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ILLINOIS BIOLOGICAL
MONOGRAPHS
Vol. VI October, 1920 No. 4
Editorial Committee
Stephen Alfred Forbes William Trelease
Henry Baldwin Ward
Published under the
Auspices of the Graduate School by
THE University of Illinois
CopnioaT, 1921 bt the Umvzssmr or Illinois
DiSTKIBUTED MaSCH 9, 1921
THE LARVAE OF THE
COCCINELLIDAE
WITH SIX PLATES
BY
J. HOWARD GAGE
Contributions from the
Entomological Laboratories of the Univerd^ of minds
No. 02.
THESIS
SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE
DEGREE OF MASTER OF SCIENCE IN ENTOMOLOGY
IN THE GRADUATE SCHOOL OF THE
UNIVERSITY OF ILLINOIS
1919
TABLE OF CONTENTS
PAGE
Intxoduction 7
Morphology 9
Head 9
Fixed Parts of the Head 9
Movable Parts of the Head 13
Thorax 17
Prothorax 17
Mesothorax and Metathorax 18
Appendages 20
Abdomen 21
Armature of the Body-wall 23
Synopsis of Larvae 27
Epilachninae 29
Coccinellinae 31
Chilocorini 32
Coccinellini 34
Microweiseini 42
Scymnini 43
Hyperaspini ., 45
Bibliography 48
Explanation of Plates 52
239] THE LARVAE OF THE COCCI NELLI DAE— GAGE
INTRODUCTION
The adults of this family are known to most persons as lady-bugs or
ladybirds. Their distinctive characteristics are the apparently three-
segmented tarsi and the broad hatchet-shaped distal segment of the
maxillary palpi. Other characteristics are: the eleven segmented anten-
nae, in which the distal segments are commonly modified to form a more
or less distinct club-shaped enlargement; the insertion of the antennae
near the mesal margin of the compound eyes; the ventral direction of the
mouth; the retraction of the head into the small transverse prothorax;
the transverse front coxae; the closed coxal cavities, except in Coccidula;
the convex elytra; and the abdomen consisting of five to seven exposed
ventral segments.
Le Baron has said, in speaking of this family, that "The Coccinellidae
occupy a remarkably anomalous and isolated position. Whilst having
the rounded form of the plant beetles, the clavate antennae of the scavan-
gers, and the dilated palpi of the fungus beetles, they agree in food and
habits with none of these, but resemble in their predaceous habits the
ground beetles and the soft-winged carnivora, all of which have their
bodies more or less elongated, their tarsi five-jointed, their antennae fili-
form, and their palpi slender or moderately dilated."
The larvae of the Coccinellidae, though they may be known to many
people, are not as a rule associated with the adult coccinellids or lady-bugs.
The most distinctive characteristics of these larvae are: their porcupine-
like appearance; elongated body which is usually striped or mottled with
red, black, white or yellow areas; small three-segmented antennae; power-
ful mandibles; and the habit of being continually on the move. From the
systematist's point of view these larvae do not show any unusual charac-
teristics such as Le Baron has noted for the adults, for they resemble in
most respects the distinctly predaceous tjrpes of coleopterous larvae.
This is true even of the Epilachninae, which are phytophagous. With
the exception of this subfamily the Coccinellidae are all more or less pre-
daceous in their adult stages, and almost entirely so in their larval stages.
The purpose of this investigation is to study the morphology of coc-
cinellid larvae and to arrange tables for the identification and classification
of a few of the more common species. The work has been limited to those
genera and species which have in the main been found in or reported from
Illinois. Specimens were collected and bred during the autumn of 1918,
and others were obtained from the collections of Dr. A. D. MacGillivray,
the University of Illinois, the Illinois State Natural History Survey, and
a specimen of Brachyacantha ursina received from Cornell University.
8 ILLINOIS BIOLOGICAL MONOGRAPHS [240
The investigation of the immature stages of insects was to a great
extent neglected by entomologists until about the beginning of the twen-
tieth century. There has been some previous work done, however, upon
the immature stages of the Coccinellidae. The works of L. Ganglbauer
(1899), Dimmock (1906), Palmer (1914), and Boving (1917) are valuable
to one pursuing a study of the immature stages of these beetles.
I am greatly indebted to Dr. A. D. MacGillivray, under whose super-
vision this work was carried on, for the interest he has shown at all times,
for the use of specimens from his collection, and for the privilege of using
the morphological nomenclature which he has devised. I am likewise
indebted to Professor S. A. Forbes for the use of material belonging to the
Illinois State Natural History Survey, and to Professor J. G. Needham
for the loan of material from Cornell University.
241] THE LARVAE OF THE COCCINELLI DAE— GAGE
MORPHOLOGY
This discussion of the general comparative morphology of coccinellid
larvae is based for the most part upon a study of Chilocorus hivulnerus.
This species represents the most generalized condition of the carnivorous
coccinellids that I have studied. A still more generalized condition occurs,
however, in the subfamily Epilachninae, the members of which are fof
the most part entirely phytophagous.
HEAD
The heads of coccinellid larvae are symmetrical and the general outline
is circular or nearly so, except in the genus Microweisea in which it is oval
or oblong. In Chilocorus and Epilachna the mouth is directed ventrad;
while in all of the genera of the Coccinellini, Hyperaspini, and Microweis-
eini it is directed caudo-ventrad. The greatest departure from the gen-
eralized condition is found in Scymnus. In this genus the mouth is directed
cephalad. For the sake of convenience the head will be considered under
two divisions; first, the fixed parts; second, the movable parts.
Fixed Parts of the Head
The fixed parts of the head consist of an external and an internal
skeleton. The external skeleton is composed of the fused front and post-
clypeus, preclypeus, veterx, labrum, and gula. The boundaries of these
sclerites are marked by distinct furrows or sutures. The internal skeleton
is made up of the floor-like tentorium, which in the Coccinellidae consists
of three parts.
In the head capsule of C. bivulnerus the epicranial suture (Fig. 6, es)
is present on the meson. It extends from the occipital foramen (Fig. 17,
of) to a point on the cephalic aspect about one-third the distance from the
occipital foramen to a line drawn through the antennal fossae. This part
of the epicranial suture is the epicranial stem (Fig. 6, es). The epicranial
stem bifurcates at its ventral end and the two epicranial arms (Fig. 6, ea)
extend latero-ventrad a short distance, then make a broad curve and
extend ventro-mesad on each side to a point where they become much
thickened. Each thickening is a pretentorina (Fig. 6, pt) and marks the
point of invagination of the pretentorium. The epicranial arms curve
broadly laterad and ventrad from each pretentorina to a point dorso-
mesad of an antennal fossa where they become obsolete. The three
sclerites included within or ventrad of the fork of the epicranial stem are
the fused front and postclypeus (Fig. 6, fc) and the labrum (Fig. 6, I).
There is an indistinct furrow which marks the position of the clypeal
10 ILLINOIS BIOLOGICAL MONOGRAPHS [242
suture (Fig. 6, cs) on each lateral margin of the head. There is a distinct
precoila (Fig. 6, pel), in which a preartis articulates, located on each side of
the postclypeus meso-ventrad of the antennal fossa at the point of origin
of the clypeal suture. The vertex occupies all of the dorsal and lateral
parts of the head capsule not included within the fork of the epicranial
suture (Fig. 6, v). There are six ocelli (Fig. 6, oc), in two groups of three.
Each group is situated on the lateral margin of the vertex dorsad and
laterad of the lateral end of an epicranial arm. The antennal fossae are
located ventrad of the ocelli on the dorso-lateral margin of the vertex.
The large somewhat oval opening in the caudal aspect of the head is the
occipital foramen (Fig. 17, of).
A primitive type of epicranial suture is found in the adults of Peri-
planeta and the larvae of Corydalis. The condition of the epicranial suture
in C. bivulnerus is very similar to that of these primitive forms, except that
the epicranial stem is not so long in proportion to the length of the epi-
cranial arms and that a portion of each arm is wanting near the antennal
fossae. This suture in Epilachna (Fig. 4, es) very closely resembles that
of C. bivulnerus, but the epicranial stem is much longer and extends almost
one-half the distance from the occipital foramen to a line drawn through
the antennal fossae. In Megilla the epicranial suture (Fig. 7, es) is present
but very short, not extending more than one-fifth the distance from the
occipital foramen to a line drawn through the antennal fossae; while in
Adalia, Anatis, Hippodamia, Coccinella, and Microweisea the epicranial
stem is not present and the epicranial arms diverge immediately from the
occipital foramen. In the adult larval stage of Hyperaspis the epicranial
suture is wanting, but it is present in the first instar. The epicranial stem
is absent in the second instar, but the epicranial arms are present; while
in the later larval instars the entire epicranial suture is wanting. The
adult larvae of Scymnus (Fig. 14) also lack an epicranial suture; no observa-
tions were made on the conditions present in very young larvae.
The epicranial arms are present in most coccinellid larvae, but are
absent in the adult larvae of Scymnus and Hyperaspis. In C. bivulnerus
the epicranial arms extend ventro-laterad from their point of origin to a
point dorso-mesad of the antennal fossae. In Epilachna the epicranial
arms extend ventrad of the antennal fossae, but do not extend as far
laterad. In Chilocorus and Epilachna the epicranial arms are not widely
divergent, but in all of the genera of the Coccinellini they diverge widely
and become obsolete slightly ventrad of the antennal fossae. In the first
larval instar of Hyperaspis they diverge immediately upon the dorso-
cephalic aspect of the head, the epicranial stem being very short, and they
extend laterad almost parallel with the caudal margin of the head, make
an abrupt turn and extend laterad and ventrad to their point of obsoles-
cence. In Microweisea (Fig. 13) the epicranial arms diverge gradually
243] THE LARVAE OF THE COCCINELLIDAE—GAGE 11
latere- ventrad and become obsolete dorsad of the antennal fossae. Due
to the extreme length of the head in this genus, these arms are very long.
In Scymnus the epicranial arms are entirely wanting.
The two proximal unpaired sclerites between the arms of the epicranial
suture in Corydalis are the fused front and post'clypeus. These areas are
separated from each other in the more generalized forms by the fronto-
clypeal suture; in the specialized forms, however, they may become com-
pletely fused. In such cases the fronto-clypeal suture is absent, but por-
tions of it may be indicated by a furrow on each lateral portion of the head.
These portions extend mesad from near the precoila. In C. bivulnerus
and all coccinellid larvae the front and post-clypeus (Fig. 6, fc) are com-
pletely fused and the fronto-clypeal suture is wanting. The clypeal suture
(Fig. 6, cs) is indicated by an indistinct furrow extending mesad from the
precoila (Figs. 5 and 6, pel). The area ventrad of this furrow in the pre-
clypeus (Fig. 6, pc) and the area dorsad of it is the fused front and post-
clypeus (Fig. 6, fc). In Epilachna (Fig. 4) the front and postclypeus are
entirely separated from the preclypeus by the complete clypeal suture;
while in all of the other genera of the family that were studied the condi-
tion of the cljqjeal suture is approximately that found in Chilocorus.
The labrum of C. bivulnerus is the distinct, slightly chitinized, shield-
like sclerite attached to the ventral margin of the preclypeus (Fig. 6, /).
The ventral margin of the labrum may be slightly emarginate and usually
bears four or six medium-sized setae. In general the structure of the
labrum in all of the coccinellids studied approximates very closely the con-
dition found in the labrum of C. bivulnerus. Its general shape, however,
varies in the different genera of the family. In the genus Epilachna the
labrum varies most widely from the Chilocorus type. In this genus it is
broadly transverse and widely but shallowly emarginate on the ventral
margin.
The vertex of C. bivulnerus (Fig. 6, v) consists of the paired continuous
areas on the cephalic and dorsal aspect of the head. In Chilocorus,
Epilachna, and Megilla the epicranial stem is present and marks the line
of separation of the two halves of the vertex; in Hippodamia, Coccinella,
Anatis, Adalia, and Microweisea the epicranial arms alone are present and
the two halves of the vertex do not meet on the meson, but a portion of the
front extends between them to the occipital foramen. In Hyperaspis and
Scymnus the epicranial suture is absent and the front, postclypeus, and
vertex are fused. The vertex is continuous on its lateral and caudo-
lateral margins with the genae (Fig. 5, ge), the region of the vertex ventrad
and mesad of the ocelli and the antennal fossae. The size, shape, and
extent of the vertex is dependent upon the location and extent of the epi-
cranial suture.
12 ILLINOIS BIOLOGICAL MONOGRAPHS [244
i4
In the head of Corydalis caudad of the antennae there is a distinct
ocularium which bears five or six ocelli. This indicates the position of the
developing compound eyes. In C. bivulnerus the ocelli (Figs. 5 and 6, oc)
are placed three on each lateral aspect of the head dorsad of the antennal
fossae. They are usually arranged in the form of a triangle. These ocelli
are undoubtedly the homologues of the ocelli of Corydalis and represent
the developing compound eye of the adult. The arrangement, number,
and position of the ocelli is fairly constant in all of the genera of the family.
There is only one exception worthy of note; the region bearing the ocelli
in Scymnus is darkened and slightly chitinized. Two of the ocelli are
equal in size, while the third is almost twice as large as either of the others.
In generalized insects the gula is present as a distinct chitinized sclerite
extending cephalad from the occipital foramen to the articulation of the
maxillae and submentum. It is bounded on its lateral margins by the
postgenae. In all of the larvae of the Coccinellidae the gula (Fig. 17, g)
is present as a more or less membraneous, rectangular, glabrous area caudad
and dorsad of the submentum.
The internal skeleton of the head of insects is formed by invaginations.
It serves undoubtedly to make the head more rigid, to support the soft
and delicate parts, and as a place for the attachment of muscles. The
entire internal skeleton of the head is known as the tentorium (Fig. 47).
It consists of two or three pairs of arms that have been invaginated from
the external skeleton. In the more primitive forms there may be only
two pairs. In all of the larvae of the Coccinellidae there are three distinct
pairs of arms.
The pretentoria, also known as the anterior arms of the tentorium
(Fig. 47, prt), are invaginated on the dorsal aspect of the head near the
point where the epicranial arms turn abruptly laterad.
The supratentoria, sometimes called the dorsal arms of the tentorium
(Fig. 47, sup), are invaginated mesad of the antennal fossae and are well
developed in the larvae of the Coccinellidae.
The metatentoria or the posterior arms of the tentorium (Fig. 47, met)
are invaginated on the ventral aspect of the head near the articulations of
the maxillae.
In many insects the occipital foramen is divided into two parts by a
distinct bridge. This bridge is the corpotentorium. In coccinellid larvae
the corpotentorium is not united to form a complete bridge, but the mesal
boundaries of the two lobes which ordinarily fuse t6 form the corpoten-
torium (Fig. 47, cp) approximate each other very closely on the ventral
aspect of the head and are located much nearer the mouth than the occipi-
tal foramen. The metatentoria extend dorsad from their point of invagi-
nation along the gular sutures toward the occipital foramen and form
important landmarks for the identification of the gula. When they
245] THE LARVAE OF THE COCCI NELLI DAE— GAGE 13
reach the occipital foramen, they diverge laterad and form a ring around
the entire opening.
The small bridge-like structure about one-half way between the lobes
of the corpotentorium and the occipital foramen is the laminitentorium
(Fig. 47, It). It is formed by the fusion of the ends of the pretentoria,
supratentoria, and metatentoria.
The small pit on the ectal surface of the dorso-cephalic aspect of the
head where each pretentorium is invaginated is a pretentorina (Fig. 4, pt).
The point on the ectal surface of the head marking the place of invagi-
nation of each supratentorium is a supratentorina (Fig. 6, supt). It is not
present as a pit in coccinellid larvae. The metatentorina is the point of
invagination of each metatentorium on the caudo-ventral aspect of the
head. The metatentorinae are not indicated by a pit or thickening.
Movable Parts oj the Head
The form, structure, and arrangement of the movable parts of the head
of coccinellid larvae are readily homologized with the movable parts of
the head of generalized insects, such as Periplaneta or the larvae of Cory-
dalis or the adults of the Carabidae. The movable parts consist of the
antennae, mandibles, maxillae, and labium.
The antennae of C. bivulnerus (Fig. 36) are about as long as wide,
slightly conical, and composed of three segments. The scape (Fig. 36, sc)
is cylindrical, wider than long, slightly chitinized, and bears a few fine setae.
The scape is attached to the heavily chitinized antennaria which bounds
the periphery of the antennal fossae (Fig. 36, ant) by a delicate membrane,
the antacoria (Fig. 36, ante). The pedicel or second segment (Fig. 36, pd)
is distinctly smaller than the scape, about as wide as long, and bears a dis-
tinct long seta on its mesal surface near the distal end. It also bears a
small number of fine setae. The flagellum, the small mound-like segment
on the distal end of the pedicel (Fig. 36, fl), usually appears to be a part of
the pedicel, but careful examination shows it to be distinctly separated
from it. The flagellum usually bears three peg-like setae which are prob-
ably tactile organs (Fig. 36, ts) and four small oval openings which are
probably sensoria (Fig. 36, se).
The antennae of Epilachna (Fig. 35) are about three times as long as
wide; the greater part of the elongation is found in the pedicel. In this
segment the peg-like seta at the distal end is clearly a part of the pedicel
and does not appear as a part of the flagellum as in the other genera of the
family. In all of the genera of the tribe Coccinellini and in the genus
Microweisea the antennae are less than twice as long as wide and the
antacoriae are protuberant and might easily be mistaken for the first
antennal segment. The condition of the remaining parts is similar to that
of Chilocorus. In Hyperaspis the antacoria (Fig. 38, ante) is more pro-
14 ILLINOIS BIOLOGICAL MONOGRAPHS [246
tuberant than in the Coccinellini, the flagellum is more mound-like, and
one of the three apical setae is much larger than the other two. Scymnus
(Fig. 39) presents a type of antennae that shows a wide variation from the
type found in the other genera of the family. The antenna of this genus
is wider than long, only slightly chitinized, scarcely elevated, and conical.
The antacoria (Fig. 39, ante) is narrow; the scape (Fig. 39, sc) is about three
times as wide as long; the pedicel (Fig. 39, pd) is about twice as wide as
long, its distal end much narrower than the proximal; and the flagellum
(Fig. 39, fl) is more or less mound-like and about twice as wide as long.
All of the coccinellid larvae examined have well-developed mandibles.
They may be of a crushing type, that is with many dentes as in Epilachna,
or they may be of a piercing type, that is with one or two dentes as in all
of the members of the subfamily Coccinellinae. Within the family there
are all stages of variation between these two extremes. The type of man-
dible found in such generalized genera as Periplaneta or Corydalis is that
with many dentes. This tends to show that in these forms the mandible
is a crushing organ primarily and not for piercing as appears to be the case
in the specialized coccinellid larvae.
The mandible of C. hivulnerus (Figs. 40, 41) is of moderate size, heavily
chitinized, somewhat triangular in outline, thick and heavy at the proximal
end, and tapering to a bidentate distal point. The proximo-mesal margin
of each mandible is provided with a distinct wedge-shaped mola (Fig. 41,
mo) and the proximo-lateral margin with a stout spine-like seta. On the
cephalo-mesal margin of each there is a preartis (Figs. 42, 41, ps) which
articulates in a precoila (Fig. 5, pel) ; on the caudo-mesal margin there is
a postartis (Fig. 41, poa) which articulates in a postcoila. The mandibles
of Chilocorus are intermediate in form between the two extreme types.
The majority of the species studied have mandibles similar to those of
Chilocorus and tend to show an evolution from the multidentate type
found in Epilachna, which are entirely phytophagous, to the unidentate
type found in Hyperaspis, Microweisea, and Scymnus, which are entirely
carnivorous. In all of the genera studied the mola is present, but it is
much reduced, is almost indistinguishable in Epilachna (Fig. 42, mo) and
shows the extreme condition of its reduction in Microweisea. The mandi-
bles of all coccinellid larvae are connected with the head capsule cephalad
and ventrad of the antennal fossae by a small and distinct membrane.
This membrane is the mandacoria (Fig. 5, mco). The extensor muscles
are attached to the lateral margin of the mandible between the preartis
and the postartis, while the retractor muscles are attached near the mesal
portion of the mandible.
The maxillae of the Coccinellidae show a greater degree of departure
from the primitive type than any other of the movable parts. They
resemble in general form the maxillae of Periplaneta and the larvae of
247] THE LARVAE OF THE COCCINELLI DAE— GAGE IS
Corydalis, but show a more striking resemblance to the conditions found
in the maxillae of carabid larvae. Though the coccinellid maxillae are
similar in form to those of Periplaneta or to those of the larvae of Corydalis,
they also show a great difference in structure and are much more specialized
than those of carabid larvae. In order to homologize the parts of the
maxilla, it was necessary to trace the development of this appendage from
the more generalized to the more specialized condition. In the tracing of
this development maxillae of the larvae of Carabidae, Lachnosterna,
Elateridae, and Curculionidae were found most useful.
Each maxilla of C bivulnerus is moderately large, slightly chitinized,
and consists of the following parts: a fused cardo and stipes, a palpifer,
a maxillary palpus, and a galea. The lacinia is apparently wanting.
In the primitive type of coleopterous larvae the cardines are composed
of two sclerites; the subcardo, which articulates with the postcoila and a
second sclerite, the alacardo, which lies between the subcardo and stipes.
This condition is found in the larvae of Pterostichus and Lachnosterna;
while in the larvae of a curculionid beetle, Phytonomus, the subcardo and
alacardo are fused. A similar condition is found in the larvae of the Ela-
teridae. In the Coccinellidae the subcardo and alacardo are not only
fused to form the cardo, but the cardo and stipes are fused and the extent
of the cardo is only indicated by small and in many cases indistinct notches
along the sides of the fused cardo and stipes. This indication of the.suture
between the cardo and stipes is found in the maxillae of all of the larvae
studied and there is but little variation shown. It will be seen from the
above that the stipes and cardo must be discussed as one sclerite. This
sclerite (Fig. 18, ss-{-ca) occupies the area at the proximal end of the maxilla
cephalad or ventrad of the gula. Its form in C. bivulnerus is that of a
rectangle. It is about as wide as long, slightly setaceous, and chitinized.
This general form of the stipes and cardo is found in all of the genera
studied except Microweisea, Scymnus, and Hyperaspis. In these genera
the stipes and cardo are narrow and elongated. In Microweisea and
Hyperaspis the proximal end is curved laterad. This curved portion
represents the cardo and the remaining part of the sclerite is the stipes.
The palpifer is present as a distinct shoulder-like area (Fig. IS, pf) near
the distal margin of the fused stipes and cardo. Its form and position are
generally constant in all of the genera studied. This area bears at its
distal extremity the three-segmented maxillary palpus. The maxillary
palpus (Fig. 18, tnxpl) is well developed in all of the genera. In C. bivuU
nerus they are stout and slightly chitinized. The first segment is cylindri-
cal, wider than long, and usually with one or two small setae. The second
segment is longer than wide and with one or two large setae. The third
segment is conical, about one-half as long as wide with two or three setae
and with a group of sensory organs at its distal end. The maxillary palpus
16 ILLINOIS BIOLOGICAL MONOGRAPHS [248
in Epilachna (Fig. 16, mxpl) is much elongated, the proximal segment is
nearly twice as long as wide, the second segment more than twice as long
as wide, and the conical third segment nearly three times as long as wide.
In all of the other genera the maxillary palpus resembles that of C. hivul-
nerus and the distal segment of each bears a group of peg-like organs,
tactile setae.
The galea (Fig. 18, go) in C. bivulnerus is the broad triangular area dis-
tad of the palpifer. Its shape and structure is more or less constant
throughout the family, the most notable exception being found in Hyperas-
pis (Fig. 27, ga). In this genus the galea is rounded and appears to be
more or less sponge-like and bears a few setae. In Epilachria (Fig. 16,
ga) the distal margin of the galea is densely setaceous, while in all of the
other genera of the family it bears only a few setae. There is a peg-like
structure on the galea that bears a striking resemblance to the distagalea;
this is nothing more than a tactile seta and cannot be interpreted as a
distagalea.
The mesal margins of the maxillae and the lateral margins of the labium
are connected in all of the genera by a distinct membrane, the labiacoria
(Fig. 18, lie). The lateral margins of the maxillae and the mandibles are
connected by a similar membrane, the maxacoria (Fig. 18, mxc). An
extension of the maxacoria connects the stipes and cardo of each side to a
postgena.
The labium of coccinellid larvae differs considerably from that found
in the more primitive forms, as the larvae of Corydalis and the adults of
Periplaneta. Many of the parts seem to be lacking. The type of labium
found in the more generalized coleopterous larvae shows but little resem-
blance to the coccinellid labium. By a study of the labia of the primitive
forms named and of the more generalized Coleoptera, one is able to homol-
ogize the parts.
The labium of C. bivulnerus is the more or less membraneous area
cephalad and ventrad of the gula and between the maxillae. It appears
to be made up of two parts, the submentum (Fig. 18, su) and the ligula
(Fig. 18, lig.). The mentum is indistinguishable or fused with the ligula.
The submentum (Fig. 18, su) is the ^arge proximal portion. It is mem-
braneous, more or less rectangular, usually with four setae, two large ones
which decussate, and two smaller ones. In Anatis, Megilla, Coccinella,
and Hippodamia there may be many setae on the submentum, the number
varying from four in Hippodamia convergens to eighteen in Anatis. The
submentum is not clearly or distinctly separated from the ligula in C.
bivulnerus but there is a distinct division between the two in Epilachna
(Fig. 16), while in Hyperaspis (Fig. 27) the division between the ligula and
submentum is entirely obsolete. In all of the other genera studied the
condition of this division approximates that found in Chilocorus. The
2491 THE LARVAE OF THE COCCINELLJDAE—GAGE 17
ligula is the distal portion of the labium. In C. bivtdnerus it is composed
of the fused stipulae, glossae, and paraglossae. Near the ventro-lateral
margin there is a distinct shoulder-like swelling, the palpiger (Fig. 18, pg),
which bears a two-segmented labial palpus (Fig. 18, lipl). There is a
heavy, semicircular, chitinous band (Fig. 18, cb) that surrounds each pal-
piger which probably serves to increase its rigidity. Each labial palpus
consists of two segments. The proximal segment is short, as wide as long;
while the distal segment is conical and bears a group of tactile setae at its
distal end. The ligula also bears four to six moderately large setae. The
type of ligula found in C. hivulnerus is remarkably constant within the
family, the only notable variation being found in Hyperaspis, where the
labial palpi have been reduced to a single dome-like segment which bears a
few tactile setae (Fig. 27, lipl).
THORAX
That part of the body caudad of the head consists of thirteen segments.
The dorsal surface of the segments is convex and the ventral surface con-
cave or flattened. They may be provided with scoli, senti, parascoli,
strumae, verrucae, or chalazae and setae. The variation in the type of
armature will be taken up later. The first three segments constitute the
thorax and the remainder the abdomen. In all of the genera of the family
the thoracic segments are distinctly separated from each other by a deep
coria, more distinct on the ventral than on dorsal surfaces.
Prothorax
The prothorax of C. hivulnerus (Figs. 1, 2 and 3, prth) is about two-thirds
as long as the mesothorax and metathorax combined. The same is true
of Epilachna, Microweisea, Scymnus, and Hyperaspis; while in all of the
genera of the Coccinellini the prothorax is about one-half as long as the
other thoracic segments together.
The tergum is usually convex and oval in outline when viewed from
above (Figs. 1 and 2, t). In C. bivulnerus the greater part of the dorsum
is covered with a dark heavily chitinized dorsal shield from which the
cone-like senti project. This shield is formed by the fusion of pinacula and
its surface bears numerous fine setae. The dorsal shield, when viewed from
above, appears to be rectangular in outline and is divided into two parts
by a slender white line extending along the dorso-meson. The senti are
arranged in three distinct groups, a cephalic group of six placed in a trans-
verse row near the cephalic margin, a lateral group of one placed near the
middle of each lateral margin, and a caudal group of one placed near the
middle of each half upon the caudal margin. In Epilachna the dorsal
shield does not cover such a large portion of the dorsum. It is more or
less oval in outline and bears only the cephalic row of four scoli. Its sur-
18 ILLINOIS BIOLOGICAL MONOGRAPHS [250
face is also covered with short fine setae. In all of the genera of the Coc-
cinellini the dorsal shield is large and covers the greater part of the dorsal
aspect. It is not so heavily chitinized as in Chilocorus and is divided
longitudinally into four plates. Two of these are adjacent on the dorso-
meson and each bears three or more large chalazae and many small
setae. The lateral plates are smaller than the mesal and bear a fringe of
large chalazae on their lateral and cephalic margins. In Microweisea and
Scymnus the dorsal shield is only indicated and very slightly chitinized.
It appears to extend over the larger part of the dorsum and bears several
large black setae. These are arranged in three transverse rows, a row on
the cephalic and caudal margins and another midway between them. In
Hyperaspis the dorsal shield is wanting and the tergum is only slightly
chitinized. It is traversed by three rows of prominent black setae arranged
as in Scymnus and Microweisea. There are in addition to these larger
setae, many smaller and inconspicuous ones.
The pleural area in C. bivulnerus is the more or less reduced area ven-
trad of the dorsal shield (Fig. 2, prpl). It extends ventrad to the sternal
area. There is a small group of setae cephalo-dorsad of the procoxacoila
in most of the genera studied. This is the only group of setae located on
the lateral aspect of the prothorax.
The prosternum of Chilocorus is the rectangular area located between
the coxal fossae. There has been much controversy among morphologists
as to the number of sclerites in this area. It is not my purpose to discuss
this question, there are, however, a few landmarks of this region that must
be considered. In all of the species studied there is a small pit, the fur-
dna, (Fig. 3, fur) found near the meso-caudal boundary of each procoxa-
coria. There is a distinct ridge extending between these pits which prob-
ably serves as a place for the attachment of muscles. The prosternum of
Chilocorus, Epilachna, Microweisea and Hyperaspis usually bears a few
small setae and chalazae just cephalad of the procoxacoria, while in all of
the genera of the Coccinellini it bears two distinct verrucae which are
located adjacent to the meson.
Mesothorax and Meiaihorax
In general the mesothorax and metathorax of coccinellid larvae are so
nearly similar that a single description will suffice for both of them. Each
of these segments is wider than long, but in other respects they are similar
to the prothorax in form.
The mesotergum in C bivulnerus (Fig. 1, mst) is distinctly longer and
narrower than the metatergum (Fig. 1, mtt). In practically aU of the
genera of the Coccinellini the mesotergum and metatergum are subequal
in length, but the metatergum is as a rule wider than the mesotergum. In
Microweisea, Scymnus, and Hyperaspis the mesotergum is longer and
251] THE LARVAE OF THE COCCI NELLJ DAE— GAGE 19
narrower than the metatergum. The mesotergum and metatergum of
C. bivulnerus do not bear dorsal shields. Each tergum bears a transverse
row of four senti, the pinacula of which are distinct and never fused.
The median senti are much smaller than the lateral ones. In Epilachna
there is a small median dorsal shield from which two scoli project, these
scoli originate so close together that they seem to be the two forks of a
single scolus. In all of the genera of the Coccinellini the dorsal shield is
present on the mesotergum and metatergum as a raised oval area which
covers the larger part of the dorsum. The mesal margin of this shield
bears a pair of small parascoli, while the lateral margin bears a large para-
scolus. The surface of the shield is densely covered with fine setae and in
the genera Coccinella, Hippodamia, and Anatis also bears chalazae. In
Scymnus and Microweisea the dorsal shield is weakly chitinized, covers
the larger part of the dorsum, and bears a small verruca on each side of the
meson. Hyperaspis, on the other hand, has no dorsal shield nor is there
any chitinization to suggest the presence of a shield. The terga bear many
long, black, prominent setae which are arranged in more or less transverse
rows; one row on the cephalic and one on the caudal annulet of each
segment.
The pleural areas of the mesothorax and metathorax of all of the species
studied are well developed. They are the lateral vertical areas between
the terga and the sterna (Fig, 2, mspl, mtpl). In C. bivulnerus the meso-
pleural area is much larger than the metapleural; this is also the case in
Microweisea, Scymnus, and Hyperaspis. In Epilachna the metapleural
area is larger than the mesopleural; while in all of the genera of the Coc-
cinellini they are subequal. The ventral portions of the mesopleural and
metapleural areas are obliquely crossed by a furrow extending caudo-
mesad from the coxacoila. In C. bivulnerus the mesopleural area bears
two subequal senti, one near the cephalic and one near the caudal margin;
the base of each of these senti is provided with a small pinaculum. The
metapleural area bears two senti, but the cephalic one is much smaller
than the caudal. In Epilachna the cephalic area bears a few small setae,
while the caudal one bears the large scolus. In the Coccinellini the
cephalic area usually bears a few small setae, and the caudal one is pro-
vided with a parascolus in Hippodamia, Coccinella, and Anatis; while in
Megilla and Adalia this area bears a struma. In Microweisea, Scymnus,
and Hyperaspis the cephalic area is smaller than the caudal area which
bears a small verruca in the two former, and a few fine setae in the latter.
The mesosternum and metasternum are so nearly similar to the pro-
sternum that no description is necessary.
In C. bivulnerus the mesothoracic spiracle is located in the small tri-
angular area lying between the mesotergum and the mesopleural area, the
protopleurite (Boving 1917). This condition is also found in all of the
20 ILUNOIS BIOLOGICAL MONOGRAPHS [252
members of the Coccinellini and Epilachna; while in Micro weisea, Sc>Tn-
niis, and Hyperaspis the mesothoracic spiracle is not located on the tergum,
or protopleurite, as Boving points out, but distinctly in the mesocoria.
The metathoradc spiracles are rudimentary or entirely wanting.
Appendages
The thorax of all coccinellid larvae bears three pairs of legs. One pair
is attached to each segment. They are well developed and fitted for walk-
ing or clasping the surface of the leaves and stalks upon which the larvae
are usually found. Since the general form and structure of each pair of
legs is the same, their only difference being that of size, a description of a
single prothoracic leg will suffice for all. The mesothoracic and meta-
thoradc legs are subequal and slightly longer and wider than the pro-
thoradc legs. Each leg is about as long as the body is wide, except in
very yoimg larvae in which they are distinctly longer. In adult larvae
they are comparatively stout and fitted for clasping. The coxal fossae
are the circular or oval holes in the sternum in which the legs are inserted.
The coxa are attached to the coxal fossae by a distinct membrane, the
coxacoria. There is a distinct coxacoila on the lateral margin of each
coxal fossa in which the coxa articulates. The procoxa (Figs. 2 and 3,
pre) is subcylindrical, short, tapering toward the distal end and bears a
few scattered but prominent setae. The protrochanter (Figs. 2, 3, prUr)
is short, triangular, the ventral surface the longer and bears a prominent
group of coarse setae. The profemur (Fig. 3, prfr) is short, about twice as
long as wide, with its dorsal, caudal, and cephalic surfaces sparsely setace-
ous. This condition of the femur is found in Chilocorus and Hyperaspis;
while in Epilachna and all of the genera of the Coccinellini, and the genera
Microweisea, and Scymnus, the femur is at least three times as long as wide.
The protibia (Figs. 2, 3, prii) is about as long as the profemur, one-third
as wide as long, and tapering distinctly toward the distal end. The
cephalic, caudal, and dorsal surfaces of the proximal two-thirds are densely
setaceous; while the entire surface of the distal one- third is densely covered
with dub-shaped setae or tenent hairs (Fig. 43, U). The tenent hairs are
very numerous in Chilocorus, Epilachna, and all of the genera of the
Coccinellini; while in Scynmus and Hyperaspis there are only a few, five
to ten, on each tibia, and in Microweisea there are only two which are very
broad, flat, elongated, and paddle-like (Figs. 44, 45). The protarsus
(Figs. 2, 3, prta) consists of a single short triangular segment which bears
a few tenent hairs. Its distal margin is provided with a sickle-shaped
daw (Figs. 4, 3, prd). This claw is provided with a short, blunt appendic-
ulated tooth on its proximo-ventral angle, and it probably serves as an
aid in clasping surfaces. There is some variation in the general shape
of the tarsal daw in the various genera of the family and this characteristic
serves as a means of separating them.
253] THE LARVAE OP TEE COCCI NELLI DAE— GAGE 21
ABDOMEN
The abdomen of all coccinellid larvae is composed of ten segments
which are connected by more or less distinct coria. This coria is usually-
more prominent on the ventral than on the dorsal aspect. The abdomen
is generally subdepressed, widest on its cephalic half, and tapering on its
caudal half. The abdomen in C. bivulnerus narrows gradually toward the
caudal end. The first three segments are about as wide as the metathorax.
This is also true of Epilachna and of all of the genera of Coccinellini in
which the first, second, third, and fourth abdominal segments are subequal
in width. The remaining abdominal segments become narrower toward
the caudal end. In Microweisea and Scymnus the first, second, and third
abdominal segments are the widest; caudad of the third the abdomen
becomes narrower. In Hyperaspis the first, second, fifth, and sixth seg-
ments are subequal in width; while the third and fourth are the widest
segments in the body.
In C. bivulnerus the first abdominal segment is slightly narrower and
shorter than the metathorax. Its tergum (Figs. 1, 2) bears four distinct
senti arranged in a transverse row. The dorsal senti are adjacent on the
dorso-meson and the dorso-lateral ones are placed on each side near the
lateral margin of the tergum. There is a small circular pinaculum at the
base of each sentus. The dorsal pinacula are brown or yellow colored and
the dorso-lateral ones are white. There is also a few inconspicuous setae
on the surface of each pinaculum. The lateral aspect is a vertical area,
almost square, and bears the large lateral sentus, at the base of which there
is a very large pinaculum nearly covering the entire surface. The para-
lateral group is wanting. The sternum (Fig. 3, st) is about as long as the
tergum. The coria between it and the metatergum is not at all distinct.
The cuticle is thin and membraneous and bears two groups of small ventral
setae adjacent to the ventro-meson. The ventro-lateral setae are wanting.
The external structure of segments two to five inclusive is similar,
however, each succeeding segment is narrower than the preceding one.
The coriae between the segments are distinct. Each tergum has four
senti arranged in a transverse row. The dorsal senti and their pinacula
are adjacent to the dorso-meson; the dorso-lateral senti are located in a
position similar to those of the first abdominal segment. All of the pina-
cula are dark brown in color and bear numerous fine setae. Each lateral
aspect is almost square and bears a distinct lateral sentus. The pinacula
at the base of these senti are small. In the second and third segments
ventrad of the lateral senti there is a solitary small seta; while in segments
four and five there is a prominent chalaza surrounded by a group of small
setae. This group represents the paralaterals. Each sternum is as long
as its respective tergum and the coriae between the sterna are distinct.
The cuticle is thin and membraneous and bears on each sternum four groups
22 ILUNOIS BIOLOGICAL MONOGRAPHS [254
of chalazae arranged in a transverse row; those near the meson are the
ventral chalazae and those on the lateral margin are the ventro-laterals.
Each chalaza is surrounded by a small group of setae.
The sixth, seventh, and eighth abdominal segments are as long as the
preceeding but narrower. The pinacula at the bases of the dorsal senti
have become fused on their mesal margins and appear to be dumb-bell-
shaped. The dorso-lateral senti are shorter than those of the preceding
segments and their pinacula are very much reduced. The lateral aspect
of each of these segments is similar to that of the preceding, except that
it is considerable smaller. There is a small lateral sentus with a very small
pinaculum on the lateral aspect of the sixth and seventh segments and
almost a rudimentary sentus on the eighth segment which is without a
pinaculum. In Epilachna the lateral aspect of the seventh segment is
provided with a distinct parascolus rather than with a scolus as in the
sixth; while the eighth segment has a struma and the ninth only a
few chalazae. There is a distinct chalaza surrounded by a group of fine
setae ventrad of the lateral sentus of each segment. This chalaza and the
setae represent a paralateral group. Each sternum is as wide and as
long as its respective tergum. The cuticle of each segment is soft and
bears four distinct chalazae arranged in a transverse row. Each chalaza
is surrounded by a group of small setae. The ventral groups are adjacent
to the ventro-meson and the ventro-lateral groups are located near the
extreme lateral margin of each segment. The coriae between the seg-
ments are distinct.
The ninth abdominal segment is small, cylindrical, slightly narrower
and shorter than the eighth. The tergum is inclined ventrad at an angle
of about thirty degrees. The dorsal senti are absent and in their place
there are distinct dorsal strumae. The dorso-lateral senti are also reduced
to strumae and located near the lateral margin of the tergum. The lateral
aspect is small and the senti are wanting, but there is a prominent chalaza
surrounded by a group of setae near the ventral margin. This group
represents the lateral group; the paralaterals are wanting. The ninth
sternum is short, deeply emarginated on the caudal margin, and bears four
chalazae arranged in a transverse row. The ventral chalazae are located
adjacent to each other and the ventro-meson and are surrounded by a
group of fine setae; while the ventro-lateral are without setae. Through-
out the family the ninth segment shows the greatest variation in its size,
shape, and structure. In Chilocorus the ninth abdominal segment is
rectangular, about twice as wide as long; its anterior margin is little if any
narrower than the eighth segment; the caudal margin is sharply rounded
and never serrated; and its lateral aspect bears chalazae and setae. The
lateral aspect of the ninth segment in Epilachna is rectangular, about
twice as wide as long, much narrower than that of the eighth segment, and
2551 THE LARVAE OP THE COCCINELLIDAE— GAGE • 23
with the caudal margin broadly rounded. Each lateral aspect bears a
small chalaza. In all of the genera of the Coccinellini the ninth segment
is more or less rectangular. The caudal margin is broadly rounded, usually
widest near the middle, distinctly narrower than the eighth segment, and
never crenulate or serrate. The lateral aspect is provided with numerous
setae. In Microweisea the ninth segment is conical, about twice as long
as wide, and much narrower than the eighth. The distal margins are
sharply rounded and the tergum is chitinized and brown. The lateral
aspect is provided with a few fine setae. In Scymnus the ninth segment
is conical, about one-half as wide as long, much narrower than the eighth
and not heavily chitinized. In Hyperaspis the ninth segment is semi-
circular, about twice as wide as long, and narrower than the eighth. The
caudal margin is broadly rounded and never crenulate or serrate. The
lateral aspect bears a few fine setae.
The tenth abdominal segment is as a rule not visible from the dorsal
aspect. It appears to be a small ring of thin membrane surrounding the
rosetted anal area. In the Coccinellini this membrane is often pigmented
and appears black or brown colored. The rosetted appearance of the anal
area is caused by the evagination of the rectum. This serves as a sucking
disk and aids the larvae in locomotion.
In all the coccinellid larvae there are eight pairs of abdominal spiracles
(Figs. 1 and 2, ahdsp), a pair situated on each abdominal segment from one
to eight inclusive. They are located near the cephalic margin of each
tergum between the dorso-lateral and lateral senti in Chilocorus and Anatis,
between the dorso-lateral and lateral scoli in Epilachna, between the dorso-
lateral and lateral parascoli, strumae or verrucae in Hippodamia, Coc-
cinella, Megilla, Adalia, and Scymnus, and between the dorso-lateral and
lateral chalazae or setae in Hyperaspis.
There is a pair of repugnatorial pores on segments one to eight (Figs.
1, 2, rp). These pores are located on each lateral margin of the tergum
in the coria between the segments. They emit a bad smelling fluid which
is of a repulsive nature and serves to protect the larva from its enemies.
On each side of the dorso-meson in segments one to eight there are two
small pits. There is another pair on each side of the tergum near the mid-
dle about midway between the dorsal and dorso-lateral senti of segments
one to nine. All of these pits are arranged in a transverse row on the
tergum. There is another pit on the lateral aspect of segments one to
nine which is located immediately ventrad of the ventral senti. These
pits are probably the points of attachment for muscles.
ARMATURE OF THE BODY-WALL
The armature of coccinellid larvae consists of spine-like setae on the
body- wall or of conical, finger-like, or mound-like projections of the body-
24 ILLINOIS BIOLOGICAL MONOGRAPHS ^S6
wall which bear setae. They are known as scoli, senti, parascoli, strumae,
verrucae, chalazae, or setae and vary greatly in form in the different
tribes and genera of the family. After a study of the larvae of the Chry-
somelidae, the probable progenitors of the Coccinellidae, one becomes
convinced that Chelymorpha shows perhaps the greatest resemblance to
the generalized coccinellids. In Chelymorpha each lateral margin of the
body is provided with a longitudinal row of long branched projections of
the body-wall. The distal end of each of these branches bears a stout seta.
Fracker in his work on lepidopterous larvae called structures similar to
these scoli. It was unfortunate, however, when he applied the same term
to a non-branched projection of the body-wall which bears setae upon its
trunk. These two structures are so widely different that they cannot be
considered as one and the same thing and for the latter the name sentus is
proposed. Fracker has also shown that the arrangement and number of
setae on the prothorax represents the generalized condition in lepidopterous
larvae. This may be true in coccineUid larvae, but there has been no
attempt made in this work to homologize the setae or the projections that
bear them. Since the arrangement of the setae in the various genera
differs, especially on the abdominal segments, this character has been
used to some extent in separating genera, and it is, therefore, necessary to
adapt a tentative nomenclature for these structures. This nomenclature
is based upon the concKtions found in the third abdominal segment and
has been applied only to the segments of the abdomen.
There is a seta or a projection bearing a seta or setae on each side of the
dorso-meson. These are designated as the dorsal group. The projection
on each lateral margin of the tergum is a dorso-lateral group, the one on
the dorsal portion of the lateral aspect is a lateral group, the one on the
ventral margin of the lateral aspect is the paralateral group, the small
group on each lateral margin of the sternum is a ventro-lateral group, and
the one on each side of the ventro-meson is the ventral group.
The scolus is a branched projection of the body-wall, usually more than
five times as long as wide (Fig. 28). Each branch of the scolus bears at
its distal end a single stout seta. The dorsal and lateral surfaces of the
thorax and abdomen of Epilachna are provided with distinct scoli.
The parascolus is a modification of the scolus in which the projection
is not more than three times as long as wide and usually not more than
twice (Fig. 30). This structure bears a few short branches which are about
as wide as long, each with a seta at its distal end. This modified scolus is
designated as a parascolus. It is found in Hippodamia, Coccinella, and
on the caudal segments of Anatis.
A sentus is an elongated, cone-Uke projection of the body-wall which
is not branched hke a scolus, but bears a few short stout setae upon its
trunk (Fig. 29). Fracker called this a scolus, but it differs decidedly in
257] THE LARVAE OF THE COCCINELLIDAE—GAGE 25
form from the true scolus and has been called a sentus. Senti are found
in Chilocorous and Anatis. In the latter genus the senti are 'short and
thick.
A pinaculum is the more or less chitinized plate which surrounds the
base of a scolus, sentus, or parascolus (Figs. 28, 29, 30). It usually bears
numerous, small, dark-colored setae. Several pinacula may become fused
to form the shield-like plates of the body.
A chalaza is a distinct but slight pimple-like projection of the body-wall.
It may be considerably wider than long and bears on its distal end a stout
seta (Fig. 33). Chalazae are present in practically all coccinellid larvae.
They are for the most part found on the sternum and in some cases on
the lateral aspect. In Microweisea distinct chalazae are found on the
dorsal aspect of the abdomen, there are also distinct chalazae on the dorsal
shield in most of the genera of the family.
A struma is a parascolus which has become shortened, usually appearing
to be nothing more than a distinct mound-like projection of the body-wall
(Fig. 31) upon which are situated a few chalazae. This structure was for-
merly called a tubercule, but this term is misleading and has been applied
by various workers to most any kind of an extension of the body-wall.
In order to avoid further confusion the term struma is proposed for this
structure. Strumae are found in the armature of the abdomen of Adalia
and Megilla.
The struma becomes much reduced in some of the more specialized
larvae so that it appears to be mound-like and bears setae instead of
chalazae (Fig. 32). This structure has also been called a tubercule by
some workers and by others a verruca. Since the term verruca is not
misleading or conflicting, it should be restricted to structures such as these.
Verrucae are found in the armature of Scymnus and very small indistinct
ones in Microweisea; the latter genus also bears chalazae.
In the most specialized of the coccinellids verrucae and chalazae have
become so greatly reduced that the setae are not elevated above the
general surface of the body (Fig. 34). Setae are found on the body of all
of the more generalized coccinellids, but it is only in the more specialized
groups that the setae constitute the only type of armature. The Hyper-
aspini is the only tribe that I have studied which has this condition.
In the niore generalized genera the armature of the body consists of
scoli arranged upon the dorsal and lateral surfaces. Such a condition as
this is found in the subfamily Epilachninae. In Chilocorous and Anatis
the scoli are replaced by senti on the dorsal and lateral aspects and on these
regions in Hippodamia, Coccinella, and to a small extent in Adalia para-
scoli are present. The projections of Adalia seem to show a stage of
transformation between parascoli and strumae; for parascoli are found on
the dorsal surface and strumae on the lateral. Strumae are found on the
26 ILUNOJS BIOLOGICAL MONOGRAPHS (258
dorsal and lateral regions of Megilla, but on the lateral aspect of the eighth
segment there is a verruca. Verrucae are found almost exclusively on
Scymnus, while in Microweisea they are very small and closely approxi-
mate the form of chalazae. Verrucae are also found on the ventral por-
tion of the ninth segment in practically all genera of the Coccinellidae, the
only exceptions being Hyperaspis and Scymnus. Chalazae are found on
the dorsal shield and on the shield-like spots of the mesothorax and meta-
thorax of Hippodamia, Megilla, Anatis, Adalia, and Coccinella. There are
also a few on the prothoracic shield of Scymnus. The same type of arma-
ture is present on the ventral parts of the thoracic and abdominal seg-
ments throughout the family and on the lateral portions of the eighth and
ninth segments of Adalia, Megilla, and Scymnus. They are usually sur-
rounded by a group of finer setae. In Hyperaspis and its allies the arma-
ture of the body has been so reduced that it is composed of setae forming a
hair-like covering over the entire surface. The ventral surface of all of the
members of the family is provided with setae rather than scoli, parascoli,
senti or strumae. Even in those genera where chalazae or verrucae are
found on the ventral surface, setae are also abundant. The type of arma-
ture of the Coccinellidae shows a steady and unbroken series of changes in
specialization from the generalized scolus to a seta through the reduction
or the disappearance of parts.
259] THE LARVAE OF THE COCCINELLIDAE—GAGE 27
SYNOPSIS OF LARVAE
In the taxonomic study of any group of organisms, the investigator
should not draw conclusions from characteristics which upon the surface
may show a high degree of specialization or generalization, without first
making a careful study of these characteristics, no matter how important
or unimportant they may seem. According to Comstock, "The logical
way to go to work to determine the affinities of the members of a group of
organisms is first to endeavor to ascertain the structure of the primitive
members of this group; and then endeavor to ascertain in what ways these
primitive forms have been modified by natural selection." With such a
view as this in mind, the taxonomist must not only study the members of
the group upon which he is working, but he must also endeavor to ascertain
the conditions that existed in the progenitors of the group. It is obvious,
then, that those conditions in the chosen group which are most nearly
similar to the conditions in their progenitors are the most generalized;
further, that those individuals possessing these primitive characteristics
are the generalized individuals of the group. There are often two or more
sets of prominent characters, and many times these may not run in parallel
lines, but seem to contradict each other or to run in opposite directions.
For instance in the larvae of the Coccinellidae that form which shows the
most generalized condition of the head sutures, the Epilachninae, have
also what seems to be the most highly specialized condition of the setae
or scoli on the body; and Hyperaspini which have the most highly special-
ized condition of the head sutures show what appears to be the most
generalized condition of the setae.
If one studies the conditions present in the Chrysomelidae, the prob-
able progenitors of the Coccinellidae, he will find in the genus Chelymorpha
conditions of the epicranial suture and scoli similar to those found in the
Epilachnini. The epicranial stem in Chelymorpha is long and the epi-
cranial arms are gradually divergent, a condition almost identical with
that of the Epilachninae. The members of the subfamily Coccinellinae
have a much shorter epicranial stem or in many genera it may be entirely
wanting and the epicranial arms diverge immediately from the occipital
foramen. Even the epicranial arms are wanting in the adult larvae of
Scymnus and H)rperaspini; while the clypeal suture, which is distinct and
entire in Chelymorpha and Epilachna, is incomplete and only indicated
28 ILLINOIS BIOLOGICAL MONOGRAPHS [260
on each margin of the head in the Coccinellinae. This fact tends to show
a complete reduction of the sutures which are distinct in Epilachna and
are entirely wanting or only slightly indicated in the Hyperaspini.
The condition of the armature of the body also shows a like reduction.
In both Epilachna and Chelymorpha scoli are present. In Chilocorus the
scoli are replaced by senti and in those forms which show a further reduc-
tion of the epicranial arms we find that there is a further reduction in the
armature of the body. In Hippodamia, Coccinella, Adalia, Anatis, and
Microweisea, forms in which the epicranial arms alone are present, the
armature consists of parascoli, strumae, verrucae, and chalazae or setae;
while in the Hyperaspini in which the epicranial suture is wanting the body
is provided only with setae. The condition of the scoli in Epilachna might
easily be taken as a highly specialized characteristic, but when one studies
the characteristics of the progenitors of the CoccineUidae, he finds such a
condition in the armature of the body as is found in the Epilachninae.
There is a further likeness between the Chrysomelidae and the Epilach-
ninae that seems also to be of importance, that is the food habits of the
two are almost identical, as 'both are phytophagous. The Epilachninae
are perhaps the only group of coccinellids that are entirely phytophagous
in both the larval and adult stages.
The fact that the setae in Hyperaspini show an apparently generalized
condition, while the epicranial suture is absent in the adult larval stages
and present in the first larval stages, surely a specialized condition, does
not interfere with this proposed classification. For specialization, as
Comstock has pointed out, may take place in two wholly different ways.
"First, by the addition or complication of parts, specialization by addition;
second, by the reduction in the number or the complexity of the parts,
specialization by reduction." The latter is considered to be the case in
the Hj^eraspini; the primitive scoli have been reduced to setae. Granting
this to be true, we can readily see that these two wholly different charac-
teristics, the condition of the scoli and the epicranial suture, show in a
very striking way the presence of specialization in Hyperaspini and of
generalization in the Epilachninae. In the first case there is the absence
of the epicranial stem, only an indication of the clypeal suture, and a
reduction of the scoli to setae; while in the latter there is the presence of
the epicranial and clypeal sutures and of scoli similar in form to those of
Chelymorpha.
Since, as Comstock has shown, there is such a thing as specialization
by reduction and since the progenitors of the coccinellids, as nearly as we
can ascertain, have an epicranial and a clypeal suture and a well-developed
system of scoli, it seems to me altogether logical, and with the evidence
at hand quite clear that the Epilachninae represent a generalized type of
coccinellid larvae, though at a first glance they may appear to be highly
specialized.
261] THE LARVAE OF THE COCCI NELLJ DAE— GAGE 29
There is a great variation in the general structure of the larvae of the
Coccinellidae. This variation ranges from the phytophagous type found
in Epilachna to the extreme carnivorous type found in Scymnus and
Hyperaspis. The larvae of the family possessing the phytophagous type
of structure can very easily be mistaken for certain chrysomelid larvae
which they resemble in general shape and in the arrangement of their
scoli and pinacula, while those of the more carnivorous types might at
first glance be mistaken for chrysopid larvae. The coccinellid larvae may
be distinguished from this latter group by the development of the mandi-
bles which are not so prominent in the coccinellids. The mandibles of all
of the carnivorous coccinellids differ from those of the Chrysomelidae in
that they are not so broad and have a smaller number of dentes; while in
Epilachna they resemble very closely the chrysomelid mandibles. All
of the coccinellid larvae examined possess three ocelli on the lateral por-
tions of the head, while the chrysomelid larvae may possess from one to
six or none on each side.
The Epilachninae are undoubtedly the most primitive type of coc-
cinellid larvae, a fact which is shown by their likeness to their chrysomelid
progenitors. Among the carnivorous coccinellids, Chilocorus is the most
primitive, while Hyperaspis, which is the farthest removed from the
chrysomelid larvae, shows the most specialized condition.
SUBFAMILIES OF COCCINELLIDAE
Epicranial suture present, epicranial stem extending one-half the dis-
tance from the occipital foramen to a line drawn through the anta-
coriae; clypeal suture present; antennae slender, more than three
times as long as wide; body with scoli Epilachninae
Epicranial suture present or wanting, when present, the epicranial stem
never extending one-half the distance from the occipital foramen to
a line drawn through the antacoriae; clypeal suture never complete,
only indicated on each side; antennae short, never more than twice
as long as wide; body without scoli Coccinellinae
EPILACHNINAE
The body is elongate, oval to fusiform. The dorsal and lateral aspects
are armed with scoli, the sternum with strumae or chalazae, and the head
with a few long setae. The epicranial stem and epicranial arms are always
present; the clypeal suture is entire and distinct. The antennae are more
than three times as long as wide, inserted more than their own length
dorso-mesad of the precoila. Each mandible is heavily chitinized, its mola
not well developed, and the distal portion with several dentes of various
lengths, the distal dentes the longest.
This subfamily is represented in North America by a single tribe, the
Epilachnini. Casey says, "This tribe is represented in the United States
30 ILUNOIS BIOLOGICAL MONOGRAPHS [262
by two or three large pubescent species belonging to the single genus
Epilachna." One species occurs in small numbers in southern Illinois.
The Epilachninae are truly phytophagous. Our common species feeds
almost entirely upon the squash plant and its relatives. The beetle is
commonly called the squash lady-bug. This subfamily is represented in
the material studied by a single species of the genus Epilachna Chevrolat.
Epilachna borealis Fab. — The prothorax is slightly chitinized with a
transverse row of four scoli on the dorsum near its cephalic margin and a
transverse row of small setae on its caudal margin; the pleural area is small
and glabrous; the prosternum is short with two distinct setae on the
ventro-meson; and the procoxacoriae are distant. The mesothorax and
metathorax are subequal in length and width, the dorsum of each with
four scoli, two on each side of the meson arising from the same pinaculum;
the mesothoracic spiracles are located in the mesocoria; the metathoracic
spiracles are wanting; the caudal portion of the mesothoracic and meta-
thoracic pleural areas are each provided with a large prominent scolus;
the mesothoracic and metathoracic sterna are each provided with a group
of setae on each side of the ventro-meson; and the mesocoxacoriae and
metacoxacoriae are distant. The coxa is short and subcylindrical; the
trochanter is triangular, about as long as the coxa and bears a few setae;
the femur is as wide as the trochanter, about twice as long and covered
with numerous small stiff setae; the tibia is about as long as the femur and
about two-thirds as wide, its dorsal and lateral surfaces are covered with
short stiff setae, its ventral surface is thickly covered with fine setae, and
the distal one-third is covered with tenent hairs; and the tarsus consists
of a single segment which bears the heavily chitinized hooked claw and
a few tenent hairs. The terga of the abdominal segments one to eight
inclusive are similar. Each tergum is provided with four scoli, the dorsal
scoli are adjacent to each other and the meson and the dorso-lateral scoli
are on the lateral margins of the tergum. The ninth tergum bears four
strumae which represent the dorsal and dorso-lateral groups; the tenth
segment is membraneous. Each lateral aspect of segments one to six is
provided with a prominent lateral scolus, of segments seven and eight with
a lateral struma, and of segment nine with a lateral verruca. The para-
lateral group is represented by a prominent chalaza surrounded by a group
of small setae ventrad of each lateral scolus. Sternum one is provided
with two prominent ventral chalazae, the ventro-laterals are wanting;
sterna two to nine bear two adjacent ventral chalazae and two ventro-
lateral chalazae located near the lateral portion of each sternum; and
sternum ten is crescent-shaped and armed with setae. The rectum is
evaginated to form a disk-like sucker. It is used as an aid in locomotion.
263] THE LARVAE OF THE COCCINELLIDAE—GAGE 31
COCCINELLTNAE
The body is elongate to oval, dorsal surface convex, ventral surface
concave or flat. The body-wall is provided with senti, strumae, verrucae,
chalazae, or evenly distributed setae. The head is provided with long
distinct setae. The epicranial stem and epicranial arms may be present
or wanting, when present, the epicranial stem never extends more than
one-third of the distance from the occipital foramen to a line drawn through
the antacoriae. The clypeal suture is never entire, but is usually indicated
by a distinct furrow which extends mesad on each side from the precoila.
The antennae are never more than three times as long as wide, consist
of three segments, and are inserted about midway between the ocelli and
the precoilae. The mandibles usually have one, two, or three teeth at the
distal end, and the proximo-mesal margin is provided with a triangular or
mound-shaped mola.
This subfamily includes all of those tribes the members of which are
carnivorous. They may also be phytophagous to the extent that they
may eat fungi and probably small amounts of pollen.
Tribes of Coccinellinae
1(6) Epicranial arms always present; epicranial stem always present
in all larval stages unless obliterated by the separation of the
epicranial arms 2
2(3) The dorsum of the body armed with senti six to eight times as long
as wide; the epicranial stem extending one-third of the distance
from the occipital foramen to a line drawn through the antacor-
iae; epicranial arms gradually divergent Chilocorini
3(2) The dorsum of the body never armed with senti six to eight times
as long as wide 4
4(5) Dorsum of the body armed with short senti, parascoli, or strumae;
epicranial stem, when present, never extending one-third the
distance from the occipital foramen to a line drawn through the
antacoriae; epicranial arms gradually or abruptly divergent
Coccinellini
5(4) Dorsum of the body provided with small verrucae or setae; epi-
cranial stem always obliterated by the separation of the epicra-
nial arms, epicranial arms diverge from the occipital foramen
Microiveiseini
6(1) Epicranial stem and epicranial arms never present in adult larvae 7
7(8) The dorsal and lateral aspects of the body with strumae or
verrucae Scymnini
8(7) The dorsal and lateral aspects of the body with fine slender setae
Hyperaspini
32 ILLINOIS BIOLOGICAL MONOGRAPHS [264
Chilocorini
The body is subovate, widest at the metathorax. The first three
abdominal segments are slightly narrower than the metathorax, segments
four to eight are successively narrower, segment nine is about as wide as
long, its caudal margin is broadly rounded and its lateral margin is pro-
vided with a distinct struma; and segment ten is small and directed ven-
trad. The dorsal and lateral aspects of the body are provided with long
senti the length of which varies on the different portions. The epicranial
stem and epicranial arms are present. The epicranial stem extends about
one-third of the distance from the occipital foramen to a line drawn through
the antacoriae, divides and forms the two epicranial arms which diverge
gradually for a short distance, then widely, and finally each extends
laterad to a point dorsad of an antacoria and ventrad of an ocellus where
it becomes obsolete. The mandibles are triangular in outline, heavily
chitinized, the apex is bifurcated, and the proximo-mesal margin is pro-
vided with a distinct mola.
This tribe is represented in the material studied by the single genus
Chilocorus Leach.
Chilocorus bivulnerus Muls. — The head is chitinized, dark, and pro-
vided with numerous setae. The epicranial stem and epicranial arms are
distinct; the clypeal suture is indicated on each lateral margin of the head.
The mouth is directed cephalo-ventrad. The prothorax is provided with
a large dark colored dorsal shield bearing ten senti, six of which are ar-
ranged in a transverse row near the cephalic margin. The lateral aspect
of the prothorax bears a small group of setae cephalo-dorsad of the pro-
coxacoria. The cephalic portion of the prosternum also bears a small
group of setae on each side of the meson. The procoxacoriae are distant.
The mesotergum is distinctly longer and narrower than the metatergum.
Each is provided with four large senti arranged in a transverse row; the
dorsal senti are distinctly smaller than the lateral ones. The pinacula are
small and bear distinct setae. The mesopleural area is longer than the
metapleural; both areas are obliquely crossed by a distinct furrow. The
cephalic portions of the mesopleural and metapleural areas are subequal
in size, triangular, and each provided with a large sentus. The meso-
thoracic spiracle is located in the mesocoria near the cephalic margin of the
mesothoracic sentus; the metathoracic spiracle is rudimentary or wanting.
The caudal portions of the mesopleural and metapleural areas are unequal
in size, that of the metapleural area is much larger than that of the meso-
pleural. The cephalic metapleural sentus is about one-half as long as the
caudal one. The mesosternum and metasternum are similar in size and
shape. The mesocoria is distinct while the metacoria is obscure. The
cephalic portions of the mesosternum and metasternum are each provided
with a chalaza on each side of the ventro-meson; these chalazae are sur-
265] THE LARVAE OF THE COCCINELLIDAE—GAGE 33
rounded by groups of setae. The legs are about as long as the body is
wide and the tibia is very densely setaceous.
The first abdominal segment is shorter than the metathorax and its
tergum bears four senti arranged in a transverse row; the dorsal pinacula
are yellow and the dorso-lateral ones are white; the lateral aspect is rect-
angular with a large lateral sentus the pinaculum of which covers the
greater part of the lateral surface and the paralaterals are wanting. The
sternum is about as long as the tergum, the ventral setae are adjacent on
the ventro-meson while the ventro-laterals are wanting. Abdominal seg-
ments two to five are similar, narrower than the first; their terga are sub-
equal in length, the dorsal and dorso-lateral senti of each segment are
arranged in a transverse row, and each sentus is provided with a distinct
pinaculum. The lateral aspect is nearly square; the lateral senti are
larger than the dorso-lateral, their pinacula are small. Ventrad of the
second and third lateral senti, there is a small paralateral seta and ventrad
of the fourth and fifth lateral senti, there is a paraleteral group of setae sur-
rounding a distinct paralateral chalaza on each segment. The sterna are
as long as their respective terga, slightly chitinized, their coriae are dis-
tinct, and each bears four chalazae arranged in a transverse row, each
chalaza surrounded by six or eight small setae. Segments six, seven, and
eight are each narrower than the preceding ones. Their dorsal pinacula
are fused and the dorso-lateral senti are shorter than the dorso-lateral senti
of segments two to five. Their lateral aspects are smaller, but are similar
in shape to those of the preceding segments, the lateral aspects of the
sixth and seventh segments are provided with small lateral senti, while the
lateral sentus of the eighth segment is much reduced and the pinaculum
is absent. Ventrad of each lateral sentus, there is a paralateral chalaza
surrounded by a group of setae. The sterna are as wide as the terga, their
coriae are distinct, and the ventral and ventro-lateral groups of setae sur-
rounding the chalazae are present. The ninth abdominal segment is
about as wide and about as long as the eighth, cylindrical, and inclined
ventrad. Its tergum is shield-shaped and has two distinct dorsal verrucae
near the caudo-mesal margin, the dorso-lateral verrucae are wanting. Its
lateral aspect is small, each with a distinct lateral verruca, ventrad of the
verruca there is a distinct paralateral chalaza surrounded by a group of
setae. Its sternum is short, deeply emarginated on the caudal margin,
the ventral and ventro-lateral groups of chalazae are present, and the
ventral chalazae are surrounded by a few setae while the ventro-laterals
are represented by a single chalaza. The tenth segment is small, not
visible from the dorsal aspect and the rectum is evaginated to form a disk-
like sucker. There is a pair of repugnatorial pores located in the coriae of
segments one to seven about one-half the distance between the dorsal and
dorso-lateral senti.
34 ILUNOIS BIOLOGICAL MONOGRAPHS [266
Coccinellini
The body is fusiform or elongate, widest at the metathorax, usually
highly colored with black, red, yellow, orange, or blue; never with senti
except in Anatis in which the senti are short and thick, but never five
times as long as wide, usually with parascoli or strumae. The abdomen
becomes gradually narrower toward the caudal end. The ninth abdominal
segment is about twice as long as wide, never wider than long as in Chiloc-
orini or Hyperaspini, never with lateral parascoli or strumae as in Chiloc-
orini, dorsum provided with a light colored, slightly chitinized shield
bearing many setae or chalazae. The head is heavily chitinized, the
epicranial stem, if present, never extends one-third the distance from the
occipital foramen to a line drawn through the antacoriae; the epicranial
arms are usually widely divergent. The mouth is directed cephalo-ventrad.
The members of this tribe are more numerous in Illinois than those
of any of the other tribes. They are almost wholly carnivorous, their
only plant food being either fungi or in some cases pollen grains.
Genera of Coccinellini
1(2) Epicranial stem and epicranial arms always present; terga one to
eight with strumae usually bearing three distinct chalazae and
densely covered with fine setae MegUla
2(1) Epicranial arms always present; epicranial stem obliterated by the
separation of the epicranial arms 3
3(4) Terga one to eight with senti, not five times as long as wide .Anatis
4(3) Terga one to eight without senti 5
5(8) Terga one to eight with parascoli or strumae which bear more than
five chalazae and are sparsely setaceous 6
6(7) Tarsal claw with a distinct appendiculated tooth at its proximal
end Coccinella
7(6) Tarsal claw without a distinct appendiculated tooth at its proxi-
mal end Hippodamia
8(5) Terga one to eight with strumae which never bear more than five
prominent chalazae and the surface of the strumae densely seta-
ceous Adalia
Megilla Mulsant
This genus is represented in the material studied by a single species.
Megilla maculata DeGeer. — The body is elongate, widest at the meso-
thorax and metathorax. Its general color is black and mottled with light
yellow or cream-colored areas. The head is heavily chitinized, the dorsal
portion dark and the ventral portion more or less white. The epicranial
stem is present as a short line or suture on the dorsal portion of the head.
It extends about one-fifth of the distance from the occipital foramen to a
2671 THE LARVAE OF THE COCCI NELLI DAE— GAGE 35
line drawn through the antacoriae. The epicranial arms diverge laterad
and ventrad and become obsolete slightly mesad of the antacoria. The
mouth is directed ventrad. The cephalic margin of the prothoracic shield
is provided with two small cream-colored spots, and the caudal margin
with a yellow boundary. The mesothorax and metathorax have a shield-
shaped cream-colored area on the dorso-meson; the cephalic portion of the
mesothoracic and metathoracic lateral aspects are also cream-colored.
There is a light colored area on the dorso-lateral portions of abdominal
segments one to eight. On the lateral portion of the first segment, and
on the tergum and lateral aspects of the fourth and fifth segments, the
cuticle is yellow. The ventral aspect of the entire larvae is tan colored.
The prothoracic shield is provided with many chalazae and numerous
setae, the terga of the mesothorax and metathorax are provided with a
large struma-like shield which bears six to eight chalazae. The lateral
aspect of the prothorax is glabrous. The mesothoracic spiracles are
situated in the mesocoria near the cephalic margin of the lateral aspect
and each spiracle is surrounded by three or four small, black setae. The
cephalic portion of the lateral aspect of the metathorax has three or four
setae, but the metathoracic spiracles are wanting. The caudal portions
of the mesothorax and metathorax are about twice as large as the cephalic
and provided with a struma-like plate which bears six to eight chalazae.
The sternum of all of the thoracic segments is provided with a small pair
of verrucae adjacent the meson near the cephalic margin of the segment.
Each tergum of segments one to eight is provided with a transverse row of
four strumae, two dorsal and two dorso-laterals. Each struma usually
bears three distinct chalazae. The lateral aspect of segments one to eight
is each provided with a large lateral struma with four to eight chalazae,
of segments two to eight with a small mound-like paralateral struma usually
with one large chalaza and many seta on each segment. Each sternum of
segments one to eight has a transverse row of four chalazae, each chalaza
surrounded by a few setae. The ventro-lateral group is wanting on the
first sternum. The tergum of the ninth segment is longer than wide,
shield-shaped, its cephalic margin slightly emarginated, darkly colored
and densely setaceous. The lateral aspect is much reduced and bears a
single large lateral chalaza, the paralaterals are wanting. The sternum
is shorter than the tergum, the caudal margin deeply emarginate with the
ventral and ventro-lateral chalazae distinct. The tenth segment is not
visible from the dorsal aspect, slightly setaceous, and appears to be cylin-
drical when viewed from the ventral aspect. The legs are long and slender,
about one and one-half times as long as their thoracic segment is wide.
Anatis Mulsant
This genus is represented in my material by a single species.
Anatis 15 -punctata Oliv. — The body is elongate, widest at the meta-
36 ILLINOIS BIOLOGICAL MONOGRAPHS [268
thorax. The dorsal portion of the head is dark colored while the ventral
part of the face is white or yellow. The epicranial stem is absent, the
epicranial arms extend cephalad and ventrad from the occipital foramen
for a short distance and diverge widely laterad and ventrad; they become
obsolete dorso-mesad of the antacoriae. The mouth is directed cephalo-
ventrad. The mesothorax, metathorax, and the first eight abdominal
segments bear short stout senti from one to five times as long as wide.
The prothorax has a distinct parascoli on the caudo-lateral margin of the
dorsal shield. There are four or five chalazae cephalad of this parascoli.
The caudo-mesal portion of the dorsal shield is provided with a distinct
yellow or white shield-shaped area, the cephalic portion of which bears two
chalazae adjacent to the meson. The mesal and lateral tergal senti of the
mesothorax and metathorax arise from the dorsal shields of their respec-
tive segments. The caudal portions of the lateral aspects of the meso-
thorax and metathorax are each provided with a small sentus. The meso-
thoracic spiracles are located in the mesocoria and the metathoracic spiracles
are rudimentary or wanting. The thoracic sterna are all similar, the
sternum of each segment has a pair of small verrucae adjacent to each other
and the ventro-meson, the coxacoriae are distant. The first to eighth
segments of the abdomen are subequal in length. Their terga are pro-
vided with four senti arranged in a transverse row; the dorsal senti of the
sixth, seventh, and eighth segments have their pinacula fused on each
segment, and the dorso-lateral senti of the seventh and eighth segments are
short and inconspicuous. The metathorax and the first abdominal seg-
ment are each provided with a small cream-colored area caudad of and
about one-half the distance between the dorsal and dorso-lateral senti.
The lateral aspects of the first and second abdominal segments are white,
each with a distinct white lateral sentus and pinaculum; the lateral
aspects of the third to eighth segments are cream-colored with their senti
and pinacula brown; the lateral aspects of the seventh and eighth segments
are without lateral senti but are provided with lateral verrucae while
ventrad of each lateral sentus or verruca, except in the first, ninth, and
tenth segments, there is a distinct chalaza surrounded by a few setae which
represent the paralateral group. The sterna of the first to the eighth
segments are similar in shape, except that they become narrower toward
the caudal extremity. The first sternum has two ventral ohalazae adjacent
to the meson, the ventro-laterals are wanting, while in sterna two to eight
the ventral and ventro-lateral groups are present on each segment. In the
eighth and ninth segments the ventral and ventro-lateral groups on each
side have become fused so that there appears to be a single pair on each
side of the meson. The tergum of the ninth segment is longer than wide,
shield-shaped, and with many setae on its surface; its lateral aspect is much
reduced and bears small and indistinct lateral verruca, the paralaterals
269] THE LARVAE OF THE COCCINELLJDAE—GAGE 37
are wanting; and its sternum is provided with ventral and ventro-lateral
chalazae. The tenth tergum is not visible from the dorsal aspect and the
greater part of its ventral aspect is hidden by the evaginated rectal disk.
The legs are long and slender, about one and one-half times as long as the
metathorax is wide.
CocciNELLA Linne
The body is fusiform, elongate and widest at the metathorax. The
head is usually light-brown colored and provided with many setae. The
epicranial arms diverge immediately from the occipital foramen and be-
come obsolete near the antacoriae. The mouth is directed ventrad and
slightly cephalad. The thoracic segments are provided with a dorsal
shield. The dorsal and dorso-lateral aspects of abdominal segments one
to eight are provided with parascoli or verrucae, the ventro-lateral aspect
with verrucae, and the ventral aspect with verrucae or chalazae surrounded
by a few setae. The dorsum of the ninth abdominal segment is shield-
shaped and densely setaceous, the ventral aspect is provided with verrucae
or chalazae, and is about one-half as long as the dorsal aspect. The legs
are well developed, stout, and extend beyond the lateral margins of the
body. The coxacoriae are distant. The basal portion of the tarsal daw
is provided with a distinct appendiculate tooth.
Species of Coccinella
Terga one to eight with distinct parascoli; prothoracic shield with two
distinct, longitudinal, dark, heavily chitinized areas on each side of
the meson C. 9-notata
Terga one to eight with strumae; prothoracic shield with a single, dark,
heavily chitinized area on each side of the meson C. sanguinea
Coccinella 9-notata Herbst. — The body is fusiform, usually stout, and
widest at the metathorax. The dorsal and dorso-lateral surfaces are pro-
vided with parascoli, and the ventro-lateral surface with small verrucae.
The general color is light grayish-brown to dark tan or brown marked with
white or lemon-yellow areas. The dorsal portion of the head is light-
brown and the face is cream-colored. The epicranial arms diverge from
the occipital foramen and the mouth is directed slightly ventrad and
cephalad. The prothorax is wider than long and oval in outline; the dorsal
shield covers the greater part of its dorsal surface and bears four distinct,
dark-colored, longitudinal areas; and the cephalic, lateral, and part of the
caudal margins are provided with chalazae. The mesothorax and meta-
thorax are subequal in size, each is about twice as wide as long and each
dorsal surface is provided with a small oval dorsal shield on each side of the
meson. There is a small dorso-lateral parascolus on the caudo-lateral
portion of both the mesothorax and metathorax. Each thoracic sterna
is provided with a pair of verrucae which are adjacent to the ventro-meson.
38 ILUNOIS BIOLOGICAL MONOGRAPHS [270
The dorsal and dorso-lateral portions of abdominal segments one to eight
are provided with parascoli. The dorsal parascoli are brown, while the
dorso-lateral and lateral ones of the first and fourth segment are white or
cream-colored, and also the area near the pinacula of these parascoli is
white. Segments two to nine are provided with paralateral verrucae,
while segment one has a small paralateral chalaza. The sterna of segments
two to nine are provided with four verrucae arranged in a transverse row
on each sternum; while the sternum of the first segment has only two ver-
rucae, the ventro-lateral ones are absent. The ventral and ventro-lateral
verrucae on either side of the meson of the ninth segment are almost fused
so that there seems to be only two on this segment. The dorsal surface
of the ninth segment is about twice as long as the ventral, shield-shaped,
the caudal margin is rounded, and densely setaceous. The legs are well
developed, long, and stout. The proximal portion of the ,tarsal claw is
provided with a distinct appendiculate tooth.
Coccinella sanguinea Linn. — The body is elongate and widest at the
metathorax. The dorsal and lateral surfaces are provided with strumae,
and the ventral surface with verrucae or chalazae. The general color of
the body is light to dark brown, mottled or spotted with yellow or cream-
colored areas. The head is light brown and the face is yellow. The mouth
is directed ventrad and slightly cephalad. The prothorax is about three-
fourths as long as wide, and the caudal margin is slightly emarginated.
The dorsal shield is light brown in color, entire, with a light yellow line on
the meson, and the cephalic and lateral margins are provided with chalazae.
The mesothorax and metathorax are subequal in size, about twice as wide
as long, each with a small oval shield on either side of the meson. There
is a light yellow area on the meson between the shields of each segment.
The mesothoracic spiracles are located in the mesocoria cephalo-dorsad
of the mesocoxacoilae, the metathoracic spiracles are rudimentary. The
caudal portion of the lateral aspect of the mesothorax and metathorax
bears a small verruca. The sterna of all of the thoracic segments are
provided with a pair of small verrucae adjacent to each other and the
meson. Abdominal segments one to eight are provided with dorsal and
dorso-lateral strumae, and each segment bears more than five chalazae
and a few scattered setae. The dorsal strumae of the fourth abdominal
segment, the dorso-lateral strumae of the first and fourth segments, and
the lateral strumae of the first, fourth, and fifth segments are white or light
yeUow. There is a small paralateral struma, which in many cases appears
to be verruca-like, located ventrad of each lateral struma, except in the
first abdominal segment where it is wanting. The sterna of segments one
to nine are each provided with ventral and ventro-lateral groups of chala-
zae, except in the first segment, where the ventro-lateral groups are
wanting. The dorsum of the ninth segment is shield-shaped with the
271] THE LARVAE OF THE COCCINELLIDAE— GAGE 39
caudal margin rounded, is twice as long as its sternum, is dark colored, and
bears many short setae. The legs are long, slender, well developed, and
the tarsal claw is provided with a distinct appendiculate tooth at its
proximal end.
HippoDAMiA Mulsant
The body is fusiform, elongate, and usually widest at the metathorax.
The head is brown to dark colored and distinctly setaceous. The epi-
cranial arms diverge immediately from the occipital foramen and become
obsolete near the antacoriae. The mouth is directed ventrad and slightly
cephalad. The dorsum of each thoracic segment is provided with dis-
tinctly chitinized shield-shaped areas which constitute the dorsal shields.
The dorsal and dorso-lateral aspects of the body are provided with para-
scoli. The tergum of the ninth abdominal segment is shield-shaped,
setaceous, and with the caudal margin broadly rounded. The ventro-
lateral aspect of the body is provided with small strumae; while the sterna
are all provided with verrucae. The legs are well developed, extend beyond
the sides of the body, stout, and with the tarsal claw not provided with a
distinct appendiculate tooth at its proximal end. The coxacbriae are
distant.
Species of Hippodamia
Dorsal shield with two brown or black chitinized plates on each side
of the meson H. convergens
Dorsal shield with a single brown or dark chitinized plate on each side
of the meson H. 13-punctata
Hippodamia convergens Guer. — The body is elongate, widest at the
metathorax, and the dorsal and lateral surfaces are provided with para-
scoli. The general color of the body is dark-brown to black, marked with
yellow, orange, red, or white. The head is dark-brown to black, the face
is lighter than the dorsal part. The epicranial arms diverge immediately
from the occipital foramen and become obsolete dorso-mesad of the anta-
coriae. The mouth is directed ventrad-and slightly cephalad. The pro-
thorax, when viewed from above, is oval in outline and wider than long.
The dorsal shield is provided with four longitudinal dark colored areas
separated by white or orange colored bands. The cephalic and lateral
margins of the dorsal shields are provided with chalazae. The dark plate-
like spots also bear chalazae and resemble strumae. The mesothorax and
metathorax are subequal in length. The dorsal surface of each is provided
with parascoli, the pinacula of which have grown together on each side of
the mespn to form a basal shield. The cephalic portion of each lateral
aspect of the mesothorax bears a mesothoracic spiracle, the metathoracic
spiracle of each side is rudimentary. The caudo-lateral aspect of the meso-
thorax and metathorax are each provided with a distinct parascolus, the
40 ILLINOIS BIOLOGICAL MONOGRAPHS [272
mesothoracic one is brown and the metathoracic one is white or cream-
colored. Each thoracic sternum is provided with a pair of verrucae ad-
jacent to the meson. The terga of abdominal segments one to eight are
provided with black or brown parascoli, except the fourth, in which the
parascoli are orange colored. The dorso-lateral parascoli of segments one
and four are also yellow or orange colored. The lateral aspect is provided
with a row of lateral parascoli, those on the first and fourth segments are
white and the remainder are brown. The lateral aspect is also provided
with a row of brown paralateral strumae ventrad of the lateral parascoli.
Sterna two to nine are provided with a transverse row of four verrucae
on each segment, sternum one bears only two ventral verrucae which are
adjacent on the ventro-meson, the ventro-lateral verrucae are absent. The
dorsum of the ninth abdominal segment is shield-shaped, twice as long as
its sternum, and provided with many setae or chalazae. The legs are well
developed and extend beyond the sides of the body. The tarsal claw is
without an appendiculate tooth.
Hippodamia 13-punctata Linn. — The body is slender, elongate, widest
at the metathorax. The dorsal and lateral surfaces are provided with
parascoli. The general color is brownish-tan to dark grayish-brown and
the head is darker than the remainder of the body. The head is heavily
chitinized, the epicranial arms do not meet but extend separately to the
occipital foramen. The mouth is directed cephalo-ventrad. The pro-
thorax is wider than long, and when viewed from above, is oval in outline.
Its entire surface is covered with a solid brown dorsal shield which has a
fine white line running along the dorso-meson. The cephalic, lateral, and
caudal margins are provided with distinct chalazae. The mesothorax is
slightly longer than the metathorax, about twice as wide as long, with a
distinctly chitinized, oval, shield-shaped struma on either side of the
meson. This struma bears ten to fifteen chalazae. The metathorax is
more than twice as wide as long and provided with a chitinized shield-like
struma on each side of the meson. This struma bears about as many
chalazae as the mesothoracic strumae. The cephalic portion of the lateral
aspect of the mesothorax and metathorax bears the thoracic spiracles.
The mesothoracic spiracles are prominent and are located in the mesocoria
cephalo-dorsad of the coxacoriae, the metathoracic spiracles are rudimen-
tary. The caudal portion of the lateral aspect of the mesothorax is pro-
vided with a brown struma, while the metathorax is provided with a white
struma. Each thoracic sterna is provided with two distinct verrucae
adjacent to the ventro-meson. The legs are well developed and extend
beyond the sides of the body. The tarsal claw is without a distinct appen-
diculate tooth. The terga of abdominal segments one to eight are pro-
vided with distinct dorsal parascoli located near the dorso-meson, those
on the fourth abdominal segment are white. The lateral margins of terga
2731 THE LARVAE OF THE COCCJNELLIDAE— GAGE 41
one to eight bear the dorso-lateral parascoli, the first and fourth are white
and the remainder are brown to dark tan-colored. The ninth abdominal
segment is longer than wide; its caudal margin is acutely rounded; its
dorsal surface is brown colored, chitinized, and bears many chalazae and
small setae. The lateral aspect of segments one to eight is provided with
distinct lateral strumae, those on segments one and four are white. There
is a small paralateral verruca ventrad of each lateral strumae. The lateral
aspect of the ninth segment bears chalazae. The sterna of segments two
to nine are provided with four chalazae arranged in a transverse row on
each segment, the sternum of the first segment has only two chalazae
present, the ventrolaterals are wanting.
Adalia Mulsant
This genus is represented in the material studied by a single species.
Adalia bipunctata Linn. — The body is elongate, oval in outline, the
third and fourth abdominal segments are the widest. The general color
is dark brown to bluish-gray, mottled with light yellow or cream-colored
spots. The dorsal part of the head is dark brown to black and heavily
chitinized, while the ventral portion of the front and clypeus is white or
cream-colored. The epicranial stem is absent and the epicranial arms
curve broadly laterad and mesad to the pretentorinae, giving the front a
more or less circular appearance, then extend laterad from the pretentorinae
toward the antacoriae near which they become obsolete. The mouth is
directed cephalo-ventrad. The prothorax is crossed longitudinally by a
median and two lateral yellow stripes. The dorsal shield is not united to
the meson of the mesothorax and metathorax and the two portions are
separated by a cream-colored area. The lateral aspect of the prothorax is
glabrous. The cephalic portion of the mesothorax and metathorax is also
glabrous; while the caudal portion is large and bears a small but distinct
struma on each segment. The mesothoracic and metathoracic spiracles
are located in the coriae between the segments. The thoracic sterna are
distinct, each sternum bears a pair of small verrucae near its cephalic
margin. The coxacoriae are distant. Abdominal segments one to eight
are each provided with a transverse row of four strumae on the tergum.
The mesal portions of the dorsal strumae of the fourth segment are white,
while the lateral portions are brown, the dorso-lateral strumae of the first
segment are surrounded by a distinct white area, and the dorso-lateral
strumae of the second to the eighth segments are surrounded by a much
smaller white area. Each struma is provided with three to five prominent
chalazae. The lateral aspect of segments one to eight is yellow and the
strumae are brown except on the fourth segment where they are light
yellow. There is a small but distinct chalaza on each segment ventrad of
the lateral strumae of segments one to eight which is surrounded by a few
42 ILLINOIS BIOLOGICAL MONOGRAPHS [274
setae representing the paralateral group. The ninth tergum is shield-
shaped, longer than wide, the caudal margin concave and provided with
numerous setae, while the cephalic margin bears only a few chalazae.
The tenth tergum is visible from the dorsal aspect as a small, brown
colored, chitinized area caudad of the ninth tergum and is provided with
a single chalaza on each side of the meson. Each of these chalaza is sur-
rounded by a group of small setae. The ninth sternum is about one-half
as long as its tergum, deeply emarginate on its caudal margin, and bears
two chalazae on each side of the meson, each of which is surrounded by a
few setae. The tenth sternum is longer than its tergum and bears a dark
colored spot near the lateral margin. The legs are slender, well developed,
and longer than the body is wide.
Microweiseini
The body is small, elongate, fusiform, depressed, widest at tlie meta-
thorax, provided with setae, and light-brown to yellowish-tan colored.
The head is dark-brown to black in color and chitinized. The mouth is
directed cephalo-ventrad and the mandibles are unidentate. The epi-
cranial arms diverge slowly from the occipital foramen and become obsolete
near the antacoriae. The spear-shaped front and post-clypeus are divided
on the meson by a distinct, dark colored, chitinized bar which extends from
the occipital foramen to the clypeo-labral suture. The prothorax is oval
and slightly wider than long; while the mesothorax and metathorax are
distinctly wider than long. The terga of the thoracic segments are longi-
tudinally crossed by a small dark-colored area on each side of the meson,
more distinct on the prothorax than on the mesothorax or metathorax.
The ninth abdominal segment is conical, narrower than the eighth, and
about twice as long as wide. The legs are well developed and extend
beyond the sides of the body; the tibiae are provided with two paddle-
shaped tenent hairs at the distal end; the tarsal claw bears a distinct appen-
diculate tooth; and the coxacoriae are distant.
MiCROWEiSEA Cockerell
This genus is represented in the material studied by a single species.
Microweisea misella Le Conte. — The body is fusiform, elongate and
light-brown to yellowish-tan in color. The head is elongate, dark colored,
and chitinized. The epicranial arms are present, but the epicranial stem
is wanting. There is a dark-brown heavily chitinized bar extending along
the dorso-meson from the occipital foramen to near the clypeo-labral
suture. The mouth is directed cephalad. The prothorax is wider than
long and somewhat oval in outline. There are two brown chitinized spots
near the middle of the dorsum adjacent to the meson. The dorsum is
provided with a very few short setae. The mesothorax and metathorax
2751 THE LARVAE OF THE COCCI NELLl DAE— GAGE 43
are about twice as wide as long; and the lateral margin of each is provided
with a fringe of fine setae. The sterna of the thoracic segments are equal
in size to that of their respective terga. The prosternum appears to be
glabrous; while there is a pair of small verrucae adjacent to the ventro-
meson of the metathorax and mesothorax. These verrucae are provided
with setae which are about as long as their segments. The coxacoriae are
distant and placed near the lateral margins of the sterna. The legs are
small, well developed, and extend beyond the sides of the body. Each
tibia is provided with two paddle-like tenent hairs placed near the distal
end of the segment, the tarsal claw bears a distinct appendiculate tooth.
Abdominal segments one to eight are similar, except that they become
successively smaller. Each tergum is provided with a few small setae
which are never as long as the segments. The dorsum of the eighth seg-
ment is dark, chitinized, and its caudal margin is not emarginate. The
sterna and the lateral aspects of segments one to eight are similar in general
size and structure. They are membraneous and provided with a few small
setae. The coriae between the segments are distinct. The sternum of the
eighth segment is deeply emarginate on its caudal margin. The ninth
segment is longer than wide, its caudal margin narrower than the cephalic.
The tergum is shield-shaped, dark-colored, chitinized, and the caudal
margin is acutely rounded and bears many setae about one-half as long as
the segment is wide. The shape of the ninth sternum is similar to that
of its tergum, it is very slightly chitinized and supplied with only a few
setae. The tenth segment is cylindrical, directed caudo-ventrad; the rec-
tum is evaginated to form a sucking disk.
Scymnini
The body is small, elongate, fusiform, widest caudad of the metathorax,
provided with verrucae or chalazae and setae, and light-yellow to light-
brown in color. The head is slightly chitinized, light colored, densely
setaceous, and directed cephalad. The epicranial suture is wanting.
The prothorax is oval, slightly chitinized, and provided with verrucae.
The mesothorax and metathorax are about twice as wide as long and are
provided with verrucae on the dorsal surfaces and with chalazae and setae
on the lateral and ventral surfaces. The abdominal segments are provided
with verrucae on the dorsal and lateral surfaces and with chalazae and setae
on the ventral. The ninth abdominal segment is cylindrical, about twice
as long as wide and the sternum is shorter than the tergum. The legs
are well developed and extend beyond the sides of the body, the tibiae
bear more than two tenent hairs, and the coxacoriae are distant.
ScYMNus Kugelann
This genus is represented in my material by a single species which could
not be determined.
44 ILLINOIS BIOLOGICAL MONOGRAPHS [276
Scymnus sp.? — The body is fusiform, elongate, flattened, Kght-yellow
or cream-colored to light-brown. The head is circular in outline, very
slightly chitinized, and provided with numerous black setae which are
about one-half as long as the body is .wide. The epicranial suture is
entirely wanting and the mouth is directed cephalad. The prothorax is
slightly wider than long, and the dorsum bears the light-brown slightly
chitinized dorsal shield. The cephalic margin is provided with six large
black setae which are about one-third as long as the segment, the caudal
margin has six setae of about the same size as the cephalic setae, but not
so darkly colored; midway between the cephalic and caudal rows there is
another of four long black setae. The mesothorax is distinctly narrower
and shorter than the metathorax which is broadly emarginate on its
caudal margin. Both the mesothorax and metathorax have an oval-
shaped verruca on either side of the dorso-meson. This verruca is pro-
vided with setae which are about as long as the verruca is wide. The
thoracic sterna are distinct. Those of the mesothorax and metathorax are
provided with a few setae; while the prothoracic sternum is glabrous.
The coxacoriae are distant and the legs are well developed, usually white
or light-yellow in color, and extend beyond the margins of the body.
Each tibia is provided near the distal end with six to eight tenent hairs. The
tergum of the first abdominal segment is distinctly longer than the second.
Each tergum of abdominal segments one to eight is provided with a trans-
verse row of four distinct verrucae. The dorsal verrucae are adjacent to the
dorso-meson and the dorso-lateral are near the lateral margin of the tergum
in each segment. The lateral aspect of segments one to seven is lobed and
each lobe bears a distinct verruca with setae as long as the segment bearing
them. The lateral aspect of the eighth segment bears two or three chala-
zae about as long as the setae on the verrucae. There are on segments two
to eight one or two distinct paralateral chalazae immediately ventrad of
the lateral verrucae and chalazae, while the first segment bears a small
paralateral seta. The sterna of segments one to eight are similar to their
terga in size and shape. The cuticle is membraneous and bears on each
segment a transverse row of small setae. The coriae are distinct. The
ninth abdominal segment is longer than wide, slightly chitinized, and
marked with a dark spot near the middle and on either side of the dorso-
meson. The tergum is shield-shaped and provided with many dark colored
setae. The lateral aspect of the segment appears to be glabrous. The
sterna is shorter than the tergum, the cuticle is membraneous and armed
with a transverse row of four small setae. The tenth abdominal segment
is cylindrical, glabrous, directed caudo-ventrad and bears the large disk-
like sucker.
277] THE LARVAE OF THE COCCINELLJDAE—GAGE 45
Hyperaspini
The body is ovate and the first to the fourth abdominal segments are
widest. The prothorax is wider than long and the caudal margin is longer
than the cephalic. The ninth abdominal segment is directed ventrad,
wider than long, the tergum very slightly chitinized and provided with a
few small setae. The body is usually white or yellow in color. The larvae
cover themselves with flocculent masses of wax-like excretion. The body
is never provided with scoli, senti, strumae, or verrucae, but is usually
armed with setae or small chalazae. In the early larval stages the epi-
cranial suture is present, while in the adult larval stages the epicranial
suture is wanting. The antennae are short, three-segmented, and slightly
setaceous. The mandibles are provided with a slightly developed mola
and the apex is never bifurcate.
Genera of Hyperaspini
Cephalic margin of the prothorax with setae about as long as the pro-
thorax is wide and lateral tergal abdominal setae are about twice as
long as the segments bearing them Brachyacantha
Cephalic margin of the prothorax with setae which are not as long as the
prothorax is wide and the lateral tergal abdominal setae are shorter
than the segments bearing them Hyperaspis
Brachyacantha Chevrolat
This genus is represented by a single species.
Brachyacantha ursina Fab. — The dorsal portion of the head is slightly
chitinized. The head is light yellow in color; the mouth is directed ventrad
and slightly cephalad. The body is elongate, widest at the second and
third abdominal segments, white to light-cream-colored, usually covered
with a flocculent wax-like excretion. The prothorax is one-half as long
as wide. The dorsal shield is wanting, but the cephalic margin of the
prothorax bears eight setae which are longer than the segment is wide, the
lateral margin bears two setae which are also longer than the segment.
The lateral aspect is poorly defined and glabrous; the sternum is small
and without setae. The mesothorax and metathorax are each about three
times as wide as long, are provided with a seta near the lateral margin
which is longer than the segment, and the lateral aspects are not well
defined. The mesothoracic spiracles are located in the mesocoria near the
cephalic margin of the segment, the metathoracic spiracle is wanting or
rudimentary. The legs are short and not well developed, rarely extending
beyond the sides of the body. Abdominal segments one to eight are
similar. The terga are strongly convex, the dorsal and dorso-lateral setae
are longer than the segments bearing them, and the spiracles are located
near the cephalo-ventral margin of each tergum. The lateral aspect of
each segment is provided with two large setae which are not longer than
46 ILUNOIS BIOLOGICAL MONOGRAPHS [278
the segments bearing them, the lateral setae are larger than the paralateral
ones. The sterna are flat; each, except the first, bears a transverse
row of four large setae. In the first segment the ventro-lateral setae are
wanting. The caudal margins of the seventh and eighth sterna are deeply
emarginate. The tergum of the ninth segment is semicircular, about as
wide as long, and the dorsal surface is densely setaceous. The ninth
sternum is small and bears four small setae. The tenth segment is small,
cylindrical, glabrous, and directed ventrad.
Hyperaspis Redtenbacher
The members of this genus are generally white to cream-colored, and
the body is usually covered with a flocculent wax-like mass. When seen
from the dorsal aspect, the body is oval to elongate in outline, the dorsum
is strongly convex and the sternum is more or less flattened. The pro-
thorax is wider than long and bears setae which are not as long as the pro-
thorax is wide. The ninth abdominal segment is semicircular, wider than
long, and usually retracted into the eighth segment. The legs are small,
short, and well developed. The mandibles are unidentate and serve as
piercing organs. The members of this genus are carnivouous living for
the most part upon aphids and soft-bodied coccids.
Species of Hyperaspis
Body elongate-ovoid and densely covered with dark hair-like setae
H. binotata
Body oval, not elongate, apparently glabrous, but with a few small
inconspicuous setae H. signata
Hyperaspis binotata Say. — The dorsal aspect of the body is brownish-
yellow to yellowish-gray; the front, vertex, and clypeus are spotted with
light or dark brown areas and provided with numerous setae. The pro-
thorax is about twice as wide as long and the cephalic and lateral margins
bear setae as long as the segment. The mesothorax and metathorax are
subequal in length, but the latter is the wider. The lateral margin of the
mesothorax is provided with a group of setae about as long as the segment,
while the lateral margin of the metathorax has a distinct chalaza on each
side which is surrounded by a group of long setae. The lateral margins
of abdominal segments one to eight are each provided with a chalaza sur-
rounded by a distinct group of setae. The dorsal surface of the thoracic
and abdominal segments are densely covered with short black setae. The
sterna and lateral aspects of the entire larva are provided with numerous
fine and inconspicuous setae. The tergum of the ninth abdominal seg-
ment is about twice as wide as long, not strongly chitinized, and the caudal
and lateral margins bear setae which are longer than the segment. The
sternum is about one-half as long as the tergum and deeply emarginate on
279] THE LARVAE OF THE COCCINELLIDAE—GAGE 47
the caudal margin. The tenth segment is small, cylindrical, and slightly
chitinized, with a few fine setae on its surfaces. The segment is usually
directed caudo-ventrad. The rectum has been evaginated to form a suck-
ing disk. The legs are short and well developed, but do not extend beyond
the sides of the abdomen in the adult larvae. The coxacoriae are distant.
Hyperaspis signata Oliv. — The general form of the body is oval, the
dorsal surface is globose. The body is usually light-yellow to yellowish-
green in color. The dorsal aspect of the head is brown, tan, or yellowish
gray. The cephalic portions are spotted with small brown or black areas.
The head is provided with many setae which are usually the longest setae
found on the entire body. The tergum of the prothorax is rectangular
and about twice as wide as long. The dorsal shield is wanting and the
lateral and caudal surfaces are provided with setae which are about as
long as the segment. The mesothorax and metathorax are subequal in
length, but the metathorax is wider than the mesothorax. The lateral
margins of both of these segments are provided with a few short setae
which are not as long as the segment. The dorsal aspect of abdominal
segments one to eight are similar and strongly convex, with a very distinct
coria between the segments, and never densely setaceous but provided
with a few setae which are never as long as the segment bearing them.
The lateral margin of the dorsal aspect of the abdomen is provided with
a series of lobes between the annulets. These lobes are provided with a
few setae which are not as long as the segment. The sterna and the
lateral aspect of segments one to eight are provided with a few short
setae which are almost invisible. The tergum of the ninth abdominal seg-
ment is more or less shield-shaped, more than twice as wide as long, with
the caudal margin broadly rounded. The lateral and caudal margins
bear setae which are not as long as the segment. The ninth sternum is
about one-half as long as the tergum, its cephalic margin is broadly convex
while the caudal margin is deeply emarginate. The sternum and the
lateral aspects are provided with a few small setae. The tenth abdominal
segment is retracted into the ninth so that, as a rule, it is not visible. The
tenth segment is small, circular, and membraneous. The rectum is eva-
ginated to form a sucking disk. The legs are small, dark brown, and well
developed, but do not extend beyond the sides of the body.
48 ILLINOIS BIOLOGICAL MONOGRAPHS [280
BIBLIOGRAPHY
Arrow, Gilbert J.
1918. Life History of Scsonnus capitatus. Entom. Monthly Mag., 54: 8-9.
Banks, Chas. S.
1906. Principal Insects Attacking Coconut Palm (Part II). Philippine Jour, Sd.,
211-228; 10 pis.
Bexjtenmueller, Wm.
1891. Bibliographical Catalogue of the Described Transformations of North American
Coleoptera. Jour. N. Y. Micros. Soc., 7:1-52.
Blatchley, W. S.
1910. An illustrated Descriptive Catalogue of the Coleoptera or Beetles (Exclusive of
the Rhynchophora) Known to Occur in Indiana. Indianapolis. 8vo. Pp. 506-533.
BOviNG, Adam.
1917. A Generic Synopsis of the Coccinellid Larvae in the United States National
Musevun, with a Description of the Larvae of Hyperaspis binotata Say. Proc.
U. S. Nat Museum, 51:621-650; 4 pis.
Britton, W. E.
1914. Some Common Lady Beetles of Coimecticut. Bull. Conn. Agr. Exp. Sta,, 181,
Ent. Series 19:1-24.
Casey, Thos. L.
1899. A Revision of the American Cocdnellidae. Jour. N. Y. Entom. Soc., 7:71-169.
Clausen, C. P.
1915. A Comparative Study of a Series of Aphid-feeding Coccinellidae. Jour. Econ.
Entom., 8:487-491.
COMSTOCK, J. H.
1893. Evolution and Taxonomy. Wilder Quarter-Century Book. Pp. 37-113. 3 pis.
CoMSTOCK, J. H., and Kochi, C.
1902. The Skeleton of the Head of Insects. Amer. Nat, 36:13-45.
COQUEREL, C.
1849. Observations entomologiques sur divers Coleopteres recuellis auz Antilles. Ann.
soc. ent. France (2), 7:441-454.
DiMMOCK, G. W.
1906. Algunas Coccinellidae de Cuba. Primer Informe Anual de la Estaci6n Central
Agron6mica de Cuba, 287-392; 3 pis.
French, G. H.
1883. Preparatory Stages of Epilachna borealis, Fab. Can. Entom., 15:189-191.
Ganglbauer, L.
1899. Die KSfer von Mitteleuropa. 8o. VoL 3. Pp. 941-1023.
Hilton, W. A. •
1902. The Body Sense Hairs of Lepidopterous Larvae. Amer. Nat, 36:561-578.
Leng, Charles W.
1903. Notes on Coccinellidae. Jour. N. Y. Entom. Soc, 11:35-45; 193-213; 2 pis.
Letzner, K.
1857. Beitrage zur Verwandlungsgeschichte der Coccinellen. Zeit f. Ent, 11:3-24;
IpL
281] THE LARVAE OF THE COCCJNELLJDAE—GAC^ 49
Lewcoce, G. a.
1893. Note on Cocdnella ocellata, L. Entomolo^t, 26:249.
Fauces, Miriam A.
1914. Some Notes on Life History of Ladybeetles. Ann. Ent. Soc. Amer., 7:213-238;
2 pis.
SlUANTON, F. L.
1916. Hyperaspis binotata, A Predatory Enemy of the Terrapin Scale. Jour. Agri.
Res., 6:197-203; 2 pis.
Weise, Juuus.
1879. Bestimmungs-Tabellen der europfiischen Coleoptem. n. Cocdnellidae.
Zeitschr. fOr Ent., n. F. 7:88-156.
283] THE LARVAE OF THE COCCI NELLI DAE— GAGE 51
PLATE I
52
ILUNOJS BIOLOGICAL MONOGRAPHS
\m
EXPLANATION OF PLATE
CHILOCORUS BIVULNERUS
Fig. 1. Larva, dorsal aspect.
Fig. 2. Larva, lateral aspect.
Fig. 3. Larva, ventral aspect.
abdcof
abdominal coria
tbdsp
abdominal spiracle
ds
clypeo-labral suture
CO
coxacoria
die
dorsal senti group
dise
dorso-lateral senti group
ds
dorsal shield
fc
front and postdypeus
fuf
furdnia
I
labrum
he
lateral senti group
mi
mandible
micot
mesocoria
mspi
mesopleural area
msst
mesostemum
mssp
mesothoradc spirade
mst
mesotergum
mieor
metacoria
mtpl
metapleural area
mta
metastemum
mU
metatergnra
mxpl
maxUlaiy palpus
oc
ocelli
pc
predypeus
flse
paralateral senti group
pma
points of musde attachment
pre
procoza
firco
procoxacoria
prd
protarsaldaw
prjr
profemur
Pfpl
propletiral area
PfSt
prostemum
prta
protarsus
prti
protibia
prth
prothoraz
prtr
protrochanter
TP
repugnatorial pore
sd
sucking disk
St
sternum
i
tergum
Ose
ventro-lateral sent! group
tse
ventral senti group
ILLINOIS BIOLOGICAL MONOGRAPHS
VOLUME VI
GAGE
LARVAE OF THE COCCINELLIDAE PLATE I
2851 THE LARVAE OP THE C0CCINELLIDA3-GAGE 53
PLATE II
54
ILLINOIS BIOLOGICAL MONOGRAPHS
[286
EXPLANATION OF PLATE
CEPHALIC ASPECT^OF THE HEAD
Fig. 4. EpUackna borealis.
Fig. 5. CkUocorus bivulnerus, lateral aspect.
Fig. 6. CkUocorus bivulnerus.
The label els for the clypso-labral suture is wanting.
Fig. 7. Megilla tnacidata.
Fig. 8. Hippodamia 13-punciata.
Fig. 9. Hippodamia corner gens.
a
antenna
els
ctjrpeo-labral suture
cs
clypeal suture
ta
epicranial arms
es
epicranial stem
fc
front and clypeus
ta
galea
g«
gena
I
labrum
UU
labialpalpus
mandible
md
mco mandacoria
mxpl maxillary palpus
oc ocelli
pc predypeus
pd precoba
pf palpifer
pt pretentorina
ss+ca fused stipes and cardo
supt supratentorina
t vertex
ILLINOIS BIOLOGICAL MONOGRAPHS
VOLUME VI
GAGE
LARVAE OF THE COCCINELLIDAE PLATE II
2871 THE LARVAE OF THE COCCINELLIDAE—GAGE S5
PLATE III
56 ILLINOIS BIOLOGICAL MONOGRAPHS (288
EXPLANATION OF PLATE
CEPHALIC ASPECT OF THE HEAD
Fig. 10.
Couindla Q-notaia.
Fig. 11.
Anatis 15-punctata.
Fig. 12.
Adalia bipunctata.
Fig. 13.
Microweisea tnisella.
Fig. 14.
Scymnus sp.
Fig. 15.
Hyperaspis binotata.
a antenna
mco
mandacoria
ch clypeo-Iabral suture
oe
ocelli ^m
cs clypeal suture
PC
preclypeat
ea epicranial arms
pa
precoila
fc front and postclypeut
pt
pretentorins
fe+v front, postclypeus and vertex
9
vertex
/ labrum
md mandible
ILLINOIS BIOLOGICAL MONOGRAPHS
VOLUME VI
GAGE
LARVAE OF THE COCCINELLIDAE PLATE III
289] THE LARVAE OF THE COCCINELLIDAE—GAGE 57
PLATE IV
58
ILUNOIS BIOLOGICAL MONOGRAPHS
1290
EXPLANATION OF PLATE
VENTRAL ASPECT OF THE HEAD
fig. 16. EpUachna borealis.
The Label Pg for the palpiger is wanting.
Pig. 17. Chilocorus bivulnerus, caudal aspect,
fig. 18. Chilocorus bimdnerus.
Fig. 19. MegiUa maculata.
The Label su for the submentum b wanting.
Fig. 20. Eippodamia 13-pundata.
The Label lig for ligula is wanting.
Pig. 21. Eippodamia convergens.
a
chidnoasband
cp
corpotentorium
ta
epicranial arms
a
epicranial saton
I
gula
ga
galea
le
gena
lie
labiacoria
lit
ligula
upi
labial palpus
mxpi maxillary paljxit
of occipital forunen
pf palpiler
pg palpiger
pt pretentorina
ss+ca fused sdpes aod cardo
nt submentum
sup supratentorium
ts tactile setae
ILLINOIS BIOLOGICAL MONOGRAPHS
VOLUME VI
GAGE
LARVAE OF THE COCCINELLIDAE PLATE IV
291] THE LARVAEfiF TEE COCCINELLIDAE—GAGE 59
PLATE V
60 ILLINOIS BIOLOGICAL MONOGRAPHS [292
EXPLANATION OF PLATE
VENTRAL ASPECT OF THE HEAD
Fig. 22. CoccineUa 9-notata.
Fig. 23. Anatis 15-punctata.
Fig. 24. Adalia bipunctata.
Fig. 25. Microweisea misella.
Fig. 26. Scymnus sp.
Fig. 27. Hyperaspis binotata.
dt clutinoas band mzc maxacoria
I gula mx^ maxillary palpua
ta galea pf palfufei
It gena ft palpiger
lie labiacoria M+ca fused stipes and caido
Ht ligula su submentom
Upl labial palpus U tactile letae
ILLINOIS BIOLOGICAL MONOGRAPHS
VOLUME VI
GAGE
LARVAE OF THE COCCINELLIDAE PLATE V
2931 THE LARVAE OP THE COCCINELUDAEr-CAGE 61
PLATE VI
^
tfl ILLINOIS BIOLOGICAL MONOGRAPHS [294
EXPLANATION OF PLATE
MISCELLANEOUS PARTS OF THE BODY
Fig. 28. Scolus, EpUachna borealis.
Fig. 29. Sentus, Chilocorus bivulnerus.
Fig. 30. Parascolus, Hippodamia convergens.
Fig. 31. Struma, MegiUa maculata.
Fig. 32. Verruca, Microweisea misella.
Fig. 33. Chalaza, Hippodamia convergens.
Fig. 34. Seta, Hyperaspis binohUa.
Fig. 35. Antenna, EpUachna borealis.
Fig. 36. Antenna, Chilocorus bivulnerus.
Fig. 37. Antenna, Hippodamia convergens.
Fig. 38. Antenna, Hyperaspis binotata.
Fig. 39. Antenna, Scymnus sp.
Fig. 40. Mandible, Chilocorus bivulnerus, lateral aspect.
Fig. 41. Mandible, Chilocorus bivulnerus, mesal aspect.
Fig. 42. Mandible, Microweisea misella.
Fig. 43. Tarsus, Chilocorus bivulnerus.
Fig. 44. Tarsus, Microweisea misella, lateral aspect
Fig. 45. Tarsus, Microweisea misella, ventral aspect.
Fig. 46. Tip of labial palpus, Chilocorus bivtt^erus.
Fig. 47. Tentorium, Chilocorus bivulnerus.
ant antennaria foa postartif
ante antacoria ^ pretentorium
at appendiculated tooth ps preartis
cp corpotentorium sc scape
ea epicranial arms ' S€ sensoria
€S epicranial stem sup supratentorium
d denies te tarsal daw
Jl flagellum te tenenthair
met metatentoria ti tibia
mo mola ts tactile setae
pd pedicel
ILLINOIS BIOLOGICAL MONOGRAPHS
VOLUME VI
GAGE LARVAE OF THE COCCINELLIDAE PLATE VI
Nrntural History 6ttnF«f
t^ Ukitarf
