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Maddie American Mpecies 21, FA ta oe Ne le epee cess creweu ne rime 25
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The Entomological Society of America

Volume VII TUNE. rots Number 2

A STRUCTURAL STUDY OF THE CATERPILLARS:
Iii, THE SOMATIC MUSCLES.

Won. T. M. Forses, Ph. D., Worcester, Mass.

In consideration of the very few dissections of the muscular
system of Lepidoptera which have been published, and their
radically different interpretation, it has seemed advisable
to study a few more forms and trace if possible the points of
disagreement.

PREPARATION.

A serious matter in such a research is the preparation of
material. The muscles are small and slender, quite difficult
to trace without a good microscope and plenty of light, easily
broken in dissection in hardened material, and almost per-
fectly transparent when fresh. Besides this they do not
differ in color from the fat with which they are intermingled,
and when preserved in formalin hardly differ in consistency.
The most satisfactory material was opened out after killing
with cyanide, and pinned out on a piece of cork; then treated
with strong corrosive sublimate (bichloride of mercury) and
dissected while immersed in a dilute solution of the sublimate.
This makes the muscles intensely white, distinguishable from
the fat by their silky luster; but the mercury attacks dis-
secting instruments badly and so most of the work was done
with material preserved in formalin (4 per cent., that is, 10
per cent. of the commercial solution). Alcohol was also
tried, and was nearly as good; the muscles being darkened

109

110 Annals Entomological Society of America [Vol. VII,

a good deal in some cases, which made them more difficult to
see in a dim light, but brought them out in contrast with the
fat. For the work with the thorax it is important to open
up the caterpillar and pin it out before hardening, as other-
wise many of the muscles will be broken. If specimens are to
be preserved whole for dissection purposes they should be
killed in hot water, or injected with a stronger solution of
preservative to prevent decay. The prothorax and last seg-
ment are particularly. hard to get in a satisfactory condition,
because of their peculiar shape, and the close connection
of the former to the head, which should be split vertically
when opening the caterpillar. It is most convenient to open
the caterpillar near, but not quite on the middorsal line.

The viscera and loose fat of the body cavity are removed
as a preliminary, as well as the wings of the heart, leaving the
heart, trunk trachea and nervous system as long as possible
for landmarks. The dissection can follow the order given by
Lyonet, to advantage, but it is often unnecessary to open any
specimens by the venter, as the dorsal musculature is com-
paratively simple, and is often uninjured on one side. Plate
XIX follows this order fairly closely in its six stages, but in
other plates less stages are shown for economy.

It soon appeared that the muscles represented by Lyonet,
for the Goat moth caterpillar, could be found in such
widely divergent forms as a Sphinx, a Noctuid and a Lasio-
campid, so his lettering is used in the figures. Lubbock’s dis-
section was hardly as perfect, while Berlese’s figures are wholly
diagrammatic and useless for the study of homologies; in fact
several of his comparisons with the muscles of other insects are
invalid. We will only be able to come to a true knowledge of
the homologies when the nerve-muscle relations are fully worked
out; and in the Lepidoptera the matter is much confused by the
anastomoses between all three pairs of nerves, and even between
successive segments.

THORAX.

To go on to details; I have not made any satisfactory dis-
section of the prothorax. Evidently however, Lyonet represents
the state of affairs much more accurately than Berlese. The
dissection figured on Plates 1 and 2 may be taken as accurate so

1914] A Structural Study of Cater pillars. 111

far as it goes, but is incomplete. The following comparisons
may be made between Lyonet’s, Lubbock’s and Berlese’s
lettering:

LYONET. LUBBOCK. BERLESE.
er 82 CxXXxXIXa
D, E 1,4,5 139-140
A 657 140b
a 22 CXXXIX
Gee al 61, 62 CXXXIV
B, y, 5, ete 35, 76, 80, 81 CXLI
a 16, 17, 21 CXXXII
¢ 19, 20, 26 CXXXIla
c 18. CXXXIIb
r, etc 77 129
f, | ete 70, 71, ete 147
u, V 57, 58 XL

The whole arrangement is complex and only in a general
way comparable to that of the other segments. Ct is interest-
ing as extending well beyond the segment line, and in Cossus
the length of two whole segments—being the longest muscle in
the caterpillar. It also crosses the middle line in Cossus, but
not in Noctua, where it is shorter. If Lyonet is correct there
are no longitudinal muscles within the nerves, (except perhaps
the aberrant A and Ct which presumably represent the great
longitudinal muscles of the following segments); and some of
the muscles are innervated from the subcesophageal ganglion,
evidently belonging to the cervical system. The spiracle is
supplied by the “bride epiniere’’ or so called sympathetic
fibre, derived from the same segment, but it runs largely in the
following one and seem to supply also some ‘of its muscles,
besides anastomosing with its first nerve. This indicates
strongly that the spiracle originally lay on the incisure, as the
rudimentary second one actually does, and that it has moved
forward. For the same reason we see that the other spiracles
have moved back, being supplied by the nerve of the preceding
segment; and all the spiracles are accounted for.

The meso- and metathorax are perfect counterparts of each
other, and strongly contrasted with the other segments in
structure. The few points of difference between them noted
by Lyonet, are further reduced by the correction of a couple of
misinterpretations, and particularly by treating the three
bellies of his a as separate muscles. In discussing these, and
the remaining segments we may use Berlese’s division of the

Li2.- Annals Entomological Society of America [Vol. VII,

segments into four annulets, illustrated on Plate IX, Figs. 1
and 2. These are the acro-, pro-, meso- and meta-tergites and
sternites, separated by the antecosta, precosta and inter-costa.
The tergopleural suture is indicated only by the position of
the rudimentary wing, while the pleurosternal one is the
strongly marked subventral fold, to which several muscles are
attached. In the abdomen these boundaries are less distinct,
but the pleurosternal suture can be traced in a general way,
and the tergopleural must be placed at least a little higher, as
indicated by the spiracle, which should lie within the pleuron.
Of the true legs we have a well marked coxa, which is mostly
membranous in the Sphinx, a strong femur, to which only one
or two small somatic muscles are attached, and the rudiment
of a trochanter, which bears the insertion of r. The insertion
of the various muscles of the leg is beautifully shown by Lyonet,
in Plate VIII, Fig. 7. In Cossus where the coxa is wholly
chitinized k, n, p, s, t, u, V, xX, €, 0, .K, Ay B, P and] tune it.
The body muscles may be divided as to their origin in the
embryo into the dorsal and ventral longitudinal systems, and
the lateral and ventral transverse ones; the last is not repre-
sented in the caterpillar by a developed muscle, but its rudiment
may exist associated with the fork of the sympathetic nerve
and the ventral diaphragm. This would be 4 of Berlese. The
lateral transverse muscles are again divided into two sets,
between which lies couple of longitudinal fibres (E of the
abdomen) and the trunk trachea, but as the trunk trachea is
not well developed in the thorax, and replaced by collaterals in
the body cavity, the muscles must there be treated as a single
unit. If we consider C, E and G as homologues of E, etc., of
the abdomen @ will be the only fibre of the deep set in the
thorax, as it is in the abdomen. It should be noted that the
spiracles can migrate without disturbing these fibres, while the
deep transverse fibres would have to be swept before them.
The muscles also differ from each other in their normal or oblique
direction, their length and their insertion. The table annexed
classifies them, and gives the relations between meso- and
metathorax, and between the various nomenclatures, omitting
those muscles which puzzle me.

1914] A Structural Study of Caterpillars. 113

LYONET LUBBOCK BERLESE
Meso-. Meta-. Meso-. Meta-.
LONGITUDINAL DORSAL. 4

Rectus (1) A Atos 1 7 37
B B 1 70 37
Oblique segmental C Cc 8 70b .37b
; E E 8’ 37c
D D 5 70a 37a
F F 5 70a 37a
Grr H 4
a (ist belly) G 6, 7
Shorter segmental I ii
To intercosta K K 61
From precosta k L, M, L, 9-11
To precosta S S 62
dy T 63
From intercosta 6) QO 12, 13
R R 14, 15
To antecosta d (part) X (part) 64, 65 (part)
INTRANEURAL (2) c c 18 ©
LONGITUDINAL VENTRAL, SUPRANEURAL
Segmental a a 21 LXXII XXX
b b 7 “ “
Bs d (16 ) « “
e,f € 22 -
d f EX NAP OORe
g g 16 (19) s =
h h “ “
To mid-ventral line behind
cox 1 1 26 +
LONGITUDINAL VENTRAL, SUBNEURAL
To mid-line in front of coxze i i IVb (3)
Crossed to front of coxa k k 24 IVa (8)
Short anterior transverse (4) Z Z 25 ree 7 (3)
From mesosternite at mid-
ventral line n n 28 54
From precosta m a (3rd belly) 77
Short fibres from leg q q 19
Vv v 58 Pee
TRANSVERSE; TO PLEUROSTERNAL LINE
Antecosta to prosternite 8 B 49
Precosta to precosta, super-
ficial 6 6 49
Precosta to prosternite, deep v v
Precosta to mesosternite Be be 72 XVIIIa (3)
Antecosta to posterior in-
cisure € Y 66 XXXIVa
Antecosta to mesosternite ny ny ‘os XVII (3)
(posterior bellies) 65 (part)
Intercosta to mesosternite Kk, & x, & 50 XVIIIa (3)
Anterior incisure to precosta vy a (2d belly) 76
To anterior incisure n n XXXVIb

w Ww 41, 42

114 - Annals Entomological Society of America [Vol. VII,

LYONET LUBBOCK BERLESE
Meso-. Meta-. Meso-. Meta-.

Across PLEUROSTERNAL LINE ;

Intercostal region to leg 6,3 e, £, 30 46, 51 5a (3) 5

Posterior incisure to leg t t 55 XVIIB (3)

Superficial on incisure ] ] 35 XXXVI XVIla

Deep on incisure x x 34,38 XXXVIa XVIIIa
BELOW PLEUROSTERNAL LINE

To leg s s 78 - Ila, 6a

To near midventral line p p 73-75
SPIRACULAR (rudiment) T

Notes: (1) Muscles droits of Lyonet.
(2) Passing between the two longitudinal connectives.
(3) Fig. 478.
(4) Certainly derived from longitudinal muscles like k. In the meso-
thorax it tends to cross the middle line.
+ Indicates that the muscle is represented but not named.

The wing is shown in the deepest layer of the dissection on
Plate XVIII, in its normal position. The only fibres closely
associated with it seem to belong to w and X, practically all the
pleural muscles being inserted far above it. Evidently the
pleurites are very slightly developed in the caterpillar stage.

The mesothorax seems to differ from the metathorax as
figured only in having a fibre or two more or less in the case
of such homologous muscles as a, b, (d); 6; Q. R. S and T.
The union between thorax and abdomen is made with only a
single disturbance of the musculature, aside from the fact that
the ante- and precosta of the first segment of the abdomen
are undeveloped; and that is that one head each of E and F
have moved forward a short distance beyond the incisure,
carrying with them a couple of fibres of @, and attaching
themselves to the insertions of G and t respectively.

MIDDLE SEGMENTS OF ABDOMEN.

One of the middle segments may be taken as typical of the
abdomen. Examples of three families are figured on Plates
XX, XXI and XXII. As compared with the thorax the most
striking difference is the weak development of the oblique lateral
muscles, and the absence of muscles that cross the median
line; both conditions correlated with the simpler movements
of this part of the body. Here also there are no such confusing
cases as a of the thorax, where several morphologically widely
separated muscles form a single functional unit. There is no

1914] A Structural Study of Caterpillars. 1

fibre piercing the nervous system, and no short dorsal anterior
muscles, though it was probably such a muscle ventrally, that
gave rise to p’, which is wholly independent of p and x, though
not well marked in Cossus and Malacosoma. In the abdomen
the principal nerve divides the ventral muscles quite centrally,
forming the most satisfactory distinguishing character between
the fibres f and g, and forming the most fundamental dis-
tinction between a and the other recti; the relation of the
dorsal muscles is more obscure, as the nerve divides up, but
A, B and C evidently lie above, and E, F and H below it.
The transverse muscles are divided into two groups as already
noted in the case of the thorax. The following classification
of the muscles gives their designation by Lyonet, Lubbock
and Berlese.

LYONET LUBBOCK BERLESE
LONGITUDINAL DORSAL
Rectus A 1 VII
B 3 “
cS 2 “
Segmental D 4 :
G 5 . “
From antecosta, oblique F(4) 8 x
H 7 “
From antecosta to antecosta E 6 IX
From precosta I 9 XI
; L 10, 11 .
From intercosta Q,R 12; 15 XVI
LONGITUDINAL VENTRAL, SUPRANEURAL.
Rectus c 18 2
b 17 ‘
d(3) 16 .
Antecostal, oblique f’ (in Malacosoma only)
Longitudinal segmental e
ff 19 z
To antecosta of following segment f 25 IV
From antecosta, oblique h a
LONGITUDINAL VENTRAL, SUBNEURAL.
Antecosta to antecosta, longitudinal a 21 III
i 22 ‘i
To antecosta of following segment,
oblique g 24 IV
Segmental, oblique g’ 25 2
From front of leg k 27 1 aa
From near midventral line, between legs p 28-30 la
From posterior side of leg t 57 V
From leg toward mid-ventral line x 56, 58 laB (1)
From near mid-ventral line to leg (p’) 46 (part) lag
SPIRACULAR M 45

TRANSVERSE SUPRATRACHEAL.
Acrosternal 0 35 XVII
At precosta 6(2) 36 3)

£16 - Annals Entomological Society of America [Vol. VII,

LYONET LUBBOCK BERLESE
TRANSVERSE SUBTRACHEAL, Above subventral fold.
From near incisure to precosta a 37, 38 XVIII
Mesotergite to mesosternite 6, € 51, 52 XVIIIa
Crossing the pleurosternal line.
Spiracle to precosta 1 39 5b
Propleurite to precosta m(sometimes) 42 5b
q(rarely) 40, 41
Mesotergite to parts of leg B, ¥ 46-50 5a, 56
Short posterior muscles Cay 33-34 XVIII
Below the pleurosternal line.
Behind precosta q, m 40,41(42) 5Sba
Across leg in front n 43 ba
Across leg behind (from wy) I 53-54 68
From fold or y to leg (part) baa
From fold or y to anterior edge of
following segment ¢ (part) 31-32
Zi 55 XVIIIB?

Notes: (1) la8 in most forms, not distinct from V in Cossus, I on legless segments
(2) The fibre of 6 which runs between the precostae is marked 4 on the
plates; it is not well defined in Cossus or Sphecodina. Fibres of
@ proper spread fan-like from the acrosternite to various points in
the acro- and protergites.
(3) To antecosta of following segment in Pheocyma.
(4) From acrosternite in Pheocyma.

The muscles which attach to the ends of the segments are
sharply divided into two groups, in the first, comprised of
A, B, C, E, a, b, c, d, the muscles of successive segments are
united, forming a single polygastric muscle, the remainder
are so inserted to leave a distinct space between muscles of
successive segments. In the case of D and i, the distinction
is striking, as compared with A and a respectively.

NERVES.

There are three pair of nerves from each abdominal ganglion.
The principal or anterior one runs almost directly out from the
anterior half of the ganglion, passing over all the muscles near
the midventral line, but soon plunging in (between a and c) and
running between the layers up to the subdorsal region, where it
ends in a longitudinal fibre, perhaps the chordotonal organ.
It supplies but little of the skin, but sends off numerous branches
to the muscles, especially the larger segmental ones. The
second or posterior pair of nerves runs obliquely downward
under i, passes between k and p, often forming the only dis-
tinction between these two muscles, and then ramifies in the
crowd of short muscles connected with the proleg, and on the
skin. The third runs directly back as a single nerve from the
posterior end of the ganglion under the fused connectives,

6

1914] A Structural Study of Caterpillars. 117

until they separate, then forks, in the substance of the very
slightly developed ventral diaphragm and runs out, often
along the incisure, to the spiracle and from there to the tip of
the wing of the heart; no muscles overlie its main stem, but
it sends down branches to anastomose with the second nerve,
and connects with the anterior one in the neighborhood of the
spiracle. It certainly supplies the spiracle and probably some
muscles, but on account of the anastomoses it is impossible to
be sure. Sometimes the fusion of the connectives is complete,
as in Sphecodina (Plate 5) and this nerve seems to arise from
the-anterior end of the ganglion after the one to which it
belongs. In the thorax tts connections are always perfectly
clear, because of the wide separation of the connectives.

On the third to sixth segments the structure is identical, as
_ described, except for purely individual variation, but the others
show various stages of reduction.

OTHER SEGMENTS OF THE ABDOMEN.

The second segment differs only in the reduction of the
muscles of the proleg, and the loss of a couple of fibres, but
they do not change their points of insertion. x is insignificant
or absent; k and p are usually unchanged, but may appear
as short parallel oblique fibres evidently homologous to I and
L dorsally. This condition appeared in an odd specimen
of Malacosoma disstria. vy, m, n, q and r are reduced in
strength, but not notably changed in points of insertion.

In the first segment the proleg is so reduced as to be unrecog-
nizeable, in a stage corresponding to Berlese’s Fig. 477 D,
which was evidently prepared from the seventh segment.
The fat pad, which causes the proleg of the second segment
to keep nearly its normal relations, is unimportant here, and
the transverse fibres (of the 1, q group) are shortened. 8,
(representing y) is merely a couple of fine fibres which barely
cross the fold. The antecosta dorsally and precosta ventrally
are also not developed and the muscles which normally end
in them are continued to the front of the segment; these are
a, i, E and f’ when present. As the trunk trachea disappears at
the first spiracle a is not distinguished from 6, whose fibres
are crossed as a result of the moving forward of E and F.
H, however, may remain unchanged, and the spiracle is only

118 Annals Entomological Society of America [Vol. VII,

a little in front of its usual position. The innermost fibres
of c nearly meet in the middle line, covering the origin of a.

Toward the posterior end reduction is progressive to the
last segment, and becomes extreme in the ninth, which however,
is unmistakably a true segment. The fibres gradually go
over to the rectus type, becoming simple and longitudinal,
and gradually decrease in number, till in the ninth segment
only five ventral ones are left.

The ganglia are fused in the seventh segment more or less
completely, but always the true seventh ganglion is recognizable
with its usual nerves,—,the sympathetic nerves from the last
spiracle run back united with the nerve for the eighth, as far
as the incisure, then run normally to the spiracle and last
wings of the heart. All the principal muscles are present,
but a, f, g, h, i and E do not extend beyond the eighth acroster-
nite, because of the disappearance of the pre and ante-costa
of the eighth segment. The deep muscles are reduced, p and
x appear clearly as longitudinal (oblique) muscles similar in
character to I, Land Q, R of the dorsal region. ¢ also appears
longitudinal, a suggestive fact, as in general it resembles more
closely the fanlike transverse muscles, but z is evidently trans-
verse. ais normal, m and q are undifferentiated, and clearly
pleurosternal; d is normal, n and r extraordinarily shortened,
while 8 (representing y) and 6 are also nearly simple—
evidently the proleg is merely the small area enclosed by
n and r.

The eighth segment has the two normal nerves running back
from its ganglion in the preceding segment. The first runs
outside c, which apparently represents part of a of other seg-
ments; it passes between f and g, defining them as usual.
Muscles a toi and the corresponding dorsal ones are all reduced
to the simple segmental type, M and 1, being connected with
the spiracle, remain normal, but @ is simplified in front and
its homologue behind is much reduced. The lateral muscles
are perfectly simple, and the longitudinal ventral ones are
reduced to a single set of sort fibres (p).

In the ninth segment with the disappearance of heart and
spiracle the sympathetic is also gone, but the two principal
nerves are easily found, (sometimes with the first arising
behind the second). The first runs between a and b, which
latter has to serve for b, c, d, e, f, ff, and h of normal segments;

1914] A Structural Study of Caterpillars. 119

e probably represents normal g, while d would be a transverse
fibre. The odd fibre e’ may be the last trace of the proleg.
The posterior nerve, with the disappearance of the proleg,
runs mainly to the skin.

The last segment is specialized to such an extent that an
embryological study would be necessary to straighten it out.
It doubtless is a fusion of at least two. Its one large nerve
runs mainly to the proleg and on its way serves as a stalk
for the nerves of the preceding joint.

COMPARATIVE ANATOMY.

Including this paper I can find only five widely separated
families represented by dissections of the muscular system,
namely:

CossID#; Cossus cossus by Lyonet.

NOTODONTID2; Pygera bucephala by Lubbock.

LASIOCAMPID=; Malacosoma americana and disstria in this
paper.

NoctTuDIpD#; Two species, probably of Noctua and Nephe-
lodes, both trifide and a Pheocyma (quadrifide) in this paper.

SPHINGID#; Delilephila lineata, briefly by Berlese; and
Sphecodina abbotit in this paper.

These few species indicate, however, that the characters
of the muscles are likely to be as well marked as of any other
part; the following points stand out most strikingly:

In the Sphingide the muscles, especially the large ones,
tend to be broken up secondarily into a considerable number
of fibres. This shows strikingly in the recti. Q and R show
a variety of lengths and tend to run to at least two secondary
annulets; x and p cross the middle line and interlace with each
other. Well marked resemblances to Cossus appear in the
broad convergent and overlapping e and ff, in the filling of the
body cavity with trachez, and the slight differentiation of 4.
The thoracic muscles are particularly massive; the longitudinal
connectives are fused unusually far in the thorax, and com-
pletely in the abdomen, throwing the apparent origin of the
sympathetic fibres back to the ganglion of the next segment.

The Cossid@e show their primitive character in the well
separated ganglia in the seventh segment, the slight differentia-
tion of 4, the fact that x is nearly longitudinal and hardly
distinct from t, even in the segments with prolegs, and the well
developed ninth segment of the abdomen.

120 Annals Entomological Society of America [Vol. VII,

The Lasiocampide, like the Sphingide, show a tendency
to increase of muscles, but it is slight. The deeper transverse
fibres alone become numerous and unstable; they are weak
and altogether much as one would expect in a form which had
passed through a lappet-bearing stage and was degenerating.
It is not unlikely that such a larva as Epicnaptera will show
a strong and highly specialized lateral system. The differences
between the two species is slight, in the first segment disstria
has only three, americana usually five ventral rectus muscles;
the lappets are much distincter in dissiria, but not really
functional, so that the difference is not noticeably reflected in
the muscles. As a whole the genus is characterized by the
massive upper fibre of [, a suggestive character in a lappet-
bearing family, and the frequent presence of the aberrant
fibre f’ (as figured) in both species. In the eighth segment
there is a large tracheal tuft, which may serve as a sort of
lung to aerate the blood where it first enters the heart.

The Noctuide are marked by their simple and normal
condition, without the primitive points of Cossus or the special-
izations of the others, and are closely similar to each other.
p and x meet on the middle line, but do not cross, as in Cossus.
In general they are much like Cossus, but possess 4, and p’.
The tracheze are reduced and inconspicuous. Of the two
species the Noctua had slenderer lighter muscles: The insertion
of p along the midventral line is a little different in character,
and as it leaves its trace on the skin may prove a help in identi-
fying in this difficult group.

A Catocaline caterpillar, apparently a Pheocyma, shows
a number of interesting specializations, apparently connected
with its extraordinary jumping power. The thorax (Plates
XVII and XVIII) is normal in general plan, but with various
oblique muscles joined end for end into long sets; four pair of
these, namely, E, a1, a2, and a3; Ct, L and G; a, C, S and L; and
c, f and C extend the whole length of the thorax and there are
several other shorter sets, while only the a system of the
meso- and metathorax was recognized by Lyonet in Cossus.
The peculiar breaking up of 5S in the metathorax at least,
caused apparently by the attachment of I through it, does
not even occur in other Noctuids, about the mesothorax I am
uncertain, as that part of my specimen was damaged in dis-
section. The principal peculiarities of the abdomen, aside

1914] _A Structural Study of Caterpillars. 121

from those resulting directly from the slender body, are the
setension of F to the acrosternite, under a and ¢, the extension
of d well into the following segment outside the other ventral
muscles and the simplified leg-muscles of A3, correlated with
the much reduced proleg; the most noticeable point in this is
that while not forming a longitudinal series of transverse
fibres as they do in the seventh segment, muscles m and q are
represented by a series of several evidently homologous fibres.
The nervous system also shows the highest specialization I
have seen in a caterpillar; as the first abdominal ganglion
is moved forward well into the thorax (Pl. 1, Fig. 1) and the
fused last ganglia have also moved forward into the preceding
segment and nearly fused with the sixth abdominal. Their
nerves, however, are normal, except. for the oblique direction.

Lubbock has notes on a few other species in his paper,
among them a Pieris.

SUMMARY.

1. Corrosive sublimate produces the most perfect material
for dissection of the muscles; four per cent. formalin is more
generally satisfactory.

2. The caterpillars should be opened out before hardening.

3. Lyonet’s work has proved fully satisfactory, and
Lubbock’s sufficiently so for almost perfect correlation of
the two.

4. The meso- and metathorax are alike, the first eight
segments of the abdomen much alike, and capable of general
correlation with the thorax.

5. The ninth abdominal segment is much reduced, but an
unquestionable segment. It is less reduced in the most gen-
eralized form.

6. The first and last segments are of a different type, and
probably compound in nature.

7. The first spiracle has moved forward, the second
has become rudimentary in situ and the other eight have
moved back, but the first rather less than the others. All
have the same innervation, in the abdomen largely from the
preceding segment.

8. The meso- and metathorax alone have muscles traversing
the nerve-cord.

— a

122 Annals Entomological Society of America [Vol. VII,

9. The anterior nerve trunk divides the longitudinal
muscles into superficial and deep sets, and the longitudinal
trachea similarly divides the transverse ones.

10. The antecosta and ventral precosta arise as speciali-
zations of the acrosternal insertions of the muscles and exist
only in specialized segments. The muscles may exist in their
absence.

11. Characteristics are pointed out of members of five
families and characteristic figures are published of a Noctuid, a
Lasiocampid and a Sphingid.

12. The following muscles are differently lettered by
Lyonet in the meso- and metathorax.

MESOTHORAX METATHORAX | | MESOTHORAX METATHORAX
Gu H only | m 3d belly of a
Ist belly of a G Y 2nd belly of a
e,f e only | € Y
d , g €, & e

It is suggested that his lettering of the metathorax be
adopted in general, but that m be used for the posterior of the
three muscles he confused as a.

13. The muscle marked E by Lyonet, and IX by Berlese
is only a segment in length, both terminations being on the
antecosta. 7

14. Cases occur of wide shifts in the origin and insertion of
muscles, both relative and absolute, notably in the primitively
longitudinal ones p, p’, and x, near the midventral line and in
d and F of Phaeocyma; they should therefore be used with
a good deal of caution in identifying sclerites or parts of the
body, at least where as in the caterpillars the sclerites are
obsolescent.

BIBLIOGRAPHY.

The following items refer to the Lepidopterous larve, Bibliography of the
other insects may be found in Berlese, 1. c. I, 460.

LYONET, PIERRE: Traité Anatomique de la Chenille qui Ronge le Bois de Saule.
La Haye, 1760.

A complete monograph of caterpillar structure based on Cossus cossus, (family
Cosside) with numerous engraved plates, illustrating, among other things,
the entire muscular system and its nerve-supply.

Lubbock, J.: Arrangement of cutaneous muscles of the larva of Pygaera bucephala.
London, 1858.
Family Notodontide.
BERLESE, ANTONIO: Gli Insetti; Milano, 1909.

A long chapter on insect muscles and their homologies, with figures of Deilephila

(Sphingide) and numerous non-Lepidopterous insects.

1914] A Structural Study of Caterpillars. 123

EXPLANATION OF FIGURES.

Plates XVII and XVIII—Dissection of the thoracic muscles of Phaeocyma
sp., following Lyonet’s lettering, and nearly following his order of dissection.
The muscles not hatched, which appear in the various layers are those which
appear more distinctly in succeeding ones. The dorsum of the mesothorax, and the
prothorax, especially in front are incomplete.

oes. suboesophageal ganglion.

gang. T!. First thoracic ganglion.

gang. T?. Second thoracic ganglion.

gang. Al. First abdominal ganglion. The ganglion of the metathorax lies

near the letter c of that segment, under the muscles c.

tr. Functional trunk-trachea of the thorax.

sp. Al. First abdominal spiracle.

D, A, B and E at the lower edge of Fig. 1, and E on Fig. 2, belong to abdominal

muscles, the latter extending forward well into the thorax.

Rudimentary crossed muscles are shown in Fig. 1, overlying the prothoracic

ganglion.

In Fig. 6 the stippled areas represent the position of the wings; they underlie

all the muscles except the long fibres of T and w.

sp. in Fig. 6 is the rudimentary second thoracic spiracle, in Figs. 4 and 5

mw indicates its trachea.

In this and most of the other figures the musculature is shown as it appears
when the caterpillar is opened and spread out, the muscles nearest the body cavity
appearing superficial, and the last layer of dissection representing the ones close
to the skin. In this figure the head was cut in several pieces to enable the pro-
thorax to be spread out flat, and the broken lines represent the mid-dorsal and
midventral lines, marked by the position of the dorsal vessel and nerve cord.
The small muscles of the legs are not shown.

Plate XIX.—Metathorax of Noctua baja (?), dissected in six stages, following
approximately Lyonet’s directions and using his lettering.

gang. Ganglion.

sy. Fork of sympathetic nerve.

ma. Rudiment of middle thoracic spiracle and its trachea.

im. Rudiment of an imaginal muscle.

The wing is shown in the deepest dissection and coarsely dotted.

Plate XX—The middle abdominal segments of Nephelodes minians (?)
similarly dissected in six stages, in the following order:
4, 3,
5, 6.
Plate XXI—Middle segments of Sphecodina abbotii, dissected in the same
order, in four layers.

tr. Trachea.

Plate XXII—Middle segments of Malacosoma americana dissected in four
layers in the following order:
yp an
oy 4.
d. v. Dorsal vessel or heart.
w. h. wings of the heart.
sp. Spiracle.
tr. trachea.
gang. Ganglion.
sy. Fork of sympathetic nerve (belonging to preceding segment).
The position of the hooks of the proleg is indicated by a series of bars.
5 is a small muscle in the planta.

124 Annals Entomological Society of America | [Vol. VII,

Plate XXIII—First two layers of muscles of the first two segments of the
abdomen of Malacosoma americana, the first layer on the right.

In all the preceding dissections the caterpillar is represented as if opened on
the dorsal line and spread out flat, so that the mid-dorsum is divided between
the two edges and the midventral line lies in the middle.

Plate XXIV—Ventral musclature of the posterior segments of Sphecodina
abbotii. The left side is shown as seen from the inner side. 1, as seen when opened
and spread out; 2, after removing the muscles hatched in Fig. 1; and 3, similarly
after removing those hatched in Fig. 2. The principal nerves are.shown as wavy
black lines; the segments are numbered and their boundaries indicated in Fig. 1
by arrows. :

Plate XXV—Diagrams of segment.

Fig. 1. An abdominal segment (left half) spread out and with the principal
annulets and regions labelled, modified after Berlese.

ACRO. acrotergite.

PRO. protergite.

MESO. mesotergite.

META. metatergite.

The four sternites are similarly placed.

sp. spiracle.

ap. tergostern. tergosternal apodeme, pleurosternal suture, or subventral
fold.

ante. antecosta (ventral end).

pre. precosta.

inter- intercosta.

pl. planta of proleg.

gy, w. Minor apodemes.

w. ringlike folds, considered by Berlese to mark rudimentary segments of
the leg.

Fig. 2. A similar diagram of the metathorax, as developed, e. g. in Sphecodina.

w. Wing-bud, marking the upper boundary of the pleurites.

pleur. pleurite, its boundaries indicated by arrowheads.

m. cx. membranous part of coxa.

ch. cx. chitinized part of coxa.

ACRO, PRO, MESO. and META.., as in Fig. 1.

Fig. 3. A superposition diagram of the muscles of the metathorax as seen
from within. Each muscle is represented by a double line, interrupted
where it passes under another, and the insertions are shown as heavy
black bars or dots. The diagram is meant to show the relative position,
insertion and direction of action of the muscles, but’not their size or
form, and is supplementary to Pl. XIX. The rudimentary spiracle and
trachea is indicated to the right of the letter 6. Middorsum at left
edge, mid-venter at right. c and k, which cross far beyond the middle
line, are only partly shown. :

Fig. 4. A similar diagram of a middle abdominal segment, serving as an
index to Plates 4-6. Dorsally the precosta and intercosta are shown,
laterally the antecosta, and ventrally the precosta, pleurosternal suture,
apodeme ¢ (running to the left of the letter ¢) and the oval indicating
the boundary of the proleg. The ganglion is dotted, and the anterior
nerve indicated as a waved line.

VOL. VII, PLATE XVII.

ANNALS E. S.A.

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VOL. VII, Plate XXII.

ANNALS E.S. A. VoL. VII, PLATE XXIII.

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ANNALS E. S. A.

VoL. VII, Plate Xxty.

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is

OBSERVATIONS ON THE LIFE HISTORY AND HABITS
OF HYDROMYZA CONFLUENS LOEW., (DIPTERA).*

By Paut S. WELcH.

During the past three summers the writer carried on some
biological investigations in the vicinity of Douglas Lake, Mich.,
in the extreme northern end of the southern peninsula of that
state. The work was done in connection with the University
of Michigan Biological Station, the facilities of which aided
materially in securing the data which form the basis of this
paper. é

Hydromyza,. one of the several small genera belonging to
the Cordyluride, has only one species (H. confluens Loew)
reported for North America. This species is northern in its
distribution and seems to have been reported only from New
Jersey (Johnson, ’04, p. 162) and Michigan (Needham, ’08,
p. 270). Needham reported it from Walnut Lake, in south-
eastern Michigan, and predicted that it would probably be
found common about Nymphaea beds in the United States.
In looking over the literature relating to this insect the writer
was surprised to discover how little has been written about it.
Aside from a brief, two-page paper by Needham (’08, pp 270-
271), nothing seems to have been written on the habits or life
history of this very interesting insect. It occurred in sufficient
numbers to make possible the accumulation of a number of
interesting facts relating to this species. The data presented
in Needham’s brief paper were tested and found to agree
with the observations of the writer in almost every respect.
Data merely mentioned by Needham have been worked out
in more detail and a number of new observations were made.
Unfortunately the writer has been unable to observe the
method of oviposition and the younger larval stages have
not been studied.

THE LARVA.

Food Plant.—Hydromyza confluens was found in con-
nection with the yellow waterlily, Nymphaea americana,
(Provancher) Miller & Standley. This is the form which has
until recently been included under the name JN. advena or N.

* Contribution from the University of Michigan Biological Station, No. 21.

135

136 Annals Entomological Society of America [Vol. VII,

inci variegata, but Miller and Standley ('12, p. 78), have

shown that the boreal species has several distinct characters
which separate it from the true advena, and they have raised
Provancher’s name, americana, to indicate this northern
species. V. americana occurs in some abundance in the protected
bays, in the beach pools, in the mouths of some of the streams,
and in sphagnum bogs of the Douglas Lake region and the
insects under consideration were correspondingly abundant.
Needham (’08, p 270), reported this insect in connection with
Nymphaea advena, but judging from his description and figures
it seems very probable that the species in Walnut Lake was
N. americana, rather than the true advena.

A careful examination of all the aquatic plants occurring
in the immediate vicinity of the Nymphaea beds was made
with the view of determining whether or not other plants were
used as food. In no case did the larva occur on plants other
than N. americana. The white waterlilies (Castalia odorata)
were entirely exempt in spite of the fact that white and yellow
waterlilies mingled in the same beds. In every case the evi-
dence pointed to the conclusion that this insect is restricted to
N. americana in the region studied.

The Gall.—The immature stages of this insect were found
in the long, constantly submerged petioles of the yellow water-
lily. A large number of plants were examined and in no case
did they occur on any part of the plant other than the petiole.

Furthermore they were confined exclusively to certain petioles, |

namely, those of the floating leaves. Special effort was made
to determine whether or not larve ever occurred on the
peduncles or on the petioles of the submerged leaves. Both
were found to be entirely free from infestation. Although
the writer has no data on the method of oviposition, the reason
for this distribution on the plant seems apparent. Needham (’08,
p. 270) suggested that ‘‘ Probably the attack of the gall maker
begins when the first leaves reach the surface in late spring;
then they have their first opportunity to reach the proper
place of oviposition by crawling down the stalk.’”’ The writer
regards this as the most feasible explanation since it will be
shown later in this paper that it is possible for the adult insect
to pass under water by crawling on a supporting surface and
that it actually does so of its own volition. The short and
wholly submerged leaves of N. americana do not at any time

1914] Life History of Hydromyza Confluens. 137

reach the surface and since this is true there would be no
opportunity for the female to deposit the eggs, hence the
constant freedom from infestation. Furthermore, in the region
of Douglas Lake, the flowers do not reach the surface of the
water until in late July and in August, a date which is much
later than the time of oviposition, and as a consequence they are
exempt from the attacks of the insect.

The gall first becomes perceptible on the petiole as a slight,
ovoid enlargement and can be detected by pulling the petiole
between the fingers. Each gall is produced by one larva only
and in no case is more than one larva found in a single gall.
It is very probable that in the early stages the presence of the
larva is not perceptible since this enlargement really occurs
at a time when the larva is well toward maturity. As the
time of pupation approaches the gall begins to turn brown,
ultimately assuming a deep brown color, thus making it easy to
detect. The shape of the exterior of the mature gall varies
somewhat, usually appearing as an ovoid swelling, although
it should be noted here that many of the galls do not increase
the diameter of the petiole. The galls also vary in size to
some extent, the length of those containing pupz ranging
from 6 to 9 mm. They are always longer than broad, the
long dimension being in the direction of the long axis of the
petiole.

The number of galls per petiole varies rather widely. Some-
times only one gall occurs on a petiole, but usually the number
is greater. In a large series of observations the maximum
number on a single petiole was found to be 14 and many of
the petioles contained as high as 13. In one of the series of
counts in which 45 infested petioles were examined, 7 was
found to be the average number of galls per petiole. No
relation was found to exist between the length of the petiole
and the number of galls. It might be assumed that the longer
petioles contain the larger number of galls, but this was not
always true. Numerous instances were observed in which a
petiole, six feet in length contained only two or three galls,
while a neighboring petiole, three feet in length, contained
10-12.

Galls occur irregularly along the petiole and may be situated
near the bottom, even in those almost six feet long. They may
be distributed along the entire length of the petiole and well

138 Annals Entomological Society of America [Vol. VII,

separated from each other, or they may be distributed in
such a way that some are widely separated and others so
closely placed that two or three may appear to be continuous.
The interior of galls containing pupz, or half to full-grown
larve, can be easily examined by removing the infested petiole
from the water and holding it between the eye and the light.
The cavities are not at all uniform in shape. Each has a

slightly elongated central chamber and one or more side chan-

nels which are usually just large enough to contain the larva.
The total space in the gall of a full grown larva is commonly
three or four times as large as the larva itself. The size of
the cavity represents the bulk of the food which the larva has
consumed during its growth and since there is no connection
with the exterior all of the excrement is deposited within the
gall. The consumed maiter, as indicated by the amount of
excrement, does not seem to decrease much in bulk so that
near the time of pupation the greater part of the gall is filled
with a brown excreta which almost surrounds the insect. This
brown excrement has much to do with the brown exterior
which characterises these excrescences and renders them
conspicuous.

The full-grown Larva.—The morphological details of the full-
grown larva will be included in a future paper which is in
preparation. Needham’s paper contains a very brief descrip-
tion of the larva and presents some of the more important
anatomical details.

Variation in Maturity—An examination of the galls on a
given petiole often showed differences in the degree of maturity
and when the larve on such a petiole were extracted and ex-
amined it was found that they too were not all exactly of
the same degree of maturity. In some cases part of the number
had pupated while others on the same petiole were still in the
active larval stage. It thus appears that some of the larve
lag behind the others in development and the question arises
as to how this fact is to be interpreted. It is conceivable
that this might result from the deposition of eggs at different
times by different females, but the writer, after examining a
large number of such cases, is skeptical of such a possible
interpretation and believes that in the majority of cases at
least, all of the galls of a given petiole are the result of eggs
laid at one time since, (1) although some of the larve lagged

1914] Life History of Hydromyza Confluens. 139

behind others in development, the differences did not seem to
be sufficiently marked to warrant the conclusion that the
eggs were deposited at different times, and (2) in some cases
those lagging behind occurred in the middle of the petiole,
while those towards the upper and lower extremities of the
petiole developed at the same time, a condition which would
not be likely to happen in case the eggs were laid by different
females. The above evidence is not entirely conclusive but
it seems to support the more feasible explanation, namely,
that the eggs on a petiole are all laid at one time, and that the
variation in the degree of development is possibly due to
internal or external factors, probably the former since the
external conditions seem perfectly uniform for all those deposited
on a given petiole.

Activity—Like many other dipterous larve, this gall
maker is very sluggish. Specimens, removed from galls,
showed only slow, squirming movements which were very
ineffective so far as locomotion was concerned. ‘The activities
of these larve could be observed to some extent by holding an
infested petiole between the eye and the source of light. The
feeding activities appeared to be very deliberate.

Effect on the food plant—As mentioned above, the first
evidence of infestation is the presence of a slight, ovoid swelling
at various places along the petiole. Later these swellings
begin to develop a brownish color, ultimately becoming a deep
brown. About this time or a little later the entire petiole
loses its green color, takes on a yellow appearance, and shortly
afterwards the entire leaf begins to turn yellow, showing signs
of deterioration. During the last week in July and the first
week in August of 1913 one could readily pick out the infested
lilies by the yellowish leaves. This work of the larve leads
to the decay of both leaf and petiole. The nature of the
injury is simple. The larve affect the plant in two different
ways: (1) The larva, as it eats out the internal cavity of the
gall, severs the vessels which connect the leaf with the root-
stalk, this alone being sufficient cause for the decay of the
leaves. It was found that the heavily infested petioles deter-
iorated no more rapidly than did the slightly infested ones
and that one larva is just as efficient as several in causing the
death of the leaf. (2) These galls produce weak spots in the
petioles so that wave action breaks them at the points of

140 Annals Entomological Society of America [Vol. VII,

infestation. Leaves with broken petioles were found floating
about early in the season before the galls had begun to turn
yellow, but the greatest havoc from this cause was produced
during the first week of August of the past year when the
larve were pupating. At this time in one of the badly infested
lily beds approximately 40% of the leaves were broken off
and were floating about in a semi-decayed condition.

The possibility of other insects playing a part in causing
this deterioration of the petioles and the change in the color
of the leaves was taken into account and, while other enemies
were present, it was possible to observe many leaves and
petioles which were infested only by the larva of H. confluens
and which showed the same effects as those which happened to
be affected by more than one enemy.

All of the lily beds in the immediate vicinity of Douglas
Lake were examined and the degree of infestation observed.
There was considerable variation in this respect since some were
only slightly infested while in others the percentage of infesta-
tion was as high as 50 or 60. None were entirely exempt.
It was found that the heaviest infestation occurred in a lily
bed which was located at the end of a point which formed one
side of a protected bay and was thus exposed to the wind and
wave action to a greater extent than any of the other lily beds.
In this particular case the infestation was almost twice as great
in this exposed lily bed as in another in the protected bay only
about one hundred feet away. The infestation in the lily
beds in the beach pools and the sphagum bogs, which are
protected from the action of the wind and waves, was very
slight. The writer is not prepared at the present time to
account for this distribution, but merely gives the facts for
this particular region, realizing that the distribution just
described may not agree with that of other regions.

It thus appears that the larva of Hydromyza confluens
is a serious enemy to Nymphaea americana since every petiole
which contains even one larva is doomed. Three summers
of work on insects infesting waterlilies in the Douglas Lake
region has convinced the writer that, although there is a rather
large number of species which attack this plant, yet Hydromyza
confluens has only one rival for first place as the greatest enemy
namely, the larve of Bellura melanopyga, one of the Noctuide,
which also plays havoc in the lily beds.

1914} Life History of Hydromyza Confluens. 141

THE PUPA.

The detailed description of this stage is reserved for another
paper which is in preparation.

Position.—The position of the pupa in the gall is Vane
the only constant feature being the fact that its long axis
always lies almost or quite parallel to the long axis of the
petiole. However, the position of the head and the caudal
end is not at all constant since in some cases the head is up
(towards the surface) and in others it is down (towards the root-
stalk). In galls which are well advanced in development and
contain pupz this point can easily be determined from the
exterior without breaking into the gall by noting the position
of the window, a feature to be described later. When the
window occurs at the lower end of the gall it is positive proof
that the head of the full-grown larva or the head of the pupa is
down; if the window is at the upper part of the gall, the head
of the larva or pupais up. Ina series of observations in which
242 pupze were examined it was found that 130 occupied a
position in which the head of each was up and 112 in which the
head of each was down. Other statistics of the same kind
showed a similar result, namely, that the majority of the
pupz lie in the galls with the head towards the surface of the
water.

The Window.—The window mentioned in the preceding
paragraph is a very interesting and unique provision for the
emergence of the adult and, as stated by Needham, is con-
structed by the larva immediately before pupation. It is
circular in outline and only large enough to allow the passage
of the emerging adult. In constructing this window the
full-grown larva works towards the exterior of the petiole until
it reaches the epidermis. Here it removes all of the surrounding
tissue (exclusive of the epidermis) from a circular area which
is destined to be the window so that the latter is composed
only of epidermis. A circular incision, which extends around
approximately two thirds of the circumference, is made along
the periphery of this area. The remaining one-third is left
intact and thus a circular lid, attached at one side, is produced.
The attached portion always has a definite relation to the
position of the pupa, namely, it is constantly on the side of

142 Annals Entomological Society of America _[Vol. VII,

a

the circumference nearest the caudal end of the pupa. The
larva evidently constructs this circular incision by rotating
the head through about 240 degrees, cutting the epidermis
with the mandibles as it goes. Needham (’08, p. 270) in dis-
cussing this matter makes the following statement: “Just
before transformation to the pupal stage the larva eats a hole
out to the epidermis and returns to the center of the cavity;
this hole is a passage of exit for the adult, which then has
only to break through the transparent epidermal window to gain
its liberty.’’ One would infer from this statement that the
window is opened at the time of the emergence of the adult
by the rupture of the epidermis, but this is not the case. Also
no mention is made of the fact that the epidermis is cut in any
way. The evidence is conclusive that the operation of opening
the window is not a mere rupture of the epidermal tissue
since: (1) Very careful examination shows that a circular
incision is actually made and that the translucent window is
merely pushed open at emergence, (2) it is an easy matter to
open one of these windows on a gall from which the adult has
not emerged by either carefully inserting the point of a needle
‘ between the edges of the incision, or by splitting the gall and
applying very slight pressure against the inside of the window,
which in either case opens as a hinged shutter with the attach-
ment constant in position, and (3) an examination of a large
number of petioles from which the adults had emerged showed
that in every case the window opened as a hinged shutter,
the attachment of which always had the above described,
definite relation to the body of the gall and to the pupa. If
the opening of the window was a matter of rupturing the
epidermis there would be no possibility of this constancy in
the form of the opened window. It should be mentioned in
this connection that the incision is not an absolutely con-
tinuous one, since a few minute portions of the tissue are left
uncut and serve to hold the window closed up to the time of
emergence. It is therefore possible, under average conditions,
to determine from .the exterior whether or not the adult has
emerged. If the window is not loose around the edges the
insect is still within the gall, but if the window is loose and
gapes slightly it is very strong evidence that the adult has
emerged. An exception to this rule may occur in a petiole

1914] Life History of Hydromyza Confluens. 143

which has been subjected to side to side strains such, as are pro-
duced by wave action, and the window has thus been broken
open.

According to the observations of the writer, pupation
occurs shortly after the window is completed and the pupa
lies with the cephalic end in close proximity to the window.
In some cases the pupa lay so close to the window that the move-
ments of the adult in escaping from the puparium would have
been sufficient to open it.

One of the characteristics of Nymphaea americana is the
shape of the petiole. It is conspicuously flattened so that
approximately one-third of the circumference is quite flat
while the remaining two-thirds are very convex. For con-
venience in discussion these surfaces will be designated as the
plane surface and the convex surface. On the former there is a
median, longitudinal ridge. Needham’s paper indicates nothing
as to constancy or variation in the position of the window
with reference to the two above-mentioned surfaces although he
figures a gall with the window on the convex surface. The
position of this window is variable, sometimes occurring on
the plane surface and sometimes on the convex, but not in
equal numbers and the writer was lead to make some observa-
tions on this point. Of 226 galls examined at random the
window in 137 occurred on the convex side and in the remaining
51 cases on the plane surface. Various other counts not
recorded in the above numbers showed similar results. Available
data does not seem to offer an explanation for the predominance
of the windows on the convex surface.

THE ADULT.

Broods.—Although the writer has no positive evidence as
to the number of broods per summer, there seems to be the
possibility of at least two. A few adults were observed about
the Nymphaea beds during the first part of July, at least three
weeks before the larve in the petioles were grown. The
maximum appearance of the adults in 1913 occurred between
August Ist and 6th. During the period, July 10-25, adults
were very rare and it may be that this represents the interval
between two successive appearances of the adults. Very
few of these insects remained in the pupal stage after August

144 Annals Entomological Society of America [Vol. VII,

6th. Adults were very abundant during August 1-6, and
it was a common thing to find numbers of them copulating.

Local distribution.—In spite of the fact that these insects
have well developed powers of flight they are not found at
any great distance from the waterlilies. They occurred
ordinarily on the leaves and flowers of these plants and when
disturbed made only short flights, seldom taking to open
water or to shore. Only in rare instances were flies found
resting on plants other than NV. americana. When undisturbed
they assembled in the open flowers or ran restlessly about
over the lily leaves, making short flights where leaves were not
contiguous.

Relation to water.—These flies are related to the water
in several interesting ways. Although the emergence of the
adult has not been observed in the field it seems safe to assume
that when the adult emerges from the pupal stage it must of
necessity push the window open and pass up through the water
to the surface, either by crawling up the petiole, or by inde-
pendent passage through the water, presumably the former.
It was found that the flies emerged and came to the surface
when the infested petioles of N. americana were brought into
the laboratory and completely submerged in water. In many
cases individuals which develop near the base of the longer
petioles must, in emerging, come up through about five feet
of water before reaching the surface.

It seems almost certain that the female deposits the eggs
by going into the water and crawling down the petiole to the
place where the eggs are deposited. Careful watch was kept
on adults for periods of an hour or more at a time and none
were observed to go beneath the surface in open water. If
perchance an individual did alight on the surface it immediately
took to wing again. Adults which fell on the surface of the
water did not sink, but appeared to be supported mainly by
the surface film. The fact that they may and do voluntarily
pass under water was demonstrated when the writer observed
a few individuals walk over the edge of the lily leaf, go under
water, and travel on the lower surface of the leaf for short
distances. None were seen to go down the petiole.

Experiments in which adults were submerged showed that
under such conditions the flies apparently have a specific
gravity less than water, and consequently they rise to the

1914] Life History of Hydromyza Confluens. 145

surface if opportunity affords. They stay below only by
clinging to some submerged object. While under water a
goodly supply of air clings to them in the form of a dense,
silvery coating. When allowed to come to the surface they
immediately lose the silvery coating and’ are apparently as
dry as if they had never been in contact with water. Experi-
ments, in which adults were subjected to forced submergence
for varying lengths of time, showed that they can remain
under water for several minutes without apparent detriment
to themselves, due without doubt to the generous coating of.
air which surrounds them. Submerged individuals usually
appeared uneasy and made vigorous effort as if seeking release.
The ease with which they apparently resist wetting and the
quantity of air which they take below with them make possible
the mode of emergence and oviposition suggested above.
Relation to the Yellow Waterlily.—The adults as well as the
immature stages have a definite and interesting relation to
the yellow waterlilies. This relation will be discussed under
two heads, (1) food relation, and (2) possible agents in pollina-
tion. Although each of these relations will be treated inde-
pendently, it will be understood that such separation is purely
artificial and also that both are operative at the same time.
(1). Food relation—The time of maximum abundance of
the adults coincided closely with the opening of the majority
of the flowers and it was very evident that the flies were deriving
food products from them. Flies swarmed in the newly opened
flowers in great numbers, congregating between the petals and
the stamens to the extent that often the interior of the flower.
was black with them. In the case of flowers which had been
open only a short time the anthers were crowded in a compact
mass under the edge of the expanded stigma or were just
beginning to spread out in a radial fashion, while the petals
had spread out widely, thus forming a cup-shaped flower
and producing a space between the petals and anthers into
which the flies crowded. Flowers frequently contained as
many as fifty adults. They regularly disposed themselves as
described above with the heads in close proximity to the base
of the anthers. It often required a distinct shake of the
flower stalk to disturb them and this proved to be an easy
way to collect adults since one of these flowers could be cau-
tiously thrust into a bottle and the inmates dislodged. This

146 Annals Entomological Society of America [Vol. VII,

habit resembles a similar one described by Fulton (11, p. 300)
for certain Diptera, the adults of which also congregate in the
flowers of a yellow waterlily. Later when the anthers became
spread out the flies found better concealment beneath them.

The conspicuous assembling of flies in the flowers is indic-
ative of some rather strong attraction which the latter have for
the former and it seems safe to assume that the flies profit
therefrom. Nectar is said (Lovell, ’02, p. 205) (Robertson,
’89, p. 122) to be secreted on the outer faces of the petals in
Nymphaea advena and it is also probably true of Nymphaea
americana. Therefore it is possible that the visits of the
flies are induced in part by the presence of nectar which forms
a source of food supply. Flies were observed in the flowers
from the time of opening to the time when the flowering parts
began to disappear.

(2). Possible agents in pollination—The information that
insects are found in connection with yellow waterlilies is not
new since species representing several orders have been reported
as occurring on these plants by Robertson (’89, pp. 122-123),
Lovell (98, pp. 60-65), Bembower ('11, p. 379) and others.
Furthermore Elliot (96, pp. 117-118) and others claim that
flower haunting Diptera are of considerable importance in the
fertilization of many of the flowers which are visited. It
is claimed that N. advena may be self- or cross-pollinated
(Bembower, ’11, p. 379) and this is probably true also of NV.
americana. There is good evidence in support of the view that
the insect visitors of yellow waterlilies (V. advena and others)
may transfer pollen from one flower to another, or from one
part to another on the same flower. The writer had occasion
to examine large numbers of the adults of Hydromyza confluens
and it was discovered that many were carrying the pollen of
N. americana. Swarms of adults taken from flowers in which
they had congregated showed that the great majority, and
often all, of the insects were dusted with pollen. Very fre-
quently pollen occurred so thickly over the body that the
insect was distinctly yellow in appearance. Adults collected
August 5-22, showed that pollen was being carried during this
entire period. While it was not demonstrated absolutely
that these flies carry pollen from one plant to another, the
circumstantial evidence seems to point definitely to these insects
as being at least one of the factors in the cross-pollination

1914] Life History of Hydromyza Confluens. 147

(and possibly the self-pollination) of N. americana and may be
summed up as follows: (1) The coincidence of the blooming
period of N. americana with the maximum appearance of the
adults of Hydromyza confluens; (2) The large numbers of
flies limited in distribution to the immediate vicinity of the
lily beds; (3) The assembling of the flies in large numbers
within the flowers when the latter have opened sufficiently
to admit them; (4) The heavy loads of pollen which are carried
by many of the flies and the almost universal presence of
varying quantities of pollen on all individuals; (5) The continu-
ous blooming of N. americana throughout the greater part of
August, so that at any given time there were flowers in all degrees
- of maturity, a fact which eliminates a difficulty due to the pos-
sibility that a given flower is proterogenous; and (6) The
behavior of the adults in preferring to pass from place to place
by crawling and by very short flights (usually the former when
possible) rather than by extended flights, which means a
maxium of contact of the insect with the various parts of the
supporting plant.

LITERATURE CITED.

Bembower, W., 1911. Pollination Notes from the Cedar Point Region. The
Ohio Naturalist, 11:378-383.

Elliot, G. T. S., 1896. Flower-haunting Diptera. Trans. Ent. Soc. London,
pp. 117- 118. (Abstract in Am. Nat. 30:760, taken from Journ. Royal
Micr. Soc.)

Fulton, B. B., 1911. The Stratiomyide of Cedar Point, Sandusky. The Ohio
Naturalist, 11:299-301.

Johnson, C. W., 1904. A Sapplementary List of the Diptera of New Jersey. Ent.
News, 15:157-163.

Lovell, ” H., a Three Fluvial Flowers and their Visitors. Asa Gray Bulletin,

-65

1902. The Colors of Northern Polypetalous Flowers. Am. Nat.,
36:203-242.

Miller, G. S. and Standley, P. C., 1912. The North American Species of Nymphea.
Contributions from the U. S. National Herbarium, 16:63-108, 12 pls.,
40 figs.

Needham, J. GC, 1908. Notes on the Aquatic Insects of Walnut Lake. Appendix
i. A. Biological Survey of Walnut Lake, Michigan, by T. L. Hankinson.
A Report of the Biological Survey of the State of Michigan, Published
by the State Board of Geological Survey as a part of the Report for
1907, pp. 252-271.

Robertson, C., 1889. Flowers and Insects. I. Bot. Gaz., 14:120-126.

Department of Entomology, Kansas State Agricultural College, Feb. 26, 1914.

SOME SPECIES OF THE BEE GENUS COELIOXYS.

By J. C. CRAWFORD.

This paper discusses only species occurring in America north
of Mexico and no table to separate the males has been included
since Prof. T. D. A. Cockereli published a table for this sex in
the Canadian Entomologist for June, 1912, pp. 167-170. The
key to the females includes all the species for the region under
consideration in which the female sex has been described. In
the table here presented the characters used for separating
rufitarsis Sm. from comstockit Cress. and lucrosa Cress. from
moesta Cress. are the characters used by Prof. Cockerell in, a
table to separate the types of the Cressonian species and some
non-Cressonian species which he consulted in the collection
in Philadelphia. The illustrations were made with a camera
lucida, attached to a Zeiss binocular microscope.

1. Last ventral segment not notched laterally, at most emarginate and

the part anteriad of the emargination not pointed.................. 2
Last ventral segment notched, the part anteriad of notch sharply pointed. .19
2. Last dorsal segment with the "end upturned into a small spicule 20s See 3
Last dorsal segment not upturned at apex. (2:2: 2.-:- 22... 2 see eee <
3. Last dorsal segment very sparsely punctured at base.:.... obtusiventris n. sp.
Last dorsal segment closely punctured at base... .2:..--=-- see i

4. Punctures of first dorsal abdominal segment separated by less than a
punctured width... 020.26 feo on thea se eles ae eee Swe 5
First dorsal abdominal segment sparsely punctured.............:...2..%- 6

5. First recurrent vein received by second submarginal cell almost half as
far from base as length of first transverse cubital in the @ and slightly
less‘in Qs. oo ta Se See ae pee ee Ree eae ern gilensis Ckll.
First recurrent vein received by second submarginal cell one-third or
less as far from base as length of first transverse cubital in 9 and
still ‘less in..d\. ja. Soak ee a ee en deani Ckll.
6. Here run modesta Sm. and scitula Cress., the descriptions affording no
points for separation.
7. Last ventral segment towards apex with a long strong fringe of hairs

along Margin. . . <5... 0.06 ss'ccelse cates ware ewe phe nie oe Sp wees 8
Last ventral segment not strongly fringed./..°. ..0.....wea + ose 12
8. - Legs black... 2... )e0.n 0 peach cee ote tie oe eee ee ee 9
Legs, except coxae, Ted... 6 6 oy caste sin 6 arm ep as sate Se 11
9. Last dorsal segment without a median carina or this only indicated at
BE CIP 2 ed cca ei swrn Se7e aie Siase Scale atmo Gente Cyan tate rere share Stat Srl Onn ae a a
Last dorsal segment with a median carina.....;............+-. angelica Ckll.
10. Penultimate ventral segment with small punctures interspersed among
the larger ones... ios ooy.2<5 Ve ene oe ee eee ree apacheorum Ckll.
Penultimate ventral segment without smaller punctures among the
OCDELE oa is Feralas lag e's tale woes WR oe ea alternata Sm.
11. Fourth antennal joint distinctly longer than third; last ventral segment
with subparallel sides and a broadly rounded Es hovs> een ret texana Cress.

Fourth antennal joint hardly longer than third; last ventral segment
with the sides converging apically and apex more narrowly rounded.
hunteri n. sp.

148

1914] The Bee Genus Celioxys. 149
12. Last dorsal segment near apex with two small flattened projections
ee Tt ATES SS aoe piercei n. sp.
Last dorsal segment without projections on disk.........................
EEE forte ew so edie occ ws ie epee lee glove dw'ou cence dawetes 13a
ee Et SS tt Soe eb ne oo os warns Rok be sk de eee eee 14
13a. Front at top of inner orbits with a swollen granular area, narrowed
centrad and extending to lateral ocelli.................. deplanta Cress.
Front without such a spot, being coarsely punctured, not different from
PRP ELS Seg eS sculptifrons n. sp.
14, Scutellum strongly triangularly produced, medially almost impunctured.
; dolichos Fox
Scutellum medially closely punctured and not strongly produced........ 15
15. Last ventral segment with the sides entire................ alternata Cress.
Last ventral segment with the sides emarginate.....................0.. 16
16. Thorax above with lines of appressed pubescence.....................02. 17
Mipremanoveswitunonlys erect Nairs: oo. oe css ec sec ccc cece cacowss 18
17. ‘‘Scutellum medially produced into a tubercle’”’................. aperta Cress.
Scutellum medially not produced into a tubercle............. grindeliae Ckll.
18. Pubescence white.......... DU eee ene ribis Ckll.
“Pubescence ochreous; basal part of third abdominal segment more
sparsely punctured than in above’’............ ribis var. kincaidii Ckll.
19. Clypeus near apex bilobed (viewed from above i. e. not emarginate at
MO ENS oe eg in a Lb thn 8 hin Seo Se ae Oe a ae eee ce enn as San 20
OE SESS aS ey ey a a ne 21
20. Transverse furrows on segments 2 and 3 deep; punctures on middle of
segment 2 basad of furrow close, separated by about a puncture width;
arcuate edge of pronotum much more strongly produced, translucent;
oll Da EA Se See Se eee eae novomexicana Ckll.
Transverse furrows on segments 2 and 3 shallow; punctures on middle of
segment 2 basad of furrow separated by much more than a puncture
width; arcuate edge of pronotum not strongly produced, black; legs
eRRMEREOAL CL RISiiea Nye (hai Kye aos ec a a spec, aceet ssc eee tls sayi Robt.
21. Clypeus medially triangularly produced and somewhat reflexed, banksi n. sp.
Clypeus apically truncate, gently round, emarginate or tuberculate...... 22
22. Basal abdominal segment at least entirely red..................2..0005- 23
Basal abdominal segment black (at most with sides red)................ 25
23. Scutellum sparsely punctured, somewhat produced medially and slightly
EG Ora ROS cate eS eaetnetc ae Birch Nb Ae eerie ee menthae Ckll.
PEMeelity CLOSELY. PUN GULed te sme © Gio ee tite teas stir aly cow he ce cee ee eee 24
24. Abdominal segments 1-3 red; wings, except extreme bases dusky, slossoni Vier.
Segment 1, only, red; wings with only apical part dusky,
slossoni var. arenicola n. var.
i LaBGS ‘RSG et Ge aeons soo epics Baca SS Se cer 26
elk PUES SEE Pies gag le ae ee a rr |
26. Scutellum with a strong median projection................... germana Cress.
pembelsin wedially ab anOSt HUMETCHIALG so... oe ee ce eee 27
27. Third joint of antenne hardly longer than second, about half as long
Bok Te Garislels ee ee eee CR iy SS ie, ns er asteris n. sp.
Third joint of antennz distinctly longer than second, almost as long as
Be eR Da ae ta
28. Last dorsal segment narrowed at almost a right angle; first abdominal
segment closely punctured laterally.................. coquilletti n. sp.
Last dorsal segment not narrowed at almost a right angle, at most at
SEMAINE RENE ISC CLIN LES pn ney Renn: HAE ch whe tkla.sko's vie Sie a’oyeuyeceecelac ed een’ 29
29. Last ventral segment very narrow, strongly bent downward; apex of

last dorsal segment cephalad of notch of last ventral by one and one-

half times the distance from notch to apex of segment........ insita Cress.
Last ventral segment broad, the sides diverging basad, not strongly

MI REN te gl Ig SENS yar a i n'a vis dines = on daly ap siak wx ONY

150 Annals Entomological Society of America _[Vol. VII,

30. Face with many erect brown bristle-like hairs among the appressed
light ones; punctures of first abdominal segment laterad less than a

puncture-width, apart... 2.0. 2-05 <.52-5esd-_ == a) eee pratti n. sp.

Face without dark bristle-like hairs; first abdominal segment sparsely
punctured laterad.....-. <2... psig eee ee ee eee octodentata Say.
31. Last dorsal segment strongly angularly narrowed....................... 32
Last dorsal segment at most roundly narrowed..................--+++--- 33

32. ‘‘Part of last dorsal beyond constriction much larger than wide,”’
rufitarsis Smith
“Part of last dorsal beyond constriction almost as wide as long”’
comstockii Cress.

33. ‘Larger, 13mm. Jone”’. 01) /.0-Eose oe ee lucrosa Cress.
Smaller, hardly 12 mm. long; abdomen more slender and more closely
punctured: ...... 5.0. his. ass ee ee ee moesta Cress.

Ceelioxys obtusiventris new species.

Length about 11mm. Black, the tegule and legs, except coxe,
ferruginous; face coarsely rugoso-punctate, vertex coarsely punctured;
the punctures separated by much less than a puncture width; scape
and pedicel (rest of antennz missing) dark, obscurely reddish; face with
white hair, dense along inner orbits, interspersed with long bristle-like
hairs; mesoscutum and scutellum with punctures as on vertex; scutellum
with a tubercle medially on posterior margin; lateral teeth long; mesono-
tum with pubescence along anterior margin and base of scutellum
(badly worn); wings infuscated, apically more deeply so; abdomen
sparsely, rather coarsely punctured; segments 1-5 with apical bands of
white hair; second, third and fourth segments with transverse furrows,
interrupted medially, apicad of these furrows there is an almost impunc-
tate line, the extreme apices of these segments with a few punctures;
last segment with a few scattered finer punctures, constricted, the apical
portion covered with erect brown hairs, the extreme tip upturned;
ventral segments, except apex of last, coarsely, closely punctured, last
ventral very broad, not notched, medially produced into a long straight
spine, with a very strong fringe of brown hair.

One specimen from the C. F. Baker collection with the rec-
ord ‘‘Florida; Palm.” |

Type specimen Cat. No. 18217, U. S. N. M.

Although the single specimen is badly rubbed it is described
since it is easily separated from the other species having the
last dorsal segment turned up at apex and by that segment
being almost impunctured. The spine at the apex of the
segment is also much longer, in the other species being hardly
more than an angulation of the apex.

Ceelioxys alternata Say.

In the table this species occurs twice since the fringe of
hairs along the margin of the last ventral segment is not very
strong and there might be some difficulty on this account if
the species were not listed under both categories

a

1914] The Bee Genus Celioxys. 151

Ccelioxys texana Cresson.

For comparison with C. hunteri camera lucida drawings of
the last ventral segment and of antennal joints 2-5 of the
female are given. (Fig. 1).

a

Fig. 1. C. texana Cress. (a) Last ventral segment
(b) antennal joints 2-5-of female.

In the antenne, the third joint is shown to be hardly longer
than the second (exclusive of bulbous base) and the fourth is
distinctly longer than the third.

Ceelioxys hunteri new species.

Female. Length about 13-15 mm. Black, with red legs; ie scape
and pedicel, tubercles, carinate lateral edges of pronotum and tegule,
reddish; lateral margins of basal abdominal segments sometimes ob-
scurely reddish; face rather finely rugoso-punctate with a median
impunctate line from in front of anterior ocellus to base of clypeus and
indistinctly indicated on clypeus; anterior ocellus enclosed by two
cresent-shaped raised impunctate areas which are finely reticulate;
upper inner orbits each with a similar sculptured spot; face with rather
abundant white hair, thicker along inner orbits and around antennz;

= t

Fig. 2. C. hunteri Cwfd. (a) Last ventral segment
(b) antennal joints 2-5 of female.

second joint of antenna (not counting bulbous base) oie shorter than
third, the third about as long as fourth; (See Fig. 2, b); vertex and
mesoscutum with large rather sparse punctures, each with an appressed

152 Annals Entomological Society of America [Vol. VII,

white hair; scutellum and its lateral spines shaped about as in texana;
wings dusky, with the apical margins more deeply infuscated; coxze
black with more or less obscure reddish at apices; tarsi mostly dark;
spines on anterior coxe short; mesonotum at base and along lateral
margins with lines of appressed, slightly ochreous hair, at base forming
two spots near middle; scutellum at base with two transverse spots of
similar pubescence; under side of scutellum at apex and metanotum
with dense subappressed white hair; propodeum and pleurze with long
white hair; abdomen shiny, with sparse rather coarse punctures, last
segment with a silky lustre ae punctures longitudinally elongate, the
last dorsal and ventral (Fig. 2, a) segments (Fig. 2, a) shaped about
as in alternata; basal margin of segment one and apical margins of seg- ~
ments 1-5 with lines of appressed white pubescence; segments 2-4 with
diagonal lateral lines of similar pubescence near bases.

Type-locality: Hearne, Texas.

Described from five females collected ‘‘at nests in bogs”’,
July 23, 1906, by F. C. Bishopp.

The species is named in honor of Mr. W. D. Hunter in
charge of the investigation from which these specimens were
obtained.

Type—Specimen: Cat. No. 18218, U. S. N. M.

This species in the structure of the apical plates is near-
texana and alternata; the last has dark legs; texana has the last
ventral segment with almost parallel sides and apically broadly
rounded; altenata and huntert have this segment narrowed
apically and consequently pointed at apex; in alternata the last
dorsal segment is shiny and with sparse small punctures.

Ccelioxys piercei new species.

Female. Length about 9.5 mm. Black, ‘including the legs, only
the apical joints of the tarsi somewhat reddish; face rather finely rugoso-
punctate; antennze black; vertex and mesoscutum closely, coarsely
punctured, scutellum slightly coarser rugoso- punctate; face and dorsum
of thorax with slightly ochraceous pubescence, more abundant on sides
of face and around antennz and forming lines along anterior and lateral
margins of mesoscutum and indistinctly so along base of scutellum;
pleure with abundant lighter colored hair; lateral teeth of scutellum
moderate in length, slightly incurved; tegule black; wings slightly
dusky with the apical margins broadly deeply infuscated; abdomen
closely, rather coarsely punctured, the last segment more closely and
finely punctured, segments 2 and 3 with a deep and segments 4 and 5
with a shallow transverse furrow; segments 1-5 with apical bands of
appressed white pubescence and segment 1 with the lateral margins
with similar hair; base of first segment with an indistinct band of slightly
ochraceous hair; last dorsal segment with a median longitudinal carina,

1914] The Bee Genus Celioxys. 153

the segment rather suddenly constricted, near apex with a flattened
projection on each side of carina (see fig. 3); last ventral segment extend-
ing a little beyond last dorsal, seen from below subtriangular in outline,
the lateral edges straight, with only weak hair and without a projecting
point.

Fig. 3. C. piercei Cwfd. Last dorsal segment of female
(last ventral indicated by- dotted line).

Described from one female from Cotulla, Texas, April 17,
1906, on Verbesina encelioides, F. C. Pratt, collector.

Type—sSpecimen: Cat. No. 18219, U.S. N. M.

The two curious flattened projections on the last dorsal
segment readily distinguish this from any species known to me.

Named in honor of Mr. W. Dwight Pierce who was actively
interested in the work which resulted in the accumulation of
the splendid collection of Texan Hymenoptera.

Ceelioxys edita Cress.

This species was described from a male. Female from
Texas which I have associated with this species are deplanata
Cress. and I am inclined to think that edita should be classed
as a synonym of this species, although the association of sexes
I have made may be incorrect.

Ceelioxys sculptifrons n. sp.

Female. Length about 11.5mm. Black, with the tegulz and the
legs, except coxe, ferruginous; clypeus rugoso-punctate with smaller
punctures interspersed, the apical margin with five short teeth; face
above insertion of antennz coarsely, closely punctured, more sparsely
so laterad of the ocelli; mesoscutum and scutellum coarsely punctured,
the punctures except on disk of scutum crowded; lateral teeth of scutel-
lum short, pointed; sides of face with dense long white subappressed
pubescence, pubescence on clypeus finer and not so conspicuous; lateral
and posterior margins of mesoscutum with indistinct lines of white
appressed pubescence; pleure with dense long white hairs; wings dusky,
with the apical margin more densely infuscated; apical margins of dorsal

154 Annals Entomological Society of Amertca [Vol. VII,

and ventral abdominal segments 1-5 with bands of appressed white
pubescence; first abdominal segment rather coarsely and closely punc-
tured: second and third segments with distinct, complete, transverse
impressions, the second with rather fine punctures basad of its impres-
sion, the punctures about a puncture width apart; apicad of the impres-
sion, the punctures sparse averaging two or more times a puncture width
apart and finer and sparser toward middle, third segment basad of
impression with the punctures somewhat wider apart than on base of
second segment; apicad of the impression the punctures about as far
apart as on apical part of second segment; fourth and fifth segments
apically punctured about as apex of third segment; sixth segment with
a distinct median longitudinal carina, basally finely punctured, the
punctures slightly more than a puncture width apart; apically the
punctures become slightly larger and crowded; near apex on each side
of the median carina a depressed area bounded laterally by an elevated
* margin which is very indistinctly irregularly carinated; ventral segments
1-4 coarsely, closely punctured; fifth coarsely punctured at base, the
apical part minutely very closely punctured; last ventral segment with
the sides emarginate near apex.

Type-locality: Ithaca, New York.

Described from one specimen with the record, July, 1-7,
from the collection of Mr. Nathan Banks.

Type—Specimen: Cat. No. 18220, U.S. N. M.

This species resembles C. deplanata, but differs as shown in
the table and also by the sparser punctures on the abdominal
segments apicad of the transverse impressions on second and
third segments and by the sparse punctures on segments 4 and 5.
In deplanata the punctures on the last segment are coarser at
base, the impressions near apex are not so deep nor do they
extend so far laterad.

This species differs from C. tmmaculata Ckll. described only
in the male sex in the punctation of the second abdominal
segment beyond the transverse impression and since the two
sexes in this genus agree very closely in such characters, I do
not think it possible for this species to be the same as the one
described by Prof. Cockerell.

_Coelioxys sayi Robertson.
C. octodentata Cresson (not Say).
The synonomy of Mr. Robertson of this species and of

C. octodentata Say (C. altilis Cress.) is adopted for it is evident
that he has correctly interpreted the original description of Say.

i |

1914] The Bee Genus Celioxys. £5!

Ceelioxys banksi n. sp.

Female. Length about 11 mm. Black, femora black, the rest of
the legs ferruginous with the middle of the tibize obscured with blackish
and the tarsi dark toward apices; face above antenne very coarsely
punctured, the clypeus medially triangularly produced and somewhat
reflexed; mesoscutum amd scutellum very coarsely punctured, the
punctures well separated on the disk; lateral teeth of scutellum rather
short, pointed; scutellum gently rounded posteriorly; middle of face
with appressed long white pubescence, clypeus with similar short
pubescence; suture between mesoscutum and scutellum with a line of
appressed white pubescence, a spot of similar hair at the posterior end
of tegule; mesopleurze with the anterior and posterior margins densely
clothed with similar pubescence, as is the region immediately in front
of and below tegule; the punctures of mesoscutum each with a long
white delicate hair; tegule dark, obscurely reddish on disk and outer
margin; wings dusky, with the apical margins more densely infuscated;
dorsal and ventral segments 1-5 with apical bands of appressed white
pubescence; first abdominal segment rather coarsely and sparsely
punctured, the second and third with transverse impressions, basad of
them the punctures about a puncture width apart, apicad of them the
punctures slightly larger, laterad about as dense as basad of impressions
but medially very sparse; fourth dorsal segment punctured about as
third, with a transverse impression which is interrupted medially; fifth
segment with the punctures finer, basally less than a puncture width
apart, apically the punctures more than their own width apart; last
dorsal segment with a median carina, the punctures close, the apical
production of the last segment almost as long as the basal part of the
segment; apex of last dorsal segment basad of notch of last ventral
segment by about one and one-half times the length of the distance from
notch to apex of segment; ventral segments 1-5 coarsely, closely punc-
tured, punctures on fifth segment decreasing in size apicad; last ventral .
long, narrow, the apical portion bent downward, the sides near apex
with a distinct notch.

Type-locality: Falls Church, Virginia.

One specimen, collected August 20, from the collection of
Mr. Nathan Banks, after whom the species is named.

Type—Specimen: Cat. No. 18221, U.S. N. M.

The peculiar form of the clypeus easily distinguishes this
species. In this table if this character be omitted the species
would run to couplet no. 33, but the two species in that couplet
both have the legs entirely black as well as the clypeus differ-
ently formed, etc.

156 Annals Entomological Society of America __[Vol. VII,

Ccelioxys slossoni Viereck.

In the collection of the U. S. Nat. Mus. are two badly
rubbed females which agree with Viereck’s original description,
one with the record ‘‘Palm Beach, Fla., 3-’00, collection C. F.
Baker’’, the other without locality from the Ashmead collection.
The Palm Beach specimen has the apex of the third segment
dusky.

Ceelioxys slossoni arenicola new variety.

Female. Length about 13.5 mm. Differs only in having the
abdomen, except basal segment, black (one has segments 2 and 3 in
part obscurely reddish) and the wings subhyaline with dusky margins.

Male. Length 10.6 mm. Similar to the female in sculpture and
color; the apex of the abdomen with 4 teeth the upper pair blunt some-
what flattened and divergent the lower pair longer pointed subparallel;
base of last segment with a tooth on each side; fifth segment not toothed.

Type-locality: Brownsville, Texas, (April 17, 1895, C. H. T.
Townsend, collector). ;

Allotype male from San Diego, Texas.

Other localities: Calhoun, Co., Texas, J. D. Mitchell,
collector, one female; Nuecestown, Texas, 4-28-—’96, C. L.
Marlatt, collector, one male; also two paratype females from
Brownsville, Texas, and one male from San Diego, Texas.

Type—Specimen: Cat. No. 18222, U.S. N. M.

It is most probable that the species recorded from Galveston,
Texas, by Brues* as menthae is this form.

Ceelioxys asteris new species.

Female. Length about 14 mm. (abdomen unduly extended).
Black with the tegule reddish-testaceous and the legs, except coxe,
ferruginous; face below antennz finely rugoso-punctate, above antennz
coarsely punctured with an impunctured but lineolate area laterad of
each lateral ocellus but none in front of and beside anterior ocellus;
third joint of antennz much shorter than fourth (see fig. 4, b,); reflexed
lateral margins of pronotum strongly developed, translucent; mesoscu-
tum with coarse, close punctures; scutellum rugoso-punctate and with
a rather indistinct median longitudinal carina; lateral teeth of scutellum
rather long, somewhat incurved, thick dorso-ventrad and carinate above
along inner edge; wings deeply infuscated, more so along apical margins;
abdomen finely, sparsely punctured, segments 2-4 each with a shallow
transverse impression broadly interrupted medially; caudad of these
furrows the segments almost impunctured; base of last segment more

*Entom. News, XIV, 83, 1903.

1914] The Bee Genus Celioxys. 157

finely and closely punctured, the apical constricted portion finely
rugoso-punctate; last ventral segment broad, notched near apex (see
fig. 4, a,); ventral segments 1-4 rather coarsely punctured, five with
similar punctures at base and fine ones at apex.

;
Fig. 4. C. asteris Cwfd. (a) Last ventral segment
b) antennal joints 2-5 of female.

Type-locality: Victoria, Texas.

The type collected Nov. 6, 1904, by Mr. A. J. Leister,
“on aster’’; a paratopotype with the same date and collector
is only about 11 mm. long.

This species resembles octodentata which has the third joint
of the antenne almost as long as the fourth.

Type—Specimen: Cat. No. 18223, U.S. N.. M.

Ceelioxys coquilletti new species.

Female. Length about 12 mm. Black with ferruginous tegulz and
legs; face below antennz finely rugose, above, coarsely so without any
smooth spots; third joint of antennz almost as long as fourth; face with
abundant pubescence at sides (rest worn ?); mesoscutum coarsely rugose
all over; with a short lateral carina on each side near tegulz; reflexed
lateral.margins of pronotum strongly elevated, translucent; an inter-
rupted line of appressed white pubescence (worn ?) at base of mesoscu-
tum, a line at base of scutellum and one at lateral margins of mesoscutum ;

Je

a

Fig. 5. C. coquilletti Cwfd. (a) Last dorsal segment
(b) last ventral segment of female,

158 Annals Entomological Society of America [Vol. VII,

scutellum slightly more finely rugose than mesoscutum, the lateral
teeth rather long and slightly incurved; propodeum and pleure with
long white hair; wings slightly dusky with the apical margins somewhat
more so; spines on anterior coxz long; abdomen with the venter largely
reddish and the edges of dorsal segments close to venter somewhat
reddish (to be seen only from below); first segment of abdomen closely
punctured, the punctures laterad separated by less than a puncture
width; segment 1 with a basal and segments 1-5 with white apical hair
bands; segments 2 and 3 deeply and 4 rather shallowly transversely
impressed; the segments basad of the impressions closely punctured,
apicad of them very sparsely so; last dorsal segment suddenly angularly
constricted, and with a median carina (see fig. 5a); last ventral notched
at sides (see fig. 5b); ventral segments 1-5 with apical hair bands and
coarsely punctured.

Habitat: Los Angeles, Co. Cal., (D. W. Coquillett, collec-
tor).

Type—Specimen: Cat. No. 18224, U.S. N. M.

In general this resembles octodentata but differs in the vertex
being rugose and without smooth areas, in the rugose meso-
scutum (in octodentata the punctures on the disk of the meso-
scutum are well separated); in the abruptly narrowed last
dorsal segment, and in the closely punctured first segment of
the abdomen. In this last character it resembles pratti but
differs in all the other characters quoted above; pratti has the
mesoscutum more closely punctured than octodentata but it
is not rugose. In the shape of the last dorsal segment this
resembles rufitarsus from which it differs in addition to the
difference in the color of the legs by the first segment being
closely punctured, by having the second and third segments
basad of the transverse impressions more closely punctured
and by having the fifth ventral, except apex, with coarse
punctures, etc.

Ceelioxys insita Cresson.
The apex of the ventral segment as illustrated and marked
on the figure is the approximate point to which the last dorsal
segment comes (indicated by a U in the sketch). (Fig. 6.)

Fig. 6. C. insita Cress. Last ventral segment of female.

1914] The Bee Genus Celioxys. 159

Ceelioxys pratti new species.

Female. Length 11.5 mm. Very similar to C. octodentata but
differs in the clypeus having many erect, brown, bristle-like hairs among
the dense white pubescence, the eyes with longer, much denser and
distinctly brownish pubescence, the punctures of the mesonotum
somewhat finer and denser, the first abdominal segment with close
punctures, those laterad separated by much less than a puncture
width; fifth ventral abdominal segment with coarse punctures only
at base.

One female with the record Kerrville, Texas, April 14, 1907,
on Marrubium vulgare, H. Durham, collector.

Type—Specimen: Cat. No. 18225, U.S. N. M.

C. octodentata has the punctures of the mesonotum well
separated medially, the first segment. with the punctures toward
sides separated by more than a puncture width and the fifth
ventral segment with coarse punctures except on the apex.

This species is named for Mr. F. C. Pratt through whose
efforts the large amount of material from the type-locality of
this species was secured.

Ceelioxys rufitarsis var. rhois Ckll.

This differs from the typical form only in having the tegulz
black and the veins of the wings darker than normal, and
would run out in the table where the typical form does.

CONNECTANT FORMS BETWEEN THE MUSCOID
AND ANTHOMYIOID FLIES.

By Cuar_es H. T. TownseEnpD,
Director of Entomological Stations, Lima, Peru.

The object of this communication is to point out certain
forms which appear to be transitional between the muscoid and
anthomyioid types, to call attention to their evident affinities,
and to suggest characters which may be used for establishing
a boundary line between these two natural groups of flies.

Girschner’s system, proposed in 1893, recognizing two groups
which he called Tachiniden and Anthomyiden, contains many
elements of truth. It has resulted in demonstrating muscoid
affinities in certain forms hitherto accepted without question
as anthomyioid. Bezzi and Stein have followed this system
in their catalogue, and Schnabel and Dziedzicki have recently
attempted to reinforce it in their treatment of the anthomyioid
flies. What concerns us chiefly in the present consideration
is the fact that Musca and its immediate allies fall in the
Anthomyiden according to this system. The concept is funda-
mentally wrong nomenclatorially, however it may be justified
otherwise. Whatever group Musca is found to fall in must
take its name from that genus. If Girschner’s group Anthomy-
iden be adhered to as it stands, its name must be Musciden
according to all accepted rules of nomenclature. But it is
certain that many characters remain to be investigated before
this grouping can safely be pronounced a natural one, for the
main separation is founded practically on a single character—
the presence or absence of hypopleural bristles.

The solution of the question practically hinges on whether
Musca is, or is not, more closely allied to Anthomyia than it
is to Calliphora. Wherever Musca goes, it will carry with it
a considerable contingent—Stomoxys, Muscina, Mesembrina,

160

1914] Muscoid and Anthomyioid Flies. 161

Glossina, all their immediate allies, and quite probably a
block of forms hitherto classed as Anthomyiide. The position
of Calliphora has never been questioned, but the other forms
are more or less connectant between Calliphora and Anthomyia,
and the affinities of Musca have long been confused with those
of the truly connectant forms. We are thus practically in
the rather paradoxical position of being unable to place taxo-
nomically the type of the superfamily Muscoidea, which seems
inclined to fall in the Anthomyioidea.

If Musca prove to be more nearly related to Anthomyia than
to Calliphora, then one of two things must result. Either
the Muscoidea in the writer’s sense must extend itself to
include all the anthomyioid flies; or the latter must be grouped
with Musca and the connectant forms into a totally different
superfamily to be known as the Muscoidea, thus completely
changing the sense of the name and leaving the Calliphorine and
higher groups to form a superfamily by themselves. It is there-
fore evident that a pressing necessity exists for fixing definitely
the status of Musca with relation to the connectant forms
that intervene between Anthomyia and Calliphora.

Certain students, not caring to proceed farther, will adopt
the former solution of the difficulty and thus dismiss the whole
subject. But this is not the correct solution, for it obscures
the real affinities of the two groups. The anthomyioid flies,
as a whole, present a far greater contrast with the rest of the
Schizometopa, which is to say Muscoidea, than do the various
groups of the latter with each other. Moreover, there are
at least two family types— Coenosia and Anthomyia—repre-
sented in the Anthomyioidea, and it is a question whether
Drymeia does not constitute a third and Fannia a fourth.
Nor can we reduce the value of the taxonomic groups one
notch, thereby considering the whole Schizometopa as one
superfamily, for such action would only compel the inauguration
of a new category farther down the line in order to preserve a
proper conception of relationships. The anthomyioid flies
constitute a superfamily of the atypic class, which is to say
that they occupy a position entirely outside the proper limits
of the superfamilies Muscoidea and Borboroidea (Holometopa
excl. Conopide), but intermediate between the two. As
such they claim separate recognition.

162 Annals Entomological Society of America [Vol. VII,

In order to fix permanently the taxonomic status of Musca,
a comparative study must be made of Anthomyia pluvialis L.,
Calliphora erythrocephala Mg., Musca domestica L., the con-
nectant forms and the main anthomyioid types, along the fol-
lowing lines:

(1) Chaetotaxy.

(2) Pilotaxy—This term is coined to designate the dis-
position of hairs and pile in the Diptera in general and the
Muscoidea in particular.

(3) Pleural and other external thoracic anatomic char-
acters.

(4) Venation.

(5) Male reproductive system.
(6) Female reproductive system.
(7) Hypopygium.

(8) Egg.

(9) First-stage maggot.

Lowne has worked out Calliphora erythrocephala quite
extensively, and Hewitt has done the same for Musca domestica.
Anthomyia pluvialis needs similar attention before exact
comparisons can be instituted. .As to male reproductive-
system characters in the connectant and anthomyioid forms,
Stomoxys has been worked out by Roubaud, and verified by
others including the writer. Othellia, Haematobia, Hypodermodes
and Morellia have been worked out by Thompson, the last two
not yet published; Muscina, Synthesiomyia, Morellia, Limno-
phora, Leucomelina, Fannia and Gen. Indet. have been worked
out by the writer. In addition to these many nonconnectant
muscoid forms have been investigated as to the male reproductive
system by both Thompson and the writer, and Auchmeromyia
and Choeromyia have been similarly worked out by Roubaud.
All of the above named connectant and anthomyioid genera
except Fannia agree with Musca in lacking the male accessory
glands. Fannia and all the nonconnectant muscoid forms
possess such glands, though they may be rudimentary in the
higher forms.

1914] Muscoid and Anthomyioid Flies. 163

The writer has worked out the female reproductive system
and egg in Stomoxys, Muscina, Synthesiomyia, Leucomelina,
Limnophora and Spilogaster, besides many mnonconnectant
muscoid genera. The first three agree with Musca and the
Calliphorinae in egg characters, but the last three differ con-
siderably from them in these characters. Available data on
the lines above specified are presented below.

CHAETOTAXY AND PILOTAXY.

Hypopleural bristles present in a more or less vertical row,
pteropleural bristles present; when 3 sternopleural bristles
present, their formula is either 2.0.1 or 1. 1. 1—All muscoid
families except Muscidz, Oestride, Cuterebride.

Higher True hypopleural bristles present, pteropleural bristles absent
Muscoidea but in their place hairs or pile; sternopleural bristles 2. 0. 1 or
1.0.1—Bengaliine (Calliphorine).

Macrochaete entirely absent; row of hypopleural hairs present
homologous with true hypopleural bristles—Gastrophilus and
Cobboldia.

(
True hypopleural bristles absent, hypopleural hairs and pile
Typical
|

absent; pteropleural bristles present, also often pteropleural
Muscoidea

hairs or pile; sternopleural bristles 1. 0. 2—Musca, Morellia
and Glossina.

Hypopleural hairs present, pteropleural’ hairs absent; sterno-
pleurals 1. 0. 2 or 0.0.2—Synthesiomyia and Graphoymia.
Hypopleural hairs and bristles both absent; pteropleural
hairs, sometimes of a bristly nature, present; sternopleurals
none, 0.0.1, 1.0.1, or 1.0.2-4—Haematobia, Hypodermodes,
Connectant Mesembrina, Eumesembrina, Pyrellia, Orthellia.
Muscoidea Hypopleural and pteropleural hairs present; sternopleurals
0.0.1—Stomoxys.
Neither hypopleural nor pteropleural hairs, pile or bristles
present; sternopleurals normally only 3 and formula 1.0.2
(abnormally 2.0.2)—Muscina, Myospila, Clinopera, Leuco-
melina, Gen. Indet., Limnophora, Aricia, Spilegaster.

(
be 3 Neither hypopleural nor pteropleural hairs; sternopleurals 3 or
Anthomyioidea { more and rarely 1.0.2—Anthomyia, Fannia, Coenosia.

PLEURAL ANATOMY

The name squamopleura is hereby proposed for the inferior
swollen lobe of the metapleura of authors, being the lower
lobe of the lateral plate of postscutellum (Hewitt). The
sclerite in question is a part of the mesothorax. The term
metapleura is thus misapplied here, since metapleura can have
no place in mesothoracic terminology. The metathorax is
represented in the Muscoidea by the metasternum, whose
lateral wings are termed the hypopleure; and by the true
metapleura which is situated behind the hypopleura, the
metanotum being practically evanescent.

164 Annals Entomological Society of America _|Vol. VII,

The squamopleura exhibits characters in connection with
the hypopleura and the posterior thoracic spiracle which are
at times of importance. It therefore requires a special designa-
tion though it is not apparently a separate sclerite. It is
sometimes bare, often pilose or hairy, sometimes bristly, while
the position of the spiracle with reference to it and the hypo-
pleura may be used in the separation of groups among the
connectant forms.

Higher Posterior thoracic spiracle behind vertical axis of squamopleura
and Typical and hypopleura—Musca, bulk of Muscoidea, Leucomelina,
Muscoidea , Limnophora, Spilogaster, Gen. Indet., Fannia.

Anthomyioidea {Posterior thoracic spiracle squarely interposed between the
Connectant squamopleura and hypopleura—Muscina, Synthesiomyia,
Muscoidea Morellia, Aricia (last judged from figures).

VENATION

Fourth vein when complete or apical crossvein when present
reaching margin at or before extreme wingtip, the hind cross-
vein always joining fourth vein well before bend of latter or
origin of apical crossvein—Musca, bulk of Muscoidea, but

Higher and including only Cobboldia among the Oestride and allies.
Typical {Fourth vein always complete and reaching margin before wingtip,
Muscoidea | the apical cross-vein not present, the hind cross-vein prac-

| tically in line with the last section of the fourth vein—Glossina,
Cuterebride, Hypodermine, Oestrine.

Fourth vein incomplete, not reaching wing margin; apical and
hind cross-veins obsolete—Gastrophilus.

Fourth vein always complete, reaching margin behind extreme
wingtip, always bowed forward apically; no apical crossvein—
Connectant Stomoxys, Haematobia, Lyperosis, Hypodermodes, Eumesem-
Muscoidea brina, Muscina, Myospila, Clinopera, Pararicia, Leucomelina;
the last three with least forward bow to fourth vein and thus

most approaching the anthomyioid type.

Fourth vein not bowed forward in any part of its extent, but
Anthomyioidea often bowed backward apically—Limnophora, Spilogaster,
Fannia and other Anthomyioidea.

Of the above Haematobia furnishes an aberrant form of the
Stomoxys type, and Glossina an aberrant form of the Musca
type. Only Clinopera, Pararicia and Leucomelina are inter-
mediate between the Stomoxys and Anthomyia types. A
number of forms are intermediate between the Stomoxys and
Musca types.

1914] Muscoid and Anthomyioid Flies. 165

REPRODUCTIVE SYSTEM.

Higher
Muscoidea

least their rudiments visible; female usually with a large
number of ovarioles—All Muscoidea down to and including

Male with accessory glands always more or less developed, at
the Calliphorine.

Male without accessory glands, with very long and curled ejacu-
latory duct whose head is developed into a very elongate
vesicula seminalis; female without uterus, with many function-
ing ovarioles—Musca, Muscina, Synthesiomyia.

Typical

Muscoidea

not over 3 to 5 times as long as the vas deferens, with more
than half of it functioning as vesicula seminalis—Morellia,
Stomoxys, Haematobia*.

Male without accessory glands; female with only one or two
functioning ovarioles, one or two maggots or eggs developing
at a time in the uterus—Glossina,* Dasyphora,* Mesembrina,
Hylemyia.*

Male without accessory glands; female with few ovarioles,

Connectant depositing large eggs which hatch shortly into maggots omit-
Muscoidea ting the second stage and developing rapidly—Hypodermodes,
Eumusca,* Myospila.*

Male without accessory glands; female with few ovarioles,
depositing a small number of large eggs—Orthellia, Grapho-
myia,* Pyrellia.*

Male without accessory glands, with very long vas deferens
communis present and long vesicula seminalis; female with few
ovarioles, depositing a small number of large eggs—Leucome-
lina, Limnophora, Spilogaster*.

Male without vas deferens communis or accessory glands, with
very short ejaculatory duct—Gén. Indet.

| Same as preceding, but ejaculatory duct of male much shortened,

Male without vas deferens communis, with accessory glands,
Anthomyioidea with bulbous vesicula seminalis at head of the very short
ejaculatory duct—Fannia.

The term vas deferens communis is here proposed for the
slender tube present in some forms extending from the union
of the two vasa deferentia to the beginning of the swollen
and more or less elongate vesicula seminalis, and apparantly
not to be interpreted as a part of the ejaculatory duct. It is
short in certain Borboroidea (Paralimna sp. for example) and
Syrphoidea (Volucella sp.), but very long in Leucomelina
and Limnophora. It seems to be the homologue of the common
oviduct of the female, notwithstanding Berlese’s homologies
in his Gli Insetti (p. 841).

*Some doubt exists as to whether the 9 starred genera agree with the male
characters given.

166 Annals Entomological Society of America {Vol. VII,

HYPOPYGIUM

Worked out by Schnabl and Dziedzicki for a large number
of connectant as well as true anthomyioid forms. The char-
acters agree in a general way throughout the connectant
forms much as do those of the female reproductive system,
not appearing to furnish variations of sufficient scope for
definite separation into two main groups, except to mark off
the Coenosiide from the other forms. But they will doubtless
be of much use in the separation of small groups. The Calli-
phorine and higher muscoid groups need the same careful
study for comparison with the excellent results of these authors
on the forms which they have investigated.

EGG

Leucomelina, Limnophora and Spilogaster differ considerably
in egg structure from Musca, Muscina, Synthesitomyia, Stomoxys,
the Calliphorine and higher groups. They deposit a small
number of very large elongate eggs, either heavily striate
longitudinally or ribbed, or very minutely scaled-reticulate,
very slightly curved, yellowish-whitish in color, with thick
chorion, translucent-enameled in appearance.

It is probable that these approximate the characters of the
eggs of Orthellia, Graphomyia, Pyrellia, Myospila, Eumusca and
Hypodermodes, all of which deposit only a small number of
very large eggs.

The egg may be expected to furnish important characters
for the separation of the connectant forms.

MAGGOT

The position of the anterior spiracle in the third-stage
maggot of Fannia is quite in contrast to its position in Musca,
being situated well forward on the third segment. Whether
this holds good for Anthomyia and Coenosia is doubtful. The
first-stage maggot characters, especially the cephalopharyngeal
skeleton and anal stigmata, should differentiate the con-
nectant forms from the true anthomyioids.

1914] Muscoid and Anthomyioid Flies. 167

It appears from the foregoing that the most serviceable
characters for defining the natural boundary between Musca
and Anthomyia will be found in the egg, first-stage maggot,
male reproductive system, chaetotaxy and pilotaxy; while the
venation, thoracic sclerites, female reproductive system and
hypopygium will furnish supplementary characters of value.

The indications from the very incomplete data which it
has been possible to present are that Musca is much more
nearly related to Calliphora than to Anthomyia, but final
judgment must be reserved until all the main types con-
cerned can be investigated and the results compared and
correlated. The present meager notes will form a starting
point for an extended study of the subject.

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“CONTENTS OF THIS NUMBER.
roicns: Wa. op Ae ime Siay of} the on
Caterpillars: _ Iu, The mame Muscles- Che oc Le
WEtcH, PauL S. AO beenvations on the ite Hiner 4
and Habits of Hodenyes Confiuens Ast, ei
(Diptera). ....- nh oe te face ae = RSME recite 2 |
: CRAWFORD, ts ‘C. = Some Species of the Bee Genus
ry Coelioxys Aces Makan ee waaay ae ees eee
TowNsEND, Cece Hh 7. — ‘Ghana Fons: ie 3
Between the Muscoid and He See Sa Plies. “a
ie The regular inal sabsanption sities a ete the ae i
as the United States, Cuba, Porto Rico, Hawaii and Mexico, $3.00;
g Canada, $3.50; other countries, $4.00. Checks, drafts or money
(: orders should be drawn payable to ANNALS -ENTOMOLOGIC.
he SoclETY OF America, and addressed to Biological Bailding,
i State bees Columbus, Ohio, v. S. A.
bi
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