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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.
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. 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:
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Amans, P. C.
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ee
4 ih!
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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. 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 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. ees eae ae
. Studies on Ammonification in Soils by Pure Gultures; by C. B. Lipman — ;
and-P: S;: Burgess. Pp. 141-172; April, 1914s. oso tae
Loughridge.. Pp. 173-274.. August, 1994 -2...-sc..ee cst celle eeceneecernennnee
. New Experiments on Alkali Soil Treatment (Preliminary Report), by RYE
Charles B. Lipman and Leslie T. Sharp. Pp. 275-290, plates 1-4.
A ft Ys Pees 2 8 eae Nn Me aN Paar aaOE cae re Glee feeeMre a setae Wim Ba, PON SS
Fundamental Interrelationships between Certain Soluble Salts and. Soil
Colloids, by L. T. Sharp. Pp. 291-339. April, 1916 .2...20.. cet
JUN 97 1916 © )
“tional Museu™
UNIVERSITY OF CALIFORNIA PUBLICATIONS
TECHNICAL BULLETINS
COLLEGE OF AGRICULTURE, AGRICULTURAL EXPERIMENT STATION
ENTOMOLOGY
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 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. 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 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— 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 :
: aaniocs PARTS OF THE STATE, BUT CHIEFLY
_ FROM THE CAMPUS OF THE UNIVERSITY 7 ae
OF CALIFORNIA, BERKELEY, ea ee oe
CALIFORNIA ; =
SE GMEssiG a = os Se ;
UNIVERSITY OF CALIFORNIA PRESS ee ee 3
"BERKELEY Pee ero Se ee ee
a =
A
= ? oa
‘i oe =
Ste ~t
a
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ENTOMOLOGY.—C. W. Woodworth, Editor.
Technical Bulletin of the College of Agriculture, Agricultural Experiment iia &
Cited as Univ. Calif. Publ. Entom.
Vol. 1. 1. The Wing Veins of Insects, by C. W. Woodworth. Pp. 1-152, 101 texk
figures. September, 1906 Pa Sa Cea s pctees 1k Sanat = teat eee Pre SNC NS So i
2. Catalogue of the Bphydridae, with Bibliography and Descriptions of
! New Species, by Burle J. Jones. Pp. 153-198. October, 1906 _...... a
3. Synoptical Keys. to the Genera of the North American Miridae, by — 7
Edward P. Van Duzee.- Py. 199-216. February, 1916 {220 sess Aa
4. New Genera and ‘Species of Orthotylini (Hemiptera), py Edward Boo See
Vans Duzees Pp. 2r7-22 72 larch, 1916. ences sa esac 2 cc raceee seca atebcengenae ee
5. Notes on Some Hemiptera taken near Lake Tahoe, California, by Bdward Ped
P; Van Duzee. . Pp. 229-249. May, 1916 conan. secetiep cent sent ct ci eceeeennnnee a
6. Life-History and Feeding Records of a Series of California Coccinelli- ~
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
>
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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. 3. { er
Ps i Me A
Si
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6. Life-History and Feeding Records of a Series of California Coccinelli- est
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versity of California, Berkeley, California, by HE. O. Essig. Pp. 301- ak
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Vol. 1. 1, The Distribution and Activities of Bacteria in Soils of the Arid =
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Transformations of Nitrogen in Soils, by W. P. Kelley. Pp. 39-49. ‘
December 1 S18 eye Oe EB ONE RT NE Ly Ge Stead
4, The Aluminum Reduction Method as Applied to the Determination of
Nitrates in ‘‘Alkali’’ Soil, by Paul S. Burgess. Pp. 51-62. May,
gC BB eR AR aR NCA Reg anton WR) Lc AVR lt Nee sy RSS
5. Studies Upon Influences Affecting the Protein Content of Wheat, by G.
W/ Shaw. Ppii63-126;-- October: 1918 0 ai i er ra
6. The Effect of Copper, Zinc, Iron and Lead Salts on Ammonification and
Nitrification in Soils, by C. B. Lipman and P. S. Burgess. Pp. 127-
EBON LARCH TOD: ek ie VRS era RS OES Ely Rat ek Ue
7. Studies on Ammonification in Soils by Pure Cultures, by C. B. Lipman
and P. S. Burgess. Pp. 141-172. April, 1914 2.0... cceccc cece cece eae i
8. Humus and. Humus-Nitrogen in California Soil Columns, by R. H.
Loughridge. Pp. 173-274. August, 1904 o-oo peice cccenccsentnenesonenenes
9. New Experiments on Alkali Soil Treatment (Preliminary Report), by
Charles B. Lipman and Leslie T. Sharp.. Pp. 275-290, plates 1-4.
WHC} TOT Pe cae NO IOS, AEP RN A AR eA OS A es a a we
10. Fundamental Interrelationships. between Certain Soluble Salts and Soil
Colloids, by L. T. Sharp. Pp. 291-339.. April, 1916 20.020.0.20ce 50
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Solution on Plants Grown in Sand Cultures, by Arthur Hugo Ayres.
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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 '
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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
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NORTH AMERICAN SPECIES.
ier PERO ag, A RIS en A a hase
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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, ¢.
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[ 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, ¢.
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