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ANNALS
OF
The Entomological Society of America
V f
VOEUME: ViV<19%4
433204
EDITORIAL BOARD
J. H. COMSTOCK, L. O. HOWARD,
IrHaca, N. Y. WASHINGTON, D. C.
Cc. J- S. BETHUNE, W. M. WHEELER,
GUELPH, ONTARIO, CANADA. Boston, MASss.
Cc. W. JOHNSON, P. P. CALVERT,
Boston, MAss. PHILADELPHIA, PA.
Vv. L. KELLOG, J. W. FOLSOM,
STANFORD UNIV., CAL. URBANA, ILLS.
HERBERT OSBORN, Managing Editor,
CoLUMBUS, OHIO.
PUBLISHED QUARTERLY BY THE SOCIETY
COLUMBUS, OHIO
CONTENTS OF VOLUME VII.
PAGE
TRIGGERSON, C. J.—A Study of Dryophanta Erinacei (Mayr) and its Gall.... 1
Cuitps, Leroy—The Anatomy of the Diaspinine Scale Insect Epidiaspis
Pract, (CELA CE as sie ait GEC ikke es SRP ici Dic! Ee BIO eae
GILLETTE, C. P.—Some Pemphiginae Attacking Species of Populus in Colorado 61
ZereK. J.——Wispersal of Musca Domestica ‘Linne.. 2.0.2. 05 6c o eect eee 70
housek o:—conwerntzia Hagent Banks. 5... ci feces be nent bole gee ees 73
Moore, Wm.—A Comparison of Natural Control of Toxoptera Granimum
MHeSOUEeAthICaL ang Unibedustatess.. 4h ase, <)cllen site seta teres iste eisiare = 77
Renmane lyr — i epore OF PaAtasit@St. ois nc425 cass coc eo vleseras ele eewaa cose 86
FERNALD, H. T.—Notes on Some Old European Collections.................. 89
MacGILLtivRay—Proceedings of the Atlanta Meeting................. Ree ntnare 97
Forses, Wm. T.—A Structural Study of the Caterpillars; III, The Somatic
IS CI AGH es ck Nisiic s Sioteoe Bone ee Ba ean ae Re ae AEA 109
We cu, PAut S.—Observations on the Life History and Habits of Hydromyza
Caniuensieocw.s (Wiptera)s. 2 .s.ecloen} sors Does Ae ee eee 135
CRAWFORD, J. C.—Some Species of the Bee Genus Coelioxys..............-++ 148
TOWNSEND, CuHas. H. T.—Connectant Forms Between the Muscoid and
Nth @ naytOLCarkull Coumreterrane seis se st el chewak estas ne eetatier are eiees 160
PETRUNKEVITCH, ALEXANDER—Spiders Collected by Mr. C. William Beebe
AAAI CESGNTC UTC Ey CLIT Ose eyes aio we Gotoh Semaai ave J a daa Saligeiaty Siete 169
SEVERIN, Henry H. P., SEVERIN, Harry C., HARTUNG, Wm. J.—The Ravages,
Life History, Weights of Stages, Natural Enemies and Methods of
Control of the Melon Fly (Dacus Cucurbitae Coq.)................. 177
PALMER, MirtAm A.—Some Notes on the Life History of Ladybeetles........ 213
ALEXANDER, CHAS. PAuL—On a Collection of Crane Flies (Tipulidae Diptera)
HROTaa HAVE dR pled SIE 06 Kee Sie Ae EA SS 3 a 239
GaHAN, A. B.—A New Species of Cheiloneurus with a Key to the Described
PEClESesnOmMeE tee W spore yes « -1- 5s Ae MRM Anse la tek avaiels ws ee ete 247
Tower, DAntEL G.—Note on the Number of Spiracles in Mature Chalcid
WAAC. Ret Piste cq taker tetris Uh: SA a Re NE oP Pairk a ath 2 249
Noyes, ALicE Ayr—The Biology of the Net-Spinning Trichoptera of Cas-
PEO SS) 2 (a eta ee 251
MosHER, EpnNA—The Classification of the Pupae of the Ceratocampidae and
JS SEPALS OC S52) cs ep i eS Cf So 277
BLOESER, WM.—Notes on the Life History and Anatomy of Siphona Plusiae.. 301
NEWCOMER, E. J.—Some Notes on Digestion and the Cell Structure of the
ieesnweseprtielutn i) Insegtse.- ge. ace-c- at cccle cece cee cece 311
BAUMBERGER, J. P.—Studies in the Longevity of Insects.................... 323
MetGrEcor,.E. A-Vour New Tetranychids.. 2) 2..5. 22... 0.6.5 ec eee 354
=n ; > iy . ), rn
a : ete rs Tiere ‘ ae oe ee 4
‘ 4 no Ws oN edt : tale: ees
‘. ¥ y fi yy p> | r >. £ , he .
2 a td . t P ¥ h a ;
> ; ‘ ad i :
Number 1.
ANNALS.
OF
The Entomological. Society of America
: MARCH, 1914
EDITORIAL BOARD
as oy COMBTocK, L. 0, HOWARD,
Eee eps ee 4 i Irmaca, N. Y. WASHINGTON, D.C.
| o J. S: BETHUNE; W. M. WHEELER,
he é GUELPH, ONTARIO, CANADA. Boston; Mass.
ag oe C: W. JOHNSON, P. P, CALVERT,
WRAL Gry Boston, Mass. PHILADELPHIA, Pa.
ee UN OLARELLOGG, : J. W. FOLSOM;
STANFORD UNIV., CAL. oy «URBANA, ILLS.
HERBERT OSBORN, Managing peters
One: OHIo.
PUBLISHED QUARTERLY BY THE. SOCI
ee) COLUMBUS, OHIO
eee
Lation a} Huse
"-Entered as sbéona class matter head H, 1908, at the Pea Office at Columbus, Ohio,
“under the Act of Congress of March 3, 1879.
The Entomological Society of America.
FOUNDED 1906.
OFFICERS 1914.
President-Ps Bs CaUVERT dij bogie Cc Kowa fait bs Philadelphia, Pennsylvania
First Vice-President—Jas. G. NEEDHAM.......... Vie siege Name be Ithaca, New York
Second Vice-President—C. GORDON HEWITT. 2.2... eee eee Ottawa, Canada
Secretary-Treasurer—A, D. MACGILLIVRAY... 2 ese e cece eee cee Urbana, Illinois
Executive Committee—THE OFFICERS and HERBERT OSBORN, W. M. WHEELER,
V. L. KELLOGG, NATHAN BANKS, E. P. FELT, J. M. ALDRICH.
Committee on Nomenclature—H. T. FERNALD, E. P. FELT, T. D. A, COCKERELL
Price List of Publications.
Annals, Vols. I, II, III, IV, V, and VI complete, each..............0eeeeees $3.00
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’ REPRINTS FROM VOLUME I.
Proceedings of first three meetings; Constitution, By-Laws and List of
I rea RIS ais sce iE as erie ttl wicca he eas ncn We 1h/Gs eid in’ CRIN Wh helace SF tas le Mani chd Aletheia wie 25
WHEELER, WM. M.—Polymorphism OP ARB ei ess. os seehind Ur naa he bieaeis cee. AO
OsBoRN, HERBERT—The Habits of Insects as a Factor in Classification..... .20
Severin, H. H. anp Severin, H. C.—Anatomical and Histological Studies
of the Female Reproductive Organs of the American-Saw Fly, Cimbex .
EPIC RNA i WOACU IE ii is VE copie gisne es kee snneall bie Wola eleahte hime nauk dey) Yow
Fett, E. P.—Some Problems in Nomenclature... .....0..0.0secedeeseeceece 10
Hammar, A. G.—On the Nervous System of the Larva of Cocytaln cornutaL .25
‘BrapvLey, J. C.—A case of Gregarious Sleeping Habits among Aculeate
REVINCHOMLETAL eave an Ul oc ogieh wi aeink nambly sealae’t p MPMinlels-o tar MAEM cae gale 10
Davis, J. J.—Notes on the Life History of the Leafy Dimorph of the Box-
elder Aphid, Chaitophorus negundinis Thos......... 00.00. bce eeeeeeees 10
HamBLetTon, J..C.—The Genus Corizus, with a Review of the North and
Middie ‘American, Species :y oc’. his e's win bie Kale wes Wiha be sla raed aialeigreaseiole wpe -25
Grrautt, A. A.—Biological Notes on Colorado Potato Beetle............... 25
Grrautt, A. A.—A Monographic Catalogue of the Mymarid Genus Alaptus.. .25
‘Severin, H. H. and Severn, H. C.—Internal Organs of Reproduction of
Nite Swe RY saliicia sins Se Galevid, sla Has Wn ehid ele cio Po kinle ety studs UW bi sigitic +15
Suitu, C. P.—A Preliminary Study of the Aranze Theraphose of California.. 75.
Davis; J Ji-—Studies on Aphididae. 300 0a dec estes Mae
Ruzy, W. A.—Muscle Attachment of Insects..........0.4. dciiadeiinaetine oe 15
NgEDHAM, J. C.—Critical Notes on the Classification of the’ Corduliine ‘
(57, tS EY Poe ter en oS ye OGRE eS eee a phere amr bird Core PR RL Ve Re 15
Howarp, L. O.—A Key to the Species of Prospaltella with Table of Hosts |
and Descriptions of Four New Species...... Be cieeia Voip uea wala alain cleats Rah tee
Hoop, J. D.—Two New Species of Idolothrips..............+ cocconeeceeesce, 010,
Address
ANNALS ENTOMOLOGICAL SOCIETY OF AMERICA,
Biological Building, State Univ., Columbus, Ohio.
ANNALS
The Entomological Society of America
Volume VII MARCH, 1914 Number |,
A STUDY OF DRYOPHANTA ERINACEI (MAYR) AND
ITS GALL.*
C. J. TRIGGERSON.
\
\
ian Ins#
‘ nsomian tuys >
os %
CONTENTS
1. Introduction.
2. The Life-History of Dryophanta erinacei. APR 23 1914
A. The agamic form. y
By The sexual form. ‘ Ce:
tonal Muse”
3. The Parasitic and Inquiline Life of the Gall.
A. Breeding Experiments and their Results.
B. Parasites and their Relation to the Agamic form otf Dryophanta
erinacei.
C. Inquilines, their Relation to Dryophanta erinacei, to the Parasites,
and to each other during Gall Formation.
4. The Stimulus to Gall Production.
A. The Relation of the Malpighian Vessels to Gall Formation.
B. The Relation of the Oenocytes to Gall Formation.
Conclusion. ,
Bibliography.
Dor
INTRODUCTION.
The Cynipidae constitute, biologically, one of the most
interesting families of the Hymenoptera. They have long
attracted attention, not only from the systematic view-point,
but also from the view-point of their life-history, the variety
of the galls they produce or inhabit, their biology, and the
cause of gall formation. The purpose of this paper is to pre-
sent an intensive study of one gall-maker Dryophanta erinacet
(Mayr), discussing its life-history, its parasites, its guests,
and the cause of gall formation.
The Oak Hedgehog Gall is rounded or oblong, with the
surface finely netted with fissures, and more or less densely
covered with spines. It varies in length from 10-15mm.,
and occurs on both sides of the White Oak leaf. The point
*Contribution from the Entomological Laboratory of Cornell University.
2 Annals Entomological Society of America [Vol. VII,
of attachment is generally on the midrib (Fig. I, Pl. I), though
it is often found on the lateral veins. When young it is yel-
lowish green, but in autumn it becomes yellowish brown,
much lighter in color than the tinting of the leaf. The gall
first appears late in June, and reaches full development about
the third week in August. It is widely distributed, having
been reported from New England, North Carolina, Iowa,
Illinois, Indiana, Kansas, Michigan, Ohio, Virginia, Canada,
and probably Florida and Colorado.
A longitudinal section through the gall shows that it contains
several chambers varying from two to eight in number. These
I have named according to their location. First to be noted
are the central cavities, (Fig. 2a, Pl. I), which measure 2mm. x
omm. and are located in the central portion of the growth.
These are occupied by Dryophanta erinacei and the parasites.
Second, there are the lateral cavities, (Fig. 2b, Pl. I), situated
at the side and base of the growth and measuring 114mm.
x 2mm. These are occupied by inquilines. Lastly, there
are to be found the peripheral cavities, (Fig. 8a, Pl. II), located
on the coriaceous portion of the gall, and covered with the
basal layer of spines. These are 1mm. in size, and are likewise
occupied by iniquilines.
The gall was first described by Walsh in 1864 under the
name Cynips q. erinacei. When Mayr in 1881 established
the genus Acraspis he included the insect causing this gall,
which therefore was known as Acraspis erinacei. The first
description of the insect appeared in a paper by Beutenmiiller
"09, entitled “Species of Biorhiza, Philonix, and their allied
Genera, and their Galls,’’ in which he places it in the genus
Philonix. As will be shown later in this discussion, the insect
belongs to the genus Dryophanta, and should be known there-
fore as Dryophanta erinacei (Mayr).
THE LIFE-HISTORY OF DRYOPHANTA ERINACEI.
The agamic form of Dryophanta erinacei emerges from
the oak hedgehog gall about the fifth of November. It varies
from 1.50 to 3mm. in length. The head is black, rufous
on both sides of the face, finely punctate, with whitish pubes-
cence; antennae black, fourteen jointed, with basal joints
rufous; thorax rufous; plurae black with rufous mark anteriorly;
all minutely punctate; parapsidal grooves distinct posteriorly,
1914] A Study of Dryophanta Erinacei. 3
obsolete anteriorly; scutellum rufous, punctate and pointed
posteriorly; metathorax black; abdomen piceous; ventral spine
and tip of abdomen hairy; legs yellowish rufous, tibia slightly
darker; wings aborted.
The insect makes its way to the leaf and flower buds of
the white oak, where oviposition takes place. On the tree
where our observations were made, it continued to emerge and
Oviposit until the twenty-first of November. The insects
are most active on cold days or early in the morning. During
the warm weather they are inactive and sluggish, hiding at
the base of the petioles, in the crotches of the young shoots,
or in the crevices of the bark. They have been taken in this
vicinity on rare occasions in early December, but usually they
succumb to the first heavy frosts at the close of November.
Its method of oviposition does not differ much from that
already described by Kieffer for other species of the Cynipidae
which attack buds. The insect clasps the apical portion of
the bud with the second pair of legs, (Fig. 3, Pl. I), and pressing
alternately with the first and third pair produces a teetering
motion which forces the ovipositor into the buds. The long
ovipositor lifts the apical edge of the outer scale, and is grad-
ually pressed down along the edge of succeeding scales, and
finally thrust into the region of the young leaf and flower.
Then there is a sudden jerk of the body which curves the distal
end of the ovipositor, turning the openings against the concave
face of the innermost scale. The insect now retains a motionless
attitude for almost four minutes, during which the egg is
deposited. The ovipositor is then withdrawn, the passage
being filled with a waxy substance for the protection of the
egg. This waxy secretion is doubtless from the accessory
glands of the reproductive system, and is homologous with
the secretions with which Corydalis cornuta, certain of the
Lepidoptera, as the Apple Tent-Caterpillar, the Tussock-
moths, and many other insects cover their eggs.
The egg, (Fig. 20, Pl. III), is an oval body 400u. x 225n.
provided with a pedicel which is lmm.inlength. It is attached
by this pedicel to the upper brown portion of the scale, falling
either against the green portion of the scale (Fig. 6, Pl. II),
or being held among the young leaves or flowers, in which
position it remains during the winter. It is worthy of emphasis
that this pedicel does not constitute the apical pole of the egg
+ Annals Entomological Society of America __[Vol. VII,
since the larva emerges from the opposite pole, and as already
indicated it serves as an appendage for attaching the egg to
the bud scale.
As will be seen later the eggs of the Chalcids are flask-
shaped, (Figs. 31, 32 and 37, Pl. IV), but here the elongate
portion is in reality the cephalic portion of the egg. This is
shown by the fact that the egg is oriented in the ovum in
such a way that the elongate portion is cephalad, also that
the larva always emerges at the base of the neck (Figs. 32
and 37, Pl. IV). This therefore, is exactly opposite to the
condition found in Dryophanta erinacet.
Buds were examined on the eighth of May and the
unhatched eggs found were very turgid, appearing slightly
enlarged. On the twelfth of May a slight swelling, at the
apex of which an empty egg shell was visible, appeared on the
lower green portion of the scale, (Fig. 9, Pl. II). This proved
to be a freshly formed gall, containing a young larva of
Dryophanta erinacei. The gall at this stage was thin-walled,
with a pebbled surface, greenish in color, and contained a
watery fluid. The egg-shell remains attached to the apex
of the gall until the latter has reached considerable size, when
it dries up and disappears. These hypertrophies develop
rapidly, as many as three appearing on one scale. The wall
of the gall has by this time changed to a yellowish brown
color, and soon becomes quite dry and brittle.
Galls also develop on the apical portion of the leaf and flower
buds, (Fig. 10, Pl. II). These are red, being similar in color
to the young leaf and flower. The wall is pebbled on the
surface, and thin. The cavity contains a single larva bathed
in a watery fluid, and similar in all respects to the one inhabiting
the scale gall. The terminal galls are of the same size as
those on the scales, varying in number from one to four, and
when mature are reddish brown. Since only the agamic
form of Dryophanta erinacei was found ovipositing on the
leaf and flower buds, and since the eggs of this species and no
other were found in the leaf and flower region, and since males
and females similar in size and character emerge from the
two galls, it is evident that they are produced by the same
insect. The difference in color in the galls is due to the normal
difference of the tissue of which they are formed.
1914] A Study of Dryophanta Erinacei. 5
Shoots were brought into the laboratory, placed in water
and covered with bell jars. Here about noon on the twenty-
first of May the first male and female emerged. They were
quite vigorous, and about four-thirty in the afternoon the
female was noticed actively moving along the midrib of the
young leaf. Suddenly she stopped, and set up a rapid nodding
motion which lasted thirty-five seconds, during which the
ovipositor was thrust into the tissue. The insect remained
motionless for a time, then withdrew the ovipositor, filling
the passage with a yellow substance which, as in the agamic
form, is probably a secretion poured forth by the accessory
glands of the reproductive system. The process was repeated
four times in succession without moving the body forward.
Each time the ovipositor was inserted the body was curved
slightly more than at the preceding puncture. The entire
time occupied by the four ovipositions was from four-thirty-
four to four-fifty, or sixteen minutes, thus allowing four minutes
to each oviposition of which a little over two minutes and a
half was occupied by the passage of the egg. Many other
observations were made, and the time in all instances cor-
responded to the first recorded.
While the first observations of oviposition were made
without having seen copulation occur, in all the following
instances it was observed. The male strikes the female several
times with the antennae, after which the latter rests quiet.
The male then clasps her thorax latero-caudad of the second
pair of wings with the second pair of legs, while the first pair
rest on the dorso cephalic portion of the thorax, and the third
pair extend slightly latero-cephalad of the abdomen; copulation
takes place, lasting for a few minutes.
The egg of the sexual form, (Fig. 25, Pl. III), is oval,
1604 x 450u. provided with a pedicel 750u. in length, which is
shorter than in the agamic form. It is always placed in the
fibro-vascular bundles, and at an angle of about 80° to the
axis of the leaf. The egg differs from that of the agamic form
only in the elongate portion being shorter.
The larva is characteristic of the Cynipidae, having a
slightly depressed head, fine needle-like mandibles, broad
thorax, and reflexed pointed abdomen. During development
the abdomen does not become as enlarged as in the agamic
form. The thorax also continues prominent throughout all
6 Annals Entomological Society of America [Vol. VII,
larval stages, which is not the case with the agamic form.
Fig. 27, Pl. III, represents a mature larva of the sexual form,
and may be compared with Fig. 19, Pl. III, which represents
a mature larva of the agamic form in the corresponding stage.
In the open the adults did not emerge until the twenty-ninth
of May, and continued to oviposit from that time until the
fifth of June. Oviposition here was as observed in the labora-
tory, the time occupied corresponding exactly to that already
noted. Fig. 4, Pl. I, shows a female of the sexual form.
The sexual form which possesses the characteristics of
the genus Dryophanta may be described as follows:
Female: Color. Head, thorax and abdomen shining black, nonpubes-
cent; mandibles yellowish brown; mouthparts yellow; antennae first
two joints yellowish, flagellum shading to black at the tip. In the
male the entire antenne are black. Length 2mm.
Head: Face opaque, surface irregular, rugose about the ocelli;
compound eyes 200yu.x300u. Distance between compound eyes and
hind ocelli 75u.; between hind ocelli 100u.; between compound eyes and
fore ocelli 500u. Distance between compound eyes and antennez 75x. ;
between antenne 75u. Width of head at temples 1.50mm.; mandibles
tridentate, mouth parts as in Figs. 21, 22, and 23, Pl. III. Antenne
fourteen jointed,(Figs. 28 and 29, Pl. III).
Thorax: Smooth, parapsidal furrows distinct posteriorly, obsolete
anteriorly, pluree smooth.
Scutellum: Rugose, becoming smooth in front, cross furrow
reduced to a shallow depression (Fig. 26, Pl. III).
Appendages: Wings hyaline, fringed with setz, veins yellowish
brown, (Fig, 24, Pl. III); legs yellowish, coxz of the third pair yellow-
ish brown.
Abdomen: Smooth, deeper than long, first segment one-third the
size of the abdomen, outline of the remaining segments as seen from the
side serrate; ventral spine and tip of abdomen hairy.
Male: Color. Same as female, length 14mm.
Head: Distance of compound eyes to hind ocelli 50u
“ between hind ocelli 150u
“ of compound eyes to fore ocelli 600u
“of compound eyes to antennz 50m
“between antennze 50u
Antenne fifteen jointed.
Thorax: Mesonotum more gibbous than in female.
Abdomen: Petiolated, longer than deep as seen from the side;
petiole cylindrical.
On the twenty-fifth of June the first evidences of gall-
formation appeared on the leaf-veins, the hypertrophied
tissue pushing through the slightly ruptured epidermis. The
1914] A Study of Dryophanta Erinacet. 7
embryoes obtained at this stage measured 125u.-130u. In
galls gathered on the first and second of July, larva were
found measuring 374. These were similar to the young
larva which give rise to the sexual form, having a slightly
depressed head, sharp pointed mandibles, broad, prominent
thorax, and pointed, reflexed abdomen. During the summer,
molts were observed after which the larva measured 500n,
750u., 144mm., 124mm., 2144mm., respectively, thus showing
five stages during the life-history. Fig. 19, Pl. III, shows a
larva 134mm., obtained about the middle of August. From
this time, the thorax does not show a great increase in size,.
but the abdomen loses its reflexed character, becomes globose,.
and increases in size until pupation. The first pupa was
obtained on the fifth of September, but the adults did not
emerge until the fifth of November. Fig. 5, Pl. I, shows a
pupa of the agamic form.
Thus we have another illustration of dimorphism in the
Cynipidae, the agamic form of Dryophanta erinacei developing
in the oak hedgehog gall on the white oak leaves, emerging and
ovipositing in the leaf and flower buds of the same tree, from
which, in scale and terminal galls, the sexual form develop.
These, emerging, oviposit on the veins of the white oak leaves,
and their offspring cause the oak hedgehog gall.
The Parasitic and Inquiline Life in the Gall.
The oak hedgehog gall is not merely the abode of the
maker, but also of several parasites and inquilines. In order
to obtain a knowledge of these, their mode of life, their relation-
ship to the maker, and to each other, we shall consider them
under the following heads:
A. Parasitic and inquiline life as shown by breeding
experiments.
B. Parasites in relation to Dryophanta erinacei and to
each other during gall development.
C. Inquilines, their relation to Dryophanta erinacei, to
the parasites, and to each other during gall development.
8 Annals Entomological Society of America [Vol. VII,
Parasitic and Inquiline Life as shown by Breeding Experiments.
The breeding experiments were of a twofold character.
First, galls were placed in cages, and the species inhabiting
them bred out. Second, larva were studied individually in
order to obtain larval characters, and then bred out and thereby
related to the adults.
I. Leaves bearing galls were gathered, separated from the
grass and other leaves, then placed in cardboard boxes at one
end of which a test-tube was inserted. In these tubes the
insects gathered, and were easily collected. The galls were
divided into two classes—those gathered when the leaves were
falling from the tree, and those subjected to snow, frost, and
general winter conditions for one and two months respect-
ively. The leaves were moistened once a week in order to keep
the galls from drying up, and thus preventing the adults from
emerging. The parasites appeared first, but the inquilines did
not emerge until the last of February.
The parasites, of which eight different species were obtained,
belonged to the family Chalcidide, and were all known to be
parasites in the oak galls. The inquilines belonged to the
Cynipide, genus Synergus. The following table will give the
various species, and the number of each obtained.
TABLE I.
Decatoma flava (Ashmead) 600 specimens
querci-lana-dorsalis (Fitch) 1 specimen
4 varians (Walsh) 30 specimens
Eurytoma studiosa (Say) 75
. auriceps (Walsh) 30 i
Ormyrus ventricosus (Ashmead) 150 “
Syntomaspis sp. 15 S
Tetrastichus sp. 10 4
Synergus erinacei (Bass. ) 70 is
II. The larve were removed from the galls, and studied,
and the larval characters determined. These specimens were
bred out in order to connect with the adult form. The method
employed was to note and tabulate such characters as the form
of the mandibles, the arrangement and size of the sete, and the
general larval form of a large number of specimens. Each
individual was then placed separately in a four dram vial which
was sealed and set in a dark place. Four species were bred
through to the adult. Both from the study of the larva in the
1914] A Study of Dryophanta Erinacei. 9
cavities, and the result of the breeding experiments, it was
evident that the parasites inhabited the central cavities, while
the inquilines, though occasionally found in this region, were
mostly confined to the lateral and peripheral cavities.
Owing to the small percentage of several of the parasites as
shown in Table I, we were able to breed out only five species.
The descriptions of the larval forms obtained are as follows:
Decatoma flava: The larval form of this Chalcid, (Fig. 34, Pl. IV),
when mature measured 14mm. It possesses slender bidentate man-
dibles, (Fig. 33, Pl. IV). The setz are short and spine-like, arising
from distinct, prominent tubercles, and are located as in diagram,
(Fig. 42, Pl. V), six on the head, ten on the prothorax, eight on the
mesothorax, six on the metathorax, and four on each of the abdominal
segments.
The egg, (Fig. 32, Pl. IV), is flask-shaped and measures 200u.xd0Ou.,
neck 560u., and pedicel 56u. It is pigmented, and becomes brownish
black on maturing. The long neck lies cephalad in the ovary of the
adult, and the larva emerges from the egg at the base of the neck,
(Fig. 32, Pl. IV). Thus the neck is not comparable to the long pedicel
of the eggs of the Cynipide. The short, crooked pedicel at the opposite
pole represents in atrophied form that found in the Cynipide.
Eurytoma studiosa and Eurytoma auriceps: The larval forms of
Eurytoma studiosa and Eurytoma auriceps are so similar that it is
impossible to determine specific characters. The one shown in Fig. 43,
Pl. V, always bred out to Eurytoma studiosa during the winter, but
during the summer larve corresponding in all respects to the diagram
bred out to both Eurytoma studiosa and Eurytoma auriceps. Hence
the general characters given here may be considered generic rather than
specific. Fig. 35, Pl. IV, gives a general view of this larva.
The mature larva measures 14mm., having bidentate mandibles
similar to that shown in Fig. 33, Pl. IV. The sete are long, slender,
with distinct tubercles, and give the body a very hairy appearance.
The general distribution of these, (Fig. 48, Pl. V), is twelve on the head,
ten on the prothorax, ten on the mesothorax, ten on the metathorax,
six on the first abdominal segment, four on each of the second, third,
and fourth segments, and six on each of the remaining segments. The
larva can be readily distinguished from the larva of Decotama flava by
the length of the sete, those of the latter being short, spine-like, and
fewer in number.
The egg is flask-shaped measuring 240u.xl44u., neck 720u., and
pedicel 64u. The neck lies eee nelae in the ovary of ae adult, and the
embryo emerges from the egg just at the base, (Fig ig. Pl TV) ihe
pedicel is short, curved, and aborted. The egg 1s apeaveeec! and
becomes black on maturity. It is quite similar in form to the egg of
Decatoma flava, but is slightly larger, and deeper in color when mature.
The egg of Ormyrus ventricosus is flask-shaped measuring 200u.x
120u., (Fig. 31, Pl. IV). In this Chalcid egg the pedicel is absent.
10 Annals Entomological Society of America [Vol. VII,
Synergus erinacei: The larva of Synergus erinacei, (Fig. 38, Pl. IV)
—summer brood—is fleshy, 1mm. in length, and possesses tridentate
mandibles, the second tooth of which is pointed like an arrow-head.
The setz are very small, difficult to locate, and without distinct tuber-
clesat the base. Their location, (Fig. 44, Pl. V), is fourteen on the head,
fourteen on the prothorax, twelve on the mesothorax, six on the meta-
thorax, four on each of the eight following abdominal segments, six on
the ninth, and eight on the tenth segment.
The egg is white, the body being kidney-shaped, 240u.x80u., and is
provided with a long neck 440u., Fig. 39, Pl. IV, shows one of these.
The larva of Synergus erinacei (spring brood) is dark, fleshy, 700.
long, (Fig. 40, Pl. IV). The mandibles are tridentate, the central tooth
blunt. The sete are minute, without distinct tubercles, and distrib-
uted as in Fig. 36, Pl. IV, eight on the head, sixteen on the prothorax,
ten on each of the meso- and metathorax, eight on each of the first three
abdominal segments, and six on each of the remaining segments.
The eggs, two of which are shown in Fig. 41, Pl. IV, are white. The
body is kidney-shaped 125yu.x56u., and provided with a neck 410u.
in length.
Parasites, their relation to Dryophanta erinacei, and to one
another during the development of the gall.
The Chalcids, Decatoma flava, Eurytoma studiosa, and
Eurytoma auriceps were observed ovipositing from June tenth
to the fourteenth. The method in all cases was similar, but the
time occupied during oviposition, and the number of eggs
deposited differed.
Decatoma flava selected a spot on the midrib where Dryo-
phanta erinacei had oviposited, thrust the long ovipositor down
alongside the same channel, and deposited an egg in contact
with that of the Cynipid. The ovipositor was then withdrawn,
and the opening sealed. This required three minutes.
Eurytoma studiosa and Eurytoma auriceps each selected a
spot about the region where Dryophanta erinacei had ovipos-
ited, and forcing the ovipositor into the fibro-vascular bundles
placed from one to six eggs near, but not in contact, with the
egg of the Cynipid. The eggs are usually laid in clusters, and
appear black in the tissue of the leaf. The opening is sealed
on the withdrawal of the ovipositor. The time consumed by
these two species in oviposition was four minutes each.
When the larva of Dryophanta erinacei emerges from the
egg, it proceeds at once to form a cavity which encloses the
eggs surrounding it. In newly-forming galls the cavity is
small, and the egg of the parasites is frequently found resting in
1914| A Study of Dryophanta Erinacet. i
the abdominal angle of the larva of Dryophanta erinacei. Here
it often hatches. The larva breaks the shell near the base of
the neck, (Fig. 37, Pl. IV.), and emerges, proceeding to attack
the host in the abdominal region. If the Cynipid larva has
just molted it is destroyed at once. If on the other hand, it
escapes the attacks of the parasites during this period, they will
live together until the next molt occurs, when the host is almost
invariably killed and eaten. Only on rare occasions have the
host and parasite been found living together in the same cavity
until both have reached 1mm. in length.
If two parasitic larve of the same or different species are
found in one cavity in the early stages, the stronger alone
survives, for | never have observed more than one adult emerge
from a single cavity. Since no Chalcid eggs are found in the
cavities inhabited by the inquilines, we may conclude that the
Chalcids are parasitic primarily on Dryophanta erinacei, and
secondarily on one another.
The larva of Eurytoma studiosa and Eurytoma auriceps
develop rapidly, and from the twenty-fourth of July to the first
of August adults emerge, thus giving a summer brood. No
adults of Decatoma flava emerge in the summer or autumn.
After the parasites have destroyed the host, it is questionable
whether they feed on the plant tissue, since the lining of the
cavity they inhabit turns brown, becoming hard and brittle
much earlier than is the case with the cavities occupied by.
Dryophanta erinacei.
It is impossible to determine absolutely the extent of para-
sitism in these galls, yet we gain some idea from the following.
During four weeks 1050 galls were examined, which showed
sixty per cent of parasitism not including the internal parasites
which had not emerged from the maker.
Inquilines, their relation to Dryophanta erinacet, to the parasites,
and to each other during the development of the gall.
The relation of the inquilines of this gall both to the host
and to the parasites is very interesting, since they are present
not only as guests, but also as parasites. The parasitic charac-
ter of certain species of Synergus has already been pointed out
by Moller and Mann, but nowhere have I found any record of
their singular action as observed in this gall. Synergus erinacei
is not only parasitic on Dryophanta erinacei, and the parasites
12 Annals Entomological Society of America [Vol. VII,
in the central cavities, but it carries its parasitic habits to the
extent of mining from cavity to cavity, and having a meal out
of the occupants. Fig. 12, Pl. Il shows where a larva of Syner-
gus erinacei has mined from A-B, also one is already breaking
down the wall at C. Fig. XI, Pl. II shows where a larva of
Synergus erinacei has mined from a lateral to a central cavity.
In all, over eighty instances of mining have been observed. On
eighteen occasions we have fed Dryophanta erinacei and dif-
ferent Chalcid larva to the inquilines, but only once were we
able to induce it to attack a larva of its own species. The
average time required by Synergus erinacei to consume a larva
was 144 hours. Hence we see that the supposed guest is not
only a plant feeder, but has shown itself to be a serious parasite
among the occupants of the gall.
The Stimulus to Gall Production.
A. The Relation of the Malpighian Vessels to Gall Formation.
Investigators have generally agreed that galls cannot be
produced apart from the presence of insects, but different
theories have been presented as to the cause of the abnormal
growth. Adler (1881) points out that in Neuroterus levius-
clus and Biorhiza aptera, the gall is first caused by the insect
wounding the surrounding cells with its fine mandibles, and that
the growth of the gall.is in some way dependent on the presence
of the larva. Cook (1903) in his publication ‘‘Galls and Insects
Producing Them,”’ states that the Cynipide stimulate the
plant to excessive growth by biting, and it is his contention
that gall formation is primarily the result of mechanical stimula-
tion. Rédssig (1904) in a paper entitled ‘‘ Von Welchen Organen
geht der Reiz zur Bildungder Pflanzengalle aus?’’ attributes
a rdéle to both oenocytes and the Malpighian vessels, and though
regarding the latter as giving off an effective secretion, he attrib-
utes the primary source of this secretion to the oenocytes.
His studies of the Malpighian vessels are purely from the
morphological standpoint, and are based on a limited and
poorly selected variety of species. He places great emphasis
on the size of the cells constituting the vessels, and on the size
of the vessels as compared with that of the larva. Moreover
he includes in his discussion species that do not produce galls
by means of any product poured forth by the Malpighian
1914] A Study of Dryophanta Erinacet. i:
vessels, since the galls are produced before the eggs are hatched.
Finally, he brings no evidence from a broad, comparative study
of the Malpighian vessels of various gall-producing species
to support his conclusion.
The condition found in the bud-gall from which the sexual
form of Dryophanta erinacei emerges is as follows: The egg
rests on the living portion of the scale. When the larva emerges
a viscous mass is adhering to it, but outside of this is a clear
fluid resembling the secretion of the Malpighian vessels both in
color and in action on glass when exposed to the air. The
young galls soon appear enclosing the larva. In this instance
the secretion of the Malpighian vessels appears to provide the
first stimulus to gall-formation. With the agamic form, where
the egg is enclosed in the plant tissue, one cannot observe the
process so easily.
On examining the galls of both the agamic and sexual forms
of Dryophanta erinacei, it was noticed that the cavities were
lined with growing tissue, abundantly supplied with chlorophyll,
also that where the larva of Dryophanta erinacei rested, both
it and the plant tissue were bathed at times with a colorless
fluid. When the larva was placed on a glass slide it at once
poured forth an abundance of this fluid, which always became
opaque, milky white, on drying. By varying the position of
the larva when placing it on a glass slide, this secretion was
seen to pour forth from the anal region, while the head and thor-
ax remained dry. About this time a study of sections of the
larva revealed two tubules consisting of four cells each, which
showed great activity. Longitudinal sections proved these to
be cells of the Malpighian vessels, (Fig. 45, Pl. VI.).
The Malpighian vessels of the agamic form of Dryophanta
erinacei consist of two long tubules containing fifty-six rounded
cells, with large nuclei, and attached to the hind gut, just at its
point of union with the mid-intestine, (Fig. 48, Pl. VI.). They
are whitish in color, the cells varying in size according to the
larval period. They reach their maximum in the fourth larval
stage. These larval tubules do not give rise to the adult
vessels, but, degenerating in the prepupal and pupal stages,
give place to the adult tubules which arise as evaginations of
the hind-intestine, just below the attachment of the larval
vessels. They are the largest glands in the body, extending
slightly ventrad along the mid-intestine, their cephalic ends
14 Annals Entomological Society of America _[Vol. VII,
reaching beyond the point of union of the mid and fore-intestine
in the region of the thorax, and are held in place by the fat
tissue. Ina longitudinal section they appear as in Fig. 47 Pl. VI
The individual cells consist of a homogeneous cytoplasm in
which vacuoles are found in the outer portion and also near the
nucleus. Secretions are sometimes seen in these. The nucleus
is irregular, and often greatly branched, sending long arms into
the cytoplasm. It is densely packed with chromatin granules.
The larva were mostly fixed in Dietrich’s fluid, and stained
with borax carmine and Lyons blue. This proved very satis-
factory for general work, but the best results were obtained
when the larve were fixed in hot Gilson’s fluid.
The cells are very active in secreting a colorless fluid duding
the period of gall-formation, which continues from the end of
June until the middle of August. At this latter time, the larva
has reached the fourth stage, measuring 134mm. There is
somewhat of an increase in the size of the cells up to this point,
which may be in proportion to the demand upon them. After
this there is a slight decrease to a constant size, which is retained
until degeneration begins. The following table shows the
increase.
TABLE II.
LENGTH SIZE OF CELLS
Larva 500u 64u.x72u. molt.
£ 750u. 64u.x80u. molt.
e lmm. 72u.x80u.
144mm. 72u.x82u. molt.
. 14mm. 72u.x88u. Z
« 134mm. 112u.x120u. molt.
o 2mm. 72u.x96u.
ae 214mm. 72u.x88u. molt.
246mm. 72u.x88yu.
As pointed out earlier in this paper, molts occur at 500u,
750n, 144mm., 134mm., and 214mm. It will be observed that the
cells reach their maximum at the fourth stage, about which
time the gall is rapidly maturing. From this time, less and less
secretion is poured out, and the linings of the cavities begin to
lose their green appearance, gradually becoming yellowish
brown, dry, and hard. Further, the larval form rapidly
changes. The abdomen increases in size, becoming globose,
while the head and thorax show only small increase. This is
in striking contrast to the early stages. A larva of the fifth
stage, measuring 214mm., when placed on glass or any foreign
substance excretes practically no fluid.
1914} A Study of Dryophanta Erinacet. 1
It was observed in all early stages that the excretion of the
fluid was under the control of the larva, being poured forth
freely when required. When a larva was found not feeding on
plant tissue the body was dry. When a parasitic larva rested
on the host both were bathed in a colorless fluid. If the larva
of Dryophanta erinacei was placed on a glass slide or foreign
substance, it immediately poured forth an abundance of fluid.
It was evident that a reserve must be retained in the tubules.
In a longitudinal section, (Fig. 45, Pl. VI.), it will be noticed
that the small proximal cells (indicated by V.) on each side of
the lumen are arranged so as to press closely against those
opposite. This formation appears constant throughout the
various stages, and we believe has a valvular function.
A number of tubules were dissected out from larva of the
earlier stages in normal salt-solution. This solution was
allowed to evaporate, and the salt crystals formed used in
grinding up the dried tubules. To the powdered mass a few
drops of normal saline were added, and when all was dissolved,
the fluid was filtered. The filtrate was treated with 85%
alcohol, and the action brought down a heavy, white, floculent
precipitate, which suggested that something of an enzymic
nature might be present. This phase of the investigation was
not pursued further at this time.
Fresh material was again obtained, the Malpighian tubules
dissected out as above, thoroughly dried, and ground with
powdered carborundum, which reduced them to a finer powder
than the salt-crystals. The powdered mass was dissolved in a
few drops of normal salt-solution and filtered. The filtrate
was injected with a hypodermic syringe into the midrib of the
white oak leaves, one drop being used to each puncture. The
operation was repeated three times on several leaves. Checks
were made, normal saline being used in these. The solution
containing the Malpighian tubule product penetrated from one-
fourth to one-half an inch in the fibro-vascular bundles of the |
midrib. The tissue was turned yellowish brown, and cracking
appeared similar to that seen in many young leaves where the
gall formation has just started, but owing to the death of the
larva, has ceased. While these experiments did not produce a
gall, they give suggestions as to the work performed by the
secretion of the tubules. Nothing of the above described
appearance was to be seen in the checks. E
16 Annals Entomological Society of America __[Vol. VII,
From a further study of Table II, it will be seen that between
the first and fourth larval stages there has been a considerable
increase in the size of the cells which constitute the tubules.
The greatest increase was between the third and fourth larval
stages, which was coincident with the greatest growth of the
gall, when it doubled in size. During this period the larva
gives off the greatest amount of secretion from the Malpighian
vessels. After this time, as already noted, the larval form
changes, and the amount of secretion diminishes rapidly, so
that a larva taken, say two weeks later, that is about the first
of September, would not pour forth any secretion when placed
on a foreign substance. The lining of the cavity is by this
time quite dry, brittle, and deep yellowish brown in color.
Now, from the development of the Malpighian vessels, and
the amount of secretion poured forth by them coincident with
the gall development, also in view of the effect of this secretion
when applied to the plant tissue in the experiments, it is evident
that the Malpighian vessels have elaborated some product
which when poured forth by the insect stimulates the surround-
ing plant tissue to rapid growth. In a few instances, we have
found urate crystals in the lumen of the tubules, but urates are
present in the Malpighian vessels of all insects, and, as Réssig
has shown, chemically pure urates do not produce galls. Hence
an additional factor is without doubt present in the secretion of
the Malpighian tubules of Dryophanta erinacei, and this
produces the effective stimulus.
The Malpighian Vessels of the Inquilines.
The Malpighian vessels of the inquilines were dissected out.
They were white in color, and consisted of two slender tubules.
These arise at the point of union of the mid and hind-intestine,
having a broader attachment than that found in Dryophanta
erinacei, (Fig. 60, Pl. VIII). The cells are smaller than those
of D. erinacei, the nuclei more regular, and the lumen quite
distinct. They show no evidence of great secreting activity,
and in a longitudinal section appear as in Fig. 61, Pl. VIII.
The larva when placed on a glass slide does not pour forth a
secretion as does Dryophanta erinacei. Further, the species,
though an inhabitant of a gall, does not emerge from the egg
until the gall has attained considerable growth. Its eggs are
1914] A Study of Dryophanta Erinacet. 17
rarely found in the central cavities, but generally in the rap-
idly growing tissue of the gall, where, after emerging, it forms a
cavity. Again the summer brood oviposit on the young galls
in July, laying their eggs just beneath the soft outer layer.
Here on hatching the larva forms a mere depression in the hard
portion of the gall, the soft outer layer forming the other wall.
It therefore gives rise to no gall formation. We have here,
then,’a species of the Cynipide, an inhabitant of a gall, appear-
ing after the stimulus to abnormal growth has been given, and
evidently not contributing toit. Its larval Malpighian tubules
are less developed than those of Dryophanta erinacei.
The Malpighian Vessels of the Parasites Inhabiting the Gall.
The Malpighian vessels of a Eurytoma larva, as dissected
out, are yellowish in color, larger than those of the inquiline,
and four in number. There are two long, clavate tubules
drawn to a point at their cephalic ends, and two short ones with
blunt ends, (Fig. 62, Pl. VIII). These arise at the union of the
mid and hind-intestine. The long tubules extend cephalad
slightly ventrad of the mid-intestine beyond the point of union
of the mid and fore-intestine in the thorax. The cells are
smaller than those of Dryophanta erinacei, the nuclei more
compact, and they do not give evidence of a high state of activ-
ity. Further, it must be remembered that the black eggs of
these parasites are found only in the central cavities, and never
in the tissue of the gall. Hence they are in a place where there
is no demand for gall formation. Again, the galls are well
developed, and the cavities of fair size before these emerge
from the egg. Therefore they do not give rise to a chamber,
as do both the maker and the inquilines. Moreover when
placed on a glass slide or foreign substance they do not excrete
a quantity of fluid, as do the larva of Dryophanta erinacei.
Finally, when they have destroyed the host, the cavity lining
loses its green, healthy appearance, passing from a yellowish
brown to a deep brown color.
Now, considering the habits of the parasites—that they do
not form a cavity, but occupy one already developed by Dryo-
phanta erinacei, and feed upon this species—also that the cells
of their Malpighian vessels do not give evidence of great
activity, we must conclude that the size of the tubules provides
no evidence that they produce a gall through their agency.
18 Annals Entomological Society of America [Vol. VII,
The larva of Decatoma flava also possesses four tubules.
Two are long, clavate, and drawn to a point at their cephalic
ends, and two are short with blunt ends, (Fig. 64, Pl. IX).
They are more slender, the cells smaller, and the nuclei more
regular than those of the Eurytoma larva. The general con-
dition stated regarding the former also applies to Decatoma
flava.
It is interesting to note that the galls that contain the high-
est percentage of parasites and inquilines, and those in which
the larva of Dryophanta erinacei have been destroyed at an
early stage never reach full development; also that the tissues
become dry, hard, and brittle. It is largely from this type of
gall that the summer brood of the inquilines, and of Eurytoma
studiosa, and Eurytoma auriceps emerge.
The Malpighian Vessels of other Gall-Forming Cynipide.
In the agamic form of Holcaspis globulus (Fitch) the
Malpighian vessels, as obtained from fresh material, are white
in color. They consist of two long tubules with large, globose
cells, and irregularly branched nuclei. They give evidence of
a high state of activity, and on contact with a foreign sub-
stance pour forth a fluid as did Dryophanta erinacei. In the
degeneration of the larval tubules and the development of those
of the adult they are similar to Dryophanta erinacei, (Figs. 54,
Pi Vil-and’ 57, and 538,7PL Vaile
The agamic form of Dryophanta polita (Bass.), which
causes the polished oak gall, possesses two Malpighian vessels
which as dissected out are white in color. They are smaller
than those of Dryophanta erinacei and Holcaspis globulus.
The cells are globose, nuclei irregular, and branched. Their
general action is similar to those already discussed. In a
longitudinal section they appear as in Figs. 55 and 56, Pl. VII.
The process of degeneration of the larval tubules, and the
development of the adult vessels correspond to that already
described for Dryophanta erinacei. Fig. 55, Pl. VII, is a longi-
tudinal section through a pupa of Dryophanta polita, which
shows the degeneration of the larval tubules and the adult
vessels forming.
1914] A Study of Dryophanta Erinacet. 19
The Malpighian Vessels of Gall-Producing Tenthredinide.
The larva of Nematus pomum (Walsh) which causes the
willow-apple gall was secured. Longitudinal sections through
the larva show cylindrical tubules arising at the union of the
mid and hind-gut, and extending caudad, (Fig. 66, Pl. IX). A
longitudinal section of the tubule is shown in Fig. 65, Pl. IX.
The cells are numerous, small, and regular, the nuclei being
symmetrical, and densely packed with chromatin. We have
no evidence here of great activity, nor does the larva secrete
any fluid when placed on a foreign substance.
Now it is known that the Tenthredinide do not produce
galls in the same manner as the Cynipide, but the stimulus is
given at the time of oviposition. Adler says “I have carefully
observed Nematus vallisnierii. The fly cuts into the tender
leaves of the end shoot of Salix amygdalina, and inserts her eggs
in the wound, frequently placing several in one leaf. At the
same time some glandular secretion from the insect flows into
the wounded leaf. A few hours after this injury, the leaf
surface presents an altered appearance, and new cell-formation
begins, freely leading to the thickening of the surrounding leaf
surface. After the elapse of about fourteen days the green and
red, bean-shaped gall is fully grown. If it is now opened the
egg will be seen lying in the cavity. Three weeks elapse before
the larva emerges from the egg.”’
Thus it is evident that the Malpighian vessels of the Ten-
thredinid larva are not factors in gall production.
The Malpighian Vessels of the Gall-Producing Diptera.
The larva of Trypeta solidaginis, from the globular gall on
the goldenrod, shows Malpighian vessels consisting of small,
round cells containing spherical nuclei, (Fig. 68, Pl. IX). The
cells show no evidence of exceptional activity, nor have we any
reason to believe that the Malpighian tubules are here factors
in gall formation.
The larva of Cecidomyia strobtloides, which causes the pine-
cone willow gall, likewise shows Malpighian vessels of a normal
type. The cells are small, very regular, and do not indicate
any unusual state of activity, (Fig. 69, Pl. IX).
20 Annals Entomological Society of America _[Vol. VII,
From a study of these two forms it seems probable that the
galls which they form are not due to any product poured forth
by the Malpighian vessels, for these neither secrete a fluid when
in contact with a foreign substance, nor do the cells show any
divergence from the normal type.
The Malpighian Vessels of Other Hymenoptera.
Since we have only considered species which form galls or
are associated as parasites and inquilines with the gall-maker,
it is necessary that we study related species that do not form
galls, in order that this comparative study may be more com-
plete. For this purpose we have selected species of Braconids
and Ichneumons.
A Braconid larva was obtained from the fall-webworm.
A longitudinal section of this larva is shown in Fig. 67, Pl. IX.
The cells of the Malpighian vessels are medium in size, and the
nuclei irregular. They are equal to those of the tubules of the
Chalcids, and larger than those in the vessels of the inquilines.
The Ichneumon larva was secured from the red-humped
apple-worm. Sections through the Malpighian tubules showed
that the cells were small and regular, the nuclei round. Here
we have a tubule the cells of which correspond in size to those
found in the tubules of the inquilines.
The Degeneration of the Larval Malpighian Tubules.
The degeneration of the larval tubules, and the development
of the adult vessels in the forms studied, are of such interest
that, though in part discussed by Rossig, they may here be
considered, and especially since the stages missed by Rossig
can be supphed.
Degeneration of the larval Malpighian tubules commences
in the prepupal stage. The cytoplasm shows huge vacuoles,
appears in shreds, and clings to the cell wall. The nucleus
becomes greatly elongate and branched, and chromatolysis
sets in. About this time small evaginations appear in the
hind-intestine, which develop into cylindrical tubules. Grad-
ually the cells of the larval Malpighian vessels break down, and
pass into the lumen of the hind-intestine, while the adult
tubules with small cells, and regular nuclei elongate rapidly.
1914] A Study of Dryophanta Erinacet. an
A series of the Malpighian vessels dissected out show this process
(Pigs, 45, 49, 52, and 53, Pls. VI, and VII).
The method of degeneration and the relation of the pha-
gocytes to this process has been in question. In the process of
degeneration as shown in Dryophanta erinacei, Holcaspis
globulus, Dryophanta polita, and in the Eurytoma larva, it is
clear that the phagocytes play no part whatever. The cells
break down and pass into the alimentary tract. Fig. 50, PI.
VII is from a cross section of a larva of Dryophanta erinacei,
and shows a fragment of a cell of a tubule found in the lumen of
the intestine. Fig. 51, Pl. VII is from a longitudinal section of
a similar larva, and shows the degenerating cell just breaking
away into the lumen of the intestine. Figs. 54, and: 57, Pls.
VII, and VIII show the degenerating cells of Holocraspis
globulus, also the adult vessels forming. Fig. 55, Pl. VII rep-
resents the same condition in Dryophanta polita.
The only instance where phagocytosis was found was in the
larva of Trypeta solidaginis. Here as shown in Fig. 76, Pl. X,
the phagocytes are present, but from a study of the slide it was
evident that chromatolysis had already set in, and the phago-
cytes were only of secondary importance in the degeneration
of the cell,
B. The Relation of the Oenocytes to Gall Formation.
We must now consider the relation of the oenocytes to the
production of the gall. Rdossig says ‘‘The oenocytes have a
certain influence, in that they in some manner break up the
blood fluid, and work it over in advance for the Malpighian
vessels.’’ This conclusion rests on the following: First, a
mere comparison of the size of the oenocytes with that of the
larva; second, ‘‘The general opinion that they are excretory
organs destined to store up urates, especially, as Verson has
shown in Bombyx, during the time when the Malpighian ves-
sells do not carry out their function, during molting and pupa-
tion. Berlese is of the same opinion.’’ Lastly, a possible
correlation in the development between the Malpighian ves-
sels and the oenocytes.
Rossig points out that the oenocytes of the various larva
which he has studied reach an unusual size. This growth is
attained within a short time, after which they shrink and
bo
bo
Annals Entomological Society of America _[Vol. VII,
gradually degenerate. Their size, in comparison with that of
the larva, is remiarkable. From some of the larva investigated
he gives the following:
LENGTH OENOCYTES NUCLEI
Larva of Biohriza terminalis A470u. 20u.
« “ Andricus ostreus 375u. 23u.
« « ~ Andricus fecundatrix 450u. 25p.
“« « Dryophanta divisa (end of June) 460y. 20-67. 25p.
“« “ Dryophanta divisa (17th July) 600u. 100u 50u
« « Dryophanta divisa (end of July) 78dy. 146-150z. 59u
From this, the author points out, first the unusual size of
the oenocytes as compared with that of the larva, and second
the three-fold increase in size of the oenocytes within one
month. He also gives measurements of oenocytes found in the
inquiline inhabiting the gall formed by Andricus globuli, Vespa
crabro, Nematus vallisnierii, Hormomyis fagi, and Aphis mali.
With one exception, the larva are all larger than those of the
gall wasps, and their oenocytes smaller. Now by a comparison
of the size of the oenocytes with that of the larva, and secondly
these oenocytes and larva with those of the gall-forming Cyni-
pide he endeavors to establish his theory.
In discussing the first point the author says “‘In no other
instance have such large oenocytes been found in so small a
larva.’’ Throughout his treatment of the oenocytes this fact
is kept continually in the foreground. Mention is made of
Kochevnikov’s discovery of a remarkable oenocyte 176y. in a
pupa of a honey-bee 15mm.—l6mm. in length, but the author
points out that, since the oenocytes in Dryophanta divisa are
so large as compared with the larva, and the latter so small .
as compared with the pupa of the honey-bee, (that is, 785.
as compared with 16mm.), great importance must be attached
to the oenocytes of the gall wasps.
Now does the size of the cell in comparison with that of the
body determine its importance? It is very doubtful if such
evidence can be used to support his conjecture regarding the
function of the oenocytes.
In the second place we must discuss the three-fold increase
of the oenocytes. This appears to be tabulated from one set
of larva in a single species, but is not shown to be constant
throughout that species. Now before a general conclusion can
be drawn, this triple increase would have to be shown to be
constant not only for a large number of larva of Dryophanta
1914] A Study of Dryophanta Erinacet. 23
divisa, but also in many species of the gall-forming Cynipide.
In the following table it will be seen that in Dryophanta erinacei
the oenocytes show no remarkable increase in size, and that
they reach their maximum long after the Malpighian vessels
have passed their period of greatest activity.
TABLE, LL:
D. ERINACEI. LENGTH OENOCYTES NUCLEI
pupa 56u.x64u. 26u.x32u -
64u.x64u. 32u.x382u-
48u.x48u. 24u,.x24y-.
larva 246mm. 80u.x120pu. 32u.x40p-.
64u.x 72u. 32u.xd2u.
48u.x 64u. 32u.x32u.
s 244mm. 64u.x 56u. 32u.x32u -
56u.x 56u. 24u.x24u-
56u.x 48u. Ay .x24y-
= 2mm. (2.x 76m. 32u.x32p.
56u.x 56u. 24u.x2Ap.
48u.x 48y. 24u.x24u.
¢ 134mm A8u.x 48u. 32u.xd2u.
40u.x 40u. 24y.x24y.
‘ 144mm. 56u.x 56u. 28u.x28y.
48u.x 48u. 24u.x24p.
“i 14mm. 48u.x 52u. 24u.x24y-.
48u.x 48u. 24u.x24y.
40u.x 40u. 24u.x24u.
= Ere, 40u.x 48u. 24u.x24u.
40u.x 40u. 24u.x24y.
s 500u. 40u.x 40u. 24u.x24u.
Eurytoma pupa 88u.x 88u. 40u. x48.
90u.x 96u. 48u.x48p.
80u.x S8Ou. 40u.x40p.
Eurytoma larva 134mm. 88u.x 80Ou. 40u.x36u.
64u.x 64u. A40u. x40u.
Synergus erinacei larva 134mm. 88u.x 72u. 32u.x32u.
56u.x 64u. 24u.x24y.
Internal parasite, larva 750y. 48u.x 42u. 24u.x24y.
40u.x 40u. 24u.x24y.
Nematus pomum larva 56u.x 40u. 24u.x24y.
40u.x 44u. 24u.x24u.
Trypeta solidaginis
larva 72u.x 80u. 32u.x32u.
60u.x 62y. 32u.xd2y.
In Table III, the measurements given are constant in a
large number of individuals for each species, and from%the
study of these it will be seen:
1. In Dryophanta erinacei the oenocytes reach their
maximum size when the- larva measures 245mm. Fig. 93,
Pl. XI shows such an oenocyte.
2. No oenocyte shows a threefold increase during the
larval development. The average of the largest oenocytes was
64u.x40u., nucleus 32u. while the smailest was 40y.x40u.
nucleus 24u.
24 Annals Entomological Society of America [Vol. VII,
3. The oenocytes of the second larval stage are as large as
those of the fourth stage.
4. Among the inhabitants of the gall the largest oenocytes,
(Figs. 83 and 84, Pl. X). were found in a Chalcid pupa, of the
genus Eurytoma, while in a larva 144mm. long of the same
genus the oenocytes were larger than those of a similar sized
larva of Dryophanta erinacei. Figs. 85 and 86, Pl. X, show
such an oenocyte. It is important to note that these occur in
a parasite, which, as pointed out earlier in this paper, does not
produce a gall.
5. A 134mm. larva of Synergus erinacei possesses oenocytes,
(Fig. 87, Pl. X), larger than those of a similar sized larva of
Dryophanta erinacei, yet Synergus erinacei is only an inquiline.
6. In an internal parasite of Dryophanta erinacei measur-
ing 750u. the oenocytes are as large as those of a similar sized
host. The parasite does not emerge from Dryophanta erinacei
until after the gall has attained full growth, and hence has no
part in the production of the gall.
7. The Tenthredinid, Nematus pomum, which develops in
a gall not produced by any product poured forth by the Mal-
pighian vessels possesses oenocytes of considerable size, while
the Malpighian vessels are normal, (Figs. 80 and 81, Pl. X).
8. The same is true of Trypeta solidaginis, an oenocyte of
which is shown in Fig. 82, Pl. X.
Now since the largest oenocytes are not found in Dryophanta
erinacei, but in a Chalcid parasite which is not a gall-maker,
since there in no triple increase in the oenocytes of Dryophanta
erinacei, and further, since there is a distinct limit within the
range of which the varying oenocytes of all species really fall, it
is clear that the conclusion of Réssig is not substantiated by the
present investigation.
“The general opinion that they are excretory organs des-
tined to store up urates, especially, as shown by Verson in
Bombyx, during the time when the Malpighian vessels do not
carry out their function, during molting and pupation.”’
In general, students of oenocytes have considered them
secreting organs, but as Perez has pointed out in “‘ Contributions
a l’ Etude de Metamorphosis’? we do not know what their
secretion 1s.
It is true that Verson considered the cells secretors of
urates in Bombyx, and Koschevnikov discusses the urate-laden
1914] A Study of Dryophanta Erinacet.
bo
Or
oenocytes of the honey-bee, speaking of them as permanent
reservoirs which could not free themselves of their products,
and had ceased activity. Berlese also describes oenocytes
containing urates in many of the species he has studied. Perez,
however, has pointed out that these workers have confounded
urate cells with oenocytes, and it is significant to note that
Berlese in his recent work ‘‘Gli Insetti’’ does not speak of
oenocytes bearing urates, but limits that function to urate cells.
The urate-bearing oenocytes that Rdossig describes in
Andricus Malpighi have the distinct marks of urate cells, and
are probably such. Further, he has shown that the injection
of chemically pure urates into the plant tissue gives negative
results, and therefore urates are not considered factors in gall
production. Suppose that we concede to the oenocytes the
function of secreting urates, have we gained anything?
According to the above quotation, the urate-secreting
function of the oenocytes is performed particularly during the
molting and pupating periods when the Malpighian vessels are
not functional. It has been shown that the oenocytes do not
secrete urates. In all the species we have studied, urate
crystals or urate globules have never appeared in the oenocytes.
In Dryophanta erinacei we have never observed any unusual
activity in these cells during the periods when the Malpighian
vessels are not active, and the same can be stated of Holcaspis
globulus, Dryophanta polita, Synergus erinacei, and the
Eurytoma larva. Moreoever we do not know the chemical
constitution of the secretion seen in the vacuoles of the oenocytes.
Therefore we have no reason for assigning to the oenocytes the
function of secreting urates.
A Possible Correlation in the Development of the Malpighian
Vessels and the Oenocytes.
In discussing this phase of the problem, Rossig states that
he cannot speak with certainty, but thinks that at least in
Dryophanta divisa a correlation exists between the development
of the Malpighian vessels and the oenocytes. He presents the
following tabulation:
MALPIGHIAN VESSELS OENOCYTES
LENGTH CELLS NUCLEUS CELLS NUCLEUS
D. divisa 460u. 50u. 36u. 50. 25 .
ee My 600u. 73y. 50pu. 100. 50 J
i Me 714y. 11dpu. 56u. 150z. 59.
26 Annals Entomological Society of America _[Vol. VII,
From this he concludes that the cells of the Malpighian
vessels are doubled in size while those of the oenocytes increase
threefold.
Now in comparing Table II and Table III, it will be seen
that in Dryophanta erinacei there is no indication of any cor-
relation existing between the oenocytes and the Malpighian
vessels. The oenocytes have reached their maximum when the
larva is in the prepupal stage, and the Malpighian vessels are
largest when the larva measures 124mm.—approximately the
middle of August. As far as our investigation has gone we have
found nothing to support the idea of any correlation in the
development of the oenocytes and the Malpighian vessels.
CONCLUSION.
The conclusions drawn from the foregoing study of Dryo-
phanta erinacei are as follows: ‘
1. From a study of the life-history of Dryophanta erinacei
we have another illustration of dimorphism in the Cynipide.
A. The agamic form of Dryophanta erinacei produces the
oak hedgehog gall on the veins of the white oak leaves, passes
through five larval stages extending over a period from the last
of June to the first of September. Pupation occurs on the first
week in September, and the adults emerge about the fifth of
November.
B. The adults oviposit on the leaf and flower buds of the
same tree.
C. The following spring the eggs hatch, and the larvae
produce galls on the leaf scale or the terminal growing points
of the buds, from which within two weeks the sexual form of
Dryophanta erinacei emerges.
D. These oviposit on the midrib and lateral veins of the
young leaves of the white oak. From the eggs deposited
emerge the young larve which produce the summer gall.
E. The sexual form belongs to the genus Dryophanta, and
will therefore be known as the sexual form of Dryophanta
erinacei, of which the insect, formerly known as Acraspis
erinacei is the agamic form.
2. The study of the parasitic and guest life shows that the
following insects inhabit the gall: Decatoma flava (Ashmead) ;
Decatoma querci-lana-dorsalis (Fitch); Decatoma varians
oa ag
1914] A Study of Dryophanta Erinacet.
bo
~J
(Walsh); Eurytoma studiosa (Say); Eurytoma auriceps (Walsh)
Ormyrus ventricosus (Ashmead); Syntomaspis sp.; Tetras-
tichus sp.; Synergus erinacei (Bass.)
A. The Chalcids are primarily parasitic on Dryophanta
erinacei, and secondarily on each other.
B. The inquiline, Synergus erinacei, is parasitic on the
entire life of the gall, mining from cavity to cavity and devour-
ing the larve they contain.
C. Eurytoma studiosa and Eurytoma auriceps, and Syn-
ergus erinacei have two broods. The spring brood appearing
June tenth to fourteenth, and the summer brood appearing
from July twenty-fourth to August first.
D. The percentage of parasites, not including the internal
parasites, is at least sixty per cent.
3. The Malpighian vessels of Dryophanta erinacei secrete
a fluid which stimulates the plant to produce the gall. This is
shown by the following:
A. The character of the Malpighian vessels of the sexual
and agamic forms of Dryophanta erinacei—their size, cellular
structure, and exceptional glandular activity.
B. The character and effect of the secretion poured forth
by the Malpighian vessels during gall formation.
C. The ultimate decline and ceasing of marked activity
of the tubules when the gall has matured.
D. The increase in the size of the cells of the Malpighian
vessels coincident with the development of the gall, and their
decrease in size when the demand upon them is withdrawn.
E. A comparison of the Malpighian vessels of Dryophanta
erinacei with those of the parasites and the inquilines found in
the gall, and particularly the lack of any abnormal secreting
activity in the latter.
F. A study of the Malpighian vessels of Holcaspis globu-
lus, and Dryophanta polita, both of which correspond in their
action, development, and degeneration to those of Dryophanta
erinacel.
G. A comparative study of the Malpighian vessels of Dryo-
phanta erinacei with those of Nematus pomum, Trypeta
solidaginis, and Cecidomyia strobiloides shows that all the
latter, though gall producers, possess tubules of normal type,
which do not pour forth an abundant secretion during gall
development, nor when in contact with foreign substances.
28 Annals Entomological Society of America [Vol. VII,
H. The study of the Malpighian vessels of species of
Braconids and Ichneumons, shows tubules with cells not larger
than those of the Chalcids and inquilines. The mode of degen-
eration however, appears similar to that found in Dryophanta
erinacel.
4. The theory of the relation of the oenocytes to gall
production as urged by Rossig is not confirmed by this study.
A. His argument is without support from the data furn-
ished by Dryophanta erinacei, Dryophanta polita, and Holo-
craspis globulus. The oenocytes of the Eurytoma pupa and
the larva, Synergus erinacei, are relatively larger than those
found in a larva of similar size of Dryophanta erinacei, while
the oenocytes of Nematus pomum and Trypeta solidaginis
were equal to those found in Dryophanta erinacet.
B. The oenocytes do not secrete urates. Perez has shown
this to be true in the ants, and Berlese appears to have now
accepted this view. In the oenocytes of the various species
studied, we have found no urate crystals or globules.
C. Since we do not know the chemical character of the
secretion in the oenocytes, and since there appears to be no
unusual activity in these cells during the molting and pupating
periods of these species under consideration, we are not con-
vinced that they take the place of the Malpighian tubules
during these periods.
Until we know the chemical character of the secretion pro-
duced by the oenocytes, we shall only deal in speculation as to
the réle of these cells in insect life.
In conclusion I wish to thank Professor Comstock and
Dr. W. A. Riley for the direction, and criticism so freely given,
and especially Dr. Riley, at whose suggestion the work was
undertaken, and whose assistance was invaluable throughout
the entire study.
2
x4
1914} A Study of Dryophanta Erinacet. 29
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1914] A Study of Dryophanta Erinacet. 31
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pp. 239-241.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Annals Entomological Society of America [Vol. VII,
EXPLANATION OF PLATES.
PiatTeE I.
Oak Hedgehog Gall attached to the midrib of a White Oak leaf.
Longitudinal section through a Gall. A. Central cavities; B. Lateral
Cavities.
Agamic form of D. erinacei ovipositing on the bud of White Oak.
Female, sexual form of D. erinacei.
Pupa of agamic form of D. erinacei.
PLATE II.
Bud scale with eggs of agamic form attached.
Bud scale with enlarged gall, egg-shell still attached.
Longitudinal section through Oak Hedgehog Gall showing Peripheral
Cavity.
Bud scale on which young gall is forming with empty egg-shell attached.
Terminal Galls.
Oblique section through Oak Hedgehog Gall showing where S. erinacei
has mined from A - B.
Longitudinal section through Oak Hedgehog Gall showing where S.
erinacei has mined from A - B, and is breaking down the wall at C.
PLaTeE III.
Mandible of Agamic form of D. erinacei.
Maxilla of Agamic form of D. erinacei. A. Cardo; B. Stipes; C. Palpus;
D. Galea.
Labium of Agamic form of D. erinacei. A. Palpus; B. Glossa; C. Para-
glossa.
Antenna of Agamic form of D. erinacei.
Two distal segments of antenna showing sensory pits.
Aborted wings of agamic form of D. erinacei.
Larva of agamic form of D. erinacei.
Egg of agamic form of D. erinacei.
Mandible of sexual form of D. erinacei.
Maxilla of sexual form of D. erinacei. A. Cardo; B. Stipes; C. Palpus;
D. Galea.
Labium of sexual form of D. erinacei. A. Palpus; B. Glossa; C. Para-
glossa.
Wings of sexual form of D. erinacei.
Egg of sexual form of D. erinace}.
Scutellum of sexual form of D. erinacei.
Larva of sexual form of D. erinacei.
Antenna of sexual form of D. erinacei.
Distal segments of antenna showing sensory pits.
PLATE IV.
Larva of internal parasite from agamic form of D. erinacei.
Egg of Ormyrus ventricosus.
Egg of Decatoma flava containing embryo.
Mandible of Eurytoma larva.
Larva of Decatoma flava.
Larva of Eurytoma sp.
Diagram showing location of setz on segments of larva of S. erinacei
(spring brood).
Eurytoma egg with larva emerging.
Larva of S. erinacei (summer brood).
Egg of S. erinacei (summer brood).
Larva of S. erinacei (spring brood).
Egg of S. erinacei (spring brood).
1914]
Fig. 42.
Fig. 48.
Fig. 44.
Fig. 45.
Fig. 46.
Fig. 47.
Fig. 48.
Fig. 49.
Fig. 50.
Fig. 51.
Fig. 52.
Fig. 53.
Fig. 54.
Fig. 55.
Fig. 56.
Fig. 57.
Fig. 58.
Fig. 59.
Fig. 60.
Fig. 61.
Fig. 62.
Fig. 63.
Fig. 64.
Fig. 65.
Fig. 66.
Fig. 67.
Fig. 68.
Fig. 69.
A Study of Dryophanta Erinacet. 33
PLATE V.
Diagram showing location of sete on segments of larva of Decatoma
flava.
Diagram showing location of setze on segments of Eurytoma larva.
Diagram showing location of setz on segments of larva of S. erinacei
(summer brood).
PLATE VI.
Longitudinal section through larva of agamic form D. erinacei showing
Larval Malpighian tubule, Valve, mid-intestine hind-intestine.
Longitudinal section through larva of agamic from of D. erinacei showing
larval Malpighian vessel, mid-intestine, hind-intestine, and hypo-
dermis.
Longitudinal section showing larval Malpighian vessel extending the
length of the mid-intestine.
Larval Malpighian tubules as dissected ovt from D. erinacei, agamic
form. R
Degenerating larval Malpighian tubules, and adult vessels forming, as
dissected out from D. erinacei, agamic form.
PLATE VII.
Cross section from D. erinacei, agamic form, showing cell of larval
Malpighian tubule in the lumen of the intestine, and adult vessel
forming.
Longitudinal section from larva of D. erinacei, agamic form, showing
cell just breaking away into the lumen of the intestine.
Larval Malpighian tubules of D. erinacei greatly reduced, and adult
vessels nearing maturity.
Larval Malpighian tubules of D. erinacei reduced to a few cells, adult
vessels well developed.
Longitudinal section of degenerating larval Malpighian vessel, with
adult tubules forming from H. globulus, agamic form.
A portion from a longitudinal section of a larva of D. polita, agamic
form. showing degenerating larval tubules, and adult vessels forming.
Longitudinal section through cells of degenerating larval Malpighian
vessels of H. globulus.
PLATE VIII.
Larval and adult Malpighian vessels of H. globulus as dissected out.
Portion of longitudinal section of larval Malpighian vessel of H. globulus.
Longitudinal section through degenerating cells of Malpighian vessel
of H. globulus.
Larval Malpighian vessels of S. ernacei as dissected out.
Longitudinal section through cells of larval Malpighian vessel of S.
erinacei.
Larval Malpighian vessels of Eurytoma larva as dissected out.
Longitudinal section through cells of Malpighian vessel of a Eurytoma
larva.
PLATE IX.
Larval Malpighian vessel of D. flava as dissected out.
Longitudinal section through cells of Malpighian vessel from Nematus
pomum.
Longitudinal section showing attachment of larval Malpighian tubules
to alimentary tract of N. pomum.
Section showing larval Malpighian vessels, and adult tubule just
appearing.
Longitudinal section through a portion of a larval Malpighian tubule
of T. solidaginis.
Longitudinal section*through a portion of a larval Malpighian tubule
of C. strobiloides.
34> Annals Entomological Society of America [Vol. VII,
Fig. 70. Longitudinal section through a portion of a larval Malpighian tubule
of D. erinacei, sexual form.
Fig. 71. Longitudinal section through a portion of a larval Malpighian vessel
of an Ichneumon.
Fig. 72. Across section of a larval Malpighian tubule of D. erinacei, agamic form.
PLATE X.
Fig. 73. Across section of a larval Malpighian vessel of the same species showing
degenerating cells. ;
Fig. 74. Cross section of an adult Malpighian tubule of D. erinacei.
Fig. 75. Cross section of larval Malpighian tubule of N. pomum.
Fig. 76. Cross section of larval Malpighian tubule of T. solidaginis showing
phagocytes.
Fig. 77. Cross section of a larval Malpighian vessel of C. strobiloides.
Fig. 78 and 79. Oenocytes from larva of N. pomum.
7
8
Fig. 80 and 81. Oenocytes from larva of internal parasite in D. erinacei.
2. Oenocyte from larva of T. solidaginis showing vacuoles.
Fig. 83. Oenocyte from thorax of Eurytoma pupa.
Fig. 84. Oenocyte from the abdomen of a Eurytoma pupa.
Fig. 85. Oenocyte from the thorax of a Eurytoma larva 134mm.
Fig. 86. Oenocyte from the abdomen of a Eurytoma larva 134mm.
Fig. 87. Oenocyte from S. erinacei containing vacuoles.
PLATE XI.
Fig. 88. Oenocytes from the abdomen of a larva of D. erinacei 14mm.
Fig. 89. Oenocytes from the abdomen of a larval of D. erinacei 244mm.
Fig. 90. Oenocytes from the abdomen of a larva of S. erinacei.
Fig. 91. Oenocytes from the thorax of a larva of H. globulus.
Fig. 92. Oenocytes from the abdomen of a larva of H. globulus.
Fig. 93. Oenocyte from the thorax of a larva of D. erinacei 244 mm.
Fig. 94. Oenocyte from the abdomen of a larva of D. erjnacei 244mm.
Fig. 95 and 96. Oenocyte from a larva of D. erinacei 2mm.
Fig. 97. Oenocyte from a larva of D. erinacei 14mm.
Fig. 98. Oenocyte from a larva of D. erinacei Imm.
ABBREVIATIONS.
rbd ge ts Re aig ee eek eae ei hs oxerte 0 Weeeh race oe NG a Cell of larval Malpighian tubule
a he Sart hage PP me ee iE Rh rks Lar am Mold Goma ai sed Foc. thos 2 E
CAE tet See Sone eee Cee ME SP. oa arin od Minin ena neigeotad op ad bso hind-intestine
(Eee ees iS Ob sa ATT GOO Sy Ey hypodermis
ETI NGS ee eeT RR ae I Oe Oo ae eee Imaginal Malpighian tubule
ands a oe hs BR a Eo ote ae eae oe leet Mandibles
eer a eee Ci een een iy Vee eer eA intact o Og oR LA ROIs SOS maxilla
aa; Ol) ee eet aS: ee re Ee etn Potro oot Tes Hes mid-dorsal
Te, PGT RE nse er eee ee tie ae ee errs rE SoS AM IAS aig jimh Oe mid-ventral
Ts ee ee ate poe serene Grin anh ac been te eto g Soe mid-intestine
set et ese UN a Ne om aN HS OSS cc SRD TOG Oar IEF ey: labium
| en awe Met ONT S Wencunee rl ae Sree Sa pO SO ote OC Oe ye CE lateral
| ee ee ee ee PORN MAE Se ORE EN Nebel Se ois ie acm set gO Thao lumen
(orale cata Sees era Ais nit sacar Paine See teach SOP a oF Larval Malpighian tubules
(0) Sa Rar a a are art ohn i vtie ae SaER t BOSS Se IRs ak eee eae enone oenocytes
pie S70) Rann ee ree ee ech 8 Say ee ee ee phagocytes
Se Bae shen Rees Ned oc tHe bei oak the tea Rea) eee scale galls
Sol Polo clea Rees Mee eet rake 8 Eine ab age Oe ee en eae sensory pits
ee en Nn oR RE Hee een ir, Arad TS. MOV ALA Boone Mey B98 sete
ie aan AA ARPES e ree er org hota ou ious J cy Chee So terminal galls
RT ae ERS ot eer APIS OER Ge ag Fe EG oO o~ vacuoles
ey
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a
THE ANATOMY OF THE DIASPININE SCALE INSECT
EPIDIASPIS PIRICOLA (DEL GUER.).
By Leroy CuHiLps, Stanford University, California.
The anatomy of several species of scale insects (Coccide)
has been studied in the entomological laboratories of Stanford
University, these species representing several different genera,
such as Physokermes, Ceroputo and Icerya. All these, however,
are of more or less generalized type and show but little marked
divergence from a common form. Comparatively little work
has been done in this laboratory or elsewhere on the anatomy
of the more specialized Coccide, the Diaspine.
It is the purpose of this paper to describe the more important
anatomical characteristics that are representative of the sub-
family Diaspinze as a whole. A knowledge of the facts of the
make-up and functions of the parts should add to the interest
of any study of the sub-family, whether the student have the
viewpoint of an economic or systematic entomologist. In
taking a particular member of the sub-family for this study, I
have chosen the species familiarly known as the Italian pear
scale Epidiaspis piricola (Del Guer). It is one commonly found
about the University and in the whole Santa Clara Valley.
In studying the anatomy of such a small and well chitinized
insect a number of difficulties of technic present themselves.
The first thing that must be considered is the method of killing.
Embedding and cutting were also features that demanded
considerable experimentation before desirable results were
obtained.
Three killing fluids were experimented with in particular;
Towers’ formula No. 2, Gilson’s, and hot water. The latter
proved to be the most useful. A fourth, Carnoy’s mixture of
six parts of absolute alcohol, three parts of chloroform and
one part of glacial acetic acid, was also used somewhat for
killing. It brought out the nervous system admirably, but
the other tissues were badly distorted or destroyed by the action
of this powerful agent.
The orienting of the material in the paraffine, ready for
sectioning, was a little problem in itself, on account of the
minuteness of the insect. The best results were obtained by
47
48 Annals Entomological Society of America __[Vol. VII,
the following means: A small paper receptacle was half
filled with melted 55% paraffine and allowed to cool consider-
ably. The insect was then taken up in melted paraffine in
a warm pipette, and dropped into the receptacle, which was
then filled. The specimen was then moved into desired position
with a warmed needle. After following this method of embed-
ding, little trouble was experienced in trimming up the block,
and the cutting could be done with accuracy in the plane
wished.
Necessary care must, of course, be taken in seeing that all
material is thoroughly embedded or the sections are very
liable to tear when the knife comes in contact with the tough,
chitinous covering of the insect. This chitin was found to
be extremely impenetrable, and all attempts at staining im
toto proved futile, though specimens were allowed to stay
in the staining fluid for days.
This paper was prepared in the Entomological Laboratory
of Stanford University, under the direction of Professor V.
L. Kellogg. I am indebted also to Professor Harold Heath
for suggestions during the work.
NERVOUS SYSTEM.
The nervous system (Plate XII, Fig. 4) of the Coccide
seems to vary little in structural characteristics in the many
widely differing groups of the family. The central system
consists of a fused, bi-lobed cephalic ganglion forming the
brain, and a prominent compressed thoracic ganglion from
‘which four pairs of lateral nerves are given off. The posterior
pair curve out towards the lateral margins of the insect and then
curve back again apparently fusing or at least giving off a
great number of smaller nerves which form a delicate fan-shaped
nerve center in the pygidium. The circum-oesophageal con-
nectives, (Plate XII, Fig. 4-a) are exceptionally long in Epi-
diaspis and, together with this lesser nerve center just mentioned,
represent the chief differences of the Diaspinz from the other
forms studied. This pushing forward of the brain and the
lengthening of the oesophageal commissures is undoubtedly
a result of specialization arising from the development of the
mouth-parts. The extremely well developed muscles that
govern the sucking apparatus are found just below the brain,
and it is undoubtedly their growth that has pushed forward
1914] Anatomy of Epidiaspis Piricola. 49
the more or less functional brain. However, a nerve can be
seen arising from either side of the lobes reaching out towards
the antennal rudiments.
Experiments were undertaken to ascertain the insect’s
sensitiveness to touch, which show that there is a decided lack
of visible response to any sort of stimulus. The only move-
ment that could be noted was that of the drawing in, or tele-
scoping, of the posterior region when touched with a needle.
This shortening takes place through the contraction of the
segments. No other movement of the body was observed in
response to other stimulants such as light, heat and water.
DIGESTIVE SYSTEM.
The Diaspine present an extraordinary arrangement of
the digestive system, diverging in this respect considerably
from the other sub-families. Dr. A. Berlese, the well known
Italian biologist, seems to be about the only man who has
done any considerable amount of work on this group, and
he reports a very novel condition of the system. He describes
elaborately in his studies, the arrangement of the organs of
digestion and assimilation, and finds that the stomach is
entirely disconnected from the intestine and the rectum.
This condition seems to be almost unbelievable. It is a con-
dition met with usually only in certain animal forms where there
occurs a regurgitation of undigestible foods. Such an action
is highly improbable among the Diaspine. Certainly no one
has ever observed this phenomenon among them and the
removal of wastes can probably be explained in another way.
The digestive epithelium of the stomach of Epidiaspis
is made up of very large cells (Plate XIII, Fig. 9-a) with cor-
respondingly large nuclei. The action of the digestive secre-
tions on the ingested plant juices is such that it reduces them
to a condition where they can be taken up by the blood-plasma
and used for food, reaching this medium by osmosis through
the walls of the blind sac or stomach. With the food also
passes that which is of no use to the insect and which is taken
care of by the exceedingly well developed Malpighian tubules,
of which there is a single very large pair, (Plate XIII, Fig. 9-b).
These excretory organs are fused, and at the very point of
fusion a short duct leads: into the rectum to which the
50 Annals Entomological Society of America [Vol. VII,
Malpighian tubes are attached by a filament (Plate XIII,
Fig.. 9-e) a short distance above the anal aperture. The
proportionately large size of these tubules indicates that their
function is not of ordinary or small proportions. They are
made up of exceedingly large granular cells, with distinct
nuclei, surrounding a thread-like lumen leading forward to
the fusion of the two tubules, from which there is a connection
into the tube leading into the rectum. These organs must
be’ considered primarily as organs of excretion and capable of
taking from the body cavity not only that material taken
in with the food, but removing from the system the waste
products of metabolism.
Dr. Berlese declares that there is absolutely no continuous
connection from mouth to intestine. However, the writer finds
some sections in his series that show what can hardly be denied
to be direct connections, (Plate XIII, Fig. 9-c). Berlese’s
theory of digestion and assimilation is quite plausible and
there is a good argument for its probability, for this connection,
at best, is very small. The greater number of the sections
that have been made—and I have sectioned several score
of specimens—show the condition as Dr. Berlese describes.
But it seemed possible that the result might be due to an
imperfect technic, so that to find a united alimentary canal
was the cause for cutting so much material. The results
seem to point to this accomplishment. I think that it is in
the killing that the trouble lies. We have to do with an insect
with a rigid, chitinous exterior with the anal aperture and
esophagus attached to chitin. The intermediate system pos-
sesses two large bodies, the stomach and the Malpighian
tubules, joined by a very delicate intestine, which, unable to
withstand the sudden shock of certain killing fluids, is ruptured,
with the natural result that many, if not most, of the insects,
show a disconnected digestive system.
This condition might possibly vary in the different Dias-
pinine genera, yet this is not probable. More work is still
to be done on the group as a whole, and should this finding
be true it will be a point of morphological importance at least,
though perhaps not altering the present accepted theory in
regard to the manner in which this system carries on its digestive
and secretive functions.
1914} Anatomy of Epidiaspis Piricola. 51
REPRODUCTIVE SYSTEM.
The female reproductive system of Epidiaspis (Plate XIV,
Figs. 14 and 15), is found to be characteristic of the usual
insect type, consisting of a pair of ovaries joined to the vagina
to form a figure much the shape of a capital Y. The vagina
is a rather long, thick duct, lined with prominent gland cells
with prominent nuclei, which undoubtedly secrete a tough,
shell-like material during the passing of the eggs. The vagina
opens on the ventral surface opposite the anal aperture. At
the junction of the two branches of the vagina is found the
minute opening of the seminal receptacle (Plate XIV, Fig.
14-b). This sperm sac is a long blind tube (Plate XIV, Fig.
16), and at the time observations were made was filled with
an exceedingly large quantity of sperm cells, which could often
be seen in the semi-cleared specimens under the microscope.
From the branches of the ovaries, masses of ovariole buds
are given off, varying in size from a mere evagination of the
egg tube to that in which the eggs are well developed (Plate
XIV, Fig. 14-a). Each of these ovarioles is capable of pro-
ducing a single egg, which, upon reaching maturity, passes
down the slender connective into the vagina (Plate XIV,
Fig. 15-c), and thence to the exterior. It is quite evident
that all of these buds do not develop, and I have noted that
the female apparently stops feeding to any great extent after
egg laying begins. Consequently after a certain number of
eggs have been deposited, she probably does not possess the
vitality to bring all the others to maturity. .
CIRCUMGENITAL GLANDS, OR SPINNERETS.
The study of the grouped glands or spinnerets (Plate XIV,
Fig. 17 to 21), and their histology and function, offered an
especial opportunity for some needed observations, and proved
to be very interesting. The histology and actual function
of these spinnerets have been subjects of much conjecture,
although little real work seems to have been done on them.
The reason for this is undoubtedly the fact that the glands
(Plate XIV, Fig. 17-a) are functional for a very short period
of time, and unless sections are made at this particular time,
no glands can be found in connection with the grouped orifices.
Sections made before the egg-laying period begins, cutting
52 Annals Entomological Society of America [Vol. VII :
squarely through these grouped spinnerets, often show a heavily
nucleated invagination of the hypoderm, but aside from this
no inkling as to the function can be ascertained. Just before
the insect commences egg-laying, however, a white powdery
substance can be found issuing from the openings, and sections
made from the material killed at this time brought out the
gland cells in their minutest detail. A slender duct (Plate
XIV, Fig. 17-c) is found to connect each of the circular openings
with the wax secreting glands. These units vary in number
of cells from one or two to six or seven, each division possessing
a prominent nucleus and uniting with the main duct by a slender,
thread-like lumen. Between these ducts are numerous elongate
supporting cells (Plate XIV, Fig. 17-b) which have no relation
with the functions of secretion, except that they may aid in
keeping the passageway open.
The spinnerets are circular, rather lens-shaped, and made
of chitin, through which a minute pore is found connecting the
cell and duct to the exterior. This opening seems always to
be uniform in its makeup—a rosette with five small parts.
Prof. E. E. Green is the author of a general rule which
can be applied to Coccids possessing these glands, that they
are for the most part ovo-viviparous, or egg-laying, while
those that do not possess these spinnerets are viviparous.
I have made observations on a number of the local species
and find the rule to hold true. I have also noted that the
embryo reaches a greater state of development before the egg
is deposited in the cases of those species in which there are
very few spinnerets, than in those that possess a large number
of grouped glands. In Aspidiotus uvae the circumgenital
glands are found in very small numbers, two or three in a group,
and the species is reported to be viviparous. The opportunity
of studying a large series of these insects might reveal some
very interesting facts. One can easily make a mistake in
examining material for this phenomenon by using adult females
which have died containing well developed eggs. The eggs
are not destroyed by the death of the mother, but from them
hatch young, around which is the shriveled skin of the mother,
and through which there is no means of escape for the newly
hatched insects. Such material when mounted impresses one
as being of a viviparous form. Circumstances of this nature
may be the explanation of some of the seeming exceptions to
the rule.
Ot
ee)
1914} Anatomy of Epidiaspis Piricola.
Upon the arrival of the egg-depositing season the female
assumes a different posture than is its earlier position, shorten-
ing itself to a considerable extent and appearing much more
rounded (Plate XIV, Fig. 21). On the insect’s taking this
shape the vulva opens directly back instead of on the ventral
aspect. With this change comes also a shifting of the normal
position of the spinnerets (Plate XIV, Fig. 19) to the position
shown in Fig. 20. In Epidiaspis the spinnerets consist of
five groups, three anterior and two posterior, all about equi-
distant from the genital aperture. This shifting from the nor-
mal, together with an enlargement and infolding of the vulva,
draws the glands to a position immediately surrounding and
lining the opening. The eggs passing through this opening
must necessarily pass over the glands, and in this passing
the moist egg picks up a quantity of the powdery secretion
that has exuded, more or less covering the newly deposited
eggs (Plate XIII, Fig. 21). This powdery substance not only
acts as a protection to the eggs, but also aids in keeping them
from drying and sticking together and thus blocking up the
limited space in which the insect has to store her ova.
THE MOUTH PARTS.
The mouthparts of the Diaspine are, as with thé other
subfamilies of the -Coccide, hard to homologize with those
of other insects. They are too minute to dissect and are
always flattened out of shape when mounted, thus making
a detailed description of the size and shape of the various
parts a difficult thing to do with accuracy. For the most part
this box-like framework of the Diaspinine mouth structure
can be homologized with that for other Coccide, as described
by Putnam, Mark, Moulton and others.
This framework is a very important structure in that it
serves aS a means of attachment for the powerful muscles
that govern the sucking and swallowing apparatus. In the
main, there are two arches (Plate XIII, Fig. 11) arcus superior
(a), and arcus inferior, (6), which fuse to make a very rigid
structure. Just below this box-like structure is a rostrum
or mentum, conical in shape and attached at the base. This
conical structure is covered with a chitinous layer and inside
of this are a few short muscles, which are undoubtedly used
54: Annals Entomological Society of America [Vol. VII,
in the manipulation of the long buccal setz, the four apparently
modified mandibles and maxillz, which are, in the case of this
group of insects, extended into a long piercing beak. These
four chitinous rods are arranged so that the plant juices pass
up through the tube formed by their union. Attached behind
the rostrum and lying free in the body cavity, is the setal
pouch, extending well back towards the posterior end of the
thoracic ganglion, into which the sete are pulled when they
are not in use.
At the base of the arcus inferior (Plate XIII, Fig. 11-b)
is found a very interesting apparatus made particularly striking
by its close resemblance to a piston valve, with all of its attach-
ments, chamber, rod and head (Plate XIII, Fig. 12). Figure
13 shows a diagramatic cross section of this valve, showing
inlet, 13-i, from salivary glands (Plate XIII, Fig. 11-d) and
outlet into oesophagus 13-o.
Here again, in the impossibility of actual observation of
the functioning, the interpretation of the actual function of
the organ has to be based upon its structural make-up and the
work that it apparently has to perform. Dr. Berlese is of the
opinion that this pump is used for drawing the saliva from
the large paired glands (Plate XIII, Fig. 11-e), and the arrange-
ment of the ducts and the openings into the cylinder would
seem to indicate this. Yet the rule for, most insects with
comparable organs is that these glands, as a result of an internal
pressure caused by a continual secretion of the cells from
within, or from muscular action, force the juices out. In
the case of this insect no muscles are to be found that could
perform this function, and, from the make-up of the glands
themselves, they seem to the writer to be admirably adapted
to operate through a pressure formed from within. Again,
from the make-up of the long, slender, four-pieced proboscis,
it would seem to be impossible to pump saliva into the plant
tissues, for pressure from the inside would disrupt the tube.
Necessarily, therefore, if there is a passage of fluid down this
setal arrangement it would have to be done with little or no
pressure. The presence of stained plant tissue at the point
of puncture possibly indicates that some fluid does pass, as
exemplified by the familiar reddish staining occasioned by the
presence of the San Jose scale (Aspidiotus perniciosus Comst.)
1914] Anatomy of Epidiaspis Piricola. 55
on various fruit trees. This possibly might be explained as
being the result of a mechanical stimulus and the disrupting
of the cellular make-up of the plant, for the long chitinous
‘ rods that compose the mouthparts pierce and destroy a great
many cells. This theory seems to be at fault however, in
that all scale insects do not cause this phenomenon; for
example, on the apple A. perniciosus causes a very distinct
reddening while EF. piricola does not.
The real function of the saliva is undoubtedly not to aid
in the taking up of the plant juices, but to act upon these
food properties after they have entered the insect’s body.
The relationship of the opening of the salivary glands to the
oesophagus would seem to point to this. The food is poured
into a common chamber at the base of the proboscis (Plate
XIII, Fig. 12-c) passing on through the slender oesophagus
(12-e) into the stomach.
The posterior part of the pharynx is a decidedly chitinized
structure, apparently valvular, to which a number of powerful
muscles (Fig. 12-f) are attached, and whose function it is
to help force the food forward, through expansion and contrac-
tion of the walls of the oesophagus. These muscles are attached
to the ventral wall of the insect, and undoubtedly act in con-
junction with the large retractor muscles found directly above
the pump-like cylinder of the mouthparts (Plate XIII,
Fig. 12-b).
In the main the long, slender oesophagus resembles a
capital letter U running forward from the chitinized pharynx
parallel to the ventral wall (Plate XIII, Fig. 12-e) of the insect
to about that point corresponding to the cephalic region of
the chitinous box-like framework of the mouth-parts. At
this point it turns toward the dorsal surface, passing between
the two circum-oesophageal commissures close to the base
Gfethe. brain. ~ Mere the tube turns parallel to the center,
emptying into the stomach at a point just posterior to a vertical
line that would run through the mentum (Plate XIII, Fig. 8).
This tube is cylindrical in outline and it may be distinguished
in sections from the surrounding tissue by the minute, circular
cells, of which it is composed.
56 Annals Entomological Society of America _[Vol. VII,
RESPIRATORY SYSTEM.
The Coccids possess a respiratory system that seems to
be nearly uniform through the entire group, and that found in ©
this species varies little from that of the larger representatives of
the family. It consists, in Epidiaspis, of two pairs of stigmatic
openings or spiracles, well guarded by hairs and spinneret-
like glands which excrete a powdery wax over the exterior
opening. These apertures (Plate XII, Fig. 6) possess
rosettes (6-b) that make them indistinguishable from those
that surround the vulva. A short tube leads from the stigma,
opening into a chamber from which numerous tracheoles’
radiate, always, however, in a definite way so that all the
individual insects of that species show the same character-
istics and design (Plate VII, Fig. 2). For the most part these
tubes are confined to the ventral surface, but can often be traced
into the body cavity, surrounding the different groups of organs.
The degree of diffusion of this system and its limit of ramification
could not be determined, as only those tubes that possess a
chitinization can be positively identified in the sections. In
the larger, less specialized forms of Coccids, taenidial rings,
the familiar characteristic of tracheal tissue are found, but
in this species no such rings were noted, though a careful
search for them was made. The characteristic trunk con-
nectives joining the spiracles are very small and as a rule
are unbranched. In some Coccide these have been found to
be wanting, undoubtedly the result of degeneration.
THE CIRCULATORY SYSTEM.
The scale insects as a whole seem to be lacking in anything
that may be called a definite circulatory system. No trace
of a dorsal vessel can be found and no movement or pulsation
of the body was noted that would indicate the presence of
any such system.
Pm Soa es
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bie
1914]
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
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12.
ig. 13.
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Anatomy of Epidiaspis Piricola.
EXPLANATION OF FIGURES.
PLATE XII. ;
Pygidium of adult female, dorsal surface with circumgenital orifice and
glands showing through.
Respiratory system of female insect, showing two pairs of spiracles.
Glandular spine, used in spinning the shell. (cross section shown in
Fig. 18-e).
Nervous system. a. Circumoesophageal commissures; b. brain; c.
thoracic ganglion.
Cross section of segments. a. chitin; b. muscle; c. muscle attachment.
Spiracle. a. opening; b. wax producing spinnerets; c. tracheal tubes.
PLATE XIII.
Sagittal-longitudinal section of adult female. a. stomach; b. malpighian
tubules; c. rectum; d. vagina; e. ovarioles; f. thoracic ganglion;
g. cephalic ganglion; h. salivary glands; i. sete (rostral); j. setal
pouch.
Digestive tract. a. stomach; b. malpighian tubules; c. intestine;
d. rectum; e. attachment filaments.
Characteristic muscle structure found under chitinous covering.
Mouth-parts and accessories. a. arcus superior; b. arcus inferior;
d. valvular pump; e. salivary gland; g. costa inferior; h. costa super-
ior; i. oesophagus (i. oesophagus drawn to one side); j. salivary ducts;
k. setae; m. muscle.
Longitudinal cross section of mouth-parts. a. sete; b. muscle; c. common
chamber at base of sete; the union point of oesophagus and duct
leading from pump; d. salivary duct; e. oesophagus; f. muscles attached
to base of oesophagus and to the body wall.
Diagrammatic cross section of pump showing inlet and outlet.
i. inlet; o. outlet.
PLATE XIV.
Ovaries, dorsal view. a. ovarioles; b. seminal receptacle; c. vagina;
d. vulva.
Ovaries, lateral view. (lettering the same as Fig. 14).
Seminal receptacle. a. sperm cells.
Sagittal section of gravid female cutting through caudo and cephalo-
lateral circumgenital glands. a. wax glands with neuclei; b. sup-
porting cells with neuclei; c. gland ducts; d. grouped gland orifices.
Cross section of spinning glands that form the shell. d. dorsal spinneret
with characteristic silk gland; e. spine-like marginal spinneret;
f. silk gland; g. gland found in connection with the silk gland and
supposed to secrete a cement-like fluid.
Normal position of the vulva and circumgenital glands of adult female.
a. vulva; b. anus.
Shift in position of the circumgenital glands of the adult female during
egg laying.
Position of adult female during egg laying. a. normal position indicated
by dotted line; b. position during the depositing of the eggs.
ANNALS E.S. A. VOL. VII, PLATE XII.
Leroy Childs.
VOL. VII, PLATE XIII.
ANNALS E. S. A.
576
Co ethe Pera
sae
PAT
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il
—
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Leroy Childs.
S £2 DRI
ANNALS E. S. A.
VOL. VII, PLATE XIV.
Leroy Childs.
SOME PEMPHIGINAE ATTACKING SPECIES OF
POPULUS IN COLORADO.
(Concluded from Vol VI, p. 493.)
By C. P. GILLETTE.
Thecabius populiconduplifolius Cowen, Plate XV, Figures 1 to 9.
Pemphigus populiconduplifolius, Cowen, Hemiptera of Colorado, Bull. 31,
Colo. Exp. Sta., p. 115, 1895. Hunter, Aphidide of North America, p. 79, 1901.
Gillette,* Jour. Economic Ent. p. 355, 1909. Jackson, Cols. Hort. Soc. Vol. 22,
p. 217, 1908. Contributions No. 29, Dep. Zool. and Ent., O. S. U., p. 217, 1908.
Pemphigus ranunculi n. sp., Davidson, Jour. Economic Ent. p. 372, 1910.
Pemphigus populiconduplifolius, Davidson, Jour. Economic Ent. p. 374, 1910.
Pemphigus californicus, Davidson, Jour. of Economic Ent. p. 414, 1911.
Pemphigus populiconduplifolius, Essig, Pom. Jour. Ent. p. 699, 1912.
Pemphigus californicus, Essig, Pom. Jour. Ent. pp. 699, and 700, 1912. Essig,
Pom. Jour. Ent. p. 827, 1912.
Pemphigus populiconduplifolius, Patch,* Bull. 218, Maine Exp. Sta. p. 76, 1913.
The above literature may be briefly summarized as follows:
The original description by Mr. Cowen dealt with the alate
fundatrigenia in the folded leaves of the cottonwoods with the
mere mention of yellow apterous individuals, all from Colorado.
Hunter lists this species only.
The writer, in 1909, recorded the species from Massa-
chusetts.
Davidson, 1910, described the alate and apterous forms
taken in California from the buttercup (Ranunculus Caltfor-
nicus) to which he gave the name ranuncul1, but which is prob-
ably populiconduplifolius, as Mr. Bragg and Mr. Asa C. Maxson
have repeatedly traced this species to the buttercup in Colo-
rado, where it seems to be perfectly at home. On page 374 of
the same paper Davidson records populiconduplifolius in the
folded leaves of Populus trichocarpa and mentions seeing the
stem mother.
*These can hardly be populiconduplifolius, as the stem females were reported
in both cases as being present in the colonies of developing lice, a condition which
we have never found in Colorado where the types of the species were taken. Fur-
thermore, I have a stem female from Massachusetts that was taken by Mr. L. C.
Bragg, and it is readily distinguishable from any of the stem females that I have
seen from Colorado by having remarkably thickened femora for all legs. The
femora are very nearly twice as great in diameter as they are in the Colorado form
and are of about the same length. Four winged migrants taken from Populus
balsamifera (Acc. No. 47-10) in Maine by Dr. Edith M. Patch are before me,
mounted in balsam. These seem to differ from Colorado examples principally
by having weaker sensoria, which are also fewer in number, on the sixth joint of
the antennz. I will suggest that this eastern form be known as Thecabius patchit,
though it does not have the typical habit of most known examples of this genus of
having the stem mother in a gall by herself.
6.
62 Annals Entomological Society of America [Vol. VII,
Jackson merely quotes Cowen’s original description.
Davidson, finding the name ranunculi preoccupied, suggests
californicus instead.
Essig lists this species both as populiconduplifolius and
californicus.
Dr. Patch records several captures of this species in Maine
on the leaves of Populus balsamifera and gives a figure showing
the distribution of the wax glands of what she took to be the
stem mother, and also an excellent figure of the antenna of the
alate fundatrigenia, or summer migrant.
This is a rather common but not abundant species in north-
ern Colorado and the writer has also taken it at Wheatland,
Wyoming, upon a broad-leaved cottonwood.
Fundatrix, Figures 1-4.
The general color of the fundatrix is yellowish olive green, lightest
over the middle area of the abdomen, more or less covered with a white
powdery secretion and a few wax threads about the lateral margins and
posterior portions of the body; head, eyes, antenne and legs, including
coxee, black or blackish; in general form, broad oval; eyes small but very
prominent; length 3.75 to 4.50, and width 2.50 to 3.00; hind tibia, .60;
length of antenna about .70; 5-jointed; joints I, II and IV subequal in
length, the fourth being a trifle the shortest; joint III barely as long as
IV and V together with the spur; spur about half as long as joint IV;
only permanent sensoria present and they are bordered with cilia. The
arrangement of the wax plates upon the head and thorax is shown in
figure 3, and is about as follows:
Head with a pair of large circular plates on the vertex between the
antenne; on the occiput a similar pair, not quite so large but somewhat
wider apart, and midway between these a smaller pair, rather close
together; on the prothorax four large plates in a transverse row, and
two small ones in front of the middle pair; meso- and meta-thorax and
joints 1-6 of the abdomen, each with a transverse row of 6 plates; joint
VII with 4 plates; joint VIII with 2, and none on joint IX. It is not
uncommon for two of these plates next each other to coalesce and so
reduce the number.
The fundatrix is never found in the leaves folded along the
midrib in which the other lice occur, but is always found in a
narrow fold on the margin of one of the first leaves to open
and upon the under side, see figures 1 and 5a. The second
generation, almost as soon as born, leave the pseudo-gall of the
fundatrix and travel to the tenderest little opening leaves at
the tip of the twig, where they locate, several to a leaf, upon the
lower or ventral surfaces where they begin to feed, causing the
ew
1914] Pemphigine Attacking Populus In Colorado. 63
leaves to fold along the midrib as shown at figure 5, b. Mr.
Maxson in a letter makes the following statement in regard to
his observations upon the early occurrence of the second brood:
“The first larve of the second generation were observed
June 18th. These were traced to the young leaves at the tips
of the branches where they located on the underside. These
leaves began to fold along the midrib and in a few days typical
P. conduplifolius galls were formed.”’
This second brood all acquire wings and leave the cotton-
woods and go to the buttercups, Ranunculus sp., so far as our
observations go. At the present writing, December 24th,
there are several thrifty colonies in the laboratory on butter-
cups where they have been since the migrants were put upon
the plants in July. They attack, not the roots, but the crown
and leaves and stems near the ground. The buttercup seems
to be a permanent food plant for this species, upon which it
seems to be able to live continuously throughout the year.
COLLECTION DATA FOR FUNDATRIX.
Specimens in the collection have been taken as follows:
Ft. Collins, Colo., 6- 2-13, L. C. Bragg, Populus occidentalis
; “ 6-12-13, L. C. Bragg, 2 -
= S “ ~ 6-16-13, L. C: Bragg, “ i
. . “6-20-13, L. C. Bragg, . A
Denver, Colo., 6-25-13, L. C. Bragg, q us
Fundatrigenia.
The winged fundatrigenia taken from the folded poplar
leaves is the form described by Mr. J. H. Cowen in Bulletin 31
of the Colorado Experiment Station, page 115, as follows:
‘““Length 1.8 to 2.2mm. Alar expanse 6.85 mm. Nearly black,
pruinose. The abdomen is deep green when the glaucus matter is
removed by placing the insect in alcohol. Antenna 1.00 mm. long,
joints slender, 5th and 6th with about six or seven annulations each;.
stigma short and broad; ungues usually with a constricted neck. Similar
to ramulorum but larger and the antennal joint not nearly so strongly
annulated.”’
In addition it might be said that the transverse sensoria (Figure 7)
_are not complete rings, many of them extending but a short distance,
and especially is this true on joint III; on joint IV the number commonly
varies between five and eight, and the same is true of joint V, while the
number on joint VI is usually six besides the terminal or permanent
one; spur, finger-like and about .05 in length, or about one-half as long
as joint II; joints I and II equal in length, the former cylindrical, the
64 Annals Entomological Society of America [Vol. VII,
latter larger at distal end, each measuring .07; near the proximal end of
- joint III on the front side is a short tooth or spine; wings (Figure 6) clear,
veins slender, stigma rather small. Described from seven types on one
slide taken at Boulder, Colorado, June 23, 1910, by Mr. L. C. Bragg,
along with a large number of co-types.
COLLECTION DATA FOR FUNDATRIGENIA.
Ft. Collins, Colo., 7-11-12, L. C. Bragg, Populus Sp.
% § to 8- (12, bb. Ca Brace,
& s “« 8 6-12, C. P. Gillette, “ a
: € een fal iia Ore 22 Gillette, . ss
Wheatland, Wyo., 7-15-13, C. P. Gillette, “ “
Boulder, Colo., 6-23-10, L. C. Bragg, c
Boulder, Colo., 7— 4-11, L. C. Bragg, $
Newcastle, Colo., 7-23-93, C. P. Gillette, “ us
Greeley, Colo., 7-21-09, C. P. Gillette, “
Lynn, Mass., 6-27-09, L. C. Bragg, € e
La Salle, Colo., 7-20-09, C. P. Gillette, “ &
Eckert, Colo., 6-26-10, C. P. Gillette, “ x
Windsor, Colo., 7-29-93, C. P. Gillette, “ «
Niwot, Colo., 7-15-08, C. P. Gillette, “ «
Grand Jct, Colo. /—21-93 Ces Gilletie; == “g
Veazie, Maine, 7- 5-10, E. M.-. Patch, “ ‘
Delta, Colo., 7-20-93, C. P. Gillette, “ «
Niwot, Colo., 7-15-08, C. P. Gillette, “ «
Paonia, Colo., 6-23-10, C. P. Gillette, “ :
Mr. Maxson has given me the following statement in
regard to the early appearance of the stem mother gall:
‘“The earliest date on which the galls were found was May
20th. On this date the lice were very small and appeared to be
in the first stage after the egg, since no shed skins were to be
found in the galls.”’
Alate Sexupara.
This form differs from the fundatrigenia by having joint II of the
antenna (Figure 8) decidedly longer in proportion to its diameter;
joint VI without transverse sensoria; joint V often without sensoria
except the permanent one, but sometimes with one, two, or even three
sensoria present; joint IV with but four or five sensoria, and joint V as
long as joint VI with the spur.
This form has been taken at Fort Collins on several different
dates during September on Ranunculus by Mr. Bragg, and
about Longmont, Colorado, by Mr. Asa C. Maxson.*
*T am indebted to Mr. Maxson for the privilege of using the data that he has
accumulated during the past two years from his studies of this insect. Mr. Maxson
has also independently traced this insect to the Ranunculus from the cottonwood
leaves.
1914] Pemphigine Attacking Populus In Colorado. 65
Young larve.
The rather young larve of this species taken on Ranunculus are
pale green in color and are heavily covered, especially over the abdomen,
with fluffy wax threads. For full descriptions see Davidson’s paper in
Journal of Economic Entomology, 1910.
Pupz on Ranunculus, pale yellow in color and very heavily covered
with wax threads, especially over the abdomen.
Compared with A ffints.
This species is very close to affinis Kalt. of Europe, but
seems to be distinct, and especially because of what seem to be
rather marked differences in the antenne. I have but a single
European example of the alate fall migrant, sent me by Mr.
J. J. Davis, from Dr. Tulgren, of Sweden.
I have found only six wax plates on the vertex of the funda-
trix in twelve examples examined, while Tulgren gives eight
for affinis; joint IV of the antenna is somewhat shorter in pro-
portion to the other joints as compared with affinis in Tulgren’s*
figures, and joint V in the fall migrant commonly has two or
three sensoria, while affinis is represented as having none.
Sexupara of affints.
We have a good example of affinis from Europe, sent by
Dr. P. van der Goot. Joints V of the antenne have each one
good transverse sensorium near the middle and one very small
sensorium besides; joint IV has four well developed transverse
sensoria on each antenna and is decidedly club shaped, being
much heavier at the distal end. The same form of condupli-
folius has joint IV more nearly cylindrical and usually with
four well developed transverse sensoria, joint V usually with
none but sometimes with one or two small sensoria, and joint
VI with none.
The antenna of the virgogenia seems to agree with that of
affinis as figured by Tulgren.
Asiphum sacculi n. sp., Plate XV, Figures 10 to 14.
I first saw the gall of this louse about eighteen years ago
when.on a mountain trip some twenty miles or more northwest
of Fort Collins. I did not meet with it again until the present
summer, when, on July 13th, I found two of the leaf pockets
characteristic of this species in Estes Park on twigs of Populus
tremuloides about six miles'apart and at an altitude of about
*Aphidologische Studinen for Zoologi, 1905, Band 5, No. 14.
66 Annals Entomological Society of America [Vol. VII,
7500 feet. On August 9th, four more of the galls were taken
near the Half-way House on Pike’s Peak, at an altitude of about
9000 feet. In two of these galls there were no living lice. In
each of the occupied galls taken (four), there was one large
stem-female present, with a large number of her offspring in all
stages of development up to the adult alate lice. There were
no adult apterous lice and all that were half grown or more gave
evidence that they were to get wings. In the breeding cages
the alate lice began at once to deposit young with long beaks.
In every case the lice were accompanied by a species of large
black ant.
The Gall. An infested leaf becomes very much enlarged,
and somewhat thickened with the edges turned in so as to make
a heart shaped pocket, and the apex of the leaf is extended and
turned back as shown in Plate XV, Figure 10. The color of the
infested leaf is yellowish green, and lighter than the healthy
foliage surrounding it.
Fundatrix. Figures 11 and 18.
A very large, oval, slatey gray louse, lightly covered with a fine
white powder, and set everywhere with delicate gray hairs above and
below; length of body about 4.50; width 4.00; antenna, .75; joint III
longest and almost as long as joints IV and V together; permanent
sensoria ciliated, joints I, II, and III with numerous delicate hairs;
beak barely attaining third coxee; hind femur .80; hind tibia .80.
Pupa. Figure 12.
The pupz are quite dark in color, the abdomen being very dark
olive green and the head and thorax a rather blue slated gray; the tarsi,
eyes and terminal joints of the antenne black. A conspicuous marking
of the larve and pupe consists of a row of five tufts of white waxy
secretion along either lateral margin of the abdomen.
Fundatrigenia.
All the young of the fundatrix, the second generation, become
winged and leave the galls. General appearance, that of a black louse;
abdomen olive green; thorax, head, and antenne blackish, or dusky;
legs yellowish; tarsi dusky; wings a trifle smoky; the veins slender with
a narrow dusky line on either side; stigma narrow, lanceolate, dusky,
fork rising about midway on the cubital vein; length of body 3.50;
wing 4.50; antenna (Figure 14) 1.00; joint III with 7 to 9 oval sensoria
and a well developed spur near the base; joint IV, with two similar
sensoria near the distal end; joints V and VI with the usual permanent
sensoria only which are ciliated about the margins; joint III longest, |
fully as long as joint VI with the spur; joint V slightly longer than joint
IV; cauda a broadly rounded lobe.
Habits for the remainder of the year unknown.
1914] Pemphigine Attacking Populus In Colorado. 67
Mordwilkoja vagabunda Walsh.,* Plate I, Figures 15 to 20.
Byrsocrypta vagabunda Walsh, Proc. Ent. Soc. Pha., V. I, p. 306, 1862.
Pemphigus vagabundus, Walsh and Riley, Am. Ent. V. I, pp. 57 and 107, 1869;
Riley, V. I, Mo. Rep. p. 120, 1869; Packard, Guide to Study of Insects, p. 524,
2nd. Ed. 1870; Thomas, Ent. Rep. Ill., V. I, p. 153, 1880; Oestlund, Aphids Minn.
p. 22, 1887; Packard, Forest Insects, p. 434, 1890; Osborn, Cat. Hemip. Ia. p. 130,
1892; Cowen, Bull. 31, Colo. Exp..Sta. p. 116, 1895; Hunter, Aphid. of N. A., p.
79, 1901; Cook, O. Nat. V. IV, p. 118, 1904.
Pemphigus oestlundi n. sp., Cockerell, Ent. News, p. 34, 1906.
Pemphigus vagabundus, Jackson, Genus Pemphigus, Cols. Hort. Soc. XXII,
p. 200, 1908.
Mordwilkoja oestlundi, Davis, William’s Aphidide of Neb. p. 4, 1911; Patch,
Bull. 213, Me. Exp. Sta. p. 100, 1913.
The galls of this louse at the terminal buds of cotton-
wood twigs have occurred in greater or less abundance in at
least one limited locality near Fort Collins, Colorado, for
the past fifteen or more years. The section referred to is
mostly rather low, moist land, along the course of an irrigating
ditch and near the river. It seems strange that the galls
should not have become more generally distributed unless the
alternate host is largely limited to the area mentioned. I am
assuming that there is an alternate host for the reason that
the lice all become winged and leave the galls rather early in
the summer. Most of them are gone by August Ist here.
The Galls, Figures 18, 19 and 20.
When growing, the galls are as green in color as the cottonwood
leaves, and are, in fact, a transformed leaf in each case. On the inside
of the green gall the main veins of the leaf are very prominent.
Apparently these galls differ from others produced by Aphids by not hav-
ing any opening to the exterior during their growth, but Mr. L. C. Bragg
has discovered a small brown scale (Figure 18 A, and 20, b) at the base
of the gall, which seems to be the apex of a folded leaf, beneath which
is an opening to the interior and through which the blade of a pen-
knife may be passed without cutting any tissue. This opening is so
narrow that the lice do not escape by it. About the time that fully
matured winged lice are developed in a gall (about July 10th to 15th,
*Professor Oestlund, in his Aphidide of Minnesota, p. 22, states that Walsh's
vagabundus is evidently something different from the louse that has since been
known to be associated with the coxcomb gall. To be sure, September is late
to take the migrants from these galls, and the measurements given by Walsh are
too large for this species. But he evidently had a louse belonging to the Genus
Pemphigus, as then understood, and in the Walsh-Riley paper published in Vol.
I, of American Etomologist, page 107, the vagabond gall was figured, and both the
winged lice and the apterous stem mothers from the galls mentioned. As Walsh
at that time considered the winged lice from these galls the same as what he had
described as B. vagabunda, it seems to me best to abide by his identification of
his own species, and especially as we do not know any other species to which
to refer his original description, which is quite inadequate for its identification
anyway. I am therefore retaining the name vagabunda.
68 Annals Entomological Society of America [Vol. VII,
at Fort Collins), little star-shaped mouths (Figures 18 and 19, 0)
appear at the apices of the more prominent lobes thru which the alate
lice escape. Many of them appear on a single gall as shown in Figure
18. The galls nearly always are from terminal buds, and whether
one or more of the leaves form a single gall I have no certain knowledge,
but apparently it is one in each case. Large galls measure as much as
SO to 95 mm. in greatest diameter, and about 50 to 60 in the greatest
thickness.
The Fundatrix. Figures 15 and 16.
One fundatrix was found in every gall opened. General color
of body yellowish green, the yellow tinge seeming to come from a large
number of small embryos within; nearly unicolorous throughout,
but a little darker along the lateral margins; legs and antennz yellowish,
the antenna blackish at tip; both antennz and legs very short; joints
of antenna 4; joint III about one-half the entire antenna in length;
spur two-thirds as long as joint IV; permanent sensoria ciliated.
Length of body 5; width 3.70; constricted a little at base of abodmen.
In general, stout, pear-shaped, the small end being at the head; covered
everywhere with a light covering of white powder.
Young lice and pupe in the galls are very pale yellowish and much
powdered. There are a great number, possibly a thousand in all,
in a gall and apparently the stem females were just in their prime and
packed full of embryos on July 11. All of the offspring get wings
when adult. ;
Fundatrigenia, Figure 17.
All alate lice with black head thorax and antenne, and dark green
abdomens, and the thorax and abdomen are heavily pruinose with
more or less cottony threads towards tip of abdomen, making a tuft.
Length of body 2.50 to 3.00 mm.; antenna .85; joint III as long as
joints IV, V and VI to base of spur; joints IV and V sub-equal; spur
about equal to joints IV and V together; a distinct spur near the base
of joint III; sensoria on transverse ridges, but not surrounding the
joints; joint III with nine to eleven sensoria; joint IV, two; joints
V and VI with permanent sensoria, only; spur with two or three sen-
soria, usually three, scattered along its length; cauda very small,
rounded.
Described from lice escaping from the galls July 11, 1918.
ANNALS E. S. A.
VOL. VII, PLATE XV
C. R. Gillette
1914] Pemphigine Attacking Populus In Colorado. 69
EXPLANATION OF PLATE XV.
(G3PACinre TE)
Plate XV. Figures 1 to 9, Thecabius populiconduplifolius Cowen: 1, stem mother
gall on margin of leaf; 2, young adult stem mother; 3, showing wax plates on head
and thorax of stem mother; 4, antenna of stem mother; 5, cottonwood twig showing
the folded terminal leaves where the young from the stem mother gall have set-
tled at b, and a stem mother gall at a; 7, antenna of fundatrigenia, or second gen-
eration lice; 8 antenna of alate sexupara or pre-sexual form; 9 antenna of apterous
form found on Ranunculus during the summer and fall.
Figures 10 to 14, Astphum sacculi, n. sp.; 10, pocket-like gall from Aspen;
11, fundatrix or stem mother; 12, pupa; 18, antenna of the fundatrix; 14, antenna
of the fundatrigenia.
Figures 15 to 20, Mordwilkoja vagabundus Walsh: 15, fundatrix; 16, antenna of
fundatrix; 17 antenna of fundatrigenia; 18, the vagabond gall containing fundatrix
and young; 18, a, brown scale beneath which there is an opening into the interior
of the gall; 19, a section of the gall showing oscula (0) that open for the escape
of the mature lice; 20 (b) the closed mouth, also shown at 18, a.
Drawings by Miss Caroline M. Preston, except Figures 1 and 2, which are by
Miss M. A. Palmer. Original.
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DISPERSAL OF MUSCA DOMESTICA LINNE.
JAMES ZETEK, Ancon, C. Z.
The latter part of May, 1913, unusual numbers of flies
appeared at the Isthmian Canal Commission hotels and com-
missary at Balboa, Canal Zone. An inspection revealed a
pile of cow manure, etc., about 800 square feet in area, located
at one corner of Ancon Cemetery, 2,500 feet distant 1n direct
line from the hotel. This place is indicated by the letter ““B”
on the accompanying map. This manure was heavily infested
with maggots, principally of Musca domestica, Hermetia allucens,
Volucella obesa, and Paralucilia macellaria. Puparia were very
abundant and adults in countless numbers. This was the only
manure pile found away from the incinerators.
EAST BALBOA
CANAL ZONE, PANAMA
SCALE
Sin sath). taker
\ vo Probeble I 10
fy
|
i (PN
fh
y y gy) Se fea eee
2
3
4
edu: IMPROB 45,
Fic. 1. Map of the region.
A pit was dug, into which was placed a small quantity of the
infested manure. A cover of earth one foot deep was added as
a protection from heat rays. Over this was placed a screened
cage. On June 6th the cage contained about five thousand
adult flies, most of them the common typhoid-fly. At 9:00
a. m. these were sprayed with an aquaeous solution of gentian-
7°
1914] Dispersal of Musca Domestica Linne. 71
violet, to which a small amount of gum tragacanth had been
added. (See Annals Ento. Soc. Am. Vol. VI, No. 1, pp. 5-21).
These marked flies were liberated at 9:30 a. m. at the same place.
At the hotels, commissary and a few private houses, tangle-
foot paper was used freely, and this was afterwards collected
by the writer and examined for the presence of any marked
flies. The method used was to wet each fly with a solution of
alcohol and glycerine. The marked fly became known by the
resolution of the tiny crusts of anilin dye adhering to its body.
Seventeen marked flies were thus recovered, viz:
East Balboa Hotel, 4 &@ Musca domestica from paper ex-
posed for 32 hours after the sprayed flies had been liberated
at the cemetery.
Profile , “B line from
breeding place to hotel.
Impro bable path
> 2500 ft.<
Fic. 2. Profile in direct line.
Spanish Mess, 9 (7 o&, 2 2) typhoid flies from paper exposed
32 hours.
Commissary, 3 o& typhoid flies, from paper exposed 32
hours.
Spanish Mess, 1 & typhoid fly, from paper exposed 75 hours
(in reality 43 hours aftér last batch of paper was collected.)
No attempt was made to catch flies outside of these screened
buildings, hence the 17 recovered adults represent only such as
had gained entrance through holes in the screening or while
the doors were opened and closed. The species found on the
fly paper were mainly Musca domestica and Hermetia illucens,
both of which breed freely in pit closets as well as in manure,
and which were very plentiful upon cooked and uncooked food.
72 Annals Entomological Society of America [Vol. VII,
The probable path taken by the flies in this particular case
was down the East and West gully along the cemetery, then
along the lowlands to the Commissary and Hotels. The
profile figured is made in a direct line from the breeding place
to the hotel, but it seems hardly probable that this was the
path actually taken. The former is the more likely one.
Thus it has been clearly shown that a mass of manure 2,500
feet from the hotels and 150 feet above them, was a menace to
these places. The experimental results were augmented shortly
after when this manure heap was completely destroyed. The
flles at the Commissary and hotel quickly diminished
in numbers.
CONWENTZIA HAGENI BANKS.
Life History Notes and Variations in Wing Venation.
By J. S. Houser, Ohio Experiment Station.
The species under consideration was first called to the
attention of the Department of Entomology, of the Ohio
Experiment Station, in November, 1912, by Mr. J. M. Keck,
of Cleveland, who submitted the overwintering cocoons for
determination. Dr. MacGillivray, to whom specimens were
referred, pronounced them as belonging to the neuropterous
family Coniopterygide. <A _ little later, reared adults were
sent to Mr. Nathan Banks, who gave the specific determination
as Conwentzia hageni Banks.
At the time the material was taken the insect was in the
larval stage, neatly encased in the double cocoon so well
described by Quayle (Bul. 234, California Experiment
Station). The gross appearance of the cocoon so closely
resembles the compactly woven webs of some of the smaller
spiders that it would be very easy to confuse the two, but as
soon as a dissection is made, the double silk formation of the
former very quickly separates them.
The cocoons were found most abundantly on a cherry
tree some ten feet in height, the greater portion occurring in
masses on the trunk. A considerable number, however, were
to be found on the upper portions of the tree, where the crotches
of the twigs seemed to be a favorite spot for their construction.
Larve only were to be found, hence, it is safe to say the winter
is passed in this stage. The three specimens upon which I
was able to get exact records, changed from the larval to the
adult stage in 16 days after being transferred to a temperature
ML. tO. lo.
Since quantities of the overwintering eggs of the clover
mite, Bryobia pratensis Garman, were found upon the tree
which bore the Coniopterygid hibernacula, it is altogether
likely that this pest supplies one of the principal sources of
food for the insect. Quayle reports Conwentzia hageni Banks,
feeding abundantly upon the citrus red spider, Tetranychus
mytilaspidis Riley. Two visits were made last summer to
Cleveland with the hope of learning something definite regard-
ing the feeding habits of the larve; an additional visit was
ig
Hh
74 Annals Entomological Society of America _[Vol. VII,
made this winter in the hope of securing more of the hiber-
nating material, but at no time were any of the insects to be
found. The colony seemed to have disappeared entirely.
So much for general notes. The chief reason for directing
your attention to the subject at this time is to point out some
variations of wing venation which the writer observed, and in
ee
Fig. 1. Forewings of Conwentzia hageni Banks.
B. From Banks Proc. Ent. Soc. Wash., Vol. VIII; Q. From Quayle Univ.
Cal. Publications, Bul. 234.
1,7,4L,2R. One wing from each of four individuals in the author’s collection.
6L and 6R, 5L and 5R, 3L and 5R. Left and right wings of three individuals
in the author’s collection.
so doing he does not wish to be misunderstood to be attempting
to break down the validity of the specific determination,
but wishes merely to record the variations from the standpoint
of scientific interest. In the accompanying illustrations, B and
Su
Or
1914] Conwentzia Hagent.
Q are copied from Banks, Proc. Ent. Soc. Wash., Vol. VIII,
and Quayle, Univ. Cal. Publications, Bul. 234, respectively,
while the remainder are from retraced photo-micrographic
prints. Tllustrations 1, 7, 4L, and 2R, are all from different
insects. 6L and 6R; 5L and 5R; 3L and 3R, are three pairs
of wings. All of my specimens were bred from the same
lot of material and illustration No. 1 is made from a mounted
slide used by Banks in making the determination.
—— ———
B 6L
Q 6R
1 5L
“S SS
7 5R
2R
Fig. 2. Hindwings of Conwentzia hageni Banks.
The corresponding hindwings of the series of forewings shown in Fig. 1.
Banks refers to the variation in the placing of the cross-
vein* from Sc to R1 and the one from the radial sector to
M1,2. The illustrations of the fore wing by Banks and Quayle
do not differ materially, excepting that the cross vein between
* Whether this is a cross vein or SC2 is an unsolved point. It is possible that
an examination of pupal wing parts would decide the matter.
76 Annals Entomological Society of America [Vol. VII,
Cu2 and the first anal vein is not shown by Quayle. In all
of my specimens the cross vein from Se to R1 is either exceed-
ingly faint, or wanting entirely; sometimes being indicated
by a thickening of R1 at the point where it might be expected
to occur. Cross vein, from -Ril::to radial sector is usually
indicated by a sharply defined stub, extending about one-
seventh of the distance upward from the radial sector, and is
sometimes indicated above by a less sharply defined stub
extending downward. The position of the cross vein from
the radial sector to M1,2 is exceedingly variable, sometimes
joining above to R2,3,4,5, and sometimes to R4,5. In some
cases, such a difference occurs between the wings of the same
insect as in the case of 5L and 5R;3Land3R. Inthe majority
of cases the cross vein between Cu2 and the Ist A is repre-
sented by a short stub only, which arises from 1st A.
The variations in the venation of the hind wings are even
more striking than in the fore wings, especially regarding the
types illustrated by Banks and Quayle, when compared with
my photographs. Banks illustrates a cross vein between
R1 and the radial sector; Quayle shows a long oblique cross
vein between Sc and R1; while in all my specimens Sc bends
sharply downward towards the distal end and joins Rl. Some
wings have a short cross vein joining Sc and R1 near the point
where the former joins the latter, forming a very peculiar
little cell.
None of the longitudinal veins of the hind wing are shown
by Banks as joining the outer margin which agrees with Ender-
lein’s generic description of Conwentzia. Quayle shows radius
and its branch; media and cubitus joining the margin. In
my specimens Cu Rl, if not joining the margin, approaches
perilously near to it, while the radial sector unquestionably
joins the margin and cubitus joins in some instances.
An additional and final comparison may be drawn between
the comparative size of the fore and hind wings as represented
by the drawings of Banks and Quayle and my photographic
series. In the entire set of reproductions, the size ratio between
the fore and hind wings is truthfully maintained, hence, by
collating wings B, Q and 1, of the fore and hind series, it will
be seen that no very great difference obtains regarding the size
of the forewings, whereas the hind wing of Quayle’s specimen
is markedly larger than either of the others.
A COMPARISON OF NATURAL CONTROL OF TOXOPTERA
GRAMINUM IN SOUTH AFRICA AND THE
UNITED STATES.
By WiLt1aM Moore,
Asst. Prof. of Entomology, University of Minnesota.
During the past year, the author, while engaged as Lecturer
in Entomology at the Potchefstroon School of Agriculture,
spent considerable time in studying Toxoptera graminum in
South Africa and its control there by natural enemies. As the
results obtained have been somewhat different from those
obtained in the United States, it is thought that a comparison
of the conditions and the results would be of interest to Ento-
mologists.
Toxoptera graminum is found over certain large areas in
South Africa but attracts the greatest amount of attention in
the Orange Free State, Basutoland, and the western portion of
the Transvaal. It seems to have been present, at least in the
Free State, for many years, since the older farmers can remember
the pest as long as they have been farming in the eastern portion
of the Free State, known as the Conquered Territory. The
earliest definite tecord was 1896, which year is distinctly remem-
bered by a farmer who lost his entire wheat crop in that year.
In the higher portions of South Africa, i. e., an elevation of more
than 5000 feet, Toxoptera is either not present or is not present
in numbers sufficient to attract the attention of the farmer.
In the lower coastal regions, it also appears to be absent as, for
example, along the coastal region of Natal. It is probable that
the increased importance of Toxoptera during later years has
been due to the destruction of the locusts which formerly
swarmed over South Africa. There is little doubt but that the
locusts destroying the grain and incidentally destroying Toxop-
tera were responsible for the prevention of what might have
been a serious attack during that year. Since the locusts have
been destroyed, Toxoptera has had a better opportunity of
showing the injury of which it is capable.
In South Africa, there seem to be two forms of Toxoptera
under normal conditions, namely, the winged migratory
females and the apterous, viviparous females, the males and the
77
78 Annals Entomological Society of America - [Vol. VII,
oviparous females apparently being absent. As these forms
are too well-known to entomologists to require a description,
only a few notes concerning their migration and food plants
will be given.
Toxoptera, during the summer time, from about the middle
of November until March, lve upon various grasses and
volunteer grain plants. The most important grasses in this
connection ate Johnson’s Grass (Sorghum halepense), Goose
Grass (Eleusione indica), Sweet Grass (Panicum laevifolium),
Teff, Millet, Indian Corn, and Kaffir Corn (Sorghum Sp.).
The Blue Grass, upon which it is found so frequently in the
Free State, is not Andropogon hirtus, as reported by C. B. Van
der Merve and mentioned by F. M. Webster, but is the Sweet
Grass Panicum laevifolium, which is usually called Blue Grass
in the Free State and Sweet Grass in the Transvaal. There
are a number of grasses in South Africa known as Blue Grass,
without a distinctive common name. From March until June
or July, Toxoptera is found upon green forage crops such as
barley, rye, and oats. These give it an opportunity to exist
from the time that the grass becomes too old for it, or is killed
by the frost, until the main grain crop of the year is up, about
June or July. It is also very abundant during the winter time
upon Rescue Grass (Bromus willdenowii).
It exists on the winter grain until about September or
October when it changes to its summer host plants. The
severe attack usually occurs either in March on the green
forage crops or in July, August, or September, on the main
grain crops. The most critical time of the year for Toxoptera
in South Africa is in October or November, when it becomes
necessary to change from the grain field to its summer grasses.
The winters in South Africa are dry and, if rains do not occur
before October or November, the summer grasses do not come
up, while, on the other hand, the grain rapidly ripens, becoming
unfavorable for Toxoptera. At such a time, no doubt, large
numbers of. Toxoptera are lost in making this change. Some
of these are always saved by their ability to live upon the roots
and underground shoots of Johnson’s Grass where they are
attended by a common, grayish-brown ant, Plagiolepsis dusto-
diens. ‘This is interesting, inasmuch as H. Maxwell-Lefroy,
Government Entomologist of British India, reports Toxoptera
1914] Natural Control of Toxoptera. 79
graminum seeking shelter in the depths of the grass roots. They
were found in this situation in South Africa in many cases all
through the summer, probably thus obtaining some protection
from the hot rays of the sun. The forms on the roots were
quite white but when such were removed to wheat plants in
the Insectary, either they or their off-spring assumed their
natural color and it was found that the colonies on the roots
tended by the ants consisted of both Toxoptera graminum and
also Aphis maidis. These two species were also frequently
found associated upon the summer grasses, especially Panicum
laevifolium, Indian Corn, and Kaffir Corn.
As to the rate of reproduction of Toxoptera in South Africa,
it might be stated that it is about the same as it is in the United
States. In South Africa, migrations over large areas as re-
ported in the United States, are not so apt to occur. Similar
migrations on a smaller scale, however, do occur. Usually, it
is from the dryer portions to the wetter portions of the country,
inasmuch as Toxoptera will kill the grain sooner, under dry
conditions. There have been no reports of extensive migra-
tions in the Transvaal but they are known in the eastern
districts of the Orange Free State. Farmers about Ficksburg,
O. F. S., state that the swarms occur coming from the west late
in September or October. As the natives of Basutoland do
not raise grain for green forage crops, and as the grass is always
dead long before the winter grain comes up, most of the Toxop-
tera injuring the grain in Basutoland must come by migrations
from the Orange Free State. It is difficult to get any definite
data on this point from the natives but they seem to know that
something of the sort occurs.
In South Africa, two internal parasites have been found
which are capable of breeding in Toxoptera. The first (Aphid-
ius phorodontis (?) is commonly bred from Toxoptera in the
field. Another species is Diaeretus rape and has been bred in
the field from both the Cabbage Aphis (Aphis brassicze) and the
Green Peach Aphis (Myzus persice), both occurring upon cab-
bages but in the Insectary this species was also bred into
Toxoptera. It is doubtful, however, whether this species
would normally be found breeding in the field upon Toxoptera
unless the grain field was quite close to a cabbage patch. Aphid-
ius phorodontis has been bred from the Green Peach Aphis
80 Annals Entomological Society of America [Vol. VII,
(Myzus persicez), the Black Peach Aphis (Aphis persice-niger),
the Corn Leaf Aphis (Aphis maidis), Yellow Aphis on Milk-
weed (Aphis nerii), a reed Aphis (Hyalopterus arundinis) and
the Black Bean Aphis (Aphis rumicis).
Aphidius phorodontis, however, seems to be capable of
destroying just as many individuals of Toxoptera as is Aphid-
ius testaceipes. The average period of development from egg
to adult seems to be about ten to twenty days. The maximum
number developed from one female was 286 which is only 15
below the maximum obtained by Mr. Parks in the United
States Department of Agriculture. It is safe to assume that
in ‘‘stinging’’ so many that it must often occur that two or
more eggs are laid in one individual Toxoptera so that it is safe
to assume that Aphidius phorodontis lays 300 or 400 eggs.
There seem to be about 70 per cent. of the parasites females if
the mother Aphidius has been fertilized. It was also shown
that one male would fertilize more than one female but time did
not permit of finding how many females might be fertilized by
one male. If the female was not fertilized, she would lay eggs
and the proportion of the parthenogenetic off-spring were
about 70 per cent. males.
When an infested grain field is examined, even though
species of Aphidius are present, one never finds a large number
of parasitized forms of Toxoptera upon a weak plant such as is
shown to be the case in the United States. Probably one
wheat plant would, at the most, not have more than ten para-
sitized forms upon it. Besides this species of Aphidius, there
are three different species of ladybirds which play an important
part in controlling Toxoptera in South Africa. The first and
most important is the Black Spotted Ladybird (Adalia flavo-
maculata), the Red Spotted Ladybird (Chilomenes lunatus),
and the Black Ladybird (Exochomus nigromaculatus). These
ladybirds are a most important factor in controlling Toxoptera.
It is seldom that one finds a field badly infested in which lady-
birds are not present and rapidly destroying Toxoptera. The
life history of Adalia flavomaculata was worked out rather
completely.
Under favorable climatic conditions, the eggs hatched in from
five to seven days from the time they were laid. The larve
feed for a short time upon the eggshells but soon begin feeding
1914] | Natural Control of Toxoptera. 81
upon the Aphids. The larval stage lasts from about ten to
thirteen days during which time they eat, on an average, 320
Toxoptera per larva,—that being about 26 to 28 Toxoptera
per day. The pupal stage lasts six to 10 days and about 30
to 35 days elapses from the time that they emerge until they
lay eggs. During this time, they eat about 825 Toxoptera per
ladybird, making an average of about twenty-five per day to
each ladybird.
In this species, all the eggs seemed to be laid during one
period which lasted for about a week to ten days, during which
time they laid about 100 to 150 eggs. After having completed
the egg-laying, they live for some time before they die. The
Adalia seems to live about three to four months,—the males
dying first. In one experiment, the larvae were hatched from
eggs laid on the 10th of October, 1912, the last ladybird died
on the 24th of February, 1913,.and the average number of
Toxoptera destroyed was 2844 per ladybird. In this experi-
ment, the ladybirds were given as many Aphids as they could
eat. When the Aphids were scarce, the ladybirds would not
pass as rapidly through the different stages and the number of
eggs laid by the adults had a direct bearing upon the quantity
of food present. When the food supply runs short, the eggs
which have already been laid will be eaten while even the larva
will eat each other and the adults will eat the larve.
In the case of the Red Spotted Ladybird, we have a larger
species, being about 3-10 inches in length and nearly as wide,—
hemispherical in shape. The adult ladybird of this species
will lay from 150 to 250 eggs during her life. The egg-laying
period seems to be divided into several different stages. In
feeding experiments with the larve of this ladybird, it was
found that during the ten days of its larval existence, each
larva eats about 440 Aphids. Another interesting point is that
the normal life of the Red Spotted Ladybird is much longer
than that of the Black Spotted,—being about four to six months.
Both of these ladybirds were found to be parasitized by a
species of Dinocampus. The parasite laid its eggs in the larva
and in some cases, probably, in the pupa. The cocoon of the
parasite is formed underneath the adult ladybird and it seems
that the ladybird always reaches the adult stage before the
larva of the parasite emerges from the ladybird’s body to form
82 Annals Entomological Society of America [Vol. VII,
its cocoon. This species of Dinocampus seems to consist
entirely of females, as no males were found, even though
individuals were bred from females in the Insectary.
The third species of ladybird is much smaller than either of
the other two but has a life-cycle similar to the others. No
exact records were obtained as to the number of Aphids eaten
but it was found that the average life, from the egg until the
adult dies is about five to six months. One peculiarity noted
was that the eggs were not laid on end as is generally the case
with ladybirds, but rested on their sides on the leaf in little
heaps. This little ladybird is very severely parasitized by a
Chalcid belonging to the genus Homalotylus. The parasite
is usually bred from the larva collected in the field. In the
month of October, about 25% of the larvae seemed to be para-
sitized. From three to seven eggs are laid by the parasite in
one larva and the ladybird larva usually hangs itself up as
though about to pupate before dying. The larva of the para-
site pupates in the dead body of the ladybird larva.
This Chalcid will also breed in captivity in both the other
species of ladybird, but has not been bred from either of them
in the field.
The little Black Ladybird is most beneficial during the
spring and autumn when the cold affects the parasite more than
it does the ladybird. During the summertime the ladybird
becomes more or less rare, no doubt due to the parasite effect-
ively controlling it.
The ladybirds seem to be of greater value in controlling
Toxoptera in South Africa than in the case with the ladybirds
which feed upon Toxoptera in the United States. The fact
that Aphidius does not immediately destroy Toxoptera but
allows it to live for a few days, during which time, if the Toxop-
tera is an adult, it will produce young, detracts from the value
of Aphidius, when compared with the ladybirds. An example
will show better what is meant. Take a plant with 200 Aphids,
50 adults and 150 young, under control conditions and introduce
a female parasite. There is then a possible chance that all the
Aphids on this plant will have an egg laid in them but this will
not always happen, inasmuch as two or more eggs would be
laid in one Toxoptera while others would escape the parasite.
Granting, however, that all of them contained eggs and would
1914] Natural Control of Toxoptera. 8
OO
die in about seven days, it is found that the fifty adults would
have produced about 500 young during the first two or three
days after the egg of the parasite had been laid in their bodies.
These young would have a very small chance of being destroyed
by the parasite and would reach maturity after the death of
their mothers or about three or four days before the offspring of
the Aphidius which had “‘stung”’ the 200 Aphids, had emerged.
During these three or four days, they would be producing
young, so that when the Aphidius emerged, there would be
between 2000 and 2500 Toxoptera on the plant.
When the offspring of this female parasite emerged, however,
the chances are that all the Toxoptera would be destroyed.
From the above, it will be seen that starting with one Aphidius
and 200 Toxoptera on one plant, there would be at the end of
about fourteen days, 2000 or 2500 Toxoptera, while the plant
would not be entirely clean of Toxoptera until about twenty
days had elapsed. On the other hand, if an adult ladybird had
been introduced with the 200 Toxoptera, every individual
would have been cleaned away from the plant in ten days at the
most, while it probably would have been within five or six days.
When the ladybirds are present in the field with Aphidius,
another fact must be remembered, namely, that if Aphidius
has “stung’’ a number of Toxoptera and a ladybird later
ate these parasitized Toxoptera, the ladybird is also destroy-
ing the parasites as well as the Toxoptera. This is even
carried further by the ladybirds, inasmuch as they will eat the
parasitized forms even when the Toxoptera is dead, and the
parasite is in the pupal stage. It is no doubt due to this
reason that one does not find wheat plants covered with para-
sitized forms in the field in South Africa, as one does in the
United States. When a number of infested wheat plants were
enclosed with wire netting so that Aphidius could gain entrance,
but a larger insect, as a ladybird could not, it was soon found
that the wheat plants were crowded with parasitized forms
similar to those described by Webster as occurring in the
United States. There seems to be but little doubt. that in
South Africa, the ladybirds are of more value in combating
Toxoptera than Aphidius. This is of particular interest
inasmuch as an attempt is now being made to introduce A phid-
tus testaceipes into British East Africa where Toxoptera is
found near Njora.
84 Annals Entomological Society of America __[Vol. VII,
If Adalia flavomaculata and Chilomenes lunatus are found
there, it is very doubtful whether the value of Aphidius testa-
ceipes will be as great as it is in the United States.
The Little Black Ladybird, Exochomus nigro-fasciatus, is
reported from the Soudan and it, no doubt, occurs in British
East Africa. Several Aphid-eating Ladybirds have also been
reported from the Soudan as being particularly beneficial in
controlling an Aphid on Kaffir Corn.
Another point in favor of the ladybirds is that they will
breed and will control Toxoptera at a lower temperature than
Aphidius. All the stages are greatly retarded, however, by
cold and the adults do not seem to lay eggs. It seems from
experiments carried out that more Aphids are required per
ladybird in their lives when it is cold than is the case at a
warmer temperature. Larve of Adalia lived for about thirty-
five days at a mean daily temperature of from 45 to 55 degrees
and ate during that time 416 Aphids per larva, as compared
with 319 in the summer time in a period of thirteen days. At
a lower temperature, therefore, the larva eats only about half
the number of Aphids per day, but feeds for about 224 times as
many days.
Besides the ladybirds, one finds a Syrphid Fly (Xantho-
gramma scutellare) does considerable good in controlling Tox-
optera. The Syrphid also incidentally destroys Aphidius by
destroying Toxoptera which contain the eggs or larve of
Aphidius. In no case, however, were they found destroying
the parasitized forms of Toxoptera. A leaf which has been
cleaned by Syrphid Fly larva will be found to have a few
parasitized forms of Toxoptera remaining on it. This, how-
ever, would not be the case if the leaf had been cleaned by a
ladybird or a ladybird’s larva.
This Syrphid Fly is also retarded in its good work by a
parasite Bassus laetatorius. This parasite very effectively
controls the Syrphid Fly when the latter becomes very abund-
ant in the field. The most beneficial work of this Syrphid
was found to be in fields which were just becoming infested
with Toxoptera migrating from some other place. This
Syrphid seems to be the first to find the Aphids in their new
home and commence the work of destruction, but as soon’as the
1914] Natural Control of Toxoptera. 85
Syrphids become overly abundant in the field, the parasites
find them and they so reduce their number that they are of
very little value.
In conclusion, it might be stated that the cause of a bad
outbreak of Toxoptera in South Africa is due to the same
causes as a bad outbreak in the United States. If the early
winter months are abnormally cold, while the middle months
of the winter are warmer than the average, followed again by a
cold spring, there is a long period of from five to six months
during which time Toxoptera breeds more rapidly than the
ladybirds or Aphidius, the result being a bad outbreak of
Toxoptera.
Another factor which sometimes tends to cause bad out-
breaks is long periods of drought since, under such conditions,
plants are not able to withstand the number of Toxoptera
which normally they could carry without showing any ill effects,
thus dying, not entirely from injury by Toxoptera, but, with
drought as a secondary factor. If such land be irrigated, the
plants are enabled to survive the attack.
REPORT ON PARASITES.
Dr. L. O. Howarp.
The work on parasites and predatory enemies of the gipsy
moth and brown-tail moth has continued along the same
lines as during the previous year, except that no attempt
has been made to import additional parasites this season.
The material imported from Europe last year has been colonized
and an effort has been made to determine the extent to which
the species secured have established themselves in the field.
Owing to the fact that one of the imported egg-parasites
of the gipsy-moth, Anastatus bifasciatus, breeds very slowly,
extensive collections were made during last winter of parasitized
gipsy moth egg-clusters from colonies that were planted in
previous years. From this material it has been possible to
liberate 1,500,000 parasites of this species, and these have
been placed in 1,500 colonies in sections where the insect had
not become established. Eight hundred colonies were planted
in towns along the western border of infestation, and the
balance were liberated in a number of towns in the northern
part of Massachusetts. During November of this year col-
lections were made in New Hampshire in the colonies of Anas-
tatus that were planted a year ago, and examination showed
that these plantings were practically all successful although
the spread has been slow. From these collections about
100,000 parasitized eggs were secured and will be used for
colonization in New Hampshire next spring.
Investigations have shown that another egg-parasite of
the gipsy moth, namely Schedius kuvane has become perfectly
established in several colonies where it has previously been
planted. During the past year there has been a decided
increase in the abundance of this parasite, and in some cases
it has spread nearly a mile and a half from the limits of its
last year’s spread. The parasites attacking the caterpillars
of the gipsy moth have been found more abundantly than
during the previous year.
Compsilura concinnata, a species of Tachinid fly, was
very abundant during the summer of 1912, especially in the
territory which was longest infested by the gipsy moth, and
86
1914] Report on Parasites. a7
continued to spread during the past summer. It has not
been so abundant in the oldest infested territories as in some
of the outlying colonies. Collections of more than eleven
hundred gipsy moth caterpillars made in four towns in central
Massachusetts show a parasitism by this species of over 40
per cent, while similar collections in the central infested area
have indicated an average parasitism of about 5 per cent. It
is probable that the decrease in parasitism 1n the old infested area,
as far as this species is concerned, is due to the fact that gipsy
moth caterpillars are not nearly as abundant as they were
during the previous year, and also because of the enormous
numbers of the American tent and forest caterpillars which
were present in this region and which are also attacked by
this parasite. :
Limnerium disparidis and Apanteles species were received
from Europe for the first time in 1911 and were planted in
several badly infested gipsy moth colonies. Both species
were recovered during the summer of 1912, which indicated
that it is possible for the insects to withstand our cold winters.
In the case of the latter species, as high as 7 per cent of para-
sitism of gipsy moth larve was found. The present summer
the Limnerium was recovered from a single locality where
the species was liberated in 1911. Although it has evidently
become established, it has not thus far shown marked ability
to increase in the gipsy moth infested area in New England.
Another species of Apanteles, namely A. Jacteicolor, an
important parasite of the brown-tail moth caterpillars, has
been recovered in large numbers and has been found to attack
gipsy moth caterpillars in widely separated regions. This
species seems to be multiplying more rapidly than any of the
other Hymenopterous parasites of the gipsy moth. In order
to colonize this species over as wide an area as possible, an
arrangement was made with the State Entomologist in New
Hampshire and the Superintendent of Moth Work in Maine
to liberate as many colonies as possible along the outskirts
of the area infested by the brown-tail moth in those states.
Small collections of gipsy moth larve were made at Melrose,
and in some cases ten per cent of the larve were killed by this
species. In several localities in New Hampshire the past
summer cocoons of this parasite were very abundant, and
88 Annals Entomological Society of America [Vol. VII,
several hundred were easily collected for experimental work.
They were taken for the most part on the foliage of trees and
attached to dead caterpillars.
The Calosoma beetle (Calosoma sycophanta) has coon
observed in large numbers in towns where bad colonies of the
gipsy moth were present. It has not been possible to obtain
definite records of the amount of benefit derived from this
species or of its abundance, except in cases where trees were
burlapped, as these bands furnish favorable hiding places for
the caterpillars and are favorite locations for the beetles and
larve to obtain food. In such cases, where caterpillars were
abundant, twenty or more of the Calosoma larve have fre-
quently been found under a single burlap band on an average
sized tree. As they feed upon the pupz as well as upon the
caterpillars, the amount of benefit derived is very great,
although it is difficult to figure the percentage of larve killed.
From collections made during the winter of 1912-13 it
was determined that Monodontomerus aereus has spread over
practically the entire territory now known to be infested by
the brown-tail moth. It was not found in as large numbers as
during the previous year. Pteromalus egregius has been found
widely scattered over the area infested by the brown-tail
moth, and its numbers are slowly increasing, judging from the
records that have been secured from sample collections.
There is thus no doubt that a number of the imported
species are thoroughly established and that they are increasing
each year, and further that many hundreds of thousands of
caterpillars were killed by them during the past summer.
NOTES ON SOME OLD EUROPEAN COLLECTIONS.
By H. T. FERNALD, Amherst, Mass.
There seems to be little on record in this country about
the older European collections of insects. Possibly the facts
are more or less common knowledge, but if so, a rather careful
search has failed to produce much of value. Yet in these
days when types are coming into such prominence as the
“court of last resort’’in our .attempts to finally establish
specific identities, the location of these collections, their state
of preservation and any facts which may enable workers to
find the specimens they desire to examine, should be on record.
The following notes are therefore offered in the hope that
they may be of some use at least, to those who expect to study
abroad.
The collection of Linné as including the first insects to
which the binary nomenclature was applied, is of much interest.
This collection appears to be now in part at Upsala and
in part at London. The material at Upsala was for many
years at the royal castle Drottningholm, but in 1803 what
remained was sent to the Academy of Science at Upsala by
Gustav Adolph IV, where it was arranged and labelled by Lin-
naeus’ student, Thunberg. This was probably the portion
which constituted the collection belonging to the Queen of
Sweden, and of which Clerck illustrated the Lepidoptera in
his Icones Insectorum, and to judge from this, and such state-
ments as are available, consisted only of Lepidoptera.
In the zoological museum of the University of Upsala
are two wooden cases of Linnaean Hymenoptera which have
recently been examined by Schulz (Berl. Ent. Zeits., LVII,
55, 1912), and reported upon.
Linné’s private collection was sold by his wife after the
death of her son in 1783, to Dr. James E. Smith, of England,
for one thousand guineas. It consisted of his books, cor-
respondence, insects and plants, and reached London in 1784.
In 1788, due largely to the influence of Smith, the Linnaean
Society was established, and this material is now in charge
of this society which is located at Burlington House, Piccadilly,
89
90 Annals Entomological Society of America [Vol. VII,
London. It is kept in the original case, in which are drawers
containing the insects, while others contain the plants and
books. Smith long retained the collection and being a col-
lector, received many specimens from friends. These it would
seem, he placed in the drawers with the Linnaean specimens,
so that as it now stands, the collection contains many speci-
mens not properly belonging with the Linnaean material.
This unfortunate condition is liable to cause confusion in any
examination, as the Linnaean specimens can only be dis-
tinguished by the handwriting on the labels.
The work of Fabricius can be found in a number of col-
lections in Europe. His personal collection is in the Zoological
Museum of the Univeristy of Kiel, but he made numerous
visits to different museums, naming the insects where he
went, and many. of these are still in existence.
At Kiel his collection is in glass-topped trays, the tops —
being loose in many cases. The trays are in two large cases,
one occupied almost entirely by beetles, while the Hymenoptera
and some other orders fill the other.
The two groups named, at least, are in fair condition, and
the Hymenoptera are arranged to correspond with the Systema
Piezatorum. Each genus has a special label, bearing besides
the name, the number of the genus, as given in the Systema,
and each species is preceded by a similar label giving the name
and number of the species. In some cases the insect itself
is missing though the label is present, and some numbers are also
absent, probably because the book enumerated species he had
seen and described, but which he himself did not have.
Probably the next largest collection of Fabrician material
is now in the British Museum of Natural History in London,
and is known as the Banksian Collection. This material
was obtained by Mr. (afterwards Sir Joseph) Banks and Dr.
Solander, who accompanied the celebrated Captain Cook in
his voyage to the South Sea Islands and around the world,
from 1768 to 1771. It was studied by Fabricius between 1772
and 1775 and the new species were published by him in 1775
in his Systema Entomologiz and are indicated by the words
‘“Mus. Bankianum,’’ or in his later writings by the words
‘“Mus. Dom. Banks.’’ This collection was for years in the
hands of the Linnaean Society, but is now in the British Museum
1914] Notes on Some Old European Collections. 91
where most fortunately it is preserved as a separate collection
bearing the original Fabrician labels, thus giving us Fabricius’
ideas of the old species even where they are not his types.
They have been worked over somewhat since, not always
fortunately, and some care must be exercised to be sure of all
the labels. Thus Frederick Smith in his Catalogue of the
Hymenopterous Insects in the British Museum records Sphex
pensylvanicus L. as being in this collection. An examination
of the specimen shows the unquestionable Fabrician label
‘“‘pennsylvanica’’ and above it a much more recent label,
‘Pennsylvania’ written by Smith, to judge from the hand
writing. So close to the underside of the body as to be easily
overlooked, is a third and very old label reading ‘“‘ Nov. Holl.”’
Now as Captain Cook on the voyage during which these insects
were collected, did not touch at any point in North America,
it is evident that Fabricius named the insect entirely from
its appearance, without reference to the locality where it
was captured, and Smith, probably failing to notice the upper
label, added a locality label of his own preparation to agree
with the name. An interesting feature of the case is that
Fabricius wrongly identified the specimen, which is not pensyl-
vanica at all, but fumipennis of Smith, described by the latter
from other material, in the same book in which he records this
insect in the Banksian collection, evidently failing to recognize
the identity of the two. An occasional case like this indicates
that care in the study of this collection is indispensable.
The National Museum at Paris also contains some speci-
mens named by Fabricius. What remains of the Bosc collection
is there, and Fabricius frequently records species from the
““Mus. Dom. Bosc.”’ which apparently came into the hands
of the National Museum at the death of Bosc in 1828. This
material is much scattered, however, and can only be recog-
nized by labels in the hand of Fabricius and the others which
read ‘‘Museum Paris Coll Bosc 1828”’ printed on green paper
and which were undoubtedly added when the collection became
the property of the Museum.
The collection of Klug is in the Zoological Institute of the
University of Berlin. It is not kept separate but has been
worked into the general collections and can now be recognized
only by the labels. These are on green paper and bear the
92 Annals Entomological Society of America [Vol. VII,
specific name, beneath which is a capital N and at the bottom
the locality, all enclosed by a black line. Frequently the middle
of the left side of the label has a slit cut in, so that it might
slip around the pin instead of being penetrated by it. In
some cases the label has been closely trimmed, but comparison
of the writing with that on the entire ones, is sufficient to estab-
lish its identity.
Klug seems to have been more liberal with names on museum
specimens than with published descriptions of these species,
at least in the Hymenoptera. Apparently later workers in
some cases found the names Klug had given to the specimens
and published them, adopting names which would otherwise
have no standing. Erichson Seems to have done this in one
case at least, and Dahlbom in several.
Though the collection of Dahlbom is at the University of
Lund, his work is to some degree in evidence at Berlin where
he studied for a time with Klug, and specimens named by him
are frequently met with there. His own collection at Lund
is kept by itself in the condition in which he left it and for
the most part in a good state of preservation.
In the Paris Museum a few specimens which seem to have
been labeled by Latreille are still preserved. These labels
have double red line borders and the names which are hand-
printed are, first the French name, and beneath this the Latin
one, the two being bracketed together on the right, beyond
which is the abbreviation ‘‘Latr.’’ In some cases at least,
these names do not appear to have been published and -there-
fore have no standing.
A few boxes of Hymenoptera in this collection are marked
‘“Brullé Collection’? on the outside, so that some of Brullé’s
species at least, are still in existence.
The Lepelletier collection is in much the same condition
and some boxes bear the label ‘‘Lepelletier Collection.’’ In
these the material named by him is probably indicated by
names written in red ink between parallel red lines on the
labels. Unfortunately many of his species are missing, and
in the case of the Sphecide none of his American species can
be found. Whether they have been accidentally destroyed,
or, most of them being from Serville, were returned and later
were lost, cannot now be determined.
1914] Notes on Some Old European Collections. 93
Though not one of the old collections, that of Achille
Costa may be mentioned. This is now at the University of
Naples, where it is kept in a room by itself. There is no
entomologist at the University and the collection is in charge
of the Professor of Parasitology, but it is in excellent con-
dition and apparently well cared for.
Many other old collections may be found in different
parts of Europe, but not having paid particular attention to
them they are not touched upon here. It is noticeable nearly
everywhere that these collections are for the most part kept
in trays so open that in this country a single year would probably
see their complete destruction by museum pests. These
nuisances do not appear to be very important abroad or the
priceless collections of Linne, Fabricius and others would long
ago have become mere heaps of dust at the bases of the pins.
DISCUSSION.
Dr. Howard said in discussion that there is so much of
value in information of the character of that contained in
Dr. Fernald’s paper, that he was emboldened to add two state-
ments: First, that the bulk of the collection of A. H. Haliday,
the brilliant Irish entomologist, is now carefully preserved by
Prof. G. H. Carpenter in the College of Agriculture, in Dublin;
Second, that the Ratzeburg types are preserved like religious
relics by Dr. Eckstein at the forest school at Ebersiwalde bei
Berlin.
THE ENTOMOLOGICAL SOCIETY OF AMERICA.
Organized 1906.
OFFICERS FOR 1914.
President.
Pei! CAWVERT. 6.2.0). University of Pennsylvania, Philadelphia, Pa.
First Vice-President.
ames (Gay NEEDHAM so. 505 x... Cornell University, Ithaca, N. Y.
Second Vice-President.
@; GORDON Hrwitt... 2). 2%. Dominion Entomologist, Ottawa, Canada.
Secretary-Treasurer.
Rana Os NUACGELIVRAY Sei. | University .of Illinois, Urbana, Ill.
ADDITIONAL MEMBERS OF EXECUTIVE COMMITTEE.
HERBERT OsporNn, Ohio State University, Columbus, O., ex officio.
Wittaim M. WHEELER, Harvard University, Cambridge, Mass.
VERNON L. KELLocG, Leland Stanford Jr. Univ., Stanford Univ., Calif.
NaTHAN Banks, United States National Museum, Washington, D. C.
E. P. Fett, State Entomologist, Albany, N. Y.
J. M. Atpricu, U.S. Bureau of Entomology, Lafayette, Ind.
COMMITTEE ON NOMENCLATURE.
H. T. Fernatp, Massachusetts Agric. College, Amherst, Mass., 1914.
E. P. Fett, State Entomologist, Albany, N. Y., 1915.
T. D. A. CocKERELL, University of Colorado, Boulder, Col., 1916.
COUNCILORS FOR THE AMERICAN ASSOCIATION FOR THE
ADVANCEMENT OF SCIENCE.
Puitire P. CALVERT, University of Pennsylvania, Philadelphia, Pa.
CHARLES J. S. BETHUNE, Ontario Agric. College, Guelph, Ontario.
EDITORIAL BOARD OF ANNALS.
HERBERT Ossporn, Managing Editor, Ohio State Univ., Columbus, O.
J. H. Comstock, Cornell University, Ithaca, N. Y., 1914.
CHARLES J. S. BETHUNE, Ontario Agric., College, Guelph, Ontario, 1914.
C. W. Jounson, Boston Society of Natural History, Boston, Mass.,1914.
VERNON L. KeEttoce, Leland Stanford, Jr., Univ., Stanford Univ., Cal.,
1914.
L. O. Howarp, Chief, Bureau of Entomology, Washington, D. C., 1914.
W. M. WHEELER, Harvard University, Cambridge, Mass., 1914.
Puitip P. CALvertT, University of Penn., Philadelphia, Penn., 1914.
J. W. Fotsom, University of Illinois, Urbana, Ill., 1914.
95
RESOLUTIONS.
ON THE DEATH OF PHILIP REESE UHLER.
Dr. Philip Reese Uhler, L. L. D., Provost of the Peabody
Institute, Baltimore, Maryland, died October, 21, 1913. ©
Dr. Uhler was an honorary fellow of the Entomological
Society of America, one of the first to be distinguished by
such election and was recognized as one of the most eminent
entomologists of the country. His contributions, especially
to our knowledge of the Hemiptera, were of great extent and
form the basis for the study of this group in America and have
had no small influence upon the development of the subject
in the world at large. They will stand as an enduring monu-
ment to his industry, skill and keen insight as a systematist.
Along with the high qualities of his scientific work he
had a most engaging personality and his memory will be
cherished by all who had the good fortune to secure his friend-
ship.
He was a man of true scientific spirit and enthusiasm,
a lover of the fields and woods, an expert collector, and most
generous in his aid to his fellow workers.
Recognizing the permanent value of his contributions to
entomology and also the high character and worth of the
individual, the members of the Entomological Society of
America desire to place on record in an enduring manner its
sense of appreciation for the man and his work and it is hereby
resolved that this testimonial be spread upon the minutes
of the Society and printed in the ANNALS.
Committee HERBERT OSBORN
PHILIP P. CALVERT,
J. H. COMSTOCK.
96
Annals E, S, A., Plate XVI.
PHIEIP” REESE: UHLER,. LL. D.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY
OF AMERICA.
Atlanta Meeting.
The Eighth Annual Meeting of the Entomological Society
of America was called to order by Dr. Philip P. Calvert in
the absence of the President, Rev. Charles J. S. Bethune, at
10:00 A. M., Tuesday, December 30th, in the rooms of the
Physiological Department of the Atlanta Medical College.
In calling the meeting to order Dr. Calvert conveyed to the
Society Dr. Bethune’s deep regret at being unable to be present .
and his hearty good wishes for its success. The meetings
were all well attended, the number was surprising considering
Atlanta’s isolation from great educational centers. The fol-
lowing committee, appointed by President Bethune, was
named:
Committee to draft resolutions on the death of Dr. Philip
Reese Uhler.——Herbert Osborn, J. H. Comstock, and Philip
P. Calvert.
The chair was directed by motion to appoint the following
committees: Committee on Resolutions; Committee on Nom-
inations; Auditing Committee.
The following papers were then read, but through the action
of the Executive Committee that all abstracts should be omitted
from the proceedings, only titles are given:—
J. T. Lloyd, Cornell University—The structure of the hind
intestine of Corydalis.
Paul S. Welch, Kansas Agricultural College—Observations
on the habits and life-history of Hydomyza confluens Loew.
Read by title.
Stanley B. Fracker, University of Illinois—New char-
acters in the classification of microlepidopterous larve.
Cornelia F. Kephart, Cornell University—The poison glands
of Euproctis chrysorrhoea Linn. Presented by W. A. Riley.
N. L. Partridge, University of Illinois—The tracheation of
the anal area of the wings of the Lepidoptera and the homology
of the veins. Read by title.
Herbert Osborn, Ohio State University—The box-elder
bug in Ohio.
|
98 Annals Entomological Society of America [Vol. VII,
V. E. Shelford, University of Chicago—The elytral trachea-
tion of the subfamilies and genera of Cicindelide.
Edna Mosher, University of Illinois—Some interesting
structures in the pupze of Lepidoptera.
W. A. Riley, Cornell University—Some sources of error in
the interpretation of insect tissue.
J. S. Houser, Ohio Agricultural Experiment Station—
Conwentzia hageni Banks, life-history notes and variations in
wing venation.
Alvah Peterson, University of Illinois—Notes on the head-
structures of Thysanoptera. Read by title.
Philip P. Calvert, University of Pennsylvania—The desira-
bility of a biographical dictionary of entomologists.
The President announced the following Committees :—
Committee on Resolutions—F. L. Washburn, Henry Skinner,
and J. G. Sanders.
Committee on Nominations—W. A. Riley, P. J. Parrott, and
G. M. Bentley.
Auditing Committee—T. J. Headlee, H. T. Fernald, and
Ris Hy Pettit:
The Society then adjourned to meet at 2:00 P. M. The
afternoon was devoted to a joint meeting of Section F of the
American Association for the Advancement of Science and
of the Entomological Society of America at which the fol-
lowing papers were presented :—
L. O. Howard, United States Entomologist—Note on the
present status of the Gipsy Moth parasites in New England.
E. L. Worsham, State Entomologist of Georgia—Some notes
regarding the natural history of the mole cricket.
H. T. Fernald, Massachusetts Agricultural College—Notes
on some old European collections.
P. J. Parrott, W. O. Gloyer and B. B. Fulton, New York
’ Agricultural Experiment Station—Studies on the Snowy Tree-
cricket, Oecanthus niveus, with reference to apple bark diseases.
Presented by P. J. Parrott.
J. Chester Bradley, Cornell University—Collecting insects
in the Okefenoke swamp. Presented by J. G. Needham.
Herbert Osborn, Ohio State University—Studies on the
geographical distribution of leaf-hoppers, especially of Maine.
Philip P. Calvert, University of Pennsylvania—The fauna
of the epiphytic bromeliads in Costa Rica.
1914] Proceedings of the Atlanta Meeting. 99
The Society adjourned at 4:45 P. M., to meet Wednesday,
December 31st, at 9:30 A. M.
The annual business meeting of the Society was held upon
reconvening and the following reports were presented :—
The committee appointed to draft resolutions on the death
of Dr. Philip Reese Uhler presented its report. It was ordered
accepted and printed.
The Secretary presented the following report for the Execu-
tive Committee, which met at the Hotel Ansley, Tuesday
evening. Dr. W. M. Wheeler and Dr. Henry Skinner were
named as additional members of the committee. Dr. Skinner
sat with the committee. '
REPORT OF THE EXECUTIVE COMMITTEE.
The following matters were considered in the interim since the last
Annual Meeting :—
The appointment of Dr. E. P. Felt to give the Annual Public
Address at the Atlanta meeting.
A request from the Secretary that the President be given permission
to call for the election of a sufficient number of persons to make a quorum
of the Executive Committee at the Atlanta meeting in case there were
not four members in attendance.
Mr. Nathan Banks presented a request from the International
Committee on Nomenclature of the Second International Congress of
Entomology, that this society should name two members to serve on
the American National Committee. President Bethune named as
representatives, Dr. E. P. Felt and Dr. H. T. Fernald.
Mr. A. W. Baker, Secretary of the Entomological Society of
Ontario, presented a request that the Entomological Society of America
should send a delegate to its Jubilee meeting, the fiftieth annual meeting,
to be held at the Ontario Agricultural College, Guelph, Wednesday,
Thursday, and Friday, August 27 to 29. The President named the
Secretary as the representative of the Society. Your Secretary had
the pleasure of attending these meetings. He had the most delightful
time, intellectually and socially, of any scientific meeting that he has
attended. There were delegates present from England, Scotland,
United States, and Canada.
The following new members were elected June 1, 1913:
W. C. Allee. David Gunn.
Frank M]. Gibson. H. A. Horton.
The following members have died during the year:
D. F. Berrenger. .» Charles W. Hooker.
A. G. Hammar. J. M. Shaffer.
PR: Uber.
100 Annals Entomological Society of America [Vol. VII,
The following resignations were presented and accepted:
Otto Bucholz. P. E. Smith.
Fred Johnson. R. I. Smith:
C. A. Shull. Anna C. Stryke.
A. G. Vestal.
The names of thirteen members were dropped from the rolls for
non-payment of dues.
The following new members were elected by the Executive Com-
mittee at its meeting last evening:—
L2@. Barber. W. M. Mann.
G. T. Bethune-Baker. E. A. McGregor.
S. W. Bilsing. Ximena McGlashan.
M. W. Blackman. J. D. Mitchell.
Josef Bruner. L. J. Nickels.
R. E. Campbell. F. B. Paddock.
Leroy Child. Phil. Rau.
E. S. Cogan. E. A. Richmond.
W. M. Davidson. L. P. Rockwood.
G. A. Dean. James Sinclair.
H. F. Dietz. M. P. Somes.
W. D. Edmonston. Dayton Stoner.
W. O. Ellis. D. T. Stevens.
}.-BY Gall, Pts Valbert:
J: E: Graf J. D. Tothill
T. E. Holloway. C. T. Vorhies.
J. R. Horton. Otis Wade.
H. L. Johnson. J. R. Watron.
Cornelia F. Kephart. H. B. Weiss.
R. J. Kewley. G. M. Wendelken.
F. H. Lathrop. F. X. Williams.
Philip Luginbill. T. S. Wilson.
F. L. McDonough. H. P. Wood
R. S. McDougall. W. C. Woods
J. R. Malloch. M. A. Yothers.
The total membership of the Society as reported at the Seventh
Annual meeting was 410, deducting the names of the persons who have
died during the year, resigned, or dropped by the Secretary and adding
the names of the new members elected in June and at this meeting,
the membership is now 439.
The Secretary was instructed to prepare a small booklet describing
the origin of the Society and its aims, as an aid in further extending
the membership.
The first suggestion of a national entomological society was
presented at the last Philadelphia meeting in 1904, of the American
Association for the Advancement of Science. Dr. Henry Skinner was
asked by the Executive Committee to prepare a history of the Ento-
mological Society of America to be read at the Eighth Annual Meeting.
1914] Proceedings of the Atlanta Meeting. 101
TREASURER’S REPORT.
RECEIPTS.
Cash on deposit in the First National Bank of Champaign, Illinois,
Weceiise rs LON mnt thee Metin Been areec Races wakes otis miele a Meany, ale 0 3% $ 40.87
Life Membership Fees deposited with the Cleveland Trust Company,
Cleveland wOhitos September 18; 1912 ss ne os eet. hoses oS avs ces 100.00
Life Membership Fee of W. T. M. Forbes deposited with the Cleveland
Trust Company, of Cleveland, Ohio, September 18, 1913.......... 50.00
Interest on Fees of Life Members, July 1, 1918........................ 4.37
Cash received from Herbert Osborn, Managing Editor of the Annals... 252.86
Cashmecollectedmaseduesea tse tian crea a eisine Gohnes Wie save ata ed tea Slaton os ¢ 726.09
$1,144.19
BALANCE.
Bills paid:
Annals and separates for December, 1912.............. $377 .47
Annals and separates for March, 1918................%.. 332.25
Bieravyine bills paid) by the Mreasurer..o0 i... own. 22. ss 66.09
Stamps and Stamped envelopes...............02.0000eee 27 .62
Printing for the Secretary-Treasurer’s office............. 28.03
SLEMOOAD Gra wee ae poe ree et eee ete hee sins oe static Mace 20.25
One-half guarantee on Cleveland smoker................. 3.67
Gheclesreitised? bymbanks ante tet ee esi ieee sees ete bee 2.00
he as
Life Membership Fees deposited with the Cleveland Trust Company,
jae Cleveland: Ohio's Decemibers8; = 191322 isn 5 sian ho veo ik eg als hone 150.00
Interest on Fees of Life Members deposited with Cleveland Trust
} Company, Cleveland; Obio,, December'$, 1913 oo. wine cncted eins ess 4.37
Cash on deposit in the First National Bank of Champaign, Illinois,
IDSCEMIO CIES mil Date one weriehirs CsA rset sie a dacsleie a Reloaeaten wareie ad as 132.44
$1,144.19
Dr. C. Gordon Hewitt, Provincial Entomologist of Canada and
Dr. William Barnes, Lepidopterist of Decatur, Illinois, were elected
Fellows of the Society.
It was voted, that in the future no abstracts of the papers presented
at the Annual Meeting should be included in the printed proceedings
of the meeting.
The following amendment to Article IV, Section 2 of the Consti-
tution was recommended, which reads:
Article IV, Section 2. The business of the Society not otherwise
provided for shall be in the hands of an Executive Committee, consist-
ing of the officers named in Section 1, and of six additional members,
five of whom shall be elected from the Fellows by the Society, and the
sixth shall be ex officio the Managing Editor. Four members of the
Committee shall constitute a quorum.
To be amended to read:
Article IV, Section 2. Executive Committee——The business of the
Society not otherwise provided for shall be in the hands of an Execu-
tive Committee, consisting of the officers named in Section 1, and of
six additional members, five of whom shall be elected from the Fellows
by the Society, and the sixth shall be ex officio the Managing Editor.
102 Annals Entomological Society of America [Vol. VII,
There shall be a meeting of the Executive Committee at each Annual
Meeting. Four members shall constitute a quorum and in the case of
the non-attendance of this number at any Annual Meeting, the Society
shall elect a sufficient number from among the Fellows in attendance
to complete the quorum.
On motion the report of the Executive Committee was
adopted.
REPORT OF THE MANAGING EDITOR OF THE ANNALS.
The Editor begs leave to report for the past vear the following items:
RECEIPTS.
Suibserip biomes shes Pe cease ee RE Oe ee $170.60
Back Numbersiney ©. See atee ea on eee 8A ee a RST A Re Re 81.91
Authorseprimakse. 2 10s se i. Sra ce. Beate Re SS ee ged tee ne 70.00
$323.11
EXPENDITURES.
ETeRa van ase Chey Sk. mee cee =o Rae el te eee ee an a ee ee ed hse ote ee $52.80
I BXofs| ey kone Meeaann ORE Ree et RO eee LET Oe att nian, od ears Nhs = ae guy ee Brews 9.65
stenosraphic Helos cos. {2 ac oe Seine aed Serge ee eae ee eee 7.80
Paid “Preasuren. ess ok oles c helene ate Seen ee es eee 252.86
$323.11
The volume for the year, which includes about five hundred pages
and fifty-nine plates, will I believe maintain the quality of preceding
volumes and it has been necessary to postpone or refuse other papers
of excellent quality because of the lack of funds for further publication.
It is hoped that the income for the coming year will permit the handling
of a number of these papers but the Editor believes that the amount
printed should be kept within the margin of receipts and if possible no
deficit be created. The sale of a few sets of back volumes would assist
much in the publication of additional matter and any assistance in
placing such sets will be very much appreciated.
The Editor desires to express his appreciation of the aid rendered
by the members of the Editorial Board, and to the Secretary for his
untiring efforts in attending to the details of the business falling to
his office. Respectfully submitted,
HERBERT OSBORN,”
Managing Editor.
REPORT OF THE AUDITING COMMITTEE.
Your Auditing Committee presents herewith its report. We have
examined the books of the Treasurer for the year ending December 8,
1913, and find them to be correct. We have also examined the accounts
of the Managing Editor of the Annals and find them to be correct.
Signed, THomAS J. HEADLEE,
H. T. FERNALD,
Rh. Peri.
ah i
ip
1914]- Proceedings of the Atlanta Meeting. 103
REPORT OF THE COMMITTEE ON NOMINATIONS.
Your committee begs leave to report the following names as nom-
inees for the respective offices for 1914:—
OFFICERS.
President: Philip P. Calvert, University of Pennsylvania, Phil-
adelphia, Pa.
First Vice-President: James G. Needham, Cornell University,
Tthaca..Ne Y.
Second Vice-President: C. Gordon Hewitt, Provincial Entomolo-
gist, Ottawa, Canada.
Secretary-Treasurer: Alex. D. MacGillivray, University of Illin-
ois, Urbana, II.
ADDITIONAL MEMBERS OF EXECUTIVE COMMITTEE.
Herbert Osborn, Ohio State University, Columbus, Ohio, ex officio.
William M. Wheeler, Harvard University, Cambridge, Mass.
Vernon L. Kellogg, Leland Stanford Jr., University, Stanford Uni-
versity, Cal.
Nathan Banks, United States National Museum, Washington, D.C.
E. P. Felt, State Entomologist, Albany, N. Y.
J. M. Aldrich, United States Bureau of Entomology, Lafayette, Ind.
MEMBER OF COMMITTEE ON NOMENCLATURE.
T. D. A. Cockerell, University of Colorado, Boulder, Colorado.
Signed, W. A. RILEY,
Pes PARROTT.
G. M. BENTLEY.
On motion, the Secretary was instructed to cast a ballot
for the officers named and they were declared elected.
REPORT OF THE COMMITTEE ON RESOLUTIONS.
The Committee on Resolutions beg Jeave to submit the following :—
Resolved, That the Entomological Society of America hereby
express their appreciation of the various courtesies extended to them at
this meeting by the city of Atlanta, by Governor and Mrs. Slaton, by
the University and Capital City clubs, by the Atlanta Medical College,
and by the local Press.
Resolved, That its thanks are also due the Atlanta Chamber of
Commerce for its hearty co-operation with the University Club and
the Atlanta Convention Bureau through whose efforts the meeting at
Atlanta was made possible; and further
Resolved, That our thanks are especially due and are hereby ex-
tended to the local Executive Committee and Mr. E. L. Worsham for
their painstaking and effective efforts in behalf of the convention.
Signed, F. L. WASHBURN,
HENRY SKINNER,
J. G. SANDERS.
104 Annals Entomological Society of America [Vol. VII,
The Committee on Nomenclature presented an informal
report which was ordered accepted.
REPORT OF THE COMMITTEE ON ENTOMOLOGICAL TYPES.
One of us (Cockerell) examined the collections of the U. S. National
Museum and the Carnegie Museum during the year. The types, so
far as seen, were found in excellent condition, but not all in systematic
order or available for study and comparison without some searching.
We know of no museum in which the entomological staff is really
adequate. At the Carnegie Museum one is amazed at the richness and
value of the collections, including materials which have been described
in many important memoirs, and enormous numbers of specimens not
yet worked over, but evidently including much of interest. The
entomological curator has succeeded in keeping everything in good
condition, but it would take a considerable staff of workers to put the
collections all in order and keep the accessions worked up. At the
U. S. National Museum one finds a large staff of well-known entomolo-
gists, many of whom work over time and on holidays in the effort to
keep the collections in order and work up the accessions. However,
the appearance of an adequate entomological staff is illusory, since
nearly all of the men belong to the Bureau of Entomology of the Depart-
ment of. Agriculture, and have to give their attention to economic
problems and routine work of various kinds. Judging by the large
amount of published work issued from the Museum, one might suppose
the number of workers to be sufficient, but this idea is soon dispelled
on examining the very large and important collections remaining
unstudied and noting the continued stream of accessions. The scien-
tific staff of the National Museum is inadequate in almost all depart-
ments, but especially in Entomology, a subject which covers a much
greater and more important field than the public imagines. The
type problem becomes part of the general problem of securing adequate
and competent curatorial assistance; not only for the proper care and
availability of the types already owned by the Museum, but also and
especially in regard to obtaining other types. The study and descrip-
tion of the new species now in the Museum would add thousands of
types to the collection; while many private workers would give or leave
their types to the institution, were entomology treated by Congress and
the authorities as it deserves.
We think, therefore, that all entomologists should make a point of
urging, whenever possible, the claims of their science to a larger share
of support in important Museums. In doing this, they may properly
point out the astonishing revelations of recent years in regard to the
importance of various insects to man, showing that a knowledge of
entomology is of prime importance for the progress of civilization.
They may also point out that in the case of large public museums, all
the major expenses have been met, and it is only necessary to add a
comparatively small amount to greatly increase the scientific output.
fteaennis
1914] Proceedings of the Atlanta Meeting. 105
Several workers in the National Museum recently agreed with the
suggestion that 5% of the total expenditure on the Museum, added to
the cost of maintaining the scientific departments, would double the
scientific out put.
Signed, T. D. A. CocKERELL,
L. O. Howarp,
HENRY SKINNER.
On motion, the report was ordered accepted and printed
and the Committee continued for another year.
REPORT OF THE SPECIAL COMMITTEE ON HOLDING A SUMMER MEETING ON
THE PACIFIC COAST IN 1915.
Your committee desires to report that there seems to be a strong
sentiment in favor of holding a summer meeting on the Pacific coast
in 1915.
It finds the officers of the Panama-Pacific International Exposition
willing to co-operate in every reasonable manner, and it is informed that
the meeting can be held on the exposition grounds or at either Stanford
University or at the University of California. One member of the
Committee suggests the desirabliity of a group of entomologists travel-
ing together and arranging to have a field meeting at some Rocky
Mountain point. There would probably be no difficulty in arranging
for the necessary stop over privileges.
The western entomologists will be enthusiastic supporters of such
a gathering and it should be attended by a number of eastern men.
We feel it highly desirable to have such a meeting and therefore recom-
mend that arrangements be consumated. We respectfully suggest ©
including in the nominations for 1915, at least one vice-president who
would be in position to serve as chairman of the meeting in case a presi-
‘dent was unable to attend.
Signed, EB P2REtr, W. M. WHEELER,
V. Lo KEELOGG, AG, Dok, COCKERELL:
A. J. Coox.
On motion, the report was ordered accepted and printed
and the committee was ordered continued as a Committee on
Arrangements with the addition of Dr. E. C. Van Dyke of the
University of California.
The following amendments to the Constitution submitted
at the Cleveland meeting of the Executive Committee were
read —
Article IV, Section 3. The President shall represent the Society
upon the Council of the American Association for the Advancement of
Science until such time as the Society shall be qualified for representa-
tion by two councilors, in which case the second councillor shall be
elected from the Fellows by the Executive Committee.
106 Annals Entomological Society of America [Vol. VII,
To be amended to read as follows:
Article IV, Section 3. Councillors to the American Association.
The President and the preceding Past-President shall represent the
Society upon the Council of the American Association for the Advance-
ment of Science. In case of the death or resignation of either or both
councillors, the vacancy shall be filled by the Executive Committee.
Article V, Section 3. Election of Officers. All officers shall be
elected by ballot at the Annual Meeting for the term of one year and
shall be eligible for re-election. The term of their office shall commence
with the first of June following their election.
To be amended to read as follows.
Article V, Section 3. Election of Officers. All officers shall be
elected by ballot at the Annual Meeting for the term of one year and
shall be eligible for reelection.
On motion, the amendments were adopted.
The Executive Committee at the Cleveland meeting sub-
mitted the following addition to the Constitution:
ARTICLE. VII.
SEcTION 1. Publication—The official publication of the Society
shall be known as the Annals of the Entomological Society of America.
Each volume shall consist of four fascicles and the first fascicle of each
volume shall contain the proceedings of the Annual Meeting.
SECTION 2. Editorial Board——The publication shall be under the
charge of an Editorial Board consisting of ten members, one of whom
shall be the Managing Editor. The Managing Editor and his associ-
ates shall be responsible for the selection of the material to be published.
SECTION 3. Election of Editorial Board.—The members of the Edit-
orial Board shall be elected by the Executive Committee. Each
member of this board, excepting the Managing Editor, shall serve for
three years or until his successor has been elected, three members
retiring annually.
SECTION 4. Report Managing Editor—The Managing Editor shall
present a report at each Annual.Meeting to the Executive Committee
and the accounts of his office shall be reported upon by the Auditing
Committee.
On motion, this additional article to the Constitution was
adopted.
Mr. H. H. Lyman read a letter from President Bethune
expressing his regret at being unable to be present at the
meeting.
Pa;
1914] Proceedings of the Atlanta Meeting. 107
On motion, the Secretary was instructed to send a Night
Letter to President C. J. S. Bethune, extending greetings
from the Society, their pleasure on the recovery of his sight,
and their regret at his inability to attend the meetings.
The following communication submitted to the Secretary
by Mr. Nathan Banks was read :—
Inasmuch as there is no independant society in this country able to
publish large works on Entomology, and since there are even now
manuscripts awaiting printing, and with time there will be more, I
suggest that the Entomological Society of America found such a soci-
ety. This Society to be known as “The Thomas Say Society.” Its
object to publish catalogues, revisions, and monographs of North
American insects. That it be authorized to solicit and collect money
for a permanent fund, the interest on which shall be used for the print-
ing of said works. That the Society shall be controlled by a board of
five entomologists, chosen by the Executive Committee of the Entomo-
logical Society of America. Each member to serve five years, the first
board to have one member for one, two, three, four, and five years,
thereafter one selected each year. That all money received for sale of
publications be added to the permanent fund. That said board of
control shall select whatever officers they deem necessary and have
authority for accepting articles for printing and disbursement of funds.
On motion, the President was directed to appoint a com-
mittee of three to consider ways and means for the establishment
of such a society. The President appointed the following com-
mittee :—Nathan Banks, Chairman, H. H. Lyman, and Morgan
Hebard.
The following papers were then read :—
James Zetek, Panama Canal Zone—The dispersal of Musca
domestica.
William Moore, University of Minnesota—A comparison
of the enemies of Toxoptera graminum in South Africa and the
United States. Presented by F. L. Washburn.
Robert Matheson, Cornell University—Life-history notes
on Psephenus lecontei and Hydroporus septentrionalis. Read
by title.
V. E. Shelford, University of Chicago—The sequence of
color changes during ontogeny in Cicindela.
R. W. Leiby, Cornell University—Notes on the external
anatomy of some Pentatomide.
L. 5S. Barber, Florida State College for Women—The
biology of Gelechia gallesolidaginis with some reference to
some of its parasites.
108 Annals Entomological Society of America [Vol. VII,
A. F. Conradi, Clemson College—A little known wire-
worm, Horistonotus uhlert.
Leonard Haseman, University of Missouri—The life-history
of a species of Psychodiz. Read by title.
A; TB): MacGillivray, University of Illinois—The Structure
of the thorax in generalized insects.
James Zetek, Panama Canal Zone—Behavior of Anopheles
tarsimaculata Goldi. Read by title.
J. T. Lloyd, Cornell University—Life-history of Elophila
magnificalis, an aquatic lepidopteron. Read by title.
The following exhibits were shown:—
J. S. Houser, Ohio Agricultural Experiment Station—
Conwentzia hagent Banks, a coniopterygid.
E. L. Worsham and J. Chester Bradley, Office State Ento-
mologist of Georgia—Collections of Coleoptera and Odonata
from Georgia belonging to Georgia State Board of Entomology.
J. Chester Bradley, Cornell University—Photographs of
the Okefenoke swamp.
S. B. Fracker, University of Illinois—Setz of microlepid-
opterous larve.
James Zetek, Panama Canal Zone—Blue-print upon which
was shown the more important data obtained in the study of
the behavior of Anopheles tarstmaculata Goldi.
The Annual Public Address of the Society was given on
Wednesday evening, December 31st, at the Atlanta Medical
College by Dr. Ephraim Porter Felt, State Entomologist of
New York, on the Subject of Gall Insects.
A smoker for the entomologists and zoologists in attendance
at the meetings was held at the University Club on Thursday
evening January Ist, Mr. E. L. Worsham and the staff of the
State Entomologists office acting as hosts.
On motion, the Society adjourned to meet in one year
with the American Association for the Advancement of Science,
at Philadelphia, Pennsylvania.
ALEX. D. MACGILLIVRAY,
Secretary.
ar
_ NOTICE TO MEMBERS AND CONTRIBUTORS.
The Annals of the Entomological Society of America, pub-
lished by the Society quarterly, includes the Proceedings of the
Annual meetings and.such papers as may be'selected by the
Editorial Board. |
Papers may be submitted to any member of the Editorial ©
Board and should be as nearly as possible in the form desired as
final, preferably typewritten, and illustrations must be finished
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CONTENTS OF THIS NUMBER.
TriccErson, C. J.—A Study of Divontieee t Erinacei
(Mayr) ane dts Gall gu ee ae Sk oe eee I
CuILps, LzRov—The Anatomy of the Diaspinine Scale
Insect Epidiaspis Piricola (Del Guer) .--.-- aS ois AT
GILLETTE, C. P.—Some. Pemphiginae fener ; |
Species of Populus in Colorado.....--.-...- ---- 61
ZETEK, J.—Dispersal of ‘Musea Domestica Linne. errr) 70
Houssr, J. S.—Conwentzia Hageni Banks...-...... : 73 2
| -Moore, Wu.—A Comparison of Natural Control of : |
oxipeers Granimum in South Africa and United
Bre ts Aas Cari Be Ue) avr ae ee ah) 74 pe
Howarp, L. Oo.” —Report on Parasites bed ete a iB are ie ‘ 86. : a Tae Dla
FERNALD, H. T.—Notes on Some’ Old European
OMECHONS ot po ATs Te Ae 89
MacGILLivrayv—Proceedings of the Atlanta Meeting Etec a, |
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Proceedings of first three meetings; Constitution, By-Laws and List of
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WHEELER, WM. M.—Polymorphism of Ants... 2.5.0 c cece cece cere eencecnsces .30
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SEVERIN, H. H. anp SEVERIN, H. C.—Anatomical and. Histological Studies
of the Female Reproductive Organs of the American-Saw Fly, Cimbex
Aimericanay Beach s cae We SEO Ra OG a els yeahh Sree -25
Feit, E. P.—Some Problems in Nomenclature. .....ccc.c. sees ecececeececcee -10
Hammar, A. G.—On the Nervous System of the Larva of Corydalis cornutaL .25
BRADLEY, J. C.—A case of Gregarious Sleeping Habits among Aculeate
PRP MIEOOD tera eases Vk so dis Hoel aw Wane fast coe deta oes eek Sela ecbtl sab wate cols 10
Davis, J. J.—Notes on the Life History of the Leafy Dimorph of the Box- .
elder Aphid, Chaitophorus negundinis Thos... 2.2.2... cect eccccccccses -10
HAMBLETON, J. C.—The Genus Corizus, with a Review of the North and
Middle American Species oe 8 Pe tsb ces obicn ei cle oe ois eae ee bps owe b's 20
Grravutt, A. A.—Biological Notes-on Colorado Potato Beetle............... .25
Grrautt, A. A.—A Monographic Catalogue of the Mymarid Genus Alaptus.. 20
SEvERIN, H. H. and Severin, H. C.—Internal Organs of Reproduction of
Male AML goo ties veh iaeb Me Tea eaiais ld hem xine ke pIacere Naw Rees “acta Caleta 15
Smit, C. P.—A Preliminary Study of the Arane Theraphose of California... .75
DAVIS, J] >—Studies on Aphididayy eis 30h. aa ls salpisien opealney ep ee cma 8 TN A
Ritey, W. A.—Muscle Attachment of Insects... 22.5... eececce ees e eee ececes 15
NEEDHAM, J. C.—Critical Notes on the Classification of the Corduliinze
COT Ea) sees eae Ge ply et orcheeins Cereie a paver oe ths © atamadetatee.-¥ ag die Gas le 15
Howarp, L. O.—A Key to the Species of Prospaltella with Table of Hosts
and Descriptions of Four New Species.......... Fae cista‘cia ss bina ays alye's rary |
Hoop, J. D.—T wo New Species of Idolothrips......... cc scceeeeeccens eephe erowee
Address
ANNALS: ENTOMOLOGICAL SOCIETY OF AMERICA,
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ANNALS
OF
The Entomological Society of America
Volume VII §-U-NoEF ES 14 Number 2
A STRUCTURAL STUDY OF THE CATERPILLARS:
lil, THE SOMATIC MUSCLES.
Wo. T. M. Forpss, 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
7,
* Po rol >"
Sion 2 | M ust Vw
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 ina 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 1s 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 1s 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
aa 4
baer
1914] A Structural Study of Caterpillars. 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.
Ce 82 CXxXXIXa
D, E 1,4,5 139-140
A On 140b
a 22 CXXXIX
(CdS al G 61, 62 CXXXIV
B, 7; 5, etc. 35, 76, 80, 81 CXLI
a, b 16517, 21 CXXXII
& 19, 20, 26 CxXXXIla
c 18. CXXXIIb
Pebes Ue
ip Wetec, (On le 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. C+ 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 C+ 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
142 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, <8, tj , -vV, x; 6, 0, x) Nie, » ane es tue etere
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 6 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.
Rectus (1) A A 1 70 37
B B 1 70 37
Oblique segmental Cc Cc 8 70b 37b
E E 8’ 37c
D D 5 70a 37a
3 F F 5 70a 37a
Gai H 4
; ; a (1st belly) G Ga
| Shorter segmental I I
To intercosta K K 61
4 From precosta L, M, Li, 9-11
i To precosta S S 62
4 ‘Ie its 63
‘ From intercosta QO O 12, 138
| R R 14, 15
To antecosta d (part) A (part) 64, 65 (part)
INTRANEURAL (2) c Ee 18 4
LONGITUDINAL VENTRAL, SUPRANEURAL
Segmental a a 21 LXXII XXX
b b 17 “ “
é d (16) «“ “
et e 22 F
d ik 20 TRO IES Ae EXER.
g g 16 (19) « «
h h 23 es
To mid-ventral line behind
coxe 1 26 Fz
LONGITUDINAL VENTRAL, SUBNEURAL
To mid-line in front of coxz i i IVb (8)
Crossed to front of coxa k k 24 Viens)
Short anterior transverse (4) Z Z 25 HEP EON (|
From mesosternite at mid-
ventral line n n 28 i
From precosta m a (3rd belly) 77
Short fibres from leg q q 79
u u 57 Wy od ila
Vv Vv 58
TRANSVERSE; TO PLEUROSTERNAL LINE
Antecosta to prosternite B B 49
Precosta to precosta, super-
ficial 6 49
Precosta to prosternite, deep v pv
Precosta to mesosternite be io 72 XVIIIa (8)
Antecosta to posterior in-
cisure € Y 66 xXXX1Va
Antecosta to mesosternite r r
te XVII (3)
art)
é k, & 50 XVIIIa (8)
a (2d belly) 76
n 40 XXXVIb
Ww 41, 42
(posterior bellies)
Intercosta to mesosternite K,
Anterior incisure to precosta vy
To anterior incisure “n
Ww
114 Annals Entomological Society of America [Vol. VII,
LYONET : LUBBOCK BERLESE
Meso-. Meta-. Meso-. Meta-.
ACROSS PLEUROSTERNAL LINE
Intercostal region to leg 6,0 e, 6,0 46, 51 5a (3)
Posterior incisure to leg t t 55 XVIIB (8)
Superficial on incisure 0 ) 35 XXXVI XVIIa
Deep on incisure x x 34,38 XXXVla XVIIIa
BELOW PLEUROSTERNAL LINE
To leg Ss Ss 78 Ila, 6a
To near midventral line p Pp 73-75
SPIRACULAR (rudiment) 1
Notes: (1) Muscles droits of Lyonet.
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.
(
(2) Passing between the two longitudinal connectives.
(
(
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 0, 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
ore ere.
Merwe
PB an boots om
1914] A Structural Study of Caterpillars. Gh
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 is
C 2 :
Segmental D 4 $
5 “
From antecosta, oblique F(4) 8 xX
7 “
From antecosta to antecosta E 6 IX
From precosta if 9 XI
L 10, 11 &
From intercosta OR 12) 15 XVI
LONGITUDINAL VENTRAL, SUPRANEURAL.
Rectus ce 18 2
b 17 *
d(3) 16 iS
Antecostal, oblique f’ (in Malacosoma only)
Longitudinal segmental e 20 2
ff 19 £
To antecosta of following segment f 25 IV
From antecosta, oblique h a
LONGITUDINAL VENTRAL, SUBNEURAL.
Antecosta to antecosta, longitudinal a 21 III
299
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) lap
SPIRACULAR M 45
TRANSVERSE SUPRATRACHEAL.
Acrosternal tf) 3 XVII
At precosta a(2) 36 ¢
116 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, € HU a2 XVIIIa
Crossing the pleurosternal line.
Spiracle to precosta if 39 5b
Propleurite to precosta m(sometimes) 42 5b
q (rarely) AO, 41
Mesotergite to parts of leg Boy 46-50 5a, 58
Short posterior muscles (ens 33-34 XVIII
Below the pleurosternal line.
Behind precosta q, m 40,41(42) 5ba
Across leg in front n 43 6a
Across leg behind (from w) Tr 53-54 68
From fold or yw to leg (part) baa
From fold or y to anterior edge: of
following segment ¢(part) 31-32
Zz 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
6 proper spread fan-like from the acrosternite to various points in
the acro- and protergites.
(3) To antecosta of following segment in Pheéocyma.
(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,
a
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 its 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. y, 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 @, 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 6 (representing y) and 6 are also nearly simple—
evidently the proleg is merely the small area enclosed by
fi arid er.
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:
Cossip#; Cossus cossus by Lyonet.
NOTODONTID2; Pygera bucephala by Lubbock.
LASIOCAMPID; Malacosoma americana and disstria in this
paper.
NocTuDID#; Two species, probably of Noctua and Nephe-
lodes, both trifidee and a Pheocyma (quadrifide) in this paper.
+ SPHINGIDA; Delilephila lineata, briefly by Berlese; and
Sphecodina abboti 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 tracheze, 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 Cosside 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 disstria, 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, EK, a;, a, and o,; C*, L-and Ge, @.s and Wend
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 S 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
ac Fe
1914] A Structural Study of Caterpillars. nee
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 A8, 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.
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
Gr H only m 3d belly of a
Ist belly of a G OY 2nd belly of a
en e only € OY
d f 6,¢ 65,9
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.
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.
Luppock, J.: Arrangement of cutaneous muscles of the larve 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 Detlephila
(Sphingide) and numerous non-Lepidopterous insects.
Sadars
2 eae
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. T2. Second thoracic ganglion.
gang. A'. 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
m 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.
a. 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:
b 2,
4, 3,
5, 6.
Plate XXKI—Middle segments of Sphecodina abbotu, dissected in the same
order, in four layers.
tie irachea:
Plate XXII—Middle segments of Malacosoma americana dissected in four
layers in the following order:
7g &
3, 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 XX V—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. a
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.
g, 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. ex. 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.
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OBSERVATIONS ON THE LIFE HISTORY AND HABITS
OF HYDROMYZA CONFLUENS LOEW., (DIPTERA).*
By Pau 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.
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,
advena 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. N. 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
as
1914] Life History of Hydromyza Confluens. fod
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 pupe 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
Los 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 pupe, 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 1n 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 variable,
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 pupz 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,
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 (8) 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 N. 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 Dzptera 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 JN.
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
def
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; (8) 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 Supplementary List of the Diptera of New Jersey. Ent.
News, 15:157-163.
Lovell, J. H., 1898. Three Fluvial Flowers and their Visitors. Asa Gray Bulletin,
6:60-65.
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.
eee from the U. S. National Herbarium, 16:63-108, 12 pls.,
40 figs.
Needham, J. G., 1908. Notes on the Aquatic Insects of Walnut Lake. Appendix
Ill. 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. Cockerell 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 comstocki 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............ 3
Bastidorsalssesment not upturmed aitvapexs ys aaa ace eee ern ee 7
3. Last dorsal segment very sparsely punctured at base...... obtusiventris n. sp.
wast dorsalesecinent closely, punctured at bases... eee eee ee eee +
4. Punctures of first dorsal abdominal segment separated by less than a
punctunedwidila yay waren he ee ions aya cue Pcie cent oe es ber tele eer SL eran mee sree 5
First dorsal abdominal segment sparsely punctured...................... 6
5. First recurrent vein received by second submarginal cell almost half as
far from base as length of first transverse cubital in the o and slightly
TeSG Pia <= piesa Sera ree Pept ies AESc ieee pie coe ea ea eect we WES 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 @ and
Sill bol Lesioks bois tite sah ek en Wiebe rt RGN Ph Ite he Se atte y 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
alone Margin... Ae osa Sake cave Fees ORE aed es De he ce ee 8
Hast. ventral segment not, strongly fringed ys.-..42c- 4.0 eee 12
ts eae 31D 25a 0) (21 6) gee a re a ae ae ALA eh RRR TE Sen Fn tere reer per aco Onatc cue 9
begs except. coxae,: ted iy: shh toss Gee EER ao nO ee 11
9. Last dorsal segment without a median carina or this only indicated
Ch AG] OP ae Roane terete ae eh ayant earns iy ih bis WR Mn eens be ceded ade. 10
Last dorsal segment with a median carina...................-.- angelica Ckll.
10. Penultimate ventral segment with small punctures interspersed among
the dargersonesy See ean acs See gee ee See Eee ee apacheorum Ckll.
Penultimate ventral segment without smaller punctures among the
Othersit Fhe Ale Peg Meirs, eels a ee cee ee EE toner at alternata Sm.
11. Fourth antennal joint distinctly longer than third; last ventral segment
with subparallel sides and a broadly rounded apex........ 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
. ie
1914] The Bee Genus Celioxys. 149
12. Last dorsal segment near apex with two small flattened projections
Sapna yy eves Whreshert vn. va ce sire SA Gia w ees Oe piercei n. sp.
Washidorsalesesmenti without projections on) disk.) oo... cae ee ieee ees 13
1S» [beesy Watley occ cgmend ok Qe blo ton) on Og eee elo e ERO ie ices Oe ee 13a
ULES CSD). 5 Sg a ects on ayta ce chee Ce ee ORTHO OREN GCR eke Cae 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
SUNG iti Chin SURANGA Aba ee ate < icc ieie ag alae aetes ee ieee he 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 appréssed pubescence......................0. 17
sonata Ome Wabi Onl ye CLEC Em Ma IiG type ccisl leit ays sel or tere-istee Dela esse dase 18
17. ‘“‘Scutellum medially produced into a tubercle”’................. aperta Cress.
Scutellum medially not produced into a tubercle............. grindeliae Ckll.
IS Gi > TPAUABYESTCVESETOE EN LAME Oi Cyne er eeone Oc eee nee eke eee teers 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
APR PL sec ee ar ae PLS AL PR Rey ght ay ene, et BTC NE RPA hile gad 20
Lop SUIS ELE STRICT (Gy SRR I Rca RE er Se ge a 21
20. Transverse furrows on segments 2 and 3 deep; punctures on middle of
21.
22.
23.
24.
25.
26.
27.
28.
29.
segment 2 basad of furrow close, separated by about a puncture width;
arcuate edge of pronotum much more strongly produced, translucent;
WEES SUNOS E of at0 bis anereae ae est na Perc Cates aT eee ea ee Recs 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
AULA Nie iaavoneey Abbe All homtol eh lke Goes eieae Mee Oe lei ai aaa Re ae ee sayi Robt.
Clypeus medially triangularly produced and somewhat reflexed, banksi n. sp.
Clypeus apically truncate, gently round, emarginate or tuberculate...... 22
Basal abdominal segment at least entirely red... 0.2.0 0.0 u. ee eee tee eats 23
Basal abdominal segment black (at most with sides red)................ 25
Scutellum sparsely punctured, somewhat produced medially and slightly
ELD BCLS TE DR ne eich Acree 2 An a menthae Ckll.
BICC Mi eeran Olese yrs pL AC HEME | otis ssa hs Gosiacs. cla Pass acy weclmtahe pve saves ine no 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.
ILDNRENTTEOULG 5g ole c'o took en © UTA UOT ONC RTE eV NS RE COSI ae a een gt 26
lEcoomaimledictueimorareGa kine. ceramic lets ices ct Sere bed tig tinele bo aces 31
Scutellum with a strong median projection................... germana Cress.
BemeIMimImedialive dh Most iihercilate nc. ici keek eesti dees what ele es eds 27
Third joint of antenne hardly longer than second, about half as long
DG MPO otek os Deeind Chih eae eC ere asteris n. sp.
Third joint of antennz distinctly longer than second, almost as long as
Se EN ae PT RA hg ryt TiS eS ears Cecio SA Ra. da Seok Pages Soden ee 28
Last dorsal segment narrowed at almost a right angle; first abdominal
Seriment closely punctured) laterallys...3..0. 20.2406: coquilletti n. sp.
Last dorsal segment not narrowed at almost a right angle, at most at
ARVER VEO DLUSSr ate CMmmry Pacer cr srs me Aer SA ee REAR NO instal s shen Sica tla Mts 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
DemiAG@ewitiwicuicl meee Merete sitet er tienen won, Se Se cee tos Suvi sPtm ah Nai chee 30
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
punctire-widithy apante os askeer ce oat wie ee Sere ee ee pratti n. sp.
Face without dark bristle-like hairs; first abdominal segment sparsely
jO(6UaYey abi geCO uae Hier g2K6 bp oe ca er eee Rn ei rh ee cet ei ae octodentata Say.
31. Last dorsal segment strongly angularly narrowed..:..................+- 32
Last dorsal segment at. most roundly narrowed.:.........0.......:2-+00- 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.
3a: .< Larger: 13 min long V0 Ale Re. PAE Foe me ree eee lucrosa Cress.
Smaller, hardly 12 mm. Mae abdomen more slender and more closely
punctured wh ne cee L eee ge Rae oOo Ne i Re eet anne 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 antennze 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 éxtreme 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
1914] The Bee Genus Celioxys. 151
Coelioxys texana Cresson.
For comparison with C. huntert 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; the 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 antenne;
a £
Fig. 2. C. hunteri Cwfd. (a) Last ventral segment
(b) antennal joints 2-5 of female.
second joint of antenna (not counting bulbous base) much 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; coxe
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 pleure with long
white hair; abdomen shiny, with sparse rather coarse punctures, last
segment with a silky lustre the 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 hunteri have this segment narrowed
apically and consequently pointed at apex; in alternata the last
dorsal segment is shiny and with sparse small punctures.
Ceelioxys 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; antenne 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;
pleurze 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—Specimen: 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 11.5 mm. Black, with the tegule 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; pleurz with dense long white hairs; wings dusky,
with the apical margin more densely infuscated; apical margins of dorsal
154 Annals Entomological Society of America [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. ammaculata 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.
Ccelioxys 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.
1914] The Bee Genus Celioxys. 155
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 tegulze; 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.
Lype—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.
Ccelioxys 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 10.5 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 antenne 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 antenne 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. ; THe
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.
a set
4
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,
‘fon 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 tegulze and
legs; face below antenne 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 tegule; 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 ;
4
ies
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 pleurze with
long white hair; wings slightly dusky with the apical margins somewhat
more so; spines on anterior coxze 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 octodentaia 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 pratiz 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.
Ccelioxys 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
Ccelioxys 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 tegule
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 CHARLES H. T. TOWNSEND,
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. Conopidez), 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 bee eh 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 nonconnectant
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 Muscide, 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.
{Neither hypopleural nor pteropleural hairs; sternopleurals 3 or
Anthomryicides 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 ts 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,
Cuterebrida, 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.
Male with accessory glands always more or less developed, at
Higher least their rudiments visible; female usually with a large
Muscoidea number of ovarioles—All Muscoidea down to and including
the Calliphorine,
Male without accessory glands, with very long and curled ejacu-
Typical latory duct whose head is developed into a very elongate
Muscoidea vesicula seminalis; female without uterus, with many function-
ing ovarioles—Musca, Muscina, Synthesiomyia.
Same as preceding, but ejaculatory duct of male much shortened,
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—Gen. Indet.
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, Synthesiomyia, 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.
a Gay Mita tan yas eh Faas
(fd }, he 4
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CONTENTS OF THIS NUMBER.
A
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o
Forbes, Wm. T. M.—A Structural Study of the
- Caterpillars: III, The Somatic Muscles. -.....-
10g
Wetcu, Paut $.—Observations on the Life History ©
and Habits of Hydromyza Confluens Loew.,
(Diptera) i Aap aL BUSS sR ken pag TN UATE RTE ct ENC HE Nd
CRAWFORD, J. C.—Some Species of the Bee Genus
VACOCHOMVS ie at et OLS as Mae A acct
TOWNSEND, CHARLES H. T.— Connectant Forms
Between the Muscoid and Anthomyioid Flies...
135
148
160
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‘PUBLISHED QUARTERLY BY THE SOCIETY.
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The Entomological Society of America. :
FO UNDED 1906.
OFFICERS 1914. .
President P. CALVER® fos eR aes 8 cyan bel Philadelphia, Pennsylvania
|. First Vice-Presideni—Jas. G. NEEDHAM... 0) 20 noe ee Ithaca, New York
Second Vice-President—C. GORDON HEWITT... <.. avant Seth eke Ottawa, Canada
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Price List of bata |
Annals, Vols. 1, Il, III, IV, V, and VI complete, each RA Se Br oa oak ote SCO
Annals, Separate Parts except as sent edch essere cae ae ge 2h ARS Oe 8 "Jo.¢ [OO
Annals, Vols. Land II, Part 3, each...2.5...... Poe apie S Aaa Neel Seely Bia Sic vate .50
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REPRINTS FROM VOLUME I.
Proceedings of first three meetings; Constitution, Bylaw and List of
NGS ME OBAD EE Hs <(Csin’s sie’ lee < Fitlon ins Wer y SuNay Oc ale eer wask Matias «Bea ale'a fe Sipinbeale groie 0285,
WHEELER, Wu. M,—Polymorphism of Ants... 2.0. ee eee cee Pole SNE Scene sh
‘Ossorn, HERBERT—The Habits of Insects as a Factor in Classification..... _.20
SEvERIN, H. H. anp SEVERIN, H. C.—Anatomical and-Histological Studies
of the Female Reproductive Organs of the American-Saw Fly, Cimbex
Americana, Leacimis say ole WOR as REN Rice ak Bala Ok ia OE on oe ice «25
Feu, E. P.—Some Problems i in Nomenclature .i5) \Paeccv ose o2 is codes -10
Hammar, A. G—On the Nervous System of the Larva of Corydalis cornutaL .25
BRADLEY,, J. CA case of Gregarious Sleeping Habits among Aculeate
WEL VIMOMOP GOLA T. 5:5 tains toe mire maaan Re wists ore akin wee bislale goa nee wee ile .10-
' Davis, J. J.—Notes on the Life History of the Leafy Dimorph of the Box-
elder Aphid; Chaitophorus negundinis Thos....... 00.0 cece de eee cecene - 10
-HamsBietTon, J. C.—The Genus Corizus, with a Review of the North and é
oy Middle American ‘Species, 4. tk ah cies Maa ss voce es pene e slows 20
-Grrautt, A. A.—Biological Notes on Colorado Potato Beetle...... Fe IPA 20.
Grrautt, A. A.—A Monographic Catalogue of the Mymarid Genus Alaptus.. .25
SEveRIN, H. H. and SEVERIN, H. C.—Internal Organs of Reproduction of
Male Sawfly. caso os Ort pis Gen An CSS, hedge aa mide a binehes ete 15
$mitu, C, P.—A Preliminary Study of the Aranze iscsi oe of California. .75
Davis, J. J.—Studies on Aphidide.......5........ CAA BEERS Sal CRE a UE SNE 32025:
‘Riey, W. A.—Muscle Attachment of Insects. 2.2.0.2... ce ceveee ee eee ewes Bia eo,
NEEDHAM, J. C -—Critical Notes on the Classification of the Corduliine
KOM Gna La YOON eC aay ES BN a Sake in oe 15
Howarp, L. O.—A Key to thé Species of Prospaltella with Table of Heats
and Descriptions of Four New Species........... NE rar ee ey bt ae BN bo
Hoop, J. D.—Two New Species of Idolothrips.. ks Shanes heeded seuee wae 10)
Address
ANNALS ENTOMOLOGICAL SOCIETY OF AMERICA,
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iin.
ANNALS
OF
The Entomological Society of America
Volume VII SEPTEMBER, 1914 Number 3
SPIDERS COLLECTED BY MR. C. WILLIAM BEEBE IN
BURMA AND BORNEO.
With Plate XXVI.
ALEXANDER PETRUNKEVITCH, PH. D.
Family Theraphoside.
Haplopelma Dorie (Thorell). One female from Kuching, Borneo.
Family Drasside.
Drassodes Drydeni n. sp. One male from Pongatong, Burma.
Drassodes ignobilis n. sp. One female from Tabu Pum, Burma.
Family Pholcide.
Pholcus phalangioides Fussl. One female from Wahsaung, Burma.
Family Theridiide.
Theridion sarapus Thorell. Two females from Pongatong,
Burma.
Dipoena tristis n. sp. One female from Tabu Pum, Burma.
Enoplognatha marmorata L. One immature female from Tabu
Pum, Burma.
Family Linyphiide.
Erigone longipalpus F. Two males and two females from Tabu
Pum, Burma.
Linyphia sp? One immature female from Tabu Pum, Burma.
Family Clubionide.
Clubiona tabupumensis n. sp. One female from Tabu Pum, Burma.
Clubiona sp? One young from Tabu Pum, Burma.
Clubiona sp? Two young from Wahsaung, Burma.
Palystes sp? One young from Wahsaung, Burma.
169
170 Annals Entomological Society of America __[Vol. VII,
Family Argiopide.
Nephila maculata F. One female from Wahsaung, Burma.
Nephila clavata L. Koch. Two females from Tabu Pum, Burma.
Leucauge tesselata (Thorell). One female from Pongatong,
Burma.
Gasteracantha arcuata F. One female from Kuching, Borneo.
Gasteracantha frontata Bl. Two females from Kuching, Borneo.
Theridiosoma sp? One young from Tabu Pum, Burma.
Araneus microtuberculatus n. sp. One female from Tabu Pum,
Burma.
Araneus Beebei n. sp. One female from Wahsaung, Burma.
Family Thomiside.
Philodromus tabupumensis n. sp. One female from Tabu Pum,
Burma.
Bomis sp? One young without abdomen, from Wahsaung, Burma.
Porrhopis sp? One young from Wahsaung, Burma.
Family Lycoside.
Lycosa stictopyga Thorell. One female from Tabu Pum, Burma.
Lycosa (Pirata) sp? One immature female from Tabu Pum, Burma.
Lycosa sp? One young from Tabu Pum, Burma.
Family Oxyopide.
Oxyopes sp? One young from Wahsaung, Burma.
Oxyopes sp? Two young from Wahsaung, Burma.
Family Salticide.
Evophris sp? One young from Wahsaung, Burma.
Evophris albopatella n. sp. One male from Wahsaung, Burma.
Cobanus Beebei n. sp. One male from Central Borneo.
Ballus tabupumensis n. sp. One female from Tabu Pum, Burma.
Attulus sp? One young from Wahsaung, Burma.
Thiania sp? One young from Wahsaung, Burma.
Description of new species.
Drassodes Drydeni* n. sp. (Plate XXVI, figs. 1 and 2).
Total length 7.4 mm. Cephalothorax and all appendages brown,
abdomen grey. Sternum oval, pointed behind, broadly truncated in
front. Lip much longer than wide. Lamine maxillares strongly
impressed. All femora slightly thickened. Cephalothorax much nar-
rower in front than in middle. Anterior row of eyes recurved, posterior
procurved. AME half their diameter apart, subcontiguous with ASE.
Eyes of second row aequidistant. ASE separated from PSE by less
than half their diameter. AME slightly larger than ASE. Anterior
row viewed from in front curved downward. Clypeus as high as AME.
First tarsus and metatarsus with a heavy scopula. Second tarsus with
a scopula only in its distal half. Second metatarsus without scopula.
* In honor of Mr. John Dryden Kuser.
1914] Spiders from Burma and Borneo. 171
Heavy spines on all legs. Femur of pedipalp with a short blunt sub-
terminal apophysis on the inside. (Plate XXVI, fig. 2). Copulatory
apparatus with an extremely long and thin embolus (Plate XXVI,
fig. 1).
One male from Pongatong, Burma.
Drassodes ignobilis n. sp. (Plate XXVI, fig. 3).
Total length 5.9 mm. Cephalothorax and all appendages brown,
abdomen dark grey. All femora slightly thickened. Tarsi without
scopule. Sternum oval, broadly truncated in front, pointed behind.
Lip longer than wide. Laminz maxillares strongly impressed. Fourth
legs with spines. All femora with two long, upright spines in median
dorsal line. No spines on other joints of first, second and third leg.
Anterior row of eyes recurved, posterior row procurved. PME about
their diameter apart. PSE about half diameter from PME. Eyes
of anterior row equal in size, contiguous. Anterior row viewed from in
front curved downward. Side eyes separated by about half their diam-
eter. Clypeus as high as AME. Epigynum as figured, much higher
than wide.
One female from Tabu Pum, Burma.
Dipoena tristisn.sp. (Plate X XVI, fig. 4).
Total length 4.8 mm. Cephalothorax and legs reddish brown,
Abdomen dark greyish brown with a narrow dark median line and two
pairs of transverse whitish bands. No spines on legs. Sternum
triangular. Anterior coxz widely apart. Lip wider than long. Ante-
rior row of eyes strongly recurved, posterior row straight, longer than
anterior. Eyes of posterior row aequidistant and equal in size. Side
eyes contiguous, equal. AME smaller than the other eyes. Eyes of
anterior row aequidistant. Clypeus as high as quadrangle. Epigynum
as figured.
One female from Tabu Pum, Burma.
Clubiona tabupumensis n. sp. (Plate X XVI, fig. 5).
Total length 7.8 mm. Cephalothorax and all appendages yellowish
brown, abdomen grey. Anterior row of eyes much shorter than posterior
row, slightly recurved. Posterior row almost straight. Quadrangle
wider than long, much narrower in front than behind. Eyes of anterior
row aequidistant, separated from each other by somewhat less than
their diameter. PME somewhat farther from each other than from the
PSE. Distance between the PME equal to about 2% their diameter.
Clypeus not more than half the diameter of the AME. Sternum a long
oval, pointed behind, truncated in front. Lip much longer than wide.
First and second tarsi and metatarsi with a thick scopula. Similar scopula
on distal half of third and fourth tarsi. First tibia with 2-2 long spines
below, first metatarsi with 2 long spines at base below. Third and
fourth tibiz with a row of 3 spines in median line below, 2 laterals inside
172 Annals Entomological Society of America [Vol. VII,
and 2 laterals outside. Third and fourth metatarsi with spines below,
above and laterals. Epigynum very small, considerably in advance
of genital slit.
One female from Tabu Pum, Burma.
Araneus microtuberculatus n.sp. (Plate X XVI, figs. 6, 7, 8).
Total length 2.8 mm. Cephalothorax high, without groove or
stria. Abdomen with two tubercles or shoulders in front and two
prominent tubercles behind. (Plate XXVI, figs. 7, 8). Integuments
soft. Side eyes subcontiguous, on black tubercles. ASE much smaller
than PSE. Both rows of eyes recurved. Quadrangle wider behind
than in front, wider than long. PME much larger than AME. Cly-
peus lower than quadrangle, about twice the diameter of the AME.
Sternum triangular, broadly truncated in front. It is also truncated
behind, between the hind coxee which in consequence are separate. Lip
triangular, very wide. Chelz smooth, inferior margin with 3 teeth,
superior with 4 teeth. Pedipalpi with a claw. One dorsal spine at end
of all patellas. One inside lateral spine on first femur and tibia. Upper
claws almost cordate, with four teeth each. Third claw smooth.
Epigynum as figured. (Plate XXVI, fig. 6). Colorin alcohol: cephalo-
thorax brown with a median white spot. Chele brown, legs brown,
femora lighter than other joints. Sternim dark brown, lip and laminz
with tips of lighter color. Abdomen above mottled with white and
brown, tubercles dark. Sides whitish with three dark lines. Below
almost black. Spinnerets brown.
One female from Tabu Pum, Burma.
Araneus Beebein. sp. (Plate XXVI, figs. 9, 10).
Total length 2.6 mm. Cephalothcorax with a somewhat recurved
groove. Clypeus not half the diameter of the AME. Quadrangle
much wider in front than behind. Anterior row strongly recurved,
posterior row slightly recurved. AME are the largest eyes. Chelz
distinctly longer than thick. Inferior margin with 3 teeth, superior
margin with 4 teeth. Abdomen oval, considerably overlapping cephal-
othorax. Sternum triangular, widely truncated in front, produced
behind between the fourth coxze which are separate. Pedipalpi with a
claw. Legs with many spines. Two rows of long bristles below all
femora, especially on those of the first and second pair. Epigynum as
figured, brown and relatively very large. (Plate XXVI, fig.9). Color in
alcohol : cephalothorax greyish brown, legs yellow, sternum, lip, lamine,
pedipalpi and chele also yellow. Abdomen (Plate XXVI, fig. 10) above
grey with white iridescent spots, a transverse anterior black band and
a large median more or less triangular black spot pointed backwards.
Below grey with iridescent white spots.
One female from Wahsaung, Burma.
1914] . Spiders from Burma and Borneo. 173
Philodromus tabupumensis n. sp. (Plate X XVI, fig. 11).
Total length 4.8mm. Cephalothorax 1.8 mm. long, 2.0 mm. wide.
Legs 2134. Anterior row of eyes shorter than the posterior row and
recurved. Posterior row very slightly recurved, almost straight.
Anterior eyes about equal in size, AME farther from each other then
from the ASE. PME smaller than the PSE, the distance between the
PME much larger than between the PME and the PSE. Quadrangle
narrower in front than behind, about as wide as long. Clypeus 1%
diameters of the AME. Sternum longer than wide, emarginate between
hind coxz which are widely separate. Spines on all segments of legs
and palpi, except tarsi. First femur 2.2 mm. long, second 2.6 mm.
Color in alcohol: cephalothorax light brown with black marginal and
three black longitudinal parallel bands. A narrow, curved black band
runs in front of the eyegroup, touches the PSE on each side of the head
and merges with the median longitudinal band at the posterior margin
of the cephalothorax. Legs yellow, spotted above with dark brown.
Abdomen above light brown with two dark brown V-shaped spots
pointed forward, sides dark brown, underside altogether light yellow.
Epigynum as figured.
One female from Tabu Pum, Burma.
Evophris albopatellan.sp. (Plate X XVI, figs. 12, 13).
Total length 2.7 mm. Legs 4312. Inferior margin of chele with
one tooth, superior with a row of eight teeth. Of these only the prox-
imal two are large, while the others are exceedingly small. (Plate XXVI,
fig. 12). Cephalic part shorter than thoracic. Eyegroup wider in
front than behind. ASE very prominent. Eyes of second row minute,
situated in middle. Iridescent white scales above AME, between
side eyes on face and along the edge of the cephalothorax which is very
dark brown. First, second and third femur, patella and tibia dark
brown above and below. Fourth femur yellow with a dark brown spot
above at distal end. Fourth patella and tibia yellow above with two
lateral dark brown lines, below yellow. All other joints of all legs
yellow. Palpi dark brown except patella which is covered with white
iridescent scales. Sternum, lip and laminez dark brown with yellow
edges. Chelze reddish brown. Abdomen above yellow with two
parallel longitudinal dark brown bands and white iridescent scales.
Below yellow with a median dark brown broad band. Tibia of pedi-
palp with a curved apophysis, copulatory apparatus as figured. (Plate
XXVI, fig. 13).
One male from Wahsaung, Burma.
Cobanus Beebei n. sp. (Plate X XVI, figs. 14-18).
Total length without mandibles 9.0 mm. Cephalothorax 3.8 mm.
long, 3.1 mm. wide. Chel long, with long, curved fang. Length
of chelz without fang 3.8 mm. Superior margin with two teeth of
174 Annals Entomological Society of America [Vol. VEL,
which the subapical one considerably larger than the proximal. Inferior
margin with two teeth. (Plate XXVI, fig. 14). Legs 3142. Cephalo-
thorax high. Eyes of second row small, situated behind the middle.
Measurements of legs: First lec—femur 4.6 mm., tibia+ patella 5.5 mm.
metatarsus+tarsus 4.8 mm.; Second leg—femur 3.3 mm., tibia+ patella
3.5 mm., metatarsus-+tarsus 3.6 mm.; Third leg—femur 5.2 mm., tibia+
patella 5.1 mm., metatarsus+tarsus 5.8 mm.; Fourth leg—femur 3.0
mm., tibia+patella 3.7 mm., metatarsus+tarsus 4.5mm. All legs with
many heavy spines. Tibia of first leg curved, with 4-4 spines below.
First metatarsi straight, with 2-2 long spines below and laterals. A
heavy comb of black hair above and below first patella, a comb of
shorter hair on distal half of femur and one of quite short hair on back
of first tibia (Plate XXVI, fig. 15). Claws as figured (Plate XXVI,
fig. 17). Sternum longer than wide, lip not reaching half of lamine.
(Plate XXVI, fig. 16). Tibia of pedipalp with a straight apophysis, copula-
tory apparatus as figured (Plate XXVI, fig. 18). Color in alcohol:
cephalothorax red-brown, with two lateral white patches and a fringe
of brown hair around the eyes. Chele above red-brown, below black.
Fang black with red-brown tip. Pedipalpi yellow. First femur and
patella very dark brown, tibia yellow with dark brown distal end,
metatarsus and tarsus dark brown. Other legs brown, fourth leg
lighter. Lip and lamine dark brown, sternum reddish brown. Abdo-
men dark grey above and below.
One male from Central Borneo.
Ballus tabupumensis n. sp. (Plate XXVI, figs. 19, 20).
Total length 5.7 mm. Legs 1423. Cephalothorax flat and square.
Eyegroup wider behind than in front. Pars cephalica longer than pars
thoracica. Eyes of second row minute, situated considerably in front
of middle. Posterior row somewhat shorter than width of cephalo-
thorax. Chelz short and heavy (Plate XXVI, fig. 20) with a humped
back carrying a row of tubercles. Inferior margin with three teeth of
which the middle one is the smallest. Superior margin with a row of
12 very small teeth. Sternum much longer than wide, posterior coxze
contiguous. Lip longer than broad, laminze very long with emarginate
outer edge, wider at end than at base, inclined over lip. First leg much
heavier than the others. First femora dilated. First tibia with 3 - 3
heavy spines below and no laterals. First metatarsus with 2 - 2 very
heavy spines below, the first pair reaching beyond the middle of the
article and the second pair almost to the base of the claws, no laterals.
Claws with a single tooth. Epigynum as figured (Plate X XVI, fig. 19).
Color in alcohol: cephalothorax, chele and first leg very dark brown.
Abdomen and femora of second, third and fourth pair brown. Other
joints of second, third and fourth leg yellow with dark spots. Under-
side dark brown.
One female from Tabu Pum, Burma.
ia a sins
-
~-
=
1914] Spiders from Burma and Borneo.
EXPLANATION OF PLATE XXVI.
Drassodes Drydeni n. sp.—male.
Fig. 1. Copulatory apparatus of male.
Fig. 2. Femoral apophysis of pedipalp.
Drassodes ignobilis n. sp.—female.
Fig. 3. Epigynum.
Dipoena tristis n. sp.—female.
Fig. 4. Epigynum.
Clubiona tabupumensis n. sp.—female.
Fig. 5. Epigynum.
Araneus microtuberculatus n. sp.—female.
Fig. 6. Epigynum.
Fig. 7. Sideview of abdomen and cephalothorax.
Fig. 8. Dorsal view of abdomen.
Araneus Beebei n. sp.—female.
Fig. 9. Epigynum.
Fig. 10. Dorsal view of abdomen.
Philodromus tabupumensis n. sp.—female:
Fig. 11. Epigynum.
Evophrys albopatella n. sp.—male.
Fig. 12. Chelez.
Fig. 18. Copulatory apparatus.
Cobanus Beebei n. sp.—male.
Fig. 14. Chele.
Fig. 15. First leg.
Fig. 16. Sternum, lip and lamine.
Fig. 17. Claws.
Fig. 18. Copulatory apparatus.
Ballus tabupumensis n. sp.—female.
Fig. 19. Epigynum.
Fig. 20. Chele.
ANNALS E.S. A. VoL. VII, PLATE XXVI.
A, Petrunkevitch.
i+
THE RAVAGES, LIFE HISTORY, WEIGHTS OF STAGES,
VI.
Vil.
VIII.
IX.
NATURAL ENEMIES AND METHODS OF CON-
TROL OF THE MELON FLY (DACUS
CUCURBITAE COQ.).
Henry H. P. SEVERIN, Ph. D. Harry C. SEVERIN, M. A.
WILLIAM J. HARTUNG, B. S.
CONTENTS.
PAGE
MELO ETOTA na eN ee PP EON tec ears, eet er Soe niche wa < tds, seNeceporetacaue a hots 178
INGE ep elLOTiene, Sern at rene clei one Chen e aeioeure cise). Ate ale 179
Desempiiom OlmVirtanty cer yes eae onesie ak eb nea tae Cella 179
icidtObsenvationssmramo ump lcm ateh.t sacs sere ee © ie buen = 180
1, Oviposiiomin stems of pumpkin vines... 0.0 [2.0.0.6 ees 181
eo SOmipOstitenen: Petre siOl |GAVESs «os si o's ioe es oes 181
3) Oviposiion in pumpicia GOWELS. «0.6.5 e ss Se sl een Sea ws 181
Bm Gas Tae AT. PIUMIAIRSIAS. | Gea 5IS Sunless cares kis slew e oaae Tate 182
Pi Smeeliny ty asuOuS Cl wacen tee meaner ick errs uals «racic tga cts eis auye 182
(CRO MLERS EEE ae ROR aes oy Ge ecient a a ae 183
Renin WO MOW CLG ses eter tn cae oe wh OG ie eM aim andare tess Sa ee 183
Ba, LAjiry, CO pumpkins 10 he eee nie yee te ane bea 184
Seemirigiaiiys GO" SUEUR, DEANS. Meet ts wea ae weston ete aie eos Nee Xe 185
OO lsat compere wate ae Nes oe oats orscr ie) sry oye wane tanetoe aperalerel as 185
10. Number of melon flies bred from the) food! plants; 4... 46 = 185
ehew shore agelt toe ee at ne roe ete Sea ets ade Seles ee nie den Nes 186.
11. Methods of inducing oviposition.......................455 187
i2iebaciiletonistleshon CHESOVApPOSLCOD: >. cnc gin ot ek cn ee eate ole = 188
fh Prdpeesian OViposttionrrs ot anne s sued reste ctuts een mee 188
14. Number of eggs deposited in a receptacle.................. 189
[ }eMINTMbDeTIOlmipe CkOSHmOVATICS a. cscka enti dae ess tite ses 189
16. Duration of the egg, larval and pupal periods.............. 189
17. The effect of temperature on the egg period............... 189
18. The effect of decay of food on the rate of development of the
IBIRGE2 ccs > GR nolo eno ee, HO IC ee Se Re oeo 189
(95s theetteet of drying foodson) the larve....2 0.5... -.2-5- 2 190:
20.” Respiration.or larves ar liquified pulp... 6.2.56... ae. ne: 191
SSURY SS] ei eee) ory ase) ot Gag: a ro ean 191
Ben SR eceuiie Hctoles Of the MELON TMESY sh) 0 Os oc. wis tatae wheres 191
Zit DommelonuiiesiiamiteSt Caters sere keel a soeiee eo Mele brows w 6 si 192
24. Number of days required before the adults begin to oviposit 192
Dee OUMinaryaOn Siagesror tie lite biShOry.. 32. -n oe tle ae ee 192
265 Number of cenerations annually: 2.2 soc)... vets settee oe 193
Weights of the Eggs, Larve, Puparia and Adult Melon Flies....... 193.
INI artsta ped Syd CLC Stes MRED Perey eet oye ate ee airs trae ia cel BY aleea ens «Zeke Soucte ass ds 197
US ERLE" SOS" GEE 0s doy Sime a a ne Sn 198
27. Experiments in destroying infested vegetables............. 198
OR ee SCRE ETITIIOY © TEL hata eer epeaie. ees oe Pec n'a Mai sac als, cisetetan eek otal) heeraae 200
ee i ga MSETN eye ads cn ea oP cch ta ii e,c5, ON da aseae aie wou elelaisinjals bis es 201
SM eto Re Wet gency one ain ieee ote ips b nen eels Powe ve 201
31. The use of vegetable, animal and petroleum oils to trap the
Balelkaal TANS AC 56 Be ae FOR OE I aC aE BR era 201
DMM Cam UNS yaegere aie mesic cl ues ioees Oe ai he oe cud tan oO sels eisuabanet 202
Som ee GISONned dhalESpLray... 6 a--.csra ss: © cette tle eA i eae ee ge 202
PE Aran ete, chee Oe temas ene Pho cont alas ee eG a Maas Shs hye oe, Owes wie wena 207
177
178 Annals Entomological Society of America [Vol. VII,
I. INTRODUCTION.
The most destructive pest of the Cucurbitacez in the
Hawaiian Islands is commonly called the Melon Fly (Dacus
cucurbitz Coq.) or the Bitter Gourd Fruit Fly in other parts
of the world. Previous to the accidental introduction of this
insect into Hawaii, melons were sold at ten cents each, but
today the consumer often pays from fifty cents to one dollar
for a watermelon. It has been estimated that the loss in the
Hawaiian Islands amounts to almost a million dollars annually,
in tribute to this fly, or a little over five cents a day for a family
s
Text Fig. 1. Melon fly, Dacus cucurbitea Cog. (After Perkins).
of four, on an estimated population of 192,000. When one stops
to consider that the Hawaiian Islands are smaller than the
state of Rhode Island, that the principal agricultural products
are sugar, pineapples, coffee and rice, one realizes what a tre-
mendous amount of injury this fruit fly causes to the limited
vegetable crops grown in the islands. This trypetid has been
allowed to play havoc unmolested for a period of sixteen years
or more, so that today in many localities swarming with the_
pest, barely do the seeds of some cucurbits germinate, when
the seedlings are “‘stung’’ by the flies; the larve which hatch
from the eggs devour the tissue of the stems and cause decay,
then the maggots penetrate into the roots and completely
destroy the plants.
1914] Melon Fly, Dacus cucurbitae. 179
II. NATIVE HOME.
Some difference of opinion exists as to the native home of
the melon fly. Muir (5, p. 17) “found that India was its original
home’ and later on, Froggatt’s (5, p. 17) investigations showed
that the melon fly was widely distributed over India and also
Ceylon. Perkins (10, p. 36) believes “its true home is no doubt
in China or Japan.’”’ In a map showing the fruit fly regions
with steamship connections to California, Compere (2, p. 728)
records the danger of introducing the pest into that state from
the following sea ports: Hawaiian Islands, Timor, Manila,
Nagasaki, Hongkong, India and Singapore. Compere claims
that this pest is rarely found in the Philippine Islands and he
informed the writers that the melon fly was originally native
of these islands. It was imported from one of the above men-
tioned countries into the Hawaiian Islands about the year 1897.
Text Fig. 2. Wing of melon fly. (After Froggatt).
III. DESCRIPTION OF MELON FLY.
The melon fly is wasp-like in its general shape and behavior
and resembles a hornet (Polistes) somewhat in color but is less
than half as large. The head is yellow in color; the thorax
is reddish yellow marked with a number of light yellow areas
and the abdomen is yellow on the first two segments and red-
dish-yellow on the others. At the base of the second, abdom-
inal segment is a transverse black line, which unites with a
black, dorsal, median line on the next three segments. A
lateral, brown spot is usually present on the fourth and fifth
segments. The transparent wings are marked with brown
bands. A brown band extends along the front margin of each
wing and ends in a large spot at the apex; another brown band
extends along a fold of the wing near the body; between the
distal ends of these bands is a transverse marking (Text fig. 2).
The legs are light yellow in color.
180 Annals Entomological Society of America [Vol. VII,
Compere (2, p. 710) recognized the melon fly as a new species
in Hawaii and sent specimens to Coquillet (3, pp. 129-130)
whose original description follows:
“Dacus cucurbite.—Head light yellow; the occiput, except
the sides and upper margin, reddish-yellow, an ocellar black
dot, front marked with a brown spot in front of its centre, and
with three pairs of orbital brown dots, a black spot on each side
of the face near the middle, and a brown spot on the middle
of each cheek; antenne, palpi, and proboscis yellow, the latter
mottled with brown; thorax, reddish-yellow, the humeri, a
median vitta on the posterior half of the mesonotum, another
on each side, above the insertion of the wings, uniting with an
irregular band which extends upon the pleura to the upper part
of the sternopleura, also a large spot on each side of the metan-
otum, encroaching upon the hypopleura, light yellow; scutellum,
except its extreme base, light yellow, bearing two bristles; abdo-
men light yellow on first two segments, reddish-yellow on the
others, the extreme base, a fascia at the bases of the second and
third segments, usually a lateral spot on the fourth and fifth, also
a dorsal vitta on the last three segments, blackish or brownish;
first segment of the ovipositor of the female slightly longer than
the fifth segment of the abdomen. Wings hyaline, the apex of
the subcostal cell, from a short distance in front of the apex of
the auxiliary vein, the marginal and submarginal cells, the
median third of the first basal cell, and a large spot in upper
outer corner of the first posterior cell, brown, this colour en-
croaching on the third posterior cell and bordering the sixth
vein almost to its apex; posterior cross vein bordered with
brown, this colour extending to the hind margin of the wing;
upper end of the small cross vein is also bordered with brown.
Halteres light yellow. Legs light yellow, the broad apices of
the femora and the last four joints of the tarsi reddish-yellow;
hind tibiz reddish-yellow or dark brown. Length, 6 to 8 mm.
Type No. 4,207 in the United States National Museum.’’
IV. FIELD OBSERVATIONS IN A PUMPKIN PATCH.
Casual observations on the injuries caused by the melon fly
have been put on record but no intensive study of its destruc-
tive work has as yet been published. In our work careful
examinations were made of different parts of pumpkin plants
to ascertain the following points: (1), where the pest deposits
1914] Melon Fly, Dacus cucurbitae. 181
its eggs under natural conditions; (2), the external indication
of egg deposition and (3), the injury caused by the larve.
Observations were also noted on other host plants injured by
this fruit fly, but these were not so extensive as those made on
pumpkin plants.
1. Ovtposition in stems of pumpkin vines.—-The melon fly
often deposits its eggs in the stems of cucurbit seedlings. In
pumpkin vines the eggs are often laid within the tender stems
near the growing ends (plate X XVII, fig. 3), but the pest is not
able to puncture the older and tougher stems with its needle-like
Oovipositor. A gummy substance exudes from the wound and
hardens in the form of a small, resinous lump on the stems
(plate X XVII, figs. 1 and 2).
2. Oviposition in petioles of leaves Occasionally the melon
fly deposits its eggs within the petioles of the leaves. The
external indication of oviposition, as in the case of the stems,
is shown by the resinous material which accumulates at the
region where the petiole was punctured by the ovipositor.
3. Ovtposition in pumpkin flowers ——Eggs were found on
the outer and inner surfaces of the corolla and its lobes (plate
XXVII, figs.4and7). When the melon fly oviposits on the inner
surface of a corolla, it punctures the closed flower, glueing the
eggs either at one end in a mass or scattering them loosely on
the inner surface of the corolla and its lobes. At that region
where the flower has been punctured it becomes discolored
(plate X XIX, figs. 20 and 21).
Dacus often oviposits within a receptacle formed by its
ovipositor in the anthers or column of the stamens of the stami-
nate pumpkin flowers. More often, however, eggs are de-
posited in the tissue of the cup-shaped disc formed by the
union of the calyx and corolla, or the eggs are simply dropped
loosely into this cup-shaped disc. Occasionally, the eggs may
also be found within the peduncle of the staminate flowers.
The pest also deposits its eggs in the stigmas and styles
of the pistillate flowers. The trypetid does not enter the male
or female pumpkin flowers to lay its eggs, but punctures the
corolla from the outside with its ovipositor. Wherever the
ovary of the pistillate flower has been “‘stung’’ by the female
fly, a resinous material covers the wound (plate XXIX, fig.
18). Within the ovary, the ovipositor forms a small receptacle
in which the eggs are laid (plate X XVII, figs. 5 and 8). Eggs
182 Annals Entomological Society of America [Vol. VII,
are often deposited in the constriction between the perianth
and ovary, as is shown by the resinous substance found fre-
quently in this region (plate X XIX, fig. 19).
Green staminate and pistillate buds are often “‘stung’”’ by
the fly, the eggs being laid in the various parts of the unopened
flowers (plate X XVII, fig. 6), as has just been described for
the mature flowers.
4. Oviposition in pumpkins—The trypetid deposits its eggs
in small, green pumpkins, but the larger, uninjured pumpkins
are immune from the attacks of the pest, because the fly is
unable to pierce a hard rind with its ovipositor. If, however,
the rind of a large pumpkin has been injured, the fruit fly will
lay its eggs within the wound. ‘The insect will oviposit in an
exceedingly small hole extending through the resinous substance
of a healed wound, such as is often due to a previous infestation
by the pest (plate XXVIII, fig. 10). On a pumpkin in the
field, eighteen melon flies were counted with their ovipositor
inserted within a crack extending through the resinous exuda-
tion of such a wound and new arrivals were coming continu-
ously to oviposit in the same place. At the end of that day
the resinous material was removed, and hundreds and hundreds
of eggs were found closely packed in the pulp beneath the crack.
After the melon fly ‘‘stings’”’ the unripe pumpkin and squash,
the tissue surrounding the receptacle in which the eggs are laid
is killed, probably by a secretion which the fly pours over the
eggs. In the further development of these cucurbits a de-
pression results (plate XXVIII, figs. 9, 11 and 12) wherever
oviposition has occurred. Small pumpkins which have been
“stung” repeatedly, may assume all sorts of abnormal shapes
in their further growth (plate XXVIII, fig. 12).
5. Injury to stems.—The recently hatched larve devour
the tissue of the tender stems of young cucurbits and cause
decay, then they penetrate the roots and destroy the plants
entirely. Several acres of watermelons under observation
were replanted a number of times and, without exception,
every plant was destroyed in this way. The maggots often
destroy the terminal shoots of old pumpkin vines by penetrating
from one internode to another and feeding on the tissue of the
tender stems. A yellowish substance, probably the excrement
of the pest, stains the undevoured fibrous tissue of the stems.
No maggots were found in the old stems or roots.
be Fast
1914] Melon Fly, Dacus cucurbitae. 183
The old stems of the pumpkin vines are often infested with
the larva of a Cerambycid (Apomecyna pertigera) which is
able to penetrate through the hard nodes. More often, how-
ever, this beetle larva was found in the roots of the pumkin plant
- (Text fig. 3.)
Text Fig. 3. Root of a pumpkin vine split open lengthwise showing the larva of a
Cerambycid (Apomecyna pertigera) which feeds on the tissue of the
roots and occasionally of the stems.
6. Injury to petioles—Melon fly larve, which had recently
hatched, were found within the petioles of leaves, but nearly,
full-grown maggots were never observed within this part of the
plant. In order to ascertain whether the pest could complete
its larval period within a petiole, ten maggots of different sizes
were placed within a half dozen leaf-stalks. All of the larvee
obtained sufficient nourishment from the wall of the petioles
to complete their development. While most of the maggots
bored out of the petioles to pupate, others pupated within the
leaf-stalks close to the node of the stem.
7. Injury to flowers.—The larve that hatch from the eggs
deposited in the anthers, first feed upon and destroy these
structures (plate XXIX, fig. 28); then they may eat out the
184 Annals Entomological Society of America [Vol. VII,
column of the stamens (plate X XIX, fig. 29); next the pests
may work their way down into, and entirely destroy the cup-
shaped disc beneath the column and finally they may penetrate
into the long peduncle. The flower often drops from its stalk
(plate XXIX fig. 30) on account of a decay which follows an
infestation. The wall of the peducle is now eaten until only a
thin, papery envelope remains, which encloses a yellowish
substance similar to that observed in the infested stems. No
evidence was found that the larve pass through the node which
shuts off the hollow peduncle from the stem, but at this node
puparia were occasionally found.
In the pistillate flowers the larvee devour the stigmas and
styles, leaving a decayed mass to which the corolla clings. The
maggots then descend into the ovary and often the withered
corolla becomes detached (plate XXIX, fig. 25) and drops
to the ground, leaving a black, flower scar (plate XXIX, fig.
22). As the ovary is devoured, decay sets in, the pulp becomes
spongy (plate X XIX, figs. 23 and 24) and the channels are filled
with wriggling maggots. After the larve bore out, the ovary
turns black and either detaches from the pumpkin vine (plate
XXIX, fig. 27) and drops to the ground, or remains adhering to
the vine as a dried, shriveled mass (plate X XIX, fig. 26).
8. Injury to pumpkins —-A green pumpkin which is in-
fested with a small number of melon fly larvee may continue to
grow after the maggots have bored out, but when attcked by a
large number of the pest, the pumpkin turns black and decays.
After the larve have bored out of a green pumpkin, the wound
becomes covered by a gummy substance (plate XXVIII, fig. 10).
During the further development of this cucurbit, the resinous
material often cracks and a second oviposition may occur in the
crevices. If a ripe pumpkin is reinfested with a large number
of maggots a rapid decay changes the pulp into a semi-liquid
mass possessing a most sickening rancid odor. The rind may now
collapse (plate XXVIII, fig. 15), and the thick, liquid contents
then oozes out. After the maggots bore out, only the rind
containing the seeds remains. A glance at plate XXVIII, figures
13 and 14, shows the remains of two pumpkins which were turned
over to show the side that had been resting on the ground. In
such cases the seeds drop to the ground and often germinate.
When a ripe pumpkin is reinfested with a small number of
larvee the rind does not collapse (plate X XVIII, fig. 16) and the
seeds within the semi-liquid pulp may then decay.
oe
1914] Melon Fly, Dacus cucurbitae. 185
9. Injury to string beans.—An examination was also made
of the injury which the melon fly larve cause to the seeds and
pods of green-podded string beans. It was found that the
maggots feed upon the seeds and also the fleshy part of the pods
(plate XXX, figs. 31 and 32). After these portions have been
consumed the inner surfaces of the pods turn black (plate XXX,
Hig, So).
Dead Dacus larve were found within the seeds and pods
of string beans. Sometimes the dried bodies of the maggots
were found protruding from the pods (plate XXX, fig. 39);
these evidently died in the process of boring out of the host.
Pupation, which normally occurs in the ground, often takes
place within the dried pods (plate XXX, figs. 36 and 37).
V. Foop PLANTs.
In the Hawaiian Islands the melon fly has been bred from
the following food plants:
VEGETABLES. FRUITS.
Cucumber Mango. Bred by Terry (11, p. 32).
Egg Plant ?Orange. Bred by Ehrhorn (4, p. 337),
Kohlrabi Papaya.
Muskmelon.
Pumpkin.
Squash.
String bean.
Tomato. ,
Watermelon.
Wild cucurbit (Sycos sp.). Bred by Van Dine. (11, p. 32).
In India, Froggatt (5, p. 18) bred Dacus cucurbite from
melons, bitter gourds and egg plants.
10. Number of melon flies bred from the food plants—To
determine whether the pest could obtain sufficient food material
from the corolla of pumpkin flowers to complete the larval
development, the corolla was removed from six staminate
flowers in the field and each corolla was then placed in a breeding
jar together with recently, hatched, melon fly maggots. The
larve during their development obtained no other food than
that furnished by a single corolla. One male and one female
melon fly succeeded in completing their life history on this
food supply.
An experiment was now performed to determine the number
of melon flies which could be bred from an entire staminate
flower including its long peduncle. A dozen, infested, stam-
186 Annals Entomological Society of America [Vol. VII,
inate flowers were cut from pumpkin vines in the field and
placed in twelve breeding jars. The following figures indicate
the number of adults which were reared from each flower:
10, 14, 19, 23, 24, 25, 31, 32, 33, 37, 63 and 64% a total of 375
or an average of 31 flies for each flower.
A similar experiment was performed with a half dozen
pumpkins. The cucurbits were of different sizes and were
taken from the field and placed in separate breeding jars. The
following figures show the size of the pumpkins and the number
of adults reared from each.
From a pumpkin 24% inches long, 277 melon flies were bred.
From a pumpkin 3 inches long, 183 melon flies were bred.
From a pumpkin 314 inches long, 378 melon flies were bred.
From a pumpkin 3% inches long, 464 melon flies were bred.
From a pumpkin 4_ inches long, 637 melon flies were bred.
From a pumpkin 434 inches long, 283 melon flies were bred.
2222 total.
From twelve, infested, green-podded string beans gathered
in the field, the following number of melon flies were reared:
4,7, El, 14, 12) 14, 16,25, 16, 46; 18 and 26:"a total-ot tGsior en’
average of 13 flies for each pod.
VI. lare History,
Although the melon fly has been very destructive during
the past sixteen years in Hawaii, the duration of the different
stages of its life history have never been determined. Clark
(1, p. 6) makes the following statements on the life history of
this pest. The fly ‘‘stings’’ not only the fruit with its ovipositor,
but also the young and tender growth of the vines, depositing
a number of eggs, which soon hatch into small, white maggots.
that feed on the tissues of the fruit causing it to decay. After
the maggot has attained its growth, it descends into the soil
where it develops into a small chrysalis of a light, yellowish
brown color, and in about ten or twelve days comes out a
perfect insect, ready to repeat its mission of destruction. I
do not know how many generations it will produce in a year,
but in the warmer and drier districts I believe it will breed the
year through, except possibly a while during the winter months,
and then its development is only retarded by the cooler weather,
which prevents the chrysalis maturing so rapidly.”
1914] Melon Fly, Dacus cucurbitae. 187
Van Dine (11, pp. 32-34) gives the following contribution
on the life history of the melon fly: The life history covers a
period of about three weeks. The number of eggs which the
female deposits varies from 5 or 6 to as many as 15. After
hatching from the egg, the larve burrow into the tissue of the
melon and feed on the interior. When fully developed the
larve leave the infested melon or vine and enter the soil directly
beneath, where at a distance of an inch or so from the surface
they pupate.
Marsh (9, p. 156) writes, “In the insectary an effort was
made to work out the life history of this fly, but little progress
was made owing to the fact that the cages in which the speci-
mens were confined were too small.”’
11. Methods of inducing oviposition—Various methods
were adopted to induce melon flies to lay their eggs in different
food plants placed in a pumpkin patch swarming with the pest.
One method followed was to cut a non-infested, ripe pumpkin
in half and the trypetids which were probably attracted by
the odor of the pulp, would visit the cut surfaces and deposit
their eggs. Even the removal of a small piece of the rind from
a pumpkin or squash would be sufficient to attract and induce
the insects to oviposit. Another method used was to make a
semicircular cut through the peel and pulp near the surface
of cucumbers, egg plants and tomatoes and the loose flap was
then pinned back like a lid. After the females had deposited
their eggs in the pulp, the lid-like peel was pinned into its normal
position again, thus covering the eggs.
In one experiment about a square inch of the rind of a
pumpkin was removed and in a short time melon flies began to
visit the injured vegetable. The flies wandered about upon
the rind until they found the exposed pulp, when they began
to feed upon the exuding juices. At times as many as twenty-
five specimens were clustered together in this small area. So
closely crowded were the insects that their wings, which are
usually held at almost right angles to the long axis of the body,
overlapped. More and more individuals were attracted to the
cut area until the newcomers were actually forced to walk
over the bodies of the earlier arrivals, some of which were now
laying eggs without apparently being disturbed, for the ovi-
positor was not withdrawn.
188 Annals Entomological Society of America [Vol. VII,
12. Tactile bristles of the ovipositor.—The melon fly often
seeks a suitable place wherein to oviposit by walking about on
the cucurbit with the tactile bristles at the end of the ovipositor
(Text fig. 4) in contact with the rind. These tactile bristles
probably assist in locating a hole in the rind, or possibly discrim-
inate between hard and soft surfaces. One specimen was
observed with its ovipositor inserted within a pin hole which
was made in a pumpkin and another female fly in orienting
itself over this wound, would take a step or two backwards,
grope around with the tactile bristles and finally push the
ovipositor into the same hole. When a piece of the rind has
been removed the flies apparently seek a soft area in the pulp
with the tactile bristles of the ovipositor. The fruit flies will
readily locate and oviposit in a slit which has been cut in the
pulp.
Text Fig. 4. Distal end of the ovipositor of Dacus cucurbite showing
the tactile bristles.
13. Process of oviposition—The process of oviposition can
readily be observed in the field with a hand lens. When once
the fruit fly has found a suitable place, the abdomen is bent at
right angles to the long axis of the body and then the distal,
needle-like part of the ovipositor moves up and down in the
proximal, tube-like portion. As the ovipositor is forced into the
food plant, the female, in endeavoring to get a firmer foot-hold,
will let go with the tarsi and claws of the middle and hind legs .
and grasp anew hold. The tube-like, proximal portion is thrust
deeper and deeper until it disappears entirely and the eggs are
then deposited. If, however, this tube has not been pushed
entirely into the host plant, the eggs can actually be seen gliding
through the ovipositor at the rate of one in about every fifteen
or thirty seconds. Two specimens were timed during oviposi-
1914] Melon Fly, Dacus cucurbitae. 189
tion and it required seven and nine minutes respectively to
complete the egg laying period.
14. Number of eggs deposited in a receptacle-—The number
of eggs which the pest lays within a receptacle varies from one
to forty. The insect often punctures various parts of the host
plant with its ovipositor and yet does not deposit any eggs.
15. Number of ripe eggs in ovaries.—-In order to ascertain
the number of ripe eggs present in the ovaries, eighteen melon
flies were captured in the field and dissected. The average
number of mature eggs found in the two ovaries was forty-
eight; the largest number of fully-developed eggs dissected from
one specimen was seventy-four and the smallest number ob-
tained from an individual was twenty-two. The number of
eggs which one female is able to lay during its natural life was
not determined.
16. Duration of the egg, larval and pupal periods.—After
melon flies had been induced to oviposit in pumpkins, squash,
egg plants, tomatoes, cucumbers and string beans in the field,
the vegetables were transferred to breeding jars and a careful
record was taken of the different periods of the life history.
The following table shows the duration of the egg, larval, and
pupal periods of Dacus:
TABLE I.
DURATION OF THE EGG, LARVAL, AND PUPAL PERIODS OF DACUS CUCURBIT.
Fost Egg period Larval period Pupal period
(hours) (days) (days)
[2xbomeholsciknt, Uae oleae olor 30 4-7 10-13
WQUASaMer. ake 32 5-7 11-14
Sonoita weet een. 2 31 5-11 12-14
AROMALTOU Re tren eee: 36 5-8 12-13
CGucumbere ssn. 2 33 5-9 12-14
Stringsbean: ayn... 31 5-10 11-12
17. The effect of temperature on the egg period.—Pumpkins
containing eggs which had been recently laid were exposed to
the hot sunshine, while others were kept in the shade under
field conditions, but no marked difference in the duration of the
egg period was observed.
18. The effect of decay of food on the rate of development of
the larve.—One important factor upon which the rate of develop-
ment of the melon fly larve depends, is the rapidity of decay of
190 Annals Entomological Society of America [Vol. VII,
the host plant which follows an infestation of the pest. A
large number of maggots in a ripe pumpkin cause decay more
rapidly than a smaller number in a similar host of the same size.
The following table shows a comparison of the rate development
of the pest in rapidly and slowly, decaying, food plants:
TABLE It.
COMPARISON OF THE RATE OF DEVELOPMENT OF THE MELON FLY LARV4Z IN RAPIDLY
AND SLOWLY DECAYING HOSTS.
Rapidly Number | Larval | Slowly Number | Larval
decaying of period || decaying ot period
hosts larve (days) | hosts larvee (days)
211 7 1 4
201 434 52 5
Pumpkins. 0.015 - +. 55 584 || Pampkim..... c.0.. 26 6
6 634 8 a
1 8
4 5 4 8
1 6 | 45
Bee planitmews: ) eee i 7 Beorplanicassee see rr a
38 8 | 1 11
Cucumber. . 2.0.02. | a Gneumberss see. 6 9
It is possible that the rap'd decay of vegetables is caused
not only by the enzymes secreted by bacteria, but also by the
enzymes of the sal va of the melon fly larve. The enzymes
change the pulp into a thick liquid and it may be possible that
the maggots absorb some of this food directly through the body
wall. Larve swallowing and possibly absorbing liquified food
probably would require less time to complete their development
than maggots feeding upon solid food.
19. The effect of drying food on the larve.—tIn string beans
which gradually dry up during an infestation, there is a marked
individual variation in the growth of the melon fly larve, even
when hatched from the same batch of eggs. In a dried, bean
pod the larval period is longer than in a decaying one. In the
laboratory the maggots often died in the seeds and the pods
of dried string beans, a fact that was also observed in the field.
In all probability, these larve died from lack of moisture due
to the drying up of the string beans.
1914] Melon Fly, Dacus cucurbitae. 191
20. Respiration of larve in liquafied pulp.—Melon fly
larve in a decayed tomato were observed to obtain a fresh
supply of air by pushing the posterior spiracles above the sur-
face of the liquified pulp. The stem of this tomato had been
cut out and many maggots were found suspended from the
surface film with their bodies immersed in the liquid. When
the finger was snapped against the tomato, the larvae wriggled
down into the liquified pulp, but sooner or later, they would
come up to the surface film to breath.
21. Jumping habit of larve——When nearly, full-grown,
melon fly larvae are removed from the host, they exhibit a
peculiar habit of jumping, but this behavior is not manifested
by the smaller maggots. The larve curl the body in a circle
(plate XXVIII, fig. 17), the jaws attach to the posterior end
of the body, and then by a sudden muscular relaxation, they
spring about six to eight inches into the air.
22. Feeding habits of the melon flies.—As a rule, Dacus feeds
during the early morning from sunrise to about ten o’clock.
During the hottest part of the day thousands of these insects
may be found at rest under large leaves of plants in or near an
infested field of cucurbits. The flies were frequently found
several hundred feet away from their breeding grounds, feeding
upon the flowers of the glue bushes, sunflowers and Chinese
bananas. Nota single fruit fly was ever seen entering and feed-
ing within the carolla of pumpkin flowers or morning glories,
but after a rain, specimens were occasionally observed lapping
up the small droplets of water on the lobes of the corolla of
morning glories. Melon flies feed upon the juices of injured
or infested cucurbits in the field. Many individuals were ob-
served feeding on the juices exuding from sweet corn. One adult
was noticed feeding on a dead and partly decomposed cater-
pillar.
When a piece of the rind of a pumpkin was removed, large
numbers of the pest were attracted to the exposed pulp and fed
upon the exuding juices. When a common house fly also
visted the pulp to feed and approach a melon fly already en-
joying a meal, the latter would dart forward and chase the former
away, but when laying eggs the fruit fly would not withdraw its
ovipositor even when the house fly crawled over its body.
192 Annals Entomological Society of America [Vol. VII,
23. Do melon flies manifest fear?-—A melon fly is keenly
aware of movements within its field of vision. When a large
Odonata flies above a melon fly at rest on a cucurbit, the try-
petid may tilt its head to follow the flying insect with its eyes.
When one approaches a pumpkin slowly and carefully upon
which a specimen is feeding, the fly may tilt its head and sway
its body from side to side by bending the legs at the tibio-
femoral joints. At such times a slight movement on the part
of the observer will cause the fruit fly to take wing. The
swaying movements of the alert insect may possibly be inter-
preted as an external indication of fear. Howlett (7, pp. 415-
416) believes that this swaying movement ‘‘seems to be asso-
ciated with ‘courtship’ in all species of Dacus that occur at
Pusa.’’ When the head of Dacus cucurbite is lowered, however,
and the insect walks about with the wings held almost at right
angles to the long axis of the body one can then approach the
pest without danger of causing flight.
24. Number of days required before the adults begin to oviposit.
—An attempt was made to determine the number of days re-
quired before the egg-laying period begins, after the melon
flies issued from the puparia. A large number of adults upon
emerging, were kept in breeding jars and fed on diluted molas-
ses, fruit and vegetable juices and on water. After having
been kept in captivity for eight days, three females were dis-
sected but no fully-developed eggs were found in the ovaries.
A daily dissection of three fruit flies was continued from now
on, and at the end of fourteen days twenty-three ripe eggs were
counted in the two ovaries of one fly, but others did not show
mature eggs in the ovaries at the end of sixteen days. At the
end of seventeen days, thirty-one eggs were dissected from the
ovaries of another specimen. In all probability, the effect of
confining the insects in breeding jars plays an important part
in the rate of development of the eggs.
25. Summary of stages of the life history—The duration of
the different stages of the life history of Dacus cucurbita may
be summarized as follows:
DAYS
Bieormertodis tasiyewaeor sce tere ae 1y4— 1%
Ikanvallpertodiag.cen cecunt sy err 3%4—l11
Pupaliperiod. cre. et errr ee 10 —14
Beodayine begins... seas. ae 14 —17 after the adults emerge.
29 —4314
1914} Melon Fly, Dacus cucurbitae. 193
26. Number of generations annually.—In the Hawaiian
Islands one brood of melon flies is followed by another through-
out the year. Since the duration of the complete life cycle
may vary from twenty-nine to forty three days, one would
expect from eight to twelve generations a year. Assuming
that a single female produces only ten descendants and that the
sexes are produced in equal numbers at the end of the year she
would be the ancestor of from 100,000,000 to 1,000,000,000,000
offspring.
VII. WEIGHTS OF THE EGGs, LARVAE, PUPARIA AND ADULT
MELON FLIEs.
Accurate weighings of the following stages in the life history
of Dacus cucurbite were taken: eggs (plate XXXI, fig. 46)
a few hours after these were deposited and again shortly before
hatching; larve after hatching and every day thereafter (plate
XXXI, figs., 47 and 52); recently, formed puparia (plate XX XI,
fig. 53) and male and female melon flies.
Pumpkins, in which melon flies had been induced to deposit
their eggs, were taken from the field to the laboratory and three
hours after oviposition, the eggs were removed and counted in
two lots, each containing one hundred eggs. Each batch of eggs
was placed upon a small piece of filter paper and weighed in a
weighing bottle. The eggs were then transferred from the
filter paper into a pumpkin and twenty four hours later, the
same eggs were removed and weighed again. The weights of
the eggs three hours and eyenty seven hours after deposition
were as follows:
TABLE III.
WEIGHT IN MILLIGRAMS OF EGGS OF DACUS CUCURBIT#Z AFTER DEPOSITION, BEFORE
HATCHING AND LOSS IN WEIGHT.
3 hrs. after 27 hrs. after Loss in wt.
deposition deposition in 24 hrs.
Wt. of first 100 eggs.... 1.2 6. 1.2
Wt. of second 100 eggs.. 7.5 6.1 1.4
013
Average wt. of l egg... .0735 .0605
The result shows that each egg on an average, lost .013
milligrams in weight, during the twenty-four hours between the
two weighings.
194 Annals Entomological Society of America [Vol. VII,
After the second weighing, the first lot of one hundred eggs
was put back into the pumpkin and the second batch was placed
between moist filter paper. These lots hatched respectively
in thirty and thirty-two hours after deposition. The hatching
of the first batch of eggs was observed under a binocular micro-
scope in order to remove the larve before they had an oppor-
tunity to feed. The weight of the maggots and the difference
in weight between the eggs and larve are recorded in the
following table:
TABLE IV.
WEIGHT IN MILLIGRAMS OF RECENTLY HATCHED LARVA OF DACUS CUCURBIT#, AND
DIFFERENCE IN WEIGHT BETWEEN EGGS AND LARV#.
Difference in wt. between
eggs and larve
Wt. of 100 recently hatched larve
3 hrs. after 27 hrs. after
deposition deposition
Hatched from first 100 eggs............ 4.2 3. 1.8
Hatched from second 100 eggs......... 5.6 1.9 15
IAWenage WibwOm IlArVicvesne sae eto .049 0245 .0115
The two hundred, recently hatched maggots weighed one-
third less than the eggs after deposition and about one-fifth less
than the eggs before hatching. This loss can be attributed,
in part, to the shedding of the chorion and vitelline membrane.
The one hundred larve which hatched from the first batch
of eggs were fed upon a hard, dry pumpkin while the other one
hundred maggots were fed on a soft, juicy pumpkin. The
following table shows the weights of the larve after they had
fed a day.
TABLE V.
WEIGHTS IN MILLIGRAMS OF LARVA OF DACUS CUCURBITZ FED UPON A HARD, DRY
PUMPKIN AND A SOFT, JUICY PUMPKIN.
Fed on dry, Fed on soft, Difference
hard pumpkin | juicy pumpkin in wt.
Wt. of 100 larve 1 day old....... 26.6 63.6 37.
Witotd tamavliday:olde ass . 266 .636 Of
It is evident from this table that the larvze which fed on
the soft, juicy pumpkin for one day weighed over twice as much
as the maggots which fed on the dry, hard pumpkin. . Further-
more, the former had increased their initial weight over eleven
times and the latter over six times at the end of one day.
1914] Melon Fly, Dacus cucurbitae. 195
Fifty of the one hundred larve which fed on the soft, juicy
pumpkin were weighed at intervals of one day throughout
their larval life. In order to remove all of the pulp, which
adhered to the bodies of the maggots, they were carefully
washed and dried with filter paper before each weighing. The
following table shows the daily increase in weight of the larve,
daily increase over the initial weight, and increase or decrease
of weight over the previous day:
TABLE VI.
DAILY INCREASE IN WEIGHT, DAILY INCREASE OVER THE INITIAL WEIGHT AND INCREASE
OR DECREASE IN WEIGHT OVER THE PREVIOUS DAY IN MILLIGRAMS OF THE
LARV# OF DACUS CUCURBIT.
Age of |Wt.50/ Wt.1 Increase in wt. Increase or loss of wt.
larvee larve | larva of 1 larva over of 1 larva over pre-
initial wt. vious day
At hatching.| 2.8] .056
I Glanve eee 31.8] .636 .58 .58 increase
2 days....| 101.2) 2.024 1.968 1.388 «
3 days....| 801.6 |16.032 15.976 14.008 «
4 days... ./1021.7 |20.434 20.378 4.402 «
46 days... .|/1004.3 |20.086 20.03 .348 loss
The daily increase in weight over the initial weight may be
stated as follows:
After feeding 1 day a larva weighs 11.3 times its initial
weight.
After feeding 2 days a larva weighs 36.1 times its initial
weight.
After feeding 3 days a larva weighs 286.2 times its initial
weight.
After feeding 4 days a larva weighs 364.8 times its initial
weight.
After feeding 414 days a larva weighs 358.6 times its initial
weight.
The minimum increase over the initial weight occurred dur-
ing the first day of the larval life and the maximum increase
took place during the third day.
The daily increase or decrease in weight over the previous
day may be put as follows:
The first day a larva acquires 10.3 times the weight at hatch-
ing.
196 Annals Entomological Society of America [Vol. VII,
The second day a larva acquires 2.1 times the weight of the
first day.
The third day a larva acquires 6.9 times the weight of the
second day.
The fourth day a larva acquires .27 times the weight of the
third day.
In 41% days a larva lost .017 times the weight of the fourth
day.
The minimum increase in the daily weight of the larve over
the previous day occurred during the fourth day and the maxi-
mum increase took place during the first day. During the last
twelve hours of the larval period, the maggots decreased in
weight. In all probability, this loss may be attributed in part,
to the evacuation of the contents of the alimentary canal pre-
vious to pupation.
After the fifty larve bored out of the pumpkin in 414 days
they were allowed to pupate in moist sand. Twelve hours
later the sand adhering to the newly-formed puparia was washed
off and the moisture adhering to them was absorbed with filter
paper. After the puparia were thoroughly dried they were
weighed in a weighing bottle. Other melon fly maggots which
bored out of pumpkins in 324 and 4 days were weighed, and
twelve hours later the newly-formed puparia were weighed.
The following table shows the average weights of the mature
larve, the puparia twelve hours after the larve bored out of the
pumpkins, and the loss in weight after pupation:
TABLE VII.
AVERAGE WEIGHTS IN MILLIGRAMS OF MATURE MELON FLY LARV4, WEIGHTS OF THE
PUPARIA TWELVE HOURS AFTER THE LARV# BORED OUT OF THE PUMPKINS
AND THE LOSS IN WEIGHT AFTER PUPATION.
Age of Average wt. Average wt. Loss in wt. Loss in wt.
larvee 1 larva 1 puparium after pupation %
416 days..... 20.086 18.95 1.136 5.6
A dayGirss: 22.509 20.336 2.173 9.6
334 days..... 20.87 19.0534 1.8166 8.7
The results show that twelve hours after the melon fly
larve bored out of the pumpkin and transformed into puparia,
there was a loss of 5.6 to 9.6% in weight.
1914] Melon Fly, Dacus cucurbitae. 197
Is there a difference in weight of the male and female melon
flies upon emerging from the puparia? The following table
shows the weights of two lots of flies shortly after the wings
were expanded:
TABLE VIII.
WEIGHTS IN MILLIGRAMS OF MALE AND FEMALE MELON FLIES SHORTLY AFTER EMERG-
ING FROM THE PUPARIA.
Number of : Tat | Difference in weight
fies Weight Weightlfly | Shao andee ay
507 47.52 9504
509 48.9 .978 .0276
407 51. 1.275
309 45.69 1.523 . 248
It is evident from this table that the female melon flies are
heavier, on an average, than the males shortly after they issue
from the puparia.
Sanit NATURAL ENEMIES.
Predaceous insects sometimes prey upon the melon fly. At
all times of the day the yellow dragon fly (Panatala flavescens
Fab.) was observed flying over a pumpkin patch teaming with
pest, and one would be inclined to believe that the melon flies
are sometimes preyed upon by this predaceous insect. A
number of dragon flies were captured in this field and the con-
tents of their alimentary canal was examined, but no remains
of the melon fly were found. A predaceous bug (Zelus peregri-
nus Kirkaldy) was observed sucking out the juices of a melon
fly onasunflower. Staphylinids or rove beetles were frequently
seen within infested pumpkins but whether or not these feed
upon the melon fly maggots was not determined.
The European, horn fly parasite (Spalangia hirta Haliday)
was bred from the puparia of the melon fly. These puparia
were gathered from beneath infested pumpkins; some of these
puparia were partly exposed from the ground; while others
were taken from one to four inches below surface of the soil,
the usual depth, however, being one to two inches. From five
hundred puparia which were collected on March 25, three
parasites emerged on May 4, 1912. It is evident that this
parasite is of little importance in the control of the melon fly. —
198 Annals Entomological Society of America [Vol. VII,
IX. METHODS OF CONTROL.
27. Experiments in destroying infested vegetables.—Van Dine
(11, p. 35) formerly stationed at the Hawaiian Agricultural
Experiment Station, recommends that all melons and vines
infested with melon fly larve should be collected at intervals of
five or six days and covered with earth to a depth of several
inches.
A number of experiments were performed to determine the
distance that melon flies, after issuing from the puparia, were
able to burrow through sand and soil. In the first experiment
several, hundred puparia were placed on two inches of dry,
sterlized sand at the bottom of a cylindrical, museum jar
(24x111% inches) and this jar was then filled with more of the
same kind of sand. A similar vessel, half filled with dry sand,
was then inverted over the top of the above mentioned jar.
This was done by placing a heavy, glass plate over the mouth
of the jar to be turned upside down, inverting the same above
the other vessel and then pulling the glass plate out from be-
tween the two jars. A similar experiment was conducted with
wet sand which had been previously sterlized. The puparia in
both experiments were arranged in a circle close to the wall of
the jars so that when the fruit flies emerged and burrowed
through the sand their paths might be seen. When the trypetids
emerged, many would bore up to the region where the jars came
in contact with one another and then escape through the small
spaces between the jars. These small spaces were due to par-
ticles of sand which rested on the rims of the jars. One could
scarcely believe that these large flies were able to flatten their
bodies to such an extent as to squeeze through such small
spaces as existed between the jars.
It was evident that some of the melon flies were not able to
burrow as far as others, for many died at the upper end of the
channels before obtaining their liberty (plate XX XI, fig. 45).
Flies would frequently bore into an excavation made by other
specimens and if the union of the channels would form a more
or less circular path, some of the individuals would continue
to burrow slowly round and round and finally die in this endless
passage. Usually, however, most of the trypetids showed
a definite orientation and bored directly upward. This nega-
1914] Melon Fly, Dacus cucurbitae. 199
tive reaction to gravity is common with many insects after
emerging from the egg or pupa.
As there was a possibility that the melon flies might have
been hindered by being against the glass, holes two and three
feet deep were drilled in hard soil with a fence-post borer. At
the bottom of these holes 500 puparia were placed. The pupa-
ria were then covered with sterlized dry or wet sand or soil.
After these pits were filled each hole was covered at the surface
with a large mouthed jar which rested tightly against the solid
earth. The following table shows the number of melon flies
which succeeded in boring through two and three feet of sand
or soil:
TABLE IX.
‘NUMBER OF DACUS CUCURBITZ WHICH BURROWED THROUGH TWO AND THREE FEET
OF SAND OR SOIL.
Number of feet puparia were buried........... 2 3
Number of puparia buried in each hole........ 500 500
Number of flies that bored through dry sand.. 34 2
Number of flies that bored through wet sand.. 8 0
Number of flies that bored through soil....... 5 0
It is evident from this table that a larger number of melon
flies were able to bore through dry sand than wet sand, and that
very few specimens succeeded in making their way through the
more lumpy soil.
Burying infested cucurbits and the vines in three feet of
soil would require a considerable amount of labor. Lime,
which is often thrown into the garbage-can to destroy the larve
of the house fly and blue bottle fly, would probably destroy the
melon fly maggots if it was buried in sufficient quantity with
the infested vegetables, but this method would increase the
cost.
In an experiment melon fly maggots were submerged in
fresh water for a period varying from two to four days, in order
to determine whether such larve would pupate and give rise
to flies. Larvze were selected which had bored out of a pumpkin
and were ready to pupate. These maggots were submerged
in seven inches of distilled water which was renewed daily.
After remaining in the water for two, three or four days, the
larvee were transferred to filter paper and after pupation, the
puparia were placed in moist sand in a breeding jar. The
following table indicates the results obtained.
200 Annals Entomological Society of America [Vol. VII,
TABLE X.
NUMBER OF MELON FLY LARV4 WHICH PUPATED AND ISSUED AS ADULT FLIES, THE
LARV4Z BEING SUBMERGED IN SEVEN INCHES OF WATER FOR A PERIOD OF
TWO TO FOUR DAYS.
Number of Days submerged | Number Number of Adults
larvee in water pupated dead pupee reared
100 2 75 28 47
100 3 16 11 5
100 | 0 0 0
From this experiment one could conclude that infested
vegetables may be submerged in a barrel or tank of water for
a period of four days and then plowed under without danger
of having melon flies emerge from this material. By following
this method a valuable fertilizer is added to the soil. Un-
doubtedly, certain chemicals could be added to the water which
would destroy the melon fly larve in the infested vegetables in
less time, but this again, would increase the cost.
Burning or boiling maggoty vegetables is somewhat expen-
sive on account of the fuel consumed. The old vines in seri-
ously, infested, cucurbit fields were often pulled out of the
ground, raked together in piles, the infested vegetables were
scattered within these piles and then all was burned. Some
kinds of infested vegetables could be boiled and then fed to
hogs.
28. Screening or netting—Attempts to grow the various
kinds of Cucurbitacee in Hawaii are carried on mostly by
Japanese and Chinese. . Many of these cultivators screen the
newly set cucurbits with pieces of gunny sac, paper bags,
newspapers or straw. Some of the growers screen their melon
and cucumber beds with cheese cloth or mosquito netting but
as Froggatt (5, p. 18) states, ‘though it kept the flies out it also
kept all the bees and small insects that, under ordinary con-
ditions, fertilize the flowers, so that very few melons ever set
and matured.’”’ Hand pollination could be resorted to, but
this method would require a considerable amount of labor.
One individual who attempted to grow pumpkins hired a
Japanese who removed from the vines, the staminate flowers
which were seriously infested with the larve. Screening the
vines is not at all practical except possibly for a few vines in a
1914] Melon Fly, Dacus cucurbitae. 201
garden. Covering the newly set cucurbits requires constant
attention and cannot be recommended, if the question of labor
is taken into consideration.
£9 Trap crops.—Marsh (9, pp. 155-156) made a test of a
trap crop by planting cantaloupes among cucumbers. “It
was thought that the cantaloupes would prove more attractive
to the flies than cucumbers, but such was not the case, as the
cucumbers were more badly damaged than the cantaloupes, and
in the end both crops were practically destroyed by the larve.”’
30. Traps——In Honolulu a Japanese glass fly trap is
used by many of the Oriental merchants to capture house
flies, blue bottle flies etc.; the insects enter the trap to feed
and are drowned in soapy water within the apparatus. As
this trap is similar to our American style of glass fly trap, a
description of the Oriental type is not necessary. A Japanese
fly trap with molasses as a bait was wired in a large orange
tree and in twelve days nineteen male and fifty-eight females
Mediterranean fruit flies and three male and one female melon
fly were found drowned in the soapy water. Two of the Ameri-
can style of glass fly traps with molasses as a bait were placed
upon the ground in a pumpkin patch but no trypetids were
caught in these. A dozen of the common, mosquito, screen,
fly traps with molasses diluted either with water or stale beer
as a bait were fastened to sticks above the pumpkin vines but
not a single fly was found in the traps during the five days that
they were kept in the field.
31. The use of vegetable, animal and petroleum oils to trap
the melon flies —Recent investigations have shown that certain
vegetable and petroleum oils attract enormous numbers of male
fruit flies of different species. Howlett (7, pp. 412 and 414) found
that citronella oil has an attraction for the males of the peach
fruit fly (Dacus zonatus Saund.) and the three-striped fruit
fly (Dacus diversus Coq.) but the attraction in the last case,
however, seems perhaps a trifle less powerful than with Dacus
zonatus. Froggatt (5, pp. 13 and 17) found that the mango
fruit fly (Dacus ferrugineus Fab.) is also attracted to citronella
oil but that the melon fly (Dacus cucurbitz Coq.) never came to
this oil. According to Illingworth (8, p. 160) the apple maggot
or railroad worm (Rhagoletis pomonella Walsh) avoids citro-
nella oil.
202 Annals Entomological Society of America _[Vol. VII,
We found that many of the oils derived from crude petroleum
attracted the males of the Mediterranean fruit fly (Ceratitis
capitata Wied.), but rarely was a specimen taken in animal
or vegetable oils. Our experiment with kerosene shows that
of every thousand Mediterranean fruit flies captured only
three, on an average, were females, the remainder being males.
It is noteworthy to mention that the Queensland fruit fly
(Dacus tryoni Frogg.) and the Mexican or Morelos orange
worm (Anastrepha ludens Loew.) are not attracted to kerosene.
The vegetable, animal and petroleum oils listed in the
following table were poured in pans and placed upon the ground
in a pumpkin patch which was swarming with melon flies. The
number of pans used, the number of days each oil was tested
and the results obtained are stated in the following table:
TABLE XI.
NUMBER OF MALE AND FEMALE MELON FLIES CAPTURED IN ANIMAL, VEGETABLE AND
PETROLEUM OILS.
Pans ‘Of 2
(Citranetla, A543 ah: aoa a eae es 2 5 days t 2
Vegetable oils {Turpentine............. PES EE 8 ee SU 1 16 hours 0 0
COCCAanUE SS Fike ee Ree 2 5 days 0 2
Animal oils Ala =) (eee eetteges Bech hare ceo ee 5 oy Sv 2 5 days 2 4
(EES 5 i afraaco oo ch te See ee eee 2 5 days 1 1
Kerosene about 120° Bé............. 3 5 days uy 3
Petroleum oils ;Gasoline about 86° Bé............. 1 16 hours 0 0
(Benzine\ aboutiiG3e Bet. 2.525 65. 1 16 hours 0 0
6 2
In all probability, the specimens were not attracted to these
oils but came within the sphere of influence by accident, became
stupefied and dropped into the oil.
32. Night traps.—As the melon flies show a strong positive
reaction to light, an attempt was made to capture the pest with
anight trap. Herrick’s moth trap was placed in a pumpkin patch
but not a specimen was caught. A seventy candle-power
hunting lamp was placed above the moth trap and the light
rays were directed towards thousands of melon flies resting
under sunflower leaves in the pumpkin patch but not a single
Specimen was attracted to the light.
33. Poisoned bait spray.—Striking demonstrations have
been made of the effectiveness of the poisoned bait spray in
the control of the olive fly (Dacus ole Rossi) in Italy and France
1914] Melon Fly, Dacus cucurbitae. 203
and the Mediterranean fruit fly (Ceratitis capitata Wied.) in
South Africa.
Recently similar control measures have been used in the
United States and Canada against the apple maggot (Rhago-
letis. pomonella Walsh), the cherry fruit flies (Rhagoletis
cingulata Loew. and Rhagoletis fausta O. S.) and the currant
or gooseberry fruit fly (Epochra canadensis Loew.). We have
tested the effectiveness of the poisoned bait spray to control
the Mediterranean fruit fly under Hawaiian conditions. The
method adopted was to wire ten kerosene traps in different
parts of an orchard containing about four hundred fruit trees.
The total number of fruit flies captured in five weeks was 10,239;
of this number 10,203 were males and only 36 were females.
During the following five weeks the poisoned bait spray was
applied to the trees about once a week. The total number of
fruit flies captured in the kerosene traps during these five weeks
was 182, of which number 93 were caught during the first
week.
As already stated Dacus cucurbite requires at least two weeks
under laboratory conditions before the egg-laying period com-
mences. Under natural conditions, the flies seek food during
this period and subsist on a variety of sweet substances already
discussed under the feeding habits of the melon fly. In capti-
vity, the adults show a fondness for diluted molasses and they
fed on this liquid until their aodomens became greatly distended.
One can readily understand that if this insect is attracted to
diluted molasses under natural conditions, that the greediness
of the fruit flies for this sweet when poisoned, would be the |
weak point in the life history to attack the pest. If this poisoned
bait is applied in the form of a spray to the food plant, when
the trypetids issue from the puparia, no doubt large numbers
would be killed before the egg-laying period begins.
The poisoned bait was prepared according to the following
formula:
Brown sugar 21 |b.
Arsenate of lead 5. oz.
Water 4 gal.
The solution was prepared by dissolving the brown sugar
and lead arsenate through cheese cloth in cold water so as to
strain out all foreign material including ants, which in the
Hawaiian Islands frequently gnaw through the paper sacs
204 Annals Entomological Society of America [Vol. VII,
containing the sugar. The mixture was thoroughly agitated
by pumping the liquid back upon itself with a common, garden,
brass spray-pump. To kill the enormous numbers of melon
flies quickly in a badly infested cucurbit field, one ounce of
a soluable poison such as potassium arsenate or sodium arsenite
dissolved in a small quantity of water, was added to the solution
instead of arsenate of lead. The pump was provided with a
rose, sprinkler nozzle which throws a fine, mist-like spray.
Shortly after sunrise the insecticide was applied to all of
the foliage within the pumpkin patch and also to the vegetation
bordering the same, such as glue bushes, algeroba trees, bananas,
sunflowers, castor oil beans, weeds and grass. As already
stated in the discussion of the feeding habits of the adults, the
pest was found feeding on flowers about a hundred yards away
from the breeding grounds. To spray all of the feeding grounds
which often consisted of dense brushes of glue bushes, would be
practically impossible. The results obtained after spraying
were rather striking. Before spraying, thousands and thousands
of melon flies could be found resting on the lower surface of
the leaves of the sunflower and castor oil plants, but after
spraying, only here and there could a specimen be found.
In all probability, these living flies had recently emerged from
puparia, or came in from the neighboring feeding grounds or
from surrounding cucurbit fields. The soluble poisons, how-
ever, burned the foliage and can not be advocated.
A few days after the application of the first spray, all of
the pumpkin vines and bean plants were pulled out of the ground
and raked together in piles. The infested pumpkins were
scattered within these piles and then all was burned.
To determine whether the melon flies coming from their
feeding grounds or from the surrounding fields of cucurbits
could be controlled, watermelon seeds were planted in a field
adjacent to the former pumpkin patch. The seeds sprouted
before we were able to make a vigorous campaign in surrounding
cucurbit fields. The watermelon plants were sprayed with
the bait, using arsenate of lead, but frequent rains washed
off the thin film of sugar and left the plants subject to
the attacks of the pest coming from outside sources. As
soon as the weather became settled, a fresh application
of the bait was made to the watermelon plants and sur-
rounding vegetation, but the tender stems of many of the
1914] Melon Fly, Dacus cucurbitae. 205
watermelon plants were already infested. Whether the pest,
which has been allowed to increase unmolested during
the past sixteen years, can be controlled under Hawaiian
conditions when one individual sprays and his neighbors
do not, is problematical. In all probability, better results
could be obtained with the poisoned bait spray in a well
isolated cucurbit field away from the valleys where rains are
less frequent during the summer months.
Marsh (9, p. 155) tested a poisoned bait spray to control
the melon fly in the Hawaiian Islands. He writes, ‘‘The baits
were prepared by sweetening water with molasses and adding to
the solution arsenate of lead or Paris green. These baits were
then applied, at frequent intervals, to the foliage of infested
cucumbers with a gardener’s syringe. With the aid of the syringe
the poisoned liquids were shot into the air above the beds of
cucumbers and allowed to fall on the foliage in fine drops. In
the experiment with Paris green the application was made
daily from September 9 until October 14. The formula used
in this experiment was as follows:
Molasses ee otianct:
Paris green 14 ounce.
Water 114 gallon.
Neither the experiment with arsenate of lead or with Paris
green proved effective. The flies were frequently observed
feeding on the poisoned liquids; but evidently they did not
relish them, and so failed to consume a fatal dose.”’
Fuller (6, p. 26) stationed in Natal, South Africa, tested
the poison bait spray to control the Mediterranean fruit fly and
melon fly. Trials which have been made in several citrus
orchards to control the Mediterranean fruit fly with the poisoned
bait spray “have been attended with remarkable effects, and
where treatment has been applied for the melon fly which
attacks, squashes, marrows, pumpkins and the like, it has
proved equally successful.’’
206 Annals Entomological Society of America _[Vol. VII,
EXPLANATION OF PLATES.
PLATE XXVII.
Fig. 1. Resinous lump which heals the wound produced by the ovipositor of the
melon fly, in the tender stems near the growing ends of the pump-
kin vines. =
Fig. 2. Resinous lump somewhat enlarged.
Fig. 3. Split stem of pumpkin vine near growing end, showing the eggs depos-
ited by the melon fly.
4. Eggs deposited on the outer surface of the corolla and its lobes.
5. Longitudinal section of the ovary of a pumpkin, showing the eggs depos-
ited within a receptacle. |
6. Green pistillate bud with. the corolla cut open, showing the eggs depos-
ited on the style and inner surface of the corolla.
Fig. 7. Eggs deposited on the inner surface of the corolla.
Fig. 8. Cross section of the ovary of a pumpkin, showing the eggs deposited
within a receptacle.
PLATE XXVIII.
Fig. 9. Squash showing depressions caused by the oviposition of the melon fly
during the early development of this cucurbit. The tissue surround
ing the receptacle in which the eggs are deposited is probably
killed by a secretion, and in the further development of the cucurbit
a depression results.
Fig. 10. Healed wounds due to a previous infestation of the melon fly. This
trypetid is unable to puncture the hard rind of the larger pumpkins
with its ovipositor, but often the resinous material covering the
wound cracks and oviposition may then occur in the crevices.
Fig. 11. Depressions and healed wound in a ripe pumpkin.
Fig. 12. Pumpkin, abnormal in shape, as a result of being ‘‘stung’’ by the melon
fly during its early development.
Figs. 13 and 14. Remains of two pumpkins which were turned over to show the
side resting on the ground. After the maggots bore out of a seri-
ously reinfested pumpkin, only the rind and seeds remain.
Fig. 15. The rind of a seriously reinfested pumpkin collapsing due to decay
caused by the larve.
Fig. 16. When the pumpkin is not seriously reinfested with the melon fly larve,
the rind does not collapse. Such partly decayed cucurbits often
serve as hosts for other insects which complete the work of de-
struction.
Fig. 17. Pumpkin cut in half showing the decayed pulp and seeds, and the larve
which have jumped out of the decayed mass.
PLATE X XIX.
Fig. 18. Resinous material which exudes from the wound produced by the ovi-
positor of the melon fly in the ovary of a pumpkin.
Fig. 19. Resinous substance in the constriction between the flower and the ovary
where the eggs are often deposited.
Fig. 20. Discoloration on the corolla indicating the region where the fly has
punctured the closed flower to deposit its eggs within the stigma
or style.
Fig. 21. Discoloration on the corolla where the pest has punctured the closed
flower to deposit its eggs on the inner surface of the corolla lobes.
Fig. 22. The corolla of one of the flowers has detached and dropped to the ground,
leaving a black flower scar. The larve in this case devoured the
stigmas and styles and descended into the ovary.
Figs. 23 and 24. Longitudinal sections of two ovaries showing a spongy decayed
pulp caused by the maggots.
Fig. 25. Corolla detaching from the ovary after the larve have devoured the
stigmas and styles and descended into the ovary.
Fig.
Fig.
Fig.
1914] Melon Fly, Dacus cucurbitae. 207
Fig. 26. Dried and shriveled pistillate bud still adhering to the pumpkin vine
after the larve have bored out of the ovary.
Fig. 27. Pistillate bud detaching from pumpkin vine after the larvze have bored
out of the ovary.
Fig. 28. Two anthers of pumpkin flowers; one anther has been partly devoured
by the maggots.
Fig. 29.. After the larve have destroyed the anthers, the pest devours the column
of the stamens.
Fig. 30. The maggots finally penetrate into the long peduncle of the staminate
flowers and feed on the wall of the stalk. The flower often drops
from its stalk due to decay caused by the larve.
PLATE XXX.
Figs. 31 and 32. String beans split open showing the melon fly larve feeding on
the seeds and flesh of the pods.
Fig. 33. After the seeds and flesh of the string beans have been devoured the
inner surfaces of the pods turn black.
Figs. 34 and 35. External appearance of the infested bean pods.
Figs. 36 and 37. Melon fly pupae inside of pods.
Figs. 38 and 39. Melon fly larve which have died while attempting to bore out
of the string beans, probably due to the drying of the pod.
Figs. 40 to 44. Dried and shriveled string beans after the melon fly larve have
bored out. These bean pods do not drop to the ground but remain
adhering to the plant.
PLATE XXXI.
Fig. 45. Channels in moistened sand made by melon flies after emerging from the
puparia. The black areas in the channels represent flies which
died in their attempt to burrow through the sand.
Fig. 46. Egg of melon fly.
Fig. 47. Recently hatched melon fly larva.
Fig. 48. Melon fly larva after feeding one day.
Fig. 49. Larva after feeding two days.
Fig. 50. Larva after feeding three days.
Fig. 51. Larva after feeding four days.
Fig. 52. Mature melon fly larva after feeding four and one-half days.
Fig. 53. Puparium of melon fly.
BIBLIOGRAPHY.
1. Clark, B. O., 1898. The Hawaiian. I, No. 27, p. 6.
2. Compete, G., 1912. Mon. Bull. State Comm. Hort., Cal. I, No. 10, pp.
709-730.
3. Coquillett, D. W., 1899. Ent. News, X, No. 5, pp. 129-130.
4. Ehrhorn, E. M., 1911. Hawaiian Forester and Agric. VII, No. 11, pp.
336-338.
5. Froggatt, W. W., 1910. Dep’t Agric. New South Wales, Farmers’ Bull.
No. 24, pp. 1-56.
6. Fuller, C., 1909-10. 7th. Gov. Ent., Dep’t Agric. Natal. p. 26.
7. Howlett, F.M., 1912. Trans. Ent. Soc. London. pt. II, pp. 412-418.
8. Illingworth, J. F., 1912. Cornell Univ. Agric. Exp. Sta. Bull. 324, pp. 126-188
9. Marsh, H. O.,1910. Rept. Div. Ent. for 1910. Bd. Agric. and Forestry,
Hawaii. pp. 152-159.
10. Perkins, R. C. L., 1902. Rept. Gov. Ter. Hawaii. p. 36.
11. Van Dine, D. L., 1907. Ann. Rept. Hawaii Agric. Exp. Sta. for 1907. U. S.
Dep’t. Agric. Office Exp. Stas. pp. 30-35.
ANNALS E.S. A. VoL. VII, PLATE XXVII.
Severin.
ANNALS E. S. A. VoL. VII, PLATE XXVIII.
Severin,
ANNALS E. S.A. VOL. VII, PLATE XXIX.
Severin.
ANNALS E. S.A.
Severin.
VOL. VII, PLATE XXX.
VoL. VII, PLATE XXXI.
ANNALS E. S. A.
12.
Sever
SOME NOTES ON LIFE HISTORY OF LADYBEETLES.*
By Miriam A, PALMER
The species treated in this paper are the more common
forms found in Colorado; namely, Hippodamia convergens
Guer., Coccinella 5-notata Kirby, Coccinella monticola Muls.,
Coccinella 9-notata Ubst., and Adalia melanopleura Lec.,
annectans Crotch, coloradensis Casey, and humeralis Say; also
incidentally, Olla abdominalis Say, Hippodamia sinuata Muls.,
and parenthesis Say, Coccinella sanguinea Linn., and Scymnus
sp. Special attention was paid to the duration of the life
cycle, and habits regarding egg-laying and feeding, with inci-
dental observations on injurious influences and other points,
also descriptions were made of the beetles in all stages.
Hippodamia convergens Guer.
This is our most common species, and may be described as
follows:
Adult: Fig. 1, Plate XXXII.
Head black, with pale frontal spot connected laterally with the eyes;
pronotum black with pale narrow border along apical and lateral mar-
gins; the two discal marks distinct and converging posteriorly; elytra
yellowish red, each with a scutellar spot and with six other spots rather
small in size and never united; the three posterior ones more developed
and constant. The spots are frequently lacking altogether and the
elytra immaculate. Legs black; length 6 to 7 mm.; width 3.5 to 4mm.
Egg: Fig. 2, Plate .©.C.6 0S
Pale to deep amber yellow, even as deep as yellow cadmium in some
cases; length 1.13 to 1.83 mm.; width .49 to .55 mm.
Larva: Fig. 3, Plate XXXII.
First instar: Entirely black with exception of pale area on lateral
and dorso-lateral margins of first abdominal segment; meso- and meta-
thorax each with two large setaceous areas and small setaceous spot
laterally; abdomen with three pairs of rows of setaceous areas. Second
instar: Same as first except that margins of pronotum are often pale
yellowish and the pale spots on the first abdominal segment are now
light orange colored, and in addition to these, faint orange spots are
seen similarly placed on the fourth segment. Third instar: Same as
before except that it is more orange colored in the light spots. Fourth
*This paper is an outgrowth of breeding cage work with the Coccinellids,
assigned me by Professor Gillette as a part of his Adams fund project on Life His-
tories of the Plant Lice and Their Enemies. Acknowledgments are also due to
Professor C. P. Gillette and Mr. L. C. Bragg for determination of Plant Lice
herein mentioned.
213
214 Annals Entomological Society of America _[Vol. VII,
instar: Same as third except for the addition of pale orange spots on
the 6th and 7th segments on dorso-lateral portion, not extending later-
ally. Some larve show quite a bluish frosted appearance over the black,
more especially toward the later stages. Length about 9.5 mm. for a
full grown larva.
Pupa: Fig. 4, Plate XXXII.
Ground color brownish yellow throughout; pronotum with black
lateral margin and an anterior and a posterior pair of black spots;
wing pads black along posterior margin and over apical half; each
segment with sublateral and medio-lateral spots. Spiracles black.
Sometimes much lighter in general color and only a dot found on wing
pad at basal third. Legs black. Length 5 mm.
Life cycle records* were made as follows:
Egg stage (22 records*) 3-7 days, mostly 3 days;
Larva stage (22 records) 10-28 days, mostly 14 days;
Pupa stage (22 records) 4-9 days, mostly 4-5 days;
Egg to adult, mostly 21 days.
Adult stage (13 records) 3-4 months for the summer generations.
A hibernating female lived 8 months and 12 days, dying May
25th, having laid eggs from April 27th to May 19th. This female
would doubtless have lived longer under natural conditions.
The weather was wet and unfavorable and the female was evi-
dently in a weak or unhealthy condition as evidenced by the
poor hatching of all the eggs. Another hibernating female
lived until July 6th. All the females which had laid eggs before
going into hibernation died either during the winter or in the
spring before laying any eggs. A male which had hibernated
lived 9 months and 8 days, dying June 13th.
The earliest egg record obtained was April 27th, the latest
record in 1908 was September 3rd, and in 1907, was October
30th. The earliest beetles reared emerged June 22nd and the
last one lived until September 22nd. Under natural conditions
and in early spring, doubtless the first generation emerges as
soon as the latter part of May or even earlier, according to the
season. There were at least three generations obtained in a
season.
The females would usually begin to lay within five days
after emerging and the longest egg-laying record obtained was
1 month and 18 days. Several egg counts were made. They
are as follows: 130 eggs (in 20 days), 199 eggs, 296 eggs, 312
eggs (in 1 month and 15 days), and 399 eggs, by different
*The term, record, in this sense shall be understood to mean, not notes made
on a single individual, but on an egg patch and the beetles resulting therefrom.
1914] Life History of Lady Beetles. 215.
females respectively. These numbers are certainly far below
’ what the beetles are able to lay under favorable conditions in a
state of nature. The number of eggs laid in a single batch was
usually about 30, though the number varies considerably both
above and below this number. When in good laying condition
a female will deposit from one to two batches a day.
Food records.—Several records were taken on the feeding
capacity of both larvae and adults for plant lice. One larva
ate 50 Prociphilus fraxinifolii Riley, another half grown, ate
33 Macrosiphum gaurae, and another half grown, ate 105 Chait.
negundinis Thos., in a single day. Entire counts of the whole
number of lice eaten during the larval period were taken in
four cases and are as follows: 264 (57 M. gaurae, 56 Chaitoph-
orus negundinis, 151 P. fraxinifolit), 309 (71 M. gaurae 137 C.
negundinis, 101 P. fraxinifolii), 585 (423 C. negundinis, 72 P.
fraxinifolii, 90 M.rosae Linn., small), and 576 (263 C. negundinis,
54 M. rudbeckiae Fitch, 114 P. fraxinifolit, 57 S. schizoneura
lanigera Hausmann, and 88 M. rosae), of all sizes, by each
larva respectively.
A pair of beetles, male and female, ate 150 Chattophorus
populifolii Fitch, in one day, and 120 Aplus setariae Thos., on
another day. A male made the following records on diferent
days respectively: 200 M. gaurae Will., 30 M. rudbeckiae, 33
A. torticauda Gill., 60 C. negundinis, a 75 M. cerast Fab.,
these lice being of all ages and sizes.
A female ate on different days, 36 M. rudbeckiae, 180 C.
negundinis, 165 M. cerasi, 110 A. heliantha Monell, and 120 A.
setariae respectively.
The amount eaten varied very considerably on different
days. For 12 hours before molting the larvae would sometimes
eat comparatively little, while for several days before pupating
they would be extremely voracious. Female adults always ate
the most when laying eggs. The temperature also affected
their appetites very remarkably, the highest records being made
on dry hot days, while in damp cool weather they often ate
almost nothing.
The state and temperature of the weather was indeed a
very important factor in the welfare of these insects. Cold
damp weather would very much retard the rate of growth of
the larvae and the egg-laying of the females and would seem to
216 Annals Entomological Society of America _[Vol. VII,
encourage diseases. Another injurious influence was the prac-
tice of cannibalism among the larvae and also the eating of eggs
and pupae by both adults and larvae. A female would often
eat her own eggs. As a rule, these habits were not practiced
unless there was scarcity of food and in some cases not even
then, while in other cases these deeds would be perpetrated with
apparently no excuse. Individuals seemed to differ widely
in this respect. Some kinds of lice proved injurious in the
breeding cage. Large lice such as M. rudbeckiae and M. am-
brosiae Thos., caused the death of several beetles by smearing
their mouths shut by means of the glue extruded from the corni-
cles so that the beetles starved to death. Probably the results
would not have proved fatal, had the beetles not already been
somewhat weakened by other injurious influences, and were it
not for the unnatural conditions of the breeding cage so that
the lice were restless and walking about constantly.
The larvae, especially in the younger stages, seemed to
prefer smaller lice in the cages, though out of doors they were
frequently found feeding on large ones with no apparent diffi-
culty. This protective function of the cornicles, however,
would probably, even in nature have more or less of an injurious
effect on the beetles.
This ladybeetle seems to be a very general feeder on plant
lice. One beetle was observed to eat a larva of Aphidoletes
marina, a dipteron, which is also predaceous on plant lice, but
whether it would make a practice of this in nature was not
ascertained, probably not, as in other cases the larvae of A phi-
doletes were rejected. An unusually hungry beetle or larva
would often chew or suck about on the leaves which were put
into the cage with the lice. They seemed to do no visible
injury to the leaves and were probably only sucking and licking
off the honey dew left by the aphids. Both the beetles and
larvae refused to eat chrysomelid eggs and membracid larvae
which were offered to them. No evidence was obtained of their
feeding, to any appreciable extent, on anything besides the living
plant lice. They do not seem to touch even newly laid
eggs of Aphididae.
The species of aphididae which were used for food in the
breeding cage were as follows: Aphis gossypii Glover, A.
oxybaphi Oest., A. carbocolor Gill., A. torticauda Gill., A. heliantht
1914] Life History of Lady Beetles. 214
Mon., A. pomi De Geer, A. cerastfolit Fitch, A. medicaginis,
Koch, Phorodon humult, Schrk., A. oenotherae Oest., A. brassicae
Linn., Chaitophorus negundinis Thos., C. populicola Thos.,
C. populifolu Fitch., Macrostphum ambrosiae Thos., M. rud-
beckiae Fitch., M. pisi Kalt (?), M. rosae Linn., Hyalopterus
arundinis Fab., M. cynosbati Oestl., Lachnus sp., Melano-
xantherium smithiae Monell, Myzus cerast Fab., Prociphilus
fraxinifoli Riley, S. lanigera Hausm, and Melanoxantherium
bicolor Oestl. .
Outdoors they were observed feeding on A. pomi, M.
rudbeckiae, C. negundinis, Myzus cerast, Mac. on Euphorbia,
A. atriplicis Linn., M. pist, and M. cynosbati. Although these
were all the lice noted, H. convergens has been found feeding on
practically all of the common plant lice; in fact, it does not
seem to be at all particular, and, both because of the wide range
of its feeding habits, and on account of the comparative hardi-
ness of constitution, probably does as much or more in con-
trolling the plant lice than any other Coccinellid.
Hippedamia sinuata Muls.
This species also is a comparatively common ladybeetle
about Fort Collins Colorado though it does not seem to rank
very high in economic importance in this vicinity.
Adult: Fig. 5, Plate XXXII.
Head black with fine apical line of whitish and median pale spot
connected laterally with the eyes. Pronotum black with pale border
along anterior and lateral margins with tendency to abbreviated acute
line before, though this is often wanting; two converging discal marks
as in convergens; elytra yellowish red with suture black from one to
three-fourths of entire length and each with four spots, the first near
humeral angle, the second, a large one, just back of the middle, near the
suture, the third slightly caudo-lateral of this, and another at apical
fourth, the second and third spots very often united and frequently
humeral spot, also, joined to second spot by means of vitta, coalescence
of third and fourth spots occasionally occurs, thus completing the entire
amalgamation of all the spots; legs black; general shape rather narrow;
length 5-6 mm., width 3-3.9 mm.
Egg: Fig. 8, Plate XXXII.
Amber yellow; length 1.4 mm.; width .6 mm.
Larva: Fig. 7, Plate XXXII.
First instar: Black except pale spot on lateral margin of first
abdominal segment and lateral and dorso-lateral of fourth abdominal
segment. Second instar: same as first except that pale spots are more
pronouncedly yellowish. Third instar: same as preceding but spots
218 Annals Entomological Society of America [Vol. VII,
have become orange and in addition the anterior and posterior borders
of the pronotum have become yellow orange (pale), also dorso-lateral
spots of yellow on posterior margin of metathorax, (which seems to be
a distinguishing mark in this species) and faint yellow spots dorso-later-
ally placed on fifth, sixth, and seventh segments, and whitish median
line on all thoracic segments; length 8.5 mm.
Pupa: Fig. 6, Plate XXXII. .
Ground color amber yellow covered with black except median line
of yellow throughout though somewhat broken at segment margins;
pronotum bordered with black; wing pads black except light spot on
shoulder, meso- and metathorax with pair of black spots, abdominal
segments black except median area and yellow orange spots on first and
fourth segments laterally placed, fainter ones similarly placed on fifth,
sixth, and seventh segments; legs black; length 5.7 mm.
Life cycle records were made as follows:
Egg stage (3 records) 3-6 days.
Larva stage (3 records) 21-23 days.
Pupa stage (1 record) 8 days.
Egg to adult, 32 to 37 days.
From these figures it would appear that the life cycle of this
species was considerably longer than in the other species studied.
This may, however, be partly due to the fact that these records
were all made in the latter part of the summer so that the beetles
were overtaken by cool weather towards the end of the larval
stage.
In nature this species does not seem to be a very general
feeder, being found, by Mr. Bragg, almost exclusively in the
grass, or more especially on the Carex, feeding on Rhop. braggit
Gill., and Callip., flabellus Sanb. In the breeding cage,
however, it readily fed on Mac. rudbeckiae, Mel. Smithiae, Aphis
cornifolit, Rhop. pastinaceae Linn., Aphis heraclet Koch, and
Chait. populicola.
Hippodamia parenthesis Say.
This species :anks about equal with H. sinuata in economic
importance.
Aduli: Fig. 9, Plate XXXII.
Head black with pale abbreviated median dash from the front and a
spot next each eye, usually connected with median line, sometimes
entire front portion of face pale; thorax black with anterior and lateral
margins and median abbreviated acute line before white, also square
spot at base; elytra pale reddish with yellowish area in middle of poster-
ior half, marked with common large spot near the base connected with
the scutel, and each with humeral spot, and pair of parenthesis shaped
1914] Life History of Lady Beetles. 219
dashes on posterior half sometimes united into a single large lunule;
legs blackish to black, tarsi brownish; general shape, somewhat pointed
posteriorly; length 4-5 mm., width 2-3 mm.
Egg: Fig. 10, Plate XXXII.
Amber yellow; length 1 mm., width .4 mm.
Larva: Fig. 11, Plate XXXII.
First instar; general color grayish or blackish; head shining black
except medio-anterior portion, setaceous spots black, as in all the other
species reared, dorso-lateral and lateral portions of first and dorso-
lateral portions of fourth abdominal segment pale, nonsetaceous portions
of thoracic segments rather pale. Final instar; head black except
medio-anterior portion, which is brown; pronotum bordered with pale,
meso- and metathorax with whitish median portion; yellow markings
as follows, lateral and dorso-lateral spots on first and dorso-lateral
spots on fourth abdominal segments orange yellow, and faint indica-
tions of dorso-lateral spots on fifth, sixth, and seventh segments pale
yellow; length 8 mm.
Life cycle records were made as follows:
Egg stage (1 record) 3 days.
Larva stage (1 record) 11 days.
Pupa stage (1 record) 6 days.
Egg to adult, 20 days.
This life cycle appears relatively short, but this is probably
due to the fact that this batch was reared in July when the
weather was specially favorable.
This species, like H. stnuata, has been found chiefly in the
grass and Carex feeding on Rhop. braggiu and Chait. fabellus,
tho in the breeding cage it seemed to thrive on Rhop. pastinceae.
Coccinella 5-notata: Kirby.
Ranking next in order, or perhaps equal with H. convergens
in economic value, is Coccinella 5-notata. This is somewhat
larger and more robust species and may be described as follows:
Adult: Fig. 12, Plate XXXII.
Head black with triangular pale spot next each eye; pronotum black
with subquadrate pale spot at the anterior angles; elytra brownish red,
marked with common subbasal fascia and a transverse spot on each
elytron before the middle, near the suture, and another at apical fourth.
Sometimes the subbasal fascia is broken into three spots, sometimes
there is also a dot on the anterior lateral portion of the elytra. Length
5.5 to 7.5 mm.; width 4.5 to 5.5 mm.
Eggs: Fig. 13, Plate XXXII.
Light to deep amber yellow in color; length 1.31 to 1.37 mm.;
width .53 to .60 mm.
220 Annals Entomological Society of America __[Vol. VII,
Larva: Fig. 14, Plate XXXII.
Indistinguishable from that of H. convergens until it reaches the third
or fourth instar when the head becomes pale, except on the posterior
and lateral margins, which remain black. Also, the yellow spots on the
abdomen are usually a stronger orange color and the dorso-lateral spots
on the 6th and 7th segments are lacking. The light blue frosted appear-
ance is sometimes more pronounced than in H. convergens. Length
about 12 mm., for a full grown larva.
Pupa: Fig. 15, Plate XXXII.
Ground color pale brownish yellow, tinged in places with pinkish,
paler, as a rule, than in the case of H. convergens. Pronotum with nar-
row black margin all around, except on median posterior portion; wing
pads black on posterior and apical two-thirds of anterior margin, also
two spots, one at basal third, and the other at apical third. These
spots often coalesce, darkening the whole apical two-thirds of the wing
pad. On the metathorax is a pair of black spots on the posterior
margin, medio-laterally placed; spots similarly placed on 2nd to 7th
segments inclusive, which segments also show another spot just within
the spiracles which often runs into the first mentioned spots along the
posterior margin. On the 3rd segment, the black often covers the entire
surface clear to the lateral edge, leaving only a pale line on the median.
Seventh segment often pale or nearly so. The amount of black varies
somewhat and is often less extensive, but the pupa can usually be dis-
tinguished from that of H. convergens by the more extensive black
markings, and by the paler ground color, but this does not always hold.
Legs black. Length, 6 mm.
Life cycle records were made as follows:
Egg stage (15 records) 3-5 days, mostly 3 days;
Larva stage (13 records) 11-19 days, mostly 13 days;
Pupa stage (12 records) 3-8 days, mostly 4 days;
Egg to adult, mostly 20 days.
Adult stage (4 records) 2-3 months for the summer generations.
The efforts to carry this species through hibernation all
resulted in failure, but seven beetles were captured before May,
19, 1908, which, in all probability, had hibernated, as no
larvae had been observed before that time or for some time
afterwards. The last of these beetles lived until August 14th,
and must have lived for nearly a year, as the beetles in the
cages which went into hibernation began to emerge August
6th. In the cage of these seven captured beetles, eggs were
found from May 19th to August 7th, a period of two months
and eighteen days.
Egg records were made as follows: 368 eggs (in 2 months
and 9 days); 469 eggs (in 1 month and 11 days), 532 eggs (in 1
month and 8 days), 539 eggs (in 1 month and 2 days) respective-
ly. Single egg batches often contain as many as 60 eggs; the
1914] Life History of Lady Beetles. = i ae
average size, however, would be nearer 30 to 50. A beetle in
good laying condition would often lay one or two batches a day,
though in the breeding cage this rate was not kept up very
long at a time.
The female would usually, under favorable conditions, begin
to lay from four to ten days after emerging and continued in one
case for one month and eleven days, and in another, the longest
record, for two months and fourteen days. The third genera-
tion was reached in a season in the breeding cage and both
2nd and 3rd generation beetles went into hibernation but none |
survived, the hibernating quarters being unsatisfactory. The
earliest eggs obtained in the spring were found May 19, 1908;
the first generation began to emerge June 17th, and their first
eggs were laid June 26th. The last of this brood lived until
September 21st. The latest record of eggs obtained was
September 14th. That year there was much cold wet weather
during August, which very considerably checked the egg
laying so that the beetles would very probably have laid for
at least a month later under favorable conditions.
Records on the feeding capacity of both larvae and adults
were taken as follows: One larva ate 595 and another 621
aphids during the entire larval period. The lice used in these
counts were of all ages and sizes. One ate C. negundinis 333,
M. gaurae 24, P. fraxinifolii 128, and M. rosae 110, total 595; the
other C. negundinis 445, M. gaurae 18, P. fraxintfoli 88, and
M. rosae 70, total 621. These lice were of all sizes. It will be
seen that C. negundinis was used more than any other one
species. One larva made the following records on single day
counts: during the first instar, 30 C. negundinis in a day; 2nd
instar, 84 C. negundinis; 3rd instar, 100 C. negundinis in a day.
These were days when all was favorable and the larva was on
full feed. An adult female when in best condition ate 200 A.
heliantht in a day.
The plant lice used for feed were the same as used for H.
convergens. This species seems to be just as general a feeder
as H. convergens and, being somewhat larger, would naturally
be expected to consume a few more lice, but it seemed to be
more delicate in constitution so that it succumbed more easily
to unfavorable conditions in the breeding cage. This species
was affected by the same injurious influences as were mentioned
for H. convergens.
222 Annals Entomological Society of America [Vol. VII,
In 1907 and 1908 this species, from casual local observations,
seemed to rank next to H. convergens, which was first in numbers,
but in 1909 it seemed to rank about fifth, with H. convergens
sixth. During this year, the first in numbers seemed to be
C. monticola which, during the two previous years, had ranked
third.
Coccinella monticola Muls.
This species is quite similar in general appearance to C.
5-notata and may be described as follows:
Adult: Fig. 16, Plate XX XIII.
Head black with triangular spot next each eye as in C. 5-notata;
pronotum black with quadrate pale spot at the anterior angles; elytra
brownish red, with common scutellar spot, and on each a broad trans-
verse oblique median fascia and a shorter subapical one black. Some-
times the median fascia is broken, leaving a small spot laterally. Legs
black. Length 5 to 7.5 mm.; width 4 to 4.75 mm.
Egg: ~ Wig!17, Plate XX XE.
Same as in C. 5-notata.
Larva: Fig. 18, Plate XX XIII.
Same as in C. 5-notata, except that in 3rd and 4th instars the head is
pale clear to the posterior margin, while C. 5-notata almost always has
a line of black along the posterior margin. Length of full grown larva
about 12 mm.
Pupa: Fig. 19, Plate XX XIII.
Ground color pale brownish yellow, usually paler than in C. 5-notata,
sometimes with pinkish spots on lateral portions of Ist and 4th seg-
ments of the abdomen where the orange spots were in the larva. Black
markings were as follows: Three spots on anterior edge of pronotum,
and one on posterior lateral margin, wing pads with two transversely
duplex spots, one at basal third and the other at apical third, and a spot
at base close to posterior lateral margin, which margin is also black;
median pair of spots on metathorax, also on 2nd to 6th abdominal
segments inclusive, smallest on 2nd; 8rd to 6th abdominal segments
also with spot within spiracles largest on third and inclined to extend to
the very margin of the segment. Knees black, remainder of legs
usually paler. Length 6.5 mm. The pupe of this species can usually
be distinguished from those of C. 5-notata by the less extensive black
markings, though the two species vary so as often to be indistinguish-
able.
Life cycle records were taken as follows:
Egg stage (17 records) 3-8 days, mostly 4 days;
Larva stage (11 records) 12-14 days, mostly 13 days;
Pupa stage (11 records) 4-8 days, mostly 6 days;
Egg to adult, mostly 23 days.
Adult stage (2 records of summer generation) 2 months and 6 days, and
53 months and 12 days respectively.
1914] . Lafe History of Lady Beetles. 223
Three beetles captured July 15, 1907 hibernated and the
last one died August 31, 1908, 13 months and 16 days from
date of capture. In this cage, during 1907, only four eggs
were laid, though it contained 9 beetles, but after hibernating,
eggs were laid rather abundantly, though they were infertile,
by one or more of the surviving three from May 11, 1908 to
July 18, 1908, a duration. of 2 months and 7 days. Three
other beetles which were captured May 15, 1908, and in all
probability had hibernated, lived until September 8, 1908, so
they must have been a year old, at least, eggs having been laid
from May 18, 1908 to August 18, 1908, a period of three months.
No egg records were taken on this species, as only captured
females laid at all satisfactorily and these had doubtless laid
a portion of their eggs while still out of doors, so a complete
record could not be gotten. The earliest eggs obtained in the
spring were found May 11, 1908, laid by females which had
hibernated in captivity; June 16, 1909, from captured females.
The latest record obtained in the fall was September 3, 1909,
but as this was the only year that a record was taken and as
August was very wet and cool, the beetles would probably be
able to lay for a month later, at least, under favorable condi-
tions. .
The earliest generation reared emerged June 24, 1908, and
began to lay July 17, 1908, and the last beetle died September
15, 1908, probably due to the unfavorable weather. Later
batches of the same generation which emerged July 4, 1908,
later went into hibernation but died during the winter. Beetles
emerging July 5, 1909, the earliest generation obtained that
year, went into hibernation.
It seems that this species may have both one and two
generations in a year, since some of what were evidently first
generation beetles went into hibernation while others of this
same generation laid eggs, infertile though they were, and died
before winter.
In feeding capacity they were about the same as 5-notata.
One larva from time of hatching to pupation ate 388 plant
lice, (23 A. torticauda, 65 C. negundinis, 165 C. populicola, 90
A. setariae, 45 M. ambrosiae); another ate 376 plant lice, (12
A. torticauda, 120 A. setariae, 174 C. negundinis, 40 C. populi-
cola, 30 M. ambrosiae). another ate 901 plant lice, (150 A.
224 Annals Entomological Society of America _[Vol. VII,
gossypi1, 480 A. helianthi, 19 A. setariae, 82 C. populicola, 140
A. pomi, 30 M. rudbeckiae); and another, 962 plant lice, (774 A.
gossypii, 108 M. pisi, 80 Myzus cerasz). One larva during the
first instar ate 30 A. gossypii in a single day; in the 2nd instar,
100 to 130 A. gossypii; third instar, 150 A. gossypit. Of course
these records were made when all conditions were favorable.
No records were taken on the adult.
The range of feed seemed to be the same as for H. convergens
and C. 5-notata. They were observed out doors feeding on
A. pomi, H. arundinis, M. cynosbati, and S. lanigera. Though
these were the only observations recorded they seem to feed on
practically every common species of plant lice.
As to injurious influences, they seemed to be affected by the
same factors as the foregoing species but seemed to stand
cool wet weather rather better. Some of these beetles got
rather badly pasted up by the glue from the cornicles of M.
rvudbeckiae but it did not result fatally. One beetle was found
with one front foot glued fast to its head, probably the result
of an attempt to clean itself for which operation the beetle was
not sufficiently vigorous, the foot sticking fast until it dried.
A nearly full grown larva was observed to attack a good
sized pupa of M. ambrosiae, biting it in the side of the third
abdominal segment. The louse immediately struck the larva
in the face with its cornicle and discharged a quantity of glue.
The larva paid no attention to this but continued until it had
finished the louse, and, inside of fifteen minutes, every trace
of glue was removed. Evidently when the beetles and larvae
are in vigorous condition this protective device of these aphids
produces little result beyond temporary annoyance.
This species seems to be peculiar in several points of its life
history as compared with the other Coccinellidae studied; first
in the fewer generations in a season, and the consequent greater
longevity of the beetles; and second, in the non-activity of the
males. Mr. Bragg, who had dissected at least 100 monticola
specimens, found only one male and he has some doubt as to
its really being this species. No monticola has been observed
in coition either indoors or out except with C. 9-notata or C.
5-notata males. The eggs of a female captured mated with
C. 9-notata all proved infertile, even though the male was kept
with the female for over two months and was frequently
1914] Life History of Lady Beetles.
bo
bo
Ort
observed in coition and the female laid a considerable number
of eggs. A male of 9-notata was introduced into a cage of
monticola which had been laying infertile eggs and though he
mated readily no fertile eggs were produced. One female
monticola was taken in coition with a male 5-notata. The eggs
were fertile and produced all monticola, but this female may have
been previously fertilized by another male so there is no evi-
dence that the 5-notata male had any effect.
There is a possibility of the males being weaker than the
females and so being killed in the struggle for existence before
they matured in the breeding cage, since about 50 per cent.
were, as a rule, lost when a number were reared together.
All the larvae from several batches were reared in individual
cages. In this way 90 percent were brought to maturity and
in the adult state all were put together in the same cage, but
they always died down to the usual 50 percent before there
was time for sexual development to be really completed, so that
there is a probability in these cases of the males having been
lost because of having a more delicate constitution than the
females. This would also account for their scarcity out of
doors.
Two batches were reared in individual cages, maturing 17
beetles from 18 larvae and 12 beetles from 23 larvae respectively.
These beetles were not put together but were dissected by Mr.
L. C. Bragg as soon as mature, the first lot proving to be 7
male and 9 female and the second lot 9 male and 3 female.
This latter case seems evidence against the possibility of the
males being weaker than the females, as in this batch little
over 50 percent matured and 75 percent were males. The
only explanation remaining seems to be a sexual non-activity
for some reason.
A good share of the females captured proved to be infertile,
when, however, a female is once fertilized it seems to suffice for
the season. Females captured May 15, 1908, laid fertile eggs
until August 18, 1908, a period of over three months, after which
they very soon died.
In spite of this seemingly weak point in its life history, this
beetle seems to rank quite high as an eneniy of plant lice.
226 Annals Entomological Society of America _[Vol. VII,
Coccinella 9-notata Hbst. :
Description of adult, Fig. 20 Plate XX XIII.
Head black with pale triangular spot next each eye, or these spots
maybe connected so as to form a broad. white band across the. face;
pronotum black with quadrate pale spot at anterior angles, and apical
margin usually pale, often broadly pale; elytra bright brownish red or.
red ochre, with common scutellar spot, and each with four other spots,
the two small ones sometimes connected by a-fine line, one in humeral
angle and the other near the lateral margin before the middle, and two
large ones, one near the suture and before the middle, the other rather
transverse and near the apex. Length 6-7.5 mm.; width 5-6 mm.
Egg: Fig. 21, Plate XXXIII.
Same as the foregoing species except, perhaps, a little smaller on
the average. ; f i
Larva: Fig. 22, Plate XX XIII. : es
First instar: Entirely black or blackish gray, except pale dorso-
lateral spots on first abdominal segment, and with setaceous tubercles
as in the foregoing species. Second instar: Same as Ist except that
median and anterior portion of head and anterior and posterior margins
of pronotum are slightly pale or dusky, the spots on the first abdominal
segment more yellow, and paler spots begin to show similarly placed on
Ath abdominal segment. Third instar: Same as last except that pale
portions of head and‘pronotum are lighter, and spots on abdomen are
more orange colored. Fourth instar: Same as last except pale portion
of head which is light yellow. The larve of this species seem to be
identical in appearance with those of 5-notata but do not grow to quite
as large a size. Length of full grown larva 9.5 mm.
Pupa: Fig. 23, Plate X XXIII.
Ground color light reddish or brownish yellow, often with pinkish
spots on 1st and 4th abdominal segments where the orange spots were
located inthe larva. © Pronotum with a pair of black dashes on anterior
margin, and a pair. of black dots on the posterio-lateral margin; a
median pair of black spots on metathorax and 2nd to 7th abdominal
segments inclusive; Spiracles black; Wing pads with posterio-lateral
margin black and a round black spot at basal third. This marking of
the wing pad is comparatively constant and nearly always distinguishes
it from all of the foregeing species, though sometimes this spot seems
to spread apically so that the pupa cannot be distinguished from 5-notata.
Length about 6 mm.
Life cycle records were taken as follows:
Egg stage (14 records) 3-6 days, mostly 4 days;
Larva stage (8 records) 10-21 days, mostly 11-14 days;
Pupa (8 records) 4-5 days, mostly 5 days;
Egg to adult, mostly 20-23 days.
Adult (7 records) 2 to 3 months for the summer generations.
1914] Life History of Lady Beetles. 227
Some of the beetles which went into hibernation were then
nearly four months old but did not survive the winter, probably
on account of improper winter quarters. Only one beetle sur-
vived. It had emerged September 14, 1908, and lived until
April 9, 1909
Four egg records were taken as follows:
435 eggs (in 1 month and 9 days); 493 eggs (1 month and 22
days); 950 eggs (in 2 months and 22 days); and 1047 eggs (in
2 months and 14 days). They laid from 40 to 68 eggs a day
when in full laying condition. Some of these beetles began
to lay as soon as one day after emerging. Others began at
7 days, and one at one and a half months after emerging. This
latter was in all probability not normal.
The 2nd generation was reached in the season when the
experiments were carried on, but, as a rather late start was
made in the spring, it is more than probable that they are able
to attain to the third generation when the spring is early.
The earliest eggs obtained were June 11, 1909 and the latest
record was September 2, 1909.
No exact feeding records were made with this species,
but judging from general observation they seemed to consume
as much as the foregoing species. Being as a rule somewhat
smaller than 5-notata and monticola, they probably ate a little
less. They seemed to take to the same range of feed as the
foregoing species in the breeding cage. This species seemed to
be susceptible to all of the injurious influences already mentioned
and besides these, several cases of parasitism were observed
in beetles captured out of doors. After emerging, the parasite
formed a silky cocoon underneath the beetle. The beetle,
though still alive, seemed unable to leave the cocoon of its
enemy and clasped it with all of its feet as though to protect it.
One was carefully examined and found to be perfectly free
from any attachment and the only reason for its remaining
there seemed to be a partial paralysis. When taken off the
cocoon, which was accomplished with considerable difficulty
because of the beetle holding so persistently with its tarsi, the
beetle seemed to be unable to walk or even stand, and when
offered food it made vain attempts to eat. It seemed absolutely
helpless from inability to co-ordinate its movements.
228 Annals Entomological Society of America _[Vol. VII,
There were no serious results in this species from the glue
thrown out from the cornicles of the large species of lice but this
was probably only accidental and not because of any specific
resistent character. Once, a small larva, apparently in the
second instar, was observed to seize a full grown apterous
Macrosiphum ambrosiae which appeared to be three times the
size of itself. It first caught it by the right hind foot. The
louse struggled, dragging the larva about for a minute or two
and extruded glue from the left cornicle. The larva then
managed to get hold just behind the right cornicle and pro-
ceeded to feed on the louse, though dragged about somewhat.
by the latter. Two and a half hours later it was found still
feeding on the louse, which was still living, but had extruded
no glue from the right cornicle on which side the larva was
holding it. Unless for some reason the right cornicle contained
no glue, this action of the louse seems to be explained only by
awkwardness or stupidity. Indeed, the cases were extremely
few when the louse seemed to make any well directed effort to
smear its enemy.
Perhaps next in order might be classed several forms of
Adalia Muls., namely, melanopleura Lec., annectans Crotch,
coloradensis Casey, and humeralis Say. As these four forms
interbreed freely and seem to be identical in life history it
seems best to treat of them here as one species, which indeed
they undoubtedly are. In the descriptions of the adults, however,.
they must be treated separately. In the larval and pupal
stages they seem to be identical in appearance but in the adult
state are strikingly different.
For full description of adults of these forms, see Annals of
. the Entomological Society of America, Volume 4, page 283.
figg: Fig. J, Plate XIX*.
Pale to deep amber yellow in color as in the foregoing species.
Length about 1 mm.; width about .56 mm.
Larva: Fig. I, Plate X1X*.
First instar: Head and pronotum shining black, rest of body dull
black or dark grayish with a median row of paler spots from pronotum
to tip of abdomen; 2nd instar, the same except lateral and medio-
lateral pale whitish spots on lst abdominal segment and a median pair
of whitish spots on 4th abdominal segment; pronotum bordered all
around with whitish and with median pale line; 3rd and 4th instars;
color often blackish gray or blue; median and anterior portion of head
*Annals E. S. A. Vol IV, Sept. 1911.
1914] Life History of Lady Beetles. 229
dusky or pale, a pair of pale spots on anterior lateral margin and three
on posterior margin of pronotum; also a pale median line, and pale
median portion of meso- and meta-thorax quite noticeable; spots on
abdomen more noticeable than in previous instars. Legs black. Length
of full grown larva 8-9 mm.
Pupa:' Fig. H, Plate XIX™*.
Ground color pale to sordid flesh color, tinged in places with brown-
ish. Pronotum with pale median line and broad lateral portion pale,
remainder dusky to black. Wing pads brownish to black, meso- and
meta-thorax dusky to black except median pale line; each abdominal
segment with three pairs of rows of dusky to black spots, median row
lighter on Ist and 2nd segments, giving a lighter appearance to that
part of the pupa. In dark specimens the markings are so extensive as
to almost coalesce, giving quite a melanic appearance. Legs brownish
to blackish. Length 4-5 mm.
Life cycle records were taken as follows:
Egg stage (84 records) 3-7 days in the spring, 2-6 in summer, mostly 3-5.
Larval stage (68 records) 11-18 days in the spring, 7-18 summer, mostly 15
spring and 11 summer.
Pupa stage (68 records) 5-9 days in the spring, 3-9 summer, mostly seven.
Egg to adult, mostly 21 to 23 days.
Adult (9 records) 1 month to 4 months, for the summer generations.
The records taken in the spring, April and May, it will be
noted, covered more days than those taken in the summer,
owing to the cooler temperature. Five beetles of first to
third generation that had laid a considerable number of eggs
and were from three to four months old, went into hibernation.
One of these lived until April 4th the next spring, aged 8 months
and 6 days. The rest died during the winter, though the hiber-
nating quarters were excellent. As there is a probability that
the first female captured, May 13, 1910, and with which the
start was made in the breeding cage, was not a hibernating
female, there is some uncertainty as to the number of each
generation. The beetles which survived hibernation were
the 4th or 5th generation and they survived in large numbers,
about 150 beetles in all.
A pair of beetles, the female of which was 3rd or 4th genera-
tion humeralis that had laid no eggs, and the male 2nd or 3rd
generation annectans, hibernated successfully, and in the spring
the female laid eggs abundantly until June 12, 1911, and they
both lived until June 16, 1911, the female aged nine months
and the male nine months and four days. Almost all of the
hibernating beetles which were cared for were still living May
230 Annals Entomological Society of America [Vol. VII,
23, 1911, aged about eight months, and a number were living
June 12, 1911, but all died soon after, some being nine months
old or more.
The females would usually begin laying about :ix days
after emerging and continue for about three weeks. One
continued for three months, by far the longest record made,
but probably the most nearly correct as most records were
doubtless cut unnaturally short on account of the great diffi- ~
culty of keeping the beetles healthy in captivity. The hiber-
nating females of all the three forms began to lay in six days
after having been brought in from hibernating quarters and
given food.
There seemed to be at least four, and possibly five genera-
tions during the season that the experiments were carried on,
though 38rd generation and possibly 2nd generation individu-
als hibernated successfully. There was no evidence of any
females which had laid eggs before winter hibernating and
laying again in the spring, though it may possibly take
place. The hibernating quarters used for these beetles were
excellent and apparently gave the beetles every chance.
The earliest eggs were obtained April 10, 1911, and the first
‘f the first generation reared emerged May 8, 1911. The
latest eggs in the fall were obtained September 16, 1910, but if
there had been no prematurely cold weather, the beetles might
have kept on laying for several weeks longer. From two or
three tests made, a female seemed not to be able to lay fertile
eggs longer than three or four weeks after being separated from
a male. If anew male was introduced, however, it seemed to
affect the eggs almost immediately and after two or three-days
none of the progeny showed any of the characters of the former
male, even when little or no interim existed between the males.
No feeding or counts were taken on this species as all the
time in the work with these beetles was used in heredity in-
vestigation * and only such life history data as took but little
time were noted. On general observation it was evident
that they eat much less than the foregoing larger species, and
in the breeding cage, at least, will not eat as large lice as the
above ladybeetles. Probably out of doors, when the lice are
quiet instead of restless as in the cages, they may feed upon
*See ‘‘Some Notes on Heredity in the Coccinellid Genus, Adalia Muls.’’
Annals of Ent. Soc. of Am., Vol IV, No. 3.
1914] Life History of Lady Beetles. 231
_ larger species. Mr. Bragg says that he found them in numbers
with Prociphilus fraxintfolii, a species which, from their large
size and great quantities of honeydew, would have certainly
brought disaster in the breeding cage, for great care had to, be
exercised in feeding these even to the larger species of beetles.
They seemed to thrive in confinement on C. negundinis, A.
setariae, A. helianthi, C. populifolit, C. populicola, M. san-
borni Gill., Myzus persicae, Sulz., Rhopholosiphum pastinacae,
Aphis heraclei Koch, M. cynosbatt. Outdoors they were
found with P. fraxinifolii. n 1910 and 1911 this species
seemed especially abundant. In the spring of 1910 the box-
elders were extremely lousy but by June 26, it was extremely
dificult to find any C. negundinis excepting the dimorphic
form which the ladybeetles seemed unable to eat. This species
of beetles, together with Syrphus larvae, seemed to be chiefly
responsible for the destruction of the lice. Again this spring,
1911, they seem to be the chief factors in cleaning up the box-
elder lice which were almost killing some trees.
These beetles seemed to have a more delicate constitution
than the other species reared and had to be tended with more
care, and even with the best care, hardly 25 percent could be
matured, and often less in the pure strains. The hybrids were
much more vigorous, and of them it was often possible to rear
50 percent to maturity. These beetles were susceptible to all
of the injurious influences which have been named, and besides,
they seemed to object to what might have been an odor left
by a certain species of ants, as the cottonwood C. populifolit,
which is abundantly attended by ants, was rejected at times
and accepted at others with no other apparent reason.
On the occasion of rejection they would turn cannibal and almost
the entire cage of larvae would be lost in one day, as though
they had been left without any feed at all. It seems, however,
that in spite of their seemingly rather frail constitution, they
hold their own in nature pretty well, as during 1909 and 1910
and the spring, at least, of this year (1911) they ranked high
in comparative numbers out of doors and doubtless have been
very beneficial.
232 Annals Entomological Society of America _[Vol. VII,
-Olla abdominalis Say.
This species is sometimes found, though it is rather rare.
One larva was found on a boxelder tree with C. negundinis June
25, 1909, and was reared to maturity. It was taken when
nearly full grown.
Larva: Fig. 25, Plate XX XIII.
General color black, except medio-anterior portion of the head
which was pale yellow to dusky; anterior portion of pronotum and
median spot on posterior margin pale; median spots of pale yellow on
meso-and meta-thorax; dorso-lateral yellow spots on first abdominal
segment, and dorsal and dorso-lateral on 4th abdominal segment;
lateral tubercles pale or partly so. Median pale spot on median post-
erior margin of each segment becoming larger toward caudal end of
abdomen and giving the appearance of a pale median longitudinal line
the entire length of the abdomen. Legs black.
This larva resembled the Adalia larvee very much but grew to be
somewhat larger than they did. Length of full grown larva 10 mm.
Pupa: Fig. 26, Plate XX XIII.
Ground color whitish tinged with yellowish and brownish in places.
Markings asin Adalia. Length 5mm.
Adult: Fig. 24, Plate XX XIII.
- Head white except two black spots on the posterior margin; pro-
notum with (M) design as in Adalia, but with the white more extensive
so that the pale spots often run together in places; elytra pale maize
yellow, each with a row of four basal spots, a row of three just before
the middle, the inner one crescent-shaped, also one subapical near the
lateral margin. Sometimes this apical spot and the middle row are all
connected into a triangular black patch. Legs brownish yellow.
Length 4-6.5 mm.; width 4-5 mm. No work has been done with this
beetle further than the rearing of the one larva found. This larva was
found with C. negundinis on boxelder.
Coccinella sanguinea Linn.
This seems to be the species designated by Casey as Cycloneda
rubripennis Casey, and by C. W. Leng as Subgenus Cycloneda
sanguinea var. rubripennis Casey.
This species is comparatively if not indeed quite rare about
Fort Collins, Colorado, but seems to occur more plentifully in
the vicinity of Boulder, Colorado.
Adult: Fig. 27, Plate X XXIII.
Head black with broad white border next each eye, often also
median pale spot and in many cases entire front of head white; prono-
tum black with rather narrow pale border along sides and extending
posteriorly and internally along the base to about lateral sixth, apical
pale line quite narrow with median acute line often rudimentary, also
isolated sublateral pale spot which is sometimes narrowly connected to
1914] Life History of Lady Beetles. 233
_the -pale apical margin; elytra deep and bright scarlet with transverse
pale spot on each side of scutellum; legs black to brownish, tarsi brown-
ish; general shape broadly oval and rather strongly convex. Length
4.5 to 5.5 mm.; width 3.5 to 4.75 mm.
Egg: . Fig. 28, Plate XX XIII. !
Pale amber yellow; length 1.36 to 1.5 mm.; width .63 to .7 mm.
Larva: Fig. 30, Plate XX XIII.
First instar: Black or dusky with pale lateral spot on first abdom-
inal segment, setaceous spots black as in other species. Second instar:
Same as first except that fourth abdominal segment shows pale lateral
and dorsal spots, and sometimes lateral pale spot appears on fifth and
sixth segments, median of meso- and meta-thorax whitish to yellow.
Third instar: Same as previous except that yellow spots on fifth and
sixth segments and on median of thoracic segments are always pro-
nounced and anterior margin of pronotum is also pale, median of
abdominal segments whitish, giving effect of light median stripe;
median and anterior portion of head brownish, while lateral and posterior
portion is deep black; legs, especially front pair unusually long. Fourth
instar: Same as preceding except that the spots are more yellow and
lateral margins of all thoracic segments are pale whitish or yellowish;
legs still longer than those of the other species studied; general color
dark bluish. This larva strongly resembles the larva of Olla abdom-
inalis.
Pupa: Fig. 29, Plate XX XIII.
Ground color pale maize yellow, darker in spots, especially in the
depressions between the abdominal segments, a broad band along
anterior and lateral margins and a round spot within lateral margin on
each side of pronotum. Wing pads dark maize to honey yellow or
sometimes quite brownish with posterior margins always brown; black
markings as follows: Pronotum with a pair of black spots on anterior
and another on posterior margin, also an anterio-lateral pair less pro-
nounced, a pair of median spots on meso- and meta-thorax and on
abdominal segments two to six inclusive, third segment with three
pairs of spots; legs brownish honey yellow. Length 5 mm.
Life cycle records were made as follows:
Egg stage (7 records) 3-5 days, mostly 4 days.
Larva stage (7 records) 9-16 days, mostly 11 days.
Pupa stage (7 records) 3-5 days, mostly 4 or 5 days.
Egg to adult, 15 to 20 days.
This species has been observed by Mr. Bragg feeding on
Mac. solidaginis, and in the breeding cages it did well on Aphis
heraclet Koch., Rhop. pastinaceae, Mac. gaurae, Mel. smithiae,
Aphis setariae, Aphis oxybaphi, and Aphis helianthi.
Some larvae of Scymnus species, determined by Major
Casey as probably a new species somewhere in the vicinity of
cockerellt Casey and consobrinus Lec., were reared but’ no life
history notes were taken.
234 Annals Entomological Society of America [Vol. VII,
Adult: Fig. 31,Plate XX XIII.
Head black, yellowish red between the eyes; pronotum black with
lateral thirds yellowish red; elytra black; legs brownish red; entire
beetle covered with hairs; length 2.5 mm.; width 1.7 mm.
Egg:
Same color and shape as foregoing species but much smaller.
Larva: Fig. 32, Plate XX XIII.
Pale brownish throughout, but covered with six large tufts of waxy
secretion on each segment so as to render the larva quite conspicuous.
In the pupa the larval skin is not pushed down as tightly as in the
other Coccinellids studied, so that it gives the pupa a white cottony
appearance and doubtless affords it considerable protection.
This species has often been found doing valuable service in
keeping <A. setariae in check on the plum. Professor Gillette
has often found it abundant with Schiz. lanigera on the apple.
GENERAL SUMMARY.
In general appearance and color pattern these species of
ladybeetles resembled each other most in the larval and pupal
stages. Coccinella 5-notata, 9-notata, and monticola resemble’
each other so much as to form one group. They of course had
some characteristic differences but they often intergraded and
merged together so as to be indistinguishable until they matured.
H. convergens, together with H. parenthesis and H. sinuata
seemed pretty distinct in these stages but an occasional indi-
vidual was found which seemed to show no distinguishing
character. The Adalia beetles, O. abdominalis, and C. sanguinea
seemed to form another group in color pattern of the larva and
pupa. The forms of the Adalia seemed to be exactly identical
in these early stages. Scymnus, of course, with its covering
of waxy secretion, was entirely different.
In life cycle periods from egg to adult, all the species thus
studied, H. convergens, C. 5-notata, 9-notata, monticola and the
genus Adalia seemed practically alike. From egg to adult
in H. convergens took, as a rule, 21 days; in C. 5-notata, 20
days; in C. monticola, 23 days; in 9-notata, 20-23 days;
Adalia, 21-23 days. The age to which the adults lived seemed
practically the same, two to three and perhaps four months for
the summer generations. In C. monticola, however, it seemed
that the first generation commonly hibernate so that these
beetles would live to a greater age than even the hibernating.
beetles of the other species in which only the beetles emerging
1914] Life History of Lady Beetles. 250
later in the season hibernated. There did not seem to be any
certain fixed generation to hibernate in any of the species but
there was no evidence of any females that had laid eggs before
winter hibernating and laying again the next spring, and none
ever hibernated in other than the adult state.
Of all the species studied, C. monticola was the only one
where there was any difficulty in breeding males in captivity;
in all other species about half of the beetles reared were males,
but here there was no evidence of any, and only females cap-
tured already fertilized, laid eggs which would hatch. Many
of these captured laid infertile eggs or none at all, and none reared
in captivity ever laid a fertile egg and most did not lay at all.
For three years, no undoubted instance of, the finding of a male
occurred either biologically by myself, or by Mr. Bragg, who
used dissection on captured specimens and had no difficulty
_ finding males in any of the other species. Finally, however,
two batches of eggs were reared with special care, one lot
producing sixteen beetles from eighteen larvae hatched, and
the second producing twelve beetles from twenty-three larvae
hatched. The first lot proved, on dissection, to consist of seven
males and nine females. The second lot consisted of nine
males and three females. Why none have been observed
mated, either in captivity or out of doors, and why Mr. Bragg
found so much difficulty in capturing males, still remains
unknown.
The egg laying periods seemed to be approximately the
same for all these species where records were taken, both for
the length of time for the individual, and for the laying season.
A female would often lay before being fertilized but not as well
as after. A female of Adalia would not seem to be able to lay
fertile eggs for more than about three weeks after being isolated
from a male. In C. monticola, on the other hand, if a female
was once fertilized, it sufficed for the season. In Adalia, when a
female was changed from one male to another, the later male
would take precedence over the former one, almost immedi-
ately, so that the eggs laid two or three days after and later would
develop the characters of the later male in every case.
In the individual egg records, C. 9-notata ranked first, the
four highest records being from 435 to 1047; C. 5-notata next,
with 368 to 539; then H. convergens with 199 to 312, these
numbers representing the four highest records of each species
236 Annals Entomological Society of America [Vol. VII,
respectively. Owing to the difficulty in getting these beetles
to lay their full number of eggs in captivity, many of the num-
-bers here may fall far short of what the beetles are able to do,
and the differences between the species may have been more or
less accidental, though C. 9-notata seemed very decidedly to be
the most prolific.
Considering the feeding capacity of the larvae and adults
of these species, there seemed little difference between them.
Aflarva of H. convergens was able to eat 100 C. negundinis in a
single day, and the highest entire record was 576 aphids of
different kinds. <A larva of C. 4-notata could eat 100 C.
negundinis in a day and the highest entire record was 620
aphids. A larva of C. monticola ate 144 to 190 A. corntfolit in
a single day, and ‘the highest entire record was 962 aphids.
An adult of H. conver.ens ate 120 A. setariae in a single day,
andjone of C. 5-notata, 200 A. helianthi.
Both in the per day, and the entire record of the larva, C.
monticola leads, and C. 5-notata ranks second. ‘These species
are on the whole practically alike in feeding habits and what dif-
ferences there seem to be may be partly accidental as counts '
were taken on only four larvae of H. convergens, only two of
C. 5-notata, and five of C. monticola, and only two counts of
C. monticola exceeded the other species, the other three being
no higher than in the others. The amount eaten per day
varied greatly with the larvae according to the weather and the
size of the larva, and in the adult it varied with weather and
egg laying.
In range of feed H. convergens, C. 5-notata, C. monticola and
C. 9-notata seemed practically alike and seemed to comprise
everything in the way of aphids except aphid eggs, though
the smaller species seemed to be preferred in the breeding cage.
Out of doors they all seemed to feed on the large species as well
as the smaller ones. For these four species of ladybeetles the
following plant lice were used as feed: A. cerastfoli1, A.
gossypi1, A. oenotherae, A. carbocolor, A. taraxici, A. torticauda,
A. oxybaphi, A. helianthi, A. setariae, A. medicaginis, A. brass-
ice, H. arundinis, Lachnus sp., C. negundinis, C. populidola,
C. populifolii, M. gaure, M. cynosbati, M. pist, Myzus cerasi,
Macrosiphum ambrosie, M. rudbeckie, Prociphilus fraxinifolit,
Phorodon humuli, S. lanigera, Melanoxantherium bicolor and
M. smithiae.
1914] Life History of Lady Beetles. Dad,
There was no evidence of any of these species feeding on
vegetable matter, though they often chew and suck about on the
leaves when very hungry and one newly emerged Adalia
beetle seems to have chewed up a portion of a half dried boxelder
leaf.
The Adalia beetles had to be fed on only the smaller species
of plant lice in the breeding cages, though out of doors they were
found by Mr. Bragg to be quite abundant with P. fraxinifolit.
Early in the spring, 1910 and 1911,'they were very abundant
with C. negundinis and together with the Syrphus larve they
cleaned these lice off the boxelder trees, though they had been
very badly infested. The species used successfully in the lab-
oratory were C. negundinis, M. sanborni, Myzus persice, A.
setari@, A. helianthi, R. pastinace, and Macrosiphum cynosbati.
The injurious influences affecting ‘these species were cool
damp weather, which C. monticola seemed to stand better than
the other species, very large lice that would extrude large
quantities of glue from their cornicles, a fungous disease re-
sulting from too much dampness in the cage or perhaps from the
decaying bodies of half eaten lice, and frequently beetles were
destroyed by a hymenopterous parasite known as Perilitus
americanus. Ants, also, seemed to be hostile, in one instance
killing a larva and in another, an adult, this even in the breeding
cage where the ants felt strange and were frightened. Much
loss was occasioned by cannibalism, eggs, larve, pup, and
even newly emerged adult beetles, while still soft, being eaten
by hungry brothers either larve or adults.
As to members in nature, the different species seem to rank
differently in different years. From casual local observations,
in 1907 and 1908, H. convergens ranked first, C. 5-notata second,
C. monticola third, C. 9-notata fourth, and O. abdominalis and
Adalia quite rare. In 1909, C. monticola was first, Adalia
annectans second, H. convergens, C. 5-notata, and C. 9-notata
not very abundant, and O. abdominalis only occasional.
238
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Sete SNA ge il Mig
Annals Entomological Society of America
.
EXPLANATIONS OF PLATES.
PLATE XXXII.
Adult of Hippodamia convergens Guer.
Eggs of Hippodamia convergens Guer.
Larva of Hippodamia convergens Guer.
Pupa of Hippodamia convergens Guer.
Adult of Hippodamia sinuata Muls.
Pupa of Hippodamia sinuata Muls.
Larva of Hippodamia sinuata Muls.
Eggs of Hippodamia sinuata Muls.
Adult of Hippodamia parenthesis Say.
Eggs of Hippodamia parenthesis Say.
Larva of Hippodamia parenthesis Say.
Adult of Coccinella 5-notata Kirby.
Eggs of Coccinella 5-notata Kirby.
Larva of Coccinella 5-notata Kirby.
Pupa of Coccinella 5-notata Kirby.
All drawings magnified five diameters.
All drawings except Fig. 31 are magnified five diameters.
PLATE XX XIII
Adult of Coccinella monticola Muls.
Eggs of Coccinella monticola Muls.
Larva of Coccinella monticola Muls.
Pupa of Coccinella monticola Muls.
Adult of Coccinella 9-notata Hbst.
Eggs of Coccinella 9-notata Hbst.
Larva of Coccinella 9-notata Hbst.
Pupa cf Coccinella 9-notata Hbst.
Adult of Olla abdominalis Say.
Larva of Olla abdominalis Say.
Pupa of Olla abdominalis Say.
Adult of Coccinella (Cycloneda) sanguinea Linn.
Eggs of Coccinella (Cycloneda) sanguinea Linn.
Pupa of Coccinella (Cycloneda) sanguinea Linn.
Larva of Coccinella (Cycloneda) sanguinea Linn.
Adult of Scymnus sp. magnified 8 diameters.
Larva of Scymnus sp.
[Vol. VII,
VOL. VII, PLATE XXXII.
ANNALS E. S. A.
Miriam A, Palmer
a Len pipers Pi (ed aa
OAL fie ot eee ‘
:
ANNALS E. S. A. VOL. Vil, PEATE xoxo nE
Miriam A, Palmer
~
ON A COLLECTION OF CRANE-FLIES (TIPULIDAE
DIPTERA) FROM THE FIJI ISLANDS.
- By CHARLES PAuL ALEXANDER, Ithaca, N. Y.*
The following crane-flies were included in material sent to
Mr. Harry H. Knight by Dr. James F. Illingworth. The only
published reference to the Tipulid fauna of these islands is
included in Skuses Diptera of Australia (vol. IV, 2nd series,
1889) and his records are mentioned herewith. I am indebted
to Dr. Illingworth and Mr. Knight for this material. The
types are in the collection of the author.
Family Tipulidae. Subfamily Limnobinae. Tribe Limnobini.
Dicranomyia saltens Doleschall.
1857 Limnobia saltens Doleschall; Nat. Tijd. Ned. Ind., vol. 14,
Dp. oon, plo 2; figs.
Two, a male and a female, from Nadi on July 27. This
seems to be the most easterly station so far made known for
this species; it has been recorded from southern India, Java,
the Philippine Islands, etc.
Dicranomyia illingworthi, sp. n.
Wings hyaline with sparse brown markings; Sc short ending just
beyond the origin of Rs.
Male, length, 4.6 --5 mm.; wing, 5.2 --6.8 mm.
Male: Rostrum and palpi brown. Antenne dark brown, the
flagellar segments globular in shape. Head dark brown.
Thorax rather uniformly dark brown, stripes on the praescutum not
well-defined, lobes of the scutum a little darker. Pleure uniform brown.
Halteres pale, stem moderate in length. Legs, coxe light brown,
trochanters brown. Wings hyaline, veins brown; a small rounded
brown stigmal spot; pale seams at Sc2, base of Rs and on the cross-veins
and deflections of veins along the cord and outer end of cell Ist Me.
Venation: (See fig. 1.) Sc short, ending just beyond the origin of Rs.
Basal deflection of Cu, before the fork of M.
Abdominal tergites slightly darkened, the sternites pale, the abdo-
men rather transparent. Hypopygium with the pleurites short, cylin-
drical; dorsal appendage chitinized short, slightly curved and acute at
apex; ventral appendage large, pale, almost white, with the outline
rounded, the inner lobe produced mesad into a cylindrical, feebly
chitinized, point that bears two bristles which are directed caudad.
(See fig. 7.) ,
*Contribution from the Entomological Laboratory of Cornell University.
239
240 Annals Entomological Society of America [Vol. VII,
Holotype, o', Nadi, Fiji Is., 7-28, 13. Paratypes, 4 o's;
with the type.
Libnotes strigivena Walker.
Limnobia strigivena Walker, Journ. Linn. Soc. Lond., V, 229, 1861.
This species is recorded by Skuse (Diptera of Australia,
Proc. Linn. Soc. N. S. Wales, IV, series 2nd, 787, 1889).
Tribe Antochini.
Teucholabis fijiensis, sp. n.
Head dark; thorax with three brown stripes, pleuree spotted with
brown; wings yellowish with brown spots.
Male, length, 7 mm.; wing, 7.4 mm.
Male: Rostrum brown, palpi dark brown. Antenne with the
basal segments of the flagellum rounded, brown. Eyes large, contigu-
ous on the vertex; head dark greyish black.
Pronotal scutum dull yellow, brown medially above; a brown spot
on the lateral end. Mesonotal praescutum light yellow with three
stripes of which the median one is longest and broadest, extending from
the cephalic margin to the transverse suture. The lateral stripes are
short and narrow, behind, crossing the transverse suture and ending on
the anterior border of the scutal lobes; the lateral margin of the sclerite
is suffused with dark brown. Scutum and scutellum brown except the
median portion of the former which is pale. Postnotum dark brown.
Pleurze dull yellow, the episternites of the pro- and mesothoraces brown.
Halteres pale. Legs, coxa, fore and middle, brown, hinder pair paler;
trochanters pale yellow; femora yellowish, brown with a broad brown
_ subapical ring; tibize and tarsi brown. Wings yellowish, stigmal spot
large, prominent, a small seam on Sc2; seams at the base of Rs, along
the cord and on the outer end of cell lst Ms. Venation: (See fig. 2.)
cross-vein r at the tip of the long R; and so placed slightly beyond the
middle of Rois.
Abdominal tergites dark brown, the caudal margin a little more
yellowish; the basal two or three sternites yellowish, the others more
brown. Hypopygium with the ninth tergite having the caudal margin
rounded and very feebly notched medially. Pleurze short, clothed
with sparse long hairs. Dorsal appendage of the pleurite jointed at the
base, pale, clothed with numerous long hairs at the base, at the tip
slightly bifid underneath: The ventral appendage is a long elongation
of the pleura, not jointed at its base, darker and more chitinized;
toward the tip it is constricted, the actual apex expanded and bearing
a few small hairs. (See fig. 8).
Holotype, &, Nadi, Fiji Is., 7-28,'13.
—_—
1914]. Tipulidae from Fiji Islands. 241
Tribe Eriopterini.
Gonomyia (Leiponeura) fijiensis, sp. n.
Thorax brown, lateral margin of the praescutum yellow; wings with
the costal margin yellow, the membrane light brown and hyaline diver-
sified, stigma lacking.
Male, length, 4.9 mm. Female, length, 4.8-5.1 mm.; wing, 4.3-
4.4 mm.
Male: Rostrum and palpi dark brown. Antenne light sulphur
yellow, the flagellar segments a little paler. The head bright sulphur
yellow with three pale brown marks, a median one on the frontal tubercle
and others on the sides of the vertex.
Pronotum light yellow. Mesonotal praescutum dark clove-brown
the lateral margin between the pseudcsuture and the transverse suture
yellow, scutum, scutellum amd postnotum dark brown. Pleure light
yellow, a broad brown lateral stripe, deepest ventrally fading into the
yellow of the dorsal pleurites above, extending from the ventral surface
of the cervical sclerites through the halteres to the posterior portion of
the mesonotal postnotum; the area between this stripe and the praescu-
tum is light yellow suffused with brown near this stripe; sternites dark
brown. Halteres light sulphur-yellow. Legs, fore cox light sulphur-
yellow except the extreme tip which is dark brown; trochariters light
brown; remaining coxe dark brown on the. basal half, paler brown
apically and on the trochanters; remainder of the legs broken off and
confused in the vials with the legs of several other species, but they are
probably uniform dark brown. Wings with the costa and the subcosta
conspicuously bright sulphur-yellow, remaining veins brown; wing
suffused with brown and variegated in places with hyaline, as in cell Ri
which completely lacks a stigma, in cell R, lst Mz and elsewhere. Vena-
tion as in figure 3.
Abdominal tergites brown, broadly edged with yellow on the caudo-
lateral margins, the brown always continuing to the caudal margin as a
narrow median line except in the Sth tergite where the lateral and
caudal margin is broadly yellow all around; sternites brown, very nar-
rowly edged with paler on the caudal margin; pleurites broadly and
conspicuously yellowish. Hypopygium with the 9th tergite short, broadly
concave, yellow. Pleurites rather short, cylindrical, yellow, clothed
with long pale hairs, bearing at the tip two appendages; the dorsal
appendage is entirely fleshy with two arms, the one directed caudad, the
other cephalad, the caudal arm densely clothed with abundant pale
hairs, the cephalic arm with a chitinized bristle at the tip and about four
smaller bristles on either side, subequal in length and evenly spaced.
The ventral appendage is a long simple curved hook, very strongly
chitinized. The 9th sternite is very high, convex and bears at its tip
two strongly chitinized forked appendages that are directed caudad, the
outer fork being cylindrical, acute, the inner fork flattened, twisted and
directed entad. The penis-guard viewed from above (fig. 9) is narrow
_at the base, broadening toward the tip, the lateral edges chitinized and
242 Annals Entomological Society of America [Vol. VII,
passing into two sharp chitinized points; viewed from the side (fig. 10)
these sharp tips are directed strongly ventrad and viewed from beneath
(fig. 12) they are seen to be decussate. Gonapophyses short, directed
dorsad at the tip which is blunt and truncated. ,
Female: Similar to the male but the head of one specimen is
entirely dark, the dorsal brown stripe on the pleure clearer and narrower
not grading insensibly into the yellow of the dorsal pleurites.
Holotype, o', Nadi, Fiji Is., 7-28, 718. Allotype, 9, and
paratype, 9, with the type.
Gonomyia (Gonomyia) varipes, sp. n.
Head yellow with a brown vertical spot; thoracic dorsum brown;
legs banded brown and white; wings with the costal margin conspicu-
ously bright yellow.
Female, length, 4.6 mim.; wing, 3.6 mm.
Female: Rostrum and palpi brown. Antenne with the two basal -
segments light. yellow above, brown on the under surface; the two or
three basal flagellar segments are yellowish, the remainder brown.
Head light yellow, a narrow transverse brown mark across the front
behind the antennz and a V-shaped brown mark on the vertex with its
point directed cephalad.
Pronotum light brown except the scutellum which is very light
yellow, a continuation of the dorsal pleural stripe. Mesonotal prae-
scutum very dark clove-brown, uniform; scutum similar except the
median portion and the outer caudal angles of the lobes which are paler;
scutellum brown, the apical two-thirds pale; mesonotum light brown.
Pleurze and sterna brown except a broad yellow line extending from the
wing-root along the dorsal pleurites to the pronotum and a second
broad whitish yellow stripe extending from the fore coxe, above the
middle coxz to underneath the halteres. Halteres uniform light
sulphur yellow. Legs,—fore legs, coxe light yellow at the base, the tip
brown; trochanters brown; femora brown; tibiz, extreme base and
apical two-fifths brown, the remainder china-white; metatarsus with
the basal half white, remainder of the tarsi brown. One other leg is
loose in the vial and belongs to either the middle or hind legs,—here the
base of the femur is yellowish passing into brown at the tip; the tibiz
all white except the very narrow base and slightly broader apex which
are brown and the metatarsus is white except the tip which is broadly
brown; remaining tarsal segments brown. Wings, costa very con-
spicuously pale sulphur-yellow, remaining veins brown; wing-mem-
brane with a light brown suffusion; cell Ri paler and containing the
oval brown stigma. Venation as shown in figure 4.
Abdominal tergites and sternites dark brown, the pleural region
paler.
Holotype, 2, Nadi, Fiji Is., 7-28, ’13.
1914] Tipulidae from Fiji Islands. 243
Erioptera (Erioptera) oceanica, sp. n.
Halteres dark at tip; wings light brown; male hypopygium with the
pleura bearing a chitinized knob at tip.
Male, length, 6.3 mm.; wing, 5.4 mm.
Male: Rostrum and palpi brownish yellow. Antenne rather
long, the flagellar segments rather elongate-oval; if bent backward the
organ would extend beyond the. wing-base; scape brown, the flagellar
segments a little paler. Head dark brown and sparsely hairy.
Pronotum brown, clothed with brown hairs. Mesonotal praescu-
tum light brownish yellow without apparent stripes but with a row of
hairs on either side of the middle line; scutum, scutellum amd postno-
tum brownish yellow, the latter with a narrow brown median line.
Pleurze light brownish yellow. Halteres rather long, pale, the knob
dark. Legs pale yellow with the two apical tarsal segments brown.
Wings with a pale brown tinge, the costal region a little more yellowish;
veins brown. Venation as in figure 5.
Abdomen long and slender, pale yellow, the seventh sternite brown.
Hypopygium with the pleurites very long and slender, densely clothed
with long yellow hairs; at the tip of the pleurite are two appendages,
the one a dorsal chitinized appendage, slender at the base, swollen at the
tip and slightly roughened apically, and a ventral, flattened fleshy lobe
that is rather truncate at the tip. (See fig. 13).
Holotype, <&, Nadi, Eiji Is., 7-28, ’13.. Paratype, #, with
the type.
Mongoma fijiensis, sp. n.
Trentepohlii group; wings subhyaline, indistinctly if at all marked;
legs without white bands.
Male, length, 6.8 mm.; wing, 5.5-5.6 mm.
Female, length, 8-8.6 mm.; wing, 6.4-6.6 mm.
Male and female: Rostrum and palpi yellowish. Antenne with
the basal segments pale yellow, the flagellar segments brownish. Head
dark brown. Neck elongate, brown dorsally, yellow beneath. Meso-
notal praescutum light yellow with three elongate brown stripes, the
median one broadest in front, narrowed behind and ending at the trans-
verse suture; the lateral stripes are narrower, beginning just back of the
pseudosutural fovezee and continue back to the scutum where they
suffuse the lobes. Scutum yellow, except the central portions of the
lobes which are brown; scutellum and postnotum dark brown except a
narrow margin of yellowish. Pleurz light yellow, the sterna a little
suffused with brown. Halteres rather short, pale yellow. Legs, coxe
and trochanters pale yellow, femora, tibiz and the first tarsal segment
brown, the remainder of the legs broken off. Wings with a pale yellow
suffusion; veins light brown; stigma rather pale; indications of slightly
darker seams along the cord. Venation: (See figure 6). Fusion of
Ist A and Cty slight.
Abdominal tergites dark brown medially, this mark in the shape of
along triangle with its point directed cephalad; sternites pale yellow.
244 Annals Entomological Society of America _[Vol. VII,
Holotype, &, Nadi, Fiji Is., 7-28, 713.
Allotype, 9, and paratype, 2, with the type.
Mongoma, sp.
A species belonging to the fragillima and australasiae group
in the Macleay collection mentioned by Skuse (Dipt. Aust.,
vol. 4, series second; Proc. Linn. Soc. N. S. W., Sept. 25, 1889;
p: 832, 8aa5)
Conosia irrorata, Wiedemann.
Sixteen females taken at a lamp at Nadi, Fiji Islands, July 28, 1913.
This series shows a great difference in size in the different individuals.
It was previously recorded from these islands by Skuse who noted a
specimen in the Macleay collection. (l. c., p. 837, 838). The reason
that this entire series consisted of females is undoubtedly due to the
nocturnal oviposition in this species. Series of photophilous craneflies
always show a preponderance of the female sex and many of these are
gravid specimens ready to deposit their eggs, the others having laid the
clutch earlier in the evening. When males occur at lamps or in trap-
lanterns it is probable that copulation takes place in the twilight or
early evening.
EXPLANATION OF THE PLATES.
PLATE XXXIV.
Fig. 1. Wing of Dicranomia illingworth, sp n.
Fig. 2. Wing of Teucholabis fijiensis, sp. n.
Fig. 3. Wing of Gonomyia (Letponeura) fijiensis, sp. n.
Fig. 4. Wing of Gonomyia (Gonomyia) varipes, sp. n.
Fig. 5. Wing of Erioptera (Erioptera) oceanica, sp. n.
Fig. 6. Wing of Mongoma fijiensis, sp. n.
PLATE XXXV.
Fig. 7. Hypopygium of Dicranomyia illingworth; dorsal aspect of the pleurite.
Fig. 8. Hypopygium of Teucholabis fijiensis; dorsal aspect of the pleurite.
Fig. 9. Hypopygium of Gonomyia (Leiponeura) fijiensis; dorsal aspect. d—dor-
sal appendage; v—ventral appendage; p—penis-guard.
Fig. 10. Hypopygium of Gonomyia (Letponewa) fijvensis; lateral aspect. p—
penis-guard; pl—pleurite; s—9th sternite.
Fig. 11. Hypopygium of Gonomyia (Leiponeura) fijiensis; ventral aspect of the
9th sternite.
Fig. 12. Hypopygium of Gonomyia (Leiponeura) fijiensis; ventral aspect of the
penis-guard.
Fig. 18. Hypopygium of Erioptera (Erioptera) oceanica; pleurite, lateral aspect.
d—dorsal appendage; v—ventral appendage.
B.S. A.
Ss
C. P. Alexander.
ANNALS E. S. A. VoL. VII, PLATE XXXV.
CaP: Alexander.
A NEW SPECIES OF CHEILONEURUS WITH A KEY TO
THE DESCRIBED SPECIES FROM THE
UNITED STATES.
By A. B. GAHAN,
(Assistant, Cereal and Forage Insect Investigations, Bureau of Entomology.)
The new species of Cheiloneurus below described makes the
ninth species to be recorded from the United States. In order
to more easily distinguish this new form from those already
described a key to the species is included.
Superfamily Chalcidoidea.
Family Encyrtide.
Subfamily Encyrtine.
Key to the Described Species of Cheiloneurus from the
United States.
PROTA. RRR CARCS e s Ee ie EN AL. SP se POR SHEN Stans cha) vc whe useburar Redo etaars ie eb ohG sive Gc ioletelasre S 2
IVIL ON SINS OR, Renee duc hy de PRR uet oh crabayt nisi sedctg code aie Manet Wins ve teas 9
2. Wings hyaline; last two funicleGjourts: palew........,<4905 55 2 diaspidinarum How.
ERO NyI MES LLISCUESRS © Sete Sree crate ee 1 hes ees cieny Soe Ny canada ccs heads Sa: dpa ogres aetavees 3
Se PbiGshiEclet OMmilonsen thaw the pedtdels 5. 2.6. ..4. anes sad. oo eka tne ee 8
Hirst fyunicle joint equal to or shorter than the, pedicel. ...........5.0::.+::. 4
4. Body wholly pale testaceous, except the middle coxz more or less of the
mesosternum, the pedicel, first funicle joint, fifth and sixth funicle joints
and the clip witch abet browHishe . 22). . 2. ei we thts es ek swezeyi Ashm.
Thorax not wholly testaceous, with a part at least of the mesonotum metal-
EMO EMMA ALOER vara COT RUA ecto: techs syclus< debeacomtata ayeic etevete aul teviae eee ope 5
5. Antennal club nearly as long as the funicle and much enlarged; pedicel as long
as the two succeeding funicle joints. .7......:.......... dactylopir How.
Antennal club much shorter than the funicle and not much enlarged; pedicel
NoOtasioueasiuhe LWwOLOllOowinorruni Cle jOimtS.. a1. 0s. oe bs cis ne lee oe 6
6. Funicle joints all white; scrobes shallow and triangular... .albicornis How.
Funicle joints not white; scrobes deeply impressed and semi-circular........ 7
7. Scape flattened and somewhat expanded beneath, brown with a whitish
stripe from base to apex; pedicel and first funicle joint nearly equal;antennze
compressed; ocelli less than their own width from the eye margin....
See eC es ce ney ia raat eerste ra PeRAA uctin Nace Ruan lod arid lineascapus Gahan.
Scape slender, pale; pedicel distinctly longer than the first funicle joint;
antenne only slightly compressed; ocelli about their own width from the
ERE Lis ra eainlenir aly Attn ata SY Ox is SRI is tines Ga Poe Se eR cushmani Crawford
8. Whole funicle as well as the club strengly compressed, all funicle joints except
be first distimetlyswider thanvlongiisss2. 0.2.02 ac. 3 amplicornis n. sp.
First and second funicle joints subcylindrical, following joints and club not
as strongly compressed; all funicle joints except the last distinctly longer
GA TIMRWHCL Cesare ee SEN cee Ree Pea teebiine x afare¥, ihoiger hs cupreicollis Ashm.
Of, Scchralllibtrayesaosineivele <1. ee ota ee eae Jee Jn Oo ene eres dubius How.
SCR eUitoteah “VE Ve re Moet et Gres, Sea ee ei eet Sie eRe aa ear ooeenig Terr Pe ieee 10
10. Marginal vein very short, scarcely longer than thick......... dactylopui How.
Marginal vein much longer, three or four times as long as thick............ 11
11. Forewing with a distinct clouded area below the marginal vein; scape not
appreciably expanded beneath....................... cushmani Crawford
Forewing without a cloud; scape distinctly though not greatly expanded
A COLGAN at Ai Week oe Re os oN Cs. hes cae Sedat, Sachem: OE es lineascapus Gahan.
248 . Annals Entomological Society of America [Vol. VII,
Cheiloneurus amplicornis new species.
Female—Length 1.5 mm.. Head finely closely punctate, from in
front much lengthened, the transfacial line about half.the length of the
facial line, malar space long, scrobes short and very shallow, frons
narrow; ocelli in an acute angled triangle, the lateral ocelli scarcely
separated from the eye-margins; scape slender, pedicel distinctly shorter
than the first funicle joint; all funicle joints strongly compressed, the
first a little longer than its apical width, following joints much wider
than long; club compressed, not quite so long as the funicle and about
equal to the two preceding funicle joints in length; mesoscutum faintly
punctuate and closely covered with short pale pubescence; scutellum
and axilla minutely sculptured and opaque, the latter with a tuft of stiff
' bristles before the apex; propodeum polished; abdomen apparently
smooth above. Head reddish testaceous, eyes and antennal flagellum
black; scape testaceous; mesocutum black, more or less metallic; scutel-
lum and axille pale orange-yellow; tegule, pleurzee and most of the
abdomen reddish testaceous, propodeum and base of the abdomen
above polished metallic green; anterior wings fuscous, a narrow apical
border and the basal one-third hyaline; middle and anterior legs con-
colorous with the pleure, posterior legs dark brown, fore tarsi and the
apical joint of the median pair brownish as are the posterior tarsi.
Type locality—Dalhart, Texas.
Type—Cat. No. 18801, U. S. National Museum.
Five specimens from the type locality reared by C. N.
Ainslie from a coccid, Eriococcus sp., infesting Bouteloua and
recorded in the Bureau of Entomology under Webster No. 5571.
NOTE ON THE NUMBER OF SPIRACLES IN MATURE
CHALCID LARVZE.
By DANIEL G. Tower, M. Sc.,
Lafayette, Ind.
During a recent trip to Washington, D. C., while discussing
the life history of Prospaltella perniciost Tower a chalcid para-
sitic on the San Jose Scale, Aspidiotus perniciosus Comst., with
Mr. J. C. Crawford of the National Museum, he called my
attention to a translation of a Russian work published in 1912
entitled, Parasitic and Hyperparasitic Insects, by Iv Chewyreuv,
and in particular to a statement made by the author on page 16,
which is quoted in full: ‘‘In the same paper the author named
gave* (25, 35) a much enlarged drawing of Dibrachys bouch-
eanus Rtzb. This figure shows not a single spiracle, as if the
larva has not got them. While as a matter of fact it does have
them, and under the magnification it was drawn, they must be
quite evident; nothing is said about spiracles in the description
either. The fact is that the arrangement of spiracles in mature
chalcid larve is very peculiar and as will be shown later, makes
it possible to recognize them at once and to distinguish them
from the larve of allied families. They have nine pairs of
spiracles, two of which are on the meso- and metathorax and
the rest on the first seven abdominal segments; hence, there
are no spiracles on the prothorax and last two abdominal seg-
ments. This is the peculiarity Howard did not bring out in
his drawing which is therefore incorrect.”’
The statement made in the above quotation that all mature.
chalcid larve have nine pairs of spiracles does not hold true in
the case of Prospaltella perniciosi, for the adult larva of this
chalcid has only eight pairs of spiracles, two pairs of which are.
thoracic and six pairs abdominal.
In tracing the tracheal system of this scale parasite through
its two larval forms, one finds the tracheal system to consist in
the first larval stage of two longitudinal main trunks lying near-
the surface, one on either side, each bearing ten short, stub-like
*(25, 35) refers in the author’s bibliography to Dr. L. O. Howard’s paper, ‘‘A
Study res Gk Parasitism’’—U. S. Dept. Agri., Div. Ent. Techn. Ser. No. 5,.
p. 35, 1897.
249
250 Annals Entomological Society of America [Vol. VII,
branches. During the growth of this form the two longitudinal
main trunks join anteriorly and posteriorly, forming an oval.
Spiracles are not developed during this stage.
In the second larval stage the tracheal system is at first
similar to that of the mature first stage larva, except that it
lies deep within the body of the larva. As this larval form
grows the first, second and fourth to ninth inclusive short
branches of each longitudinal main trunk grow rapidly and
terminally at the surface of the body develop spiracles during
the last stages of this instar. The third and tenth branches
remain short and do not develop spiracles.
The above shows the manner in which the eight pairs of
spiracles originate, thus proving that the statement made by
Iv Chewyreuv does not hold true.
Pe =
scree sy
. = aes pi is on ee
s, ore Vitis Mer Wak sos - b= all \
“ heey ap eet ~ re t ren uy . te
ies wake bee . pe TS ¢ ~ ‘ 4 “3 Ss
ei Saas Sa te ' : . he ; }
> ‘a - ryt - ¥
Ser
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ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA,
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CONTENTS OF THIS NUMBER. |
PETRUNKEVITCH, ALEXANDER — Spiders Collected by
Mr. C. William Beebe in Burma and Borneo-. -.-
SEVERIN, Henry H. P., SEVERIN, Harry C., Hart- |
uNG, WitLIAM J.—The Ravages, Life History, °
‘Weights of Stages, Natural Enemies and Methods |
of Control of the Melon Fly paeye Cucurbitae
HG Re epee aha ae Be eeepc iad hk cow SET AGE 4
PALMER, Miriam A. Vere Notes on the Life History,
Obviyady Beetles) aie ee eh ees
ALEXANDER, CHARLES Paut—On a Conlestien: of
Crane Flies (T SAH, Diptera) from the Fiji
Tslands #.3%).35° SEY, Oya MIA! air MSM LEGS Ome ie samy oda
V77
GAHAN, A, B. —A New Sictas, of Ghodoand a: with a
Key to the Described Species from the United
ise 11 as gue a SM aM Ey NARs AUTO a ge Oh
Tower, Daniet G.—Note on the Nuniber ‘of Paitacis
in Mature Chalcid saith PVaREN Me Nr ah ce ee
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Proceedings of first three meetings; Constitution, By-Laws and List of
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OsBorN, HERBERT—The Habits of Insects as a Factor in Classification. . '.20
SEVERIN, H. H. anp Severin, H. C.—Anatomical and: Histological Siadies
of the Female Reproductive Organs of the American-Saw Fly, eae
AMEHIGANA; LEAGH se55 NIG Ae os ONE SO ER A epee eR Boa 25
: ‘Fett, E. P.—Some Problems in Nomenclature......0cc0.0cececsevees: PRON a» 10
Hammar, A. G.—On the Nervous System of the Larva of Corydalis cornuta L. .25
‘Brapiey, J. C.—A case of Gregarious Sleeping Habits among Aculeate
FFVDICNOPVETAN Sac SP The Va iiice ee eA Lek EME later ne Malaiwe deere 10
Davis, J. J—Notes on the Life History of the Leafy Dimorph of the Box-
elder Aphid, Chaitophorus negundinis Thos... 0.2... cca ee ee bees -10
HAMBLETON, J. C.—The Genus Corizus, with a Review of the North and é
hMiddte: American jSpecres . oid SeSGAies EA Wicd wea Uwths an Lin Oneal oe aan ~ 25
Grrautt, A: A.—Biological Notes on Colorado Potato Beetle:..... Hy eee -25
» Grravtt, A. A.—A Monographic Catalogue of the Mymarid Genus Alaptus... .25
SEVERIN, H. H. and Severin, H: C.—Internal Organs of Reproduction of ;
JE CSS A GS Aen Re Ron mt i Per A aN RAD RUBLE Mn AR e tg IO Ia Nags fr IO we MAREE ST 15
SmitH, C..P.—A Preliminary Siuly of the Arane Theraphose of California.. .765
Davis, Ji J.—Studies on’ Aphididas) yoy eat Pcie ohne Cee NU ee eee OD! 7 3
“Rie: W. A.—Muscle Attachment of Tnsentey LaRue nea peas da Cad so phe peer 15
NEEDHAM, J. .C.—Critical Notes on the Classification of, the Corduliing :
(donate et Toh 2 os ah heleieg nh dinly Mah dice nts KO's pel RE pa Sey ag Bes tis
Howarp, L. O.—A Key to the Species of Prospaltella with Table of Hosts
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Hoop, J. D.—T wo New Species of Idolothrips..........2..... op ett Pr rene |
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ANNALS ENTOMOLOGICAL SOCIETY OF AMERICA,
State University, Columbus, Ohio.
ANNALS
OF
The Entomological Society of America
Volume VII DECEMBER, 1914 Number 4
THE BIOLOGY OF THE NET-SPINNING TRICHOPTERA
OF CASCADILLA CREEK.*
Miss AticE Ayr NOYEs.
CONTENTS.
PAGE
TREE YGIGEIORD fh at eel” S SP ge pa cane cae oe SER Te gee CEI a es 251
II. Catching-nets of the Families Hydropsychide, Philopotamide,
Polycentropide.
feria am btn OOM Sy, Cll Gl ao nm se tereman i aM Sata oak ocd blac oe 252
ipa yaa SONI PARC ES Goto cts ohn ables tse Seas W sates be Dae eo 259
Pammhyge Oly CenpLOmidass aes eos cece A victen Seed be Parthia tree env 260
Meee eM NG chi Ce OLUCa LOM spepan tet sn?2\s oo eet Mew oc byte ale ys se caw bets 265
EMC en a RO Le Mae Re tN ee Se ele wie a als bone arls a nds #4 O98 Lvs we 266
Venvexperimental work onklydropsyche Nets:..c.. 0... 2.2. ees te cee 268
WIL, Lull Eroyesreeyoy ny75 Sees yes lice tere nenin CDs eR etre 271
I. INTRODUCTION.
The net-spinning Trichoptera are confined to the members
of the old Family Hydropsychide, which has been subdivided
(Ulmer 1909) into four families, Hydropsychide, Philopota-
midz, Polycentropide, and Psychomyide. Nets of some of the
genera of the first three families have been described, but
- as far as is known, larvae of the Psychomyide spin no catching-
nets. (Wesenberg-Lund, 1911). The only nets described in
this country up to this time have been of the genus Hydropsyche.
Most of the work on the catching-nets has been done within the
last six years and almost entirely by Danish investigators.
Contribution from the Limnological Laboratory of Cornell University.
251
252 Annals Entomological Society of America _[Vol. VII,
This paper is a preliminary study of the net-spinning
Trichoptera of Cascadilla Creek with special reference to the
nets—the method of construction, their efficiency as a plancton-
catching apparatus, and the closely related problems of food
and feeding habits.
Since the net-spinning caddis-worms are found in still as
well as swift flowing water, and their nets are such interesting
and beautiful structures, it seems strange that they should have
been overlooked for so long a time. I have found nets repre-
senting some of the genera of the families Hydropsychide,
Philopotamide, and Polycentropide, and will treat them by
families in that order. In each family I have first given
extracts or a brief summary of the work published on the
nets up to this time and have then added my own observations.
This work was carried on under the supervision of Prof.
James G. Needham, to whom I am especially grateful for
his valuable suggestions and encouragement at all times.
II. CATCHING-NETS OF THE FAMILIES HYDROPSYCHIDAE,
PHILOPOTAMIDAE, POLYCENTROPIDAE.
FAMILY HYDROPSYCHID.
The first description of a catching-net is found in Dr.
F. Muller’s work (1881). He describes and figures the net of
one of the Hydropsychidz, a southern Brazilian sp. of the
Genus Rhyacophylax. He always found the houses on the
upper side of stones, made of irregularly interwoven plant fibres
or of small stones. Each house has a funnel-shaped vestibule
or verandah, whose sidewalls are generally constructed out of
interwoven fibres. These serve as a covering for a very delicate
silken net with square meshes, generally from 0.2 to 0.38 mm. in
diameter. The entrance to the vestibule is always directed
up stream, so that the water coursing through it catches and
holds back organisms which serve as food for the caddis-worm.
The larvae rarely live alone, but generally construct their
houses very close to one another so that sometimes continuous
rows of them are formed.
In the year following, the first work on the nets: of the
Genus Hydropsyche appeared in this country in an article by
Miss Cora Clarke. In a later paper, 1891, she mentions these
larvae and their nets again, but gives no additional data.
1914] Biology of Net-Spinning Trichoptera 258
To quote from the first article: ‘“‘The typical form of the
structure resembles a tunnel attached to the surface of a stone,
having at its mouth a vertical framework with a net stretched
across it. An open mouth or entrance to the case is always
close to this net on the side towards the current, so that without
wholly leaving its house the larva can remove from the net
anything eatable, which the current may have lodged there.
The mode of building varies considerably. The case is usually
about half an inch long and a little curved, loosely attached
to the stone by its edges and without any bottom. It may be
composed entirely of sand or of bits of plants or both combined.
The supporting framework of the net is always formed of
vegetable bits, and is sometimes a simple arch, sometimes a
complete ring, and sometimes a short cylinder. It is occasion-
ally stayed or held in position by silken cords stretching from
it to suitable points on the stone. It is stiff enough to stand
erect even when removed from the water. When it is in the
shape of a cylinder or broad arch the net is always stretched
across that end of it which is down stream and the entrance
usually opens under the shelter of the arch. * = = if
In a stream in Brookline, Mass., are large communities of these
larvae. The stones in the stream are covered with mud,
leaves and rubbish.
Sometimes a stick which has fallen into the brook has a
row of cases and nets built upon it. Often a stone will have
a row of them side by side along one edge, or there may be
only a few of these structures scattered separately upon its
surface.’’ She mentions having received a net and larva
from Mt. Desert, Me.
In 1886 L. O. Howard found similar nets of a Hydropsyche
larva on the Simulium-covered rocks in the swift water of
Rock Creek, near Washington. ‘‘The cases varied greatly
in size, the mouth of the funnel in some instances not more
than 3 mm. in diameter and in others reached fully 10 mm.
The tube of the funnel was in every case bent at nearly right
angles with the mouth and the larva ensconced within it
waited for its prey to be caught in the broadened mouth.
The broad funnel-shaped expansion was woven in fine
meshes with exceedingly strong silk and was supported at
the sides and top by bits of twigs and small portions of stems
of water plants. The central portion was so open as to allow
the water to pass through readily.”’
254 Annals Entomological Society of America [Vol. VII,
In 1888 the same author mentions finding the larvae in a
similar situation in Ithaca, the nets and cases being very
abundant on the Simulium-covered rocks. « ‘‘The nets differed
from those found at Washington and the species is probably
different.’’
In his Insect Book (1901) we find the following: ‘The
cases were preferably placed at the edge of slight depressions
in the rocky surface so that the tubular portion was protected
from the full force of the current. On the surface of a rock
about 18 inches in diameter 166 of these nets were counted.”’
Adele M. Fielde (1887) writes from Swatow, China, of a
net similar to those previously described. ‘During last
January I found on the level surface of the coarse sand which
covers the bottom of an aqueduct near here, under an inch or
two of clear running water, little structures resembling a
gray net spread to catch fish or a tiny cave with a gauze awning
stretched over the entrance. The arches had a span of from
an eighth to half an inch and always opened towards the
current. They were to be seen in scores with a buttress of
coarse sand in the rear, and a minute aperture in the floor.
The occupant of the wee grotto was in every case a caterpillar
not more than five-eighths of an inch long. It burrowed in
the sand of the floor, stretched its head forth vertically, and
fed upon what had been caught in the delicate roof of its den.”’
Comstock, J. H., (1895) in speaking of Hydropsyche larvae
says: ‘‘Stretched between two stones near by can be seen his
net. Thisis made of silk. It is usually funnel-shaped, opening
up stream; and in the center of it there is a portion composed
of threads of silk extending in two directions at right angles
to each other, so as to form meshes of surprising regularity.
These nets occur in rapids between stones, but in many places
they are to be found in greater numbers along the brinks of
falls. Here they are built upon the surface of the rock, in the
form of semi-elliptical cups, which are kept distended by the
current. Much of the coating of dirt with which these rocks
are clothed in summer is due to its being caught in these nets.”
Betten, C. (1901), says of a Hydropsyche sp. (near phalerata,
Hagen), that there was ‘‘no larval case, only strands of silk
between the rocks.”
or
1914} Biology of Net-Spinning Trichoptera 25
Comstock, Anna B., (1903), in describing the snare of
Hydropsyche writes, “‘It is formed like a dip-net and fastened
with silk to a frame of leaves or pebbles, so that its distended
mouth is directed up stream. Near the frame it consists of
fragments of vegetation woven into a silken tapestry and
is finished at the end with a bag of coarse, even mesh. The
regularity of this bit of netting is beautiful to behold, and
its use shows the cleverness of the builder. This large mesh
allows the water to flow through freely, and thereby leave
entangled in the seine any little creature not small enough to
pass through. * * On the side of this tiny seine
toward the current of the stream is a little passage which leads
to the seine-builder’s house.”’
The work on the net-spinning Trichoptera was next taken
up in Europe and it is to the Danish investigators that we
owe our most extended knowledge of the various kinds of nets,
and whose work stimulates a desire to carry their efforts
farther. E. Petersen, in 1908, found the catching nets of
Hydropsyche instabilis in a brook north of Silkeborg (Den-
mark). The larger stones were completely covered with
Potamogeton pectinatus, Fontinalis antipyretica and Junger-
mannia sp., and on them the trumpet-shaped catching nets
were placed in rows and connected with one another. The
nets were small, being only 8-10 mm. in diameter at the mouth,
and their depth about 7 mm. The nets were always supported
by the plants and parts of these were often wovenin. In many
cases one net was placed a little behind the others and con-
nected with them by a strong web. At the base of the net
lurked the larva.
In 1909 Ussing described a catching-net of H. instabilis
that he had found in Hornbek brook in the vicinity of Randers.
Being unable to obtain this paper, I have translated an extract
from it, which was quoted in Wesenburg-Lund’s (1911) article.
“The nets are placed obliquely in front of the opening of the
larva’s tunnel, built of very fine, square meshes (0.2 mm.
in diameter), propped up by bits of plants. The dwelling of
the larva is built out of mud and half decayed fragments of
plants; the tube is spun fast to a stone on the bottom of the
brook. I have often found whole rows of these dwellings
with nets placed between the separate occupants. The nets
turn their expanse against the stream, which is always very
256 Annals Entomological Society of America __[Vol. VII,
swift and in spite of their delicate construction, they stand
the considerable pressure of the water very well. I have
never noticed that the dwellings or nets protruded above
water.”’
One of the most interesting of the descriptions of Hydro-
psyche catching-nets comes from Dr. Wesenberg-Lund. In
his paper (1911) he has compiled the records of all known
cases of net-spinning and extended our knowledge greatly by
personal observations. He studied the nets of H. pellucidula
and H. angustipennis, and gives a very full description of the
beautiful structures of the latter which he studied in July,
1909, in the outlet of Foenstrup pond in Gripwalde. The
larvae had utilized the leaves of Lemna triscula in the con-
struction of their dwellings, and chains of these, arranged in
rows, were placed obliquely across the stream. Every chain
was composed of the dwellings of a number of larvae. Each
house had a funnel-like entrance facing up stream which led
into a vestibule about 1-2 cm. long and of the same height,
covered with Lemna leaves. In the farther corner was the ©
entrance to the larval dwelling which was 2-3 cm. long. This
is always laid obliquely to the principal course of the chain,
and was made of small bits of decayed wood and pebbles
interwoven in the silken mesh and covered with Lemna leaves.
In the wall of the vestibule towards the entrance to the dwelling
was a circular window about 1 cm. in diameter covered by a
beautiful screen. This served as the larva’s catching-net
and was woven of strong threads crossing nearly at right
angles and of wonderful regularity in the centre, but irregular
and of a coarser mesh toward the framework, which was made
of small pieces of straw finely fastened together. The cases
were submerged in the water, but the upper part of the vesti-
bule and window projected over the surface of the water. I
have copied a diagramatic figure of a Hydropsyche house
from his paper. (See Plate XXXVII, Fig. 1).
In regard to the seasons when the nets are found he gives
the following data: ‘‘Up to December, beautiful larval dwell-
ings and nets are found; from December to the last of April
no nets observed. During this period the larvae were found
rolled up under stones or in crevices in boards, probably taking
only a little food. At the beginning of May and during the
whole of June the nets were put up again.”’
1914} Biology of Net-Spinning Trichoptera 2a
His observations agree completely with those sent him by
Ussing, who made regular observations on the nets of H.
instabilis during the winter of 1909-10 at ‘“‘Hornbek brook.”
“On the 24th of October, 1909, the nets were very numerous; on
November 7, beautiful catching nets; on January 2, and Jan-
uary 19, 1910, none. The animals build no catching-nets in
winter. The larvae lie rolled up in a spiral and are not active,
moving reluctantly. They do not live in the usual case, but
in an irregular net with small pebbles interwoven. He believes
that the Hydropsyche larvae lie in a dormant condition and
take no food in winter.
My observations were started the latter part of October,
1912, and at this time, although the nets were numerous every-
where in the creek, they were inconspicuous, owing to a thick
coating of diatom ooze and silt, and they were badly torn
by the large numbers of fallen leaves swept along by the cur-
rent. Only rarely during November did I find a perfect net,
and during the winter months no nets at all. Heavy rains
the last week in March made any observations impossible, as
the turbid water rushed along in torrents. On my next visit,
however, on April 12, it was as if the stones had been touched
by some magic wand, for nets had sprung up everywhere.
They were found on the upper surface of stones or shelving
rocks wherever there were irregularities or crevices against
which the cases might be built; on submerged twigs, on the
underside of stones, and between stones on the bottom. The
nets were also thick along the edges of the stream, many dis-
tended pockets being found in the tangle of roots which floated
out int@sthe current. In July similar nets were observed in the
mats of Cladophora, but these were generally the tiny pockets
of very young larvae. I could find no definite dwelling tube
in either of these instances, but the larvae were found crawling
among the roots or algal filaments.
On the brinks of the waterfalls were rows of vertically
placed nets, so that a continuous stream of water was pouring
down their open mouths. On the creek bottom the nets were
generally fastened between two stones, some being of the
‘“‘dip-net’’ type, while others formed a horizontal net. In
both instances, however, the net was composed of coarse, ir-
regular mesh at its entrance and a fine regular mesh behind.
258 Annals Entomological Society of America [Vol. VII,
Although there are six species of Hydropsyche larve com-
mon in Cascadilla Creek, I have not been able to find any
specific differences in their nets, so will describe them col-
lectively. The case in which the larva lives, I found as de-
scribed by others, except Miss Clarke, to be made of vegetable
bits, pebbles, or a combination of both woven into an irregular
cylindrical tube. In front of this, opening toward the current is a
net. Mrs. Comstock’s word ‘‘dip-net’’ best describes its shape.
Beginning at the entrance and generally extending for a little
more than one half the depth of the net is a very irregular,
coarse silken mesh, the bottom of the net being composed of a
wonderfully beautiful, regular mesh. This latter is the catch-
ing surface proper from which the larva feeds. The tube in
which it lives extends a very short distance into the net, so that
its entrance opens under the fine mesh. The tube opens into
the net either from the right or left side, and is found either
extending back in a straight line with the net or almost at right
angles to it. When the stones are taken from the water, some
of the nets stand upright owing to the supports of plant tissue
woven into the coarse mesh. Sometimes there is a complete
supporting arch, but often there is only an oblique prop on
either side, anchored to the stone by silken guy lines. The
threads of the catching surface are somewhat distensible, and
when seen in the water it is concave, but when removed, it
appears as a flat, almost circular disc in its supporting frame-
work. In many cases, however, the nets collapse completely
when there is no current to distend them, there being no sup-
porting bits of any description. See Plate XXXVII, fig. 2.
In summer many of the nets have long green streamers of
Cladophora filaments, which have become entangled in the
nets and float back several inches behind them.
The average expanse of the nets at the entrance is about
8 mm. although some of the largest ones have an expanse of
20 mm., with a depth of 15 mm., while those of the very young
larve have an expanse of 144 to 2 mm. and a depth of 1 mm.
These nets and dwellings I have always found completely
submerged, and the true catching surface placed at the end of
the vestibule, instead of in its sidewall as in that of H. angusti-
pennis described by Wesenberg-Lund (1911). The threads
are very firm so that they may withstand the force of the
current and there is no difficulty in seeing the meshwork with
the naked eye.
1914] Biology of Net-Spinning Trichoptera 259
FAMILY PHILOPOTAMIDZ.
The only descriptions of catching nets of this family are
those of Thienemann. He gave a brief account of a net of
Philopotamus ludificatus in 1906; as I was unable to obtain this
paper, however, I will summarize a fuller description which
appeared in 1908.
Two similar species P. ludificatus McL., and P. montanus
Don., are found in great numbers in the swift mountain brooks.
of Middle Europe. These build dwellings which are very
much alike. The house is a broad sac-like structure of loose
mesh about as long as one’s finger. At the front end where the
opening is found, it is fastened to a stone on the bottom of the
brook. The blind end of the sac floats freely; and in the bot-
tom of it is found the larva which can feed on organic particles
caught in the net. Occasionally the larve also stretch their
houses between two neighboring stones and so construct for
themselves, in this way, a kind of catching-net. Only one
larva is found in each net.
No descriptions have appeared before of nets of the Genus.
Chimarrha. The nets of Chimarrha aterrima, which I found,
are long, narrow pockets built entirely of a very fine mesh of
delicate silken threads. (See Plate XX XVIII, Fig. 1). The
average size of the net of the growing larva is about 25 mm.
long and 3 mm. wide. The nets are rarely found singly, but
generally placed five or six in a row. Sometimes the front
edges of these are joined together, but in most cases each net
is entirely separate from that of its neighbors. There is a
large opening at the end facing the current, and a tiny opening
at the hinder end just large enough for the larva to slip through
and make its escape when alarmed. This opening is very hard
to see, not only because of its size, but owing to the fact that
the nets are generally brown with a coating of diatoms, etc.,
over much of their surface. The nets are fastened at the en-
trance by their entire lower edge, the rest of the sac floating
freely, and kept distended by the current. They are found
fastened to the underside of stones or to their upper surface
when they are covered by other stones. I have also exposed
them on the upper surface of the shelving rocks by pushing
aside the covering mats of Cladophora. The orange or yel-
lowish larva, of which there is only one to a sac, is usually seen
260 Annals Entomological Society of America [Vol. VII,
toward the hinder part of the net. It does crawl around,
however, feeding over the whole surface of the net. It does
not use its front legs to assist it in getting its food which is
entirely of microscopic plants. All observations must be
carried on in swift water, for the net collapses into a brown
slimy mass when the pressure of the current is removed. The
separate threads of the net are only clearly seen with the highest
power of the microscope when it is seen that the units of the
mesh are rectangular in shape, one dimension being about
eight times the other. The double nature of the silken threads
is not recognizable, as is that of the Hydropsyche’s, when ex-
amined with a microscope.
At times grains of sand and small pebbles are found on and
about the large nets. I believe this to be a preparation for
pupation, as the pupal cases are constructed of these.
FAMILY POLYCENTROPID2.
The nets of five genera in this family have been observed
and described.
Neureclipsts.
In the year 1900 Wesenberg-Lund first noticed the plancton-
catching-nets of Neureclipsis bimaculata in Western Jutland.
Later he also found them at three different places in Zealand.
They were not described, however, until 1907 when E. Petersen
wrote an account of them. His observations were extended
by Wesenberg-Lund (1911). The nets of this larva are trumpet-
shaped, from 69 to 90 mm. long; the expanded mouth is 25 to 35
mm. broad, and the hinder end about 10 mm. In some cases
the hinder end of the tube is attached to some object, in other
cases it floats freely. The nets show a regular variation in
color due to the plancton caught in their meshes; in the spring
they are brown or grayish from diatoms, in the summer bluish-
green from the Cyanophycee. The net is kept distended by the
force of the current and collapses into an unrecognizable mass
when taken from the water. The plancton-organisms Bos-
mina, Daphnia and the remains of Cyanophycez become caught
in its walls as the water filters through and serve as food for
the larva which is generally at the hinder end of the net. Many
thousands of these nets span the stream in Hennebach so that
a greater part of the water filters through them.
1914] Biology of Net-Spinning Trichoptera 261
These writers believe that the imago probably lays her eggs
in loose, web-like masses which are doubtless a conglomeration
of old nets and that the young larve live together in them for a
long time.
Plectrocnemia.
Miall (1895) gives a description of a Plectrocnemia net
written by Mr. T. H. Taylor. ‘‘Plectrocnemia finds its home
in streams where the water flows swiftly over a stony bed. Ifa
stone be lifted out, the under side is often found to be covered
with patches of mud from which brown larve emerge and begin
to crawl over the surface. The muddy particles are evidently
held together by some binding substance, and the whole forms
the retreat of the Caddis-worms, corresponding to the cases of
Phryganea. When a larva is placed in a vessel of clear water,
it at once begins to explore its new quarters, and eventually
selects a site for its dwelling. This is made of silken threads
secreted by the large silk glands, and when completed the
structure consists of a tube considerably longer and broader
than its occupant and open at both ends. It is supported and
strengthened by a meshwork of silken threads, which spread
out for a considerable distance, and are attached to the sur-
rounding objects.
From time to time the larva turns round in its case and
even leaves it for a short space. Generally, however, it re-
mains quiet inside, apparently on the alert for prey. If a.
Chironomus or other small aquatic larva approaches, it is
almost certain to get entangled in the network of silken threads.
At once the Caddis-worm in its retreat perceives the presence
of a possible victim. The long hairs which cover the body are
possibly tactile, and receive slight disturbances of the silken
network. The Plectrocnemia then proceeds warily to de-
termine the cause of the disturbance. Should the Chironomus
be entangled near the middle of the tube, the Caddis-worm does
not hesitate to bite its way through the side, and its jaws very
soon quiet the struggles of the prey.
There is some resemblance between the snare of the Plec-
trocnemia and the web of a spider, but the Plectrocnemia is
effectually concealed by the mud which clings to its retreat.”’
The net of Plectrocnemia conspersa Curt. is described by
Wesenberg-Lund (1911). The larve—at least from April until
262 Annals Entomological Society of America [Vol. VII,
June—build large, flat catching-nets about a square decimeter
in size. In the centre is an opening, (about 8 to 10 mm.,) which
leads into a funnel about 5 to 6 cm. long—the hiding place of
the larva. This is hidden under a stone or leaf. The mesh is
very coarse on the outer edges of the net. The water being
very shallow in the brook, the nets lie nearly horizontally on the
stones. The organisms caught in the nets by the larve are
principally gnat-larve, Asellide, etc., which are swept along
by the stream into the net.
Polycentropus.
The net of Polycentropus flavomaculatus Pict. was first
described and figured by Petersen (1907). The nets resemble
swallows’ nests and are about 30 mm. long, 20 mm. wide at the
entrance, and about 15 mm. high. They are found singly on
the bottom of slowly flowing brooks attached by their fore
corners to small stones. The mouths face the current and are
held expanded by the water. The larve are always found in
the bottom of the net. When found on vertical banks, the
mouths stand perpendicularly. The nets are also found on
the wave beaten shores of the larger lakes. The nets are
bluish-green in color.
Holocentropus.
The larve of the genera Holocentropus and Cyrnus live in
quiet water, principally among alge and water plants of the
smaller lakes and pools. The nets, which are hard to observe,
have been completely overlooked before Wesenberg-Lund’s
paper (1911). He first saw the net in June 1909.
He figures three forms of nets which he found made by
Holocentropus dubius. One type of net is in the form of a
shallow funnel attached by silken threads to Sium leaves. In
the centre is an opening which lands into a thick web-like
tube which extends to the main stem of the plant. In this
passage the larve live and may escape through an opening at
the hinder end. The second type is found where there are thick
mats of filamentous alge, as Spirogyra. In this loose mass
may be seen perpendicular tunnels 8 to 10 cm. long, covered
with spinning; these are open below and also open at the sur-
face in the middle of a shallow funnel-like net of very fine
mesh. The larve sit at the bottom of the funnel-like recess
1914] Biology of Net-Spinning Trichoptera 263
watching for prey. There is often at least one side passage
branching off from the main tunnel. Another type is a funnel-
like net spun between the angles of the main stalk of the grasses
and the side shoots, and fastened at the upper end to leaves as
Potamogetons.
On account of lack of light the study of the nets of Holo-
centropus picicornis Steph. was unsatisfactory. The deep
brown color of the nets was due to a thick deposit of iron
bacteria.
The larva of Cyrnus flavidus McLach. lives in lakes in
the Chara-and Elodea-zone at a depth of about 4m. In summer
the larva spins funnel-shaped nets to the leaves of Potamogeton
lucens when it reaches the surface. In the autumn long threads
emerging from the plants are seen floating about in every
direction. Plancton organisms become entangled in the threads
and the larva runs along these and siezes its prey. In October
and November the larve sink down with the Potamogeton on
to the Elodea and Chara zone again.
In a little arm of Cascadilla Creek (See Plate XXXVI,
Fig. 1) where the water is rather quiet and from 11% to 2 feet
deep, larve of two genera are found belonging to this family.
The larva of Cyrnus pallidus (?) is small—8 mm. long by 1.2
mm. broad—very rapid in its movements. The body is whitish,
dorso-ventrally flattened; the head yellowish with a large
brown spot covering almost the whole dorsal surface. In the
centre of this spot is a yellow cross-shaped figure and eleven
yellow dots around the margin. The yellowish pronotum is
brown posteriorly with yellow dots. On removing a stone
from the water the dwellings of this larva might easily be
overlooked, for they resemble patches of sediment clogging
the crevices. If placed in a pan of water, however, and ex-
amined under the microscope, they prove most facinating
objects for study. Stretched across crevices in the stones,
preferably along its edges, but also occasionally on the upper
and lower surfaces is the roof of the larval dwelling. The tube
of a full grown larva, is about 9 mm. long by 3 mm. broad and
is dorsoventrally flattened. (See Plate XXXVIII, Fig. 2).
It is spun of fine silken threads so closely woven that it has a
felty texture. It is always brown with a coating of diatoms.
At either end a little flap hangs from the roof which acts as a
264 Annals Entomological Society of America [Vol. VII,
stopper, closing against the opening when the stone is removed
from the water. Radiating in all directions from the floor
of its retreat, at either end, may be seen threads of silk about
7 mm. long. These are fastened to the stone at their outer
ends and a microscope reveals the fact that they are connected
with one another by a loose irregular mesh which floats up from
the surface of the stones and entangles many small organisms.
The larva lurks in its little cave, and welcomes visitors gladly
at its front or back door. Any movement on the silken strands.
in front of its doors causes it to dart out the front part of its.
body with lightning-like rapidity, sieze the intruder and draw
back again, all in the twinkling of an eye. Large numbers of
Vorticella and other Ciliates, rotifers, Chaetonotus, Chirono-
mids and diatoms were found entangled in the meshwork.
The larva of the Polycentropus sp. (?) is large and more
deliberate in its movements. It is 19 mm. long and 2 mm.
broad; the head and prothorax yellowish-brown with many
small, brown dots, and the abdomen of a pinkish-lavender color,
iridescent when the sunlight strikes it. It sometimes looks.
bluish. The larva lives on the under side of stones in a deli-
cate silken dwelling which falls together into an unrecognizable,
brown slimy mass when removed from the water. It was not
until I had examined a large number of these nets that I was.
able to detect a trace of any definite form. The larva lives in a
very delicate, silken tube fastened to the stones along its whole
undersurface. It is shaped like a flattened cylinder and slightly
curved. (See Plate XXXVIII, Fig. 3). The tube is 21 mm.
long and 5144 mm. wide with an expanded opening at either end.
Connected with each opening and along either side is a mass of
tangled, silken threads, about 20 mm. square and loosely
attached to the stone. This tangled mass may float partially
over the tube and so obscure it.
I have never observed the larva feeding but do not doubt
that Mayfly nymphs and Chironomid larve become entangled
in the meshes as they crawl about over the stones, for remains.
of these forms are abundant in the stomach contents.
1914] Biology of Net-Spinning Trichoptera 265.
III. THE AQUATIC SITUATION.
All of my collecting and observations on the net-spinning
Trichoptera were confined to a very limited area in Casca-
dilla Creek, not exceeding a half mile in extent. For a pre-
liminary study this presented advantages, one of the most
important being an abundance of material within a few min-
utes walk from the laboratory. This made it possible to ob-
serve conditions frequently and to spend more time in the field
than would have been possible had the Creek been at a distance.
The use of the Fish Hatchery, situated on the bank of the Creek,
also offered opportunities for studying things to the best ad-
vantage, for all necessary equipment as microscopes, instru-
ments and glassware could be kept there. It also furnished a
place where rearing and experimental work might be carried
on, undisturbed and under natural conditions.
The depth of the Creek varies from a few inches, where it
spreads over the large, flat rocks, to two and a half feet or
more in the middle of the stream. The creek-bed averages.
from ten to fifteen feet in width but broadens out in places to
thirty feet or more, where the larve abound, the bottom is rocky
and of two types—loose stones, both large and small, (See Plate
XXXVI, Photo 1), and continuous shelving rocks with gradual
descents of a few inches to steep descents of five feet or more.
(See Plate XXXVI, Photo 2). In early spring and fall the
water rushes along in torrents over the rocks, but by midsum-
mer the swift water is confined to the middle of the creek-bed.
Large areas of the broad, shelving rocks remain dry and where
there is water it does not exceed an inch in depth.
Most of the typical swift-water forms of insect nymphs.
and larve were found associated with the Hydropsychids.
Of the Trichoptera, Rhyacophila, Helicopsyche, Silo, and a
Hydroptilid sp.; of the Diptera, Simulium, Chironomid and
Blepharocera larve were very abundant on the upper surface
of the stones; of the Mayflies and Stoneflies, the nymphs of
Heptagenia, Chirotonetes and Neoperla were found in numbers.
on the under side of the stones. The rocks presented various
colors—the browns of diatom ooze, large black patches of Simu-
ium larve, and in places thick green carpets of Cladophora.
The swift water and great abundance of food made it an ideal.
situation for the larve.
266 Annals Entomological Society of America [Vol. VII,
IV: Foon:’:
In most of the literature one finds the larve of the old
family Hydropsychidze spoken of as carnivorous, but Siltala
(1907) gives the following general statements. ‘The larve of the
true Hydropsychide are less exclusively carnivorous than
those of the other campodeoid larve. Both animal and
vegetable food are found, remains of insects, Crustacea, algal
filaments, pieces of moss and phanerogam leaves, also pollen
grains of Conifers.’”’ In an earlier paper (1910) he speaks of
their ability to utilize hard vegetable stuff, gnawing grooves
nearly 8 cm. deep in the logs of a bridge.
“The data were insufficient in the case of the Family Phil-
opotamide to form any judgment. The Polycentropide are
purely carnivorous, eating insects, Cladocera and Ostracods.”
He also points out that a relation exists between the struc-
ture of the mandibles and the kind of food. He extends
Ulmer’s (1902) observation that forms with blunt-toothed
mandibles are herbivorous and those provided with sharp teeth
are carnivorous, and points out the importance of the presence
or absence on the mandibles of a median tuft of hairs. All
forms with the median tuft on both mandibles are herbivorous;
those lacking it are either exclusively carnivorous or at least
eat as much animal as vegetable food; larve with the tuft
only on the left mandible vary in respect to their food and among
them are found carnivorous, herbivorous and omnivorous forms.
My results in regard to the food of the larve are based
entirely on observations upon freshly killed animals taken from
their natural habitat. The alimentary canal was removed
immediately after the collecting trip and examined at once, or
placed in four per cent formalin for later study.
Collections of Hydropsyche larvae were made on Novem-
ber 14, 1912, November 21, November 30, January 31, 1913,
February 18, March 20, April 12, May 6, June 2 and July 7. ©
As many as five specimens were always examined, and in some
cases as many as ten. The food as stated by Siltala was made
up of both animal and vegetable matter. There was, however, a
seasonal difference; in the fall and winter diatoms formed the
bulk of the food, and in the spring and summer animal food
predominated; while at all times algal filaments were present
in moderate amount.
1914] Biology of Net-Spinning Trichoptera 267
Of the diatoms, Gomphonema, Cocconema and Navicula
were the most abundant forms, though Synedra, Melosira,
Encyonema and Fragillaria appeared in smaller numbers.
Ulothrix and Oedogonium and Cladophora of the green
alge were found all of the year, and in the spring and summer
Merismopedia and Cilyndrospermum of the blue-greens ap-
peared.
Heptagenia nymphs, and Chironomus larve made up the
bulk of the animal food, although Simulium larve and Ostracods
were abundant. Difflugia shells were found a few times.
These results do not agree with the statements of Wesenberg-
Lund (1911) and Ussing (1907) that the larve are inactive,
lying rolled up in a spiral and taking little or no food in the
winter. The collection made in February came at a time when
the Creek was covered with ice. The larve were found on the
underside of stones in the strearh, either in a case of pebbles
loosely held together or a mass of roots spun into a tubular
form. When the stones were removed from the water and
placed on the bank, the larve came out of their tunnels at once
and crawled about over the stones. There was also an abund-
ance of food in the stomach in every case.
In examining the contents of the stomach of Chimarrha
aterrima (Family Philopotamidz) one is greatly surprised to
find vegetable food exclusively. The mandibles are strongly
developed, with sharp teeth, and lack the median tuft com-
pletely, which, according to Siltala, would point to an ex-
clusively carnivorous form. Examinations were made on
November 14, November 30, June 11 and July 14. On the first
three dates, the stomach contents consisted of diatoms ex-
clusively, the same forms as were found in the Hydropsychids.
On the last date, however, Euglena was very abundant, as
were the simple green alga Scenedesmus and other Protoccocales;
also desmid zygospores. In every instance there was very
much silt mixed in with the food.
Only one examination of food was made on the two larve
of the Family Polycentropide. This was on July 14, when
ten specimens of each species were examined. The food of
Polycentropus sp. was made up entirely of insects, Chirono-
mids being the principal diet, and Heptagenia nymphs quite
numerous.
268 Annals Entomological Society of America [Vol. VII,
Except for one Chironomid head there were no recognizable
contents in the alimentary tract of the Cyrnus sp.—only a
dark brown fluid exuded. After watching it feed, however, on
the soft bodied forms of microscopic organisms, one can
account for this fact.
V. EXPERIMENTAL WORK ON HYDROPSYCHE NETS.
To one who tries to study the method of construction of the
nets, feeding habits, etc., in the field, the following difficulties
present themselves. The threads of the net quickly become cov-
ered with diatoms, silt and algae which obscure the mesh to some
extent; the ripples on the surface of the water make it necessary
to work with a water-glass which cuts out some of the light;
also the nets are so low down that one can only view them
satisfactorily from above.
Although the Hydropsyche larva will construct its dwelling
tube in a dish of water in the laboratory, it builds no
catching-net. The larvae, however, made perfect nets in a
trough supplied with a steady stream of partially filtered water
from Cascadilla Creek. The trough stood on a framework
three feet high and was tilted slightly, the end nearest the water-
pipe being the higher. The side boards of the trough were
grooved (14 in. by 4 in.) their entire length, and the stream of
water striking the end board was carried down into the
grooves as well as into the trough. On each side, at the
point where the water from the groove met the overflow
from the trough (See Plate XXXVI, Photo 3.) the current was
the swiftest. As might be expected these spots were chosen in
preference to others as building sites. The only caution
taken was to induce the larva to begin its spinning very near
the end of the groove so that the net would come within the
focus of a lens held in front of the groove. The making of
the larval dwelling could best be observed from above, but
observations on the construction of the net and the feeding
habits could be seen to best advantage when one knelt in front
of the groove so that the eye came on a level with it. In all
cases a glass slide was placed over the groove to smooth the
surface of the water.
By the above methods, the following results have been
obtained.
1914] Biology of Net-Spinning Trichoptera 269
1. Time of building—Many Trichoptera larve build their
dwellings chiefly during the night, but these build their tubes
and nets at all times, during the day as well as at night.
2. Time required for building—On watching the con-
struction of several larval dwellings, I found the average time
for the completion of the tube and net to be from two and a
half to three hours. The larva spent about an hour in spinning
‘its tube and the remainder of the time on its net.
3. Different species of Hydropsyche larve placed in the
trough built similar dwellings.
4. There were no temporary construction threads in the
net as described for the web of orb-weaving spiders, (Comstock,
J. H., 1895, p. 37), all of the threads being permanent.
1 2 a
Text Figs. land2. 1. Diagram showing usual method of crossing of threads
to form the regular mesh of the net. 2. Attachment of threads. (a) at beginning
of thread; (b) continuation of same thread at point of departure from supporting
surface.
5. There seemed to be no definite order in which the
threads of the net were laid down. Sometimes the coarse, irregu-
lar mesh was spun immediately after the building of the larval
tube, while at other times the fine, regular mesh was spun first.
The larva at times left its work on the net and went back to
add a few threads to the case. In general the catching surface
was formed of threads crossing each other in the fashion
shown in figurel. Threads were fastened in the manner shown
in figure 2, the double thread being split for a short distance and
each half attached separately.
6. I have never observed the larve cleaning their nets
with the dorsal tuft of hairs on the anal prolegs, a function
which Lund (1911) stated as a probable one. They have
always removed particles from the net with their mouth-parts.
270 Annals Entomological Society of America [Vol. VII,
I believe that the thick cluster of bristles on the outer edges of
the labrum are used in removing the microscopic plants
from the meshes.
7. The larva used its front legs in combination with the
mandibles for seizing, and holding in position until fastened
with silk, any bits which it might wish to weave into its tube
or use as supports for the net.
8. The position indicated in Plate XX XVII, Fig. 5 is the
one usually assumed by the larva in spinning its net and in
feeding. Since no pebbles or vegetable bits were placed in the
groove, the larva spun its tube entirely of silk, and so its position
could be clearly seen. The larva rested ventral side up with
the hooks of the anal prolegs fastened in the roof of the tube.
Usually only the head and thorax protruded from the entrance,
but if the larva needed to reach out farther than the stretching
of the abdomen would permit, the body was moved forward
in the tube. The front legs were directed forward, and were
used chiefly for clinging on to the net during its construction.
The tarsal claws were passed rapidly along a thread near to the
one which was being spun. The second and third pairs of legs
were also used for holding on, being stretched out on either
side and shifting only as the movements of the larva demanded
it.
9. Feeding Habits——The larva never was so intent upon
finishing its net but that it stopped and picked off particles of
food adhering to the threads, ate them and then continued its
labors. As soon as it finished its net, and while the mesh was
practically clean, I put insect food (Simulium larve and Hep-
tagenia nymphs) into the groove. One specimen was used at
a time, and the net was effective in holding back food as the
water filtered through. The larva siezed any intruder almost
immediately with its front legs and mandibles and pulled it
down toward the mouth of its tube. It was not without a
struggle that its victims were subdued, sometimes as long as
five minutes being required. The larva seemed to swallow its
food whole, with little chewing of it, and shoved it into its mouth
with its front legs. Perfect or only slightly mutilated speci-
mens of Chiromid larve and Heptagenia nymphs found in the
cesophageal region seemed also to point to this method of
feeding.
1914] Biology of Net-Spinning Trichoptera 2a
VI. BIBLIOGRAPHY.
1901. Betten C. Aquatic Insects in the Adirondacks. Trichoptera. N. Y.
State Mus. Bull. 47, pp. 561-572.
1882. Clarke, Cora H. Description of two Interesting Houses made by Native
Caddis-fly larvae. Proc. Bost. Soc. Nat. Hist., Vol. 22, p. 67.
1891. Clarke, Cora H. Caddis-worms of Stony Brook. Psyche, Vol. VI, pp.
153-158.
1903. Comstock, Anna B. Ways of the Six-Footed. pp. 133-138.
1895. Comstock, J. H. A Manual of the Study of Insects. pp. 188-190.
1887. Fielde, A. M. On an aquatic larva and its case. Proc. Acad. Nat. Sc.
Phil. p. 298.
1886. Howard, L.O. Report of the Entomologist, Wash., p. 510.
1888. Howard, L. O. Insect Life. Vol. I, No. 4, pp. 100-101.
1901. Howard, L. O. Insect Book, pp. 204-205.
1905. Kellogg, V.L. American Insects, p. 242.
1895. Miall, L.C. The Natural History of Aquatic Insects. pp. 265-267.
1881. Muller, F. Uber die von den Trichopterenlarven der Provinz Santa
Catharina verfertigen Gehause, Zeitschr. z. wissenschaftl. Zool. Vol.
35, pp. 47-53.
1907. Petersen, E. Om planktonfangende, fangnetspindende Hydropsychidlarver
i Danmark. Meddel fra den naturh. Foren i Kbhyn. pp. 137-148.
1908. Petersen, E. Bidrag til Kundskab om planktonfangende fangnetspindende
Trichopterlarver i Danmark, II, ibid, pp. 123-126.
1895. Sharp, D. Insects, Cambr. Nat Hist..Vol. V, pp. 482-483.
1903. Siltala, A. J. (Silfvenius) Ein Fall von Schadlichkeit der Trichopteren
Larven, Meddel Soc. F. Fl. F., Vol. 29, pp. 54-57.
1907. Siltala, A. J. Uber die Nahrung der Trichopteren. Acta Soc. pro Fauna
et Flora Fennica. Vol. 29, pp. 1-34.
1903. Simpson, C. B. Photographing of Nets of Hydropsyche, Proc. Ent. Soc.
Wash. Vol. V, pp. 93-94.
1908. Thienemann, A. Die Fangnetze der Larven von Philopotamus ludificatus,
M. L. Zeit. f. wiss. Insektenbiol. Vol. IV, p. 378.
1909. Ussing, H. Om Hydropsyche-Larvens Fangnet. Flora og Fauna, Silke-
loYoreies ov (ile
1911. Wesenberg-Lund, C. Biologische Studien uber netzspinnende Trichop-
terenlarven. Internat. Rev. der gesamten Hydrobiol. und Hydrogr.
pp. 1-63.
272 Annals Entomological Society of America -[Vol. VII,
PLATE XXXVI.
Photos by J. T. Lioyp.
Photo 1. Cascadilla Creek below Fish Hatchery. Taken in midsummer when
the water is low to show character of creek bed. Earlier in the season,
the Hydropsyche nets stretched between the stones on the boiiom are
very numerous. In the quiet water at the left the larvae of the Family
Polycentropide are found.
Photo. 2. Cascadilla Creek in the spring as it rushes over the shelving ledges
beside the Hatchery. A favorite spot of the Hydropsyche larvae.
Photo 3. Trough where experiments were carried on. Water entered through
pipe above, and spilled over at corners at lower end, through grooves in
the sides, where the Hydropsyche larvae built perfect catching-nets.
Under the trough is a water-glass used in field work, and beside it, a fold-
ing bench for use while making observations in the stream.
PLATE XXXVII.
Fig. 1. Diagrammatic figure of a house of Hydropsyche angustipennis, copied from
Wesenberg-Lund (1911) (Plate IV, Fig. 22). At the left is the tube in which
the larva lives. In front of it is a vestibule with a catching surface of fine
mesh in its side wall. Near this net is the opening of the larval tube.
Fig. 2. A typical Hydropsyche dwelling in which the coarse, irregular mesh-
work of the net is not strengthened by any supporting bits. Enlarged x 2.
Fig. 3. End view of one of the grooves of the trough with the Hydropsyche
dwelling built in it.
Fig. 4. Hydropsyche dwelling built in trough, and viewed from above.
Fig. 5. Usual position assumed by the Hydropsyche larva in spinning its net
or in feeding.
PLATE XXXVIII.
Fig. 1. Catching-net of Chimarrha aterrima. Natural size.
Fig. 2. Dwelling of Cyrnus pallidus (?). Larva lives in the tube, and at either
opening is the catching-net. This is composed of radiating strands of
silk fastened at their outer ends to the stone, and connected with one
another by anirregular mesh. Enlarged x 2.
Fig. 8. Dwelling of Polycentropus sp. (?) Delicate silken tube in which larva
lives, slightly curved, and surrounded on all sides by a delicate irregular
mesh which functions as a catching-net. Enlarged x 2.
VOL. VII, PLATE XXXVI.
Noves
VOL. VII, PLATE XXXVII.
ANNALS E.S. A.
[ AS \\ AE
Nez)
Noyes
ANNALS E. S.A
VOL. VII, PLATE XXXVIII-
Nal
Noyes
THE CLASSIFICATION OF THE PUPAE OF THE
CERATOCAMPIDA AND HEMILEUCID.
EpNA MOosHER.
The pupae belonging to the superfamily Saturnioidea may
be identified by the following characteristics: Fifth and sixth
abdominal segments free in both sexes; body surface hard and
firm, always with setae, but these rarely long enough to be
observed with the unaided eye; face-parts never with distinct
sutures; antennal suture obsolete; labial palpi or maxillary
palpi never visible; distinct cases for the mandibles never
present, these structures often represented by an elevation
or a distinct tubercule adjoining the caudo-lateral angles of
the labrum; antennae usually showing distinct pectinations,
the width at least one-fifth the length and usually much wider,
the stem of the flagellum distinctly raised above the level
of the pectinations, or if the stem of the flagellum is not dis-
tinct, then the body with the cephalic margins of the movable
segments produced into distinct flange-like plates; maxillae,
measured on the meson, seldom more than one-sixth the length
of the wings, if longer, then the body surface without visible
setae; third pair of legs very seldom visible; pupae usually
more than an inch in length.
The pupae of this superfamily are found either in thick
silken cocoons or thin ‘‘papery”’ ones, or in the ground. More
than twenty genera are found in North America; of these, the
pupae of only sixteen genera were available for study.
The material on which the following descriptions and tables
are based was borrowed in part from the Illinois State Labora-
tory of Natural History. A large series of pupae was pur-
chased from the American Entomological Co., the Kny-
Scheerer Co. and Ward’s Natural Science Establishment, with
funds provided by the Graduate School of the University of
Illinois.
Dr. A. D. MacGillivray has given many helpful suggestions
as to the preparation of this paper, for which I wish to express
my appreciation.
* Contribution from the Entomological Laboratories of the University of
Illinois, No. 44.
277
278 Annals Entomological Society of America [Vol. VII,
The superfamily Saturnioidea may be divided into three
families as follows:
A. Pupae with the movable segments provided with flange-like plates
which prevents their being telescoped, their lateral margins distinctly
tapering caudad and each segment noticeably srnaller than the segment
cephalad of it; wings never elevated dorsad above the surface of the
body; a distinct cremaster always present; stem of the flagellum of
the antenna never elevated and distinct.
B. Pupae with a distinctly bifurcate cremaster; body usually rough-
ened with spines on the exposed surface of the thorax and
abdomen; metathorax with prominent oblong tubercles on each
side the meson extending one-third or more of the distance
between the meson and the margin of the first pair of wings;
pupae always found in the ground............. Ceratocampide
BB. Pupae without a distinctly bifurcate cremaster; body never
roughened with spines on the exposed surface of thorax and
abdomen; metathorax never with prominent oblong tubercules:
pupae found either in cocoons or in the ground ....Hemileucide
AA. Pupae with the movable segments never provided with flange-like
plates which prevent their being telescoped, the lateral margins approx-
imately parallelso that the segments appear of equal size and are
usually telescoped so that only the caudal margins of the segments
are visible; wings prominently elevated dorsad above the level of the
body, the caudal portion of the mesonotum and metanotum always
depressed adjacent to the wings; a distinct cremaster rarely present;
stem of the flagellum of the antenna always elevated and distinct.
Saturniidz
THE FAMILY CERATOCAMPIDA.
Body with the margins of the free abdominal segments
usually bearing a row of spines, and the exposed surface of the
thorax and abdomen usually roughened with spines; antennae
never broadly pectinate throughout, but broadly pectinate
and almost parallel for about one-half the length, then narrowed
rapidly to about half the greatest width, tapering gradually
to a pointed tip, the stem of the flagellum never distinct, the
surface convex and the central axis of the antenna usually bear-
ing one or two rows of small spines; maxillae, measured on the
meson, never less than one-fourth the length of the wings; tips of
the tarsi of the second pair of legs meeting obliquely on the meson,
never lying adjacent on the meson; proleg scars very promi-
nent on abdominal segments five and six, the scars for the anal
prolegs often very conspicuous; first pair of wings with the
anal angles broadly rounded, usually located at the cephalic
margin of the fourth abdominal segment and never reaching
ventrad to the caudal margin of the fourth segment; second
pair of wings never produced below anal angle of first wing and
1914] Pupe of Ceratocampide and Hemileucide 279
never visible in ventral view; metathorax with distinct tuber-
cules more or less oblong in outline on each side the meson and
extending more than one-third the distance from the meson
to the margin of the wing; the suture between the seventh
and eighth segments never deep with distinct crenulations on
its margins; cremaster always present, usually long and bifurcate
at tip. Five genera of this family have been described. One
genus, Syssphinx, consisting of three species, was not available
for study. The remaining genera of Ceratocampide can be
separated by the following table:
A. Surface of pupa never spinose; cremaster broader than long, broadly
and shallowly bifurcate, never over 2 mm. in length.......... Citheronia
AA. Surface of pupa spinose; cremaster at least twice as long as broad,
bifurcate at tip, always more than 2 mm. in length.
B. Thorax rugose with short isolated spines, abdominal segments
not spinose, but bearing a row of spines along both cephalic
and caudal margins of segments 1 to 7, the spines along the
caudal margins of segments 5 to 7 much longer than the spines
CUM MCEP ACH ROW Sera ta sate rai tte ee ees teree tee Basilona
BB. Thorax and abdominal segments densely spinose; abdominal seg-
ments 1 to 7 with a row of spines along both cephalic and caudal
margins, the spines in the cephalic rows on abdominal segments
5 to 7 usually much longer than the spines in the caudal rows;
maxilla, measured on meson, one-fourth the length of the
wings.
C. Usually with prominent scattered spines on the thoracic
segments, at least four times as long as those covering
the segments; antennae with the central axis bearing
a row of prominent spines curved caudad; if without
prominent spines on the thoracic segments and antennae,
then the maxillae, measured on meson, one-third the
length of the wings.
D. Eighth abdominal segment never with a prominent
transverse ridge in the middle of the segment
bearing a row of spines; glazed eye-piece always
lighter in color than the remaining surface of the
body; species always more than an inch in length.
Adelocephala
DD. Fighth abdominal segment always with a prominent
transverse ridge in the middle of the segment bear-
ing a row of spines; glazed eye-piece always the
same color as the remaining surface of the body;
species never more than an inch in length. .Dryocampa
CC. Without prominent scattered spines on the thoracic seg-
ments, the longest never four times the length of those
covering the segments; antennae with the central axis
never bearing prominent spines, the spines never curving
caudad; maxillae, measured on meson, always one-fourth
Cements OleiiemWilOS. soins cts Saaace rae oes oe Anisota
280 Annals Entomological Society of America [Vol. VII,
Genus Citheronia Hubner.
Face-parts and appendages not at all elevated; body surface
not roughened with spines; eye-pieces both present; invagina-
tions for the anterior arms of the tentorium small but distinct;
clypeo-labral suture present; labrum a little wider than long;
maxillae, measured on the meson, about two-fifths the length
of the wings, but little longer than the greatest width, tri-
angular in outline; tips of the tarsi of the first and second
pair of legs meet obliquely on the meson; median line distinct
on all thoracic segments; mesothorax with a few minute tuber-
cules at the bases of the wings; metathorax with a prominent
oblong tubercule or plate, irregularly sculptured at the sides,
on each side the meson, extending more than half the width
of the segment and nearly its whole length; cephalic margins
of abdominal segments 5 to 7 produced into thin, plate-like
ridges; spiracular line curved slightly ventrad; cremaster
short and bifurcate at tip.
This genus is found principally east of the Mississippi
and consists of two species, C. regalis and C. sepulchralis.
Specimens of the latter were not available for study. The
pupae of C. regalis have a peculiar odor somewhat resembling
laudanum.
Citheronia regalis Fabricius. Color dark brown, almost
black; body surface usually polished, occasionally roughened
with indeterminate transverse striations; antennae in both
sexes with the length more than four times the greatest width
and reaching about half way along the exposed portion of the
second pair of legs; face parts with a slightly raised line on each
lateral margin of the clypeus extending cephalad from the
proximo-lateral margins of the labrum to the proximal ends of
the antennae; labrum variable, five-sided, pointed at the
distal end; maxillae much longer than broad, the proximal
margin sinuate; prothoracic spiracle with elevated margins, the
cephalic margin forming a prominent rounded ridge; meso-
thorax with a small tubercule on each side the meson on the
caudal half of the segment, a tubercule scar laterad of each
tubercule and in line with it, and a smaller tubercule near the
caudal margin of the alar area on each side; abdominal seg-
ments 2 to 7 with a row of punctures near the cephalic margin, in
1914] Pupe of Ceratocampide and Hemileucide 281
the movable segments, at the caudal margin of the ridge and
extending all around the segment; segmentation in abdominal
segments 8 to 10 hard to determine; the eighth segment usually
polished, its dorsal cephalic margin roughened and plate-like,
with a row of punctures along the cephalic margin of the plate
and opening cephalad; abdominal segments with two dorsal
rows of tubercule scars and one ventral row; cremaster short,
never exceeding two millimeters in length, broader than long
and broadly and shallowly bifurcate at tip. Length 134’-21%’;
girth about equal to length.
Genus Basilona Boisduval.
Face parts slightly elevated above the level of the append-
ages; body surface roughened with spines; eye-pieces both
present; invaginations for the anterior arms of the tentorium
small and indistinct; clypeo-labral suture present; labrum
with the length and breadth approximately equal; maxillae,
measured on the meson, with the length twice the greatest
width and one-half the length of the wings, triangular in out-
line; tips of the tarsi of the first pair of legs usually meeting on
the meson, but sometimes falling short so that the tips of the
maxillae lie between them; tips of the second pair of legs
always meeting obliquely on the meson; median line distinct
on prothorax and mesothorax and sometimes showing on the
cephalic half of the metathorax; metathorax with a prominent
oblong roughened tubercule with fluted edges on each side the
meson, extending half the distance between the meson and the
margin of the first pair of wings; cephalic margins of abdominal
segments 5 to 7 never with any indications of a plate or ridge;
spiracular line curved slightly ventrad; cremaster long, bifurcate
at tip. .
This genus contains a single species, Basilona imperialis,
found in the states east of the Mississippi.
Basilona imperialis Drury. Color dark brown; body sur-
face with indeterminate sculpturing and roughened with
spines; antennae with the length four times the greatest width,
the central axis set with a row of short spines directed caudad;
face parts roughened with spines irregularly arranged, with
the exception of a row extending cephalad from each proximo-
lateral angle of the labrum to the proximal end of the antenna,
282 Annals Entomological Society of America [Vol. VII,
sometimes confused with the general sculpturing; labrum
variable, usually five-sided, pointed at the distal end; maxillae
with the length twice the breadth, each half quadrilateral;
prothorax slightly wrinkled, with a row of spines around entire
margin except in the region of the spiracles; mesothorax with
fine indeterminate transverse striations and very small spines,
a spinose area extending from the meson to the alar area, a
few small spines at the base of the wings; wings with the
venation outlined with short spines; abdominal segments
1 to 8 with an interrupted row of very small spines along the
cephalic margin dorsally, and with many large semicircular
to ovate punctures caudad of the spines, distributed over
the cephalic third of the segment and the spiracular
region, the remainder of the segment sparsely covered with
smaller circular punctures; caudal margins of all abdominal
segments with a row of small curved spines directed caudad,
the spines larger than those on the cephalic rows, the largest
on segments 8 to 10; lateral cephalic margins of abdominal
segments 5 to 7 cephalad of the spiracles with three prominent
transverse ridges with distinct furrows between; cremaster
from 5 to 7 millimeters in length, a smooth dorsal concavity
at the cephalic end, then strongly rugose to the bifurcate tip.
Length 134” to 2”; girth about equal to length.
Genus Adelocephala Herrich-Schaeffer.
Face parts very slightly raised above the level of the append-
ages; body surface roughened with spines; eye-pieces both
present, the glazed eye-piece always lighter in color than the
remaining body surface; invaginations for the anterior arms of
the tentorium small, but distinct; clypeo-labral suture present;
labrum broader than long; maxillae, measured on the meson,
never less than one-fourth the length of the wings, triangular in
outline; distal two-thirds of the tarsi of the first pair of legs
adjacent on the meson, the tips of the tarsi of the second pair
of legs meeting obliquely on the meson; median thoracic line
distinct on prothorax and mesothorax; metathorax with an
oblong tubercule on each side the meson, not prominently
elevated, but slightly rugose and polished; cremaster long,
bifurcate at tip.
“=
1914] Pupe of Ceratocampide and Hemileucide 283
This genus contains two species, bicolor, found in the
Mississippi Valley and the Southern Atlantic states, and
bisecta, found in the Ohio Valley. The species may be sepa-
rated as follows:
A. Antennae with prominent spines; spines of cephalic margins of abdominal
segments 5 to 7 larger than those on the other segments........ bicolor
AA. Antennae without prominent spines; spines on the cephalic margins of
abdominal segments 5 to 7 not larger than those on the other seg-
FALETUG SEN Wee 5 a occa aut eee aPC I OT eee Scie Shouts chase teh eee ttle bisecta
Adelocephala bicolor Harris. Color dark reddish brown;
head, thorax and appendages finely spinose; abdominal seg-
ments both punctate and finely spinose; antennae with the
length four times the greatest width, strongly convex, with
two rows of spines, the outer row, large, prominent and curved
caudad, the mesal row minute; face parts with an elevated
spiny ridge on each side extending cephalad from the proximo-
lateral angles of the labrum to the proximal end of each antenna,
bearing a prominent spine near the cephalic end and a smaller
one half way between this and the labrum; epicranial area with
two prominent spines on each side the meson at the proximal
end of each antenna; sculptured eye-piece with a prominent
spiny tubercule; labrum usually six-sided, broader than long,
maxillae with length and greatest width equal, each half
quadrilateral, the length measured on meson, one-fourth the
length of the wings; first and second pair of legs elevated and
convex; cephalic portion of prothorax prominently elevated
on meson sloping gradually to lateral margins, the larger
spines on the elevation pointing dorsad, a slight elevation
with larger spines near the meson at caudal margin on each
side the meson; prothoracic spiracles with cephalic margins
arcuate; mesothorax with a slightly elevated ridge each side the
meson with at least two bifid spines, a prominent spine at the:
base of each wing and another half-way between these spines
and the meson; abdominal segments 1 to 4 with rows of minute
spines along the cephalic and caudal margins of the exposed
portion; abdominal segments 5 to 7 having the cephalic margins.
dorsad between the spiracles with sharp transverse ridges and
distinct furrows between, ventrad with large circular punctures,
the margins produced into flange-like ridges set with broad,
flat, erect spines, many of them bifid; the caudal margins of
abdominal segments 5 to 7 with similar but very much smaller
spines, the spines of both cephalic and caudal rows much smaller
284 Annals Entomological Society of America [Vol. VII,
on the venter; abdominal segments 8 to 10 thickly punctate,
the eighth segment with a distinct lateral protuberance on
each side and a prominent tubercule on the meson; ninth and
tenth segments with some larger spines on the lateral margins;
cremaster with a smooth V-shaped area on the proximal end
at dorsum, with the point of the V prolonged down the middle
of the cremaster, the remainder of the surface irregularly
rugose and bifurcate at tip for about one-fourth the length,
the tips divergent. Length 114” to 134”, cremaster one-seventh
of total length; girth slightly less than length.
Adelocephala bisecta Lintner. Color dark reddish brown;
head, thorax and appendages very finely spinose; antennae
with the length about three times the greatest width, sometimes |
exceeding this, slightly convex and without prominent spines;
face parts without prominent ridges or spines; labrum some-
what six-sided, tuberculate; maxillae with the length greater
than the breadth, the length measured on meson, one-third
the length of the wings; thorax without any prominent spines;
abdominal segments 1 to 8 with rows of minute spines along
the cephalic and caudal margins of the segments; segments
9 and 10 with rows of spines near the caudal margins, and
without any prominent lateral spines; cremaster very rugose,
bifurcate at tip for less than one-fourth its length, the tips
but slightly divergent. Length 114” to 134”, the cremaster
about one-ninth the total length; girth about equal to length.
Genus Dryocampa Harris.
Face parts elevated above the level of the appendages;
body roughened with spines; antennae with a row of prominent
spines curving caudad on each central axis; eye-pieces both
present; invaginations for the anterior arms of the tentorium
small, but distinct; clypeo-labral suture present; labrum
a little wider than long; maxillae, measured on meson, one-
fourth the length of the wings, triangular in outline; about
half the exposed portion of the first pair of legs lying adjacent
on the meson; tips of the tarsi of the second pair of legs meeting
obliquely on the meson; median line elevated on prothorax
and distinct on mesothorax, represented on the cephalic two-
thirds of the metathorax by a clear elevated area; metathorax
with a prominently elevated, polished tubercule on each side
the median elevation, slightly rugose and extending at least
one-third the distance from the meson to the margin of the
1914] Pupe of Ceratocampide and Hemileucide . 285
first pair of wings; cephalic margins of abdominal segments
5 to 7 produced into prominent flange-like ridges directed
cephalad and set with spines; abdominal segments 9 and 10
with prominent lateral spines; cremaster long, over one-seventh
the total length of the body, bifurcate at tip.
This genus includes a single species, Dryocampa rubicunda,
found east of the Mississippi.
Dryocampa rubicunda Fabricius. Color dark brown to
black; exposed surface of head, thorax and appendages finely
spinose, the abdominal segments both punctate and spinose;
face parts with an elevated spiny ridge on each side extending
cephalad from the proximo-lateral angles of the labrum to the
proximal end of each antenna, bearing two or three prominent
spines; epicranial area with a prominent laciniate spine on each
side the meson at the proximal end of each antenna directed
cephalo-laterad and giving the pupa a horned appearance;
glazed eye-piece usually one-third or more the entire width,
the sculptured portion bearing at least one prominent spine;
labrum six-sided, usually slightly sunken, pointed at distal
end; maxillae with the greatest width and length approximately
equal, each half triangular; prothorax with a few slightly
larger spines on each side the median line; prothoracic spiracles
with the cephalic margins arcuate; mesothorax with two
prominent spines along cephalic margin near the meson, a
large scattered group at base of wing and half way between
these two groups on each side the largest thoracic spine;
abdominal segments 1 to 4 with a row of minute spines along
both cephalic and caudal margins; abdominal segments 5 to 7
with the margins punctate, produced into flange-like ridges
directed cephalad and bearing a row of large sharp spines
occasionally bifid or trifid and about one-third the length of the
segment, the caudal part of these segments with a distinct
furrow near the caudal margin separating the cephalic spinose
portion from a narrow smooth portion, with a row of small
spines between it and the transverse conjunctiva; eighth
abdominal segment with a row of large spines dorsally on the
summit of a median transverse ridge, extending laterad and
becoming indistinct on the ventral aspect; abdominal segments
9 to 10 with prominent lateral spines curving caudad; cremaster
irregularly, longitudinally rugose, bifurcate at tip with the
points widely divergent. Length 7-8” to 1”; girth less than
length.
286 Annals Entomological Society of America _[Vol. VII,
Genus Antsota Htibner.
Body with the cephalic margins of abdominal segments.
5 to 7 produced into flange-like ridges directed cephalad, and
set with spines; exposed surface of head and thorax spinose,
the abdominal segments both spinose and punctate; both
eye-pieces present, the sculptured portion spinose; invagina-
tions for the anterior arms of the tentorium small, but distinct;
clypeo-labral suture present; labrum variable, small, never
twice as broad as long; maxillae, measured on the meson,
always one-fourth the length of the wings, triangular in outline;.
tarsi of the first pair of legs adjacent on the meson, tips of the
tarsi of the second pair meeting obliquely on the meson; meta-
thorax with a prominent oblong tubercule on each side the
meson, extending more than one-third the distance between
the meson and the margin of the first pair of wings; cremaster
always long and bifurcate at tip.
This genus includes at least five species commonly found
in the United States, one of these, A. skinnerz, is reported from
Arizona, the other four from the states east of the Mississippi.
These five species can be separated by means of the fol-
lowing table:
A. Cremaster one-eighth or more of the total length of the body; spines
on the epicranial area at the proximal end of each antenna large and
prominent, extending beyond the margin of the body in ventral view
and giving the pupa a horned appearance.
B. Cremaster more than one-eighth the total length of body and
bifurcate for less than one-fourth its length; small species,
lesssthanwone inch inysleneihe ein. ene virginiensis.
BB. Cremaster about one-eighth the total length of the body and
bifurcate for one-fourth its length; species one inch or more in
length.
C. Face parts prominently elevated above the level of the
appendages; mesothorax with at least one laciniate
spine on each side the meson near the cephalic margin.
stigma
CC. Face parts not elevated above the level of the appendages;
never with a laciniate spine on each side the meson near
the icephiahte mare: 2h. tu yaeees cos Miele ya aes ot oe: senatoria.
AA. Cremaster less than one-eighth the total length of the pupa; spines
of the epicranial area at the proximal end of each antenna never
extending beyond the margin of the body in ventral view, so that
the pupa does not present a horned appearance.
Each metathoracic tubercule very prominently elevated, its
length more than half the length of the segment and extending
at least half the distance from the meson to the margin of the
firstspainiob wines colon blacks. yee ee ee skinneri
BB. Each metathoracic tubercule somewhat diamond shaped, never
very prominently elevated, its length never as much as half the
length of the segment, and never extending half the distance
between the meson and the margin of the first pair of wings;
color*bright reddish browne sete oe eee consularis.
1914} Pupe of Ceratocampide and Hemileucide 287
Anisota virginiensis Drury. Color dark brown to black;
abdominal segments 1 to 4 and 8 to 10 with few spines and more
large circular punctures as compared with the remainder of
the surface; each antenna with two rows of minute spines on
the central axis, the length three times the greatest width;
face parts prominently elevated above tHe level of the ap-
pendages, an elevated densely spinose ridge extending cephalad
from the proximo-lateral angles of the labrum to the proximal
end of each antenna with a large spine at its cephalic end;
epicranial area with one large spine and several smaller ones
on each side the meson near the proximal end of each antenna;
labrum variable, usually six-sided, with two small tubercules
or spines, the width greater than the length, pointed at distal
end; maxillae with the length and breadth approximately
equal, each half quadrilateral; median thoracic line distinct
on all segments; prothorax with the median line slightly ele-
vated; mesothorax without prominent spines, usually with two
tubercule scars on each side the meson, sometimes spine-like,
seldom all prominent; metathoracic tubercules wedge-shaped,
irregularly impressed, black and polished, each extending
less than half the distance from the meson to the margin of
the first pair of wings; abdominal segments 1 to 3 with an
indistinct row of minute spines along, both cephalic and caudal
margins of the segment; abdominal segments 5 to 7 with the
cephalic margins punctate and produced into flange-like
ridges projecting cephalad and set with stout spines less than
one-sixth the length of the segment; caudal margins of segments
4 to 7 with a slight depression, the elevation adjacent to the
transverse conjunctiva set with two rows of minute spines;
eighth segment with a transverse ridge in the middle of the
segment set with spines, with slightly larger spines on the
lateral margins of the segment; ninth abdominal segment
with prominent lateral spines and the tenth with a prominent
hooked spine on each side the base of the cremaster; cremaster
longitudinally rugose, bifurcate for less than one-fourth its
length, the tips divergent. Length 7-8”; cremaster about
one-seventh the total length; girth less than length.
Anisota stigma Fabricius. Color dark reddish brown;
antennae in both sexes with the length about three times the
greatest width, central axis bearing a row of minute spines;
288 Annals Entomological Society of America _[Vol. VII,
face parts prominently elevated above the level of the append-
ages, an elevated ridge extending cephalad from each proximo-
lateral angle of the labrum to the proximal end of each antenna,
bearing a large laciniate spine near its cephalic end; epicranial
area with a stout curved spine on each side the meson near
the proximal end of the antenna; labrum variable, usually
hexagonal, with two small tubercules or spines and pointed at
the distal end; prothorax with the median line generally ele-
vated, more densely spinose on each side adjacent to the meson
than on the remainder of the segment; mesothorax with one
and sometimes two laciniate spines on each side the meson
near the cephalic margin with sometimes one or two smaller
spines, a scattering group of spines at the base of each wing
and one spine on each side, half-way between the base of the
wing and the meson, which is larger than those covering the
segment; metathoracic tubercules rugose, somewhat diamond-
shaped, each extending about half the distance from the meson
to the margin of the first pair of wings, subadjacent on the
meson; abdominal segments 1 to 3 with a row of minute spines
along both cephalic and caudal margins of the segment; cephalic
margins of abdominal segments 5 to 7 punctate and produced
into flange-like ridges directed cephalad, bearing a row of
prominent, erect, triangular spines, less than one-fourth the
length of the segment; caudal margins of abdominal segments
4 to 7 with a furrow near the caudal margin of the segment
and a row of spines on the elevation at the junction of the seg-
ment and the transverse conjunctiva, these spines about one-'
third the size of the spines in the cephalic rows; abdominal
segments 8 to 10 with fewer spines and more punctures on
the surface; the eighth abdominal segment with a prominent
transverse ridge in the middle of the segment, with a slight
protuberance on each lateral margin, the transverse ridge set
with spines similar to those along the caudal margins of seg-
ments 4 to 7, a smaller row along the caudal margin of the seg-
ment; ninth abdominal segment with two rows of spines near
the caudal margin with two or three prominent ones along each
lateral margin; tenth segment with two or three prominent
spines along each lateral margin at the proximal end of cre-
master; cremaster with a smoother, triangular depressed area
dorsad at proximal end, the remainder of the surface rugose
with wavy longitudinal ridges, the caudal end bifurcate for
1914] Pupe of Ceratocampide and Hemileucide 289
less than one-fourth of the length, the tips divergent. Length
1”—1 1-8”; cremaster about one-ninth the total length; girth
equal to length.
Anisota senatoria Smith and Abbott. Color dark brown to
black; antennae scarcely convex, each central axis with two
rows of minute spines, length about three times the greatest
width; face parts slightly elevated above the level of the
appendages; no prominent ridge extending cephalad from
each proximo-lateral angle of the labrum, but a prominent
curved spine on each side the cephalic part of clypeal area
adjacent to the proximal end of each antenna; epicranial area
with a prominent curved spine at the proximal end of each
antenna and usually one or two smaller ones; labrum usually
six-sided, broader than long, usually with two small tubercules,
slightly pointed at the distal end; maxillae with the length
slightly greater than the greatest width, each half quadri-
lateral; prothorax with a dense row of slightly larger spines on
each side the median line; mesothorax with a tubercule scar
on each side the meson indicated by a small polished area;
mesothorax without prominent spines; metathorax with the
tubercules oblong, slightly rugose, black and polished, each
extending less than half the distance from the meson to the
margin of the first pair of wings; abdominal segments 1 to 3
with a row of minute spines along the cephalic and caudal
margins of each segment; cephalic margins of abdominal -seg-
ments 5 to 7 with one distinct furrow dorsally and punctate
around entire segment, produced into flange-like ridges bearing
stout spines about one-fourth the length of the segment;
abdominal segments 4 to 7 with a distinct depression near the
caudal margin of the segment and with a caudal row of small
spines between the segment and the transverse conjunctiva,
with an interrupted row of smaller spines just cephalad; eighth
abdominal segment with a distinct median transverse ridge
bearing spines similar to those on the cephalic margins of seg-
ments 5 to 7, a row of small spines along the cephalic margin
of the ninth abdominal segment with two rows of spines near
its caudal margin and several prominent lateral spines; tenth
abdominal segment with one or two prominent lateral spines
at the proximal end of the cremaster, smaller than those
on the ninth segment; cremaster with a slightly depressed
290 Annals Entomological Society of America _[Vol. VII,
heart-shaped area at the proximal end with fine longi-
tudinal ridges, about three-fifths of the remaining length
finely rugose, the distal end smooth, bifurcate for about one-
fourth its length, the tips slightly divergent. Length 1 1-8”’—
114”; cremaster about one-ninth the total length; girth less
than length.
Anisota skinneri Biederman. Color dark brown to black;
antennae with the length three times the greatest breadth,
a row of minute spines on the central axis of each antenna;
face parts slightly raised above the level of the appendages, the
ridge extending cephalad from each proximo-lateral angle of
the labrum scarcely indicated, a medium sized laciniate spine
on the face parts near the proximal end of each antenna;
epicranial area with a long laciniate prominence or ridge,
which is never horn-like, with a small spinose tubercule caudad
of it on each side the meson near the proximal end of each
antenna; labrum variable, usually five-sided, broadly rounded
or slightly pointed at the distal end; maxillae with the length
and breadth approximately equal, each half quadrilateral;
prothorax more densely spinose on each side adjacent to the
median line; mesothorax without any especially prominent
spines; metathoracic tubercule strongly elevated, ovate, irregu-
larly impressed, almost adjacent on the meson, and extending
half the distance from the meson to the margin of the first pair
of wings; abdominal segments 1 to 4 with a row of minute,
closely set spines along both cephalic and caudal margins of
the segment; cephalic margins of abdominal segments 5 to 7
dorsad with sharp transverse ridges with distinct furrows
between and punctate around entire segment, produced into
flange-like ridges set with spines only about one-eighth the
length of the segment; abdominal segments 4 to 7 with a dis-
tinct furrow near the caudal margin of the segment and two
distinct rows of minute spines between the segment and the
transverse conjunctiva; eighth abdominal segment with a
slightly elevated transverse ridge in the middle of the segment
set with small spines and another row at the caudal margin
of the segment; ninth abdominal segment with two rows of
spines at the caudal margin of the segment, some spines slightly
more prominent at each lateral margin; tenth abdominal
segment with a small lateral spine on each side the cremaster;
1914] Pupe of Ceratocampide and Hemileucide 291
cremaster with a small, triangular, slightly depressed area at
the proximal end of cremaster dorsad, but rugose much like
the remainder of the surface, bifurcate at tip for less than one-
fourth the length, the tips not divergent. Length 1 3-8’—
1 5-8”; cremaster about one-tenth total length; girth exceeding
length.
Anisota consularis Dyar. Color bright reddish brown;
antennae with the length about four times the greatest width;
face parts slightly raised above the level of the appendages, an
elevated ridge extending cephalad from each proximo-lateral
angle of the labrum to the proximal end of each antenna and
bearing several prominent spines; epicranial area with a large
spine on each side the meson near the proximal end of each
antenna; labrum variable, usually five-sided, broader than
long and bearing two minute tubercules or spines, slightly
pointed at the distal end; maxillae with the length greater than
the breadth, each half quadrilateral; prothorax with a larger
spine on each side the median line near the middle of the
segment; mesothorax without any especially prominent spines,
a few longer ones at the base of each wing; metathoracic tuber-
cules irregular, somewhat diamond-shaped, black and polished,
irregularly impressed or punctate, each tubercule extending
less than half the distance from the meson to the margin of
the first pair of wings; abdominal segments 1 to 4 with a row
of very minute spines on each cephalic and caudal margin;
abdominal segments 5 to 7 with the cephalic margins punctate
and produced into flange-like ridges directed cephalad and set
with spines less than one-sixth the length of the segment, a
smooth band at the caudal margin of the segments and a row
of small spines along the segment adjacent to the transverse
conjunctiva, almost wanting on the seventh segment; eighth
segment with a row of spines on a slight transverse ridge in
the middle of the segment, becoming indistinct in ventral view,
the caudal row of spines indistinct dorsad, but very distinct
laterad and ventrad; ninth abdominal segment with a caudal
row of spines, a prominence on the lateral margin set with
longer spines; the tenth segment with two prominent lateral
spines on each side of the cremaster; cremaster with a small,
triangular depressed area, much smoother than the remainder
of the surface, which is longitudinally rugose, bifurcate for about
292 Annals Entomological Society of America [Vol. VII,
one-fourth the length, the tips divergent. Length 1 1-8”—1 3-8’;
cremaster less than one-eighth the total length; girth equal to
length.
-THE FAMILY HEMILEUCIDE.
Margins of the free segments never with a row of spines; the
body surface never roughened with spines; antennae with the
stem of the flagellum never distinct, the central axis never set
with spines, the antennae tapering gradually from the part
with the greatest width; maxillae measured on the meson never
more than one-sixth the length of the wings; proleg scars
seldom prominent on abdominal segments five and six and
rarely with the anal proleg scars visible; first pair of wings
with the anal angles broadly rounded, usually at the cephalic
margin of fourth abdominal segment, and usually reaching
the caudal margin of the fourth abdominal segment ventrally;
second pair of wings never produced below the anal angles of
the first pair of wings and never visible in ventral view; meta-
thorax never with prominent tubercules; abdominal segments
5 to 7 with their cephalic margins produced into thick oblique
flange-like plates directed caudad; cremaster short, never
bifurcate at tip.
Altho not usually included with the Hemileucide the genus
Automeris is placed in this group owing to the very evident
relation of the pupae to those of the genera Hemileuca and
Pseudohazis. Morphologically they seem to be more nearly
related to the Hemileucide, but they are found in cocoons
like the Saturniide.
The description of this family is of necessity very incomplete
owing to lack of material. According to our available knowl-
edge of the subject the three genera may be separated as
follows:
A. Cremaster bearing setae arranged in a transverse row and spreading
oUub: fam-litees on. 5 vids ete «cee eee Meee peter eee ee mem Pseudohazis
AA. Cremaster never with setae, either with curved spines or without
spines or setae of any kind.
B. Cephalic part of segment above the flange-like plate either smooth
or with fine longitudinal striations; pupae found in ground.
Hemileuca
BB. Cephalic part of segment above the flange-like plate with sharp,
transverse ridges, deep furrows between; pupae found in cocoons
Automeris
Trl,
1914} Pupe of Ceratocampide and Hemileucide 293.
Genus Hemileuca Walker.
Face parts slightly elevated above the surface of the body;
antennae with the stem of the flagellum indistinguishable from
remainder of surface, entire surface flat to uniformly convex,
tapering gradually to a point at the distal end; invaginations
for the anterior arms of the tentorium distinct; eye-pieces.
both present; clypeo-labral suture generally distinct; maxillae,
measured on meson, never more than one-sixth the length of
the wings, each half quadrilateral; less than half the exposed
tibiae and the tarsi of the first pair of legs with the tips of the
second pair of legs adjacent on the meson; second leg visible
for almost entire tibial and tarsal length; median thoracic
line always distinct on prothorax and mesothorax, seldom on
metathorax; first pair of wings with the anal angles broadly
rounded near cephalic margin of fourth abdominal segment;
second pair of wings visible along entire dorsal margin of first
wing, its margin entire, but never produced beyond anal angle
of first pair of wings and never visible on the ventral surface;
spiracular line almost straight; cephalic margins of abdominal
segments 5 to 7 produced into thick, oblique flange-like plates;
suture between the seventh and eighth abdominal segments
deep, both margins usually strongly crenulate, the crenulations
of the two sides fitting together like a set of teeth; cremaster
short, pointed, never exceeding two millimeters in length.
This genus includes at least nine species found in the United
States, only three of which are described here. The most
common species is H. maia, which is found from the Atlantic
states westward to the Rocky Mountains. The others are
reported from the western states. These moths spend their
pupal life in the ground. The species described can be sepa-
rated by the following key:
A. Suture between the seventh and eighth abdominal segments very deep,
the edges distinctly crenulate.
B. Clypeal region strongly convex; labrum strongly elevated; max-
illae short, inconspicuous, each half triangular in outline and
length on meson less than a millimeter; mesothorax with a
tubercle on each side the meson outlined by a depressed ring.
burnsi
BB. Clypeal region not strongly convex; labrum not elevated; maxillae
conspicuous, each half quadrangular in outline and meeting
on meson for at least a millimeter; mesothorax without tuber-
CUleswoOnmeaGhisscid Cather MMeSOME er tnn tec. os cicte soh Pate, ohare yee a= maia
AA. Suture between the seventh and eighth abdominal segments not very
deep, the edges without distinct crenulations................... olivie
294 Annals Entomological Society of America _[Vol. VII,
Hemileuca maia Drury. Color dark brown; face-parts and
appendages with fine transverse striations, remainder of surface
shagreened, excepting abdominal segments 8 to 10; face-parts
without a prominent convexity in clypeal region; antennae
in male with length four times the width, the sides parallel for
at least the proximal two-thirds of their length and then
tapering rapidly to a point, reaching just below the tips of the
first pair of legs; clypeo-labral suture sometimes indistinct;
labrum about twice as broad as long; quadrate and broadly
truncate at distal end; maxillae, measured on meson, one-sixth
the length of wings, its median length less than its greatest
breadth; first pair of wings with their anal angles at the cephalic
margin of fourth abdominal segment; abdominal segments
1 to 4 and 7 to 8 with distinct furrows between, their margins
wavy, more apparent on the cephalic margins of the segments;
abdominal segments 5 to 7 with their cephalic margins produced
into thick flange-like plates covered with fine longitudinal
striations and a distinct smooth furrow at the caudal margin
of the segment, adjoining the transverse conjunctiva; cre-
master nearly two millimeters in length, indefinitely rugose,
triangular in outline, pointed at distal end, which bears many
hooked spines. Length, abdomen retracted, about 1”, girth
about °114”.
Hemileuca maia var. lucina Hy. Edwards. Specimens
of this variety from the New England Entomological Exchange,
collected in New Hampshire, show little general difference
from H. maia. They are much smaller, however, varying from
9-16” to 34” in length.
Hemileuca burnsi Watson. Color dark brown; face-parts
and appendages with fine, transverse striations, the remainder
of the body surface shagreened; face-parts with a prominent
convexity in the clypeal region; antennae of male with length
three times the width, tapering from the region of greatest
width to form a long, pointed tip at distal end, ending opposite
the tips of the first pair of legs; clypeo-labral suture distinct,
labrum elevated, somewhat shield-shaped, rounded at distal
end; maxillae very short, scarcely visible, each half of maxilla
triangular, much broader than long; prothoracic spiracles
with strongly elevated margins; mesothorax with a prominent
tubercule on each side the meson, outlined by a depressed ring;
1914] Pupe of Ceratocampide and Henuleucide 295
first pair of wings with their anal angles nearly opposite the
caudal margin of the fourth abdominal segment; sutures
between abdominal segments 1 to 4 distinct, margins of adjoin-
ing segments crenulate, suture between segments 7 and 8 very
prominent, the dorsal cephalic margin of the suture with
longitudinally corrugate ridges, the caudal margin crenulate;
abdominal segments 5 to 7 with their cephalic margins produced
into a prominent, flange-like plate, with longitudinal striations,
never more than indications of a furrow at caudal margins of
segments, an elevated roughened line between the caudal
margin of the segment and the transverse conjunctiva; cremaster
short, not more than a millimeter in length, triangular, rugose,
ending in a blunt tip at distal end, without spines. Length
about 7-8”; girth about 1”.
Described from one male specimen, for which we are in-
debted to Dr. Wm. Barnes, of Decatur, Illinois.
Hemileuca oliviz Cockerell. Color dark brown; surface
of body with interrupted transverse striations or impressions;
face-parts slightly elevated, but without a prominent con-
vexity in clypeal region; antennae in male with length a little
more than three times the width, the sides parallel for at
least two-thirds of the distance and then tapering to form a
blunt, rounded tip, ending opposite tips of second pair of legs;
clypeo-labral suture distinct; labrum with length and breadth
approximately equal, five-sided, with a sharp point at distal
end; maxillae, measured on meson, about one-seventh the
length of the wings, each half the maxilla quadrilateral, distance
between the parallel sides about equal to the length on meson;
prothoracic spiracles with slightly raised roughened margins;
first pair of wings with their anal angles nearly opposite the
caudal margin of the fourth abdominal segment; sutures between
abdominal segments 1 to 4 distinct, cephalic margin of sutures
approximately smooth, caudal margin of sutures irregularly
corrugated and on the fourth segment depressed, suture between
segments 7 and 8 not deep, the caudal margin of the seventh
segment slightly raised above the eighth segment; abdominal
segments 5 to 7 produced into thin flange-like plates, the mar-
gins slightly undulate, a distinct furrow at the caudal margin
adjoining the transverse conjunctiva, cremaster triangular, the
296 Annals Entomological Society of America _[Vol. VII,
distal end covered with sharply recurved spines. Length
7-8’—1"; girth about 114”. .
Described from one male specimen, for which we are indebted
to Dr. Wm. Barnes, of Decatur, Illinois.
Genus Pseudohazis Grote and Robinson.
Median thoracic line distinct on the prothorax and meso-
thorax, faint on the metathorax; first pair of wings with the
anal angles broadly rounded, near the cephalic margin of the
fourth abdominal segment; second pair of wings visible along
entire dorsal margin of first wing, its margin entire, but never
produced beyond the anal angle of first pair of wings and never
visible in ventral view; spiracular line straight; cephalic margins
of abdominal segments 5 to 7 produced into thick, oblique,
flange-like plates directed caudad; suture between the seventh
and eighth abdominal segments deep, the cephalic margin
with distinct crenulations along both margins, the cephalic
margin with quadrangular depressions, the caudal margin
with deep longitudinal furrows; cremaster short, bearing a
fan-shaped group of long straight setae.
This genus and species have been described from a single
specimen kindly loaned by the American Museum of Natural
History through the kindness of Mr. J. A. Grosbeck. Unfor-
tunately the specimen had lost its prothorax, face-parts, and
all appendages except. the wings. These descriptions are in-
cluded, however, to show the relationship of this genus to the
genus Hemileuca. Little is known of its life history, but
it spends its pupal life in the ground. There are three species
named in Dyar’s ‘‘List of North American Lepidoptera,”
all from the western part of the United States.
Pseudohazis eglanterina Boisduval. Color dark reddish
brown; exposed surface of thorax, wings and abdomen coarsely
shagreened; abdominal segments 5 to 7 with their flange-like
plates shagreened like the remainder of the segment, except
for a few faint longitudinal striations near the margin; abdom-
inal segments 4 to 8 with a raised transverse line near the caudal
margin of the segment; cremaster about one millimeter in length,
indefinitely rugose, conical, bearing a fan-shaped group of
coarse, straight setae. Length, abdomen expanded, about
1 1-8”; girth 114”.
1914] Pupe of Ceratocampide and Hemileucide 297
Genus Automeris Hiibner.
Face-parts not noticeably elevated above the body surface;
antennae pectinate throughout, tapering gradually to a point
at the distal end, the stem of the flagellum never noticeably
raised above the level of the pectinations; sexual differences,
if any, very slight; invaginations for the anterior arms of the
tentorium obscure; eye-pieces both present; clypeo-labral
suture usually distinct; maxillae, measured on meson, never
more than one-sixth the length of the wings, triangular in out-
line; less than half the exposed tibiae and the tarsi of the first
pair of legs and tips of the second pair adjacent on the meson;
second leg visible for almost entire tibial and tarsal length;
median thoracic line faint, and seldom found on all segments;
first wing with anal angle broadly rounded, near the cephalic
margin of fourth abdominal segment; second wing visible
around the entire dorsal margin of first wing, its margin entire
and produced around anal angle of first wing to form a promi-
nent angle on the fourth abdominal segment, scarcely visible
in ventral view; spiracular line slightly curved ventrad; cephalic
margins of abdominal segments 5 to 7 with sharp, transverse
ridges having distinct furrows between, and produced into an
oblique flange-like plate, generally hidden when segments are
retracted; abdominal segments 8 to 10 taper gradually to caudal
end; cremaster always distinct and set with hooked spines.
This genus includes perhaps more than a dozen species in
North America of which four species are described here. These
all spin coccoons. Our common species, A. 70, which is found
all over the Eastern United States and Mexico, spins a thin
brown ‘“‘papery’’ cocoon much like Tropaea luna, but thinner
and more shapeless. They are found on the ground, usually
with a protecting leaf attached and are thin enough so that the
pupa may usually be seen through the cocoon. A. pamina is
described from Arizona and Mexico. Its cocoon is much like
that of A. 20, with many small leaves securely fastened to it.
The cocoon of A. tncarnata of Mexico is very similar to the
preceding forms, but thicker and covered with leaves. The
cocoon of A. Jeucana is shaped much like that of Samia cecropia
and covered with small pieces of bark. It is also a Mexican
species. These four species can be separated by using the
following table:
298 Annals Entomological Society of America [Vol. VII,
A. Cremaster triangular, at least two millimeters long, with a transverse
row of hooked spines curving dorsad; cephalic margins of abdominal
segments 5 to 7 produced into an oblique, flange-like plate with an
undulate margin produced into prominent curves dorsad of the
spiracular: Iie soins Se See nay Poe eee an ee ene ee leucana
AA. Cremaster never triangular, usually only a button-like constriction
with a thickly set group of strongly recurved spines, the tips curving
outward in all directions; cephalic margins of abdominal segments
5 to 7 produced into an oblique flange-like plate with its margin
entire, never produced into curves dorsad of the spiracular line.
B. Mesothorax with fine indeterminate transverse striations; body
SEUACiCONSPICUOUS), a. crs Se areca ee eee ee ee re io
BB. Mesothorax never with fine indeterminate transverse striations;
body setae inconspicuous.
C. Mesothorax rugose; a small tubercule each side the meson
on the metathorax and first three abdominal segments.
pamina
CC. Mesothorax tuberculate with blunt conical projections;
never with small tubercules each side of the meson on
the mesothorax and first three abdominal segments.
incarnata
Automeris pamina Neumoegen. Color dark brown; body
setae inconspicuous, light brown, few in number; face parts
and appendages with fine, indeterminate transverse striations;
exposed surface of thorax rugose, remainder of surface finely
shagreened; length of antennae in both sexes more than four
times the breadth and ending in line with the tips of the first
pair of legs; labrum variable, length and breadth approximately
equal, usually six-sided and pointed at distal end; maxillae,
measured on meson, about one-sixth the length of the wings,
triangular in outline, median length greater than the greatest
width; cephalic margins of abdominal segments 5 to 7 with
fine ridges, becoming indistinct on the meson of both dorsal and
ventral surfaces, the margin produced into a flange-like plate
with its margin entire, never produced into prominent curves;
dorsal surface of abdominal segments 4 to 7 with a smooth,
elevated line just cephalad of the junction of segment and
transverse conjunctiva, extending laterad and ending beyond
the spiracles on ventral surface; dorsal and lateral surfaces
of tenth abdominal segment rugose with irregular, longitudinal
depressions at the base of cremaster. Cremaster short, con-
stricted slightly at base and forming a rounded protuberance
with a closely set group of strongly recurved spines, the tips
turning outward in all directions. Length, abdomen expanded,
from 1 1-8” to 114"; girth about 134”.
1914] Pupe of Ceratocampide and Hemileucide 299
Automeris io Fabricius. Color dark brown; body setae
conspicuous, light brown, sparsely distributed over entire
surface excepting appendages, most numerous on thorax;
body often noticeably depressed; face parts, appendages,
except the wings, and exposed surface of thorax with fine,
indeterminate, transverse striations, remainder of surface
shagreened, with the projections in transverse rows; antennae
in both sexes with length three times the width and quite
reaching the tips of the first pair of legs; labrum variable,
broader than long, usually five-sided and pointed at the distal
end; maxillae, measured on meson, about one-sixth the length
of wings, median length always less than the greatest width,
each half the maxilla quadrilateral, sometimes modified so that
entire maxilla appears heart-shaped; median thoracic line
narrow, usually visible on all segments; abdominal segments
5 to 7 with the cephalic margins covered with sharp transverse
ridges, with distinct furrows between, the furrows becoming
shallower at the meson on the ventral surface, the flange-like
plate with its edges entire; abdominal segments 4 to 7 with a
distinct furrow of varying width between the segment and the
transverse conjunctiva, which becomes indistinct in the region
of the proleg scars on the ventral surface, its cephalic margin
being indicated by a raised line; abdominal segments 8 to 10
with segmentation distinct; dorsal surface of tenth abdominal
segment with deep, longitudinal ridges at base of cremaster;
tip of cremaster with a small group of closely set, sharply
recurved spines, the hooks turning outward in all directions.
Length, abdomen retracted, 7-8’—114", expanded, 1”—1 3-8’;
girth 134”—2”.
Automeris leucana Hubner. Color dark brown; body
setae light brown, inconspicuous; face parts and appendages.
with indeterminate, transverse striations, exposed surface
of thorax rugose, with interrupted transverse ridges; remainder
of surface coarsely shagreened; antennae in both sexes with
the length more than four times the breadth, not extending
as far caudad as the tips of first pair of legs; labrum variable,
length and breadth approximately equal, pointed at tip, usually
five-sided; maxillae, measured on meson, about one-seventh
the length of wings, the greatest width about one and one-half
times the median length, each half the maxilla quadrilateral;
300 Annals Entomological Society of America [Vol. VII,
median thoracic line very narrow, only distinct on the meso-
thorax; abdominal segments 5 to 7 with the cephalic margin
ridged, produced into an oblique flange-like plate with an
undulate margin having prominent curves dorsad of the spiracu-
lar line, the median line of cephalic margin indicated by oblique
ridges, a slightly raised, smooth line cephalad of the junction
‘of the segment and the transverse conjunctiva; tenth abdominal
segment having the dorsal and lateral margins of the cre-
’ master with semi-longitudinal ridges at base of cremaster;
cremaster at least two millimeters in length, triangular in out-
line, tapering rapidly to a pointed tip with a transverse row of
sharply recurved spines, the tips curving dorsad. Length,
abdomen expanded, 114”—1 5-8"; girth about 134”.
Automeris incarnata Walker. Color dark brown to blackish,
transverse conjunctiva lighter; body setae light brown, incon-
spicuous; face parts and appendages with wavy, indeterminate,
transverse striations,. exposed surface of thorax tuberculate
with blunt, conical projections; antennae in both sexes with
length about four times the width and ending opposite the tips
of the first pair of legs; labrum variable, broader than long,
usually five-sided, pointed at distal end; maxillae, measured on
meson, about one-sixth the length of the wings, median length
less than the greatest width, each half quadrilateral, lateral
margins concave, basal half sculptured and roughened; median
thoracic line wanting except on metathorax; dorsal and lateral
portions of cephalic margins of abdominal segments 5 to 7
with fine, sharp, transverse ridges becoming indistinct in the
region of the proleg scars, the cephalic margin narrower in this
region and produced all around segment into a very narrow,
flange-like plate with a distinct longitudinal impression at
meson; abdominal segments 4 to 7 with a raised line cephalad
of the line between the segment and the transverse conjunctiva;
tenth abdominal segment rugose at base of cremaster; the
cremaster short, rounded, constricted at base and set with a
small group of closely set, sharply recurved spines, the tips
turning outward in all directions. Length, abdomen con-
tracted, about 1”, expanded, about 1 1-8”; girth about 114”.
NOTES ON THE LIFE HISTORY AND ANATOMY OF
SIPHONA PLUSLE Cog.
By WILitAM BLOESER, Stanford University, Calif.
LIFE HISTORY.
The Tachinid fly, Stphona plusie, was described by Coquillet
in 1897. It was bred from a cut-worm. The specimens that
I have obtained, however, were parasitic in the larve of Phry-
gamdia californica, gathered from oak trees at Stanford Uni-
versity.
The Phryganidians were more than plentiful during the
fall of 1913, and consequently there was an abundance of par-
asites. Sztphona is only one among a dozen or more parasites
that are nursed in their infancy by the accommodating Phry-
ganidian, but notwithstanding the ravages of all these parasites,
and the scourge of a fungus disease, which killed nearly one
third of the caterpillars, there were still many left, sufficient to
insure a great number of moths again in the following spring.
The following notes on Szphona plusie are the result of
observations made in the fall of 1913:
The Egg. The adult female fly lays one or more eggs on
the outer body wall of the Phryganidian larva. The dipterous
parasites are not as careful as the hymenopterous parasites, and
they lay their eggs indiscriminately, often laying three or four
eggs on one host.
The Larve. After the eggs have hatched the young larve
make their way into the body cavity of the Phryganidian,
where they remain from ten days to two weeks, feeding on their
host until fully grown, when they measure about five-sixteenths
of an inch in length. They have eleven segments; well devel-
oped mouth parts, in the form of great hooks; two large pos-
terier spiracles and two smaller anterior ones.
The larve are loosely attached or held in a sort of cicatrix,
in the body of the host, by several rows of small hooks that
encircle the tenth and eleventh segments. From this position
the head and anterior portion of the body are free to swing in
the body cavity. Some larve are found, however, moving
301
302 Annals Entomological Society of America [Vol. VII,
about freely in the body cavity, while those that were attached
could be easily removed or could themselves change their
position.
About one hundred Phryganidia were dissected and ten
Siphona parasites were found, three of these being taken from
a single caterpillar. It would be hard to estimate with much
accuracy the probable percentage of parasites, but ten per
cent, I believe, would not be too high an estimate.
Some of the Phryganidia were kept alive in a cage, and from
these there issued several fly larvee, which pupated in about two
hours. In no instance did the parasites issue from Phryganidia
pup, but all seem to leave the Phryganidia while the host is
still in the larval stage. After freeing itself from the host the
larva soon begins the period of pupation. It begins by drawing
itself together and changing to a darker color, and within a
couple of hours it is a brown segmented pupa about three-
sixteenths of an inch in length. One pupa remained from the
sixth of October to the twenty-fourth, a period of eighteen
days, before the imago finally appeared. Other larve were
alllowed to pupate, but from eight pupz only the one fly issued,
while from the seven others, there issued hymenopterous
hyperparasites, which have not yet been determined. These
issued somewhat later, taking twenty-three to twenty-five
days to come from the pupa cases. .
This percentage of hyper-parasites is almost certainly more
than the average, as they came from Phryganidia that were
gathered from a single oak tree situated in a flower garden. It
is to be hoped that further investigation will reveal a smaller
percentage of hyper-parasites, as their abundance will greatly
check the beneficial work of Szphona plusie, which has so greatly
aided in controlling the Phryganidia in California, especially
in the Santa Clara Valley.
The Adult. The adult has been described by Coquillet,*
but practically nothing of the life history has been heretofore
given. The general characteristics of the adult are shown in
text-fig. 2, a special character being that the proboscis has.
two geniculations, one near the base and the other near the
middle.
* Canadian Entomologist, Vol. 27, p. 125.
a7
1914] Life History and Anatomy of Siphona Plusie 303
ANATOMICAL NOTES.
External Appearance. The larve are white and nearly
translucent, and the colors exhibited are at either end. The
great hooks (text-fig. 1) which form the most important
part of the mouth structure, are jet black. On the last segment
there are two large posterior spiracles, which are of a deep
brown color. There are also several rows of little dark colored
hooks around the tenth and eleventh segments (text-fig. 1).
Fig. 1.
Dorsal view of full grown larvae.
Ventral view of first segment; a, antennae (?); h, great hooks.
Dorsal view of last two segments, showing rows of little hooks and s, posterior
spiracles.
Lateral view of first segment; h, great hooks.
Dorsal view of pupa.
ie eee.
The opening of the two large trachee at the anterior end
are less plainly visible. The main tracheal trunks narrow to-
wards the head, and each branches out into two fine tubes which
terminate in small spiracular openings at about the beginning
304 Annals Entomological Society of America [Vol. VII,
of the second segment. These, however, disappear after the
larva has made two or three moults, and there are no longer
any anterior spiracles.
At the extreme tip of the first segment, on either side of the
great hooks, there is a pair of wartlike processes, as shown in
text-fig. 1. These are probably rudimentary antenne.
Internal Anatomy. The alimentary canal and Malpighian
tubules, (shown in Plate XL, Fig. 1.) are quite characteristic, in
their many turns and loops, of Dipterous larve in general,
especially those of the Muscid kinds. The parasitic life of the
larva seems to have resulted in no considerable structural
modification of the digestive system.
Fig. 2.
Adult fly, dorsal view, showing general characteristics; also lateral view of
head, showing proboscis with two geniculations, one near the base and the other
near the middle. (Greatly enlarged).
The oesophagus, starting at the mouth, extends backward
as a narrow cylindrical tube, passing through the supracesopha-
geal ganglia, or brain, and then passing above the ventral
ganglion and entering the proventriculus, which lies in about
the fourth segment of the body. From the proventriculus, the
mesenteron, or portion of the canal from the proventriculus to
1914} Life History and Anatomy of Siphona Plusie 305
the Malpighian tubules, is a nearly uniform tube of considerable
size, the anterior portion being the chyle stomach and the
posterior portion, the intestine.
The Malpighian tubules in this insect are particularly
interesting in regard to their position in the body cavity. They
arise from the alimentary canal, as shown in the drawing, as
two lateral tubes, each of which divides into two tubes. The
two from the right side swing forward and the two from the left
side run towards the posterior end. This is somewhat different
from what would be expected, and is a departure from the
general rule. The usual number of tubes is four in the dipterous
larve, but all four either turn and run posteriorly, keeping to
their respective sides, as in the blow fly, or else the right and
left branch, each sending one tube forward and one backward.
The portion of the alimentary canal from the entrance of the
Malpighian tubules to the anusis the metenteron. This portion
is considerably smaller and shorter than the mesenteron and
has a thick muscular coat.
The dorsal blood vessel or heart; the tracheal system; the
nervous system, and the salivary glands, are shown in plate
XL Pip. <2.
The heart is a thin-walled muscular tube which extends
nearly the length of the body, lying in the pericardial cavity
just beneath the dorsal wall. It tapers from a good-sized sac
to a fine tube as it runs forward.
The tracheal system is composed of two main trunks with
large spiracles opening on the posterior segment. Branches
are given off from the two main trunks at each body segment
and these finer tubes wind in about the alimentary canal. The
anterior spiracles are wanting, except in the very young larve.
The salivary glands, which extend from the mouth, starting
as a single narrow duct, branch out beneath the pharynx and
extend, one on either side of the alimentary canal, for more
than a third of the length of the body.
The brain and body ganglion, shown in the same figure,
compose the nervous system of the larva. The hemispheres
encircle the oesophagus just forward of the proventriculus, and
the main body of the body ganglion extends backward on the
ventral side nearly the same distance that the salivary glands
extend on the lateral sides.
306 Annals Entomological Society of America [Vol. VII,
The muscles and fat cells are conspicuous, but do not differ
particularly from those of other dipterous forms.
I limit my description of the anatomy to the fewest words
possible, as the figures and plates tell the story sufficiently. The
interesting thing about the anatomy of the larva is that it is so
little different from that of any free-living, outside-feeding dip-
terous larva of Muscid type. Either the parasitic habit makes no
less demands on alimentary canal, respiratory, circulatory and
secretory systems than the free life habits, or this insect has so
recently adopted a parasitic habit that no considerable struc-
tural changes in its organs have yet been brought about in
connection with it.
This paper was prepared in the Entomological Laboratory
of Stanford University.
1914]
Life History and Anatomy of Siphona Plusie 307
EXPLANATION OF PLATES.
Abbreviations used:
Big,
Fig. 2.
Bigs:
Fig. 4.
Fig. 5.
Fig. 6.
Bigs 1.
Fig. 2.
PRTIGEMM GE Ct. Snakes Siem a wea Metenteromes ss... .s:.. ssn Ye
JENVALB TS tt Abe Eten gad ant es A a A IMISG eyAe cee 5. oo Se L
Rattler ons wanna eto BodysGangliany. 2. .ce N
Hate Cell ses eo eres F @esophdcuste 4.25. O
(Greate Hooks: nn. .5 50 IRroventiiculticnes. 5-502
Peart erp ns oso H Salivary Glands.........G
imeasinalmDiscsase 4) yt 5.1 DDIDACLES roca nce aot oie
Malpighian Tubules......M bra chaerne see othe acer 1p
IMesenterome. se.n\a.aic) srs x
PLATE XXXIX.
Cross sections through larvae.
Section through anterior portion, about the second segment; T, trachea;
I, imaginal discs; G, salivary glands; O, oesophagus; L, muscles.
Section through neuroblast, about the third segment; H, heart; T, trachae
O, oesophagus; B, brain; N, sub-oesophageal ganglion; G, salivary
glands.
Section through proventriculus, about the fourth segment; H, heart;
F, row of fat cells; T, trachea; P, proventriculus; N, ganglion, G,
salivary glands.
Section through many folds of the alimentary canal, about the fifth
segment; H, heart; T, trachea; M, Malpighian tubules; C, alimentary
canal; L, muscles.
Section showing Malpighian tubules branching from alimentary canal,
about the eighth segment; H, heart; T, trachea; M, Malpighian tubules;
C, folds of alimentary canal.
Section near posterior end; T, trachea; M, Malpighian tube; C, alimentary
canal; A, anus.
PLATE XL.
Horizontal longitudinal view of larva, showing, O, oesophagus; P, pro-
ventriculus; X, folds of the mesenteron; Y, the metenteron; M,anterior
and posterior Malpighian tubules.
Vertical longitudinal view of larva, showing, S, spiracle; G, salivary
glands; B, brain; N, ganglion; F, row of fat cells; T, trachea; H, heart:
L, muscles.
ANNALS E. S. A. VOL. VII, PLATE XXXIX-.
Wm, Bloeser
Vou. Vil, PRATE 2,3
ANNALS E.S. A.
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aaa =e ae
7
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met ese YE ETAEAEE
Be 6B) YD DB
Wm. Bloeser
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Ue
SOME NOTES ON DIGESTION AND THE CELL STRUC-
TURE OF THE DIGESTIVE EPITHELIUM
IN INSECTS. (Plate XLI.)
By E. J. NEwcomer, Stanford University, California.
The ultimate structure of the cell, as it is understood in
animal and plant life, is still largely a matter of theory among
biologists. A cell appears as a tangible unit, apparently
definitely set off from its fellows, and easily discernible with
low powers. Yet its exact constitution and its exact relation-
ship to the surrounding cells are not known. So far, scientists
have had to imagine how such a structure, with the known
functions that it possesses, ought to be composed; it has not yet
been possible by actual observation to prove this composition. .
Many things enter into this difficulty. Cells are very small
structures and cannot be viewed with the naked eye. The
interposition of a lens or lenses increases the possibility of error.
Especially when using lenses of high power is there very little
certainty about what is seen. The difficulty of seeing cells in
their natural state is great, and resort is had to material that is
killed and fixed. Here, even though one is sure of what he
sees, he cannot be certain that it is the same as in life. Again,
the diversity of cells in different organs and organisms is il-
limitable, and it is known that even in the same cell the cyto-
plasm can change its appearance: so that the problem of getting
at the typical and final structure and the behavior of this cyto-
plasm is by no means an easy one.
It was with the hope of possibly finding out some facts that
might have a bearing on the general problem of animal cell
structure and behavior that I have undertaken a special study
of the make-up and behavior of the digestive epithelium in
insects, which study permits me to offer the statements and
illustrations embodied in the present paper. The digestive
epithelium of insects is notable as a cell layer in which rapid
changes occur, with a continual production and destruction
of cells. Hence it may be presumed to be a tissue in which
cell origin and growth may be advantageously studied.
205
al2 Annals Entomological Society of America [Vol. VII,
Incidentally, the behavior of the digestive canal of insects
is only imperfectly known, and though I have nothing to add
either to the various cell-structure theories or to the theories
concerning digestion, yet these notes recording what seem to be
the actual histologic conditions of the digestive epithelium in a
number of insects, and suggesting some possible significance of
these conditions, may have a little interest.
First, let us take up the matter of cell structure and cell
relationship. The old idea that cells were like so many bricks,
each to be considered separately, has had to be discarded; and a
multicellular organism can no longer be compared to a brick
building. At first sight this may seem to be a proper com-
parison, but reflection will show that the differences are great.
For the cells of an organism are by no means constant; they
are continually being built up and destroyed. A cell is injured,
or it completes the work for which it was created: it disappears,
and a new one takes its place. Cell walls are not mere rigid
boundaries; they are elastic, permeable, capable of radical
change. These things all go hand in hand with a specialization
and consequent interdependence of cells, which makes impera-
tive a study of the behavior of many cells rather than of single
cells.
A study of insect digestive cells soon makes it apparent that
here a definite and constant cell structure does not exist. Diges-
tive cells are extremely active. Two important types of cell
secretion may be distinguished, called by Haseman the holo-
crine and merocrine types. In the former the whole cell con-
tents is discharged at once, and new cells arise. In the latter
the discharge is gradual and continuous and the cell remains.
active for along period. I shall take up this matter of secretion
more fully in connection with the subject of digestion. The
important thing here is to note the marked difference in be-
havior between the two types of cells.
The holocrine type is very well represented by the digestive
cells of the dragon-fly (Needham), or by those of almost any
predaceous beetle (Fig. 6). Here the cells are formed in nests
or nidi, using Needham’s term, and gradually develop and
increase in size until they are capable of secretion. Upon the
introduction of stimulus in the form of food, the largest cells
burst, pouring out their contents to mix with the food, and the
1914] Digestion and Digestive Epithelium in Insects 313
cells in size take their places. Thus we have a regular progres-
sion of cells from the very small and scarcely distinguishable
ones in the nidi to the full-grown secretive cells. The most
interesting point here in connection with the study of cell
structure and behavior is the existence of the nidi. What
these are and how they originate is a question. They have
been variously called “‘cryptes,’’ ‘“drusenkrypten,”’ and “epi-
thelial buds.’’ None of these terms is very specific, and the
idea of these bodies being glandular can hardly be retained, fora
gland which secretes nuclei or cells is inconceivable. The nidus
appears as a group of nuclei, exactly like those in the fully-
developed cells except smaller, crowded together, and with
very little protoplasm about them. (Figs. 1 and 2, n). In
some instances this group of nuclei is enclosed in a sort of sac
protruding out into the muscles surrounding the alimentary
canal (Faussek, Frenzel, Rengel), and to this type in particular
the term ‘‘drusenkrypt’’ has been applied. More often, how-
ever, the nidus is an integral part of the wall of the canal, and
there seems to be no special limiting membrane. Are these
cell “‘anlagen’’ of which the nidus is composed split off from a
mother cell? If so, where is this mother cell? Each nucleus
of the nidus looks exactly like each other nucleus. Or is the
nidus as a whole a cell which produces these nuclei, perhaps by
division of its own nucleus? But this approaches the gland
idea. As I look at it, it is simply impossible to apply the
ordinary theories of cell constitution and cell existence to this
structure, this nidus. We must look elsewhere. Mobusz,
quoting Adlerz, mentions the presence of a network of proto-
plasm between the basement membrane and the cell bases, from
which new cells arise. We may have to advance a theory of
something similar to this to account for the origin of the nuclei
in the nidi. If they are not formed by division from others,
can they by any possible means be formed from a net of
protoplasm? A further and more careful study of these nidi
is essential, and will undoubtedly throw light on the general
question of cell origin.
Let us turn to the merocrine type of digestive cells, that is,
the type where the cell contents is only partly dicsharged as a
digestive fluid. This type is to be seen in the alimentary canal
of an insect that feeds continuously, thus demanding a con-
314 Annals Entomological Society of America [Vol. VII,
tinuous flow of digestive fluid. Haseman describes the larva
of Psychoda as possessing this type of cells. It is also the type
present in Lepidopterous larve, such as the silkworm (Fig. 11),
or in the Coccide (Fig. 4), which after once settling down,
remain attached to their food-plant, and continuously suck in
the plant juices. A study of these cells makes it clear that,
although no nests of nuclei are present from which the cells are
replaced, nevertheless: the cells are replaced. Haseman has
very carefully worked this out in the case of Psychoda and
finds that the cells are replaced at the molting period. The
old cells degenerate and slough off, and new ones, which he
calls regenerating cells appear along the basement membrane.
Haseman describes the growth of these cells, but makes no
attempt to explain their origin. At once Adlerz’ notion of a
basal protoplasmic network suggests itself. For, to judge from
Haseman’s drawings the old cells degenerate completely and
no part of them composes the new cells, except, as Haseman
mentions, that some of the old material may be absorbed by
these cells. We must look elsewhere, then, for their origin,
and it is not incomprehensible that some sort of basilar proto-
plasm may exist, from which these cells spring. My prepara-
tions of the silkworm are unfortunately not numerous enough
to show all stages of this degeneration and replacement, but
it seems evident that it takes place here, in a measure as it does
in Psychoda. Fig. 7 is from a sagittal longitudinal section of a
young silkworm killed just before molting. There are many
large, distended cells (d. c.) which appear to be pouring out
their whole contents, but as none of these protruding droplets
has ever been found detached, it seems more reasonable to
suppose that the cells are degenerating, and that the protrusion
is an artifact produced by improper killing. Between these
cells are others (ab) with a basal nucleus and a clear lumen.
This lumen I believe also has been produced artificially, but
aside from this these cells are quite different from the others,
and may perhaps be absorptive or mucous cells. Along the
basement membrane are numbers of small cells (r. c.), each with
a nucleus or occasionally two. These cells I take to be the
regenerating cells. Fig. 8 shows a cross section near the base-
ment membrane of this same epithelial layer. The three
types of cells appear distinctly.
ie
meha €-
1914] Digestion and Digestive Epithelium in Insects 315
In Fig. 11 we have in section a portion of the epithelium of a
larva that has just molted. Here the large, loosely composed
cells are not in evidence, and many of the small basal cells have
grown out until they reach the intima. The other type of cell
is present also, but is not shown in the drawing. The nodules
projecting from the cells here and in Figs. 9 and 10 are interest-
ing in that they seem to have pushed through the intima in-
stead of having stretched it as appears in Fig. 7. They may be
artifacts, or more likely they are drops of digestive fluid, such
as van Gehuchten has described and figured in Ptychoptera,
though I have never found them floating free as he has shown
them.
The larva of Dendroctonus, a Scolytid beetle, which bur-
rows into the living wood of pine and other coniferous trees,
affords a good example of an insect which feeds continuously,
and hence must possess digestive cells which gradually and
continuously pour out their secretions. Here the cells are
exceedingly regular, each one like the next. There are no nidi
to be seen, and no protruding portions are present. The
secretion evidently oozes gradually through the intima in small
droplets. The only good preparation that I have shows the
basal half of these cells to be very compact and darkly staining
while the distal half is open and loose. The nuclei are situated
just at the bounadry between these two halves of the cells.
This particular larva appears, to judge from the condition of
the cuticle, to be upon the point of molting, and this division
of the cells may be similar to that which Folsom and Welles
have described in Collembola.
The digestive epithelium of the Coccide, as represented by
Lecanium, is very simple (Fig. 4). It consists of a row of more
or less regular cells, with here and there one which is greatly
enlarged. These large ones are evidently the active, secreting
cells, while the smaller ones are developing. Frequently these
contain two nuclei, indicating that they are formed by direct
division. It would be interesting to see what happens to these
cells at molting time, but as the Lecaniums only molt twice
(Quayle), and these moltings come while they are still quite
small, it would be rather difficult to get preparations.
The digestive phenomena of various insects have been
mentioned above briefly and I now propose to take up in order
316 Annals Entomological Society of America [Vol. VII,
the insects I have studied, and give more fully some notes
regarding this process, and describing the epithelial cells.
I shall consider, first, however, the Isopods, which are not
insects but Crustaceans.
Murlin, in an excellent paper on the digestive system of the
Isopods, shows that here most of the digestive fluid is secreted
in a separate organ, the hepatopancreas, the giant cells of which,
however, pour out the secretion in much the same manner as
those of the digestive epithelium of many insects. The Isopods
have proved to be a very interesting and valuable group with
which to begin such a study, both because of the simplicity of
their organization, and ease of preparing material, and also
because of the very large size of the cells.
The Aptera, the lowest group of insects, will always be the
source of a great deal of information regarding digestion, and a
thorough study of such forms as Lepisma and Japyx would be
valuable. Campodea, I found, has a digestive apparatus very
similar to Collembola, as worked out by Folsom and Welles,
even down to the apperance of the cells. Japyx (Fig. 1) is
quite different. Here the mid-intestinal cells are very open,
and have a somewhat alveolar appearance. The cell contents
is irregularly granular, and contains numerous large clear
vacuoles of varying size, which sometimes compose nearly the
entire contents. Here and there are scattered dark granules
which probably are concretions. The nuclei are small and
basally situated, and stain almost black with iron hematoxylin,
while in preparations stained with Ehrlich’s acid haematoxylin
they are nearly invisible. The cells evidently arise from nidi,
although the latter appear to contain more definite cells than
is usual. The intima is very thin, and sometimes slight amounts
of secretion may be observed in the digestive lumen along the
intima.
Lepisma has an extremely interesting digestive system,
including a remarkable muscular crop, the posterior end of
which protrudes into the mid-intestine. Here the same con-
dition exists which Needham describes as occurring in the
Odonata, but in lesser degree (Fig. 2). The active secreting
cells, two to four in a group (s. c.) are very sharply marked
off from the young, forming cells (y. c.), staining much darker,
particularly with the iron hematoxylin stain. At the base of
1914] Digestion and Digestive Epithelium in Insects al?
the lighter cells the nidi (n) are to be found, mere rough groups
of nuclei containing each a nucleolus and many granules or
perhaps alveoles of varying sizes. The developing cells stain
very slightly and frequently contain vacuoles. The active
cells are longer, and, on account of the secretion which they
contain, stain darkly. Both kinds have a fibrillar or palisade-
like appearance basally, which extends as far as the nucleus.
The inner portions of the active cells are alveolar, or possibly
composed of a network, and contain many small highly re-
fractive concretions. There are no distinct cell walls, but there
is a periodical thickening of the fibrils, which give the cells a
distinct appearance. The intima is moderately thick and
traversed by pore canals. Frequently numerous droplets of
secretion (sec) may be seen between this and the peritrophic
membrane, and sometimes the secretion appears to be streaming
from the active cells. In the region just behind the large crop
the cells are smaller and more compact.
The termites have a very peculiar digestive epithelium which
perhaps can be correlated with their habit of feeding on dead
wood. The stomach is bordered with from ten to twenty
lobe-like projections, one of which is shown in Fig. 3. Each
of these has at its base a nidus of many nuclei, and extending
from this to the inner tip of the lobe, the cells overlap each
other in a very curious scale-like manner which is evidently
only a variation of the typical holocrine method of cell-forma-
tion. It is noteworthy that this method should occur in the
termites, which live in the wood they feed upon, and-at least
have the opportunity of feeding continously, whether they
actually do or not.
In the order Hemiptera, I have only studied the rather
abnormal Coccide (Fig. 4), which I have already mentioned.
It looks here as though the large cells discharged their contents
and were replaced by the smaller ones, which are formed by
simple cell division, though I have not observed the process.
In contrast to this arrangement we find in Myrmecophila, a
small degenerate cricket (Orthoptera) inhabiting ants’ nests
(Fig. 5), the typical nidi (n), with the regular wave-like ar-
rangement of nuclei between them. Here, besides the intima
with its pore canals, there are what appear to be cilia, and at
their ends are small droplets of secretion.
318 Annals Entomological Society of America [Vol. VII,
I have described the appearance of the epithelial cells in one
beetle larva, Dendroctonus. Another larva which I have
sectioned, that of a Carabid, is entirely different. The cells”
are arranged in lobe-like groups with a nidus (n) at the base
between each two groups, from each side of which the cells
arise and gradually grow and migrate until they become full-
sized, when the contents is discharged and others replace them.
The larger cells are extremely vacuolate, and irregularly granu-
lar basally and distally. The nuclei are fairly large and deeply
staining.
The Diptera have been studied by various investigators.
Van Gehuchten, in his complete work on Ptychoptera, a Tipulid,
was one of the first to point out the method of digestive secre-
tion in insects. Haseman’s recent paper on Psychoda describes
the conditions occurring in another group, with habits not un-
like the Tipulids. It is apparent that in larve such as these
which live practically submerged in their food, the merocrine
type of secretion prevails, and the arrangement of the cells
secreting in this manner is manifestly entirely different from
that representing the holocrine type. This latter, which we
have seen in the Carabid larva, and elsewhere, demands cells
capable of storing up the digestive fluids until such time as
food may be taken into the canal, for predatory insects neces-
sarily get their food irregularly.
The silkworm is distinctly a continuous feeder. Hence we
should expect to find no nidi or nuclei present. At first glance
it seems otherwise (Fig. 7) but a closer scrutiny reveals the fact
that these apparent groups of nuclei are quite different. In the
first place there are fewer nuclei in a group than is usual, and
then they are strictly not groups of mere nuclei, but groups of
small cells. When we realize, too, that this particular insect
was just upon the point of molting, we conclude, as I have pre-
viously shown, that these are the new cells which form to re-
place the old ones sloughed off at the molting period.
But let us examine a just-hatched larva, which has taken no
food except the portions of the egg-shell devoured in hatching.
The cells are exceedingly regular, and none of the small basal
cells to be observed. Most of the cells are deeply staining,
granular, with an elongate, central, granular nucleus, and distal-
ly containing a few small vacuoles. Frequently another type oc-
curs, lighter, more homogeneous, with basal, rounded nuclei.
1914] Digestion and Digestive Epithelium in Insects 319
These I believe are the absorptive cells and correspond to the
cells marked ab. in Fig. 7. Of course this larva has had no plant
food yet, but we must suppose that these cells contain some-
thing besides food products. As a matter of fact they are
much leaner than in older larve except for the basal part
containing the nucleus, being scarcely distinguishable between
the secretory cells. These are beginning to form the secretion,
though none of them has any protruding droplets. Larve
sectioned twenty-four hours after beginning to feed, show some
food in the alimentary canal, and some of the secretory cells
are giving off small drops of fluid. As the larva grows these
droplets increase in size until they appear as in Fig. 10 or 11.
The contents of the absorptive cells is homogeneous, and takes
the Orange G stain precisely as does the contents of the silk
glands, which indicates that it is a product of digestion, as the
liquid silk is merely this same product somewhat transformed.
The changes in the cells during molting I have already des-
cribed.
As a killing fluid I found nothing superior to Carnoy’s
fluid, which is a mixture of 6 parts of absolute alcohol, 3 parts
of glacial acetic acid, and one part of chloroform. For general
insect work it is entirely satisfactory. The chloroform very
quickly dissolves any wax or grease, and allows the insect to
sink. It acts quickly, and is very simple to use. The speci-
mens are merely dropped into it, left for a couple of hours, and
transferred to alcohol, first 70% and then 85%. There was
some distortion and occasionally shrinkage, but I am inclined
to believe that this latter was due to some fault in the subse-
quent treatment. Tower’s, Gilson’s, hot water, etc., gave no bet-
ter results, and are not so easily handled. I tried injecting a
number of silkworms with the killing fluid, just after dropping
them into it, but could see no difference between these and those
not injected. The specimens were run through alcohol and im-
bedded in the ordinary way. Sections were cut as thin as
possible. I tried a variety of stains and found combinations
either of Ehrlich’s acid hematoxylin or iron hematoxylin with
Orange G to be the most satisfactory. Occasional preparations
stained with carmine and Orange G brought out points not
observable in others. The Orange G is a very desirable second-
ary stain, as it stains tissues which otherwise would remain
almost colorless.
320 Annals Entomological-Society of America _[Vol. VII,
The work here recorded was done and the paper prepared
in the Entomological Laboratory of Stanford University by
me as holder of the Bernard Scholarship for 1913-1914 in
Insect Histology.
BIBLIOGRAPHY.
Bernard, H. M. Some Neglected Factors in Evolution. 1911.
Bordas, L. L’Appareil Digestif des Orthoptéres, Ann. Sci. Nat. Zool. ser. 8,
t. 5, 1898, pp. 1-208, plate.
Cuenot, L. Etudes physiologiques sur les Crustacés Decapodes, Arch. Biol.
t. XIII, 1893, pp. 245-303, 2 pl. :
Cuenot. Etudes physiologiques sur les Orthoptéres. Arch. Biol. t. XIV, 1896,
pp. 293-341, 2 pl. :
Eberli, J. Untersuchungen am Verdaungs-traktus von Gryllotalpa vulgaris.
Vierteljahrschrift der naturforsch. gesell. in Zurich. 37 Jahrg. 2 Heft, 1892.
Faussek, V. Beitr. z. Histologie des Darmkanals der Insekten. Zeit. wiss. Zool.
Bd. 45, 1887, pp. 694-712, 1 pl.
Folsom, J. W. Entomology with Reference to its Biological and Economic
Aspects. 2nd. ed., 1913, p. 96, fig. 151. :
Folsom, J. W., and Welles, M. U. Epithelial Degeneration, Regeneration, and
Secretion in the Mid-Intestine of Collembola. Univ. of Ill. Univ. Studies,
vol. II, no. 2, 1906. Plates.
Frenzel, J. Einiges uber den Mitteldarm der Insekten sowie uber Epithel-regener-
ation. Arch. Mikr. Anat. Bd. 26, 1886, pp. 229-306, 3 taf.
Van Gehuchten, A. Recherches Histologiques sur l’Appareil digestif de la larve
de la Ptychoptera contaminata. La Cellule, t. VI, fasc. 1, 1890, pp. 185-289.
Plates.
Van Gehuchten, A. Contribution a I’Etude du Mecanisme de I’Excrétion Cellu-
laire. La Cellule, t. [X, fasc. 1, 1898, p. 95-116, 1 pl.
Grassi, B. I Progenitori dei Miriapodi e degli Insetti. Reale Ac. dei Liucei.
Mem. VII. Anat. Comp. dei Tisan (1888), Acc. Gioenia, Mem III. Anat.
Machilis (1885), Acc. Gioenia Mem. III. Anat. Japyx and Campodea
(1885). Torino, Morf. Scolopendrella (1885).
Haseman, L. The structure and Metamorphosis of the Alimentary Canal of the
Larva of Pyschoda alternata. Ann. Ent. Soc. Am. vol. III, No. 4, 1910,
pp. 277-313, 5 plates.
Miall & Denny. The Cockroach, 1886, pp. 121-122.
Mobusz, A. Ueber den Darmkanal der Anthrenus-Larve, nebst Bemerkungen zur
Epithelregeneration. Inaug. Diss. Univ. Leipzig, 1897.
Murlin, J. R. Absorption and Secretion in the Digestive System of the Land
Isopods. Proc. Acad. Nat. Sci. Phil. May, 1902.
Needham, J. G. The Digestive Epithelium of Dragon-fly Nymphs. Zool. Bul.
vol. 1, No. 2, 1897.
Quayle, H. J. The Black Scale. Cal. Agr. Exp. Sta. Bul. 228, 1911.
Rengel, C. Ueber die Veranderungen des Darmepithels bei Tenebrio molitor
wahrend der Metamorphose. Inaug. Diss. Fried. Wil. Univ. Berlin, 1896.
1914} Digestion and Digestive Epithelium in Insects 321
EXPLANATION OF PLATE XLI.
(All are camera lucida drawings except Figs. 3, 4 and 5).
Fig. 1. Epithelial cells of mid-intestine of Japyx. Longitudinal section. X 1160.
Fig. 2. Epithelial cells of mid-intestine of Lepisma. Longitudinal section.
X 1160
Fig. 3. One lobe of mid-intestine of termite. Cross section.
Fig. 4. Portion of mid-intestine of Lecanium. Cross section.
Fig. 5. Epithelial cells of Myrmecophila. Cross section. X 760.
Fig. 6. Epithelial cells of Carabid larva. Longitudinal section. X 760.
Fig. 7. Portion of mid-intestine of silkworm, just molting. Longitudinal sec-
tion. X 740.
Fig. 8. Cross section through same on line A-B, Fig. 7. X 740.
Figs. 9 and 10. Epithelial cells of silkworm. x 720.
Fig. 11. Portion of mid-intestine of silkworm just after molting. Longitudinal
section. X 720.
ABBREVIATIONS.
ab., absorptive cells. n., nidi.
b. m., basement membrane. nuc., nuclei.
c. m., circular muscles. Tr. c., regenerating cells.
con., concretions. s. c., secreting cells.
d. c., degenerating cells. sec., secretion.
int., intima. vac., vacuoles.
1. m., longitudinal muscles. y.c., young cells.
VOL. VII, PLATE XLI.
ANNALS E. S.A.
J. Newcomer
E
STUDIES IN THE LONGEVITY OF INSECTS.
J. PERcY BAUMBERGER.
The investigation of the effect of temperature upon insects
discussed below was undertaken at the suggestion of Professor
C. W. Woodworth of the Entomological Department of the
University of California in the fall of 1911. The purpose of the
original problem was to obtain data on the length of the imago
stage of the different orders of insects without food and under
different temperature conditions.
The present article represents the extension of the original
simple experiment to a more comprehensive study of the effects
of temperature. The specific problems studied and the con-
clusions arrived at are as follows:
Part 1. Longevity as affected by different constant temperatures:
(a) -is not correlated with systematic groups,
(b) differs inversely with these temperatures,
(c) is approximately proportional with these temperatures, and
(d) is primarily dependent upon physiological factors.
Part 2. Longevity as affected by exposure to two different tem-
peratures:
(a) is increased when the temperature of first treatment is high or
low, and
(b) is decreased when the temperature of first treatment is normal.
Part 3. Hibernation as affected by exposure to two different tem-
peratures:
(a) is not brought to a close when the temperature of first treat-
ment is normal and the temperature of second treatment is
high,
(b) is brought to a close when the temperature of first treatment
is low; is continued for from ten to twenty-one days and is
followed by a second treatment at a high temperature, and
(c) is not brought to a close when the first treatment at a low
temperature is continued for a period longer than three
weeks and then followed by a second treatment at a high
temperature.
323
324 Annals Entomological Society of America [Vol. VII,
METHODS.
The collecting of large numbers of insects which were
brought alive from the field to the laboratory was greatly
facilitated by the use of a net which Professor Woodworth
invented some time ago. The advantage of this net is that
insects may be procured by sweeping, even in damp weather,
without the injuries which are usually the result of such col-
lecting. The making of this net has been previously des-
cribed by Mr. E. T. Cresson, Jr., who uses it continually for
collecting small flies along the sand. Since, however, the net
is not as widely known as its advantages deserve, another
description will be in place:—
A strong piece of iron wire, three feet, eight inches long, is
bent into a circle with a one foot diameter—the ends are then
bent at right angles so as to lie adjacent and parallel to each
other. These ends are inserted into the small end of a six
inch ferrule and soldered fast. A short two foot handle will
be found best for sweeping. The net consists of white muslin—
a conical bag about eighteen inches deep. The tip is cut off
where the circumference of the bag measures about three
inches and is replaced by a small cloth bag four by six and a
half inches. This small bag is sewed to the point at which the
circumference of the large net is- four inches, thus leaving a
sleeve which hangs down into the small bag—this small bag will
just hold a quarter pound paper bag. ‘The sleeve of the large
net ‘fits into the paper bag. When filled from a minute’s
sweeping, the paper bag is pinched at the opening, taken out
of the net and placed in a botanical can. Upon the return to
the laboratory, the bag is opened at a well lighted window and
the contents picked over for specimens.
When insects of one species were found in sufficient number
to make it desirable to keep a number of them under observa-
tion as a unit, sets of capsules were bound together in tens as
devised by Prof. Woodworth for his insecticide experiments.
A piece of small iron wire two and a half inches long, sharpened
at one end is thrust through the base of a gelatin quinine capsule
so that the capsule is on the left of the wire with open end
upward—a twist is made in the wire to hold the capsule on—
then on the right side with open end in similar position, another
2.
1914} Longevity of Insects 325:
capsule is threaded upon the wire. In like manner, four more
pairs of capsules are threaded on. The advantages are that
the holes formed by the wire give ventilation and that the sim-
ilarity of the position of the capsules makes a numbering system
possible. The left hand first capsule bears the number of the
set of ten written in ink on its face—the other capsules count
up to ten in logical sequence from left to right towards the other
end.
All insects were placed separately in capsules. If the in-
sects were sufficiently duplicated in collecting, sets of capsules.
as above described were used. Otherwise the capsules were
placed in envelopes, bearing data as to date of collecting,
locality, temperature of collecting and temperature of treat-
ment.
The envelopes or sets of capsules were placed on shallow
wooden trays at different temperatures, room 62°F.—hot
room 72°F or ice room 42°F. Each day the capsules were
opened and examined, thus permitting a further change of air.
Any insects that had died were removed and a number corre-
sponding to the datum recorded was placed in the capsule.
The specimens were generally simply classified to the family.
The results are shown in the following table :—
326
Annals Entomological Society of America
Table 1. Longevity by Families.
Order
Family
[Vol. VII,
Temperature
High 72°F. || Medium 62° F.
Longevity in Days.
| Low 42° F.
.
v4
|Ave.|Min | |Diff.|Max|Ave.|Min.
\Diff.|Max|Ave.|Min.
Diptera....
‘Coleoptera..
Hymen-
optera. .|
Hemiptera...
“Orthoptera
Detien aes
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Acalyptre......
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Cecidomyide...|
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Chalcidide......
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1914] Longevity of Insects aAl
Table 2. Longevity by Orders.
Temperature
Number of | High 72° F. ||Medium 62°F.|| Low 42° F.
Order |
Specimens Longevity in Days.
Max|Ave.}Min.)|Max Ave.|Min | Max|Ave. |Min.
eee a ae ia he lhigy heo.5| To \lOe | 4 hd
Coleoptera.........| 64 15 6.6} 1 || 23 6.5) 2 || 39 | 20.0) 5
Hymenoptera......| 50 15 5 1 i AnD a 20 styan eal OS rs
Hemiptera..........| 24 6 25a ho 5 1 || abs: OCI) all
Orthoptera: .. 2.5%. a 1 1 1 i Oviteo 7 6.3] 5
Lepidoptera........ 3 7 aha ae tee ie prec tee toca Pee ONE | pr a [ 9
Aphaniptera....... 3 cd ee ae: hal Re alia eal lene
Thysanoptera...... 3 1 RYE beet 15 | escalate | caer
Wetiroptera....55 2. : 2 SARE a eae | ae oe ane ea eee fe LL teen lect
MSEC Ac, «s)he ss 359 15 4.8) 1 || 23 i re da |
Wrnehnida 2s... 5. 26 15 SO eae ie (Ft 1b Ds S| 8.8| 3
In most cases the data are too few to be very significant as
to individual groups but are sufficient to draw certain conclu-
sions, Viz:
1. That as regards longevity, the taxonomic divisions
show little or no comparable variability. That is to say that
the amount of variation in an individual species may be as
great as the variability of the genus or family or even order
making it appear that the average longevity of a large number
of insects of one species would give the same results as the aver-
age of the same number of many species.
328 Annals Entomological Society of America _[Vol. VII,
The following table in which the maximum, minimum and
average longevity at each of the three temperatures is recorded
for the order excluding the family with which it is compared,
will show the above statement to be correct.
Table 3. Taxonomic Groups and Longevity.
Temperature.
Order Family High 72° F. ||Medium 62°F.|| Low 42° F.
Longevity in Days.
Max|Ave.|Min ||Max|Ave.|Min_||Max|Ave. |Min.
Muscide...| 3 2 1 es bas 6 3 || 15 8 3
@©fher Wipteray os e.tlie sea eee - 4 126) ls 2.5| 1 || 2¢ 4 1
Curcu-
lionide....| 15 9.9| 4 || 25 6:1) 2 33 ) T38)
Other! Coleoptera calles. sess 7 5.8) 1 a Z 5 || 39 | 24.6} 6
Cynipide...| 15 5 1 ee 5) Sa) Ma hl il gl Lore 2
Other Hymenoptera]............ 15 5.2] 1 8 5 3 || 15 i) oe
Chalcidide | 15 Bt yee] ae | 9.8] 3
Other Hymenopteral............ is: |) Bal 1 7 egies 17-2 ab
2. That the longevity of insects in general is lengthened by
a decrease in temperature and shortened by an increase in
temperature (when these temperatures are between 42° and 72°
F.) |
Table 2 proves this to be true in all except two cases: (a)
Coleoptera in general have a slight increase in longevity at.
high temperatures over that of the medium temperature.
(b) Fleas in the three specimens tested show increase in
length of life as the temperature increases. (c) (Arachnida
have the greatest longevity at medium temperature).
3. That the difference in longevity of a species at different.
temperatures corresponds roughly to the difference in tempera-
ture. Table 4 shows that the greatest difference in length of
life is between the longevity at Low and the longevity at Me-
dium temperatures—this corresponds to the greater difference
between Low 42° F. and Medium 62°F. as compared with the
difference between Medium and High 72°F.
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1914} Longevity of Insects 329
Table 4. Proportion Between Temperature and Length of Life.
Temperature
brdé- Family _ | High 72° F. Med. 62° F.| | Low 42°F.
Longevity in Days
Ave. Diff. Ave. | Diff.| Ave.
Lo sie Sie ee eee 1.78 af 2.5 It? 4.2
Muscidae... .: 16 9 | eo 1.5 +
BEMETOPIGCEA.: |S: 2cec 202.0 2 t= 3% 6.6 ed 6.5 13.5 20
Latridiide.... 9 9.9 14.5 oot 22.2
Curculionide.. 3.7 1.3 off 17.6 24.6
MraAS@@tiaind 6 llr. er ois smcisce EM ees 4.8 1e2 6 4.9 10.9
The most important conclusion arrived at is that longevity
is not correlated with systematic groups. Table 3 (Taxonomic
Groups and Longevity) upon which this conclusion is based
was compiled in each case from the family in which the greatest
number of specimens had been included in the experiment.
It is not probable that the greater variation in a family than in
the average of the other families of that order as is apparent in
the table, is due to any greater adaptability to temperature
changes in that family than in the others. For a comparison
of the maximum and minimum number of days that the rep-
resentatives of the different families lived will show that indi-
vidual variation within the family, in the majority of cases
where a number of specimens of one species were used in the
experiment, is as great as individual variation for the group.
This great individual variation is probably due to the physiologi-
cal conditions of the individual. For example, in the Capside;
of the five specimens of one species placed at a high temperature,
all died in one day except one which moulted and lived for six
days. Apparently the longevity in this case was due to indi-
vidual physiological conditions and not to any inherent temper-
ature adaptability. Such cases could be multiplied.
We may therefore come to another conclusion, viz:
4. That longevity at different temperatures is due to
individual physiological conditions and that any attempt to
determine the temperature longevity of the species would be
confused by the variability of the results unless these physiolog-
ical factors were brought into account.
330 | Annals Entomological Society of America [Vol. VII,
It has been the general belief among entomologists that many
insects of the orders Diptera, Lepidoptera and Hymenoptera
in the imago stage take no food. Recent experiments (Doten
15) have shown that some parasitic Hymenoptera take food in
the adult stage. Closer observations may prove this to be the
case with many of the insects which are at present, thought to
abstain from food. However, most insects do not feed after
the eggs are fully developed. Whether or not, starvation is a
factor in this experiment, must therefore be left undecided for
the present.
PAR
EFFECTS OF EXPOSURE TO TWO DIFFERENT TEMPERATURES ON
LONGEVITY.
It was found in Part 1 of these experiments that longevity
varied greatly according to the physiological conditions of the
individual—in order to obtain further data on the nature of
these physiological conditions, the following experiment was
performed:
It was thought probable that temperature could produce
certain of these physiological conditions—therefore, an attempt
was made to find if exposure to a certain temperature for a short
time would result in a condition that would be evident in its
influence on the longevity of the insect at a secondary and
different temperature. The insects used as objects upon which
to experiment were the larve of the very common oak tree
moth (Phryganidia californica). The larvae were placed sep-
arately in capsules, wired together in sets of tens as explained
under ‘‘Method”’ in Part 1 of this paper. The sets of capsules
were then placed in wooden trays at medium or room tempera-
ture at high or the temperature of a bacteriological incubator
or at low, the temperature of an ice room, six by twelve by five
feet. After two days’ preparation at one of these temperatures,
the larve were transferred to one of the other temperatures
where they were kept until starvation resulted in death. The
larvee were examined each day and the date of death recorded.
‘“Experiment A’’ represents the results on one hundred
young larve of the first brood of 1913. “Experiment B”’
represents the results with eighty-four older larve of the second
brood of 1912.
1914] Longevity of Insects 331
Chart I records the results of these two experiments. The
abscissa of each of the points marked with circles is the longevity
of the larve at the constant temperature represented by the
ordinate. Each arrow leaving one of these points runs to a
point indicating in the same manner the longevity resulting
from the treatment at the two temperatures.
Chart I shows that from any change in temperature there
results an increased longevity of the larve, as follows:
Two Days . Longevit
Treatment at | Followed by Results in at acnetane
98° F. 58> BE. Same longevity as.............. OSnHe
98° F. 68° F. Increased longevity over....... 68° F.
68° F. 98° F. Decreased longevity below..... 98° F.
68° F. 58° F. Decreased longevity below..... 58° F.
58~ EF. 98° F. Increased longevity over....... 98° F.
58° F. 68° F. Increased longevity over....... 68° F.
82° F. 64° F. Increased longevity over....... 64° F.
64° F. 82° F. Decreased longevity below..... 82° F.
64° F. 46° F, Increased longevity over....... 46° F.
46° F. 82° F. Increased longevity over....... 82° F.
46° F. 64° F. Increased longevity over....... 64° F.
The exceptions to this rule are in three cases: (a) where 68°
F. is the preliminary two day treatment, there is a decrease,
(b) where the change is from 98°F. to 58°F. there is no increase
and (c) where 64°F. is followed by 82°F. there is a decrease in
the longevity.
From the data recorded on Chart I, we may form two con-
clusions; (1) that the life of the larve of Phryganidia californica
will be lengthened at any temperature (when starvation is a
factor) by placing the insect for two days at either a high or a
low temperature; (2) that the life of the larvae of Phryganidia
californica will be shortened at any temperature (when starva-
tion is a factor) by placing the insect for two day at a medium
temperature.
The temperatures that have been found to have the char-
acteristic effects described are: ‘‘High’’ 98-82°F.
Above 98°F. will probably have other characteristics as 98°
F. already shows a transition in that it does not cause an in-
crease in longevity when followed by a low temperature.
““Medium”’ 68-64°F.
332 Annals Entomological Society of America [Vol. VII,
CHAR TE HE,
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1914] Longevity of Insects 333
Between 64°F. and 58°F. there must be a temperature with
another characteristic for 64°F. holds a transitory position in
that it gives an increase in longevity when followed by a low
temperature and a decrease in longevity when followed by a
high temperature.
“Low” 58-46°F.
These temperatures show the characteristics ascribed to
“‘Low Temperatures.”
Since death by starvation is the end of the phase we are
studying in this experiment, it was thought that probably a
measure of the rate of growth would determine this rate of
senescence. Sanderson recommends the following method of
obtaining a temperature growth curve, viz: that a ‘“‘definite
valuation’”’ be found “‘in relation to the accumulation of temper-
ature necessary for any stage of growth” in the following
manner: if, at a certain temperature, it requires x days to go
through certain phases and this development be considered
equal to one unit, then each day’s growth at this temperature
is equal to (1l+x)%. Using this method, the following tabu-
lated growth valuations were found:
Table 5. Temperature Growth Valuations per Day.
Growth E | | Growth Growth
Exp.|No. Days) Temp. Value No. Days Temp. | Value Value
Ist.Temp Ist Temp. [24d Temp. 2nd Temp. | Total
A 9 | High a 9 5 2a bce] a pons Dare ee 1.0
9 Me Reln te | wreMeO PON more, Slere tk eats. De ge A eae AE 1.0
38 Low HO ZS EXM OOP ae ected tes pea seed ae Sete Soc ree 1.0
2 Med. 222 5.3 | High 147 1.0
2 Med. 222 33.1 | Low .0235 1.0
2 | High 229 7.2 | Med 108 1.0
2 | High . 222 36 Low .0216 1.0
2 | Low .052 ioe) ashe 4) .0837 1.0
2 | Low | .052 12 | Med. .079 1.0
B POU etsy ee OGG) yes. bse [ace eee fees eee ses 1.0
13.4 | Med. ALEC) Sc: SIU oe |e Me a 1.0
22 Low Ra ee fegee eM en ee eS rca wae a [a asenPyaval ov bm eae XO
2 Med. | .1492 24.5 | Low .0347 1.0
2 Med. ~}..1492 13.6 High .0625 | 1.0
a | High | .1204 ee ae Med. 0733) EO
2 | Low — .0908 i -ia.2 | Med. .0689 | 1.0
2 | Low 0908 | 27.8 | High 0323/5 «5250
334 Annals Entomological Society of America [Vol. VII,
The irregularity of the results given in the next to the last
column show that there is some other factor involved in the
determination of the longevity of starving larve at different
temperatures The most probable factor is the rate of metabolic
processes for it is the most closely connected with temperature
and nutrition, of any of the vital processes. Since the rate of
growth and the rate of metabolism will determine how long the
insect can live on the reserve material in its body. If the data
of Table 5 is plotted in a similar manner to the data of Chart I,
the graph on Chart II is obtained.
But if it is true that the rate of growth and of Metabolism
determine the longevity it is necessary to bring another factor
into consideration before we can explain why a two days treat-
ment at a low temperature will decrease this rate when the
insect is placed at a high temperature.
Growth changes in rate with advance in age but is not the
process that results in death for while growth is due to the
establishment of a constant relation between the nucleus and
the cytoplasm and therefore must finally reach a stage where
the growth is stopped, senescence always results in a decrease
in weight which cannot be accounted for by any theories of
growth according to Robertson (43, 44). Still a fall in the
rate of metabolism accompanies old age—hence, we must con-
clude that there is another factor than growth that determines
this rate of metabolism. It has been determined that the speed
of the metabolic processes decreases with age—therefore, it
may be determined by a measure of senescence. The progress
of senescence has been defined variously by several investi-
gators. Minot (35) basing his theories on certain truths,
which others have used in supporting the theory that a nucleus
can control but a limited mass of protoplasm (Sachs and
Boveri), has measured the rate of senescence by growth. This
has been shown to be improbable as before stated by Robert-
son and by Loeb (30) and Moore (36) who found that the
temperature co-efficient of growth (2.8) is very different from
the temperature co-efficient of longevity (1000).
Minot finds that senescence results from a gradual shifting
of the ratio between nuclear and cytoplasmic substance (Kern-
plasma relationship) to the side of the cytoplasm and from the
differentiation of the cells which accompany this change. This
1914] Longevity of Insects 335
differentiation, he claims, is irreversible. He therefore makes
no provision for rejuvenation in the Metazoa. C. M. Child
(10) has recently constructed a theory, which I will describe
shortly, based on certain experiments and upon our present
knowledge of the cell activities.
Cells go through two processes—one constructive and bene-
ficial or life-giving, i. e. metabolism—the other destructive,
katabolism. Both are necessary to life and a balance is main-
tained between them—when however, this equilibrium is upset
in the direction of the katabolic processes, senescence is the
result and finally death. The true measure of senescence then,
may be taken to be inversely the rate of metabolism.
In the life processes, many compounds are formed which
cannot be made in the laboratory without the use of great heat
or chemicals which are incompatible with life. It is believed
more and more generally that a study of the physical conditions
of the life substance, protoplasm, would throw great light on
these processes. Alsberg in his recent paper (2) on the mechan-
ism of cell activities, has given a resumé of present day knowl-
edge and conjecture on the subject.
The nature of protoplasm has been found to be similar to
that of colloidal solutions and to emulsions. It is made up of
substances that tend to concentrate at surfaces—this concen-
tration and reduction of the size of the phase results in an
enormous surface energy, which increases in immense propor-
tion to the smallness or roundness of the surface of that phase
or chemical locality. The very general composition of proto-
plasm, i. e., 80% water, 15-20% solid and 5% fats, would make
its rigidity impossible were it not for some emulsified condition.
It being a well known fact that emulsions often show great
rigidity.
Since these substances have a tendency to form phases or
localizations of chemico-physical conditions and since all these
phases are in contact and all differ more or less in permeability,
it is very possible that they act as a long series of interacting
yet separate, semi-porous test tubes. A reaction may go toa
certain stage then penetrate into the next phase and while
being isolated and going through another reaction, may still
influence the first phase—thus making it possible to complete
a very complicated and apparently impossible chemical change.
336 Annals Entomological Society of America __[Vol. VII,
Since the substances of this colloidal or emulsified solution have
a tendency to collect at surfaces and when once out of solution
(according to Loeb (81) are very difficult to bring into their
former condition, permanent, more or less impermeable bars
to the process of metabolism may be set up. These may be
broken down by a change in the chemical process or a change
even in their rate, due to exterior causes of temperature or food
quantity.
Childs in some experiments on Planarians finds that the
toxicity of alcohol which he uses as a measure of the rate of
metabolism varies inversely with the age of the animal, i. e.,
metabolic processes are being lowered and katabolism is gaining
the upper hand. He finds however, that rejuvenation is pos-
sible by a change in the rate of these metabolic processes.
Since metabolic processes are carried on through alveolar
walls of phases in the protoplasm of the cell and since the
longer this process of metabolism is carried on at the same rate
and in the same chemical nature, the more permanent these
walls become, a lowering of metabolic processes, i. e., senes-
cence due to the establishment of alveolar walls which have
through their permanency become bars to the action of metabo-
lism, is the result. He finds however, that a change in these
processes will result in an increased rate being possible for them.
If an animal is starved for a short time and then fed, its ability
to withstand the alcohol is greatly increased—this can be
explained by the probability that the processes have gone on
in spite of the lack of food and that the actual accumulation of
cytoplasmic alveolar walls of obstruction have been destroyed
and the cell thus brought into a younger stage of differentiation.
If the animal is starved for only a few days, this increased
resistance is very small, upon again refeeding.
The rejuvenation has not gone on to as great an extent,
therefore the resistance is less than that of the animal starved
for a longer time. A similar result is obtained with animals
that have been forced to regenerate parts—the larger the piece
is that has been regenerated, the greater the increase in resist-
ance to alcohol. In the case of regeneration, direct visible data
has been given by Godlewski (19) showing that regeneration
actually leads to a simplification of cells and a reverse process
of cytomorphosis that Minot did not take into consideration in
1914] Longevity of Insects 337
the formulation of his theory. It was also found by Child
(11) that the older a Planarian is, the more likely fission i. e.,
formation of a new individual from a part of the old is to take
place. This is probably due to the greater isolation that the
tail region of the animal has, because of the clogged condition
of the cells as age advances.
An application of these results of Childs, Godlewski and the
late experiments of Loeb (32) and Lillie on permeability of
membranes will make possible an explanation of all the results
of these experiments.
It must be remembered in the first place, and above all that
one factor of the experiment was starvation—second, that the
insects were placed first for two days at a preliminary tempera-
ture and then at a different temperature until they died. Since
the result of starvation at a temperature is to clear the cell of
cytoplasmic obstacles to a certain degree. The preliminary
treatment of an insect with starvation at a temperature will
determine to a great extent the results of treatment at a second
and different temperature. On the accompanying Chart II,
I have therefore plotted, the rates of senescence. They were
obtained by finding the value of each day at a certain tempera-
ture for completion of a phase but since the end of this period
was death, they may serve as the measure of the degree of sen-
escence.
Since death will finally be the result of physiological sen-
escence, due to lack of food, we must bear in mind the distinc-
tion between this and natural death which is the result of
morphological senescence, the reverse of which is taking place
in this case.
Reference to this chart then, will show the degree to which
any treatment of temperature will result in combined morpho-
logical rejuvenation and physiological senescence. It will be
seen :—
1. That preparation at a high or low temperature will
result in a combination of physiological oldness and morpho-
logical youngness which will make the insect more liable to
live, if it be placed in any other temperature, longer than if it
had been living constantly at this secondary temperature.
2. That preparation at a medium temperature will render
the insect older, both morphologically and physiologically and
338 Annals Entomological Society of America [Vol. VII,
therefore less liable to live, if it be placed at any other tempera-
ture, longer than if it has been living constantly at this secondary
temperature.
The rapid starvation at the high temperature has morpho-
logically rejuvenated the insect but has rendered it physiologi-
ally old. This slowing down and probably also change in
function has rejuvenated and removed the cytoplasmic ob-
stacles while morphological age, due to destruction of reserve
products, has gone on to a less extent than at the high tempera-
ture. At medium temperatures, there is no change in rate nor ~
a great enough degree of starvation to remove these inactive
substances—therefore the cell is not rejuvenated morphologic-
ally and is physiologically old. In other words, the insect is
older than the insects prepared by either of the other two
methods.
PART 3:
EFFECTS OF EXPOSURE TO TWO DIFFERENT TEMPERATURES ON
HIBERNATING INSECTS.
In part 2 of. this article, certain studies of the effects of
temperature upon the longevity of starving insects were made.
In this part, I propose to further substantiate the statements
made by the results obtained from certain experiments on the
hibernating brood of the Codling moth larve (Carpocapsa
pomonella L.)
The experiment was started with larve collected from wind-
fall apples gathered under the trees and sent by the courtesy
of Mr. Frank Perry of Sebastapol, Sonoma County, California,
where the insects were collected. These insects were taken
in the late part of July, 1913, and many of them pupated. Be-
lieving these to be of the earliest second brood, the experiment
was abandoned and begun over again with larve that were
collected in the cocoon—all the two hundred and fifty larve
of the second experiment were collected in one mass of cocoons
under a packing house. There could be no doubt then as to
their hibernating condition and as to the similarity of their
exposure to temperature, humidity and disease.
The larve were handled in the following manner: the co-
coons were opened and two larve dropped into each clean test
tube which was then plugged with cotton. The test tubes
1914} Longevity of Insects 339
were mixed to avoid the possibility of having a set of larve
from the same part of the mass of cocoons. The test tubes
were placed in round paste board boxes which gave room for
seven of them and insured perfect darkness—a long strip of
paper was placed in the box upon which was kept a complete
record of the temperature treatment.
The insects were kept at three temperatures—room temper-
ature as a check, low temperature in a refrigerator, usually
about 43° F., or high temperature 86 to 96° F., maintained by
an electric light.
The first experiment performed was to place a set of test
tubes at the high temperature—the larve of this experiment all
died in twenty-six days except one, which pupated, but did not
hatch. An attempt was then made to bring the larve out of
their hibernating condition by first chilling and then heating.
Sets were placed in the refrigerator for varying lengths of time
—it was found that an exposure to cold of from seven to four-
teen ‘days greatly lengthened the life of the larve and raised
the percentage of pupation and of hatch. This percentage is
much higher than that obtained by heat without previous
chilling or by exposure to room temperature as in the check.
After fourteen days it will be seen by reference to Chart I
that the longevity does not increase and that no pupation occurs.
Four conclusions can be drawn from Chart I—
1. That pupation of hibernating Codling moth larve is not
usually brought about by heat.
2. That exposure of these larve to a low temperature for
from one to two weeks followed by heat results in pupation,
hatch and increase in longevity of those larvae which do not
pupate.
3. That after twenty-one days exposure to low temperature,
heat does not result in pupation nor is the longevity increased.
4. ‘The number of days which the larve that die, live at the
high temperature, is approximately equal to the total number of
days, the other larve take to pupate.
In order to arrive at some conclusion about these experi-
ments, first, let us consider the nature of hibernation. Hiber-
nation takes place in many forms of insects, fish, Amphibia,
Mollusks, birds, Mammals and even in man. Peasants of
Russia, according to Cleghorn (12-13) with the approach of
340 Annals Entomological Society of America [Vol. VII,
famine, build a fire in a huge stove which serves as a resting
place and lying upon this, keep as quiet and warm as possible
and thus reduce their need of food. Among the Mammals,
the marmot has been the most studied of the hibernating forms.
Cleghorn lists a number of animals that hibernate—he states
that bats of different species hibernate at different times of the
year—that when disturbed for a time, they breathe almost
normally and then again, the respiration goes down almost to
zero. If awakened suddenly by great heat, death always
ensues. He says that bears are as fat after hibernation as when
they go into it in the fall and that female bears even raise their
young while not obtaining any food and still show very little
change in condition. Bears and badgers of the North do not
go into any true state of hibernation but sleep lightly through
the winter. The black bear, however, is aroused with difficulty
from the winter sleep—the woodchuck of Canada, the European
hedgehog, chipmunks and ground squirrels, all hibernate.
Frogs hibernate in mud at the bottom of pools and if awakened
by warmth can remain much longer under water without being
drowned than during the active season. Some fish survive long
draughts by burial in the mud. - Baker (5) states that during
some seasons of draught, Lymnzidze bury themselves and form
an epiphragm inside the outer lip as is common with Helix
during hibernation and estivation. Plants have a similar
phenomenon also known as hibernation which is closely con-
nected with lowering of temperature and shows itself in the
decreased rate of the metabolic processes.
The physiology of hibernation has best been studied by
Bellion in the European edible snail (1’ escargot). Bellion (6)
finds that the moisture content of the air and not temperature
is the essential external factor of hibernation—when the mois-
ture content is low, and epiphragm is formed in spite of low or
high temperature and the snail is plunged into a condition of
lethargy. If moisture content is high, no epiphragm is formed
and activity is at its height even at a low temperature. Car-
bon dioxide content of the tissues increases towards the end
of hibernation while the oxygen content diminishes in propor-
tion. Dubois (16) has found in the marmot that when carbon
dioxide is present in a certain proportion in the blood, torpor
sets in. At moment of awakening, carbon dioxide is high—
it is very probable that the carbon dioxide and rehydration
1914] Longevity of Insects 341
awaken the snail, as the carbon dioxide and dehydration
plunges it into sleep. The amount is the essential to sleep or to
awakening. Janichen (25) believes that the theory of autonar-
cosis of carbon dioxide should be held for all cold blooded
animals.
The histological changes of hibernation have been studied
in the hedgehog by Carlier (9). Plasma cells with deeply
staining granules and with lightly staining nuclei are present
in great numbers in the base of the tongue—they have the
appearance of overfed cells although the fact that they are not
found far into the digestive tract, seems, he states, to contradict
this appearance. During hibernation the granulations dis-
appear and the tissues of the tongue are less stainable. Num-
bers of the wandering white blood corpuscles are destroyed by
macrophags and their number is recuperated all during hiber-
nation by karyokenetic division in the lymph glands. During
this period, some liver cells increase in size followed by an en-
largement of the nucleus until the latter, having overstretched
the nuclear network, ruptures and disappears—this Carlier
believes to be the natural death of the cell.
Insects usually hibernate towards the end of summer when
the temperature is falling but they are also known to go into this
condition even though placed at a high temperature. Tower
(53) found in his experiments with the potato beetle that he
was unable under any laboratory conditions of high temperature
to bring the beetles into hibernation at an unusual time. Sand-
erson (47) found that tent caterpillar eggs will not hatch if
placed in a green house before being exposed to low temperature,
while those which stay out of doors until the temperature falls
will hatch rapidly at green house temperature. Merrifield
(34) concluded from his experiments with seasonal dimorphism
that there is probably a strong tendency for individuals to take
either the winter or the summer form in spite of all temperature
treatments.
Weismann found that summer forms could be obtained in
winter (55), by chilling a pupa and then subjecting it to heat,
while on the other hand, if the pupz were put immediately at a
high temperature, they did not hatch until summer. There is.
further data to show that low temperature is in many cases not
the only factor in hibernation. Foster (*) states that of seven-
* Life History of the Codling Moth, U. S. D. A. Bur. Ent. Bul. 97, Part 2,
Foster.
342 Annals Entomological Society of America _[Vol. VII,
ty-eight Codling moth larve collected on July 17, at Walnut
Creek, Cal., thirty-eight pupated, twenty hibernated as larvae
and twenty-eight died. The temperature at Walnut Creek
during July in 1909 actually increased three degrees over
the mean temperature of June. Most larve of the second
brood leave the fruit by the first of September and ninety-five
per cent. hibernate as larvaee—yet the temperature in September
is 3.3° F. higher than the temperature of June. According to
Simpson (52), at Grand Junction, Colo., of 33 Codling moth
larve collected July 16-23, 1900, but one hibernated while of
192 collected from August 30 to September 4, 192 hibernated.
The mean temperature of June was 63.3°F. of August 67.8°F.
and of September 61.7°. Yet the percentage of larve that
hibernated had gradually increased from June to September.
Sanderson (47) finds that some Lepidoptera of the North
when introduced into the South, do not have an increased
number of broods as would be expected nor do southern forms
have more than the one hibernating period, which is common
to them in their warmer clime when introduced into the North.
He bases this statement on the fact that the following insects
have but one generation in the South: tent caterpillar, peach
borer, plum curculio, canker-worm, gypsy-moth, brown tail
moth, and insects effecting native trees, all of which are in-
digenous to the North. Newell (39) claims that the cotton-boll
weavil enters hibernation after the first hard freeze and not due
to a mean average temperature of 60° F. or even of 43° F.
This is contradicted by Sanderson (46) who claims that weevils
hibernate when the average temperature falls below 60° F.
Hunter and Hinds (24) agree with Newell in saying that hiber-
nation begins after the first hard frost—though if the insect be
deprived of food, it will go into hibernation when the mean
average temperature is below 60° F.; at a temperature of 60
to 65° F. however the adults will starve.
Moisture may also be a controlling factor of hibernation as
has been shown in the case of the snail in estivation and hiber-
nation and also in the case of zestivation in the fish and in the
Lymnezide.
Frogs also go into estivation during summer as do plants
and probably all animal life in arid countries. Loeb (81) points
out that lack of water may act similarly to a low temperature—
this may account, he says, for the fact that seeds can be kept
Ce EO ENE ge es We ee ey ee
CE BET EAG TG eS FT! BAF HOE ES Oy PEP BOTA Por: aa ae”
Sy ten ge Met
1914] Longevity of Insects 343
alive for so long. The effect of ether on plants is similar to
hibernation and since the action of ether is probably a drying,
one, this may throw light on the importance of moisture in
hibernation. Hunter (24) has found that dryness is desirable
for hibernation—he finds that more weevils die during hiberna-
tion from exposure to moisture than from cold, on the other
hand, high temperature and moisture are the best conditions
for weevil larve to develop. Sanderson quotes Tower as
keeping potato beetles in hibernation for eighteen months in a
dry atmosphere. Immediately when placed at a normal
humidity, they immerge from hibernation. Donaldson (14)
finds that frogs differ in the rate of reabsorbing water during
summer and hibernation—it being more rapid in the former—
he also finds that the water content of the spinal cord varies
with the season—during the growth period (May 30 to July 1)
it is high and gradually diminishes towards the end of the season.
Rulot (45) has found that during hibernation, the production
of metabolic water sometimes falls to zero in the bat. Hatai
(21) has found that the effect of partial starvation on the
nervous system is to decrease the percentage of water by 24 per -
cent. upon returning to normal diet, the water content is found
to be higher than in the check. Abbe (1) has found that soaking
seeds in water before planting accelerates germination but that
germination is greatest in dry soil.
Tower states that during hibernation, the cells take on a
definite appearance due to loss of water, being shrunken and
flattened. In all cells, the protoplasm takes on a colloidal
granular appearance which is retained throughout the whole
period. The nuclei have an extremely vegetative appearance—
it often being impossible to show the presence of chromatin in
cells which later will have abundant and active chromatic
conditions. There is a twenty-seven per cent. loss of weight
due to the emptying of the malphigian tubules of a red fluid and
a three per cent. loss of weight due to the emptying of the
alimentary canal that takes place just before hibernation in the
potato beetle.
Tower believes that this lowering in water content makes the
maximum and minimum at which protoplasm can survive
change in temperature in either direction, greater. Upon
emergence from hibernation the reverse of the process of pre-
paration for hibernation takes place—there is a rapid gain of
344 Annals Entomological Society of America [Vol. VII,
water—the cytoplasm becomes more watery, vacuoles appear,
the cells become larger and more turgid and the chromatic
elements stain deeply and increase in size, thus presenting all
the signs of intense activity. The preparation in the potato
beetle for estivation is similar to that of hibernation—the ani-
mal remaining underground until first rains. Tower states.
here that the reduction of water gives an increased capability
of meeting higher temperatures.
Hibernation usually follows a period of great feeding—
whether this is what makes hibernation possible or whether it is.
the controlling factor of hibernation or not is unknown. In the
marmot, there is a definite storage gland called the hibernation
gland and Cleghorn includes in his definition of hibernation, the
formation of reserve fat to be used during that period. In the
potato beetle, the great period of feeding takes place before
hibernation and estivation (a little less in the latter) this
oversupply of food is stored up in the fat body and is used to a
certain extent during hibernation for there is a decrease in
weight of the insect during that period. The spermophile and
the marmot according to Cleghorn go into hibernation imme-
diately after having laid up the last layer of fat. This occurs.
at a period when their food is most plentiful. The frog accord-
ing to Holmes (23) goes into hibernation immediately after a
period of great feeding. There is. some evidence that over-
feeding takes place just before hibernation, in the Codling moth
for example: Hammar (20) has found that the feeding period
of the larve of the first brood (transforming directly into pupz)
lasted 24.7 days while that of the first brood which hibernated
lasted 28.9 days and the whole second brood (hibernating)
34.2 days. Inthe next year (1911) he found that the first brood
which was to transform had a feeding period of 21.2 days.
while that part of the first brood that was to over-winter as.
larve fed for 28.2 days. Jones and Davidson (28) find that the
second brood feeds twenty days longer than the first and at a
higher mean temperature. Jenne (26) finds in like manner
that the over-wintering brood of larve fed a longer time (.8 of a
day) than the transforming brood.
Morgulis (38) has found that during hibernation, the
nucleus is nourished by the cell—during starvation on the con-
trary, the nucleus at first loses volume rapidly though it re-
mains more or less unaffected after it has attained a certain
1914] Longevity of Insects 345
minimum size. It is possible that by diminishing the volumne
it increases its absorbing capacity. Hibernation is also unlike
starvation in its characteristic quiescence, for animals when
starved are very active. In hibernation also,there is no regen-
eration of tissues while in starvation this often occurs.
Hibernation seems to have a close connection with the
maturation of the reproductive organs. Tower has found that
those potato beetles that have gained sexual maturity, do not
succeed in passing through the hibernating period successfully.
Sexual maturity is seldom gained before hibernation in the
second brood of this insect. This activity is greatest imme-
diately following hibernation. He finds that the germ cells
remain in the female as oocytes during hibernation and develop
rapidly after hibernation. There. are two generations in all
climates—it would be supposed, Tower says that at high
temperatures, breeding would go on continually but every
alternating brood has a rest period before breeding goes on—
this rest period is estivation or hibernation depending on cli-
mate. All grape leafhoppers that have reached sexual maturity
are unable to pass through the period of hibernation success-
fully—only the very immature males and females live through
the winter to produce the next brood (according to Johnson
27).
Morgulis quotes the case of the Rhein salmon which makes
a sojourn of from six to nine and a half months in the Rhein,
remaining without food, developing in the meanwhile, its sexual
elements at the expense of fat and proteids accumulated before
hand. Holmes states that the period of great feeding preceding
hibernation supplies food for that period and for the develop-
ment of the reproductive organs which are to come into full
activity immediately after hibernation. Hibernating insects
seldom arrive at sexual maturity before this period is over.
Newell found that the female cotton boll weevils which have
hibernated continue to deposit eggs for a much longer time than
the others. Morgulis claims that insufficient feeding effects
the ovaries the most; since these organs seem to often develop
during hibernation, it is very improbable that inanition takes
place during this period. Loeb quotes Giard and Caullery as
having found that a regressive metamorphosis occurs in Synas-
cidians and that the animals hibernate in this. condition. The
muscles of the gills of these animals are decomposed in their
346 Annals Entomological Society of America [Vol. VII,
individual cells. The result is a formation of a parenchyma.
which consists of single cells and of cell aggregates resembling a.
morula. It is probable that a similar disintegration of parts
takes place during hibernation and it is certain that it takes place
during pupation. According to Sharp, when the larva of an
insect has attained its full growth, many internal tissues disin-
tegrate and rudimentary sex organs reabsorb the products of
disintegration and with the other regenerative buds produce
the perfect imago. On the contrary Jordan claims that the
longer duration of the period of oviposition in the newt as com-
pared with many other Amphibia may perhaps be correlated
with the absence of the ‘‘fasting habit’’ (29).
The foremost essential factors of hibernation judging from
the above observations seem to be temperature and moisture
conditions, over-feeding and maturation of the reproductive
organs. It is often stated that the loss of water makes it pos-
sible for the cell to withstand freezing temperature—for other-
wise, as is claimed to be the case in plants (Vines 54) the ice
crystals formed would rupture the cells. It is a known fact
however that if cooled very slowly cells in which ice crystals.
have been formed, will again become normal. Tower and
Sanderson state that the loss in water of the protoplasm makes.
it possible for this substance to stand greater variation in
temperature for the concentration of salts makes the freezing
point lower. But it is a known fact that the freezing point of
sols is but sl ghtly lowered by an increase in the concentration of
a salt dissolved. They also believe that it makes the protoplasm
more able to withstand the high temperature but Loeb and
Bachmetjew (3 and 4) have found that the point of coagula-
tion of colloidal substances varies inversely with water content.
This may account for the great killing of hibernating insects.
which Wright (56) ascribes to a rather warm winter.
Most animals that hibernate do so at a period just following
great feeding and often at a time when their food is at its
greatest abundance, as for example the cotton boll weevil,
according to Sanderson (46). In some cases there is cytological
evidence of overfeeding—for example, the overfed plasma cells.
in the hedgehog and the vegetative staining quality of the cells.
in the potato beetle and as I have found in the Codling moth
larve. Overfeeding leads to increased number of molts or to
hypermetamorphosis according to Sharp (51) who claims that
= —
Rn eo eos ae lle tte
1914] Longevity of Insects 347
ecdysis is an extra excretory process. Quaintance and Brues
(42) found that highly nutritive foods caused less molts but
insufficient and disagreeable food resulted in more molts.
Sharp says that many hibernating larve have an extra
molt. This may be either a sign of over-feeding or of feeding
on some less nutritive substance in larger quantities.
I found in my first experiments with Codling moth larve
in windfall apples that those larvze which were about to hi-
bernate remained inactive in the apple for some time (two days
to a week) without eating before leaving the fruit to form a
cocoon.
If it is granted that there is a condition of over-feeding
in the larve before hibernating, it will be seen that there are
many similarities between this stage and the condition before
and during the molt. Before the molt, there is a period of
great feeding—then a short period of quiescence, then the his-
tolysis begins.
The process of histolysis is one of rejuvenation—in the
second part of this paper, a résumé of the present day knowledge
of the process of senescence was given—the most up-to-date and
I think the best of these theories is the one advanced by Child.
Child, basing his theory on the alveolar nature of protoplasm
and on the nature of the metabolic processes and their tendency
to lead to structural differentiation in the establishment of
cytoplasmic alveolar walls, formulates the following law:
‘Senescence in nature consists physiologically in a decrease in
the rate of metabolism and this is determined morphologically
by the accumulation in the cell of structural obstacles to
metabolism, e. g., decrease in the permeability, increase in
density, accumulation of relatively inactive substances, etc.
Rejuvenescence consists physiologically in an increase in the
rate of metabolism and is brought about in nature by the re-
moval in one way or another of the structural obstacles to
metabolism.’’ Since in the process of pupation, the tissues pass
through a more or less complete process of histolysis which is
aided by phagocytes, and new tissues often arising from germ-
inal buds absorb this old material (Sharp, Packard (41)
and Ganin (18), the cells of these tissues are probably less com-
plex in their cytoplasm. Sharp says that the physiological con-
ditions of the later larval life are different from those of the earlier
348 Annals Entomological Society of America _[Vol. VII,
life, possibly as the direct result of a mere aggregation of matter—
such a histolysis as above described, would reabsorb and re-
distribute this extra matter in such a way as to clear the cells of
all inactive substances. During hibernation in the frog (Mor-
gulis quotes Leonard) a similar process of histolysis and shifting
of the nucleocytoplasmic relation in favor of the nucleus takes
place. Without doubt, the cells are rejuvenated in the frog
during hibernation—the case of Synascidians has already been
stated. Lillie has found that fresh water Planarians if ex-
posed to starvation, ultimately return to an embryonic form.
These experiments have been confirmed by Schultz (50).
Childs found in his experiments on Planaria that starvation
and regeneration both lead to rejuvenation—starvation differs
from hibernation in that the life processes go on at a high rate
in the former while they are sunk almost to zero in the latter.
Starvation does not lead to the lowering of the water content
as hibernation does, except in the nervous system. ‘The con-
ditions of the cells in the hibernating or in the starving
insect are quite different. In the hibernating animal, the
condition is one of overfeeding and probably of old age—
that is, the accumulation of inactive substances in the cell
is very great. - In the starving animal on the other hand,
the conditions are morphologically extremely young and
physiologically old (underfed). | Child compares cells in the
overloaded condition to an ovum and the starved young
cell to a spermatozoan. Loeb (32) has found that fertiliza-
tion increases the permeability of membranes. The action of
fertilization is the same as rejuvenation. A similar rejuvena-
tion may take place by change in feeding as Calkins (8) has
found to be true in his experiments with Paramoecium—where
no conjugation took place if a change in feeding were made at
the proper time. This agrees with Child’s theory that rejuven-
ation can be brought about by a change in the chemical process
of metabolism.
One characteristic of overfed Planarians according to Child
is the physiological isolation of parts due to the overloaded
condition of the cells with inactive bars to metabolism in the
cytoplasm. This isolation leads to fission or to a senescence,
i. e., a lowering of the rate of the metabolic processes. In
Codling moth larve that are about to hibernate, I have found -
very similar conditions to exist—first, the vital processes are
1914] ~ Longevity of Insects : | 349
at a low ebb—second; there is apparently a physiologically iso-
lation of parts—this isolation is evident'in the following ways:
the larvze often become entangled or bound by a thread of a
spinning larva close by. The bonds which are thus tied
about them become so tight that the insect is almost cut in two.
I have often observed that the posterior half of the insect may
have died from the effect of this isolation and decay set in while
the anterior part may remain-unaffected for many days. Dis-
ease also has been observed.in these experiments to spread
very slowly through the, insect—this also can be accepted as
evidence of the overfed and senescent condition of the larva
which is about to hibernate.. This has generally been found to
be the case in hibernating mammals on exposure to disease.
(Carlier and Dubois (17).
It seems probable then, that the. overfed condition of the
insect and the ‘‘old”’ state of the cell has reduced the permeabil-
ity to a great degree and as a result, the rate of metabolic pro-
cesses is greatly lowered. The loss of water probably results
in the alveolar walls going out of solution and being cast out.
In starving Planarians and in those which have undergone
regeneration according to Nussbaum and Oxner (40), granules
are present throughout the tissues. These granules are absorbed
‘by phagocytes from the body wall—a similar process takes place
in the potato beetle during hibernation, according to Tower
‘and is characteristic of the process of histolysis, according to
Henneguy (22).
During the molt, pupal period, and apparently in hibernating
Codling moth larve (as I have observed in my experiments)
these granules are very abundant.
From these considerations, it is possible to formulate certain
working hypotheses which will serve as guides for further
experimentation and consideration of which may throw further
light on the nature of the processes of hibernation. These
hypotheses are:
1. That temperature is but a single factor and not neces-
sarily the controlling one in hibernation.
2. That hibernation is usually concomitant with overfeed-
ing and may be a result of that condition or the result
of accumulation of inactive substances in the cyto-
plasm of the cell due to feeding on innutritive food.
350 Annals Entomological Society of America _[Vol. VII,
3. That the loss of water which is general in hibernation
probably results in a discharge of insoluble alveolar
cytoplasmic structures which have accumulated and
produced this premature senility with an accompany-
ing lowering of the rate of the metabolic processes.
4. That starvation during hibernation together with this
loss of water may result in rejuvenation, when aided
by histolysis, and in increased permeability.
5. That this rejuvenated condition and increased per-
meability will, if stimulated to activity by heat,
permit pupation in Codling moth larve, which in this
case is the termination of the hibernating condition.
If we remember that the temperature at which colloidal
substances coagulate lowers with decrease in water content and
that long exposure to cold may result in this decrease in water
as well as exposure to high temperature and also the following
observations of Bachmetjew, we can explain that the result
of a long exposure to cold is the same as the result of a short
exposure to heat and that the intensity of the cold, shortens
the length of the period (Henneguy) :
(a). The relation of the point of coagulation varies with the
water content and the point of protoplasmic rigor is also low-
ered by hunger.
(b). Hunger lowers the critical point in direct proportion
to the number of days of its duration.
(c). The intensity of cold shortens the time necessary for
cold rigor.
The use of the hypotheses just outlined makes possible an
explanation of the results of this experiment. If hibernating
insects are placed at a high temperature directly, before being
exposed to a low temperature, the characteristics of starvation
rather than those of hibernation will set in—in other words, the
nuclear material will decrease in greater proportion than the
cytoplasmic material. On the other hand, if the insect is
placed at a low temperature the characteristic enlargement in
the nucleus at the expense of the cytoplasm and due to the
low temperature, according to Boring (7), will take place.
With the lowering in the rate of metabolism, due to low
temperature, the inactive conditions of the cells and their
i a
1914} Longevity of Insects 351
enlargement in nuclear material is the ideal condition for dis-
integration. This has been shown to be the case in the liver
cells of the hedgehog by Carlier.
In my experiments, I have found that the tissues of hibernat-
ing Codling moth larve show the presence of granular sub-
stances, immediately after the larve have been exposed to the
low temperature. Probably these granules indicate cytoplasmic
obstructions which due to the disintegration and inactivity
of cells have been thus disposed of, leaving the cell in a rejuve-
nated condition. Tower found in the potato beetle these same
granules present in hibernation and immediately after hiber-
nation, a resumption of the activities of the cell, a loss of the
vegetative unstaining quality and a more watery and less
differentiated appearing condition of the cell. If the insects
that are hibernating are exposed for increasing lengths of time
to a low temperature and then placed at a high temperature,
the tissues will have become rejuvenated and therefore with an
increase in temperature, acceleration of the metabolic processes
and of growth can take place.
However, if this exposure to cold is of too long a duration,
either too much of the water content will have been lost and
coagulation corresponding with permanent heat rigor, will set
in at a lower temperature than after but a short exposure to
cold or disintegration of tissues will have gone on to too great
an extent.
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Winterschlaf bei Raupen. Insek. Boerse., Vol. 16, 1900.
26. Jenne, E. L. The Codling Moth in the Ozarks. U.S. D. A., Bur. Ent.,
Bull. 80, part 1, 1909.
27. Johnson, F. The Grape Leafhopper in the Lake Erie Valley. U.S. D.A.,
: Prof. paper, No.'19, Jan. 24, 1914.
28. Jones and Davidson. Life History of the Codling Moth in.the Santa Clara
Valley. U.S. D.A., Bur. Ent., Bull, 115. .
29. Jordan, E.D. Development of the Newt. Jr. Morph., vol. 8, page 269.
30. Loeb, J. Ueber den Temp. koeff. f. die Lebensdauer kaltblutige Tiere und
ueber die Ursache des naturlichen Todes. Arch. ges. Phys., Bonn., vol.
124, 1908, page 411.
31. ————. Dynamics of Living Matter. Col. Univ., 1906.
32. —————. The Mechanistic Conception of Life. Univ. Chicago, 1912.
33. Melander and Jenne. The Codling Moth in the Yakima Valley. Wash.
; Agr. Exp. St., Bul. 77, 1906. ;
34. Merrifield, F. Systematic Temperature Experiments on Some Lepidoptera
in all their Stages. Tr. Ent. Soc. London, 1890, page 131. ; ~
35. Minot, C. S. The Problem of Old Age, Growth and Death.
36. Moore, A. R. ‘The Temperature Coefficient of the Duration of Life in Tubu-
laria. Arch. Ent.-mech. Org., vol. 29, page 287.
37. Morgulis, S. Contribution to the Physiology of Regeneration, I. Jr. Exp.
Zool., vol. 7, page 595.
38. —————. Studies of Inanition in its Bearing on the Problem of Growth.
Arch. Ent.-mech Org., vol. 32, page 160.
39. Newell, W. The Boll Weevil. La. Crop Pest. Comm. Cir. 9, 1906.
40. Nusbaum and Oxner. Studien ueber die Wirkung des Hungers unsw. I.
Arch. Ent.-mech. Org., vol. 34, 1912, page 386.
41. Packard, A.S. Guide to the Study of Insects.
42. Quaintance and Brues. The Cotton Boll Worm. U.S. D. A., Bur. Ent.,
Bull. 50, 1905.
43. Robertson, T. B. Further Remarks on the Normal Rate of Growth of an
Individual and its Biochemical Significance. Arch Ent.-mech. Org., vol.
26, page 108. :
err
1914] Longevity of Insects 353
44, . On the Normal Rate of Growth of an Individual and its Bio-
chemical Significance. Arch. Ent.-mech. Org., vol. 25, page 581.
45. Rulot, H. Note sur l’hibernation des Chauves-souris. Arch. de Biol. 1901.
46. Sanderson, E. D. Some Observations on the Mexican Cotton Boll Weevil.
U.S. D. A. Bur. Ent., Bul. 52, 1904.
47. —————. The Relation of Temperature to the Hibernation of Insects.
Jr. Econ. Ent., Vol. 1, page 56, 1908.
48. —————. The Relation of Temperature to Growth of Insects. Jr. Econ.
Ent., vol. 3, 1910, page 111.
49, ———.. The Influence of Minimum Temperature in Limiting the Northern
Distribution of Insects. Jr. Econ. Ent., vol. 1, page 245.
50. Schultz, E. Ueber Hungerscheinungen bei Planaria. Arch. Ent.-mech.
Org., vol. 18, page 555.
51. Sharp, D. Insecta. Camb. Nat. His.
52. Simpson, C.B. The Codling Moth, U.S. D.A., Div. Ent., Bul. 41, 1903.
53. Tower, W.L. Evolutionin Chrysomelid Beetles. Carnegie Inst., No. 48, 1906.
54. Vines, S. H. Physiology of Plants. Camb. 1886.
55. Weisman, A. Essays on Heredity. 1889.
56. Wright. Butterflies of the West Coast, San Francisco, 1905.
FOUR NEW TETRANYCHIDS.
By E. A. McGreoor, Bureau of Entomology.
The following species of phytophagus mites from the
Southeast are of considerable economic importance and are
herein described for the first time.
Tenuipalpus bioculatus sp. nov.
Female. Body crimson, with two rather well-defined eye-like spots
on cephalothorax. Widest at posterior corners of cephalothorax, two-
thirds as wide as long. The cephalothorax is narrowed considerably
anteriorly, and the abdomen tapers to a rounded tip. The body is
armed with a pair of weak spines on the anterior body margin medially,
similar spines immediately before and behind the emarginate eyes, six
at the posterior tip of the abdomen, a few along the body margin, and
scattered ones dorsally. The cephalothorax is hardly half as long as
broad, with the anterior margin convex; the palpi greatly resembles the
Tetranychus type, the penultimate joint bears a strong claw, and the
terminal joint (thumb) bears a “finger”. The legs are relatively
Fie. 1.
Tenuipal pus bioculalus. Right leg I, dorsal view (enlarged 650 times).
stout, crenulated; forelegs in length three-quarters the width of cephal-
othorax; four anterior tarsi blood-red in life; all legs bearing several
lateral hairs, and a terminal bristle in length equalling the three distal
segments; the trochanter of the four anterior legs with a lamellate hair
placed dorsally; the tarsi with several terminal appendages including
a pair of closely appressed claws, a very long bristle, and the four capi-
tate hairs, so frequently seen in Tetranychus.
Length, 0.235 mm.; width (hind margin of cephalothorax), 0.149 mm.
The egg is thickly elliptical in linear outline, and measures .096 mm.
by .067 mm. It is blood red in color from the first. The eggs are depos-
ited with the long axis perpendicular to the leaf, closely packed (like
those of Coccinellids), often comprising clusters of several hundred.
Type No. 19090, U. S. Nat. Mus.
NOTES.
The six posterior spines are much more conspicuous in the
younger stages of the larva than in the adult. The molt takes
place through a transverse rupture (at the suture between the
354
.
.
_
4
:
b
7"
2.) arg Ses
tea
al 3 x
rae 8
water? Oe
1914] Four New Tetranychids 300
cephalothorax and abdomen) quite similar to that of the red-
spiders. The male is decidedly smaller than the female, and
the abdomen is suddenly constricted behind the cephalothorax
and decidedly more attenuate than is the case with the female.
The legs of the male are relatively longer, colorless, and the
hairs and bristles are more conspicuous.
From Batesburg, South Carolina, on privet (Ligustrum
amurense), .Rumex acetosella, Oxalis stricta and garden mint
(Mentha spicata), collected by Mr. F. L. McDonough and the
writer, and from Baton Rouge, La., on privet and strawberry,
collected by Prof. E. S. Tucker. At Batesburg this pest has
been observed frequently to inflict severe damage to privet
hedges. Several adjacent bushes are often entirely defoliated
which may result in the death of several yards of hedge. The
pest attains its greatest abundance and destructiveness during
the fall months. Several insecticides were thoroughly tested
during the present (1914) season against this species. Schnarr’s
Insecticide gave complete mortality, with lime-sulphur pract-
ically as good. Following are the results of the test.
SPRAYS MORTALITY
Selim aiergemnse Chel ClG Cums creas ctiracs ees lee ees cea 100%
Lime-sulphur (Thomsen Chem. Co.)........:..... 99%
PerrASS i MAMGIA lads be. Se. forsy ston So ;x,ees.ws)d Sarstae Ses aoe 90%
molecu lel erie mieten cnr, costes traehie aie ore oes at Sea Less than 5%
Tetranychus yothersi sp. nov.
Predominating color a rusty-red, arising mainly from large internal
structures occurring on each side and connected centrally by a narrow
isthmus, a shield- or saddle-shaped pale pinkish-amber area includes
most of the cephalothorax; a narrow clear or translucent area extends
medially from behind almost to the thoracic suture. Eyes crimson,
each set at inner border of a groove overlying coxe I and II. Coxe
and femora of a greenish hue; tibize I and tarsi I salmon-color. Palpi
salmon-color. Dorsal bristles colorless, not arising from tubercles.
Body of female sphero-elliptical, widest equatorially: male subcuneate,
widest across cephalothorax which is somewhat truncate in front,
abdomen tapering to acute point posteriorly: bristles in four rows,
averaging in length two-fifths the width of the body. Mandibular
plate less than twice as long as broad, somewhat tapering anteriorly
with a distinct emargination. ‘‘Thumb” of palpus much reduced
longitudinally, bearing at its tip a relatively large, slightly clavate
“finger” whose base is almost as wide as the tip of the “thumb”’; on
its upper distal corner are two pseudo-fingers, not greatly thicker than
hairs, on upper side about midway to base is a ‘small “finger”? and
356 Annals Entomological Society of America [Vol. VII,
between this and base are two short stout hairs; the claw on the penul-
timate joint reaches to the middle of the “‘thumb’’; a hair arises laterally
from the center of the ‘‘thumb’’, and another from a similar position on
the penultimate joint. The legs are relatively short, barely as long as
width of body; femur only half again as long as wide—exactly equalling
tarsus, tibia a trifle longer than patella which equals the trochanter:
tip of tarsus bears a claw which is nearly straight for two-thirds its
length and then bent to nearly a right angle; a second claw, arising
from the other at its point of origin from the onychium, is almost
straight and forms with the first an obtuse angle; four strong spurs
(corresponding to the usual 4-cleft claw) have their origin in common
with the claws; the usual series of four capitate hairs arise by the sides
of the claws from the tip of the onychium.
The egg is globose-lenticular and bears a stalk which varies in
development from a length equalling the height of the egg to a mere
rudimentary papilla; guy fibrils are occasionally seen connecting the
egg with the leaf; the color is smoky-amber.
Type No. 19088, U. S. Nat. Mus.
The type material is from Orlando, Florida, August 28,
1914, from the upper surface of camphor leaves, collected by
W. W. Yothers. The species is evidently nearest T. mytil-
aspidis, Riley, from which it is easily distinguished through its
lack of dorsal tubercles, marked difference in the detail of the
palpal characters, emarginate mandibular plate, entirely dif-
ferent proportion one to the other of the leg joints, and through
the novel arrangement of the tarsal appendages.
An extensive series of measurements of material on Euca-
lyptus and camphor from Florida, and on oak, elm and pecan
from South Carolina have yielded the following averages:
ADULTS
| LENGTH
(not including WIDTH FORELEG
palpi)
Mem aller gigi carte ne Woe ere ce ee | .307 mm. .237 mm. .232 mm.
IE) eins eae Meee se, eh rae eras pte kt .225 mm. .152 mm. .222 mm.
EGG
| STALK
DIAMETER HEIGHT (when well
| developed
127 mm. | 082 Gum: | 077 mm.
aa a ie
1914] Four New Tetranychids 357
NOTES. °
It is of interest to record that, whereas the common red-
spiders have long been known to feed almost exclusively on the
under surface of the leaf, this species confines its activities
entirely to the top of the leaves.
To date, the species has been recorded upon camphor
(Camphora officinale) and Eucalyptus sp. at Orlanda, Florida,
arid upon two varieties of elm, the willow oak (Quercus phellos),
the white oak (Quercus alba), and the pecan at Batesburg,
South Carolina. Mr. Yothers states that at cetrain times it is.
everywhere present on the camphor tree causing a reddening
of the leaves and a reduced vitality of. the tree.
The species has been exceedingly abundant the past season
(1914) on the foliage of the small-leaved elm (Ulmus Ameri-
cana) to which as early as late June, it imparted a rusty appear-
ance. Trees thus injured have been observed at Batesburg
and Columbia, South Carolina, and Laurinburg, North Caro-
lina. During the seasons of 1911, 1912 and 1913 of the Bates-
burg investigations no evidence of the occurence of this species
had been seen. This indicates that the operation of certain
factors of natural control must have been suspended during or
just prior to the present season. Another observation of
interest, is that in spite of the exposure of this species on the
top of the foliage very little control seems to be exerted through
rains.
Tetranychus banksi sp. nov.
Color rusty-red, from underlying paired organs which occupy all
of the dorsal region excepting a median abdominal area and a clear area
containg the mandibular plate. Eyes (in mounted material) trans-
lucent, directly over suture between coxee I and IJ. The usual series
of dorsal bristles is lacking, but a series of 18 spatulate-serrate hair-like
appendages are distributed on the dorsal aspect of the body as follows:
One at either side of mandibular plate anteriorly, one just mediad of
each eye, one just overlying each coxa II, six forming a fringe at hind
margin of body and three along each side of abdomen. Body of female
rhombic-ovate, widest across cephalothorax, exceedingly obese for the
size of the legs; cephalothorax rounded generally anteriorly with a slight
concave border overlying the palpi: male almost sagitate in outline,
conspicuously reduced in proportion to the legs. Mandibular plate
about half again as long as wide, tapering somewhat anteriorly, with a
distinct emargination and with a superimposed chitinized ridge anter-
iorly. ‘“‘Thumb” of palpus subconical, upper surface twice trans-
versely depressed with an intervening dilation, bearing at its tip a long
358 Annals Entomological Society of America [Vol. VII,
slender ‘‘finger”’ which is over four times as long as thick; on its upper
side arising between middle and tip are two stout hairs, and near the
base of upper side arise a reduced “‘finger’’ and two stout hairs; the claw
of the penultimate joint reaches only to the basal ‘finger’; a hair
arises ventrally from the ‘‘thumb”’’, and another laterally from the
penultimate joint. Legs of female are of average length barely equal-
ling length of body; those of male are about twice as long as body:
femur between four and five times as long as thick—three-quarters
again as long as tarsus, tibia somewhat longer than patella which is
over twice as long as trochanter: relative length of joints as follows:
coxa 9, trochanter 3.75, femur 14, patella 8.75, tibia 10.9, tarsus 8: tip
of tarsus not provided with a claw—it being reduced to a vestigial
protruberance; the customary series of four capitate hairs arise from
the usual point.
Type No. 19089, U. S. Nat. Mus.
The type material from Orlando, Florida, August 16, 1913,
from the under surface of castor beans (Ricinus communts) and
velvet bean leaves. Collected by W. W. Yothers. Evidently
allied to T. latus of Europe.
NOTES.
Mr. Yothers states that the species is an important pest of
the castor bean plant in Florida but that at times it is controlled
by a predaceous mite (Sczulus sp.) and by the Coccinellid
Stethorus sp. Larve and pupe of Arthrocnodax carolina have
been observed on infested castor bean leaves from Orlando,
Florida.
An ample series of measurements of material on castor bean
from Orlando, Florida, have yielded the following averages:
| LENGTH
| (exclusive WIDTH FORELEG
of palpi)
Bemaler® -S6. ures ste ae tk nee .805 mm .267 mm. .295 mm.
INTER ret te, 0 ee Ah ne et ae .220 mm .197 mm. .407 mm.
Tetranychus quinquenychus, sp. nov.
There are a number of types of coloration but the general ground-
color is reddish-chestnut with the cephalothorax decidedly paler; the.
prevailing design consists of a large lung-shaped blackish area on each
side toward base of abdomen which coalesce medially toward the front,
a similar but smaller spot on each side near posterior end of abdomen:
legs and mouthparts pale. Body broadest midway between legs II
at
Js
ra
.
4
;
1914]. Four New Tetranychids 359
and III, tapering sharply forward to the narrow, slightly convex frontal
margin, also tapering considerably behind, twice as long as broad:
bristles rather long and fine, seven each in the dorsal rows and six each
in the sublateral rows, frontal pair half as long as subfrontal pair which
are placed just in front of the eyes. ‘‘Thumb” of palpus very short
and stout, on its tip is a blunt “finger” the basal width of which exceeds
its length, midway on the upper side is a “finger” equalling the terminal
“finger”? in length but very slender, at the upper distal corner are two
short hairs and two others occur at the upper proximal corner. Man-
dibular plate of average length with subparallel sides and convex at
tip with no emargination. Legs of moderate length; femur I two and
one half times as long as broad; tibia I somewhat longer than patella I;
tarsus in length equalling tibia and patella together, the tarsal append-
ages consisting of the usual series of four capitate hairs and a claw which
is sharply bent at middle at which point arises distally a strong spur
and proximally the usual four claw divisions. There is but a single
eye on each side which is set in a shallow submarginal socket directly
over coxa II.
Type No. 19087, U.S. Nat. Mus.
Collected at Orlando, Florida, September 28, 1914, on castor
bean (Ricinus communis), by Mr. W. W. Yothers. This
species appears to resemble somewhat J. tumidus Banks in the
character of the palpus but differs substantially as follows:
T. tumidus,—body moderately broad: subfrontal bristles not
twice as long as frontal pair: only 1 hair on palpal ‘‘thumb’’: sides
of mandibular plate narrowed toward tip and concave, tip
emarginated: terminal tarsal claw four-cleft. 7. quinquenychus,
body unusually narrow: subfrontal bristles twice as long as
frontal pair: four hairs on palpal ‘‘thumb’’: sides of mandibular
plate subparallel, tip not emarginated: terminal tarsal claw
five-cleft.
MEASUREMENTS OF FEMALE.
z ISCrimt Hee warty rea Satine Seatac: 455 mm
MUG Hells pth covehla & al S29) hate i Sere eee 5 eee 228 mm.
MOrele rene enn ee eo eae .do2 mm.
The relative lengths of the leg joints are as follows: trochan-
ter 10, femur 25, patella 18, tibia 19, tarsus 37.
360
Annals Entomological Society of America [Vol. VII,
EXPLANATION OF PLATES.
PLATE XLII.
Tenuipalpus bioculatus:
Fig.
1
D> OTs 09 bo
Front margin of cephalothorax: O, ocular spines; m, median spines;
e, eyes (greatly enlarged).
Female, dorsal view (enlarged 130 times).
Mouth parts showing left palpus (greatly enlarged).
Lateral outline of female (enlarged 130 times).
Tarsal appendages of left leg I, lateral view (greatly enlarged).
Hind margin of body, dorsal, showing series of 6 spines (greatly
enlarged).
All figures were drawn with aid of camera lucida, and figures 3 and 5
were drawn with oil-immersion lens.
PLATE XGLITI.
Tetranychus yothersi:
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
Tetranychus
Fig. 1.
Fig. 2.
Fig. 3.
Fig. 4.
Fig. 5.
BOG ete setae LOSING
Adult female (from Florida), dorsal view, enlarged 183 times.
Egg (lateral view) with stalk (from Florida), enlarged 196 times.
Extremity of left palpus (viewed from outside) showing ‘‘thumb’’,
“fingers’’, claw, and other appendages, greatly enlarged.
Outline and dorsal pattern of female (from Batesburg), enlarged
151 times.
Egg (lateral view) without stalk (Batesburg extreme form), enlarged
196 times.
Tarsal appendages (lateral view) showing onychium, claws, spurs
and capitate hairs, greatly enlarged.
Tarsal appendages (dorsal view), greatly enlarged.
Adult male (Batesburg form), outline and dorsal pattern, enlarged
129 times.
Figures 3, 6 and 7 were drawn with oil-immersion lens and camera
lucida.
PEATE Cl bV.
bankst:
Tarsal appendages, a, do1sal view; b, lateral view; greatly enlarged.
Extremity of right palpus (viewed from outside) showing ‘‘thumb’’,
‘fingers’, claw and other appendages, greatly enlarged.
Adult male, dorsal view, enlarged 156 times.
Adult female, dorsal view, enlarged 138 times.
Front margin of cephalothorax.
All figures drawn with the camera lucida; figs. 1 and 2 drawn with
oil-immersion lens.
PEATE GL Ve
Tetranychus quinquenychus:
Fig.
Fig.
Fig.
Fig.
Fig.
Fig.
il
Poo Ny
Tarsal appendages, ventral view.
Extremity of palpus showing ‘‘thumb’’, terminal ‘‘finger’’, dorsal
“‘finger’’, ‘‘thumb’’ hairs and penultimate claw.
Tarsal appendages, lateral view.
Left eye, seen from above.
Right foreleg, dorsal view.
Female, dorsal view (leg bristles not shown).
(Figs. 1, 2, 8 and 4 drawn with oil-immersion lens and camera lucida.
Fig. 6 enlarged 150 times.)
VoL. VII, PLATE XLII.
ANNALS E.S. A.
EB.AA. McGregor.
VOL. VII, PLATE XLIII.
ANNALS E. S.A.
£. A, McGregor.
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ANNALS E. S. A.
E. A. McGregor.
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—
INDEX OF VOLUME VII.
Adalia annectans, 213, 228, 229.
coloradensis, 213, 228.
flavomaculata, 84.
humeralis, 213, 228, 229.
melanopleura, 213, 228.
Adelocephala, 279, 282.
bicolor, 283.
bisecta, 283, 284.
Alexander, Chas. P. Article by, 239.
Anastatus bifasciatus, 86.
Anastrepha ludens, 202.
Anatomy of Siphona Plusiae, 301.
Anatomy of the Diaspinine Scale Insect
Epidiaspis Piricola (Del Guer), 47.
Anisota, 279, 286.
consularis, 286, 291.
senatoria, 286, 289.
skinneri, 286, 290.
stigma, 286, 287.
virginiensis, 286, 287.
Antochini, 240.
Anthomyia, 162, 164, 166, 167.
pluvialis, 162.
Anthomyiden, 160.
Anthomyioid Flies, Connectant Forms
Between the Muscoid and, 160.
Apanteles, 87.
lacteicolor, 87.
Aphidius testaceipes, 83, 84.
Aphidoletes marina, 216.
Aphis
atriplicis, 217.
brassicae, 217, 236.
carbocolor, 216, 236.
cerasifolii, 217, 236.
cornifolii, 218.
gossypii, 216, 224, 236.
heraclei, 218, 231, 2383.
helianthi, 215, 216, 221, 224, 231, 236.
medicaginis, 127, 236.
oenotherae, 217, 236.
oxybaphi, 216, 236.
pomi, 217, 224.
setariae, 215, 228, 224, 231, 233, 236.
taraxici, 236.
torticauda, 215, 216, 228, 236.
Apomecyna pertigera, 183.
Aptera, 316.
Araneus beebei, 170, 172.
microtuberculatus, 170, 172.
Aspihum sacculi, 65.
Aspidiotus perniciosus, 54, 55, 249.
piricola, 55.
Atlanta Meeting, Proceedings of, 97.
Attulus, 170.
Auchmeromyia, 162.
365
Automeris, 293.
incarnata, 297, 298, 300.
i0, 297, 298, 299.
leucaca, 297, 298, 299.
pamina, 297, 298.
Ballus tabupumensis, 170, 174.
Basilona, 279, 281.
imperialis, 281.
Baumberger, J. P., Article by, 323.
Bee Genus Coelioxys, Some Species of
the, 148. :
Beebe, Wm. C., Spiders collected by,
169.
Bellura melanopyga, 140.
Biology of the Net-Spinning Tri-
choptera of Cascadilla Creek, 251.
Bloeser, Wm., article by, 301.
Bomis, 170.
Bryobia pratensis, 73.
Byrsocrypta vagabunda, 67.
Calliphora, 160, 167.
erythrocephala, 162.
Calliphorinae, 166.
Calosoma sycophanta, 88.
Campodea, 316.
Carex, 218.
Castalia odorata, 136.
Caterpillars, Structural Study of
the, 109.
Cell Structure of the Digestive Epithe-
jium in Insects, 311.
Ceratitis, 202.
Ceratocampidae, Classification of the
Pupae of the, 277.
Chalcid Larvae, Note on the Number
of Spiracles in Mature, 249.
Cheiloneurus amplicornis, 248.
Childs, Leroy, article by, 47.
Chaitophorus negundinis, 215, 217, 221,
223, 231, 232, 236.
populicola, 217, 218, 223, 224, 231, 236.
populifolii, 215, 217, 231, 236.
Cheiloneurus, A New Species of, 247.
Chilomenes-lunatus, 84.
Chimarrha, 259.
aterrima, 259, 267.
Choeromyia, 162.
Citheronia, 279, 280.
regalis, 280.
sepulchralis, 280.
Classification of the Pupae of the
Ceratocampidae and Hemileucidae,
21.
Clinopera, 164.
Clubiona tabupumensis, 169, 171.
366
Cobanus Beebei, 170, 173.
Coccidae, 315.
Coccinella it
5-notate, 213, 219, 222, 224, 227, 234,
235.
9-notata, 213, 224, 226, 234, 235.
monticola, 213, 222, 224, 225, 227,
234, 235.
sanguinea, 213, 232.
Coccinellid, 217.
Coelioxys, 148, 166.
alternata, 150, 152.
altilis, 154.
asteris, 156.
banksi, 155.
comstocku, 148.
coquilletti, 157.
edita, 153.
hunteri, 151, 152.
insita, 158.
lucrosa, 148.
moesta, 148.
obtusiventris, 150.
octodentata, 154.
piercei, 152.
pratti, 159.
rhois, 159.
tufitarsis, 148.
rufitarsis varribois, 159.
sayi, 154.
sculptifrons, 153.
slossoni, 156.
slossoni arenicola, 156.
texana, 151, 152.
Coenosia, 161, 166.
irrorata, 244.
Connectant Forms Between the Mus-
coid and Anthomyioid Flies, 160.
Collembola, 315, 316.
Compsilura concinnata, 86.
Conwentzia Hageni, Banks, 73.
Cordyluridae, 135.
Cossidae, 119.
Cossus, 121.
Crane-flies, On a Collection of, 239.
Crawford, J. C., article by, 148.
Cycloneda
rubripennis, 232.
sanguinea, 232, 234.
Cyrnus, 262.
flavidus, 263.
pallidus, 263.
Dacus
cucurbitae, 177.
oleae, 202.
tryoni, 202.
Decatoma flava, 9.
Deilephila, 122.
Deilephila lineata, 119.
Dendroctonus, 315, 318.
Tree of Volume VII.
Diaspinine Scale Insect, Anatomy of
the, 47.
Dibrachys boucheanus, 249.
Dicranomyia illingworthi, 239.
saltens, 239.
Digestion of Insects, 311.
Dipoena tristis, 169, 171.
Diptera, 318.
Dispersal of Musca Domestica Linne,70.
Drassodes Drydeni, 169, 170.
ignobilis, 169, 171.
Drymeia, 161.
Dryocampa rubicunda, 279, 284, 285.
Dryophanta Erinacei and Its Gall, A
Study of, 1.
Enoplognatha marmorata, 169.
Epidiaspis Piricola, Anatomy of the
Diaspinine Scale Insect, 47.
Epithelium, Digestive, in Insects, 311.
Epochara canadensis, 203.
Erigone longipalpus, 169.
Eriococcus, 248.
Erioptera oceanica, 243.
Eriopterini, 241.
Eumusca, 166.
European Collection, Notes on Some
Id, 89.
Eurytoma auriceps, 9.
studiosa, 9.
Evophris albopatella, 170, 173.
Exochomus nigro-fasciatus, 84.
Fernald, H. T., article by, 89.
Fannia, 161, 162, 166.
Forbes, Wm. T. M., article by, 109.
Fundatrigenia ramulorum, 63.
Gahan, A. B., article by, 247.
Gasteracantha arcuata, 170.
frontata, 170.
Gillette, C. P., article by, 61.
Glossina, 161, 164.
Gonomyia fijiensis, 241.
varipes, 242.
Graphomyia, 166.
Haematobia, 162.
Haplopelma doriae, 169.
Hemileuca, 292, 293.
burnsi, 298, 294.
maia, 293, 294.
maia var. lucina, 294.
oliviae, 298, 295.
Hemileucidae, Classification of the
Pupae of the, 277.
Hemiptera, 317.
Hermetia illucens, 70, 71.
Hippodamia convergens, 213, 217, 219,
220, 221, 222, 224, 235, 236.
sinuata, 218, 217, 218, 219, 234.
parenthesis, 213, 218, 234.
Holocentropus, 262.
dubius, 262.
Index of Volume VII. 367
Houser, J. S., article by, 73.
Howard, Dr. L. O., article by, 86.
Hyalopterus arundinis, 217, 236.
Hydromyza Confluens, Life History
and Habits of, 135.
Hydropsyche instabilis, 255.
Hydropsychidae, 252.
Hypodermodes, 162.
Hypodermodes, 166.
Insects, Longevity of, 323.
Inquiline Life in the Gall of Dryo-
phanta erinacei, 7, 11.
Isopods, 316.
Lachnus, 217, 236.
Ladybeetles, Some
History of, 213.
Lasiocampa americana, 120.
disstria, 120.
Lasiocampidae, 119, 120.
Lecaniums, 315.
Lepisma, 316.
Leucauge tesselata, 170.
Leucomelina, 162, 163, 164, 165, 166.
squamopleura, 163.
Libnotes strigivena, 240.
Life History and Habits of Hydromyza
Confluens, 135.
Life History of Dryophanta Erinacei, 2.
Life History of Ladybeetles, Some
Notes on, 213.
Life History, Notes and Variations in
Wing Venation of Conwentzia
Hageni Banks, 73.
Life History of Siphona Plusiae, 301.
Limnerium disparidis, 87.
Limnobinae, 239.
Limnobini, 239.
Limnophora, 162, 163, 165, 166.
Linyphia, 169.
Longevity of Insects, 323.
McGregor, E. A., article by, 354.
Macrosiphum ambrosiae, 216, 217, 223,
228, 236.
cerasi, 215.
cynosbati, 217, 236.
gaurae, 215, 221, 232, 236.
pisi, 217, 236.
rosae, 215, 217, 221.
rudbeckiae, 215, 216, 217, 218, 224,236.
Malacosoma americana, 119.
disstria, 119.
Melanoxantherium amithiae, 236.
bicolor, 236.
Melanoxantherium bicolor, 217.
fraxinifolii, 217, 221.
smithiae, 233.
Melon Fly, The Ravages, Life History,
Weights of Stages, Natural Enemies
and Methods of Control of the, 177.
Mesembrina, 160.
Mongoma fijiensis, 243.
Notes on Life
Monodontomerus aereus, 88.
Moore, William, article by, 77.
Mordwilkoja oestlundi, 67.
vagabunda, 67.
Morellia, 162.
Mosher, Edna, article by, 277.
Musca Domestica Linne, Dispersal
of, 70.
Musciden, 160.
Muscina, 160, 162, 163, 166.
Muscoid and Anthomyioid Flies, Con-
nectant Forms Between the, 160.
Myospila, 166.
Myrmecophila, 317.
Myzus cerasi, 217, 224, 236.
persicae, 231.
Nephila clavata, 170.
maculata, 170.
Net-Spinning Trichoptera of Cascadilla
Creek, Biology of, 251.
Newcomer, E. J., article by, 311.
Noctuidae, 120.
Notodontidae, 119.
Natural Control of Toxoptera Grami-
num in South Africa and the
United States, 77.
Nephelodes, 119.
Neureclipsis, 260.
bimaculata, 260.
Noyes, Alice Ayr, article by, 251.
Nymphaea, 135.
advena, 1385, 146.
advena variegata, 136.
americana, 135, 136, 140, 1438, 144,
146, 147.
Observations on the Life History and
Habits of Hydromyza Confluens
Loew., (Diptera), 135.
Odonata, 316.
Officers, 1914, Entomological Society of
America, 95.
Olla abdominalis, 213, 232, 234.
Othellia, 162.
Orthellia, 166.
Oxyopes, 170.
Palmer, Miriam A., article by, 213.
Palystes, 169.
Panatala flavescens, 197.
Paralimna, 165.
Paralucilia macellaria, 70.
Pararicia, 164.
Parasites, Report on, 86.
Parasitic Life in the Gall of Dry-
ophanta erinacei, 7, 11.
Pemphiginae Attacking Species of Pop-
ulus in Colorado, Some, 61.
Pemphigus californicus, 61.
oestlundi, 67.
populiconduplifolius, 61, 63.
ranunculi, 61.
vagabundus, 67.
368
Petrunkevitch, Alexander, article by,
Phaeocyma, 119, 121.
Philodromus tabupumensis, 170, 173.
Philopotamidae, 259.
Philopotamus ludificatus, 259.
montanus, 259.
Pholcus phalangioides, 169.
Phorodon humuli, 217, 236.
Phryganidia californica, 301.
Plagiolepsis dustodiens, 78.
Plectrocnemia, 261.
conspersa, 261.
Polycantropus flavomaculatus, 262, 264.
.Polycentropidae, 250.
Populus in Colorado,Some Pemphiginae
Attacking Species of, 61.
Populus balsamifera, 61, 62.
occidentalis, 63.
tremuloides, 65.
trichocarpa, 61.
Porrhopis, 170.
Proceedings of the Atlanta Meeting, 97.
Prociphilus fraxinifolii, 215, 221, 231,
236.
Prospaltella perniciosi, 249.
Pseudohazis, 292, 296.
eglanterina, 296.
Pteromalus egregius, 88.
Pupae of the Ceratocampidae
Hemileucidae, 277.
Pygaera bucephala, 119, 122.
Pyrellia, 166.
Ranunculus californicus, 61.
Ravages, Life History, Weights of
Stages, Natural Enemies and
Methods of Control of the Melon
Fly (Dacus Cucurbitae Coq.), 177.
Report on Parasites, 86.
Resolutions on Death of P. R. Uhler, 96.
Rhagoletis cingulata, 203.
fausta, 203.
pomonella, 203.
Rhopholosiphum braggii, 219.
pastinaceae, 219, 231, 238.
Rhyacophylax, 252.
Samia cecropia, 297.
Saturniidae, 278.
Saturnioidea, 277.
Schedius kuvanae, 86.
Schizonerura lanigera, 215, 236.
Severin, Henry H. P., article by, 177.
Siphona Plusiae, Notes on the Life
History and Anatomy of, 301.
and
Index of Volume VII.
Somatic Muscles, Caterpillars, 109.
Some Old European Collections, Notes
on, 89.
Some Species of the Bee
Coelioxys, 148.
Spalangia hirta, 197.
Species of the Bee Genus Coelioxys,
Some, 148.
Sphecidae, 92.
Sphecodina abbotii, 119.
Sphex fumipennis, 91.
pensylvanicus, 91.
Sphingidae, 119.
Spiders Collected by Mr. C. William
Beebe in Burma and Borneo, 169.
Spilogaster, 163, 166.
Spirogyra, 262.
Stimulus to Gall Production, 12.
Stomoxys, 160, 162, 163, 164, 166.
Structural Study of the Caterpillars,109
Synergus erinacei, 10.
Synthesiomyia, 162, 163, 166.
Tachiniden, 160.
Tenuipalpus bioculatus, 254.
Tetrachychids, Four new, 354.
Tetranychus banksi, 557.
mytilaspidis, 73.
quinquenychus, 358.
yothersi, 355.
Teucholabis fijiensis, 240.
Thecabius patchii populiconduplifolius,
61
Genus
Theridion sarapus, 169.
Theridiosoma, 170.
Thiania, 170.
Tipulidae, 239.
Toxoptera Graminum in South Africa,
Comparison of Natural Control
Obedite
Tower, D. G., article by, 249.
Townsend, Chas. H. T., article by, 160.
Treasurer’s Report, 101.
Trichoptera of Cascadilla Creek, Bio-
logy of the Net-Spinning, 251.
Triggerson, Ci jq, arcicle bygale
Tropaea luna, 297.
Uhler, P. R., Resolutions on Death
of, 96.
Volucella obesa, 70.
Welch, Paul S., article by, 185.
Xanthogramma scutellare, 84.
Zelus peregrinus Kirkaldy, 197.
Zetek, James, article by, 70.
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ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA,
State University, Columbus, Ohio.
CONTENTS OF THIS NUMBER.
‘Noves, Miss Arice Avr—The Biology of the Net-
Spinning Trichoptera of Cascadilla Creek......-. 251
Mosuer, Epna—The Classification of the Pupae of
the Ceratocampide and Hemileucide...-....-.. 277
Brann Wi.Liam—Notes on the Life History and
eAnatomy of Siphona Phisize Cogice Sees 301
NEwcoMER, HE. J.—Some Notes on Digestion and the
Cell Structure of the Digestive leckarees in
Insects. cir late: KLE) eer as ea STY
| BAUMBERGER, J. Percy—Studies in the i anigévily of
SESOCIS AO are ee Tea aut ee Satie tar 323
McGrecor, EK. A.—Four New T ctranychids. BRS A st 54
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