! 225 ee ee ae eee ee ee ea 4
Ocelli large, the posterior pair close to eye-margin, coloration vivid
orange syellow ees eee eae es eres ee eee A. texanus (Howard)

4. Frontovertex about three-fourths longer than wide, the ocelli in a right
angle, coloration chrome lemon yellow .................... A. pallidus, n. sp.
Frontovertex wider, about one-half longer than wide, the ocelli in
slightly more than a right angle, coloration pale cadmium yellow

Se Ss nS cea cs Sona ed ee SS A A. gutierreziae Timberlake

MALES
1. Wangs: hyaline: “throughout 25... e lo as ee 2
Wings with a smoky band across the disk ............ A. fasciipennis, n. sp.
2. Coloration pale yellow, the abdomen usually more vivid than thorax __... 3

Coloration orange yellow, the abdomen concolorous .............---.--.--0-00-----
Se Se eee DY See SE ec ane OPES Nebel RO ed a A. texanus (Howard)

3. Abdomen without a dark blotch behind the vibrissae ~.................----.----- +
Abdomen with a conspicuous blackish brown blotch behind the vibrissae
eee ee Ree dee eee ere A. notativentris (Girault)

4. Thorax and abdomen pale cadmium yellow......A. gutierreziae Timberlake
Thorax gamboge yellow, the abdomen more orange yellow especially at
apex and with two dusky transverse bands across the middle -.........

REE PAR Aer SN et eA Ee Dec bk eT Ae Re a Sl pe aD A. pallidus,-n. sp.

Acerophagus fasciipennis, n. sp.
FEMALE

Frontovertex about one-half longer than wide; the ocelli in
a right-angled triangle, the median one situated considerably
nearer the occipital margin than in A. gutierreziae, or very no-
ticeably behind the center of the frontovertex, the posterior pair
less than their own diameter from the eye-margin and about twice
as far removed from the occipital margin; eyes not pubescent ;
antennae practically as in the other species of the group, except
that the scape and club are proportionately considerably shorter
(see table of measurements below) ; wings nearly uniformly eili-
ated, the speculum? extremely narrow above or practically ob-
literated, abruptly widening below and connecting with a branch
of the hairless streak on the posterior margin; ovipositor slightly
longer than in A. gutierreziae, or about two-fifths the length of
abdomen, its sheaths pubescent. Length, exclusive of ovipositor,
0.75 mm.

Coloration a little paler than in A. gutierreziae or more of a
elay yellow, the underparts still paler, but the abdomen decidedly
brighter or more orange yellow; appendages concolorous with

2 The term adopted here is one used by Thomson for the oblique hair-
less streak.

Vou. 1] Timberlake.—New Genera and Species of Encyrtinae 349

underparts, the antennae except club, however, a little deeper

yellow than legs; the ovipositor black at apex; wings with a

conspicuous tegumentary smoky band across the disk, its distal

margin opposite the end of the stigmal vein and shghtly convex,

the basal fourth of wing with a brown dot on lower margin at

the very base; hind wings also slightly smoky across the middle.
MALE

Frontovertex about one-third longer than wide, the anterior
ocellus at the center; otherwise structurally about as in female.
The coloration slightly paler, more of a chrome lemon yellow, the
abdomen orange yellow, and the wings banded as in the female.
Length, 0.8 mm.

Deseribed from one female, one male (type and allotype)
reared from early larval stages of Pseudococcus craww (Coquil-
lett), Uplands, California, November, 1914 (C. P. Clausen),
Clausen no. 53.

Acerophagus notativentris (Girault)
Inifes al
Pseudaphycus notativentris Girault, Descrip. Hym. Chaleid. variorum
cum Observ., vol. 3, p. 10, Glendale, Md., May, 1917.

FEMALE
Frontovertex nearly twice as long as wide, finely granulose
about as in A. teranus and minutely punctulate; ocelli forming
an angle of about 85°, the anterior one a little behind the center

Fig. 1. Acerophagus notativentris: A, antenna of female; B, right
mandible of female, exterio-dorsal view; C, same, anterior view.

300 University of California Publications. [#NToMoLoGy

of the frontovertex, the posterior pair about their own diameter
from the occipital margin and less than half as far from the eye-
margin; face considerably shorter than length of eyes, the scrobes
as in the other species; eyes with a rather thick, but fine, short
pubescence ; funicle joints of antennae (fig. 1, A) proportionately
a little wider for their length than in other species; dorsal tooth
of mandible (fig. 1, B and C) much smaller and more basal than
usually; wings uniformly ciliated, the speculum narrow, not
interrupted and reaching nearly to the posterior margin; ovi-
positor protruded about one-fourth the length of abdomen, its
sheaths pubescent. Length, 0.5-0.8 mm., exclusive of ovipositor.

Coloration about cadmium yellow, the face, cheeks, and under-
parts a little paler; a transverse spot on concealed part of
pronotum blackish; the metanotum, but not propodeum, slightly
brownish ; a spot on sides of abdomen before the vibrissal plates,
the vibrissal plates and the margin of the segments in the center
of the dorsum of abdomen brown; antennae dusky yellowish,
the scape somewhat purer yellow, the club more whitish; coxae
and femora yellowish white, the tibiae and tarsi more yellowish
with apex of last tarsal joint dusky; exserted portion of ovi-
positor dusky; wings hyaline, the veins pale yellowish; pubes-
cence of head and thorax dusky.

MALE

Similar to the female in all characters; the frontovertex a
trifle wider, the ocelli more nearly in a right angle; the abdomen
a little shorter than thorax instead of as long. Coloration some-
what brighter or more of a chrome lemon yellow, the face and
underparts more vivid, and the dorsum of abdomen behind the
vibrissal plates blackish brown except the sides. Length, 0.42-
0.6 mm.

Redescribed from thirteen females, nine males, reared Octo-
ber, 1914, and during the summer of 1915, from a Pseudococcus
species on grape, Fresno, California (H. 8S. Smith), California
State Insectary no. 675, also compared with Girault’s types.

Acerophagus pallidus, n. sp.
Fig. 2
FEMALE
Frontovertex about three-fourths longer than wide, about as

closely granulose as in A. texanus or almost punctulate; ocelli
in a right-angled triangle, the anterior one noticeably behind the

Vou. 1} Timberlake-——New Genera and Species of Encyrtinae 351

center of the frontovertex, the posterior pair nearly their own
diameter from eye-margin and twice as far removed from occip-
ital margin; face a lttle shorter than length of eyes, the head
being a little thinner anteroposteriorly than in teranus or
gutierreziae, the scrobes about the same; eyes with a very fine,
short, sparse pubescence; antennae (fig. 2, A) about as in other
species of the teranus group, the pedicel, however, proportion-
ately longer and slenderer, or a little longer than the first four
funicle joints combined; mandibles (fig. 2, B and C) normal;
wings uniformly ciliated, the speculum somewhat wider below
and nearly connecting with the hairless streak on the posterior
margin; ovipositor about one-third the length of abdomen, its
sheaths pubescent. Length, 0.9 mm., exclusive of ovipositor.
Coloration almost uniformly pale yellow, about chrome lemon,
the appendages concolorous, with apex of ovipositor black ; wings
hyaline, the veins pale yellowish; pubescence on head and body
about as abundant as in the other species of the group, but not
conspicuous except on face, where it is whitish or silvery.

MALE

Similar to the female, but the following differences may be
noted: Frontovertex slightly wider or about one-half longer
than wide, the ocelli in a slightly acute or almost equilateral
triangle. Coloration the same, except that the dorsum of the
abdomen at apex is orange yellow, separated from the paler base
by a narrow, transversely arcuate dusky band between the
vibrissae. Length, 0.7 mm.

Deseribed from two females, two males (type, allotype and
paratypes a and 0) reared from Pseudococcus yerbasantae Essig,
May 12, 1915, Sespe Cafion, California (C. P. Clausen), Clausen
no. 67.

TABLE OF FEMALE ANTENNAL MEASUREMENTS (LENGTH X WIDTH) OF
ACEROPHAGUS IN MICRONS

eed Pedicel Funicle Funicle Funicle Funicle Funicle Club

radicle 1 2 3 4 5
. fasctipennis 141x30 60x20 15x16 12x19 12x20 16x23 20x28 105x38
. pallidus 180x36 82x25 18x18 18x20 18x20 18x25 24x30 129x54
. gulierreziae TiGeR, Sys; ails) IPA) TIS) es ee
.notativentris 157x388 67x30 12x20 12x23 15x28 20x33 23x38 136x61
. texanus 159x386 64x28 15x23 18x25 20x28 20x31 20x38 136x59

302 University of California Publications. [ENToMoLoGyY

Stemmatosteres,* n. gen.
FEMALE

Head very thin anteroposteriorly, a little wider than thorax, |
the greater portion of frontovertex and face lying in one plane,
which is very shghtly convex; seen from in front, the outline is
rounded, the occipital margin appearing truneate, the length
and width about equal; in lateral view (fig. 3, B) the outline is
narrowly and irregularly oval, the greatest thickness just below
the eyes; in dorsal view (with head held vertically) only the
vertex visible, sloping gently backward to the occipital margin,
which is abruptly rounded, not sharp, its outline now appearing
concave; occiput with a rather shallow, semicircular excavation
on the dorsal half into which the prothorax fits, the neck being

Fig. 2. Acerophagus pallidus: A, antenna of female; B, right mandible
of male, exterio-dorsal view; C, same, anterior view.

inserted near the dorsal margin of the excavation, so that the
head may be held either vertically or horizontally, the former
position apparently being the normal or resting position in life;
frontovertex considerably wider than long, the oeelli absent, the
eyes small, oval, coarsely faceted; face abruptly rounded off to
the oral margin from just above the antennal sockets, the scrobes
entirely absent; cheeks a little longer than length of the eyes;
antennae (fig. 3, C) inserted rather far apart and close to the

3 From oreupatoorepys, deprived of stemmata, in reference to the lack
of ocelli.

Vou. 1] Timberlake—New Genera and Species of Encyrtinae 353

oral margin, a little less than half as long as body, distinctly
clavate; the scape slender, compressed ; pedicel as long as the
first three funicle joints; the funicle five-jointed, all joints short,

increasingly transverse, the fifth nearly twice as wide as long;

club large, oval, not much wider than the last funicle joint and
as long as funicle and pedicel combined, composed of two joints,
the basal one being much shorter; mandibles (fig. 38, E) with
three long, acute teeth, the ventral one far more basal, the upper
two at the apex, with the middle one slightly longer. Thorax
small, the visible portion of dorsum hardly more than one-half
as long as abdomen; pronotum very large, fully as long as the
mesoscutum and seutellum combined ; mesoscutum very short and
transverse, about four times wider than long; seutellum also short
and transverse, a little longer than the scutum, the axillae coa-
lesced with it, the sutures not at all visible unless insect is
mounted in balsam; metathorax and propodeum concealed by
the base of the abdomen, which overlaps the posterior part of
the thorax so that the apex of the first abdominal segment and
the posterior margin of the mesopleura le in the same vertical
plane; wings either absent or vestigial, the front pair visible in
some specimens as minute, linear scales, no longer than the scu-
tellum, bearing two bristles at apex and one on the side, hind
pair of wings apparently entirely absent; legs short, normal, the
middle tibial spur slender, tapering, a little shorter than the first
tarsal joint, hind tibiae with two spurs, the smaller one half as
long as the other. Abdomen large, oval, depressed, the first seg-
ment shortest, the second longest ; ovipositor shortly protruded,
otherwise enclosed by the fifth sternite, which reaches to the apex
of the abdomen but does not become at all compressed ; vibrissal
plates situated at the apical fourth, the vibrissae reaching about
to the apex. Surface of head and body nearly smooth, or very
finely alutaceous, the frontovertex with extremely minute, shal-
low, scattered setiferous punctures; pubescence short, very
delicate, pale colored, and hardly visible except under high
magnification ; coloration yellow, non-metallic, but with a pearly
luster.

MALE

Much like the female, except that the abdomen is consider-
ably shorter and wider, the antennal scape Gig So Dimas latcle
wider and the club considerably longer and solid.

354 University of California Publications. [ENToMoLoGy

TYPE OF GENUS
Stemmatosteres apterus, n. sp.

This genus is anomalous in the absence of ocelli, in the strue-
ture of the antennal club, the reduction and pecuhar configura-
tion of the thoracic sclerites and the overlapping of the abdomen
upon the posterior part of the thorax. The wings are probably
never fully developed on account of the reduction of the mesono-
tum. The genus appears to be most closely allied to Acerophagus
Smith.

Stemmatosteres apterus, n. sp.
Fig. 3
FEMALE

Head and body chrome lemon yellow, the dorsal surface of
the head with a pearly luster (in life) which reappears to a less
degree on other parts of the body; antennae dusky yellowish,
purer yellow at the base of the scape; legs concolorous with body,
dusky only at tip of the last joint of the tarsi; a linear mark on
each side of pronotum at the neck, and the exserted portion of
the ovipositor sheaths blackish; the vibrissal plates brown.

9

Fig. 3. Stemmatosteres apterus: A, female, dorsal view; B, lateral
view of head, female; C, antenna of female; D, antenna of male; EH, right
mandible of female, ventral view.

Vou.1] Timberlake-—New Genera and Species of Encyrtinae 355

Length, 0.5—0.6 mm. with head held vertically, 0.6—0.7 mm. with
head extended horizontally, exclusive of ovipositor.

MALE

Colored like the female, except that the yellow is decidedly
more of a chrome orange tint especially on the abdomen. Length,
0.4-0.46 mm. with the head vertical, 0.49-0.55 mm. with head
horizontal.

Described from eight females, two males (type, allotype and
paratypes a to h) reared from Pseudococcus timberlaker Cock-
erell, November 1—3, 1915, Millbrae (in salt marsh), California
(P. H. Timberlake), Salt Lake Laboratory no. 1830 A.

Pseudococcobius Timberlake

This genus recently characterized (Proc. U. S. Nat. Mus.,
vol. 50, p. 563, May, 1916) appears on further study of the geno-
type and the additional species deseribed below much more closely
allied to Aphycus than was at first supposed, and the discovery
of a few more species might well bridge the relatively shght gap
between the two groups. The species belonging here are prob-
ably all parasitic in mealybugs, and the differences separating
them from Aphycus appear to be an adaptation to their soft-
bodied hosts. The abdominal character used in my recent paper
on Aphycus to separate the two groups proves to be unreliable,
as I have discovered since that the fifth sternite offen reaches
to the apex of the abdomen even in Aphycus, although there is
no marked tendency towards compression of the abdomen as seen
in Pseudococcobius. The mandibles, however, so far as they
have been studied carefully, always have acute teeth in Pseudo-
coccobius and blunt teeth in Aphycus, but otherwise do not differ
much. The antennal club of P. terryi is not actually solid, as
stated in my paper, but composed of three closely fused joints,
as a study of fresh material proves, although it seems nearly
if not quite impossible to distinguish them under ordinary cir-
cumstances.

Aphycus oxacae Howard belongs in the Pseudococcobius
group, as a reéxamination of the type discloses the fact that the
mandibular teeth are acute. This species is similar to P. fwm-
pennis, described below, in type of coloration but is quite distinct.

Pscudococcobius ehrhorni is not econgenerie with ferryi, as
the mandibles are distinctly bidentate and considerably more
narrowed toward the apex. The antennae also are considerably

396 University of California Publications. [ENToMoLoey

different. It is one of the apparently rather numerous forms
that make a gradual transition between the more typical members
of the Ectromatini and Mirini. The accession of fresh material
of ehrhornt permits a better understanding of its structural
pecularities, and it is consequently elevated to generic rank.

Pseudococcobius fumipennis, n. sp.
Fig. 4
FEMALE
Head as seen from in front hardly wider than long, not so
thick anteroposteriorly as in P. terryi; frontovertex about one-
half longer than wide; ocelli in an equilateral triangle, the pos-
terior pair about one-fourth their own diameter from the eye-
margin, the median one at the center of the frontovertex; eyes
large, not pubescent, in outline nearly circular except for the
dorsal orbits being parallel; cheeks moderately long, or about
four-fifths as long as the diameter of the eyes; antennal scrobes
moderately deep, meeting above in an acute angle; mandibles
(fig. 4, B and C) with the teeth acute and nearly equal. An-

a

Fig. 4. Pseudococcobius fumipennis: A, antenna of female; B, right
mandible of male, exterio-dorsal view; C, same, anterior view.

tennal scape (fig. 4, A) flattened and expanded below, nearly
one-half as wide as long; pedicel about as long as the first three
funicle joints combined; the funicle increasing gradually in
width so that the sixth joint is a trifle more than one-half wider
than the first, the first three, and especially the second, shorter
than the last three joints, all increasingly wider than long; club
large, oval, rounded at apex, a little longer than the last three
funicle joints combined, and after collapsing one-third as wide
again as the last funicle joint. Wings thickly ciliated, the cilia

Vout. 1] Timberlake.

New Genera and Species of Encyrtinae 357

a little weaker at apex; speculum narrow, reaching two-thirds
of the distance to the posterior border of wings and well separ-
ated from a cut-off portion which is equally distant from the
posterior wing-margin; bristles on the submarginal vein a little
more developed than in terryi, but still weak. Length, 0.9-
1.2 mm.

Lower part of face and cheeks, almost the entire upperparts
of the body, and the venter of abdomen dull black, the latter
with a small white spot on each side near margin at the base;
frontovertex very pale ochraceous brown, narrowly margined on
all four sides and narrowly striped down the middle or at least
in the ocellar region with brownish black; upper portion of face
and cheeks creamy white to flesh tinted, the pale area on face
divided into two transverse bands by a blackish band of about
the same width which stretches between the lower corners of the
eyes; on the cheeks the pale area barely touches the lower corners
of the eyes and is separated from the eye-margin posteriorly by
a triangular blackish area continued forward as a narrow line
and cutting across the lower corners of the eyes to unite with
the black facial band; occiput pale ochraceous brown; collar of
pronotum narrowly whitish with a blackish dot on each corner ;
tegulae pale brownish at the center and on the posterior margin,
whitish on the anterior and lateral margins; underparts of thorax
dull brownish black with the anterior corners of the mesopleura,
and the prepectal plates whitish. Antennae brownish black,
with an oval spot on dorsal, apical corner of seape, and the apical
half of pedicel, with the last two funicle joints white, the elub
slightly paler at apex. Legs dull brownish black variegated with
white as follows: apical portion of all coxae, the trochanters,
the dorsal margin of front femora with apex and a small spot
on ventral margin near apex; base and apex of middle and hind
femora and a narrow band near apex of each, on the middle pair
connecting on lower margin with the apical area; base of all
tibiae and two narrow cross bands on each, less distinct on front
pair; all the tarsi except last joint on front and middle pair and
last two joints on hind pair, and the tibial spur white. Fore
wings conspicuously smoky, due to integumentary pigmentation,
gradually becoming clearer toward the apex and the base, with
a small clear spot at the apex of the submarginal vein and a
more deeply stained area at the juncture of the marginal and
stigmal veins; hind wings hyaline.

3598 University of California Publications. (ENToMoLoGyY

MALE

Entirely similar to the female in structural and colorational
characters, except that the last two funicle joints are brownish
instead of white. Length, 1.2 mm.

Described from two females and two males (type, allotype.
and paratypes a and b) reared from Pseudococcus solam (Cock-
erell), Uplands, California, June, 1914 (C. P. Clausen), Clausen
no. 23.

Pseudococcobius clauseni, n. sp.

Fig. 5

MALE
Head rather thin anteroposteriorly, as seen from in front
about one-fifth wider than long; frontovertex about twice as long
as wide, the dorsal orbits of eyes parallel; ocelli in an acute-
angled triangle, the posterior pair about one-half their own
diameter from the eye-margin, and twice their own diameter
from the occipital margin; eyes large, about one-fifth longer

Fig. 5. Pseudococcobius clauseni: A, antenna of male; B, right man-
dible of male, exterio-dorsal view; C, same, anterior view.

than wide, thinly pubescent with short pile; cheeks as long
as the width of the eyes; face with a moderately developed
prominence between the antennae, the scrobes rather shallow,
narrow and meeting above; mandibles (fig. 5, B and C) rather
wide at apex with three sharp, nearly equal teeth, which are
longer than in P. fumipennis. Antennal seape (fig. 5, A) flat-
tened, moderately expanded below, about one-third as wide as

Vou. 1] Timberlake—New Genera and Species of Encyrtinae 359

long; pedicel a little longer than the first four funicle joints
combined ; funicle increasing in width distad so that the sixth
joint is over one-half wider than the first; the first four joints
short, the first two a trifle longer than the third and fourth,
each except the first about twice as wide as long, the last two
joints nearly twice as long as the preceding, yet still wider than
long; club solid, no wider than the funicle, a little longer than
the preceding five joints combined, and obliquely rounded at the
apex dorsoventrad ; first five funicle joints with comparatively
long and coarse pile, the sixth joint and club with a much thicker,
finer and shorter pubescence. Wings with a narrow, curved
cross band of weaker, paler cilia at the end of the stigmal vein ;
speculum very narrow but uniform, not quite reaching the stig-
mal vein nor more than two-thirds of the distance to the posterior
margin; bristles on submarginal vein well developed. Abdomen
about two-thirds as long as the thorax, triangular in outline,
depressed, the base cordate. Length, 0.8 mm.

The frontovertex and upperpart of face to lower margin of
eyes pale clay yellow, more cadmium yellow next to the orbits
and dusky or brownish on vertex behind the ocelli; oceiput,
cheeks and lower part of face blackish brown, a paler brown,
transverse spot above antenna; thorax and abdomen dull black
with a shght brownish tint; the pleura more brownish, with
most of the propleura, the prepectal plates and the anterior
margin of the mesopleura dusky whitish; collar of pronotum
whitish with a brown dot on each corner; tegulae blackish brown
with anterior and exterior margin whitish. Antenna black, with
the sixth funicle joint and the club white. Anterior legs pale
brownish with most of the trochanter, the basal, inner, ventral
margin and band close to the apex of the femora, the base and
apex of tibiae and the tarsi yellowish white, the first and last
two tarsal joints, however, dusky ; middle trochanters and femora
dusky white with a brownish streak on the under side of the
latter, the tibiae whitish with base and apex narrowly blackish
and with two brownish black annuli near their middle, the tarsi
and tibial spur dusky yellowish, the last tarsal joint darker ;
hind legs similar to the front pair except that the tibiae have
an additional, narrow white annulus at their middle, and the
femora have their upper, inner margin whitish. Basal half of
wings deeply smoky to the end of the stigmal view, with the
apical margin of the area convex, and separated by a rather
narrow, curved pale band from the slightly dusky apex of the

360 University of California Publications, [ENToMoLoGy

disk, the basal area being integumentarily stained although
strengthened by the dense, dark colored ciliation, the apical dusky
area, however, entirely ciliary in origin. Pubescence of head and
body whitish and conspicuous on the dark parts.

Described from one male reared from an Erium species on
cactus, Riverside, California, October, 1914 (C. P. Clausen),
Clausen no. 10.

Cirrhencyrtus,* n. gen.

FEMALE

Head moderately thick anteroposteriorly, the dorsal surface
convex, passing over into face by a rather abrupt curve, the
frontovertex moderately wide, the ocelli in a right-angled tri-
angle, the posterior pair close to the eye-margin and much farther
removed from the occipital border; seen from in front, the head
is about as wide as long, the cheeks a little longer than width of
eyes and converging toward the broadly truncated oral margin ;
seen from the side, the head is subtriangular in outline; occiput
uniformly but not deeply concave, the neck inserted at the center ;

a
Fig. 6. Cirrhencyrius ehrhorni: A, antenna of female; B, antenna of

male; ©, left mandible of female, exterio-dorsal view; D, same, anterior
view.

+From Ky pfés, tawny yellow, and ELncyrtus.

Vou. 1] Timberlake.—New Genera and Species of Encyrtinae 361

eyes moderate in size, oval, one-third longer than wide, shghtly
pubescent; mandibles (fig. 6, C and D) narrowed toward apex
with two nearly equal, blunt or rounded teeth, of which the
dorsal is more apical; antennal scape (fig. 6, A) eylindrieal,
moderately long, reaching considerably beyond plane of front,
pedicel as long as the first four funicle joints ; funicle six-jointed,
increasing slightly in width distad, all joints short, transverse ;
elub large, oval, somewhat longer than the funiecle, in life hardly
wider than funicle, but after death collapsing so that it 1s much
wider, its sutures plainly visible, the last joint longest with a
rounded area of softer tissue at its apex. Thorax of normal
structure, the pronotum arched, the mesoscutum transverse, the
axillae meeting medially, the scutellum flat, triangular with apex
slightly rounded, its sides abruptly declivous; wings large, uni-
formly ciliated, with oblique hairless streak or speculum; mar-
ginal vein punctiform, stigmal moderately long, its apex triang-
ularly enlarged, postmarginal nearly as long as the stigmal, but
rapidly tapering and becoming transparent, bristles on submar-
ginal vein reduced ; costal cell of hind wing narrowed distad but
extending to the hooklets; legs longer and slenderer than nor-
mally, especially the middle pair, the middle tibial spur moder-
ately long and slender, shorter than the first tarsal joint. Ab-
domen rather subquadrate in shape, its sides nearly parallel, the
apex subtruneate ; excluding the ovipositor, which is slender and
prominently protruded, its length is about equal to two-thirds
of the thorax; the venter not at all compressed, the fifth sternite
reaching to the apex. Sculpture alutaceous, the frontovertex
more coarsely so; vestiture moderately abundant, dark colored
on the thorax; coloration flavous, the coilar of pronotum, tegulae
and appendages concolorous.

MALE

Very similar to the female, but the frontovertex is wider, the
ocelli larger, the antennae (fig. 6, B) a little shorter with slightly
longer, thicker pubescence, the club solid and slenderer, the post-
marginal vein considerably shorter, and the abdomen more ovate.

co
op)
bo

University of California Publications, [ENToMoLocy

TYPE OF GENUS

Pseudococcobius ehrhorni Timberlake
Cirrhencyrtus ehrhorni (Timberlake)

Pseudococcobius ehrhorni Timberlake, Proce. U. 8. Nat. Mus., vol. 50,
p. 564, 1916.
Fig. 6

The male hitherto unknown differs from the female as follows:
Frontovertex about one-fourth longer than wide; antennae a
little smaller, more pubescent, the club obliquely acuminate on
the upper side. Coloration the same, except that the black dots
on venter at apex are absent, the metapostnotum and the ab-
domen above at center blackish brown, and the antennae pale
brownish with base of scape alone yellowish. Length, 1.1 mm.

One male and female reared from Pseudococcus ryani (Co-
quillett) Berkeley, California, August 20, 1914 (C. P. Clausen),
Clausen no. 21.

Tribe ECTROMATINI

Tanaomastix,’ n. gen.
FEMALE

Head as wide as thorax, submenisciform, thin anteroposte-
riorly, the face to a slight degree inflexed; seen from in front,
subeireular in outline with cheeks converging toward mouth at
nearly a right angle, about as wide as long; seen from the side,
in outline subtriangular, the facial side longer than the dorsal
side, the latter being convexly rounded; occiput coneave, the
neck inserted near the center, the oecipito-vertical margin acute ;
frontovertex rather broad, as wide as long, the ocelli in a right
angle, the posterior pair farther removed from eye-margin than
from the occipital margin; eyes large, broadly oval, a little wider
on lower half, the dorsal orbits nearly parallel, posteriorly reach-
ing to the occipital border; face reaching considerably upward
between the eyes, a little longer than wide, the serobes distinct
but not deep, converging above but not meeting, separated by
a very slightly arched ridge, which is hardly visible in side view
of head; eheeks short and narrow, hardly longer than one-half
the width of the eyes; mandibles normal for tribe, long and
narrow with two acute teeth; antennae (fig. 7, B) inserted close

5 From ravaés, tapering, long, and udoré, whip, in reference to the
antennae.

Vou. 1] Timberlake-—New Genera and Species of Encyrtinae 363

to a line drawn between the lower corners of the eyes, about
equidistantly from each other as from eyes and eclypeal margin,
in form long and slender as in Leptomastix (Forster) Mayr, the
scape flattened but hardly expanded, the pedicel about equal to
the first funicle joint, the funicle filiform; all the joints much
longer than thick, not increasing in width distad, the club after
collapsing a little thicker than the funicle and a little longer
than the last two joints combined. Thorax short and thick, the
dorsum but slightly convex; pronotum short, arcuate; mesoscu-
tum about twice as wide as the median length; axillae lying in
same plane with scutellum, meeting or nearly meeting medially ;
seutellum flat, abruptly declivous at the sides, the apex rounded ;
metanotum and propodeum both short, almost divided medially,
the side pieces triangular, the propodeum also very abruptly

Fig. 7. Yanaomastix: A, female antenna of 7. claripennis; B, same of
T. abnormis.

deflexed and overlapped by the abdomen; legs long and slender,
the middle tibiae noticeably longer than their femora, the spur
a little shorter than the first tarsal joint, which is almost as long
as the following four combined, hind tibia with one small apical
spur; wings narrow, elongate, the costal cell very narrow, mar-
ginal cilia short, marginal vein about three times as long as thick,
the stigmal slightly longer, straight and enlarged at apex, the
post-marginal considerably longer than the stigmal; hind wings

364 University of California Publications, |#NtToMotocy

linear, the costal cell extremely narrow or evanescent, the vein but
slightly bent. Abdomen as seen from above acutely triangular
in outline; the dorsum deeply concave, the venter strongly com-
pressed ; the ovipositor entirely enclosed within the fifth sternite
which reaches to the apex; the vibrissal plates situated at the
base and withdrawn within the dorsal concavity.

Seulpture entirely alutaceous, the frontovertex much more
coarsely so or closely granulose, pubescence on head scanty and
very fine, on mesonotum more evident and arising from micro-
scopic punctures. Coloration non-metallic.

MALE

Not differing noticeably from the female except in the strue-
ture of the antennae, which are somewhat longer, the scape about
the same, the pedicel much shorter than the first funicle joint
or hardly longer than thick, the funicle joints all elongate, slen-
der, incised at each end above except at base of first joimt, each
elothed with rather long erect hairs arranged in three whorls,
the apical whorl somewhat incomplete on lower side, club solid,
no thicker than the funicle, and about one-half longer than the
preceding joint; the abdomen also does not become sunken in,
and the dorsum is more coarsely sculptured than in the female.

TYPE OF GENUS
Paraleptomastix abnormis Girault

The two other species so far known show but little deviation
from the above description based entirely on P. abnormis, the
chief differences residing in the shape of the head, and the length
of the middle tibial spur in comparison with the first tarsal joint
as brought out in the following descriptions and notes.

The genus is most closely related to Leptomastix (Forster )
Mayr, which differs in having the head menisciform, the face not
at all inflexed, the scrobes meeting above and separated by a
strongly arched ridge prominently visible in side view of head;
the antennae inserted higher up or between lower corners of
eyes; the pedicel not over one-half as long as the first funicle
joint ; the funicle clothed with stiff bristle-like hairs in the female,
and with longer hair in the male arranged in three whorls on
basal three joints and in two whorls on apical three; the club in
male with a fringe of short bristles on lower margin of basal
half; thorax more convex; wings wider and the postmarginal

Vou. 1] Timberlake —New Genera and Species of Encyrtinae 365

variable, in P. histrio and P. dactylopw not longer than stigmal,
but in an undescribed species considerably longer. (The above
characters based mainly on L. dactylopi: Howard.)

Paraleptomastix Girault does not seem to be so closely re-
lated, as it is a large, robust form of different sculpture and
metallic coloration; the head menisciform, the scrobes very short
(no facial ridge mentioned by Girault), the antennae slenderly
clavate, the funicle joints becoming shorter and thicker distad,
the sixth being a little wider than long, the pedicel less than half
as long as the first funicle joint; wings probably much wider
than in Tanaomastix, as the costal cell is wide; stigmal vein
eurved, longer than the marginal; the hind legs compressed, the
femora narrowly exfoliate.

The species of Tanaomastix are parasitic in mealybugs of the
genus Pseudococcus so far as known. In life the wings are held
semivertically erect over the back, at least in abnormis, a habit
quite unique among the Eneyrtinae.

Tanaomastix claripennis, n. sp.
Big. 7 A
FEMALE

Head wider than in 7. abnormis, the frontovertex about one-
fourth wider than long and coarsely alutaceous or granulose ;
ocelli large, in an obtuse-angled triangle, the posterior pair about
their own diameter from the occipital margin and fully twice
as far from the eye-margin; eyes about one-fifth longer than
wide, with a fine, sparse pubescence ; cheeks about as long as the
width of eyes; face as wide as long, the antennal scrobes narrow,
rather deep and not uniting above, the facial ridge hardly ele-
vated. Antennae (fig. 7, A) noticeably less slender than in
abnormis ; the scape somewhat expanded below, its greatest width
just distad of the middle a little more than one-fourth the length
exclusive of the radicle ; pedicel as long as the first funicle joint ;
funicle joints about equal, all about twice as long as wide; club
hardly wider than funicle, as long as the two preceding joints
and one-half of the next combined, its three joints subequal, the
apex pointed. Wings considerably wider than in abnormis, uni-
formly ciliated; the speculum wide, not quite reaching stigmal
vein nor more than three-fifths of the distance to the posterior
margin. Legs as in abnormis, except that the middle tibial spur
and the first tarsal joint are each considerably shorter than the
last four tarsal joints combined. Length, 1.0 mm.

366 University of California Publications. [ENToMoLoGyY

Frontovertex orange yellow, with the occipital margin dusky ;
face, cheeks and sides of head yellowish white, the cheeks shghtly
dusky on the sides; the face with a narrow, dark brown band on
each side connecting the lower corner of the eye with the base
of the antenna; occiput dilute blackish brown; the notum of
thorax and apical half of abdomen above and below brownish
black, the lateral margin of mesoscutum and the axillae pale
orange yellow, the metanotum, propodeum, metapleura and basal
half of the dorsum of abdomen pale brown; underparts of thorax
otherwise yellowish white, and most of the basal half of the venter
dusky white. Antennae dark brown, the scape with a white
triangular spot on the outer and inner surface beginning almost
at the base, not quite reaching the dorsal margin but reaching
to the middle on the ventral margin, with a narrow, oblique,
eurved band running from the dorsal margin a trifle beyond the
middle to the ventral margin just before the apex. Legs yellow-
ish white, the middle coxae dark brown, the fore and hind tibiae
on dorsal margin, all femora on dorsal margin at apex, and the
fore and hind tarsi, pale brownish or dusky, the middle tibiae
and tarsi very dilute brownish yellow with the first tarsal joint
and the spur whitish. Wings hyaline, the veins brown. Pubes-
cence of thorax pale brown and inconspicuous.

Described from two females (type and paratype) reared from
Pseudococcus ryani (Coquillett), Pasadena, California (C. P.
Clausen), Clausen no. 56.

Tanaomastix abnormis (Girault)
Nee, ff, AB

Leptomastia sp. Viereck, Monthly Bulletin, Cal. State Comm. Hort.,
vol. 4, p. 208, fig. 36-38, April, 1915.

Paraleptomastix abnormis Girault, Entomologist, vol. 48, p. 184,
August, 1915.

Paraleptomastix abnormis H. S. Smith, Jour. Econ. Ent., vol. 10,
p. 262-68, figs. 9-13, pls. 13-14, April 21, 1917.

As compared with 7. claripennis this species has the fronto-
vertex no wider than long, the face longer than wide, with the
facial ridge more elevated, the antennae (fig. 7, B) much slen-
derer, each funicle joint being about three times longer than
thick, and the middle tibial spur considerably longer. It has
about the same coloration, excepting the following points: the
frontovertex paler orange yellow, the face without dark marks,
the notum of thorax varying from dark to pale brown, the base

Vou. 1] Timberlake.—New Genera and Species of Encyrtinae 367

of the abdomen more whitish, the legs paler, the scape of the
antenna except the dorsal margin and the apex of the pedicel
white, and the wings conspicuously trifasciate, the bands due
both to integumentary pigment and to the ciliation.

A female reared from a Pseudococcus on Citrus (not Pseudo-
coccus citrt Risso), Okitsu, Japan, July, 1914 (Kuwana), Cali-
fornia State Insectary no. 783, is entirely similar to specimens
from Sicily, except that the blackish streak on the dorsal margin
of the scape does not quite reach the apex. This species has
been introduced into southern California from Sicily to help
reduce the citrus mealybug (Pseudococcus citri Risso), and, ae-
cording to the latest reports, has become established.

Tanaomastix albiclavata (Ashmead)

Aphycus albiclavatus Ashmead, Proce. U. 8. Nat. Mus., vol. 29, p. 404,
1905.

This species as compared with 7’. abnormis has a thicker, nar-
rower head, with the frontovertex nearly twice longer than the
posterior width, the dorsal orbits of eyes slightly converging be-
hind, the ocelli in an acute angle, the posterior pair close to the
eye-margin, the scape subexpanded below, the scutellum more
acute at apex, and the spur of the middle tibiae fully one-half
as long as the tarsi.

The above notes are based on one female from the Philippine
Islands (undoubtedly from the vicinity of Manila), reared from
a Pscudococcus species, and received from the California State
Insectary. It agrees with Ashmead’s type specimen, which is a
female, and not a male, as stated in the original description.

at righ fae tae : :

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Vol. 1, No. 9 pp. 369-464, plates 2-16 November 21, 1922
INCLUDING A MONOGRAPH ON THE
NORTH AMERICAN SPECIES.
ier PERO ag, A RIS en A a hase
JAMES CHESTER BRADLEY
UNIVERSITY OF CALIFORNIA PRESS 0 8)
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Vol. 1, No. 9, pp. 369-464, plates 2-16 November 21, 1922

THE TAXONOMY OF THE MASARID WASPS,
INCLUDING A MONOGRAPH ON THE
NORTH AMERICAN SPECIES

BY
JAMES CHESTER BRADLEY

INTRODUCTION

Fifteen years ago, having before me all the described North Amer-
ican masarid wasps, I prepared tables for their identification. This was
an interesting and easy task, as their characters are distinct and easily
described. The manuscript remained unpublished while I awaited
opportunity to determine the relations of American species of Masaris
with the North African vespiformis, the type species of the genus.
Later, at the suggestion of Dr. Joseph Bequaert, I renewed the study
of this group, rewrote the tables, and added descriptions of all the
North American species.

I have studied the wings, mouth parts, and male genitalia of all
the North American species, and of species of several exotie genera.
The results are of interest particularly in establishing clearly the re-
lations of American species with Masaris of North Africa and with
Trimeria. In addition, sufficient facts have been disclosed to make
advisable a tentative revision of the classification of the family, which
I present in the form of the subjoined tables. In view of the scanti-
ness of my material, I do not presume to think that this classification
will have more than a temporary value, but I do believe that it is an
improvement, taxonomically speaking, over what has preceded, and
it will at least correct certain errors in the existing keys.

370 University of California Publications [ENTOMOLOGY

Since the days of de Saussure two keys to the genera of Masaridae
have been published. The one by Ashmead in the Canadian Entomol-
ogist, volume 34 (1902), page 219, will lead the unwary user astray
for the following reasons: (1) a primary division (category 3) reads

‘labrum extensible’’

where ‘‘lgula’’ is meant, and if not corrected
is unintelligible; (2) Paraceramius, Ceramius, Cerannoides, Trimeria,
and Jujurhta [sic] are described as having the ‘‘labrum [i.e., hgula]
not extensible,’’ which is incorrect for these genera; (3) the character
used in category 8 (marginal cell with or without an appendage) will
not serve to distinguish between the groups for which it is intended;
(4) the claws of Masaris are incorrectly described as being simple, and
several other characters are thus incorrectly described and wrongly
applied. The second key is a compilation by Dalla Torre published
in the Genera Insectorum, 1904, fase. 19, and contains most of Ash-
mead’s errors and some additional ones.

I wish to express my acknowledgments and gratitude to Dr. Joseph
Bequaert, who has contributed fertile suggestions and has taken much -
interest in the prosecution of this work, and has loaned me, with
permission to dissect it, a female of Celonites, as well as specimens of
Gayella and Trimeria. Acknowledgments are further extended to Dr.
F. E. Lutz, for the loan of specimens of Paragia from the American
Museum of Natural History, with permission to dissect them; to Dr.
Henry Skinner, for the loan of several species of Pseudomasaris from
the collection of the American Entomological Society, and to the
authorities of the United States National Museum for courtesies ex-
tended to me while visiting that institution.

TAXONOMY OF THE MASARID WASPS?
Tur MoutrH Parts AND THEIR VALUE IN CLASSIFICATION

The remarkable retractile ligula of most of the genera of Masaridae
has been both figured and described by de Saussure, and I do not need
to dwell upon it here. When withdrawn, which is accomplished by a
process of introsusception, only the tip of the ligula is exposed, the

1 Haplanation of the text.—Under each genus is listed all of the species known
to belong to it, but references to literature are given only since the time of
Dalla Torre’s Catalogus Hymenopterorum.

The color nomenclature is chiefly that of Ridgway’s Color Standards and Color
Nomenclature.

Vow. 1] Bradley.—The Taxonomy of the Masarid Wasps Biff

median parts being coiled in a great loop which may at times enor-
mously distend the membrane of the neck. The character is a very
positive one, involving the entire shape and structure of the ligula,
and certain chitinized basal plates. These plates seem to be developed
for the purpose of assisting in effecting the invaginating process, and
there can hardly be any transition between the two conditions. This
ligula is the character to which Ashmead has reference in his key to
genera, writing, however, by a slip of some sort, ‘‘labrum extensile’”’

9?

or ‘“‘labrum not extensile.’’ The character may be usually made out
under a binocular microscope without dissection, and very readily and
positively by dissecting out the mouth parts; but it is a curious fact
that de Saussure, owing to the poorer optical instruments available
in his day, has described the condition wrongly in several genera.
These errors have stood in literature until the present time.

Dr. Bequaert suggested to me that the number of palpal segments
is of very doubtful generic value in the diplopterous wasps. The last
joints, he says, are very apt to drop off on the emerging of the adult,
or later, and furthermore there are all sorts of variations within the
limits of a single genus, such as Odynerus. Granting that, I can not
believe that the case is entirely similar in the Masaridae. My dissec-
tions agree perfectly for the most part with the figures and descriptions
of de Saussure except in the case of Celonites. I can not believe that
apical segments could be lost without leaving indications of the fact.
The shape of an ultimate segment is different from that of a penulti-
mate. I have examined all the North American species, and find
within the genus Psewdomasaris no variation in the maxillary palpi
(which are always rudimentary), and no variation in the labial palpi
of the females, which are of a peculiar type, 3-segmented, but very
different from the palpi of the males, even when the latter are 3-seg-
mented. The latter vary from 1- to 3-segmented, showing progressive
reduction by coalescence of the segments, as is perfectly apparent from
an examination of the series. In this ease it would be futile to use the
number of segments of this sex as a generic or even subgeneric char-
acter. Still more surprising is the similar sexual difference in both
the pairs of palpi in Celonites, and especially that it should have ap-
parently escaped detection. Analogous sexual differences may, of
course, exist in other genera, and within some genera there may be
variation in the number of segments, as, for example, in the labial

312 : University of California Publications | ENToMOLOGY

“=

palpi of males of Pseuwdomasaris; but I think that the variation is not
likely to be such as to affect the validity of the general arrangement
here laid down, although it may of course affect the details. Such
variations as occur are most probably to be found within the middle
groups, not at either end of the series, and especially not within the
final group of three genera in which the maxillary palpi are reduced

to tubercles.

THE WINGS AND THEIR TAXONOMIC SIGNIFICANCE

The wings display taxonomic characters of considerable importance
which have been largely overlooked by previous authors. The char-
acters that have been used are three: (1) the number of submarginal
cells, (2) whether the first receives both recurrent veins, or the second
and third each receive one, and (3) the presence or absence of an anal
lobe in the hind wings. These characters are all of value, but the story
told by the wings is still far from being finished.

THE POSTERIOR LOBE

In a forthcoming paper on the wings of Hymenoptera I shall dis-
cuss the nature of the posterior (i.e., anal) lobe of the hind wings
throughout that order.. In the more primitive Hymenoptera it is a
large area separated from the rest of the wing membrane by a mar-
ginal notch, the axillary excision. This lobe and the excision are not
confined to the wings of Hymenoptera, but have their homologues in
other orders. The position of this notch is always at the apex of
the second anal furrow, which les between the second and the third
anal veins. The notch may be retained long after both fold and veins
disappear.

Anterior to the axillary excision is another notch, the preaxillary
excision. It is always situated at the apex of the first anal fold, which
lies invariably just cephalad of and close to the first anal vein (anal
vein), crossing M, (the submedian vein) shortly before its union with
1st A. Between the preaxillary and the axillary excisions is an area
which I shall call the preanal lobe. Very often the preaxillary ex-
cision is reduced to a mere undulation of the wing membrane, but its
position and that of the anal furrow are the most characteristic fea-

tures of the topography of the hind wings.

VoL. 1} Bradley.—The Taxonomy of the Masarid Wasps 373

In the more primitive members of each of the major groups of
aculeate Hymenoptera (1.e., Formicoidea, Vespoidea, Scolioidea, Sphe-
eoidea, and Apoidea) both axillary and preaxillary excisions are pres-
ent, and accordingly both posterior and preanal lobes are well devel-
oped. Each have been lost in some or most of the specialized members
of each superfamily. The tendency of specialization throughout the
aculeates has been toward a reduction of the anal area of the hind
wing, and especially of the posterior lobe.

With this tendency to reduction in mind, let us turn to Huparagia,
representing the Euparagiinae, and consider the wing (fig. 100). The
posterior lobe is large, two-thirds the length of the cell M, + Cu -+ Cu,.
In all other subfamilies except the Vespinae the posterior lobe is re-
duced to a small round or oval flap at the extreme base of the wing,
from one-fifth to one-third the length of the cell M, + Cu+ Cu,. In
the Vespinae the posterior lobe has been so completely reduced that it
is altogether wanting. In this respect, as in others, Huparagia stands
out alone as ancestral, while the Vespinae take their place as most
specialized.

In all cases except the Vespinae the preaxillary excision is an in-
significant notch, a mere undulation, or is altogether lacking. In the
Vespinae it is deeper. Taking the order as a whole, it would seem
that the primitive condition of the preaxillary excision is a distinet
notch, but not a deep incision. The precise condition seems subject to
variation within narrower limits than in the case of the axillary.

In the Vespinae the anal area of the hind wing is greatly reduced,
so that the wing is subpetiolate.

THE CENTRAL AREA OF THE HIND WINGS

The venation of the central area of the hind wings is of consider-
able taxonomie importance. By referring to figures 97-101 and 104,
plates 13, 14, 15, the modification it has undergone will be seen.
Figure 98 represents a primitive condition (for aculeates) in which
M, is transverse, m and M, are inserted below its middle, and the
eaudal part of M, is interrupted just before its union with M, + Cu,,,
+ Ist + 2d A by a bulla. The bulla is formed by the crossing of the
Ist anal furrow, which is present, and which is lying immediately
cephalad of 1st A and its amalgamates. The furrow ends in a pre-
axillary excision.

374 University of California Publications [ENTOMOLOGY

In Euparagiinae (fig. 100) this primitive condition is maintained
except that m has migrated cephalad along M, to the point of separa-
tion of the latter from M,. The crossing of M, by the first anal furrow
is represented by a bulla, and the very faintest trace of the apical
section of Ist A and its amalgamates is to be seen.

The subfamilies, which have ordinarily been classed as Eumenidae
and Vespidae proper, are represented by figure 99. The vein M, is
strongly angled, receiving m and M, at the angle. Both the latter
vein and the apical section of Ist A and its amalgamates are ordinarily
retained as fully formed veins, in some eases only as traces, but they
are never wholly lost. First A ends, as always, in the preaxillary
excision.

The Masaridinae have been derived from the type of figure 99 by
the complete dropping out of the apical section of Ist A and its amal-
gamates. The wings of this subfamily are represented by figure 101,
and what has happened to them is diagrammatically shown in figure
104. In the latter case the lost apical section of 1st A and its amal-
gamates and the anal furrow are represented respectively by a dotted
and a dashed line.

In all of the Masaridinae the bulla indicating the crossing of M,
by the first anal furrow has been lost, and as the vein formed by the
union of M, with M, + Cu,,, + 1st + 2d A has straightened out it is
impossible to point out the exact spot where it occurs.

The evidence for this interpretation of the hind wing of the Masa-
ridinae lies in the position of the preaxillary excision. It is not the
obvious interpretation from a casual examination of the wing, and in
the keys I have referred to the condition as ‘‘M, apparently straight
and m lacking.”’

In the Gayellinae (fig. 97) both the apical section of Ist A with its
amalgamates and m and M, are fully preserved, but the vein m and
M, has migrated apicad along m-cu to a point remote from M.,, a con-
dition not infrequent in other aculeate Hymenoptera, but not oceur-
ring elsewhere in the Vespidae. M, is transverse and straight, there-
fore primitive, as in Huparagia, while the sidewise migration of m
and M, must be considered a specialization.

Vor. 1] Bradley —The Taxonomy of the Masarid Wasps 375

THE MEDIAL AND CUBITAL REGIONS OF THE FOREWING

Coincident with the longitudinal plaiting of the wings of many
Vespidae there has been a shifting and realignment of the veins in the
central part of the wing, through which Ist A and Cu, + Ist A, M,,
M,,,, M,,., R; + M,,., and R, + M, together form a more or less un-
broken straight bar obliquely across the wing. The plaiting occurs
just caudad of this bar. At the same time M, + Cu, is greatly short-
ened, and is situated much farther basad than usual; m-cu is inserted
basad of M,-+ Cu, and the intervening section of Cu, is deflexed to-
ward Ist A and Cu,. The cell M, is triangular, elongated basally, and
pointed, the apex truneate. The condition just described is exemplified
by the wing of Mischocyttarus. While varying in detail it is the gen-
eral condition throughout the subfamilies having longitudinally plaited
wings.

In the Masaridinae it is evident that the same tendencies of special-
ization have been operative, but they have not proceeded in any ease
to so high a degree of perfection. Of the genera figured, Paragia,
Celonites, Cerannoides, Ceramius, and Trimerva approach the condition
most nearly. In each M, + Cu, is situated apicad of m-cu, the inter-
vening sector of Cu, being deflexed. In these genera the cell M, is
narrowed or pointed at base (Paragia), and triangular (Paragia,
Celonites, Trimeria), or somewhat triangular (Ceramius and Parace-
ramius) inform. In Masariella, Masaris, and Pseudomasaris M, + Cu,
is longer than in the other genera, and is opposite m-cu (Masariella),
somewhat basad of it (Masaris), or its own length basad of m-cu
(Pseudomasaris). Consequently, in these genera, the cell M, is less
narrowed (though still unusually narrow) at base, and less triangular
in form; and the veins Cu and Cu, instead of Ist A and Cu, + Ist A
form a straight bar with M,. In other words, these genera are in this
respect more primitive than others, and show different stages in the
type of specialization that goes hand in hand with longitudinal
plaiting of the wings, and culminates in the venation exemplified by
Mischocyttarus.

The apex of the cell M, is closed by the vein M,, which may be
straight, or slightly curved, as in all wasps with longitudinally plaited
wings (see fig. 98, of Mischocyttarus), as well as in some Masaridinae
(Paragia, Celonites, Trimeria, Masaris, Pseudomasaris) ; or it may be
angled at the point of receiving m and M,, the angle being (on the

376 University of California Publications [ENTOMOLOGY

side of the cell M.) always less than 180°. This is the case in other
Masaridinae (Ceramioides, Ceramius), and in Gayellinae. In Eupa-
ragiinae the angle is about 125°, the upper part of the cell M, being
produced apicad to a point closer to the cell R, (measured along
M,,,) than the length of the cephalic section of M,, a condition not
obtaining elsewhere in the Vespidae, and giving to the wing of Eupa-
ragia a peculiar facies. In the Gayellinae the condition in this region
of the wing is almost identical with that found in Ceramoides—
M,-+ Cu, being a little longer than in that genus.

In Euparagiinae, apart from the prolongation of the upper apical
angle of the cell M,, the condition of this region of the wing is more
primitive than obtains elsewhere. M,-+ Cu, is not abnormally far
basad in the wing, is opposite m-eu and two-thirds as long as that vein,
so that the cell Cu + Cu, is wide at apex. By reason of the fact that
the veins M,-+ Cu, and M, approach each other, the caudal margin
of the cell M, is seareely longer than the basal width of the cell, a
condition very far from being the ease in any other Vespidae.

THE RADIAL REGION OF THE FOREWING

The most primitive condition of the radial region of the forewing
occurs in Kuparagiinae, Gayellinae, and Raphiglossinae, in which cells
R, and R, are distinct, receiving, respectively, veins M, and M,,,.

In Zethinae, Eumeninae, Stenogastrinae, Epiponinae, Rhopalidii-
nae, Polistinae, and Vespinae the base of the free part of the vein M,
has migrated basad toward M,,, until they both are opposite the cell R..

In Masaridinae these two veins are in the position just indicated, or
in an intermediate position, but an additional step is found in the loss
of the vein R., throwing the cells R, and R, together.

In the Euparagiinae, Masaridinae, and Gayellinae an appendiculate
cell is always present, and in the first two of these subfamilies the cell
2d R, + R, does not extend farther toward the wing apex than does
the cell R,. In the few genera in which this is not the case both cells,
and, in Gayellinae and in most of the genera of Vespidae with plaited
wings, the cell 2d R, + R, extend closer to the wing apex than they do
in Masaridinae. In Stenogastrinae this condition is carried to an
extreme.

In the Vespidae with folded wings an appendiculate cell is rather

exceptional.

~]

VoL. 1] Bradley —The Taxonomy of the Masarid Wasps 37

THE PREANAL EXCISION OF THE FOREWINGS

Most Hymenoptera have a notch (the preanal excision) in the
margin of the forewing opposite the tip of M,,,-+Cu,,. + 1st + 2d
-+ 3d A. In most Vespidae this is present except in Euparagiinae,
where it is lacking.

THE RANK OF THE ‘‘MASARIDAR”’

Not accepting the eumenid wasps as a family distinct from the
Vespidae, I can still less consider the masarid wasps as a separate
family. Certain tendencies develop within the masarid line which are
sufficiently distinct from what we find in the other solitary Diploptera
and social wacps, but in every instance such characters are developed
within the group, and we ean always find genera displaying conditions
of the same organs which do not differ from those of other Diploptera.
It is worth while to briefly review the organs that have been relied
upon for distinguishing between the three so-called families.

The mouth parts of the more highly specialized Masaridinae, by
reason of the retractile ligula, differ completely from those of other
Diploptera. The tendency to great or total reduction of the number
of palpal segments and of the size of the palpi.is also an important
group characteristic. But in Huparagia and Paragia the ligula is not
retractile, and neither it not the other mouth parts differ otherwise
from the simple conditions found in the Eumeninae.

The antennae in the more highly specialized Masarinae have seg-
ments 8 to 13 of the male fused into a club, the divisions between them
discernible except that between segments 12 and 13, which is not to be
discovered unless the specimen be boiled in caustic potash, in which case
the minute segment 13 may sometimes be observed. The antennae of
the females have a similar club, but usually differing in shape. In
Euparagia, however, the male has 13 distinct and entirely separated
segments without a club, the apical part of the flagellum being merely
slightly thickened. In Paraceramius lusitanicus the antennae of the
male have the apical segments hooked, as in many male Eumeninae,
not clavate, and consisting (as in a few Eumeninae) of 12 segments.

378 University of Califorma Publications [ ENTOMOLOGY

The wings of Masaridinae are ordinarily not longitudinally plaited,
and this is used by some authors as the one character for separating
the ‘‘family’’ from other Diploptera. In Celonites, however, the wings
are as completely plaited as in any other Diploptera.

The majority of Masaridinae have cells R, and R, of the forewing
united (two closed submarginal cells), whereas in most other Diplop-
tera these cells are separate (three: closed submarginal cells). In
Euparagia, however, cells R, and R, are separate, and also in one
or two of Cameron’s genera (if they be really Masaridinae). While
most other Diploptera have both the veins M,,, and M, arising from
the cell R., a few genera (Raphiglossinae) agree with Euparagia in
having M, arise from the cell R,.

While most Masaridinae have toothed claws, hke many eumenid
Diploptera and unlike most vespid Diploptera, certain genera, as
Pseudomasaris, ete., have simple claws, as has also Euparagia.

The vespid Diploptera usually have two spurs on the middle tibia;
the Eumeninae proper have only one, but several species usually classed
as Eumenidae have two. Many genera of the Masaridinae have two
spurs, several genera have one, and sometimes the number varies within
the genus.

I am familiar with the male genitalia of only a few genera of
Vespid and of Eumenid Diploptera. In these the squama is acute at
apex and separate from the ramus. In Masaridinae it is more often
blunt and almost always fused to the ramus. But in EFuparagia it is
separate as in other Diploptera. The genitalia of some genera of
Masaridinae differ more radically from the more primitive condition
found in other Masaridinae than do the latter from those of Vespa or
Odynerus.

In many of these characters Euparagia is the connecting link
that breaks down the distinctions between other ‘‘Masaridae’’ and
‘‘Humenidae.’’ As a result, the only tenable rank for the masarine
wasps, it seems to me, is as one or two subfamilies, along with the
several subfamilies into which the old families Vespidae and
Eumenidae may best be divided, of the single comprehensive family,
Vespidae.

Vou. 1] Bradley.—The Taxonomy of the Masarid Wasps 379

KEY TO THE SUBFAMILIES OF VESPIDAE2

A. Transverse median vein (M,) of the hind wings straight or curved, not
angled; the discoidal vein (m) usually wanting, or vague, in which latter
case’ the median vein, the submedian and the cubitus and the discoidal
veins (m, M,, M;, and m-cu) meet at a common point (Huparagia), or the
discoidal vein is completely formed, arising from the cubitus far apicad
of the transverse median vein (Gayella) ; only in Gayella are the discoidal
vein and the apical sections of radius, cubitus and the anal vein (R,,
Ry; + My, m and M,, and Ist A) all distinetly formed veins. In all other
cases one of these is entirely wanting and the rest are usually only indi-
cated as traces. Forewings with two or three submarginal cells, in the
latter case the second and third each receiving a recurrent vein (veins
M, and M,,, arising opposite cells R, and R,, respectively). Anal lobe of
the hind wing always present, but the preanal excision absent or indistinct.

B. Anal lobe of the hind wing elongate, more than one-half the length of
the submedian cell (M,-+ Cu-+ Cu,). Second discoidal cell of the
forewing (cell M;) of irregular shape, not pointed at base, its apex
greatly produced above toward the apex of the wing; the section of
the discoidal vein between the Ist recurrent and the subdiscoidal veins
(M, cephalad of m) almost longitudinal in position, longer than the
first recurrent vein (M;,,) and forming an angle of more than 180°
with its section caudad of the subdiscoidal vein (with M, caudad of
M); the transverse median vein of the forewings (M,-+ Cu,) a long
crooked vein, longer than the first transverse cubitus (r-m and Rs);
three submarginal cells present in the front wing. In the hind wing
a trace of the discoidal vein (m and M,) may be seen, arising from
the point of union of the cubital, median, and transverse median veins,
but there is no trace of the apical section of the anal vein; only a few
(9 or 10) costal hooks present. Forewings without a distinct pre-
axillary excision. Antennae of the male composed of 13 distinct seg-
ments. Mouth parts primitive, without an elongate retractile tongue,
with 6 segmented maxillary and 4-segmented labial palpi....Euparagiinae.

BB. Anal lobe of the hind wings a small circular or oval flap, much less than
one-half the length of the submedian cell (cell M, + Cu+ Cu,). In
the forewings the second discoidal cell (M,) wedge-shaped, narrowed
or often pointed at base, its cephalo-distal angle not greatly produced
toward the wing apex; the section of the discoidal vein between the
1st recurrent and the subdiscoidal veins (M, cephalad of m) transverse

2 This key does not distinguish between the subfamilies Zethinae, Eumeninae,
Stenogastrinae, Epiponinae, Rhopalidiinae, and Polistinae, as these groups are
not treated in the present paper. Their characters are stated by Bequaert in his
“*Revision of the Vespidae of the Belgian Congo,’’ Bull. Amer. Mus. Nat. Hist.,
vol. 39 (1918), pp. 13-17.

3In reality, in all the Masaridinae, it is the apical section of the anal vein
that is lost, and at least a trace of the discoidal vein is preserved. This, how-
ever, is not apparent from inspection; any one examining the wings casually
would take the reverse to be the case. An explanation of the true condition is
to be found under the description of the wings of Masaridinae.

380 University of California Publications | ENTOMOLOGY

or somewhat oblique in position, always shorter, often greatly shorter
than the first recurrent vein (M;,,), and forming an angle of 180° or
less with the section of the discoidal vein caudad of the subdiscoidal
vein (M, caudad of m); the transverse median vein short and straight,
often almost lacking, always shorter than the Ist transverse cubital
vein (r-m plus Rs). In the hind wings there is usually no trace of the
discoidal vein (m and M,), but in Gayella it is a fully formed vein,
inserted far apicad of the transverse median vein (M,); apical section
of the anal vein always present, usually only as a trace; numerous
costal hooks (20 or more) usually present.

C. In the hind wings apparently no trace of the discoidal vein (m and
M.) is present, and only trace of the apical sections of the radial,
cubital, and anal veins’ (R,, R,,;, Ist A). Mouth parts (except in
Paragiini) specialized, with a retractile ligula which is often greatly
elongate, and showing progressive reduction in the number of palpal
SC OTC MGS) seh ete e resect eae sees Nee ae Seen mcr ..Masaridinae.

CC. In the hind wings the discoidal vein (m and M,) is present and fully
formed, arising from cubitus far apiead of the submedian vein.
Mouth parts primitive, the ligula not elongate, the labial palpi 4-
segmented and the maxillary palpi 6-segmented (figs. 4, 97)

Ube Rast RE oats Dae ais Ma Ee MPs eS See as ea EA EE _...-.-Gayellinae.

AA. Transverse median vein of the hind wings (M,) angled for the reception
of the base of the discoidal vein (m) which is almost invariably a fully
formed vein, or at least a distinct indication of the discoidal vein and the
apical section of radius, cubitus, and the anal vein (R,, R,y,; + M,, m and

M., Ist A) in the hind wings and these very generally all well formed

veins. Forewings with 3 submarginal cells; the second discoidal cell (M,)

ordinarily wedge-shaped, pointed at the base and broadly truncate at apex.

It varies somewhat from this condition but not greatly, being always

strongly narrowed at base; while the apex may be somewhat sinuate, it

is never greatly extended toward the wing apex; anal lobe of the hind
wings very short and circular, oval or wanting.

B. Forewings with the second and third submarginal cells each receiving a
recurrent vein (vein M, arising opposite the cell R,, and vein M,,,
TISING 4Opposite ablienc elicits) cess a5 eee eee ee Raphiglossinae.

BB. Forewings with the second submarginal cell receiving both recurrent
veins (veins M, and M,,, both arising opposite the cell R;).

C. Posterior lobe of the hind wings present; preaxillary excision, if
present, a mere undulation or shallow notch; the apical section of
the anal vein not strongly curved caudad, the hind wing not sub-
petiolate Cig: 98)" seca ieee Te ee ee ee ee Zethinae.
Eumeninae, Stenogastrinae, Epiponinae, Rhopalidiinae, Polistinae.

CC. Anal lobe of the hind wing wanting; preaxillary excision‘ present,
deep, the apical section of the anal vein strongly curved caudad;
the hind wings subpetiolate, by reason of contraction of the anal
UTC. (GRAD Oi Pee eee a eee Sone renee ee eee Vespinae.

4 The preaxillary excision is a notch at the apex of the 1st anal vein of the
hind wing; it is not the notch that cuts off the anal or posterior lobe, when
present. Its significance is discussed in a forthcoming paper by the present
writer on the wings of Hymenoptera.

VoL. 1] Bradley —The Taxonomy of the Masarid Wasps 381

A WORKING Key For IDENTIFYING THE GENERA OF EUPARAGIINAE anp
MASARINAES5

(This key does not pretend to present the natural relations of the genera.)

1. Abdomen strongly petiolate, the first segment elongate, formed as in

JAAS ieee tee eae oe eee Plesiozethus and Paramasaris Cameron.é
Abdomen sessile, or the short first segment with a small anterior neck,
[OMB TAO CNG UM AS eT OU SIal WSS rer ee Ra ree ey eee reer ere 2

bo

. Forewings with cells R, and R,; separate (3 closed submarginal cells) ......
HOES See Fo te RS Euparagia Cresson.

Forewings with cells R, and R,; coalesced (2 closed submarginal cells) ... 3
3. The first abdominal segment with an anterior neck therefore somewhat
TOLER CMON ENG er ce eam ee Ceramiopsis Zavattari.
First abdominal segment broad and sessile, as in Vespa ........ Se eee 4
4. Postscutellum not covered by the scutellum, but produced caudad and
Vey aUSL Gon FEN OED acs ee ere Masaris Fabrians.
Postseutellum rounded and more or less covered by the scutellum ............ )
5. Sides of abdomen margined, serrate, venter concave; wings plaited long-
UCU CLUE eI ayaureas ae) °C S 9 Cleaner me nes eens eee ones Celonites Latreille.
Sides of abdomen not margined, venter convex or nearly flat; wings long-
ree btiraeMll yy joleMnexol @palbye sir OGRA ae cece ee ee eee 6

6. Tegulae short, ovate or semicircular, scalelike, not covering the base of
the scutellum; clypeus produced anteriorly, its margin truncate, in the

SEQUENT SKOPIMEN AMEN THO) ENCES eee eee ee a ee ree iH
Tegulae elongate and usually acute posteriorly, covering the base of the
scutellum; clypeus emarginate or trilobed at apex ..-.........-2..22--.-200-2-0------ 7

7. Clypeus with its apical border trilobed; m=cu opposite M, + Cu, ........

seth A OREN ak SO eI ee IONE A Jugurtia Saussure.
Clypeusiwait bnitsmapicall borden) te miar omer ey yce ce stece aetna cena ee 8

8. In the forewings the mediocubital cross-vein (basal vein) attached oppo-
site to or basad of M, + Cu, (submedian vein); third ventral segment
of males unarmed; larger spur of posterior tibiae not always bifid —.. 9

In the forewings the mediocubital cross-vein attached to M, far apicad
of M,-+ Cu,; anterior trochanters of the male unarmed; third ventral
segment of the male with a process; larger spurs of posterior tibia
{OH CGF Sec ie eee ere cane POE Res ae Se nee Pseudomasaris Ashmead.

9. Anterior trochanters of the male armed with a prominent claw or lamella;
apical segments of the antenna of the females somewhat incrassate,
but not forming a distinct ovate club; habitat South America ..............
eer MR I Se ce seas fe eras ess cues Sass stakes Lan eth Sas oa ae coe Trimeria Saussure.

5 The synoptic tables which follow later are intended to suggest the natural
relationships of the genera. They are not readily applicable for purposes of
identification, and I have therefore prepared this artificial identification key,
which I think may be used by any one easily and with certainty as to its mean-
ing. At the end is a short working key to North American genera.

6 Cameron distinguishes between these two genera on the grounds that in
Plesiozethus there are only 2 closed submarginal cells, both M,,, and M, arising
from the cell R,,;, while in Paramasaris cells R, and R; are separate; M,,, arising
from cell R; and M, from cell R, Zavattari maintains that Cameron is wrong,
and that both genera show the latter condition. In that eventuality it is prob-
able that they will have to be united under the name Paramasaris.

382 University of Califorma Publications [ ENTOMOLOGY

10.

Lett:

13.

14.

Anterior trochanters of the male unarmed; apical segments of the an-
tenna of the female forming a distinct ovate club; habitat South
BASES © sale eA CLS © U1its Hae e ae EU eee e

Pedicel greatly enlarged, globose, nearly as large as the scape; antennae
not as long as the distance between the eyes; size small, 4 mm. ............
Be ea oie freee Sa ee ae toa neg eee eee ea Quartinia Gribodo.

Pedicel not enlarged nor globose, less than one-half as long as the scape;
antennae of the male much longer than the distance between the eyes;
size) lange yi. mms OT) ON CTS tece sean eee nese es cee eee Masariella Brauns.

Scutellum semicircular, very strongly elevated, its top flat, the sides ab-
rupt, venter of the male with a process on the second segment, and
with 8 exposed segments, the seventh small, simple, the eighth with a
basal) process; -Anstrailn aa: occ ccses: asec see cen ccce crete cee eecnee een

Scutellum elongate and more or less obtusely pointed behind, not strongly
elevated, but moderately convex, or flattened on top, its sides not
abrupt, venter of the male usually unarmed, with 7 exposed segments,
the seventh large and truncate or subtruncate at apex; eyes emargi-
nat at least slightly, usually deeply; European or African

. Eyes emarginate; parapsidal furrows absent....Metaparagia Meade Waldo.

Eyes not emarginate; parapsidal furrows distinct........ Paragia Shuckard.
Hyes deeply emarsinate; veniter of male marie d sce seen ere
Eyes very broadly and shallowly emarginate, scarcely more than sinuate;
third ventral segment of the male armed with one or two tubercles;
middle tibiae with one apical spur; antennae of the sexes dissimilar,
those of male with the apical segment elongate, tapered and recurved;
mandibles of the male without a large basal tooth, clypeus of male
longer than broad; anterior trochanters of male produced at apex into
an elongates call @} chee. ssencce eee ee eee Ceramioides Saussure.
Middle tibiae with a single apical spur; antennae of the sexes dissimilar,
those of the male rolled at apex, the last segment much longer than
broad; mandibles of male without a large basal tooth; clypeus of the
male longer than broad; anterior trochanters of male unarmed ..............
$e Sah nT poten cee esr ON ees CS SU SN Paraceramius Saussure.
Middle tibiae with two apical spurs; antennae of the sexes similar, not
recurved at apex, the last segment broader than long; mandibles of
the male with a very large tooth at base; clypeus of the male broader
than long; anterior trochanter of the male produced and acute at
BLP Oy oltre tree crate tes epee Sey tees we ns eet cpa eaceeneteree eee Ceramius Latreille.

10

12

13

14

KEY TO THE GENERA OF KHUPARAGIINAE anp MASARIDINAE Known To Occur

bo

IN NortH AMERICA

. Forewing with cells R, and R, separate (3 submarginal cells) -...................

Forewing with cells R, and R,; coalesced (2 submarginal cells) -..............---
ea se te A rk ter eae Pseudomasaris Ashmead

Ab domientsessileistac. te. ern eee eee Euparagia Cresson

Abdomen! petiolate; sas) any Ziet 1s) eee see an nace nee eee sees aeeee nace ese ccens een eeneneneeeesraceene
pase a 4 ae tee a Oil ea Paramasaris Cameron and Plesozethus Cameroné

Vou. 1] Bradley.—The Taxonomy of the Masarid Wasps 383

NEw SuBFraMILy Euparagiinae

Genus Euparagia Cresson

1902. Huparagiini, tribe Ashmead. Canadian Entomologist, vol. 34, p. 218.

1879. Huparagia Cresson. Proceedings of the Academy of Natural Sci-
ences of Philadelphia, Entomological Section, vol. 6, p. vi.

1904. Plesiomasaris Cameron, Transactions of the American Entomolog-
ical Society, vol. 30, p. 267.

1905. Odynerus Cameron. Transactions of the American Entomological
Society, vol. 31, p. 380.

1909. Psiloglossa Rohwer. Entomological News, vol. 20, p. 357.

Type—Euparagia scutellaris Cresson; genus monobasic.
Habitat —Southwestern North America.

&. Head wider than the thorax; eyes large, nearly 3 times farther
apart than are the posterior ocelli, emarginate; an inconspicuous
tubercle between the antennae; clypeus longer than broad, its anterior
margin medially produced and bidentate; temples margined poste-
riorly by a fine carina, reaching to the mandibles; occiput bordered
above by a second fine carina, caudad of the one bordering the temples;
mandibles ending in two nearly equal teeth; ligula broad, not retrac-
tile, flat, its apex deeply acutely notched, but little longer than the
paraglossae; labial palpus long, about as long as the stipes, 4-seg-
mented; maxillary palpus longer than the stipes, 6-segmented. An-
tenna consisting of 13 segments, the scape about twice as long as the
pedicel, equal to segment 3, segments 3 to 6 longer than broad, seventh
about equal in length and breadth, 8 to 13 broader than long, together
slightly fusiform.

Humeri weakly prominent; parapsidal furrows wanting; tegula
semicircular, sealelike, not covering the base of the scutellum, its outer
margin entire; angles of propodeum marked only by a ridge.

In the forewing m-cu attached to the junction of Cu, and M,.
Tarsal claws simple; apex of the front trochanter with a long inferior
claw; anterior femur somewhat contorted, with a prominent inferior
tooth at base; the tibia normal, about as long as the tarsus, with an
illy formed strigil; the metatarsus a little shorter than the following
4 segments together, the fourth segment as long as broad; middle
femur unarmed, the apical half of the under surface flat; the tibia
unarmed, a little compressed, sometimes bearing a single apical spur,
in other cases with two, a trifle shorter than the tarsus; metatarsus
two-thirds as long as the remaining segments together, the fourth
segment as long as broad; posterior tibial spur acute, the apical half
of the inner margin oblique and armed with 3 large basal and 2 minute
apical teeth ; the tibia about four-fifths as long as the tarsus; the meta-
tarsus a little shorter than the following segments together.

384 University of California Publications [ ENToMoLOoGY

Abdomen sessile, unarmed, the venter flat, the last ventral segment
nearly semicircular. Squama forming an acute upturned hook, not
fused with the ramus; sagitta of irregular shape, bearing an appar-
ently movable oval process, the apical part of which is scarcely chiti-
nized and bears a patch of setae; uncus broad, flat, obtuse, not barbed
at base, but with a minute tooth about the middle of either side.

2. Eyes not so large as in the male, their emarginations less sharp,
the face, between the eyes, much broader than in the male, the bases of
the antennae much more distant from the eyes; clypeus less deeply
bifid at apex than in the male; mandibles blunt; with a tooth on the
inner margin before the apex; antennae as in the male, but 12-seg-
mented.

Trochanter, femora and other segments of the legs simple. The
middle tibia show sometimes one and sometimes two apical spurs.

The venter is less flat than in the male.

The generic identity of Psiloglossa simplicipes with Euparagia was
suggested to me by Dr. Bequaert. Acting upon this suggestion, we
together established the certainty to our mutual satisfaction.

Kxry TO THE SPECIES oF EUPARAGIA

Vertex with two prominent smooth tubercles behind the ocelli; vertex and front
coarsely, irregularly punctate; pronotum coarsely punctured; mesonotum
rugose; posterior face of propodeum with transverse rugae at the angles,
almost smooth in the middle, but with a few scattered irregular large pune-
tures, its lateral faces smooth medially, finely punctate below and shghtly
aciculate above and behind; clypeus of the female with minute punctuations
and! ‘scattered (Coamser pum etme Ss) soe seees-cencenscnaeeeeseeenee een sncee maculiceps Cameron.

Vertex simple without tubercles, vertex and front uniformly granular punctate;
pronotum finely punctate; mesonotum evenly, finely granular-punctate;
propodeum closely evenly punctulate; clypeus of the female longitudinally

AeTC laters Ae Es ee RRR Le ROCs Sarak) ene eee ee maculifrons Cresson.

Euparagia maculiceps (Cameron)

1904. Plesiomasaris maculiceps Cameron, ¢. Trans. Amer. Ent. Soc., vol.
30, p. 267.

1905. Odynerus simplicipes Cameron, g. Trans. Amer. Ent. Soce., vol. 31,
p- 380. (See Meade Waldo. Ann. and Mag. Nat. Hist., [8],
vol. 14 (1914), p. 404.

1909. Psiloglossa simplicipes Rohwer, 2. Ent. News, vol. 20, p. 357.

New Mexico: Las Cruces, August 31, 1 2 at flowers of Solidago
canadensis, type of simplicipes Rohwer (C. H. T. Townsend). °

Mexico: [types of maculiceps Cameron and simplicipes Cameron,
British Museum]; Guerrero, 3000 feet (Godman & Salvin), [British
Museum, recorded by Meade Waldo].

Vou. 1] Bradley —The Taxonomy of the Masarid Wasps 385

Euparagia scutellaris Cresson
Figures 1, 2, 11, 14, 26, 27, 42, 68-74, 93, 100

1879. Euparagia scutellaris Cresson, %, 2. Proc. Acad. Nat. Sci. Phila.,
Ent. Sec., vol. 6, p. vi.

J. Stout, form somewhat Oxybelus-like. Black, the following parts
amber yellow: clypeus except borders, mandibles except base and apex,
spot at summit of each eye, dorsal surface of pronotum, except postero-
lateral margins, tubercles, tegulae at base and at apex, spot in front
of scutellum, large pentagonal spot at apex of seutellum, claw on front
trochanter, apical third of anterior and tips of middle and posterior
femora, tibiae except for irregular reddish and brown blotches, meta-
tarsi, apical border of dorsal segments 1 to 6 laterally dilated, and on
segments 2 to 6 also medially dilated, and a median spot on each of
ventral segments 2 to 7; the four apical segments of all tarsi reddish
brown; flagellum exeept base of Ist segment, reddish brown, cream
colored beneath. Head and thorax silvery sericeous, mesotergum
brown-sericeous, abdomen somewhat yellowish sericeous toward the
apex.

Head closely, clypeus more sparsely, regularly and rather coarsely
punctulate; dorsum similarly sculptured ; seutellum with a longitud-
inal fossa on each side; pleurae more finely and sparsely punctured
than the dorsum; propodeum with a median channel, shallowly ru-
gosely punctate; the postero-lateral angles forming a ridge but not
earinate. Length, 7 mm.

?. Colored as in male except that the entire head, except spot above
each eye, the antennae, legs, except knees and venter, are black, the
mandibles piceous. The eclypeus is longitudinally aciculate. Other-
wise like the male.

NevapDa: 2 3, 2 2 [types, American Entomological Society].

CALIFORNIA: Claremont, 3 4, 1 2 (C. F. Baker), [Pomona College,
Cornell University, Jos. Bequaert]; mountains near Claremont, 1 ~¢
(C. F. Baker), [Pomona College]; Santa Clara Co. (C. F. Baker),
[Cornell Univ.] ; Sobre Vista, Sonoma Co., 1 9, May 12, 1910 (J. A.
Kusche), [Calif. Acad. Scei.].

Plesiozethus Cameron

21901. Paramasaris Cameron. Transactions of the American Entomolog-
ical Society, vol. 27, p. 312.

1904. Zethoides Cameron. Transactions of the American Entomological
Society, vol. 30, p. 93 (not Fox).

1905. Plesiozethus Cameron. Entomologist, vol. 38, p. 269.

1906.. Metazethoides Schulz. Spolia hymenopterologica, p. 213.

1907. Plesiozethus Cameron. Entomologist, vol. 40, p. 62.

1912. Plesiozethus Zavattari. Archiv fiir Naturgeschichte, vol. 78, pt. A,
no. 2, p. 62.

386 University of California Publications [ENTOMOLOGY

Type.—Plesiozethus flavolineatus Cameron; genus monobasic.

Habitat—Panama; Colombia.

I have not seen a specimen of this genus, and consider its position
very doubtful. I am even doubtful that it is a masarine wasp. Za-
vattari has pointed out the probable identity with Paramasaris, main-
taining that Cameron is incorrect in stating that Plesiozethus has only
2 submarginal cells.

List oF SPECIES oF PLESIOZETHUS

flavolineatus Cameron, ¢, 9. Panama; Colombia.

1904. Zethoides flavolineatus Cameron, g. Transactions of the American
Entomological Society, vol. 30, p. 93.

1905. Plesiozethus flavolineatus Cameron. Entomologist, vol. 38, p. 269.

1906. Metazethoides flavolineatus Schulz. Spolia hymenopterologica, p. 213.

1907. Plesiozethus flavolineatus Cameron. Entomologist, vol. 40, p. 62.

1912. Plesiozethus flavolineatus Zavattari, g, 9. Archiv fiir Naturge-
schichte, vol. 78, pt. A, no. 2, p. 64 (description of female).

Paramasaris Cameron

1901. Paramasaris Cameron. Transactions American Entomological So-
ciety, vol. 27, p. 311.
21904, 1905. Zethoides, Plesiozethus, Cameron.

Type.—Paramasaris fuscipennis, Cameron, genus monobasic.

Habitat—New Mexico.

As already indicated, if Zavattari is correct in his characterization
of Plesiozethus, it is probably identical with this genus.

Paramasaris fuscipennis Cameron

1901. Paramasaris fuscipennis Cameron, 9. Trans. Am. Ent. Soc., vol. 27,
p. 312.

‘‘Black, covered with a white pile; the underside of the antennae
brownish, the apex of the petiole and of the second segment pallid
yellow; the wings fusco-hyaline; the radial cellule smoky; the stigma
and nervures black. 9. Length, 7 mm.

‘‘Antennae shorter than the thorax; the joints of the club clearly
separated, the thickening commencing from the fourth joint. The
front, vertex and the upper part of the eye incision distinctly pune-
tured ; the elypeus is shining and less strongly and closely punctured ;
its apex is distinetly depressed and the sides are roundly narrowed.
The sides of the thorax are more densely covered with a silvery pile

Vou. 1] Bradley—The Taxonomy of the Masarid Wasps 387

than is the upper part; the pro- and mesothorax are closely and dis-
tinctly punctured; on the apex of the mesonotum, in the center, are
two short deep furrows, which are deep, and are wider at the apex
than at the base. The scutellum is more strongly and somewhat more
widely punctured; its basal furrow bears 7 stout longitudinal keels.
The median segment, except on the base and the lower part of the
pleurae, bears large, round, deep punctures; the center in the apex is
smooth, shining and depressed; down the middle of the segment is a
narrow, transversely striated band; the transverse striae being weaker
at the base, and at the apex they are stouter and fewer in number.
The base of the prothorax is keeled all round; behind this keel is
another less distinct one, which curves backwards above to near the
end of the pronotum; on the pleurae the space between the two keels
is striated. Mesopleurae obscurely and sparsely punctured; in front
of the centre are 7 large foveae, the upper 4 are round and deep and
the uppermost is in front of the others; the lower ones are larger and
deeper, are separated by stout keels, and are placed somewhat in front
of those in the middle so that the row of foveae forms a curve; the
apex 1s bordered by a narrow crenulated furrow. On the lower half
of the base of the metapleurae are four deep foveae separated by stout
keels; on the upper half, behind the middle, is a row of smaller foveae ;
the apical half is irregularly closely reticulated. Legs pruinose, black ;
the tarsi with a fuscous tint; the calearia are testaceous. The radial
cellule is distinctly appendiculated, the appendicular cellule being
longer than broad; the second cubital cellule at the top is not one-
fourth of the length of the third, at the bottom, half its length; the
first recurrent nervure is received near the base, the second close to
the middle of the cellule; the transverse basal nervure is interstitial.
Abdomen pruinose; the petiole distinetly longer than the second seg-
ment and punctured ; the punctures at the base more pronounced than
elsewhere ; the apex is depressed and narrowed; near the base of the
narrowed neck is a row of depressed furrows.’’

New Mexico: ‘‘Santa Fé Mts.’’ (I have not been able to learn of
any mountains bearing this name. )

SuBFAMILY Masaridinae
Synoptic TABLE OF THE TRIBES OF MASARIDINAE

Glossae not retractile, short, scarcely exceeding the length of the paraglossae,
the membranous part much shorter than the stipes and quite broad; the max-
illary palpus 6-segmented, of normal length, the labial palpus 4-segmented;
antenna of the male incrassate toward the apex but without a club....Paragiini.

Glossae retractile, very elongate, far exceeding the length of the paraglossae,
at least as long as the stipes and usually many times as long, narrow and
forming a sucking tube; maxillary palpus reduced in size, at most 4-segmented
(except in Ceramiopsis, where it is 6-segmented), often reduced to a mere
tuberele; labial palpus consisting of from 1 to 4 segments; antenna of the
male with the apical segments often fused into an indistinctly segmented
CSU 6) ec Ce ao on ene Masaridini.

388 . Unversity of Californa Publications [ENTOMOLOGY

SyNoptTic TABLE TO THE GENERA OF THE TRIBE PARAGIINI

Eyes emarginate; parapsidal furrows absent .............. Metaparagia Meade Waldo.
Eyes not emarginate; parapsidal furrows distin¢t .................... Paragia Shuckard.

SynoprTic TABLE OF THE GENERA OF THE TRIBE MASARIDINI

A. Maxillary palpus consisting of 6 segments, the labial of 4 ........................--.---
Be aS a Ne ete eae Ceramiopsis Zavattari.

AA. Maxillary palpus consisting of 4 segments, the ligula beneath with many
transverse scale-like appendages.

B. Middle tibia with 2 apical spurs, shorter than the first 4 seaments of the
tarsus united, the fourth segment longer than broad; antenna of the
male like that of the female, gradually incrassate toward the apex,
the penultimate segment much broader than long, the ultimate segment
short, conical, not hooked; mandible of the male slender, acute, with
two serrations on the inner margin somewhat removed from the apex
and a strong basal tooth; clypeus of the male broader than long;
second ventral segment of the male unarmed; squama slender and
acute;7 the uncus reduced to a small basal piece,? not readily observ-
able; the sagitta and the volsella much enlarged and fused with those
Of thevopposite) Sic et ees eect cere ere case eee eee ee Ceramius Latreille.

BB. Middle tibia with only 1 apical spur,’ longer than the first 4 tarsal
segments together,’ the fourth segment broader than long; antenna
of the male unlike that of the female, the penultimate segment on its
outer margin much longer than broad, the ultimate segment long, digiti-
form, forming a hook; mandible of the male obliquely truncate with
3 strong apical teeth but no basal tooth; clypeus of the male longer
than broad; second ventral segment of the male unarmed; squama
clavate;8 uncus elongate;8 sagitta and volsella small and not united
with those of the opposite side8..................---------- Paraceramius Saussure.

BBB. Twelfth segment of the antenna of the male forming a large hook;
clypeus of the male longer than broad; mandible of the male obliquely
truncate and terminating in 3 or 4 short teeth; abdominal segments
constricted at base, the second ventral segment of the male bearing a
tubercle: sx:sccot se Ree hed ae reese nes cena neteearaatens Ceramioides? Saussure.

AAA. Maxillary palpi consisting of 2 or 3 segments.
B. Scape elongate, cylindrical or curved, more than twice as long as the
pedicel.
C. Antenna of the female composed of 7 distinct segments, short; that
of the male composed of 12 distinct segments, strongly inerassate but
not forming a globular club, convex beneath........ Jugurtia Saussure.

7 These characters are drawn from an examination of the male of the type
species, fonscolombei, only.

8 These characters are drawn from an examination of the male of only one
species, lusitanicus, the genotype.

9T have not seen the genotype of Ceramioides, so state the characters indi-
cated by Saussure.

Vou. 1] Bradley —The Taxonomy of the Masarid Wasps 389

CC. Antenna of the female composed of 12 segments, incrassate apically,
but not forming a club; that of the male elongate, reaching the
scutellum, segments 3 to 6 lnear, 7 to 10 incrassate, concave be-
neath; 11 and 12 scarcely separable .................... Masariella Brauns.

BB. Scape and pedicel both globular, the latter at least one-half as long as
the former.

COE WY SEAN EEN: COO SDS Gas ak eee Quartinia Gribodo.

OKO, WiGiHeTe THEN GIP OOM CONES see ee ee Celonites Latreille.

AAAA. Maxillary palpus consisting of but a single very short segment, a mere
tubercle easily entirely overlooked.

B. Middle tibia with 2 apical spurs, spur of posterior tibia not bifid; labial
palpus of the male 4-segmented, the apical segment very short; seventh
ventral segment with a median apical notch, not deep and by no means
reaching to the sixth segment; antenna of the male long, the third,
fourth, and fifth segments linear, the fifth somewhat thickened, the
sixth more strongly so, the seventh to twelfth segments almost indis-
tinguishably fused into a club which is convex beneath; claws with a
small median tooth; last dorsal segment, seen from the side, acute, from
NOONE: Cra baayer atin, SACOM NGL ee eee Masaris Fabricius.

BB. Middle tibia with 1 apical spur; spur of posterior tibia bifid; labial
palpus 1- to 3:segmented; apical margin of seventh ventral segment
of the male either truncate or with a very deep quadrate notch reach-
ing to the sixth segment; antenna of the ¢ variously formed; claws
simple; last dorsal segment, seen from the side obtusely curved, or
truncate; in the latter case the truncature is margined by two strong
inferior and two strong superior teeth, in the former case there are
no teeth, but, seen from above, it is weakly notched at apex.

C. Antenna of the male and of the female dissimilar, that of the male
much the longer, the apex in each sex always with a club, and at
least the first segment of the flagellum and usually more, much
longer than broad; labial palpus in the male consisting of from 1 to
3 segments, in the latter case the last segment shorter than the
preceding; in the female consisting of three segments, the first
long and flattened, the second short, the third much longer than
the wiatee) alee very slender and acute, and ending in two stout
spines; the last dorsal segment is much curved ventrad, its apical
portion vertically truncate, the truncature bordered above and
below by a pair of strong teeth or tubercles, the latter placed
closer together than the former pair; apical margin of the seventh
ventral segment with a very deep, usually rectangular emargina-
tion, reaching basad to beneath the sixth segment; squama always
lamelliform and obtuse, never ending in a spine or hook; uncus
never broad and flat, but slender and usually acute, and often de-
CADE CEX6 GFT Fe eh) Oe ei ce ae A OE een Pseudomasaris Ashmead.

CC. Antenna of the male and of the female similar, except that that of
the latter has 11, of the former, 12 segments, but slightly incras-
sate, the segments of the flagellum all as broad and mostly broader

390 University of Califorma Publications [ ENTOMOLOGY
than long; labial palpi 3-segmented in both sexes;19 last dorsal
segment hood shaped, rounded to meet the venter, without apical
teeth or truneature, but slightly transversely emarginate at apex;
apex of the seventh ventral segment truncate; uncus very broad
and strongly depressed; squama ending in a sharp, strongly de-
GUTAVC CLL Okage eee nae oe eee Trimeria Buysson.

Genus Paragia Shuckard
Figures 3, 13, 24, 25, 43-45, 75, 76, 91, 92, 94, 101.

2. General form like Vespa. Head large; the occiput immargined ;
the temples broad, immargined, eyes not incised, but the inner margins
sinuate, reaching the mandibles, separate from one another above by
a distance equal to their own length; ocelli close, in a small triangle
which is slightly broader than high; front moderately prominent ;
clypeus moderately prominent, the anterior margin produced and trun-
eate; labrum short, bilobed; mandibles short and stout, the imner
margin with two teeth.

The ligula is short and not retractile, composed of the two strap-
like glossae, which are about the length of the labial palpus, strongly
divergent, united only for a short distance at their bases. The para-
glossae are similar in appearance to the glossae and but little shorter.
The glossae and paraglossae are all tipped with a chitinous button,
such as is often found in Eumeninae. The dorsal hind margin of the
glossa bears a series of elongate and very broadly transversely flattened
setae, analogous to the seales found in Ceramius and Paraceramius,
and the anterior dorsal margin is fringed with smaller and less con-
spicuous setae, somewhat flattened in the opposite diameter. At the
base of the glossa, mesad of the paraglossae is a membranous lobe armed
with a group of minute tubercles, and between the two of these, in the
median line, is a heavily chitinized tongue-shaped piece, the tip of
which is turned upward ; the labial palpus is 4-segmented, non-elongate,
the first segment stout and widened at apex, the last half as long.
Between the two palpi, in the median line, an anterior tongue-like
extension of the heavily chitinized mentum and submentum is thickly
set with sensory setae. The point of this process is very acute, turned
upward and may be distinctly seen from the dorsal surface. On the
dorsal side at the base of the glossa and bearing this at their apex are
two chitinized plates, with a partly lateral, partly dorsal surface,
joined at their bases, and formed like a letter V with expanded arms.
Laterad of these, and somewhat enclosing them, is on each side a chiti-
nized roughly triangular piece, with acute apex situated at the bases
of the paraglossae. The inner margins of these two pieces are fringed
with a double comb of spines; a chitinized band on either side between
these pieces passes forward beneath the combs to near their apices, then
ventrad between the glossae and paraglossae articulating on the ventral

10T have seen only the male, but Saussure states that they are 3-segmented
in the female.

VoL. 1} Bradley —The Taxonomy of the Masarid Wasps 391

surface with the tongue-shaped chitinized piece which has been de-
seribed as lying at the base of the glossae. The maxillae present
features of considerable interest. The cardo is of the normal form,
bent at right angles to the stipes; the inner surface of the latter meets
the ventral in a sharp ridge, crested apically with a comb of bristles.
The palpi are 6-segmented, the last two segments together equalling
the third in length. The apical portions are turned, so that from a
strictly ventral view one observes the edges of the lobes, rather than
their surfaces. From a somewhat external aspect, opposite the base
of the palpus, there is a triangular sclerite, projecting dorso-entad,
and bearing a few spines. This is ordinarily interpreted as the lacinia.
Very closely and broadly attached to its base is a large lobe extending
cephalad, and to the inner upper ‘margin of this is attached a second
narrow lobe. The latter has on its upper margin a still narrower third
lobe. These three lobes seem to correspond to what usually together
pass for the mesal lobe of the galea, but this insect would suggest that
they may really be part of the lacinia. From an inner view of the
maxilla there is seen attached to the apex of the ecardo a prominent
but searecely chitinized oval lobe, margined dorsally with a thick fringe
of bristles, and with a few longer setae on the lower margin. This
lobe is present in all Masaridae that I have examined, and may repre-
sent the basal lobe of the galea. Apicad of it is a small ear-shaped
lobe, very prominent because of being more heavily chitinized than
the other parts, and which probably is the outer lobe of the galea.

The seape is elongate, slightly compressed ; the pedicel very short ;
the flagellum is inerassate toward the apex but without forming a club,
composed of 10 distinct segments, the first almost equal in length to
the following 4 united.

Humeri prominent, slightly angled ; parapsidal furrows distinct ;
the tegula small, seale-like, oval, the outer margin entire, by no means
reaching the base of the seutellum; this is prominently elevated, but
with its surface flat, covering the postseutellum; propodeum sloping
directly to its apex, i.e., without dorsal surface, and without any lateral
angles or even ridges.

Cells R, and R, united (i.e., two closed submarginal cells) and
embracing both veins M,,, and M,; m-cu arising from Cu,, which at
that point is deflected to meet M,. Hind wing with a small but dis-
tinct anal lobe. Anterior trochanter unarmed; all the segments of the
legs with regular convex or slightly flattened surface, without ridges
or tubereles; spur of anterior tibia broad, acute, with a tooth on its
convex margin; the middle tibia has two apical spurs; the larger spur
of the hind tibia has its apex obliquely tridentate, the margin basad
of the inner tooth pubescent ; all claws are large and with a large sharp
tooth at base.

The abdomen is like that of a Vespa.

J. General appearance of a Monobia or Eumenes; head transversely
rectangular, the temples broad, not margined behind; ocelli in a low
triangle, distant from the eyes; the latter with their inner margins
sinuate but not emarginate ; clypeus prominent, with anterior margin

392 University of California Publications [ENTOMOLOGY

strongly produced and abruptly truncate, concealing or nearly con-
cealing the small labrum ; mandibles broad, with two large teeth before
the apex. Antennae long and slender, of 12 segments, the scape long,
the third segment still longer, the eleventh and twelfth segments some-
times incised beneath (tricolor).

Humeri rounded; parapsidal furrows distinct, scutellum elevated,
flattened on top; propodeum shghtly concave, the lateral angles
rounded, without tooth above (genotype) or with a blunt tooth in some.

Forewing with two submarginal cells; m-cu attached to Cu,, which
is at that point deflected caudad a very short distance to join M,.
Anterior trochanter unarmed ; all femora and tibia with even surfaces ;
middle tibia with two apical spurs; posterior tibial spur as in the
females; claws large; much curved, with a strong tooth beneath near
the base.

Abdomen shaped as in Vespa, the last dorsal segment ending in
two lobes, with a shallow notch between; second ventral segment with
an acute median prominence behind.

Habitat—Australia.

List oF SPECIES oF PARAGIA

australis Saussure, 3%, 9. Australia.
bicolor Saussure, ¢, 9. Australia.
bidens Saussure, @. Australia.
calida Smith, g. Australia.
concinna Smith, 9. Australia.
deceptor Smith, 9. Australia.
decipiens Shuekard, ¢, 9. Australia.
excellens Smith, ¢, 9. Australia.
hirsuta Meade Waldo, ¢. N. Queensland.
1911. Paragia hirsuta Meade Waldo, g. Ann. and Mag. Nat. Hist., (8), -
vol. 8, p. 749.
magdalena Turner, 9. Queensland.
1908. Paragia magdalena Turner. Trans. Ent. Soc. London, (1908), p. 89
moroso Smith, 9. Australia.
nasuta Smith, 9. Australia.
odyneroides Smith, g. Australia.
perkinsi Meade Waldo, 9. Queensland.
1911. Paragia perkinsi Meade Waldo, 9. Annals and Magazine of Natural
History, (8), vol. 8, p. 750.
praedator Saussure, 9. Australia.
saussurtt Smith, 9. Australia.
sobrina Smith, 9. Australia.
tricolor Smith, ¢, 9. Australia.
venusta Smith, 9. Australia.
vespiformis Smith, g, 9. Australia.
walkeri Meade Waldo, ¢. Australia.
1910. Paragia walkeri Meade Waldo, ¢. Annals and Magazine of Natural
History, (8), vol. 5, p. 33.

Vou. 1] Bradley.—The Taxonomy of the Masarid Wasps 393

K&ryY TO THE SPECIES

Meade Waldo: Ann. and Mag. Nat. Hist., (8), vol. 5 (1910), p. 31. The following
species not included: australis, bicolor, hirsuta, perkinsi.

Genus Metaparagia Meade Waldo

Paragia auctores, pars.
1911. Metaparagia Meade Waldo. Annals and Magazine of Natural
History, (8), vol. 8, p. 748.

This genus I have not seen.
Type—Paragia pictifrons Smith, by original designation.

SPECIES oF METAPARAGIA

doddi Meade Waldo, 2. N. Queensland.
1911. Metaparagia doddi Meade Waldo, 9. Ann. and Mag. Nat. Hist.,
(8), vol. 8, p. 748.
maculata Meade Waldo, ¢, 9. Australia.
1910. Paragia maculata Meade Waldo, ¢, 9. Ann. and Mag. Nat. Hist.,
(8); voli 5, p: 32.
1911. Metaparagia maculata Meade Waldo. Loe. cit., (8), vol. 8, p. 749.
pictifrons (Smith) Meade Waldo, ?. Australia.
1857. Paragia pictifrons Smith.

Key To Species or METAPARAGTA
Meade Waldo. Ann. and Mag. Nat. Hist., (8), vol. 8 (1911), p. 749.

Genus Ceramiopsis Zavattari

1910. Ceramiopsis Zavattari. Annali del Museo ecivico di storia naturale,
Genova, (3), vol. 4, p. 533.

1912. Ceramiopsis Zavattari. Arch. f. Naturgeschichte, vol. 78, pt. A,
no. 2, p. 60. Figure of abdomen and description.

This genus I have not seen.
Type.—Ceramiopsis gestrot Zavattari, genus monobasic.
Habitat. Brazil.

SPECIES OF CERAMIOPSIS

gestrot Zavattari, 2. Brazil.
1910. Ceramiopsis gestroi Zavattari, 9. Annali del Museo civico di storia
naturale, Genova, (3), vol. 4, p. 533.
1912. Ceramiopsis gestroi Zavattari, 9. Arch. f. Naturgeschichte, vol. 78,
pt. A, no. 2, p. 60.

394 University of California Publications [ ENTOMOLOGY

Genus Ceramius Latreille
Figures 9, 12, 15

1904. Huceramius Dalla Torre. Genera Insectorum, fase. 19, p. 5.

¢. Head broad, quadrate; clypeus broader than long, produced
medially and truneate; glossa retractile, but short, when fully ex-
tended but little longer than the stipes; both palpi 4-segmented ; the
maxillary palpus small.

Humeri rounded ; parapsidal furrows present but not deep; tegula
not elongate, scale-like and without coarse punctures, the outer margin
entire; angles of propodeum entirely rounded.

Medio-cubital cross-vein attached to Cu,; spur of anterior tibia
flattened, arched, with a transparent upper margin and bifid tip;
anterior trochanter alate at apex; front femur 3-sided, twisted ; middle
and hind femora and tibiae compressed, their surfaces regular; an-
terior and middle claws with a small median tooth, that of the hind
claw minute; middle tibia with two nearly equal spurs; longer spur
of hind tibia acute, simple.

Sixth sternite with a very deep median notch exposing a smooth
and highly polished area of the seventh; the apex of the seventh pro-
duced into a thickened and truneate lobe.

Genitalia as described in the table and illustrated in figures.

The above characters are drawn from the type species. I have not
seen a female.

In its genitalia this departs more radically from the usual type of
the family than does any other genus which I have examined. The
peculiar series of transverse erect scales beneath the ligula I have not
observed elsewhere except in Paraceramius.

Type—Ceramius fonscolombet.

Habitat—Afriea, Southern Europe, Caucasus.

SPECIES OF CERAMIUS1?1

beyeri Brauns, ¢, 9. Cape Colony.
1903. Ceramius beyeri Brauns, ¢, 9. Zeitsch. f. systemat. Hymenopter-
ologie u. Dipterologie, vol. 3, p. 69.
caffer Saussure, 9. Cape Colony (probably a variety of lichtensteinit).
capensis Saussure, 9. Cape Colony.
[capicola Brauns, ¢, 2. Cape Colony. See Ceramioides. |
caucasious Andre, @. Caucasus.
consobrinus Saussure, 3, 9. Cape Colony.
1913. Ceramius consobrinus Brauns, ¢, 2. Entomologische Mitteilungen,
vol. 2, p. 194. (First description of male.)
fonscolombei Latreille, g, 9. Mediterranean subregion.
[fumipennis Brauns, 3, 9. Cape Colony. See Ceramioides. |

11 Although these species all stand in literature under this genus, they many
of them doubtless belong to Ceramioides or Paraceramius.

Vou. 1] Bradley—The Taxonomy of the Masarid Wasps 395

hispanicus Dusmet, 3, 9. Spain.
1908. Ceramius hispanicus Dusmet. Mem. Pri. Congr. Nat. Espan., p. 180.
karrooensis Brauns, g. Cape Colony.
1902. Ceramius karrooensis Brauns, g. Zeitsch. f. systemat. Hymenop-
terologie u. Dipterologie, vol. 2, p..282; vol. 3, p. 68.
lichtensteinii Klug, @, 9. Cape Colony.
1906. Ceramius rufomaculatus Cameron. Trans. South Afriean Philos.
Soc., vol. 16, pt. 4.
1913. Ceramius lichtensteinti Brauns. Entomologische Mitteilungen, vol.
2 Joe lB}, yall, Ph sites IL.
var. macrocephalus Saussure.
1908. Ceramius macrocephalus Brauns, 3, 9. Zeitsch. f. systemat. Hymen-
opterologie u. Dipterologie, vol. 3, pp. 65, 68.
1908. Ceramius lichtensteini var. macrocephalus Brauns, ¢, 9. Entomolo-
gische Mitteilungen, vol. 2, p. 193.
[ macrocephalus Saussure. See lichtensteinii var. macrocephalus. |
oraniensis Saussure, ¢, 9. Algeria.
peringueyt Brauns, 9. Cape Colony.
1913. Ceramius peringueyi Brauns, 9. Entomologische Mitteilungen, vol.
2, p. 194.
rex Saussure, 9. Cape Colony. (Probably a variety of lichtensteinii.)
[rufomaculatus Cameron. See lichtensteinii. |
[schulthessi Brauns, @, 9. Cape Colony. See Ceramioides. |
vespiformis Saussure, 9. Cape Colony.

Genus Paraceramius Saussure
Figures 36, 102

&. Head quadrate, not as broad as in Ceramius; clypeus elongate,
not as squarely truneate as in Ceramius; the glossae retractile, elong-
ate, forming a tubular ligula, with a peculiar series of transverse erect
scales beneath, as in Ceramius; palpi as in Ceramius.

Humeri entirely rounded; parapsidal furrows wanting; tegula
small, seale-like, without coarse punctures, the outer margin rounded,
entire; angles of propodeum entirely rounded.

Venation as in Ceramius. Spur of anterior tibia as in Ceramius
except that it ends in a lobe and a spine instead of two nearly equal
spines; anterior trochanter simple, the femur with a sharp crest be-
neath; all claws with a large basal tooth; middle tibia with one apical
spur; the larger spur of hind tibia with 3 short spines before its tip.

Sixth and seventh sternites as in Ceramius.

Genitalia as described in the table and illustrated in figure 8.

The above characters apply to P. lusitanicus (Klug). I have not
seen a female.

The single spur on the middle tarsus, the difference in the spurs
on the anterior and hind tarsi, the simple front trochanters, the great
genitalic and other differences make it impossible to include this group
any longer as a subgenus of Ceramius.

Habitat—Korea, Southern Europe, Africa.

396 University of California Publications | ENTOMOLOGY

SPECIES OF PARACERAMIUS

koreensis Radoszkowski, 9. Korea.

linearis Klug, ¢. Cape Colony.

lusitanicus Klug, g, 9. Southern Europe.

var. luteoclypeata Dusmet, gf. Spain.
1908. Ceramius lusitanicus var. luteoclypeata Dusmet. Mem. Pri. Congr.

Nat. Espan., 1908, p. 180.

[nigripennis Saussure. See Ceramioides. |

spiricornis Saussure, ¢, 9. France and Spain.

Genus Ceramioides Saussure

¢. Eyes very distant from each other and from the ocelli formed on
the vertex, their inner margins very broadly and shallowly emarginate,
but little more than sinuate; clypeus flat, longer than broad, its an-
terior margin produced and squarely truneate ; mandibles rather broad
and flat, two teeth on the inner margin before the apex, labial palpi
4-seomented. Antennae long and slender, of 12 segments, a very little
widened before the apex, the eleventh segment longer than broad, the
twelfth more than twice as long as the eleventh, tapered and recurved,
forming an apical hook.

Humeri entirely rounded; parapsidal furrows weak anteriorly but
forming two deep grooves near the middle line just before the scutel-
lum; tegulae short, scale-lke, not covering the base of the scutellum,
impunctate ; dise of seutellum flat, posteriorly broadly rounded, cover-
ing and concealing the rounded postscutellum ; posterior face of propo-
deum small, flat, rounded into the sides below, but superiorly sharply
truncate.

In the forewings m-cu inserted shortly basad of M,-+ Cu,, Cu, at
the point of insertion of m-cu turning caudad for a short way to meet
M,. Anterior trochanter produced at apex into an elongate scale,
tibiae and femora with even surfaces except that the anterior femur
is widened beneath before the middle; middle tibia with a single spur
at apex; larger posterior tibial spur with two spines and a tooth on
its margin toward the apex; claws with a small tooth on the inner
margin at its middle.

Abdomen subeylindrieal, flattened beneath, the second, third, and
fourth dorsal segments somewhat constricted basally; the last dorsal
segment rounded and unarmed; the third ventral segment armed with
two tubercles; the seventh ventral segment posteriorly produced ven-
trad and pointed.

2. Differs from the male in the following particulars: elypeus with
its anterior margin broadly rounded, indistinctly separated from the
front, antennae much shorter, weakly incrassate from the third seg-
ment to the apex, the apical segment as broad as long, no longer than
the preceding, the third segment more than twice as long as the pedicel,
longer than segments 4 and 5; tooth of the claws larger ; second ventral
segment unarmed and last ventral apically simple and rounded.

Vor. 1] Bradley —The Taxonomy of the Masarid Wasps 397

Generic description drawn from capicola Brauns. I have not seen
the genotype.

Nigripennis Sauss. (det. Brauns) agrees in all respects except that
parapsidal furrows are distinct throughout and 2 tubercles are on third
instead of second segment.

Type.—Ceramius cerceriformis Saussure, genus monobasic.

Habitat —South Africa.

SPECIES OF CERAMIOIDES

cerceriformis Saussure, ¢. Cape Colony.
capicola Brauns, ¢, 2. South Africa.
1902. Ceramius capicola Brauns. Zeitsch. f. systematische Hymenopter-
ologie u. Dipterologie, vol. 2, p. 278; vol. 3, p. 68. (On p. 280,
lines 14—48 apply to fuwmipennis instead of to capicola. In line 17,
p- 279, ‘‘dorsale’’ should: read ‘‘ventrale.’’)
fumipennis Brauns, ¢, 9. Cape Colony.
1902. Ceramius fumipennis Brauns, 3, 9. Zeitsch. f. systematische Hyme-
nopterologie u. Dipterologie, vol. 2, p. 275; vol. 3, p. 68. (On
p- 280, lines 15-48 apply to this species.)
schulthessi Brauns, ¢, 9. Cape Colony.
1902. Ceramius schulthessi Brauns, 9. Zeitsch. f. systemat. Hymenopter-
ologie u. Dipterologie, vol. 2, p. 182.
1913. Ceramius schulthessi Brauns, gf, 2. Entomologische Mitteilungen,
vol. 2, p. 1196, pl. 2, fig. 6.
nigripennis Saussure, ¢, 2. Cape Colony.
1913. Ceramius nigripennis Brauns, J, 2. Entomologische Mitteilungen,
vol. 2, p. 201, pl. 2, fig. 3. (First description of the male.)

Genus Jugurtia Saussure

1894. Jugurthia Dalla Torre. Catalogus Hymenopterorum, vol. 9, p. 5.

°. Shape and general appearance of female of Psewdomasaris, eyes
very widely separated above, deeply incised, the incision broadly
rounded ; clypeus seareely convex, its apical border trilobed; labrum
semicircular ; mandibles acute, two teeth on the inner margin; ligula
retractile, labial palpi 4-segmented, the fourth segment, however, a
minute tubercle, the third about equal in length to the second, and
bearing three stout curved spines. Antennae as in Pseudomasaris.

Humeri rounded dorsally, margined laterally ; parapsidal furrows
absent ; mesonotum flattened but not depressed before the slightly ele-
vated scutellum, tegula elongate, pointed, covering the base of the
scutellum, posteriorly punctate, its outer margin entire; apical part
of scutellum with a weak depression, the apex weakly bifid ; seutellum
covering and concealing the rounded postscutellum ; propodeum poste-
riorly flat, its lateral angles weak, neither dentate nor mucronate.

398 University of California Publications [ENTOMOLOGY

The medio-cubital cross-vein attached opposite to M,-+ Cu,.
Middle tibia with two apical spurs beneath, a short one above, posterior
tibia with its larger apical spur bifid.

Abdomen as in Pseudomasaris, the second ventral segment with a
transverse ridge, the last segment broadly rounded at apex.

Habitat —Asia, Southern Europe, Africa.

SpEcIES oF JUGURTIA

chlorotica Morawitz, 9. Transcaspia.
escalerae Meade-Waldo, 9. Persia.
1910. Jugurtia escalerae Meade-Waldo, 9. Ann. Mag. Nat. Hist., (8),
vol. 5, p. 33.
[neotropica Mocsarya. See Trimeria neotropica. |
numida Saussure, g. Algeria.
oraniensis Saussure, ¢, 9. Spain, Algeria.
simpsoni Meade-Waldo, 9. Gambia.
1911. Jugurtia simpsoni Meade-Waldo, 9. Ann. Mag. Nat. Hist., (8),
vol. 8, p. 448.

TABLE TO THE SPECIES OF JUGURTIA

Meade-Waldo: Ann. Mag. Nat. Hist., (8), vol. 8 (1911), p. 449 (chlorotica
omitted).

Genus Masariella Brauns

Figures 5, 40, 41, 106
Masaris auct. pars.
1905. Masariella Brauns. Ann. Hist. Nat. Musei Nat. Hungarici, vol.-3,
p- 223.

3. Head transverse rounded, the posterior margin of the vertex
somewhat concave; temples moderately broad, margined posteriorly ;
ocelli distant from the eyes, these very deeply emarginate, the emargi-
nation broadly rounded at apex; clypeus like that of Pseudomasaris
gibbous, or merely convex, the anterior margin deeply and broadly
coneave; the labrum prominent, short, rounded at apex; mandibles
acute, with one or two teeth within; ligula elongate, retractile, like
that of Masaris; labial palpi, 4-segmented, the apical segment, minute ;
maxillary palpi said by Brauns to be 2-segmented. Antennae of the
genotype consisting of 12 segments, gradually enlarged into a long
oval club from the sixth segment to the apex, flattened but not concave
beneath, the club terminating in a slight hook and not distinetly de-
mareated from the rest of the flagellum, the divisions between all seg-
ments distinct, except that the one between the last two is largely
obliterated, the scape a little longer than the third segment, more than
twice the length of the pedicel; the antennae of saussuret are different,
the club short, broadly ovate, convex, commencing with the ninth
segment.

Vou. 1] Bradley —The Taxonomy of the Masarid Wasps 399

The rather long neck, and the shape of the head and prothorax are
suggestive of Yiphydria; the humeri entirely rounded, parapsidal fur-
rows absent; scutellum in the genotype with a median furrow and
ending in two tubercles, in sawssuret, however, simply longitudinally
coneave; tegula elongate, bluntly rounded, not pointed posteriorly,
covering the base of the scutellum, its outer margin entire, coarsely
punctured ; propodeum concave posteriorly, its side rounded, not eari-
nate, but superiorly mucronate or dentate.

Medio-ceubital cross-vein opposite M,-+ Cu,; anterior trochanter
unarmed ; the tarsal spur arcuate, simple; the tarsus not much short-
ened and flattened; femora and tibiae with simple surfaces: middle
tibia in the genotype with one large apical spur beneath and an addi-
tional short stout spine at apex on the fuels side ; in saussurez. there
are two stout spurs beneath and one above; the posterior tibial spur,
bifid as in Pseudomasaris ; claws with a small tooth beneath near the
base.

Abdomen eylindrieal, the last dorsal segment unarmed, the apical
margin broadly emarginate; ventral segments unarmed, the seventh
of the genotype with a deep depressed pocket at base extending be-
neath the sixth segment, its apex except in saussuret broadly, not
deeply rectangularly emarginate, the apical border in the middle of
the emargination produced into two teeth with a deep rounded notch
between.

Genitalia not examined.

?. Differs from the male in the following particulars: elypeus
convex but not gibbous; antennae much shorter, the segments of the
flagellum short, forming a compact oval club, not unhke that found in
females of Pseudomasaris; spur of anterior tibia shorter and broader ;
anterior tarsus shorter and flattened ; abdominal segments as in female
Pseudomasaris, the apical segments unarmed and broadly rounded at
apex. I have not dissected the mouth parts, but the labial palpi have
three segments, the third equal to the second, and bearing one or more
stout curved spines near the apex, and there may be a fourth segment
represented by a minute tubercle; the maxillary palpi I cannot make
out.

Type—Masaris alfkent (Du Buysson), genus monobasie.

Habitat—South Africa.

Kery TO THE SPECIES OF MASARIELLA
Males
Club of antenna elongate, oval, not sharply differentiated from rest of flagellum,
slightly hooked at tip and flattened beneath; seventh ventral segment with
a conspicuous deep basal pocket, extending beneath the sixth, its apical
margin broadly rectangularly emarginate, in the middle with two teeth separ-
ated by a deep and rounded median notch ...........-........ alfkeni (Du Buysson).
Club of antenna very short, broadly ovate, convex beneath, shaped as in Pseudo-
masaris texana, the tip bluntly rounded, not hooked; seventh ventral segment
slightly produced and truncate at apex, considerably obscured by numerous
long silky hairs at its base with a weakly marked shallow pocket extending
bene athe ben staat eS e om emit peesns ces csne cere w een nee encase roener nese saussurei Brauns.

400 University of California Publications | ENTOMOLOGY

Females

Propodeum with posterolateral angles weakly angled above but not dentate;
scutellum with a discal depression, not ending in two tubereles —.........00.......
oa Ae gs caan te eee io cee at ae Seared woe FOS a es Saas us eaee eS eetcmne ee San neat ee oe ee ene ee saussurei Brauns.

Propodeum with its posterolateral angles dentate above; scutellum with a longi-
tudinal median fossa and ending in two tubercles........ alfkeni (Du Buysson).

M. saussuret undoubtedly is more closely related to alfkeni than it
is to the genotype of Masaris, in which genus it has heretofore stood,
as the following considerations will show. In Masaris the ventral
segments two and three are both armed with strong processes, in
alfkeni is probably more lke it than that of sawsswrev; the ocelli of the
ment is elongate, tapered, and ends in two prominent tubercles or teeth,
while in both alfkent and saussurez it is short and rounded, the apical
margin broadly emarginate; in both alfkent and saussurei the larger
spur of the hind tibia is bifid, but not so in Masaris. Neither the an-
tenna of the male of alfkent or sausswret is like that of Masaris, but
alfkent is probably more like it than that of sawsswrez, the ocelli of the
male of Masaris are close to the eyes, those of both sawssuret and
alfkent much more distant, this character being due to the much closer
approach of the eyes to each other on the vertex in Masaris; the post-
scutellum of Masarvs is not covered by the scutellum, and is bifid, while
in both saussuret and alfkend it is entirely covered by the seutellum
and rounded, and finally, the subalate lateral angles of the propodeum
are of a very different type from that which obtains in both alfkeni
and saussuret.

It is probable that the other South African species now placed in
Masaris will go in Masariella also, but as I have not seen specimens I
eannot say. It is further probable that after the species have been
thus studied it may become desirable to erect a separate subgenus for
saussurei and probably others on the basis of the difference in the
antennae and seventh ventral segment of the male.

SPECIES

alfkent (Buysson) Brauns, g, 2. South Africa.
1904. Masaris alfkeni Buysson, 9. Bulletin de la Société Entomologique
de France, p. 144.
1905. Masariella alfkeni Brauns, ¢. 9. Annales Historico-Naturales Musei
Nationalis Hungarici, vol. 3, p. 223.
saussurei Brauns, 3g, 2. Cape Colony.
1905. Masaris saussurei Brauns, %, 9. Annales Historico-Naturales Musei
Nationalis Hungarici, vol. 3, p. 219.

Vou. 1] Bradley.—The Taxonomy of the Masarid Wasps 401

Genus Quartinia Gribodo

1904. Quartinia Cameron. Zeitschrift fiir systematische Hymenopterol-
ogie und Dipterologie, vol. 4, p. 89.

3. Head broad, transverse, the eyes very distant from each other
and from the ocelli; somewhat, as in the females of Pseudomasaris,
deeply incised, the incision broadly rounded, elypeus convex; its apex
broadly emarginate ; mandibles acute, a tooth on the inner margin near
the apex. Antennae shorter than the distance between the eyes on
the top of the head, resembling those of Pseudomasaris females; the
scape is scarcely longer than broad, the pedicel very large, globose,
nearly as large as the scape, the third and following segments minute,
the eighth and twelfth united into an oval club, convex on all surfaces.

Humeri rounded, parapsidal furrows wanting, mesonotum flattened
but not depressed in front of the slightly raised scutellum ; tegula very
large, considerably larger than the seutellum, very broadly rounded
rather than pointed posteriorly, covering the base of the scutellum,
with a few coarse punctures on the posterior part, this convex, rounded
at apex where it covers the rounded postscutellum ; propodeum poste-
riorly deeply concave, the margins thereof forming a sharp ridge sep-
arating the posterior from the lateral surface, but not carinate and
without spine or tooth.

Forewings completely plaited longitudinally as in Celonites or
Vespa; the medio-cubital cross-vein attached opposite to M, + Cu, ;
in variegata veins, M, and m appear as a mere trace, but not in the
genotype, a completely enclosed and very large triangular appendicu-
late cell present. Anterior trochanter unarmed ; middle tibia with two
weak apical spurs. I cannot make out the posterior tibial spur of
variegata, but in deleta 9 it appears to be slender and acute; claws
small, apparently with a minute tooth beneath.

Abdomen shaped as in Vespa, the last dorsal segment short and
rounded, its apex margined and deeply bifid; apical margin of last
ventral segment sinuate with a broad median tooth.

°. Except in the broadly rounded last abdominal segment the
female does not differ from the male.

The generic description is drawn chiefly from Q. variegata Brauns,
but the 2 of the genotype was compared with it. It is not clear, how-
ever, that in the latter the wings are longitudinally plaited, and the
extent of the wing margin caudad of cell M, is much less than in
variegata.

Type—Quartinia dilecta Gribodo, genus monobasic.

Habitat —Africa, India.

402 University of California Publications [ ENTOMOLOGY

SPECIES oF QUARTINIA
capensis Kohl. Algo Bay.
1898. Quartinia capensis Kohl. Termeszetradji Fiizetek, vol. 21, p. 365.
dilecta Gribodo, ¢, 9. Tunis.
indica Cameron. Deesa.
1904. Quartinia indica Cameron. Zeitschrift fiir systematische Hyme-
nopterologie und Dipterologie, vol. 4, p. 89.
major Kohl. Oran.
1898. Quartinia major Kohl. Termeszetradji Fiizetek, vol. 21, p. 365.
paradoxa Brauns, g. Cape Colony.
1905. Quartinia paradoxa Brauns. Annales Historico-Naturales Musei
Nationalis Hungarici, vol. 3, p. 324.
parvula Dusmet, %. Spain.
1908. Quartima parvula Dusmet. Mem. Pri. Congr. Nat. Espan., 1908,
p. 183.
thebaica Buysson. Egypt.
1902. Quartinia thebaica Buysson Bulletin de la Société Entomologique
de France, 1902, p. 141.

Genus Celonites Latreille
Figures 7, 22, 34, 35, 52, 87-90, 104
1906. Coelonites Du Buysson. Revue entomologique, vol. 25, p. 103.

3. Head transverse, posterior surface flat; occiput margined;
temples wanting; eyes deeply emarginate, the emargination broad and
rounded, distant from each other by three-quarters the length of the
scape; ocelli in a very broad triangle, situated well up on the vertex,
the posterior much closer to the eyes than to each other; front convex,
without tubercles; clypeus strongly convex, shield-shaped, emarginate
anteriorly ; mandibles acute, with two small preapical teeth on the
inner margin; ligula very long, slender and retractile; the labial
palpus consisting of a single segment, the apical portion of which is
partially marked off as a short incomplete second segment; maxillary
palpus consisting of two short, slender segments. Antenna a little
shorter than the thorax, strongly clavate; the scape and pedicel of
nearly equal length, globose, the third segment cylindrical, a little
longer than the pedicel, nearly as long as segments 4 to 5, these as
broad as long, 6 and 7 broader than long, segments 8 to 12 fused into
a solid, large, oval club, convex above and slightly flattened below, the
divisions between the segments distinct beneath; the ninth and tenth
segments in a depression beneath bear the cupuliform organs described
by Saussure.

Humeri angulate; parapsidal furrows wanting; tegula long, cov-
ering the base of the scutellum, pointed, the outer margin weakly
sinuate ; scutellum rather flat ; propodeum raised on each side near its
base, the sides horizontally strongly alate.

Vow. 1] Bradley —The Taxonomy of the Masarid. Wasps 403

Wings longitudinally plaited as in Vespa; cells R, and R, of the
forewing united ; m-cu arising from Cu,. Anterior trochanter simple,
the front tibia with a ridge beneath, but otherwise the femora and
tibiae are without irregular or carinate surfaces; anterior tibial spur
of a peculiar shape, stalked at base, the apical portion triangular,
acute ; middle tibia with two apical spurs; larger apical spur of poste-
rior tibia bifid at tip; all claws with a minute tooth well toward the
base of each.

Abdomen entirely sessile, fitting close against the alate angles of
the propodeum, convex above, tapered toward apex, coneave beneath,
the sides strongly margined; the posterolateral angles of segments 1
to 6 produced into a flattened rounded tooth, giving the margins a
serrate aspect; last dorsal segment with its margin notched, resulting
in 4 teeth; last ventral segment with its apical margin shallowly
concave.

Genitalia of the peculiar type shown in figure 52.

? differs from the male as follows: club of the antenna more
slender, convex beneath and without the ecupuliform organs; mandible
blunt, the inner margin near the apex indistinctly serrulate; labial
palpus of three segments, the first stout, curved, with a ventral row of
four apical setae, the second short, with two setae of which one is
very prominent, the last segment about one and one-half times as long
as the second, curved, slightly widened toward the apex, obtuse, with
a row of setae extending obliquely across the apex, of which four or
five are stout. Wings strongly plaited as in the male; anterior tibial
spur curved, slender throughout, or very slightly widened toward the
tip ; hind tibial spur as in the male; last dorsal segment with its margin
merely weakly sinuate; the last ventral segment large, obtusely pointed
at apex, with a median longitudinal ridge.

Type—Masaris apifornis Fabricius; genus monobasic.

Habitat Mediterranean subregion, Africa.

The difference between the sexes in the labial palpi closely parallels
that found in Pseudomasaris, the condition in the corresponding sexes
being very similar in each genus. The labial palpi are described by
Saussure as of four segments, but this is true of neither sex. That
author did not observe the sexual disparity in the palpi, nor has it
been previously recorded by any author, so far as I am aware. In
respect to the bifid spur of the posterior tibia this genus is also similar
to Pseudomasaris, but of course in many other characters it 1s very
different.

The generic description is drawn from a study of the type species
alone, and it is of course possible that other species may modify it.

404 University of California Publications [ ENTOMOLOGY

List oF SPECIES OF CELONITES

abbreviatus (Villers) Saussure, ¢, 9. Mediterranean subregion.
1793. Masaris apiformis Fabricius.
var. hungaricus Mocsarya, g, 9. Hungary.
andrei Brauns, fg, 9. Cape Colony.
1905. Celonites andrei Brauns, 9. Annales Historico-Naturales Musei
Nationalis Hungarici, vol. 3, p. 228.
1918. Celonites andrei Brauns, f. Entomologische Mitteilungen, vol. 2,
p-. 206. Description of male and of nest.
capensis Brauns, 3, 2. Cape Colony.
1905. Celonites capensis Brauns, 9. Annales Historico-Naturales Musei
Nationalis Hungarici, vol. 3, p. 231.
1913. Celonites capensis Brauns, g. Entomologische Mitteilungen, vol. 2,
p. 205.
clypeatus Brauns, 9. Cape Colony.
1913. Celonites clypeatus Brauns, 2. Entomologische Mitteilungen, vol. 2,
p. 206.
crenulatus Morawitz, 9. Transcaspia.
cyprius Saussure, g. Cyprus.
fischeri Spinola, ¢, 9. France, Algeria.
1906. Celonites fischeri Du Buysson. Revue Entomologique, Caen, vol. 25,
p. 108.
jousseaumei Du Buysson. Obock.
1906. Coelonites jousseaumei Du Buysson. Revue Entomologique, Caen,
vol. 25, p. 104.
mongolicus Morawitz, g, 2. Mongolia.
montanus Moesarya.
1906. Celonites montanus Mocsarya. Annals and Magazine of Natural
History, vol. 4, p. 198.
osseus Morawitz, 9. Transcaspia.
promontorii Brauns, 3g, 9. Cape Colony.
1905. Celonites promontorii Brauns, 9. Annales Historico Naturales Musei
Nationalis Hungarici, vol. 3, p. 232.
1913. Celonites promontorii, g. Entomologische Mitteilungen, vol. 2,
p- 205.
purcelli Brauns, ¢, 2. Cape Colony.
1905. Celonites purcelli Brauns. Annales Historico-Naturales Musei Na-
tionalis Hungarici, vol. 3, p. 226.
1913. Celonites purcelli Brauns. Entomologische Mitteilungen, vol. 2,
p. 205.
rothschildi Du Buysson. East Africa.
1906. Coelonites rothschildi Du Buysson. Revue Entomologique, Caen,
vol. 25, p. 105.
savignyi Saussure, ¢, 9. Egypt.
wheeleri Brauns, ¢, 9. Cape Colony.
1905. Celonites wheeleri Brauns. Annales Historico-Naturales Musei Na-
tionalis Hungarici, vol. 3, p. 230.

Vow. 1] Bradley—The Taxonomy of the Masarid Wasps 405

var. immaculatus Brauns.
1905. Celonites wheeleri var. immaculatus Brauns. Loe. cit, p. 230.
1913. Celonites wheelert var. immaculatus Brauns. Entomologische Mit-
teilungen, vol. 2, p. 205.

TABLES TO SPECIES
South Africa.
Brauns, Hans. Entomologische Mitteilungen, 1913, vol. 2, p. 207.
Mediterranean subregion.
André, Edmond. Species des hyménoptéres d’Europe et d’Algérie .. ., vol.
2, pp. 826-829. Tabulates abbreviatus, fischeri, and cyprius.

Genus Masaris Fabricius
Figures 18, 28, 29, 30, 49-51, 83, 95, 107.

3S. Head transverse ; the occiput immargined ; temples narrow ; eyes
large, distant from one another above by little more than the distance
between the hind ocelli, with a triangular emargination, the apex of
which is obtuse; ocelli in an equilateral triangle, crowded forward,
distant from the occiput, the hind pair almost touching the eyes ; front
comparatively flat, with neither tubercles nor a ridge; elypeus moder-
ately convex, its anterior edge deeply emarginate; mandible acute, the
inner edge with two preapical teeth, the more apical one fair sized ;
labial palpus with four segments, the apical segment very short;
maxillary palpus reduced to a single segment represented by a mere
tuberele. Antenna longer than head and thorax united, gradually
widened into an oval club, which is convex on all sides; the scape sub-
globular; the pedicel short, segments 3 to 5 elongate, cylindrical, 6
and 7 gradually evenly widened, longer than broad; segments 8 to 12
fused, but the divisions distinguishable.

Humeri marked by a ridge; parapsidal furrows absent; tegula
elongate, reaching over the base of the scutellum, the apex obtuse, the
outer margin weakly and broadly emarginate mesally ; scutellum con-
vex; postseutellum prominent, bifid at apex; propodeal angles hori-
zontally subulate, forming acute angles.

Forewing not plaited ; lacking R,; m-cu arising from M,. Anterior
trochanter unarmed; all femora and tibiae with regular surfaces, some
of them shghtly flattened and with a weak ridge beneath, but devoid
of tubercles and earinae; anterior tibial spur slender, shehtly eurved,
the tip acuminate and bent outward ; middle tibia with two stout spurs ;
longer spur of the posterior tibia pectinate at apex (see fig. 95) ; tarsal
claws.each with a minute tooth on the middle of the inner margin.

Abdomen sessile, slender, broadest at base, the first segment from
a dorsal view somewhat concave anteriorly, the second to fifth dorsal
segments somewhat contracted at base, the last bidentate and deeply
notched at apex; second and third ventral segments each with a process,
that of the second acute, of the third larger and transverse ; last ventral
segment with the apical margin weakly concave.

406 Unversity of California Publications [ ENTOMOLOGY

Squama simple, obtuse; sagitta thick, short, roughly trigonal;
uncus slender, acute, decurved. The genitalia are similar in type to
Pseudomasaris.

I have not seen a female. The description is drawn from a male
of the type species, subspecies aegyptiacus.

Type.—Masaris vespiforms Fabricius, by designation of Latreille,
1810.

Habitat —A friea.

Species or MASARIS

discrepans Brauns, 3, 9. Cape Colony.
1913. Masaris discrepans, Brauns, gf, 2. Entomologische Mitteilungen,
vol. 2, p. 203, pl. 2, fig. 9a.
[ saussurei Brauns, J, 9. Cape Colony. See Masariella. ]
spinolae Saussure, 9. Cape Colony.
vespiformis Fabricius, J, 9. Egypt, Algeria.
1911. Masaris vespiformis Meade-Waldo. Annals and Magazine of Nat-
ural History, (8), vol 8, p. 445, illus.
subspecies aegyptiacus Meade-Waldo, ¢. Egypt.
1911. Masaris vespiformis subspecies aegyptiacus Meade-Waldo, ¢. An-
nals and Magazine of Natural History, (8), vol. 8, p. 447, illus.

Genus Pseudomasaris Ashmead
Figures 6, 10, 19-21, 31-33, 96, 108.
1902. Pseudomasaris Ashmead. Canadian Entomologist, vol. 34, p. 221.

3. Eyes deeply emarginate; mandibles acute, with two minute
denticulations on the inner margin; clypeus convex, the apical margin
broadly emarginate; glossae very elongate, retractile, about 7 times
as long as the paraglossae; labial palpus usually 2-segmented, rarely
1- or 3-segmented, in which ease the first segment is much longer than
the following, and the third if present is stout and shorter than the
second, the second segment may be indistinctly separated from the
first; maxillary palpus reduced to a single segment, usually a mere
tubercle, sometimes exceedingly minute; a distinet lacinia present, with
a ciliate inner margin, also a subgalea and sometimes what probably
represent two lobes of the galea, sometimes only one. Segments 8 to
_ 12 of the antennae closely fused into a elub, which may be coneave or
convex beneath, and is of varying shape; scape short, nearly globular,
pedicel short, segments 3 to 5 linear, the sixth and seventh sometimes
widened, occasionally much so.

Tegula elongate, the outer margin notched; parapsidal furrows
absent.

Anterior trochanter unarmed; tarsal claws simple; middle tibia
with one apical spur; posterior tibial spur bifid. Forewing with cells
R, and R, coalesced (2 submarginal cells) ; m-cu arising from M,.

Abdomen sessile, the second, third, and fourth dorsal segments con-
stricted at base, the last dorsal decurved, its apex, seen from the side,

Vo. 1] Bradley —The Taxonomy of the Masarid Wasps 407

truncate, the truncate portion from a caudal view nearly rectangular,
margined above and below by a pair of strong, tooth-like processes, the
inferior pair closer together than the superior, sometimes an additional
pair of tubercles cephalad of the upper pair; second ventral with a
low tubercle, the third with a strong, variously shaped, process ; seventh
ventral segment deeply divided by a usually square broad notch.
Squama and ramus fused, the former lamelliform, rarely thickened,
eurved, often densely ciliate beneath ; sagitta and volsella small, closely
applied to the ramus; uncus usually long and slender, deeurved at
apex, with a pair of barbs at base, sometimes thick and without barbs
at base.

9. Eyes deeply emarginate, the emargination wide and rounded at
apex, eyes at least 3 times as far apart above as the distance between
the hind ocelli; mandibles bluntly rounded or truneate at apex, with
2 denticulations on the inner margin; labial palpi 3-segmented, the
first flattened, the second short, bent at right angles to the second, the
third much longer, faleate, very slender, tipped with 2 stout spines.
Antennae shorter than the width of the head; scape more than twice
as long as pedicel; third segment linear, as long as 3 or 4 following
segments united, segments 4 to 7 increasingly thickened, the seventh
broader than long, segments 8 to 12 fused into an oval club, convex
above and below, the divisions distinguishable, but that between the
eleventh and twelfth sometimes very indistinctly so.

Abdomen unarmed. The segments not constricted at base, the
seventh tergite and sternite with rounded apical margins. In other
respects similar to the male.

Type—Masaris occidentalis Ashmead, by original designation.

Habitat —Southwestern United States.

KEY TO THE SUBGENERA OF PSEUDOMASARIS

A. Posterior metatarsus of the male arcuate, produced at apex on the inner side
into a lobe bearing a prominent crest of ciliae; segments 6 and 7 of the
antennae of the male much longer than wide, the club shaped like the
inverted bowl of a spoon, concave beneath; anterior tarsus of the male
ciliate; anterior tibia and middle femur and tibia of the male contorted,
of the female merely flattened beneath, the tibiae constricted at base;
posterolateral angles of the propodeum alate; last dorsal segment of the
male with 6 tubercles; female with a transverse carina between the an-
tennae; squama very thick, the apex rugose ....................-.-- Toryna, n. subg.

AA. Posterior metatarsus straight or nearly so, without an apical lobe; club of
the male convex beneath, or if concave it is short ovate, not spoonlike,
the seventh segment as broad as long, or the sixth and seventh segments
form part of the club, the inner margin of which is serrate. and the
seventh segment much broader than long; female without a carina be-
tween the antennae; last dorsal segment of the male with 4 tubercles;
squama laminate or slightly thickened, the apex even.

B. Eyes of the male reaching the posterior margin of the head, touching
each other on the vertex, or separated by a distance less than that
between the posterior ocelli; these touching the eyes, 3 times as far

408

University of California Publications | ENTOMOLOGY

from an imaginary line connecting the posterior borders of the eyes
as from each other; seventh segment of antenna of male slightly
widened at apex, much longer than broad, the club solid, ovate, shorter
than segments 6 plus 7, convex above and below, slightly flattened at
base beneath; middle femur and tibia with even surface, rounded or
flattened but not concave beneath and without irregularities —..............
aa Seat deate bu obsug te ceuanace siete ten sasatesbes du. Seneeet con puue foe eee eee Holopticus, n. subg.

BB. Eyes of the male not reaching the posterior margin of the head, distant

from each other by at least twice the distance between the posterior
ocelli; these not touching the eyes, distant from an imaginary line
connecting the posterior borders of the eyes by not more than twice
their distance from each other; antennae of the males of different
forms; either the middle femur or tibia or both of the male irregularly
contorted, grooved or armed beneath, sometimes also that of the
female, but less strongly than in the male.

C. Antennal club of the male solid, ovate, thick, sometimes hollowed be-

-

neath, as long as or slightly longer than segments 6 plus 7, these
not forming part of the club, the seventh segment at least as long
as broad, usually longer; inner surface of squama of male not
fra fo Tah GC eee Se ees Pseudomasaris Ashmead.

CC. Sixth and seventh segments of the antenna of the male forming part

of the club, the remaining segments indistinguishably fused, re-
curved like a half-closed hand, concave beneath, this part of the
club scarcely longer than the sixth segment, posterior margin of
the club strongly serrate, by reason of the irregularly projecting
margin of the sixth and seventh segments, the seventh segment
more than 3 times as broad as long, less than one-third the length
of the sixth; inner surface of the squama of the male fimbriate ......
og ecto cic aut Been ae ere eh mee cae ine Cotyledon, n. subg.

A Ke&y T0 THE SPECIES OF THE GENUS PSEUDOMASARIS
Males

1. Last dorsal segment with 6 tubercles; posterior metatarsus curved and

with a ciliate lobe at apex within; antenna resembling an inverted
PS) OCCU ee ee ne (Toryna) vespoides (Cresson).

Last dorsal segment with 4 tubercles; posterior metatarsus without a

VO Tee as he eG 2 Sa ee CUE ok See

. Sixth and seventh segments of the antenna forming part of the club, the

seventh flattened, over 3 times as broad as long, their irregular edges
making the posterior margin of the club strongly serrate; front with
a prominent tubercle between the emarginations of the eyes ................
(Cotyledon) edwardsii (Cresson).

Sixth and seventh segments of the antenna not forming part of the club,

or if so the seventh is cylindrical and but little widened, the seventh
segment as long as broad, or longer, club short ovate and thick, rarely
hollowed beneath, the posterior margin not serrate; front flat, or with

‘arcwieak: MEd 1a PTO CLC Oe eeowecese ceases w cece src we eee c eet oer eee ee (3)

Vou. 1] Bradley.—The Taxonomy of the Masarid Wasps 409

3.

|

10.

Eyes reaching the posterior margin of the head, closer together than or
about as close together as the distance between the posterior ocelli, these
touchinie ithe) eyes\or Very Weary SOs... c-c.scccencceeceeee see eneee (Holopticus). (4)

Eyes not reaching the posterior margin of the head, at least twice as far
apart as the distance between the hind ocelli; these not touching the
CASTES ee pe (Pseudomasaris).(8)

. Eyes separated by from three-quarters to one and one-half times the

diameter of a posterior ocellus; fourth segment of antenna linear;
process of third ventral segment with a broad longitudinal fossa on
its summit, without two teeth in front, and without a posterior tooth....(6)
Eyes separated by twice or three times the diameter of a posterior
OCC Ii Saeco s Me oe Nes cee Pee Soe ceca Pee ee seco ree nee ee (5)

. Process of third ventral segment with a narrow longitudinal groove on

its summit, two teeth in front, and a large sharp tooth pointing caudad
behind; eyes separated by twice the diameter of a posterior ocellus......
SSS SE ae eR ee bariscipus, n. sp.
Process of third ventral segment with a broad fossa on its summit much
widened anteriorly, the process without teeth in front and obtuse be-
hind; eyes separated by three times the diameter of a posterior ocellus
pt ee) ae Se ie en ee eee ee Nhaceliaemiohwers

. Eyes separated by less than the diameter of a hind ocellus ........................ (7)

Eyes separated by about one and one-half times the diameter of a hind
ocellus:etront temum blacks amdl yell wa 22-sssssseerscersssseeeseeseaes rohweri, n. sp.

. Anterior femur brown with a white area beneath at apex; sides of pro-

podeum weakly angled, not alate nor mucronate......albifrons Rohwer.
Anterior femur usually entirely red or reddish yellow without an apical
white spot; side of propodeum subulate and mucronate -..........-...2-.--..----
Be Ear fees het ta Sate a ces ce sec Susate se aasisuees sanacatoevnavuatcseesbabdesicccoeuseee texanus (Cresson).

. Sixth and seventh antennal segments not flattened beneath, the sixth

eylindrical or slightly widened at apex, the club convex beneath,
middle femur and tibia or the tibia only strongly contorted and con-
CRS LOST Ea A ey a dA fn ee ere pn (9)
Sixth and seventh antennal segments flattened beneath, strongly widened,
the club concave beneath; middle femur concave beneath, with a
tubercle near the apex, the tibia slightly irregularly concave beneath
pene ee een ee ee ee ee. Ee ah ee ead marginalis (Cresson).

. Middle femur and tibia both strongly contorted and concaved beneath;

seventh segment of the antenna three-quarters as wide as long............ (10)
Middle femur with the inferior surface evenly rounded, the tibia on its
outer lower edge strongly produced and angled on the basal third;
seventh segment of the antenna one-half as wide as long .....................-
eee eee eRe a SIN ace orn ne ae eek Re occidentalis (Cresson).
Antenna evenly and not strongly clavate from the sixth segment to the
apex, forming a slender oval club; middle tibiae weakly ridged be-
TICE REN, caegiee Matt Sean | Sed Sanne oie Log eRe ke est aes a Rane eT coquilletti Rohwer.
Antenna with the club broadly ovate, strongly differentiated from the
rest of the flagellum; middle tibia strongly ridged beneath ~...................
SS ee ne zonalis (Cresson.)

410 University of California Publications [ENTOMOLOGY

Ie

Females

A sharp transverse carina between the antennae; clypeus coarsely trans-

VGTS Oly" MPU GOS Co secret eee eae eects seinen ome (Toryna) vespoides (Cresson).

No carina between the antennae; clypeus not rugose, but chagreened or
FOLUNTY Gh h = Pace le Serge gee ne Aare Cree ve mabe or ee PoP es rear eee oe each soa leeten (2)
. Angles of propodeum mucronate or dentate -.......----------<--------------c-eceeoeeseneen (3)

Angles of propodeum rounded, not dentate —.......--------------cs¢o---ce0ceeeneeeeeeeeeen es
Ai Bole Sih Ne het TS ee at (Pseudomasaris) marginalis (Cresson).
Middle tibia, seen from above, inflated beyond the middle; middle femur
with its inner posterior margin carinate and more or less sinuate;
color black and bright yellow............ (Cotyledon) edwardsii (Cresson).
Maddie tibia. seen trom above aor uth eit ely essence eee nee eeeeneree serene (4)
Middle femur scarcely flattened and not ridged beneath; color of the
body partly tawny, ferruginous or rufous, or at least the legs beyond
the coxae all tawny except sometimes for a yellow spot on the femora..(5)
Middle femur flattened beneath, its anterior lower border marked by a
ridge; color black and saffron or lemon yellow, without rufous or
UR 1h 1 02 ed ea OR 8 RnR eS an eee ee ica (10)

. Clypeus very coarsely and densely punctate, almost rugose; tawny, in-

verted V-shaped spot on front, mesonotum except next to the tegulae,
mesoventer, sometimes part of pleura and lower part of propodeum,
mesal spot on dorsal segments 1, 2, and 3, and a mesally dilated basal
border on dorsal segments 4 and 5, and a narrow basal border on
ventral segments 4, .5,,and 6, black. == el dec tate eer a a
PANE SECO eae te erage ee es (Pseudomasaris) occidentalis (Cresson).
Clypeus weakly and shallowly punctate, sometimes punctulate with scat-
tered coarser punctures; black, with yellow and usually with rufous
EN gc ee ee (Holopticus) (6)

. Middle of mesonotum just in front of scutellum very densely and finely

granular-punetate; second dorsal segment closely, rather finely and
@veri ly, mie bettie seek Ge ce crc meme (7)
Middle of mesonotum in front of scutellum coarsely, confluently, almost
rugosely punctate, this area of the mesonotum strongly depressed;
second ventral segment with separated coarser punctures, the abdomen
and pronotum more or less polished and shining; color black, saffron
yellow and red, with a strong preponderance of yellow on the abdomen
2 SAE cee ce er ees Parte i eee eee eee eee phaceliae Rohwer.

. Mesonotum just in front of scutellum with a strongly depressed area, not

reaching the lateral groove of either side, this area finely granular,
while the raised area at the sides and anteriorly is more coarsely and
sparsely but still densely punctate; front always with a triangular
white area, legs beyond coxae entirely reddish’ -......_.-----.------------------- (8)
Mesonotum in front of scutellum flat, but without a median depressed
area, the sides and anteriorly finely and densely punctate, but not
granular as in the middle just before the scutellum; front usually
mi ATLHL CON Ue Hee? Wiad A SPS) 0101 eee ae ae eer eee ee Net ee bes ee er eee ee ecco (9)

. Thorax and abdomen with a large amount of red. ............2.---2.--c--ceiecsreeeseeeoe==

Se SE RR he PR Se mrs eo eee ee eee texanus texanus (Cresson).

Vow. 1] Bradley—The Taxonomy of the Masarid Wasps 411

horas and-abdomen without red marking 22022202)
Be eee texanus neomexicanus Rohwer.

9. Front without a median white triangle; knees white; three basal ab-
dominal segments with their ground color dorsally mostly red —.........

SEE SE Ee SPE RE PE ee SEE TE Rp ee Ee oe basirufus Rohwer.
Front with a median white triangle; legs beyond coxae entirely rufous;
two basal segments of abdomen with ground color partly red ..............

SESE EEE OOL oe ER Se en ee ee maculifrons (Fox).

10. Seen from behind, the middle tibia strongly contracted near its base;
cephalic margin of cell R,,; less than one-half the distance between

M,,, and M, on its caudal border; humeral ridge well marked; abdomen
elongate; propodeum except the angles, femora except their apices,
Postseubellumeanidemao St otaciliyayeusy bil aici ka gerne tces ones oeruce screen eet oaners

Se ee ee ee See oR CEO rs eee Ae (Pseudomasaris) zonalis (Cresson).
Middle tibia gradually widened from base to apex; cephalic margin of
cell R,,; more than one-half the distance between M,,, and M. on its
caudal margin; humeral ridge almost obsolete; abdomen short, ovate;
propodeum, except two spots behind, femora, except basal half to
three-quarters of posterior surfaces, postscutellum at apex, and most

of clypeus, saffron yellow .............- (Pseudomasaris) coquilletti Rohwer.

Toryna, new subgenus

¢. Eyes deeply emarginate, the emargination obtuse, the eyes more
than twice as far apart above as the distance between the hind ocelli;
labial palpus 2-segmented, the first more than twice as long as the
second; maxillary palpus very short and conical. Antenna a little
longer than the head and thorax united, the scape short, globose, the
pedicel less than one-half its length, segm. 3 to 7 distinct, 3 to 6 linear,
the seventh widened, about one and one-half times as long as thick at
apex, segm. 8 to 12 fused to form an oval club, shaped like the inverted
bowl of a spoon, convex above and coneave beneath, the divisions be-
tween these segments only faintly apparent.

Anterior femur normal, its undersurface convex, the tibia some-
what contorted, obliquely constricted at base, as long as the tarsus, the
latter with a lateral fringe of rather long, silky hairs; metatarsus as
long as the following segments united, the fourth segment broader than
long; middle femur with irregular ridges and fossae beneath, the tibia
from a lateral view constricted at base and strongly expanded toward
the apex, the expansion beneath with a fossa which fits over the femur ;
tibia one-fifth longer than the tarsus, fourth segment of the latter much
broader than long; posterior tibia as long as the metatarsus; meta-
tarsus one and three-fifth times as long as the remaining segments
united, plainly curved, its apex produced on the inner side into a
rounded lobe extending well beyond the base of the following segment,
and bearing a prominent crest of stout setae; hind tarsal segments 2
to 4 with a prominent inner fringe of setae; the fourth segment about
as long as wide; second and third ventral segments both with tubercles ;

412 University of California Publications [ENTOMOLOGY

last dorsal segment, in addition to the four tubercles bordering its
truncate apex, has a pair of tubercles on the dorsal surface.

2. A sharp transverse carina between the antennae; clypeus very
convex and rugose; labial palpi 2-segmented, the first more than twice
as long as the second, maxillary palpus very short and conical.

Anterior femur flattened beneath, the tibia with a weakly irregular
undersurface as long as the tarsus; the latter without a fringe of
ciliae ; the metatarsus one-half as long again as the remaining segments
united; the fourth segment broader than long; middle femur flattened
beneath; the tibia slightly flattened beneath, not irregular, but con-
stricted at base, the outer surface bearing scattered short spines, very
slightly shorter than the tarsus, bearing 1 apical spine; metatarsus as
long as the following segments together, the fourth about as long as
broad ; posterior tibia about one-third shorter than its tarsus, the apical
spur bifid ; metatarsus one-half as long again as the remaining segments
united, not noticeably curved and without an apical lobe, the fourth
segment about as long as broad.

Abdomen unarmed.

Type—Masaris vespoides Cresson.

Pseudomasaris (Toryna) vespoides (Cresson)
Figures 21, 65, 66, 67, 86, 96

1863. Masaris vespoides Cresson, 3, 9. Proceedings of the Entomological
Society of Philadelphia, vol. 2, p. 287, pl. IV.

1904. Pseudomasaris vespoides von Dalle Torre. Genera Insectorum, fase.
19, p. 8

1913. Masaris vespoides Davidson. Bulletin Southern California Academy
of Sciences, vol. 12, p. 17 (life history).

1913. Pseudomasaris vespoides robertsoni Cockerell. Proceedings of the
Entomological Society of Washington, vol. 15, p. 107.

¢. Front rugosely punctured, raised below the front ocellus, with
a transverse ridge between the antennae, which is polished and impune-
tate; clypeus very convex, its dise polished and with few large punc-
tures, its sides with close, smaller punctures and somewhat wrinkled.

Humeri prominent, but not angled, without a distinct humeral
ridge; mesonotum anteriorly closely punctate, and with a median
ridge posteriorly with two lateral ridges between which it is depressed,
smooth and polished, with only minute shallow and sparse punctua-
tion ; scutellum prominent, polished and practically impunctate.

Basal abdominal segments strongly constricted at base, closely and
finely punctate, a median area at base of each except the first two,
impunctate, polished ; medial punctures finer toward the apex of each
segment; second ventral segment with two anterior tubercles and two
weak ones behind; third ventral segment with a strong tricuspid
prominence.

Vou. 1] Bradley —The Taxonomy of the Masarid Wasps 413

Color black and pinard yellow, the latter distributed as follows:
elypeus, labrum, mandibles except base and apex, inner orbits above
emargination, line behind eyes, broad humeral stripe, tegula, small
spot in front on tip of pronotum and within on mesonotum in front of
scutellum (sometimes wanting), large spot below tegula, two short
stripes on mesonotum behind (usually absent), large or small apical
spot on scutellum, usually the angles of the propodeum; legs beyond
middle of femora, sometimes also base of middle femora behind and
spot on front and middle trochanter and coxae behind ; subapical band
on dorsal segments 1 to 5, varying in width and nature of its lateral
incisions, the fifth usually enclosing a black spot on each side, most of
sixth and apical half of seventh segments, spot at side of second and
third ventral segment, sometimes prominence of third posteriorly, and
nearly all of the fourth to sixth ventral segments; antennae yellow
grading into deep chrome above, and marked with reddish brown
beneath.

Wings stained yellow, the veins yellow (Mars yellow). Length,
17-22 mm.

The punctuation is variable, the posterior part of the mesonotum
and the secutellum being sometimes punctate throughout. The color
is also somewhat variable.

. Clypeus coarsely rugose at base; similar to the male, but yellow
less extensive, the elypeus and labrum except two small spots, on each,
most of mandibles and scape, hind angles of prothorax, mesonotum
except spot next to tegula, secutellum except tip and angles of propo-
deum except the tooth, more of femora, broader basal parts of ab-
dominal segments, nearly all of last ventral segment black, the last
dorsal segment black with a large yellow spot on each side; antennal
club fuscous above, Sanford brown beneath. Length, 15-19 mm.

This is our largest and most handsome species. Its biology has
been described by Davidson (loc. cit.). It builds clay nests.

The subspecies deseribed by Professor Cockerell from California
does not appear, on comparison with other specimens from California
and elsewhere, to represent more than an individual variation.

Types.—Lectotype, ¢: American Entomological Society, no. 2095.
Allotype: American Entomological Society.

SoutH Daxota: Lead City, 12 [American Entomological Society ].

IpaHo: Lewiston, 1 ¢, 2 2; Craig’s Mt., 1 3 1 9 [American Ento-
mological Society |.

Couorabo: Pikes Peak, 1 4, 2 9 (W. J. Howard), [types, American
Entomological Society]; Garden of the Gods, July 13, 1877 [U. 8S.
National Museum] ; Florissant, July 21, 1 2 on flowers of Pentstemon
(T. D. A. Cockerell), [American Museum of Natural History].

New Mexico: Jemez Springs, May 20, 1913,.1 9; June 2, 1913,
6400 feet, 1 J (J. Woodgate), [Cornell University].

UraH: Pronotetali, August 21, 1906, 1 9 [Cornell University] ;
Silver Lake, July 14,1 ¢,1 9 (H. Skinner), [American Entomological
Society ].

414 University of California Publications [ENTOMOLOGY

Nevapa: [American Entomological Society]; 1 ¢ [American Mu-
seum of Natural History].

CALIFORNIA: Claremont, 2 4, 2 2 (C. F. Baker), [Pomona College
and Cornell University] ; Redlands, 1 9 (G. Robertson), [type of sub-
species robertsoni Cockerell, U. S. National Museum, Cat. no. 15529] ;
Pasadena, June 12, 1895, 1 9 (R. W. Doane), [Cornell University] ;
Los Angeles [U. S. National Museum]; summit of Sierra Nevada, 1 9
[American Museum of Natural History]; Strawberry Valley, El
Dorado Co., 7 2, August 9, 18, 1912 (KE. C. Van Dyke), (Calif. Acad.
Sci. and Cornell Univ.]; Fallen Leaf Lake near Lake Tahoe, 1 J,
July 19, 1915 (L. S. Rosenbaum), [Calif. Acad. Sci.]; Carrville,
Trinity Co., 1 3, 1 9, June 29, 1918 (HE. C. Van Dyke), [Calif. Acad.
Sei. and Cornell Univ.].

Holopticus, n. subgenus

&. Eyes deeply emarginate, emargination very narrow or acute,
eyes reaching the posterior margin of the head and almost touching
above, or separated by less than the distance between the hind ocelli;
labial palpi with three distinct segments, or with two, the apical portion
of the second contracted but not discrete. Scape short, barrel shaped,
segments 3 to 6 cylindrical or with apices nodose, seventh widened at
apex, twice as long as wide, 8 to 12 indistinguishably fused in a solid
ovate club, convex above and below, not as long as segments 6 to 7.

Angles of the propodeum dentate——Anterior tarsus with ciliate
hind margin; all femora with surface regular and convex throughout ;
anterior tibia regular; middle tibia somewhat depressed and flattened
beneath but not contorted nor with irregularities, two-thirds as long
as the tarsus, the metatarsus two-thirds to four-fifths as long as the
remaining segments united, the fourth as long as wide ; hind tibia three-
quarters as long as the tarsus; the metatarsus straight, without an
apical lobe, as long as the following segments united; the fourth seg-
ment longer than wide.

Second ventral segment with two tubercles unarmed, the third with
a large process, of variable shape, but bearing a longitudinal groove
on its summit; last dorsal segment with four tubercles, the inferior
pair small and close together.

2. Front without a carina between the antennae, clypeus not rugose.

Legs as in the male, except the middle metatarsus is about equal
to the remaining segments united.

Type.—Masaris texanus (Cresson).

Pseudomasaris (Holopticus) texanus (Cresson)
Figures 56, 57, 58

1871. Masaris texanus Cresson, 6,9. Transactions of the American Ento-
mological Society, vol. 3, p. 348.
1904. Pseudomasaris texanus Dalle Torre. enera Insectorum, fase. 19,

p. 8.

VoL. 1] Bradley.—The Taxonomy of the Masarid Wasps 415

¢. Spots on clypeus, spot on each side of pronotum, legs, except
coxae, small median spot on mesonotum, spot on pleura, most of border
of first dorsal segment, spot on each side of second and third dorsal
segment, and the second and third ventral segments chestnut ; segments
4 to 7 beneath except at apices, club of antenna beneath at base, spot
on clypeus, upper part of front and inner orbits, upper margin of
pronotum, spot in middle of mesonotum, 2 small anterior spots and
2 posterior touching the tegulae, apex of seutellum, angles of propo-
deum, 2 spots in red border of first dorsal segment, sides and spot in
middle of second and third dorsal segments and borders of fourth to
sixth, and band on the fourth and fifth ventral segments yellow.

Punctuation of front and clypeus fine and close; of mesonotum
coarser, but close; the posterior part of dise of mesonotum depressed
and closely punctate; scutellum closely punctate, propodeal angles
mucronate. Dorsal segments moderately depressed and but seantily
punctate at base, the apices finely punctate; second ventral segment
without tubercles.

Squama moderately thick, obtuse, densely pubescent on the inner
side, and with a small pubescent patch on the outer side at tip; sagitta
trigonal with obtuse tip and sharply carinated angles, about half as
long as the uncus; this long, slender, strongly but gradually decurved
toward the tip, with a carina but not a barb beneath near the base.

9°. Antenna, apex of clypeus, pronotum, legs except coxae, spot on
seutellum, stripe on first dorsal segment (with included yellow spot),
large spot on each side of second segment, chestnut; spot on elypeus,
on face, orbits, narrow upper border of pronotum, tegulae, spot on
pleura, on apex of scutellum, angles of propodeum, sides and middle
of apex of first dorsal segment, apical margins of remaiming dorsal
segments and of the ventral segments yellow, the yellow on the abdo-
men obscure; front. except on the white spot and vertex with regular,
round, dense but separated punctures, irregular and smaller behind the
ocelli; clypeus with minute punctuation in addition to coarser pune-
tures; dorsum punctured like the front, but in the depressed area
before the seutellum densely granular punctate; abdomen opaque,
densely finely and evenly punctate.

Types.—Holotype: ¢. American Entomological Society, no. 2100.
Allotype: American Entomological Society.

Texas: 3 ¢, 2 9 (Belfrage), [American Entomolgical Society] ;
Austin, 1 9, May 5, 1901 (C. T. Bruers), [| Jos. Bequaert].

ARIZONA: Phoenix, 1 ¢ [Dr. Jos. Bequaert].

SuBsPEciES Neomexicanus Rohwer

1912. Pseudomasaris zonalis neomexicanus Rohwer, 2. Proceedings United
States National Museum, vol. 41, p. 452.

Q. Black with the following parts straw yellow: wedge-shaped spot
on clypeus, triangular spot on middle of frons, spot filling incision of
eyes, posterior orbits narrowly to a point opposite incision of eyes,

416 University of California Publications [ENTOMOLOGY

large oval spot on sides and narrow posterior border of pronotum,
tegulae, lateral posterior spot on mesoscutum above tegulae, superior
spot on pleura, apical spot on seutellum, spot on angles of propodeum,
broad apical bands on tergites one to five inclusive, incised laterally
on tergites three, four, and five, interrupted preapical band on sixth
tergite, interrupted apical band on second, complete on third and
fourth and four apical spots on fifth sternites; the following parts
burnt sienna: mandibles, antennal club beneath and legs below coxae;
following parts piceous: flagellum except where mentioned and apical
margin of sixth sternite; wings slightly smoky, the costa castaneous,
the other veins dark brown; hair of head and thorax blackish.

Front dull, with large separate punctures which are much closer
medially ; clypeus finely punctulate with a few large punctures dor-
sally. Humeri rounded; pronotum with separated punctures ; mesono-
tum with large, sometimes confluent punctures which are closer and
finer in the flattened posterior part; seutellum with large, distinct
punctures laterally and with a rather narrow granular area medianly ;
sides of the pronotum mucronate, not carinate. Abdominal segments
evenly, finely and closely punctate; bases of second and third tergites
somewhat contracted. Length, 10 mm.

Type.—vU. S. National Museum, no. 14145.

New Mexico: Aztec, May 4, 1899, 1 9 at flowers of Astragalus
[type, U. S. National Museum].

Mr. S. A. Rohwer has very kindly sent me the redescription of
the type here published.

Pseudomasaris (Holopticus) albifrons Rohwer

1912. Pseudomasaris albifrons Rohwer, g. Proceedings United States
National Museum, vol. 41, p. 451.

‘“%. Length about 12 mm. Very like texanus (Cresson), from
which it may be separated by the following characters: very few
large punctures on the front and these not sharply defined; posterior
part of mesosecutum uniformly punctured (in teranus the depressed
area is more closely punctured) ; punctuation of abdomen finer ; second
dorsal segment depressed by fully half of its entire length (in texanus
it is hardly depressed) ; third segment hardly depressed (in texanus
it is depressed by fully one-third) ; punctures of the apical dorsal seg-
ment more widely separated ; second cubital cell on the radius longer,
being in the type greater than the distance between the recurrent
veins (in texanus it is much less) ; markings whitish; clypeus except
apical part pale; wings slightly yellowish in stigmal area, otherwise
hyaline.’’

Holotype.—uU. 8S. National Museum, no. 14144.

New Mexico: Las Cruces, 25 March, 1896, on plum (T. D. A. |
Cockerell), [type, U. S. National Museum].

UraH: 1 ¢ (Palm), [U. 8. National Museum].

Vor. 1] Bradley—The Taxonomy of the Masarid Wasps 417

Pseudomasaris (Holopticus) rohweri, n. sp.

&. Black, the following parts chestnut to Sanford brown; most of
the fifth, sixth, and seventh antennal segments above, scape toward
apex, front tibia except spot on outer side at base, front tarsus, hind
and middle legs except coxae and trochanters, wing veins, second dorsal
segment except for 3 yellow spots, third dorsal segment except for 3
yellow spots and base, 2 spots and median apical border on fourth
dorsal segment, second ventral segment, third ventral segment except
near base; following parts pale chalcedony yellow; large spot on
clypeus above, emargination of eyes, upper part of front and vertex,
narrow line behind the eyes, fifth and sixth antennal segments beneath
except at extreme apex, antennal club beneath at base, pronotum above
except posteriorly, tegulae, large spot on pleura beneath, triangular
spot on each side of mesonotum in front, small median spot, spot on
each side next to tegulae, apex of scutellum, angles of propodeum, spot
at apex of anterior femur and base of tibia, broad subapical band on
first dorsal segment, 3 transverse spots on second and third, sinuate
subapical bands on fourth, fifth, and sixth dorsal segments and sub-
apical bands on fourth and fifth ventral segments. Wing stained with
yellow along costal margin; head and thorax rather noticeably pubes-
cent.

Front prominent, clypeus gibbous, emargination of eyes very nar-
row, almost acute, front and eclypeus shallowly and rather obscurely
punctured, labial palpi with 3 segments, the last rather closely fused
to the preceding. Scape globular, third segment shorter than the
fourth, this not enlarged at apex, short pubescent throughout, fifth
and sixth but slightly enlarged at apex, third to seventh segment some-
what compressed, the seventh much widened at apex, about twice as
long as wide; club short, ovate, somewhat flattened below at base, about
as long as segments 6 and 7.

Mesonotum rather coarsely and closely punctate, the depressed area
more closely ; the seutellum more finely and closely punctate, also with
large scattered punctures, pleura with coarse punctures.

Anterior tarsus ciliate, segments of legs with regular surfaces;
middle tibia flattened beneath and from an external view considerably
widened medially ; middle tibia about two-thirds as long as the tarsus ;
metatarsus slightly longer than the remaining segments united; the
fourth segment slightly longer than broad; hind tibia five-eighths as
long as the tarsus; metatarsus as long as the following segments united ;
the fourth segment slightly longer than broad. Distance from r to R,
as great as that from r-m to M,.

Segments 2 to 5 of abdomen considerably contracted at base, dis-
tinetly punctate, punctures growing finer toward the apex; superior
processes of last segment acute, prominent, curved; inferior margin
of the segment with 3 minute tubercles, second ventral segment with a
transverse prominence and a median groove, prominence of third seg-
ment bearing a longitudinal fossa, much broadened in front, on its
summit, sloping away posteriorly, its anterior face notched at apex.

418 University of California Publications | ENTOMOLOGY

Genitaha differing from those of bariscipus as follows: squama
shghtly falcate, blunt; sagittae much longer, more slender, and with
obtusely rounded apex; sagitta more slender.

Type.—American Entomological Society.

ARIZONA: Quartzite, April 14, 1903, 8 3 (G. S. Hutson), [Amer-
ican Entomological Society]; Phoenix, 1 ¢ [American Entomological
Society ].

Pseudomasaris (Holopticus) bariscipus, n. sp.
Figures 77 and 85

3. Black, following parts chestnut to Sanford brown; segments 3
to 7 of antenna at base above, club of antenna except at base beneath,
apices of mandibles, anterior tibia except base on outer side, anterior
tarsus, middle femur infuseated apically, middle tibia except spot at
base on outer side, middle tarsus, hind leg except coxa, narrow median
apical border of first dorsal segment, second dorsal segment except for
three yellow spots, third dorsal segment except for three yellow spots
and three black areas, median apical band on fourth segment, second
and third ventral segments, and area on apex of seventh dorsal seg-
ment; the following pale claleedony yellow: spots on mandibles, large
spot on clypeus, inner orbits fused above, apex of scape, of third an-
tennal segment above, bases of fifth, sixth, and seventh segments below,
base of antennal club beneath, narrow line behind the eyes, humeri
broadly, tegulae, large spot on pleura, one median and two anterior
spots on mesonotum, preapical spot on scutellum, angles of propodeum,
spot on front, and small spot on middle knees, broad band on first
segment, 3 transverse spots on second and third, and a sinuate line
near apex of fourth, fifth, and sixth dorsal segments, band on fourth
ventral and three spots on fifth ventral segments ; wings stained slightly
yellowish, veins, ferruginous to brown.

Emargination of the eyes narrow, obtuse; front convex but not
prominent; clypeus gibbous, front and clypeus closely but shallowly
punctate with few large punctures; labial palpi with 2 segments, the
second with its apical portion contracted but not discrete. .Seape short,
subglobular; third segment linear, shorter than fourth; this much
widened and pubescent at apex; fifth and sixth segments also clavate
but less strongly than the fourth, seventh segment companuliform,
nearly twice as long as wide, club ovate, nearly as long as segments 6
and 7.

Mesonotum moderately coarsely and closely punctate, depressed
area more closely punctate, disk of scutellum closely and more finely
punctate, angles of propodeum acute but not mucronate.

Distance between r and R, in excess of that between r-m and M,.
Anterior tarsus ciliate; femora and tibiae all regular, without uneven
surfaces; middle tibia flattened beneath, seen from above gradually
widened in middle ; middle tibia about two-thirds as long as the tarsus ;
metatarsus nearly as long as the following segments united; fourth

|

Vot. 1] Bradley—The Taxonomy of the Masarid Wasps 419

segment about as long as broad; hind tibia five-eighths as long as the
tarsus; metatarsus as long as the following segments united; fourth
segment slightly longer than broad.

Second ventral segment with two blunt tubercles; process of third
segment compressed, with a narrow groove on its summit and two
anterior teeth, posteriorly it ends in a tooth directed caudad; last

* dorsal segment with 2 anterior processes sharp, the 2 apical ones very

small and close together.

Squama obtuse, rather densely ciliate on the inner side and also
exterlorly at apex; sagitta trigonal, its angles carinate, its apex acute,
about one-half as long as the uncus; uncus slender, shorter than in
texanus and less decurved.

Holotype, American Entomological Society.

ARIZONA: Quartzite, April 14, 1903, 8 § (George S. Hutson),
{American Entomological Society].

Pseudomasaris (Holopticus) phaceliae Rohwer

1912. Pseudomasaris phaceliae Rohwer, 4, 9. Proceedings United States
National Museum, vol. 41, p. 450.

“¢Y. Length about 13 mm. Labrum obtusely pointed; elypeus
strongly uniformly convex, arecuately emarginate in apical middle,
finely punctured; front with rather large, separate punctures; scape
not one and a half times as long as broad, third joint distinctly
shorter than fourth; anterior ocellus large, subreniform; pronotum
polished, with well-separated small punctures; mesonotum with dis-
tinet, well-separated punctures, which are somewhat closer in the
depressed area; mesopleura and scutellum similarly punctured ; propo-
deum normal; abdomen polished, with widely separated punctures
which become smaller posteriorly ; seen from above the first dorsal is
arcuately emarginate anteriorly ; second, third, and fourth dorsal seg-
ment depressed basally by about one-third the length of the entire
segment ; processes of the third ventral and,apical segments essentially
as In texanus; second cubital on the radius slightly longer than the
distance between the recurrent veins. Black except where mentioned ;
apical half of scape, flagellum (except apical spots on fourth and fifth
joints and greater part of club), face above level of antennae (the
lower margin has three indentations of black), most of elypeus, pro-
notum, large circular spot below tegulae, tegulae, spot above, two fan-
shaped spots on anterior part of mesoseutum and a small spot in front
depression, spot on scutellum and angles of propodeum yellow; abdo-
men with broad dorsal and ventral bands on apex of all segments
reddish yellow (due in part to potasium cyanide?) ; legs reddish-
yellow, knees yellowish; wings vitreous, hyaline, slightly yellowish in
stigmal region; venation pale brown.”’

9. Black except as follows: antennae rufo-piceous; posterior orbits
dorsally, narrow line on inner orbits up to and filling the eye emargi-
nation, large spot on clypeus, spot above, posterior margin of pro-

420 University of California Publications [ENTOMOLOGY

notum narrowly, and an elongate lateral spot, large spot below tegulae,
tegulae, spot above, large spot in front of depression on mesonotum,
most of scutellum, angles of propodeum, dorsal and ventral (except
first) abdominal segments apically yellow; pronotum (except where
mentioned), band on scutellum, elongate spots on second, third, and
fourth segments rufous; legs rufous; wings dusky, especially near the
veins; stigma and costa reddish brown, veins dark brown.

Clypeus convex, broadly emarginate, very finely granular, with
large punctures intermingled; front with large distinct punctures,
which are more widely separated on the eye margins and vertex; hind
ocelli equidistant from the eyes and from each other ; scape short, third
segment of antenna as long as the three following.

Pronotum with well separated large punctures; mesonotum with
punctures the size of those of pronotum, but closer and especially so
in the depressed area; mesopleura more closely punctured than the
mesonotum ; scutellum punctured like mesopleura, with an indistinct
carina medially.

Second cubital cell on the radius as long as two-thirds of the dis-
tance between the recurrent veins.

Abdomen with well separated, distinct punctures, which become
smaller posteriorly ; first dorsal segment shightly emarginate anteriorly
when seen from above; second and third dorsal segments depressed
basally for about one-third their length; apical ventral segments with
the large punctures well separated. Length, 12 mm.

Types—Holotype, ¢#: U. 8. National Museum, no. 14148. Allo-
type: U.S. National Museum.

New Mexico: Albuquerque, May 138, 1910, ¢ on Phacelia neomeaxi-
cana (J. R. Watson), [type, U. 8S. National Museum]; Mesilla, May
29, on flowers of Phacelia (T. D. A. Cockerell), [U. S. National
Museum]; Fillmore Canon, 9 (T. D. A. Cockerell), [U. S. National
Museum |.

Pseudomasaris (Holopticus) maculifrons Fox

2 black, the following parts chestnut to Sanford brown: mandibles
at tip, antenna beyond the fourth segment, legs, apex of first dorsal
segment except for 3 yellow spots, median apical band on second dorsal
segment enclosing a yellow bar, narrow median band on third and
fourth segments, and the wing veins; the following pale chalcedony
yellow: large spot on clypeus above, triangular spot on front, emargi-
nations of eyes, line behind eyes, humeral angles, narrow hne along
posterior border of pronotum, outer half of tegulae, large spot on
pleura, small spots on mesonotum touching tegulae, preapical spot on
scutellum, two narrow lines on postseutellum (absent on type), pro-
podeal angles, 3 transverse bars on first and second dorsal segments,
sinute subapical band on third, fourth, and fifth dorsal segments,
widened laterally, lateral spots on sixth dorsal segment, transverse
interrupted and obscure band on second and third ventral segments

VoL. 1] Bradley—The Taxonomy of the Masarid Wasps 421

(only 2 lateral spots on second segment of type), transverse band
occupying most of fourth ventral segment (in type 2 lateral and 2
median spots).

Front closely punctured, clypeus closely and minutely punctulate,
with scattered larger punctures, posterior ocelli as far from each other
as from the compound eyes.

Humeri rounded ; pronotum with well separated coarse punctures ;
mesonotum with close coarse punctures, posteriorly finer, closer and
less regular causing surface to be chagreened ; seutellum similarly pune-
tured but with coarse punctures more seattered ; pleura a little more
densely punctate than. pronotum.

Abdominal segments densely punctate at base, punctures finer to-
wards apex of abdomen; second and third segments slightly contracted
at their bases. Length, 10 mm.

Description drawn from a specimen from Arizona.

Holotype.—California Academy of Sciences.

Lower CauirorNiA: El Paraiso [Cal. Acad. Sci.].

ARIZONA: Quartzite, April 14, 1903, 1 9 (G.S. Hutson), [American
Entomological Society |.

Pseudomasaris (Holopticus) basirufus Rohwer

1912. Pseudomasaris zonalis basirufus Rohwer, 9. Proceedings United
States National Museum, vol. 41, p. 452.

2. Black, the following parts straw yellow: spot on mandibles,
trilobed spot on eclypeus, narrow line around the eyes, interrupted
below the incision, mesad of the summits and posteriorly below, tri-
angular spot on sides and narrow posterior border of pronotum, supe-
rior spot on pleura, tegulae, small spot near posterior border of meso-
notum, apical spot on scutellum, spot on angles of propodeum, apex
of front femora and extreme base of their tibiae externally, small spot
on apex of middle femora, spot at base of middle tibiae, lengthened
laterally, small anterior stripe at base of posterior tibiae, apical stripe
on each side of first and second tergite, preapical mesal stripe on same,
narrow apical border of third, fourth, and fifth tergites, incised later-
ally, broadly interrupted stripe of sixth tergite, preapical spots on
second and fifth, and apical bands on third and fourth sternites; the
following parts burnt sienna: flagellum except base, all of legs except
yellow spots and bases of coxae, apical third to two-thirds of first three
tergites, excluding yellow portions, first and second sternites and trace
on third; wings slightly smoky, slightly violaceous, the veins ecasta-
neous to black.

Front closely punctured, punctures more sparse toward the eyes;
clypeus finely closely punctulate, the yellow spot indistinetly so.
Humeri rounded; pronotum with separated punctures; mesonotum
closely, granularly punctate, more finely on the flattened posterior
part; seutellum granularly punctate, more densely in the middle than
on the sides; sides of propodeum mucronate, not carinate. Abdominal

422 University of California Publications [ENTOMOLOGY

segments evently and finely, rather densely, punctate; bases of second
and third segments somewhat contracted. Length, 9 mm.

Type.—vU. 8S. National Museum, no. 14146.

CALIFORNIA: Death Valley, April, 1891, 1 9 (A. Koebele), [type,
U.S. National Museum].

ARIZONA: Quartzite, April 14, 1903, 3 9 (G.S. Hutson), [American
Entomological Society |.

Subgenus Pseudomasaris Ashmead

1902. Pseudomasaris Ashmead. Canadian Entomologist, vol. 34, p. 221.

¢. Eyes never as close together as the distance between the hind
ocelli, these not touching the eyes; the temple narrow but distinct ;
incision of the eye sometimes subacute but usually broadly rounded at
apex; front with a low tubercle between and slightly above the bases
of the antennae; clypeus slightly or strongly convex, broadly emargi-
nate anteriorly. Antennae as long or longer than the head and thorax
united ; seape subglobular ; pedicel about one-third its length ; segments
3 to 6 cylindrical or compressed and somewhat widened at their apices,
the third always linear, the seventh slightly or strongly widened, some-
times more than twice as long as wide; segments 4 to 7 in one species
flat beneath ; elub short, not or but slightly exceeding the length of the
sixth and seventh segments, ovate, convex above and below or, as in
one species, hollowed beneath.

Humeri entirely rounded or more or less prominent; parapsidal
furrows absent; angles of propodeum dentate or only sharply carinate.

Anterior leg not contorted; surfaces of the segments uniform;
femur and tibia more or less flattened beneath; tarsus in one species
ciliate; middle femur with its undersurface convex and regular or
contorted and concave; tibia with its undersurface convex and regular
or dilated, contorted and coneave, but both segments never with uni-
form undersurfaces in the same species; middle tibia from two-thirds
to four-fifths as long as the tarsus; metatarsus two-thirds to three-
quarters as long as the remaining segments together; fourth segment
as broad as long or almost so; hind tibia from .6 to .8 as long as the
tarsus; metatarsus without an apical lobe, approximately equal to the
remaining segments united, fourth segment longer than broad.

Basal abdominal segments considerably ‘contracted above at their
bases ; abdomen much decurved at apex, the last segment with its apical
surface truncate, the truneature margined with 2 superior sharp or
obtuse prominences and 2 inferior smaller tubercles placed closer to-
gether; second ventral segment with 2 low tubercles; process of the
third high, posteriorly with an acute tooth directed caudad.

Squama obtuse without setae beneath or with a small patch of
bristles; sagitta trigonal, the angles carinate ; uncus usually short and
blunt but in one species elongate, slender, curved, and acute.

Vo. 1] Bradley.—The Taxonomy of the Masarid Wasps 423

. The face without a ridge between the antennae. Humeral angles
sometimes subcarinate ; angles of propodeum obtuse or dentate. Middle
femur usually flattened beneath, sometimes ridged; the basal third of
the tibia, seen from in front sometimes distinctly contracted beneath.

Type—Pseudomasaris occidentalis Cresson (by original designa-
tion).

Pseudomasaris (Pseudomasaris) occidentalis Cresson
Figures 19, 62-64, 81

1871. Masaris occidentalis Cresson, 9. Transactions American Entomo-
logical Society, vol. 3, p. 348.

1872. Masaris occidentalis Cresson, ¢. Loc. cit., vol. 4, p. 231.

1902. Pseudomasaris occidentalis Ashmead. Canadian Entomologist, vol.
34, p. 221.

é. Color black, the following parts yellow ochre to ochraceous
orange: face except at base of antennae, line between these, line in-
eluding the ocelli, clypeus, labrum, mandibles, line behind the eyes,
scape except at base, pedicel, third antennal segment except at sides,
antennal club except apical three-quarters beneath and posteriorly,
pronotum except the collar and spot in front of tegulae, tegulae, large
spot below, 2 small spots on mesonotum anteriorly, most of seutellum,
angles of propodeum, legs except coxae, apex of first dorsal segment,
second abdominal segment except black area in middle above, third
segment except dark, basal, dorsal area, fourth and fifth segments ex-
cept bases, sixth dorsal except its base, sixth ventral entirely, and a
band before the prominences of the seventh dorsal segment.

Emargination of the eyes, narrow, triangular, subacute at apex;
hind ocelli removed from the compound eyes by considerably less than
their diameter’s length; front with an elevation below the anterior
ocellus, below this a small longitudinal tubercle, only slightly convex
with rather strong and separated punctures; elypeus very strongly
convex, the punctures irregularly confluent, weaker toward the apex;
labial palpi with two segments, the second segment little over one-half
as long as the first, its apical part contracted and almost discrete,
forming an incomplete third segment; the segment of the maxillary
palpus a little longer than usual, not a mere tubercle. Segments 3
to 6 of antenna not at all enlarged at their apices; the seventh shghtly
enlarged at apex but more than twice as long as broad; the club oval,
convex above and below, a very little longer than segments 6 + 7.

Humeral angles slightly prominent but not at all angled; pro-
notum rather closely punctate, medially impunctate; mesonotum with
coarse and well separated punctures in front, becoming finer and closer
as they approach the middle; posterior portion of the mesonotum flat-
tened but not depressed, polished, with minute and scattered pune-
tures: scutellum rather strongly convex, the slight anterior median
ridge minutely obscurely punctuate, somewhat shining; angles of the
propodeum sharply dentate; pleura coarsely punctate.

424 University of California Publications | ENTOMOLOGY

Front femur and tibia slightly flattened beneath, their surfaces not
irregular, the tarsus ciliate; middle femur with regular and convex
inferior surface; the tibia seen from in front with its inferior edge at
the basal third strongly dilated and angulate; tibia three-quarters as
long as the tarsus; metatarsus seven-tenths as long as the remaining
segments together; fourth segment about as long as broad; hind tibia
four-fifths as long as the tarsus; the metatarsus as long as the remain-
ing segments united; the fourth segment slightly longer than broad.
Radial cross-vein a little less distant from R, than is M,,, from M, on
the borders of the cell R,,..

First segment of the abdomen seen from above deeply coneave ; the
abdomen closely, rather coarsely punctate at base, sparsely and more
minutely at apex; truncate surface of last segment extensive, the
superior processes acute, the inferior small blunt tubercles; process of
the third ventral segment with a median groove on its summit, ante-
riorly with 2 blunt teeth.

Squama with a patch of setae on the outside but with only short,
appressed and inconspicuous pubescence within; the sagitta short and
stout, trigonal, the angles sharp, the apex moderately obtuse; uncus
deflexed, contracted just beyond the apex of the sagitta, with a ridge
but no barb beneath.

2. Colored as in the males, except the front is black or mostly so,
the mesopleura, metapleura, and propodeum more largely tawny, the
mesonotum posteriorly with a medial ferruginous wash. Front and
vertex closely, rugosely, behind the ocelli sparsely, punctate; elypeus
very coarsely and deeply and densely punctate; mesonotum densely
punctate on the flattened but not depressed posterior portion, punctu-
late but more or less obsoletely so in the middle, and with scattered,
coarser punctures; scutellum with a weak median carina at base, sub-
obsoletely punctulate, sides of propodeum sharply mucronate ; abdomen
opaque, very densely punctured.

Types.—Lectotype, 9 [American Entomological Society, no. 2098] ;
allotype [American Entomological Society].

Texas: 2 ¢, 4 9 [including types; American Entomological So-
ciety] ;2 ¢, 22 [U. 8S. National Museum].

Pseudomasaris (Pseudomasaris) marginalis Cresson
Figures 20, 59, 60, 61, 79, 80
1864. Masaris marginalis Cresson, 9. Proceedings of the Entomological
Society of Philadelphia, 3:677.
1904. Pseudomasaris marginalis Dalle Torre. Genera Insectorum, fase. 19,
105 (Sh

¢. Black, markings on body light chaleedony yellow and on the
legs and antennae amber yellow, as follows: most of elypeus, large spot
above, labrum, mandibles obscurely, inner orbits above the emargina-
tion very narrowly, narrow line behind the eyes, tip of scape, third

Vow. 1] Bradley —The Taxonomy of the Masarid Wasps 425

antennal segment, fourth to seventh antennal segments except a black
spot beneath at the apex of each, undersurface of club except two black
spots, line on humeri, posterior border of pronotum, tegulae, small
spot on angles of propodeum, legs beyond the femora and tips of these,
irregular subapical band of first, third, fourth, and fifth dorsal seg-
ments; three subapical bands on second and sixth dorsal segments ; the
sixth and seventh antennal segments and club stained reddish; wings
stained with yellow, the veins yellowish.

Posterior ocelli removed from the eye by considerably less than
their diameter’s length ; emargination of the eyes broad, not triangular,
broadly rounded at apex; front scarcely depressed beneath the median
ocellus, elevated into a weak median tubercle above the bases of the
antennae, its surface with separated punctures; clypeus moderately
convex, its surface weakly shining, rather obsoletely punctate; labial
palpi with 2 segments, the first elongate, the second about one-half as
long. Fourth, fifth, and sixth segments of the antenna strongly wid-
ened at their apices, the seventh about as wide as long, the fourth and
fifth flattened beneath at their apices, the sixth and seventh flattened
beneath; the club short, irregularly ovate, equal in length to segments
6 and 7, convex above, slightly hollowed beneath.

Humeri rounded, pronotum laterally with well separated, rather
coarse punctures; mesonotum anteriorly similarly punctate, more
closely punctured in the middle, posteriorly flattened but not de-
pressed and closely almost rugosely punctate; scutellum closely pune-
tate, moderately convex, posterior angles of propodeum carinate but
not dentate; pleura with well separated punctures.

Anterior femur and tibia with regular surfaces, somewhat flattened
below, the tarsus not ciliate ; middle femur coneave and ridged beneath
with a prominent tubercle near its apex; middle tibia with regular
surface, flattened beneath, as seen from in front gradually widened
from base to apex, two-thirds as long as the tarsus; metatarsus ap-
proximately equal to the remaining segments united; the fourth seg-
ment about as long as broad; hind tibia three-fifths as long as the
tarsus; metatarsus equal to the remaining segments united; fourth
segment slightly longer than broad. Radial cross-vein distant from
R, by little less than the distance between M,,, and M, on the margins
of the cell R,,..

Punctuation of the abdomen fine and close ; the surface between the
processes of the last segment hardly truncate; superior process blunt,
flattened, the inferior sharp, close together; prominence of third ven-
tral segment with its summit anteriorly flattened.

Exterior surface of squama with a patch of very short, incon-
spicuous setae, the inner surface glabrous except for a group of 15
or 20 stout spines and a less conspicuous group of 3 or 4; sagitta acute,
trigonal, the angles sharp; uncus rather long and slender, decurved
and slightly widened at apex, the tip mucronate.

®. Colored as in the male, except that the face and clypeus are
black, with a triangular yellow spot between the antennae, the latter
are yellow to ferruginous with most of the club fuscous.

426 University of Californa Publications [ ENTOMOLOGY

Head closely granular, punctate, clypeus, closely punctulate with
minute and coarser punctulations; dorsulum densely punctate. Pro-
podeum with rounded, only slightly prominent angles.

Anterior tibiae short and inflated except at base, the cther leg
segments with even surfaces.

Dorsal segments much less constricted at their bases than in the
male, opaque, and densely minutely punctulate.

The male has not been previously described.

Types.—Holotype, 2, American Entomological Society, no. 2097.
Allotype,.%, American Entomological Society.

CoLoRApDo: 6 3, 6 @ [including type; American Entomological So-
ciety]; 4 9 [U. S. National Museum].

New Mexico: June 30, 1902, 9 (H. L. Viereck), [American Ento-
mological Society |.

Pseudomasaris (Pseudomasaris) zonalis Cresson

1864. Masaris zonalis Cresson, ¢, 9. Proceedings Entomological Society
of Philadelphia, vol. 3, p. 674.
1904. Pseudomasaris zonalis Dalle Torre. Genera Insectorum, fase. 19,

pS:

3S. Black, following parts Naples yellow: line entirely surrounding
eyes except beneath; elypeus, labrum, mandibles, large spot on front,
scape except at base, spot on pedicel, entire flagellum except reddish
infuseated area at apex of club, humeri, spot on pronotum below, pos-
terior border of pronotum, tegulae, large spot below, small transverse
spot at apex of secutellum, angles of propodeum, spot on anterior and
middle femur and trochanter beneath, anterior femur beneath and at
apex above, middle and posterior femur at tip, all tibiae and tarsi, the
latter reddish at apex, slightly incised subapical stripes on all dorsal
segments, spot and truneature of the last dorsal segment, spot on sides
of ventral segments 2 to 5.

Posterior ocelli distant from the compound eyes by the length of
their diameter; front but slightly impressed beneath the anterior
ocellus, raised above the bases of the antennae to a low tubercle, rather
closely but weakly punctate; the elyneus moderately convex subobso-
letely punctate; labial palpus a single long slender segment without
sign of subdivision; the maxillary palpus a very small chitinized bulb.
Antennal segments 3 to 5 eylindriecal, not widened at their apices, the
sixth slightly widened at its apex, the seventh more strongly but longer
than wide; the elub ovate, a little shorter than the sixth and seventh
segments together, strongly convex above and below.

Humeri marked by a feeble ridge; pronotum rather closely punc-
tate ; mesonotum densely punctate, the posterior flattened portion more
coarsely ; seutellum moderately convex, closely punctate; pleura with
large separated punctures; angles of the propodeum mucronate.

Under surfaces of anterior femur and tibia flattened, regular ;
tarsus not with fringe of cilia; middle femur with under surface con-

Vow. 1] Bradley —The Taxonomy of the Masarid Wasps 497

cave and contorted, as also the middle tibia, the latter with its inferior
margin seen from the front inflated medially ; middle tibia four-fifths
as long as the tarsus; the metatarsus two-thirds as long as the remain-
ing segments united; the fourth segment about as long as broad; hind
tibia five-eighths as long as the tarsus; metatarsus nearly equal to the
length of the remaining segments; fourth segment longer than broad.

Distance between r and R, considerably less than that between
M,,, and M, on the margin of the cell R,,;.

First dorsal segment, seen from above, with its basal surface plane ;
abdomen rather finely and closely punctate, superior processes of last
segment acute, the inferior, small, acute; surface between, somewhat
concave; process of the third ventral segment with its summit ante-
riorly flattened.

Squama exteriorly with only very short inconspicuous pubescence,
a small patch of setae within on the inferior margin; sagitta trigonal,
long, slender, very acute, slightly curved; uncus moderately slender,
slightly widened before the depressed and deflexed tip; the latter
sharply acute.

2. Colored as in the male except as noted; tip of the mandible’
dusky ; labrum black; yellow on elypeus confined to median bar and
two lateral spots; only a spot on seape and third segment of antenna
yellow; humeral band confluent with spot below; pronotum, larger .
apical stripe on scutellum, spot on posterior coxa and more of hind and
middle femora, yellow; broader stripe on each dorsal segment, broad
apical stripe on second and fourth ventral segments, spots on side of
sixth ventral segment yellow.

Hind oeelli little closer than their diameter’s length from the com-
pound eyes; front closely, rather coarsely punctate ; clypeus shagreened
with sparser, shorter, coarser, punctures. Humeri marked by a rather
sharp transverse ridge ; pronotum densely punctured ; mesonotum very
densely punctate; scutellum convex and densely punctate. Middle
femur with under surface flattened, its lower anterior margin marked
by a ridge; middle tibia, seen from in front, with the basal third con-
tracted. Dorsal segments 2, 3, and 4 somewhat depressed basally ;
dorsal surface of the abdomen opaque, minutely, densely punctate.

Types.—Leectotype: ¢, American Entomological Society, no. 2099.
Allotype: American Entomological Society.

Ipano: Craig’s Mt., 1 ¢ [American Entomological Society ].

Contorapo: 2 4, 11 2 [including types; American Entomological
Society] ; 3 9 [U. S. National Museum]; 3 9 (H. H. Smith), [Cornell
University |.

UraH: Salt Lake City, June 13, 1897, 1 ¢ (H. Skinner), [Amer-
ican Entomological Society |.

Nevapa: 7 4, 3 2 [American Entomological Socicty ].

CALIFORNIA: Giant Forest, in the Sequoia National Park, 6000-
7000 feet elevation, July 21-26, 1907, 3 2 (J. C. Bradley), [Cornell
Univerity] ; Fallen Leaf Lake near Lake Tahoe, 4 6, 2 2, July 12, 15,
1915 (E. C. Van Dyke, L. 8. Rosenbaum), [California Academy of
Sciences, Cornell University]; Pyramid Peak, El Dorado Co., 1 2,

428 University of California Publications | ENTOMOLOGY

July 8, 1912, 8000 feet altitude (E. C. Van Dyke), [ Calif. Acad. Scei.] ;
Carrville, Trinity Co., 2 9, June 6, 1913 (EH. C. Van Dyke), [Calif.
Acad. Sci. and Cornell Univ.] ; Nash Mine, Trinity Co., 1 9, June 18,
1913 (KE. C. Van Dyke), [ Calif. Acad. Sci.].

Pseudomasaris (Pseudomasaris) coquilletti Rohwer
Figure 6
1911. Pseudomasaris coquilletti Rohwer, 3, 9. Proceedings United States
National Museum, vol. 40, p. 555.

3. Black, the following parts wax yellow: mandibles except tip,
labrum, clypeus, stripe on front broadened at base of antennae, line
all around the eyes, stripe on maxillae, antennae (shaded apically with
ferruginous, the club dusky beneath), pronotum except anterior stripe
and stripe in front of tegulae, greater part of mesopleura, 2 lines on
mesonotum, interrupted medially, apical half of scutelluam, narrow
line on postseutellum, large lateral spot including angles of propo-
deum, front coxae beneath, trochanters, tibiae except basal spot above,
front tibiae (shaded at tip with ferruginous), middle and posterior
coxae, trochanters and femora, the latter with basal black stripe above,
tibiae and tarsi (the apical segments ferruginous), broad apical stripe
on each tergite, that on the first six separated medially from the apical
margin by a narrow black stripe, two spots on first sternite and most
of rest of the venter; wings stained shghtly yellowish, the veins tes-
taceous.

Emargination of eyes narrow, linear, obtuse; posterior ocelli re-
moved from the compound eyes by less than their diameter front with
a slight fovea below the anterior ocellus, terminating in a weak tubercle
above the base of the antennae, closely and shallowly punctulate, the
punctulation larger on the black area; clypeus strongly convex with
small seattered punctures; segments 38-5 of antennae eylindrical,
scarcely enlarged at their apices, six and seven each widened uni-
formly from base to apex, not flattened, forming the base of the
slender club which is more or less convex above and below; the remain-
ing segments fused, a little truncate at tip, together slightly shorter
than segments 6 and 7.

Humeral angles rounded, marking the termination of a weak trans-
verse ridge; pronotum with small sparse punctures; mesonotum very
densely, finely, punctate, matte; posterior medial portion slightly flat-
tened, punctate like the rest of mesonotum ; mesopleura with scattered
punctures; scutellum convex, densely punctate, with a median basal
ridge; propodeum laterally alate and sharply dentate, the posterior
margins of the alae sinuate; posterior surface of propodeum densely,
finely, punctulate, its lateral surfaces impunctate, matte.

Front femur widened at basal third, tibia flattened and somewhat
irregular beneath ; middle femur with fossa, ridge and nodule beneath ;
tibia flattened beneath, the undersurface more irregular than that of
front pair. Radial cross-vein about as far from R, as M, is from M,,,
on the borders of the cell R,,..

Vou. 1] Bradley.—The Taxonomy of the Masarid Wasps 429

Second to fourth dorsal segments constricted basally ; first segment
almost flat basally; tergites densely and very finely punctate in the
middle, becoming imperceptibly punctulate laterally and more sparsely
but distinctly punctate basally; four processes of last segment acute,
the inferior ones smaller and closer together; second sternite raised
at base, the raised portion divided by a median longitudinal depressed
line; process of third segment with a truncate molar-like anterior
surface, posteriorly produced into an acute tooth.

?. Black; the following parts yellow: mandibles except tip, clypeus
except 2 crescent shaped spots, labrum, large triangular spot above
elypeus, inner orbits convergent behind the ocelli, broad line behind
the eyes, spot on scape and third antennal segment, humeri very
broadly, posterior margin of pronotum broadly, tegulae, most of meso-
pleura, spot on metapleura, 2 longitudinal lines on mesonotum, apical
half of scutellum, postscutellum, propodeum except for 2 triangular
spots, all coxae and trochanters in front, femora in front and at tips
behind; the anterior femur except at base behind, all tibiae and tarsi,
broad subapical band on first dorsal segment, incised anteriorly, sub-
apical band on second, third, and fourth, occupying entire segment at
sides, fifth dorsal segment, sixth at base, spots on first ventral segment,
second ventral segment except at base, third, fourth, and fifth ventral
segments, and the sixth except at base and apex; antennal club ferru-
ginous, yellow beneath; third, fourth, fifth, and sixth antennal seg-
ments shading from yellow into ferruginous; wings stained yellow;
veins, yellow to ferruginous.

Front granular, clypeus finely so, hind ocelli equally distant from
each other and from the eyes.

Humeral angles marked by a feeble ridge; dorsum granular, more
- finely so on the flattened portion of the mesonotum ; seutellum granular,
with larger scattered punctures; pleura with shallow punctures.

Middle femur flattened beneath, the anterior lower margin with a
ridge; the tibia seen from in front gradually widened from base to
apex, three-quarters as long as the tarsus; metatarsus nearly as long
as the following segments united ; posterior tibia three-quarters as long
as the tarsus; this slightly exceeding the remaining segments united.
The radial eross-vein distant from R, by a little over one-half the
distanee between M.,, and M, on the margin of the cell R,...

Abdomen opaque, closely punctured dorsally; second and third
dorsal segments depressed at base.

Types —d,? [U. S. National Museum, no. 13734].

CauiFoRNIA: Los Angeles Co., April, 3, 2 (D. W. Coquillett),
[types, U.S. National Museum] ; Claremont, <4, 2 [Cornell University ] ;
Southern California, 4 2 [American Entomological Society]; Sierra
Nevada, 2 9 [American Museum of Natural History] ; Soboba Springs,
Riverside Co., 2 9, June 1, 1917, on Eriodyction crassifolium (EK. P.
Van Duzee), [California Academy of Sciences] ; Southern Sonoma Co.,
1 4, April 16, 1911 (J. A. Kusche), [California Academy of Sciences].

Mr. Van Duzee informs me that he observed 4 specimens at Soboba
Springs, all visiting Yerba Santa, Hriodyction crassifolium, but was
able to collect only two of these.

430 University of California Publications | ENTOMOLOGY

Subgenus Cotyledon, new subgenus

3. Eyes more remote from each other than the distance between
the hind ocelli, these not touching the eyes; emargination of the eye
moderately narrow, the apex rounded; front with a median prominence
above the base of the antennae; clypeus much broader than long, con-
vex, deeply emarginate anteriorly; labial palpi of 2 segments, the
second less than one-half the length of the first and indistinetly dis-
erete therefrom; maxillary palpi a very minute, searcely chitinized,
bulb bearing one heavy seta and 2 more slender ones.

Seape globose; pedicel about one-third its length; segment 3 cylin-
drical ; segment 4 a little longer, slightly and suddenly widened before
its apex; segment 5 slightly gradually widened; segment 6 greatly
widened toward its apex, nearly as wide as long, apical portion concave
beneath and forming part of the club; the seventh segment about 3
times as wide as long; the remaining segments fused, together about
as long as the sixth and seventh segments; the club formed of the
sixth, seventh, and remaining segments, strongly concave beneath and
with the shape of a partly closed hand.

Undersurfaces of front and middle femora and tibiae concave, con-
torted, and tubereulate ; middle tibia and tarsus about equal in length;
metatarsus as long as the remaining segments united; the fourth seg-
ment much broader than long; hind tibia about three-quarters as long
as the tarsus; the metatarsus about equal to the remaining segments
united, without an apical lobe; the 4th segment about as broad as long.
Distance between r and R, equal to one-third the distance between
M.,, and M, on the margin of the cell R,,..

Basal abdominal segments considerably constricted ; the last dorsal
segment without a definitely truncate apical area but with 4 acute
processes of which the inferior are the larger, closer together and
slightly closer to the superior processes than to each other; second
ventral segment with two rounded tubercles; the third with a promi-
nent transverse ridge replacing the ordinary process, armed with a
sharp median tooth directed caudad. —

The squama densely ciliate within ; sagitta short, trigonal, acute, the
upper margin very deeply triangularly notched; uncus slender, de-
flexed, with two small barbs at base.

°. Posterior ocelli about equidistant from the eyes and each other ;
eyes broadly emarginate. Humeri fitting closely around the head and
marked by a prominent ridge; posterior angles of propodeum mucro-
nate. Middle femur seen from in front, with its inferior margin
sinuate, slightly inflated in the middle; middle tibia, from an external
view distinetly inflated at about the middle, about three-quarters as
long as the tarsus; metatarsus as long as the following segments
united; fourth segment nearly as long as broad; hind tibia nearly
three-quarters as long as the tarsus; metatarsus about as long as the
remaining segments together; the fourth segment as long as broad.

Vow. 1] Bradley.—The Taxonomy of the Masarid Wasps 431

The radial cross-vein opposite R, or separated therefrom by a distance
not greater than that between r-m and M,,,. Dorsal segments 2, 3, and
4 slightly depressed at base.

Type.—Masaris edwardsii Cresson.

Pseudomasaris (Cotyledon) edwardsii Cresson
Figures 10, 31-33, 78, 108

1872. Masaris edwardsii Cresson, g, 9. Transactions American Entomo-
logic Society, vol. 4, p. 87.

1904. Pseudomasaris edwardsi Dalle Torre. Genera Insectorum, fase. 19,
p. 8.

¢. Black; the following parts mustard yellow: elypeus, labrum,
mandibles, except tips, large spot on front, interior orbits, line behind
the eyes, antenna except stripe on segments 38 to 5 above, and posterior
part of the club above, humeri, spot below, posterior border of the
pronotum, tegula, large spot below, narrow subapical line on seutellum
interrupted medially, two small basal spots on propodeum, propodeal
angles, spot below, spot on all coxae, anterior and middle trochanters
beneath, anterior and middle femora except above at base, posterior
femur at apex, extending nearly to the base in front, all tibiae and
metatarsi, subapical band on first dorsal segment, three subapical bars
on second, subapical band deeply incised on third and fourth, apical
half of fifth, two-thirds of sixth, and half of seventh dorsal segment,
except tubercles, band on second ventral segment, on the third, apical
two-thirds of the fourth, fifth entirely banded, and irregular marks on
sixth ; wings hyaline, slightly infuscated in the cell 2d R, + R.; veins
ferruginous.

Front densely but rather finely punctate with an obsoletely punce-
tate tubercle above the antennae, strongly depressed before the base
of the clypeus; this obsoletely punctate, moderately convex.

Pronotum closely and rather coarsely punctate, mesonotum densely
punctate, the posterior portion flattened with coarser and confluent
punctuation ; scutellum slightly convex, anteriorly finely, posteriorly
more coarsely punctate; pleura with separated punctures. |

Anterior femur with a median tubercle and ridge beneath; the
posterior margin of the tibia seen from above, sinuate; tarsi depressed
and broad, without a prominent fringe of cilia, segments 2 to 4 very
short and broad; middle femur irregularly concave, carinate, and
tubereulate beneath; middle tibia contorted, concave, and irregularly
earinate beneath and in front, the anterior margin from an external
view with a strong median tooth, the apex with a group of short spines
in front; hind femur with a brush of dark hairs beneath at base, the
surface slightly irregular.

Abdomen rather finely densely punctate; tubercles of the second
ventral segment polished and shining; the surface of the fourth, fifth,
and sixth ventral segments short tomentose.

432 University of California Publications | ENToMOLOGY

Squama blunt, the inner surfaces with long coppery ciliae; uncus
about two-thirds the length of the squama, slender, acute, deflexed.

?. Coloration differing from the male as follows: labrum and two
bars on the elypeus black, yellow spot on scape and segments 3 and 4
of the antenna; humeral stripe broader, confluent with spot below;
pleural spot much larger; a spot on metapleura, two lines on mesono-
tum, apical half of seutellum, propodeum entirely except for posterior
V-shaped black band, yellow; anterior coxa and trochanter and most
of middle trochanter black; middle and posterior coxa yellow in front,
more of posterior femur yellow; broad apical band on each dorsal
segment, that on first bearing two black spots; spot on first ventral,
most of second ventral, broad apical bands on following three segments,
and two large spots on sixth ventral segment, yellow.

Front and elypeus granular. Humeri fitting closely behind the
head, marked by a distinet transverse ridge; pronotum closely pune-
tate; mesonotum densely punctate, posterior flattened part granular ;
scutellum slightly convex, finely granular; propodeal angle ending in
a rather long spine. Abdomen opaque, densely, closely punctate.

Types.—Leetotype, ¢: American Entomological Society, no. 2096.
Allotype, 2: American Entomological Society.

UtaH: Logan Cajon, July 24, 1906, 1 ¢ (Dr. P. B. Homer), [Cor-
nell University].

NevApDA: 3 4, 6 2 [American Entomological Society |.

CALIFORNIA: Felton, Santa Cruz Mts., May 20-25, 1907, 2 J 3 9,
between 300 and 500 feet elevation (J. C. Bradley), [Cornell Univer-
sity] ; Claremont, 1 #7, 3 9 (C. F. Baker), [Pomona College and Cornell
University]; 3 ¢, 3 2 [including types, American Entomological So-
ciety] ; Fallen Leaf Lake near Lake Tahoe, 2 3, 2 9, June 29 and July
11 and 12, 1915 (KE. C. Van Dyke), [California Academy of Seiences
and Cornell University]; Carrville, Trinity Co., 2 9, June 6, 29, 1903
(KE. C. Van Dyke), [California Academy of Sciences].

WASHINGTON: Cheney, June 30, 1908, and May, 1906, 2 9 [Cornell
University |.

A number of specimens of this species were taken by the author
on different days within a very short area along an abandoned, over-
grown road leading south from the residence of Mr. Trotz at Felton,
California. They were flying leisurely over grass, close to the ground.

Vou. 1] Bradley.—The Taxonomy of the Masarid Wasps 433

Trimeria Saussure
Figures 23, 38, 39, 53-55, 84, 105

1912. Trimeria Zavattari. Archiv fiir Naturgeschichte, vol. 78, pt. A,
no. 2, p. 58. Redescription.

&. Head transversely quadrate ; the temples narrow, margined pos-
teriorly, rectangular below; eyes triangularly emarginate, the apex of
the incision slightly obtuse, distant from one another above; the ocelli
in an equilateral triangle; vertex prominent; front nearly flat, with-
out tubercles; elypeus slightly convex, shield-shaped, its anterior mar-
gin broadly shallowly emarginate; mandibles broad, obliquely triden-
tate toward the apex; hgula elongate and retractile; labial palpus
completely 3-segmented ; maxillary palpus reduced to a transparent,
very short, conical tuberele. Antenna much shorter than the thorax,
of 12 distinct segments, not clavate but the flagellum slightly fusiform ;
seape long, cylindrical, pedicel less than one-third as long, remaining
segments short, as broad or broader than long, the seventh to the tenth
dilated a little mesally beneath.

Dorsal surface of pronotum anteriorly transversely margined;
parapsidal furrows wanting; tegula small, oval, not covering the base
of the seutellum, the outer margin not sinuate, or scarcely so; seutellum
rather flat, in some species posteriorly margined with a reflexed rim
overhanging the postsecutellum; posterior surface of propodeum slop-
ing, the angles mucronate.

Forewing not plaited, R, absent, the cells R, and R, therefore
coalesced ; m-cu attached to Cu,, which from that point turns down-
ward to meet M,. Anterior trochanter armed at apex with a process
with two upturned lamellate edges; anterior femur with its posterior
inferior edge somewhat angled and sinuate; middle femur flattened
beneath; tibiae with regular surfaces; anterior tibial spur strongly
eurved, slender, acute; middle tibia with a single apical spur; larger
posterior tibial spur bifid at apex; tarsal claws simple.

Abdomen sessiie, the basal dorsal segment squarely truneate at
base, the apical segment short, hoodlike, strongly decurved toward and
weakly notched at the apex; second and third ventral segments un-
armed; last ventral segment with its apical border truncate.

Squama ending in an upturned acute hook, a tubercle on the inner
side apicad of the sagitta, which is reduced to a larger tubercle borne
on the inner side of the squama; uneus very broad and flat, obtuse,
beneath with two long sharp barbs at base.

I have not seen a female.

Type—Trimeria americana Saussure, genus monobasic.
Habitat —Brazil and Argentina.

434 University of California Publications | ENTOMOLOGY

LIst OF SPECIES

americana Saussure, 9. Brazil.
1853. Erynnis americana Saussure. 9. Bull. Société Entomologique de
Mrancey (3) Lexx) lana
(3) ps =xanns 2
buyssoni Brethes, g, 9. Argentine, Paraguay.
1904. Trimeria buyssoni Brethes, 2. Anales del Museo nacional de Buenos
Aires, (3), vol. 2, p. 371.
1905. Trimeria buyssoni Du Buysson, §. Bulletin de la Société Entomo-
logique de France, 1905, p. 10.
1912. Trimeria buyssoni Zavattari, g, 9. Archiv fiir Naturgeschichte,
vol. 78, pt. A, no. 2, p. 59.
howardi Bertoni. Argentina.
1912. Trimeria howardi Bertoni. Anales del Museo national de Buenos
Aires, (3), vol. 22, p. 104.
joergenseni Schrottky, 3, 9. Argentina.
1909. Trimeria joergenseni Schrottky, 9. Revista del Museo del la Plata,
ViOls LG. posi.
1910. Trimeria joergensent Brethes, g. Anales del Museo nacional de
Buenos Aires, vol. 20, p. 285.
1912. Trimeria joergenseni Zavattari, g, 2. Archiv fiir Naturgeschichte,
VOLS sDiyeAy MOA aso.
neotropica (Moesarya) Du Buysson, 0, Q.
1906. Jugurtia neotropico Moesarya, g. Annales Histoico-Naturales Musei
Nationalis Hungarici, vol. 4, p. 197.
1910. Trimeria neotropica Du Buysson, g. Zoologische Jahrbiicher, Abt.
fiir Syst., vol. 49, p. 241.
1912. Trimeria neotropica Zavattari, J, 9. Archiv. fiir Naturgeschichte,
Vole 785) ptarAy MO. 2.0.00.

KEY TO THE SPECIES *

Zavattari, Edoardo. Archiv fiir Naturgeschichte, 1912, pt. A, no. 2, p. 59.

a)

oy

EXPLANATION OF PLATES

PLATE 2

Fig. 1. Euparagia scutellaris Cresson, 9.
Fig. 2. Euparagia scutellaris Cresson, 3.
Fig. 3. Paragia decipiens Shuckard, ¢.

[436]

UNIV CARIES PUBIE ENTOMOL. VOL.

7

[ BRADLEY |

PLATE

2

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

Bo) oes) SO iS

PLATE 3

Gayella ewmenoides Spinola, ¢.
Masariella alfkeni (Du Buysson), J.
Pseudomasaris coquillettt (Rohwer), ¢.
Celonites abbreviatus (Villers), ¢&.
Paraceramius lusitanicus (Klug), &.
Ceramius fonscolombei Latreille, ¢.

[438]

UNIV. CALIF. PUBL. ENTOMOL. VOL. 1 LP BRADEBY | PLATE <s

So

3
i
i)

ra

PLATE 4

Pseudomasaris edwardsii (Cresson). Genitalia.
Euparagia scutellaris Cresson. Genitalia.
Ceramius fonscolombei Latreille. Genitalia.
Paragia tricolor Smith. Trophi, 9.

Euparagia scutellaris Cresson. Trophi.
Ceramius fonscolombei Latreille. Trophi.

. [440]

UNIV, CALIF. PUBL. ENTOMOL, VOL. 1 DBRADERY IN PREATE =4:

PLATE 5

Fig. 16. Paraceramius lusitanicus (Klug). Trophi.
Fig. 17. Paraceramius lusitanicus (Klug). Central portion of tongue, much
enlarged.

Fig. 18. Masaris vespiformis Fabricius. Trophi.

[442 ]

UNIV. CALIF. PUBL. ENTOMOL. VOL, 1 [ BRADLEY] PLATE

aie

Pad

j PLATE 6

Fig. 19. Pseudomasaris occidentalis (Cresson). Trophi.
Fig. 20. Pseudomasaris marginalis (Cresson). Trophi, the tongue retracted.
Fig. 21. Pseudomasaris vespoides (Cresson). Trophi.

[444]

UNIV. CALIF. PUBL. ENTOMOL. VOL, 1 [ BRADLEY] PLATE 6

PLATHS7

Fig. 22. Celonites abbreviatus (Villers). Trophi.
Fig. 23. Trimeria buyssoni Brethes. Trophi.

[446] |

t

PLATE

[ BRADLEY |

1

BUBIE ENTOMGE, VOL,

UNIV. CALIF.

PLATE 8

Paragia decipiens Shuckard. Right tegula.

Paragia decipiens Shuckard. Right lateral view of scutellum and
postscutellum.

Euparagia scutellaris Cresson. Right lateral view of scutellum and
postscutellum.

Euparagia scutellaris Cresson. Right tegula.

Masaris vespiformis Fabricius. Lateral view of right side of pos-
terior part of thorax and first abdominal segment.

Masaris vespiformis Fabricius. Dorsal view of right side of poste-
rior part of thorax and first abdominal segment.

Masaris vespiformis Fabricius. Left tegula.

Pseudomasaris edwardsii (Cresson). Right lateral view of scutellum
and postscutellum.

Pseudomasaris edwardsii (Cresson). Dorsal view of scutellum and
postscutellum.

Pseudomasaris edwardsi (Cresson). Right tegula.

Celonites abbreviatus (Villers). Right lateral view of thorax and
basal segment of abdomen,

Celonites abbreviatus (Villers). Dorsal view of thorax and basal
segment of abdomen.

Ceramioides capicola (Brauns). Right lateral view of scutellum and
postscutellum.

Paraceramius lusitanicus (Klug). Left tegula.

Trimeria buyssoni Brethes. Right lateral view of scutellum and
postscutellum.

Trimeria buyssoni Brethes. Right tegula.

Masariella alfkeni (Du Buysson). Left lateral view of posterior
part of thorax and first abdominal segment.

Masariella alfkent (Du Buysson). Left tegula.

[448]

UNIV, CALIF..PUBL. ENTOMOL. VOL. 1

[ BRADLEY] PLATE 8

PLATE 9

Euparagia scutellaris Cresson. Genitalia, dorsal view.

Paragia tricolor Smith. Genitalia, dorsal view.

Paragia tricolor Smith. Genitalia, ventral view.

Paragia tricolor Smith. Genitalia, right lateral view.
Paraceramius lusitanicus (Klug). Genitalia, right lateral view.
Paraceramius lusitanicus (Klug). Genitalia, dorsal view.
Paraceramius lusitanicus (Klug). Genitalia, left internal view.
Masaris vespiformis Latreille. Genitalia, right lateral view.
Masaris vespiformis Latreille. Genitalia, dorsal view.

Masaris vespiformis Latreille. Genitalia, left internal view.
Celonites abbreviatus (Villers). Genitalia, dorsal view.

[450]

UNIV SCALE PUBS ENTOMOL. VOI, 4 [ BRADLEY] PLATE 9

PLATE 10

Trimeria buyssoni Brethes. Genitalia, right lateral view.

Trimeria buyssoni Brethes. Genitalia, dorsal view.

Trimeria buyssoni Brethes. Genitalia, left internal view.

Pseudomasaris
Pseudomasaris
Pseudomasaris
Pseudomasaris
Pseudomasaris
Pseudomasaris
Pseudomasaris
Pseudomasaris

Pseudomasaris

texanus (Cresson). Genitalia, right lateral view.
texanus (Cresson). Genitalia, dorsal view.
texanus (Cresson). Genitalia, left internal view.
marginalis (Cresson). Genitalia, right lateral view.
marginalis (Cresson). Genitalia, dorsal view.
marginalis (Cresson). Genitalia, left internal view.
occidentalis (Cresson). Genitalia, right lateral view.
occidentalis (Cresson). Genitalia, dorsal view.

occidentalis (Cresson). Genitalia, left internal view.

[452]

UNIV. CALIF. PUBL. ENTOMOL

[ BRADLEY ]

PEATE 10

PAE!

Fig. 65. Pseudomasaris vespoides (Cresson). Genitalia, right lateral view.
Fig. 66. Pseudomasaris vespoides (Cresson). Genitalia, dorsal view.

Fig. 67. Pseudomasaris vespoides (Cresson). Genitalia, Left internal view.
Fig. 68. Huparagia scutellaris Cresson. Face, 9.

Fig. 69. Euparagia scutellaris Cresson. Face, 2.

Fig. 70. Huparagia scutellaris Cresson. Antenna, dg.

Fig. 71. Euparagia scutellaris Cresson. Antenna, 9.

Fig. 72. Euparagia scutellaris Cresson. Maxilla.

Fig. 73. Euparagia scutellaris Cresson. Labium.

Fig. 74. Euparagia scutellaris Cresson. Posterior aspect of right anterior
trochanter.

Fig. 75. Paragia tricolor Smith. Ultimate dorsal segment.
Fig. 76. Paragia tricolor Smith. Clypeus.

[454]

iE BiRADEEY 1) PLATE "14

UNIV. CALIF. PUBL. ENTOMOL. VOL. 1

Fig.

ig. 81.

Tle
. 78.

5 Wo)

b tellb

oo
. 83.
. 84,
SCs
g. 86.

PLATE 12

Pseudomasaris bariscipus. Face. Drawing by Carol Bradley.

Pseudomasaris edwardsti (Cresson). Inferior and lateral views of
club of antenna. Drawing by Carol Bradley.

Pseudomasaris marginalis (Cresson). Face. Drawing by Carol
Bradley.

Pseudomasaris marginalis (Cresson). Antenna. Drawing by Carol
Bradley. ;

Pseudomasaris occidentalis (Cresson). Antenna. Drawing by Carol
Bradley.

Paraceramius lusitanicus (Klug). Base of antenna.
Masaris vespiformis Fabricius. Antenna.

Trimeria buyssoni Brethes. Antenna.
Pseudomasaris bariscipus. Antenna.

Pseudomasaris vespoides (Cresson).

[456]

UNIVGSCARIE PUBES ENTOMOL; VO, 4 PBRADEEY | PEATE, 12

84

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

87.
88.
89.
90.
oie
92.
93.
94.
95.
96.
lo

PLATE 13.

Celonites abbreviatus (Villers). Labium.

Celonites abbreviatus (Villers). Maxilla.

Celonites abbreviatus (Villers). Labial palpi, 9.
Celonites abbreviatus (Villers). Maxillary palpi, 9.
Paragia decipiens Shuckard. Anterior tarsal spur.
Paragia tricolor Smith. Anterior tarsal spur.
Euparagia scutellaris Cresson. Posterior tarsal spur.
Paragia decipiens Shuckard. Posterior tarsal spur.
Masaris vespiformis Fabricius. Posterior tarsal spur.
Pseudomasaris vespoides (Cresson). Posterior tarsal spur.
Gayella ewmenoides Spinola. Wings.

: [458]

UNIV’ (CAELE: PUBL, SENTOMO@E, VOLE. 1 [ BRADLEY] PLATE 13

=

oy,

PLATE 14

Fig. 98. Mischocyttarus labiatus. Wings. Veins labeled: p.l., posterior lobe;
ax, exc., axillary excision.

Fig. 99. Vespa diabolica. Wings. Cells labeled: pr. exc., preaxillary excision.
Fig. 100. Huparagia scutellaris Cresson. p.l., posterior lobe.

[460]

UNIV. CALIF. PUBL. ENTOMOL. VOL, 1 [ BRADLEY] PLATE 14

Fig.
Fig.
Fig.
Fig.

101.
102.
108.
104.

PLATE 15

Paragia decipiens Shuckard. Wings.
Ceramioides capicola (Brauns). Wings.

Para Ceramius lusitanicus (Klug). Forewing.
Celonites abbreviatus (Villers). Wings.

[462 |

UNIV. CALIF. PUBL. ENTOMOL. VOL, 1

Fig. 105.
Fig. 106.
Fig. 107.
Fig. 108.

PLATE 16

Trimeria buyssoni Brethes. Wings.
Masariella alfkeni Du Buysson. Wings.
Masaris vespiformis Fabricius. Wings.
Pseudomasaris edwardsi (Cresson). Wings.

[464]

TOMOL. VOL. 1

—

INDEX

UNIv. CaAtir. Pusu. ENTOM., Vou. 1

Titles of papers and names of new systematic units in boldface.

Acerophagus, 354.
fasciipennis, 347, 348-349, 351.
gutierreziae, 348, 351.
notativentris, 347, 348, 349-350,
351.
pallidus, 348, 350-351, 352 (fig.).
texanus, 347, 348, 349, 350, 351.

Acer platanoides (Norway maple),
host for Drepanaphis and Dre-
panosiphum, 321, 322.

saccharinum (silver maple), 321,
322.

Achillea millefolium (yarrow), host
for Macrosiphum solanifolii, 329.

Acknowledgments, 3, 370.

Adalia bipunctata, 252, 284-288, 293,
295, passim.

Adenostoma, 227.

Aeschna, 31, 79.

Agallia californica, 248.

Agrion, wing of, in flight, 23, 26.

Air sacs, in bodies of flying insects,
29, 30.

Aligia jucunda, 248.

inseripta, 249.

Alisma plantago (water plantain),
host for Rhopalosiphum nym-
phaeae, 331.

Alnus rhombifolia (white alder), host
for Eucallipterus flavus, 323.

Alopecurus pratensis (meadow fox-

tail), host for Macrosiphum
granarium, 328.

Alydus pluto, 231.

Amaranthus retroflexus (pigweed),

host for Aphis middletonii, 340.
Ampelodesma tenax, host for Macro-
siphum granarium, 328.
Amphorophora latysiphon, 329.
rubi, 329,
Angelica tomentosa, host for Aphis
cari, 318, 320.
Anise, wild or sweet, host for Sipho-
coryne capreae, 342.
Anthocoris antevolens, 235.
bakeri (ornatus), 235.
melanocerus, 235.
Aphalara calthae, 249.
Aphididae of California, 301-346.

Aphis albipes, 336, 337.
angelicae, 318.
avenae, 337, 340.
bakeri, 337.
brassicae (cabbage aphis), 263, 337.
eardui, 338.
cari, 317-321.
ceanothi, 338.
gossypii (melon aphis), 263, 338.
hederae, 339, 341.
helianthi, 339.
heraclei, 339.
maidis, 340.
malifoliae, 339.
medicaginis, 340.
middletonii, 340.
nerii (oleander aphis), 263, 340.
oenotherae, 341.
persicae-niger, 341.
pomi, 341,
rumicis (bean aphis), 263, 341.
salicicola, 342.

‘ sambucifoliae, 342.
Aphrophora permutata, 245.
Aphycus, resemblance to

coccobius, 355.
oxacae, 355.
Apocremnus anchorifer, 243.
nigerrimus, 243.
politus, 243.

Apple, host for Aphis malifoliae,
339, Aphis pomi, 341, and Erio-
soma lanigera, 345.

Aquilegia chrysantha, 317.

truneata (native columbine), host

for Mysus aquilegiae, 317.
Aradus behrensi, 233.

borealis, 2338.

debilis, 231.

falleni, 234.

hubbardi, 232.

insolitus, 233.

persimilis, 232.

Arbutus menzeisii (madrone), host
for Rhopalosiphum arbuti, 330.

unedo (strawberry tree), 330.

Archilestes, 97.

Arctostaphylos manzanita (manza-
nita), host for Rhopalosiphum
arbuti, 330.

Pseudo-

[465]

Index

Argyrocoris femoratus, 225.
seurrilis, 225, 226.

Artemisia heterophylla (worm-wood),
host for Macrosiphum Judovici-
anae, 328.

Asclepias speciosa (milk-weed), host
for Aphis gossypii, 338, and for
Aphis nerii, 341.

vestita, 339, 341.

Asters, roots of, hosts for Aphis mid-
dletonii, 340.

Astragalus, flowers of, Neomexicanus
upon, 416.

Atractocerus, 129.

Baecharis douglasii, host for Rho-
palosiphum persicae, 332.

Balelutha impicta, 249.

Bamboo, host for Myzocallis arund-
inariae, 303, 305, 310.

Banana, host for Pentalonia nigro-
nervosa, 335.

Banasa sordida, 231.

Belostoma, 52, 75.

Berberis vulgaris, host for Liosom-
aphis, 342.

Betula alba (European white birch),
host for Calaphis betulaecolens,
322.

papyrifera, 323.
populifolia laciniata, 322.

Blattidae, 1138-116.

Bolteria picta hirta, 244-245.

Bradley, J. C., 369-464.

Cabbage, host for Aphis brassicae,
337.

Caenia bisetosa, 161.

Caenis, 13.

Calaphis betulaecolens, 322.

Callipterus arundicolens, 302.

Calotermes, 6, 8.

Camellia japonica (camellia), host
for Toxoptera aurantiae, 330.

Camtobrochis, 237, 238.

fulvesecens, 237.
nigrita, 238.
nitens, 237, 238.
validus, 237.

Capnia, 105.

Capsus ater, 237.

Carum kelloggii (wild anise), host
for Aphis cari, ete., 317, 320, 321,
342.

Catalogue of the Ephydridae, with
Bibliography and Description of
New Species, 153-198.

Catalpa, host for Aphis gossypii, 339.

Catonia grisea, 246.

majusculus, 247.
nemoralis, 246.
succinea, 247.

Cauliflower, host for Aphis brassicae,

337.

Ceanothus (wild lilac), host for Aphis

ceanothi, 338.
thrysiflorus, 338.

Celonites, 370, 371, 375, 378, 389, 401,
402-405; species of, 404—405.
Centranthus ruber (Jupiter’s beard),
host for Macrosiphum rosae, 329,
and for Rhopalosiphum persicae,

332.
Ceramioides, 375, 376, 388, 396-397;
species of, 397.
nigripennis, 397.
Ceramiopsis, 388, 393; species of, 393.
gestroi, 393.

Ceramius, 375, 376, 388, 390, 394;

species of, 394-395.
cerceriformis, 397.
fonscolombei, 394.

Cerataphis lataniae, 342.

Chalcid flies, mosquitoes breeding
ground for, 159.

Chermes cooleyi, 346.

Cherry, host for Myzus cerasi, 333,
334, 335.

Chicorium intybus (chicory), host for
Macrosiphum lactucae, 328.
Chromaphis juglandicola (walnut

aphis), 263, 274, 325.

Chrysanthemum (Shasta daisy), host
for Amphorophora latysiphon,
329.

Cicada, 122, 124.

tibicen, 35, 74.
Cicadula laeta, 249.
sexnotata, 249.
Cirrhencyrtus, 360-361.
ehrhorni (fig.), 360, 362.

Cirsium lanceolatum (bull thistle),
host for Aphis cardui, 338.

Cixius basalis, 248.

Clarkia elegans, host for Rhopalo-
siphum persicae, 331.

Clausen, C. P., 251-299.

Clematis brachiata (clematis), host
for Myzus varians, 335.

Coccinella californica, 251, 253-258,
293, 295, passim.

trifasciata, 251, 258-262, 293, 294,
295.

Coccinellidae, most important aphis-
eating species in California, 251—
299.

Coleoptera, 126-129.

Coquillettia insignis, 240.

Corizus erassicornis, 231.

hyalinus, 231.
indentatus, 231.
scutatus, 231.

Corn, host for Aphis avenae, 340.

Corylus maxima (filbert), host for
Myzoeallis coryli, 324.

Cotoneaster franchetii, host for Aphis
pomi, 341.

Index

Cotyledon, 408, 430-431.
edwardsii, 408, 410.

Crataegus oxyacantha (English haw-

thorn), host for Aphis pomi, 341.

Cycloneda sanguinea, 252, 288-292,
293, 297, 298.

Cyrtomium falecatum (holly fern),
host for Idiopterus, 329.

Dacerla inflata, 239.

Deltocephalus vanduzei, 249.
punctatus, 249.

Deraeocoris fraternus, 238.
ingens, 237, 238.

Dermestes, 128.

Dichrooscytus elegans, 237.
irroratus, 236.
speciosus, 236.
suspectus, 236.

Dicraneura carneola, 249.

Diecyphus agilis, 240.
ealifornicus, 240.
crudus, 240.
vestitus, 240.

Diploptera, 115.

Dipsacus fullonum (fuller’s teasel),
host for Macrosiphum rosae, 329,
and for Rhopalosiphum persicae,
332.

Diptera (flies), venation in wings of,
130-134.

Drepanaphis acerifolii, 321-322.

Drepanosiphum platanoides, 321.

Duzee, HE. P. Van, 199-216, 217-227,
229-249,

Eetopiocerus anthracinus, 235.

Elevator muscles, in flight of insect,
34.

Elidiptera pallida, 246.
septentrionalis, 245, 246.
variegata, 246.
woodworthi, 245,

Elymus (wild rye), host for Macro-

siphum granarium, 328.

Elytroptera, venation in wings of,
113-129.

Embiidae, venation in wings of, 106—
107.

Empoa commissuralis, 249.

Eneyrtinae, new genera and species
of, 347-367.

Ephemeridae (mayflies), venation in
wings of, 98-101.

Ephydra californica, 157.

cinerea, 159-160.
gracilis, 157.
halophila, 157.
hians, 159.
millbrae, 155-159.

Ephydridae, study of, 153-198; bibli-
ography of, 161-168; catalogue,
synopses, key to subfamilies and
to genera of, 168-198; in Tahoe
region, 230.

Epilobium adenocaulon occidentale,
host for Aphis oenotherae, 341.
Eriodyction crassifolium (Yerba
santa), 429.
Eriosoma, 345.
lanigera, 345, 346.

Errhomenellus maculatus, 248.
Eseallonia pulverulenta, host for
Rhopalosiphum persicae, 332.

Eschna. See Aeschna.
Essig, E. O., 301-346.
Eueallipterus flavus, 323.
tiliae, 323.
Euceraphis betulae, 322-323.
Eugerion, 123.
Eugeron, 111.
Eugnathodus abdominalis, 249.
Euparagia, wing of, 373, 374, 378;
mouth parts of, 377; antennae
of, 377; simple claws of, 378;
male genitalia of, 378; genus,
383-384.
maculiceps, 384.
scutillaris, type species of, 385.
Euparagiinae, key for genera of, 381—
382; description of, 383.
Euphyllura arbuticola, 249.
nevadensis, 249.
Eurygaster alternatus, 231.
Euscelis exitiosus, 249.
striolus, 249.
Evening primrose, host for Aphis
oenotherae, 341.
Fagus tricolor, host for Phyllaphis

Fallen Leaf Lake, California, collec-
tion of insects from, 229, 230.
Flight, of insects, different from that
of birds, 21, 22; motion of wing
in, 23-25; air pressure, factor in,
25; wing muscles in, 26; bobbing
motion in, 28; function of air
saes, 29, 30, of elevator muscles,
34.

Forfieulidae (earwigs), 116-119.

Fulgorina, 123.

Gayella, 370.

Gegenbaur, theory of, in wing genesis,
6, 7, 8, 14.

Geocoris pallens decoratus, 234.

Gerarina, venation in wings of, 103—
104.

Gerhardiella delicata, 243.

Gerris gilletei, 235.

orba, 235.
Gypona angulata, 248.
Hadronema militaris, 241.
robusta, 241.

Harmostes reflexulus, 231.

Hawaiian Sugar Planters’ Experiment
Station, 347.

Hedera helix (English ivy), host for
Aphis hederae, 339.

Index

Helianthus annuus (sunflower), host
for Rhopalosiphum persicae, 332,
and for Aphis helianthi, 339.

Helochara communis, 248.

Hemeristina, venation in wings of,

103. ,
Hemerobidae, venation in wings of,
109. :
Hemiptera, taken near Lake Tahoe,
notes on, 229-249. See also
Orthotylini.

Hemipteroidea, fossil insect, 111-112.

Hepialis, 139.

Heracleum mantegazzianum, host for
Aphis heraclei, 339.

Heteroptera, 120-122, 231-245.

Hexagenia bilineata, 99, 100.

Hippodamia ambigua, 251, 25
274, 293, 294, 295, passim.

convergens, 251, 252, 258, 263-269,

274, 293, 294, 295, passim.

Holopticus, 408, 409, 410, 414.

Homoemus bijugis, 231.

Homoptera, 122-124, 245-249.

Homothetidae, fossil insects, venation
in wings of, 101-102.

Hyalochloria bella, 217-218.

caviceps, 218.

Hyalopterus arundinis (plum aphis),
263, 335.

Hydrellia scapularis, 160.

Hymenoptera, venation in wings of,
134-138.

Hyoidea grisea, 220.

Idiocerus amoena, 248.

Idiopterus nephrelepidis, 329, 335.

Iinacora malina, 219.

Insects, Wing Veins of, 1-152.
also Thorax; Wings.

Ischnorrhynchus resedae, 234.

Jones, B. J., 153-198.

Juglans regia (English walnut), host
for Chromaphis juglandicola, 325.

Jugurtia, 388, 397-398; species of,
398.

9 9

oy, 4

70—

See

Koebelia ealifornica, ,248.
irrorata, 248.

Labopidea, 218, 222.
atriseta, 221.
chloriza, 221.
nigripes, 219, 220, 242.
sericata, 220, 221.
simplex, 220.

Labops hesperius, 240.

Laceocera obesa, 248.

Lachnus pini-radiatae, 327.
tujafilinus, 327.

Lake Tahoe, California, Hemiptera

taken near, 229-249,

Largidea grossa, 238.

marginata, 238, 239.

[468]

Laurestinus, host for Aphis pomi,
341.

Lavatera assurgentiflora, host for
Rhopalosiphum persicae, 331.

Leg, development of, in insects, 18,

19%

Lemon, host for Aphis medicaginis,
340.

Lepidoptera, venation in wings of,
138-142.

Leptomastix, 364.
dactylopii, 365.
Liburnia magnifrons, 248.
Life-History and Feeding Records of
a Series of California Coccinel-
lidae, 251-299.
Ligyrocoris diffusus, 234.
Lilium elegans grandiflorum, host for
Myzus circumflexum, 335,
speciosum rubrum, 335, 338.
Liosomaphis berberidis, 342.
Liriodendron tulipifera (tulip tree),
host for Rhopalosiphum persicae,
331.
Lonicera (honeysuckle), host
Aphis gossypii, ete., 339, 342.
Lopidea nigridea, 241.
Lumen, in insects’ wing veins, 42,
43, 51.
Lupinus latifolius (lupine), host for
Macrosiphum albifrons, 328.
arboreus, 328.
Lygaeus bicrucis,
reclivatus, 234.
truculentus, 234.
Lygidea rebucula obscura, 237.
Lygus campestris, 237.
plagiatus, 237.
pratensis, 237.
Macrosiphum albifrons, 328.
californicum, 328.
granarium, 328, 331.
lactucae, 328.
ludovicianae, 328.
rhamni, 328.
rosae (rose aphis), 253, 261, 263,
267, 278, 282, 287, 329, 331.
solanifolii, 329.
stanleyi, 329.
Macrotyloides,
apicalis, 223.
vestitus, 222-223.
Macrotylus angularis, 224.
infuseatus, 242.
lineolatus, 242.
multipunctatus, 242.
verticalis, 225.
vestitus, 222.

Masarid wasps, taxonomy and North
American species of, 369-464.
Masaridinae (Masaridini), synoptie

tables of, 387-390.

for

234,

999

oon.

a

Index

Masariella, 375, 389, 398-400; key to
species of, 399-400.
alfkeni, 399, 400.
saussurel, 399, 400.
Masarinae, key for genera of, 381-
382.
Masaris, 375, 389, 398, 400, 405-406;
species of, 406.
aegyptiacus, 406
alfkeni, 399.
apiformis, 403.
occidentalis, 407.
texanus, 414.
vespiformis, 369, 406.
vespoides, 412.
Mealybugs (Hymenoptera), parasites
of, 347-367.

Megascopteridae, ancient insect group,
venation in wings of, 89-90.
Melianthus major, host for Rhopalo-

siphum persicae, 332,

Metaparagia, 393; species of, 393;

key to, 393.

Mezira moesta, 234.

Microphylellus alpinus, 242.
bicinetus, 243.
modestus, 242
rubrieans, 242.

Micropteryx, 139.

Midas, 133.

Miridae, North American, keys to

genera of, 199-216; explanation
of terms, 200-201.
Mischocyttarus, wing of, 375.
Monterey pine, host for Chermes
cooleyi, 346.

Mosquito. See Ephydridae.

Myzaphis abietina, 342.
rosarum, 342.

Myzoeallis arundicolens, 305-310.
arundinariae, 302, 303, 306.
castaneae, 323-324.
coryli, 324.
essigi, 324.
pasaniae, 324.
quereus, 324.
ulmifolii, 324.
woodworthi, 324.

Myzus aquilegiae, 314-317.
cerasi, 333, 334.
circumflexum, 335.
fragaefolii, 335.
varians, 334, 335.

Nabis ferus, 235.

Nasturtium, host for Rhopalosiphum

persicae, 331.

Nectarosiphum rubicola, 327-328.

Nemura trifasciata, 104.

Neomexicanus, 415.

Neoptera, venation in wings of, 129—

142.
Nephrolepis (Boston ferns), host for
Idiopterus, 329.

Neuroptera, fossil form, type of vena-
tion in wings of, 85.
New Genera and Species of Encyrt-
inae from California Parasitic in
Mealybugs (Hymenoptera), 347—
367.
New or Little Known Genera and
Species of Orthotylini (Hemip-
tera), 217-227.
Noltea africana (soap bush), host for
Aphis ceanothi, 338.
Notes on Some Hemiptera Taken
near Lake Tahoe, California,
229-249.
Notophila bellula, 153.
sealaris, 160.
varia, 153-154.
virgata, 160.

Nysius californicus, 234.
ericae, 234.
ericae minutus, 234.

Odonata, venation in wings of, 92-
97.

Odynerus, 371, 378.

Oenothera biennis, host for Aphis
oenotherae, 341.

grandiflora, 341,

Okanagana ornata, 245.

rimosa, 245.

Oleander, host for Aphis nerii, 340,
341.

Oliarus fidus, 247.

Olla abdominalis, 252, 274-279, 293,
294, passim.

oculata, 252, 279-283, 293, 294, 295,
passim.

Oneopsis pruni, 248.

viriabilis, 248.

Orgerius rhyparus, 245,

Orthoptera, 119-120.

Orthopteroidea, fossil insects, vena-
tion in wings of, 112-113.

Orthotylini, 217-227.

Orthotylus affinis, 241.

compsus, 217.
ecuneatus, 242.
formosus, 241.
insignis, 241.
ovatus, 241.
tibialis, 241.
uniformis, 241.
viridicatus, 241.

Paleopterina, venation in wings of,
102-103.

Paleozoic, insect fossils of, 7, 8, 9, 85,
86, 88, 89, 97, 102, 103, 104, 105,
NOGS MOS hdl AOS 2 O22:

Palephemeridae, Devonian remains,
venation in wings of, 98.

Panorpidae, venation in wings of,
109-110.

Papaver (garden poppy), host for
Aphis rumicis, 341.

Index

Parabolocratus viridis, 248.

Paraceramius, 375, 388, 390, 394, 395;
species of, 396.

lusitanicus, 377, 395.

Paragia, 370, 375, 377, 390-3893; list

of species of, 392; key to, 393.
pictifrons, 393.

Paragiini, synoptic table to genera
of tribe of, 388.

Paraleptomastix, 365.

abnormis, 364.
dactylopii, 365.
histrio, 365.

Paramasaris, 386

fuscipennis, 386-387.

Paraproba hamata, 241.

Parthenicus picicollis, 226.

Parydra appendiculata, 161.

aurata, 154-155.
bitubereulata, 154.

Pasania densiflora (tanbark oak),
host for Myzocallis pasaniae,
324.

Peach, host for Myzus varians, 335,
and for Aphis persicae-niger,
341.

Pear, host for Aphis pomi, 341.

Pemphigus betae, beet louse, 263.

populitransversus, 343-344,

Pentalonia nigronervosa, 335.

Pentstemon spectabilis, host for Rho-
palosiphum, 331, and for Myzus
circumflexum, 335.

Perlidae, venation in wings of, 104—
105.

Phlepsius occidentalis, 249.

ovatus, 249.

Phorodon humuli (hop aphis), 253,
263, 267, 330.

Photinia arbutifolia (mountain holly),
host for Rhopalosiphum, 330.

Phryganeidae, 110-111.

Phthanocorus, 111.

Phylaria, 200, 201.

Phyllaphis fagi, 321.

querci, 321
quercifoliae, 321.
Physopoda, 125-126.
Phytocoris eximius, 235.
inops, 235.

Phytophthires, 124-125.

Picea excelsa (Norway spruce), host
for Myzaphis, 342.

Pieris, 26.

Pinus ponderosa (yellow pine), host
for Lachnus pini-radiatae, 327.

Pittosphorum eugenioides, 332, 341.

tenuifolium, 341.
tobira, host for Macrosiphum solani-
folii, 329, and for Rhopalosiphum
persicae, 331.
undulatum, 332.
Plagiognathus, 224, 243.

Platylygus luridus, 237.

Platypteridae, Paleozoic insect fos-
sils, venation in wings of, 104.

Platytylellus intercidenda, 235.

Plesiozethus, 385, 386.

flavolineatus, 386.
Ploiariodes, 234.
Poeciloscytus uhleri, 235.

venaticus, 235.

Pontia rapae, white butterfly, 23, 27.

Poplar, host for Pemphigus, 344, 345.

Potamogeton natans (pondweed),
host for Rhopalosiphum nym-
phaeae, 331.

Primula (cowslip), host for Amphora-
phora latysiphon, 329.

Propteticus, 102.

Protephemeridae, ancient insect fam-
ily, venation in wings of, 97-98.

Protodonata, ancient insect group,
venation in wings of, 90-92.

Protoperla, venation in wings of,
104.

Prune, host for Hyalopterus arund-
inis, 336.

Psallus, 243.

Pseudococeobius, 355.

clauseni, 358—360.
ehrhorni, 355, 356, 362.
fumipennis, 355, 356-358.

terryi, 355, 356, 357.

Pseudoecoceus, study of, by C. P.
Clausen, 347; Tanaomastix para-
sitie in, 365, 367.

eitri, 367.
erawili, 349.
ryani, 362.
solani, 358.
timberlakei, 355.

yerbasantae, 351.

Pseudomasaris, 370, 371, 372, 375,
389, 397, 398, 399, 401, 403, 406—
411; keys to genera and species
of, 407-411.

albifrons, 416.

bariscipus, 409, 418-419.

basirufus, 411, 421-422.

coquilletti, 409, 411, 428-429.

edwardsii, 431-432.

maculifrons, 411, 420-421.

marginalis, 409, 410, 424-426.

occidentalis, 409, 410, 423-424.

phaceliae, 409, 410, 419-420.

rohweri, 409, 417-418.

texanus, 399, 409, 410, 414-415,

416, 419.

vespoides, 412.

zonalis, 409, 411, 426428.
Pseudopsallus, 224.

angularis, 224-225.

vertiealis, 225.

Psiloglossa simplicipes, identity with
Euparagia, 384.

Index

Psocidae, venation in wings of, 107—
108.

Pterocomma floeculosa, 325, 326.

populifoliae, 327.
Quartinia, 389, 401; species of, 402.
deleta, 401.
dilecta, 401.
variegata, 401.

Quercus agrifolia (coast live oak),
host for Symydobius agrifoliae,
SUZ oie

lobata (valley oak), 321.
robur (English oak), 324.

Quilnus, 234.

Raphiglossinae, 378.

Rhamus ealifornicus (coffee berry),
host for Macrosiphum rhamni,
328.

purshiana (casecara sagrada), 328,
338.
Rhopalosiphum arbuti, 330.
howardii, 330.
lactueae, 331.
nervatum, 330, 331.
nymphaeae, 331.
persicae, 331, 335.
rhois, 331, 334.

Rhus diversiloba (poison oak), host
for Rhopalosiphum rhois, 334.

Rhyparochromus angustatus, 234.

sodalicus, 234.

Rithogena manifesta (mayfly), 12.

Rosa californica (wild rose), host for
Macrosiphum rosae, 329.

Rose, most serious pest of (Myz-
aphis), 342.

Rubus (cultivated blackberry), host
for Amphorophora rubi, 329-330.

parviflorus (thimbleberry), host for
Nectarosiphum rubicola, 328.
vitifolius (wild blackberry), 330.

Rumex occidentalis (western dock),
host for Aphis rumicis, 341.

Saldula interstitialis, 235.

Salix laevigata (red willow), 327.

lasiandra (yellow willow), 327,
328.

lasiolepis (arroyo willow), host for
Pterocomma flocculosa, and for

+ TThomasia salicicola, ete., 325,
326, 327, 328, 342.

longifolia (longleaf willow), 327.

Sambucus glauca (blue elderberry),

329, 342.

racemosa callicarpa (red elder-
berry), host for Macrosiphum
stanleyi, ete., 329, 342.

Scalophila hamifera, 161.

Schizoneura lanigera (woolly apple
aphis), 263, 288.

Scolopostethus thomsoni, 234.

Sialidae, primitive venation in wings
of, 108-109.

Siphocoryne capreae, 342.
xylostei, 342.

Sonchus oleraceus (sow thistle), host

for Rhopalosiphum lactueae, 331.

Sphragisticus nebulosus, 234.

Staphylinus, 128.

Stemmatosteres, 352-353.
apterus, 354-355.

Stenodictyopteridae, fossil insects,
venation in wings of, 101.

Stenodema vicina, 235.

Stictocephala, pacifiea, 245.

Strawberry, cultivated, host for Myzus
fragaefolii, 335.

Subulicornes, 146.

Symphoricarpos racemosus (snow-
berry), host for Aphis albipes,
336, 337.

Symdobius agrifoliae, 311-313.

albasiphus, 3138.

Synoptical Keys to the Genera of the
North American Miridae, 199-—
216.

Tanaomastix, 347, 362-364, 365.
abnormis, 363 (fig.), 365, 366-367.
albiclavata, 367.
claripennis, 363 (fig.), 365-366.

Taxonomy of the Masarid Wasps, in-
cluding a Monograph on the
North American Species, 369-—
464,

Telamona barbata, 245.

obsoleta, 245.

Termitidae, venation in wings of,
105-106.

Thamnotittix longiseta, 249.

Thomasia salicicola, 325-326.

Thorax, in insects, specialized for
attachment of organs of locomo-
uayel, Ilr, alsy, Pale

Thuya occidentalis (arborvitae), host
for Lachnus tujafilinus, 327.

Thyanta custator, 231.

Thyreocoris anthracinus, 231.

Tilia americana (American linden),
host for Eucallipterus, 323.
tomentosa (silver linden), 323.

Timberlake, P. H., 347-367.

Tollius eurtulus, 231.

Toryna, 407, 411.
vespoides, 408, 410.

Toxoptera aurantiae, 330.

Tracheae, presence or absence of, in

veins of insects’ wings, 43-47.

Trachael gills, in wing genesis, 4-14.

Trichopepla atricornis, 231.

Trifolium pratense (red clover), host
for Aphis bakeri, 337.

Trimeria, 369, 370, 375, 390, 433; list
of species of, 434.

americana, 433.

Tuberolachnus viminalis, 327.

Index

Ulnus americanus (water elm), 329.
campestris (English elm), 324, 345,
346.

Umbellularia californica (California
laurel), host for Rhopalosiphum
persicae, 331, and for Myzus cir-
cumflexum, 335.

University of California, campus of,
new species of Aphididae from,
301-846.

Veins, in wings of insects, distribu-
tion of in wing-root formation,
32-38, 36-37, 38, 39; nature of,
40-42; characters in recognition
of: shape, 42, 43, trachea in, 43—
47, size of, 47, pigmentation of,
47-49, hairs on surface of, 49,
56, effect of, on pattern, 49, re-
lation of folds to, 50; develop-
ment of, 51-59; mechanical neces-
sity for, 62; alignment of, in
wasps, 375.

independent, 68-69.
convex and concave, 70.
CLOSS, Wily MSs) Mon SO, WlaO:

Venation, of insects’ wings, origin of
60-73, 85-89; mechanical theory
of origin of, 62, 72; methods of
modification of, 73-84; types of,
85-142.

Vespa, 378, 390, 391, 392, 401, 403.
Vespidae, key to subfamilies of, 379-
380.

Viburnum tinus, host for Aphis pomi,
341.
Vigna catjang (blackeye bean), host
for Aphis medicaginis, 340.
Wasps, masarid, taxonomy and species,
369-464.

Wilson’s bonnet gall-louse (Eri-
soma?), 345.

Wing Veins of Insects, 1—152.

Wings, importance of, in classifica-
tion, 1; origin of, 4-14, 53; need
for development of, 9; relation
of, to body, 14-20; function of,
in flight, 21-30; articulation of,
30; codrdination of, 73-76; sys-
tem of folds of, 76-77; of wasps,
taxonomic characters of, 372—
377, 378. See also Venation;
Veins.

Woodworth, C. W., 1-152.

Wooly aphis (Eriosoma lanigera) , 345,
346.

Xenoborus canadensis, 237.

Xenoneuridae, Paleozoic fossil insect,
venation in wings of, 103.

Xerophloea viridis, 248.

Xiphydria, 399.

[472]

UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued)
10. Fundamental Interrelationships between Certain Soluble Salts and Soil

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Vol. 2. 1. Studies in Juglans I. Study of a New Form of Juglans Californica
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2. Studies in Juglans II. Further Observations on a New Variety of
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ellipsoideus), by S. K. Mitra. Pp. 63-102. November, 1917 ...............

6. Changes in the Chemical Composition of Grapes during Ripening, by
F. T. Bioletti, W. V. Cruess, and H. Davi. Pp. 103-130. March, 1918

7. A New Method of Extracting the Soil Solution (a Preliminary Com-
munication), by Chas. B. Lipman. Pp. 131-134. March, 1918 —-...:.....

8. The Chemical Composition of the Plant as Further Proof of the Close
Relation between Antagonism and Cell Permeability, by Dean David
Waynick. Pp. 135-242, plates 13-24. June, 1918) ...22.0021..-2 ee

9. Variability in Soils and Its Significance to Past and Future Soil In-
vestigations. I. A Statistical Study of Nitrification in Soil, by Dean
David Waynick. Pp. 243-270, 2 text figures. June, 1918 -.....2--2...

10. Does CaCO, or CaSO, Treatment Affect the Solubility of the Soil’s
Constituents?, by C. B. Lipman and W. F. Gericke. Pp. 271-282.
Bb bs (2 S00 G2 fey ea) ips a eaten SIE 2 Wi Ves ie a on eee Delle pan Eee
/11.An Investigation of the Abnormal Shedding of Young Fruits of the
Washington Navel Orange, by J. Eliot Coit and Robert W. Hodgson.

Pp. 283-368, plates 25-42, 9 text figures. April, 1919 ............-.-------.------

12. Are Soils Mapped under a Given Type Name by the Bureau of Soils
Method Closely Similar to One Another?, by Robert Larimore Pendle-

ton. Pp. 369-498, plates 43-74, 33 text figures. June, 1919 -.................

Index, pp. 499-509.

.60

10

1,00

UNIVERSITY OF CALIFORNIA PUBLICATIONS— (Continued)
Vol. 4. 1. The Fermentation Organisms of California Grapes, by W. V. Cruess.

Pp. 1-66, plates 1-2, 15 text figures. December, 1918 oo. cee “75
_ 2. Tests of Chemical Means for the Control of Weeds. Report of Progress,
by George P. Gray. Pp. 67-97, 11 text figures 2.00.20 .00

3. On the Existence of a Growth-Inhibiting Substance in the Chinese
Lemon, by H. 8S. Reed and F. F. Halma. Pp. 99-112, plates 3-6. ~ :
February; OTe ea as p BeSiRas etin: Soc ny Ne aati ot Reais RAR ee 25

4, Further Studies on the Distribution and Activities of Certain Groups of
Bacteria in California Soil Columns, by Charles B. Lipman. Pp. 113- :
POO APE ROTO 3-7. 28s ee ee ek IS ae. i Se eee 10

5. Variability in Soils and Its Significance to Past and Future Soil In-
vestigations. IT. Variations in Nitrogen and Carbon in Field Soils
and Their Relation to the Accuracy of Field Trials, by D. D. Way- *
nick and L. T, Sharp. Pp. 121-139, 1 text figure. May, 1919 -......... «20

6. The Effect of Several Types of Irrigation Water on the PH Value and
Freezing Point Depression.of Various Types of Soils, by D. R. Hoag-
land and A. W. Christie. Pp. 183-232. November, 1919 _.......-20 2 .25
7. A New and Simplified Method for the Statistical Interpretation of Bio-
metrical Data, by George A..Linhart. Pp. 159-181. September, 1920 .25

8. The Temperature Relations of Growth in Certain Parasitic Fungi, by
Howard S. Fawcett. Pp. 183-232. May, 1921 _..000 oe Ay 3)

9. The Alinement Chart Method of Preparing Tree Volume Tables, by
Donald Bruce. Pp. 233-243.° December, 1921-2202 «20

10. Equilibrium Studies with Certain Acids and Minerals and their Probable
Relation to the Decomposition of Minerals by Bacteria, by Douglas —
Wrisht,: St? Pp.245-387.WMlareh: W922 5 a ee eae $1.25

11. Studies on a Drained Marsh Soil Unproductive for Peas, by P. S.
Burvess.:, Pp. 339-396... Panes AOD Ss ee ee 65

AGRICULTURE.—The Publications of the Agricultural Experiment Station consist of Bul-
letins and Annual Reports, edited by Professor Thomas Forsyth Hunt, Director of
the Station. These are sent gratis to citizens of the State of California. For detailed —
information regarding them address The Agricultural Experiment Station, ‘Berkeley,
California, p

Other series in American Archaeology and Ethnology, Botany, Classical Philology,
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Roman Archaeology, History, Mathematics, Modern Philology, Pathology, Philosophy,
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UNIVERSITY OF CALIFORNIA CHRONICLE.—Issued quarterly. Price, $2.00 per year,
50 cents a copy. A magazine of general interest for intelligent people.

ADMINISTRATIVE BULLETINS OF THE UNIVERSITY OF CALIFORNIA.Tnciudes
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colleges.

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