Insect Morphology

MORPHOLOGY AND EVOLUTION
OF THE INSECT THORAX

RYUICHI MATSUDA
Entomology Research Institute
Canada Department of Agriculture
Ottawa, Canada
MEMOIRS OF
THE ENTOMOLOGICAL SOCIETY OF CANADA- No. 76
Editor: D. P. Pielou
To my Mother
Publishedby THE ENTOMOLOGICAL SOCIETY OF CANADA, OTTAWA
1970
CONTENTS
PAGE
INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
I. GENERAL DISCUSSION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
1. The External Structures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
The Neck Region . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
The Tergum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
The Sternum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
The Intersegmental Relationship . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
The Origin of the Pleuron . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Homologies of the Pleural Parts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
The Wing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
The Coxa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Positions of the Thoracic Spiracles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
2. The Musculature . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Thoracic Musculature in Lepisma saccharina . . . . . . . . . . . . . . . . . . . . . . . . . 48
Thoracic Musculature in the Pterygota . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
Major Evolutionary Features of Thoracic Musculature . . . . . . . . . . . . . . . . 76
II. SPECIAL DISCUSSION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
The Protura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
The Collembola . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
The Diplura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
The Thysanura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
The Ephemeroptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111
The Plecoptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 122
The Embioptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132
The Phasmida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
The Blattaria . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143
The Isoptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 154
The Mantodea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157
The Grylloblattodea . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 167
The Orthoptera
169
The Dermaptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187
The Coleoptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 199
The Zoraptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
The Psocoptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222
The Mallophaga . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 232
The Anoplura ............................................ ......... 239
The Thysanoptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 241
The Hemiptera (Homoptera) ...................................... : 248
The Hemiptera (Heteroptera) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267
3
4 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA
PAGE
The Neuroptera ..................................................... 283
The Mecoptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 293
The Diptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 304
The Trichoptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 326
The Lepidoptera (Zeug!optera and Dacnonypha) . . . . . . . . . . . . . . . . . . . . . . 334
The Lepidoptera (Monotrysia and Ditrysia) . . . . . . . . . . . . . . . . . . . . . . . . . . 339
The Hymenoptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 354
The Strepsiptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 373
The Siphonaptera . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382
The Odonata . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386
Literature Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 402
Index to Terminologies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 427
Index to Authors . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 429
MORPHOLOGY AND EVOLUTION OF THE INSECT THORAX
RYUICHI MATSUDA
Entomology Research Institute, Canada Department of Agriculture, Ottawa
INTRODUCTION
This is the second of a series of works in which the structural evolution of insects is
studied. This volume deals with the evolution of the thorax throughout the class Insecta.
As in the first volume, dealing with the head ( 1965), the primary objective is to establish
homologies of the external structures and muscles.
Homologizations of structures require, among other things, a clear understanding
of the phylogenetic relationships of groups, and vice versa. Whenever this work deemed
to throw new light, therefore, the phylogenetic relationships among orders of insects
are also discussed, and in doing so terminologies proposed by Hennig ( 1950, 1953,
1966) are followed.
Realizing that a thorough study covering all major problems of the insect thorax is
hopelessly time consuming, I have left aside certain important aspects such as the wing
venation, the legs, and so on. Flight mechanism in each order is discussed pTimarily
from the anatomical viewpoint, so long as the physiological factors do not override the
anatomical considerations in importance. The pattern of innervation is also omitted,
since it has become increasingly clear in recent years that at the present state of our
knowledge and techniques the innetvation pattern cannot be a primary landmark in
determining homologies of muscles and external structures.
The material on which the present work is based has been derived mainly from
published works, although a considerable number of actual dissections were necessary
to fill the gaps in knowledge. The result of this work will show where we stand today in
our knowledge of the insect thorax and what is to be studied next. The last careful
search of literature was made in the summer of 1968.
ACKNOWLEDGMENTS
This study was started in the summer 1965 at the Museum of Zoology, University
of Michigan, Ann Arbor, under a grant from the National Institute of Health. The
work was continued at the Entomology Research Institute, Ottawa, since February 1968,
and was completed in April 1969. In the summer of 1966 this work was carried out at
the Zoological Institute, University of Mainz.
Assistance came from various sources during the course of this study. Besides the
gifts of specimens from my colleagues in Ann Arbor and Ottawa, I owe the material for
dissection to Prof. G. W. Byers (Lawrence), Dr. A. B. Gurney (Washington, D.C.),
Dr. R. Kinzel bach (Mainz), Dr. G. Mickoleit (Ttibingen), Prof. S. Miyamoto
(Fukuoka), Dr. E. S. Ross (San Francisco), Dr. L. M. Roth (Natick), and Dr. P.
Wygodzinsky (New York).
Sections of the manuscript were read and valuable criticisms and suggestions were
received from Dr. S. Asahina (Tokyo), Dr./ R. McAlpine (Ottawa), Dr. J. M. T
Campbell (Ottawa), Dr. G. Mickoleit (Tiibingen), Dr. M. Parsons (Toronto), Dr.
W. R. Richards (Ottawa), Dr. E. G. Munroe (Ottawa), Dr. J. Sharplin (Edmonton)
and Dr. J. R. Vockeroth (Ottawa). Mr. J. A. Downes (Ottawa) reviewed the whole
manuscript and gave criticism and advice from an editorial viewpoint. I should also
mention useful conversations with Dr. J. Barlet (Liege) and with Dr. G. Mickoleit
(Ttibingen).
s
I. THE GENERAL DISCUSSION
In this section emphasis is placed on finding the primitive organization of thoracic
structures, to enable homologizations of structures in the higher groups. Since, how-
ever, this section cannot be independent of the following special discussion (Part II),
which deals with each order of insect, it serves also as a summary of the next section.
General principles of structural evolution throughout the class Insecta are also dis-
cussed here.
1. THE EXTERNAL STRUCTURES
THE NECK REGION
The origin of the neck region: The Origin of the neck region, whether it derives
from the head or from the prothorax or from both, has been a matter of contention in
the past. During the embryonic development of Si/pha, according to Smreczinski
(1932), the ventral and lateral parts of the labial segment participate in the formation of
the head, and the rest (the dorsal portion) connects the head and the thorax. This
observation suggests that in the adult the definitive ventrolateral part of the labial segment
must have been completely integrated into the head. It follows, then, that the ventral
part of the neck region must be prothoracic in origin, and that only dorsally can the
neck region be labial in origin. Holmgren (1909) also said that a part of the neck
membrane probably belongs to the labial segment.
The muscles occurring in the neck are at least partly labial in origin. In Carausius
Wiesmann ( 1926) showed that two of the three labial embryonic coelomic diverticula
(medial and laterodorsal) give rise to the ventral longitudinal muscles (of the neck
region), the dorsal longitudinal muscles, and the rotator of the head, and that only the
ventral coelomic diverticulum gives rise exclusively to labial muscles. In Silpha
(Smreczinski 1932) the labial coelomic sac gives rise to the labial muscles, part of the
flexor muscle of the mandible, the rotators of the head, part of the midgut musculature,
etc. It is evident that at least some neck muscles (rotators of the head) are formed
from the labial segment in Silpha and Carausius. Other muscles occurring in the neck
region are, therefore, perhaps prothoracic in origin.
Nerve distribution is sometimes used in delimiting segments, on the premise that a
nerve innervates the segment to which the ganglion from which the nerve arises belongs.'
Matsuda (1956), Schmitt (1962) and Mark! (1966), among more recent workers, have
pointed out, however, that this criterion cannot always be reliably used, because two
neighboring nerves may fuse in varying degrees or a muscle may receive more than one
nerve; furthermore, individual variation in the distributional pattern of a nerve may
sometimes be considerable. Yet if the general distributional pattern of the nerves is
more or less constant in various groups of insects, and if the above difficulties are
relatively unimportant in interpreting the anatomical area in question, innervation can be
used as an additional criterion in determining a segment.
Thus, in Dissosteira (Fig. 1), as Schmitt (1959) showed, a purely suboesophageal
nerve (second cervical nerve of Schmitt) innervates muscles op-t 1, 2, 3 ( 47-49 of
Snodgrass, 1929a) and cv(d)-t 1 (56). The fusion of the anterior ganglionic connective
and the prothoracic anterior dorsal nerve innervates t-p 2 (58), t-cv 3 and t-s(cv) 1 (52),
t-cv 2 (53), and s I, 2 (55). p-t 1 and 2 are also innervated by a prothoracic nerve.
In Periplaneta (Pipa and Cook 1959) a purely stiboesophageal nerve (SGNGL of Pipa
and Cook) innervates op-t I and op-t 2 (52, 53 of Carbonell, 1947). In Periplaneta the
most anterior prothoracic nerve (nerve 2 of Pip a and Cook) fuses with the anterior
ganglionic connective arising from the interganglionic connective (nerve 1 of Pipa and
Cook), and the fused nerve diverges further and is joined by a lateral nerve arising from
the mesothoracic ganglion. Determination of the segmental origin of muscles innervated
by the nerves of such complex origins cannot be made, unless the fibers of each individual
nerve are distinguished histologically. Nevertheless, it is clear that in Periplaneta, as in
1
The known developmental facts supporting this premise are very few and fragmentary. The relationship
between nerves and muscles must be studied from their earliest developmental stages, and in many forms
before it can be more reliably used. '
2ND CERVICAL _ ____,.
NERVE
MATSUDA: TilE INSECT TIIORAX
~ - - - - SUBOES. GANGLION
r--- ANT. GANGL. CONNECTIVE
t-cv 2
t-cv 3,
t-s 1
7
FIG. I. Diagram of some cervicothoracic nerves in Dissosteira carolina (from Schmitt).
Dissosteira, some dorsal pronotal muscles of the head receive innervation from a purely
suboesophageal nerve.
Schmitt (1962) reviewed the literature on the pattern of innervation of the second
(posterior) cervical nerve in various insects that were studied by Holste ( 1910), Maki
(1936), Marquardt (1939), Nliesch (1954), Wittig (1955), Pipa and Cook (1959), and
Schmitt (1959). Schmitt concluded that the second cervical nerve generally provides
innervation to the pronotal muscles of the head, and in some insects the first and second
tergal muscles (op-t 1, 2) receive innervation from, both the second cervical muscle and
the prothoracic dorsal nerve.
These facts suggest that the dorsal muscles of the head are at least partly labial in
ongm. Here the neurological facts and the embryological facts pertaining to the origin
of the external dorsal area and some dorsal muscles are in general agreement. Most of
the other neck muscles are probably of prothoracic origin, although possibly the ventral
longitudinal muscle (s 1 or 2) is labial in origin, as Wiesmann (1926) found embryo-
logically in Carausius. These neurological and embryological facts indicate further that
the cervical area is a composite area resulting from the fusion of the dorsal labial and
ventral prothoracic segments. As shown in Part II of this work, however, this generaliza-
tion does not apply to the entognathous Apterygota, in which the qorsal part of the
neck region is probably a part of the prothorax.
The origin and evolution of the cervical sclerites: The origin of various cervical
sclerites, whether they are labial or prothoracic in derivation, has been disputed. The
lateral cervical sclerite, which occurs in most orders, is considered first.
In Lepismachilis (Fig. 35), as Barlet ( 1967) has shown, what was called the
processus postgenal evidently corresponds to the condyle that articulates with the
anterior end of the lateral cervical sclerite, because it is continuous with the lateral end of
the tentorium. The plate that Barlet called sclerite anapleural superieur articulates with
the postgenal process. In Lepismachilis therefore it is clearly the pleuron that articulates
with the postgenal process.
The postgenal process has another process that projects backward (Fig. 35), which
Barlet ( 1967) called sclerite postlabiale. Two muscles of Barlet, 50 and 54, are inserted
on this postlabial sclerite and the postgenal process; judging from their positions, these
muscles are probably homologous with t-cv 2 and t-cv 3 in the Pterygota, which are
inserted on the anterior end of the lateral cervical sclerite.
Henry (1958) ignored this possibility of double innervation of a muscle.
The above similarity in position and in the associated musculature between the
sclerite anapleural superieur and the lateral cervical sclerite leads us to suspect strongly
that the sclerite anapleural superieur in Lepismachilis and the lateral cervical sclerite in
some pterygotes are probably homologous. The next question is, "What part of the pro-
pleuron corresponds to the lateral cervical sclerite, and in what insects?"
In Schizocerus (Fig. 155B) and other Hymenoptera there is no isolated lateral
cervical sclerite. Instead, the anterior end of the preepisternum articulates directly with
the head, just as in Lepismachilis. Therefore, the lateral cervical sclerite in other insects
can be a detached preepisternum.
In Micropteryx (Fig. 144A) the position of the lateral cervical sclerite relative to
the neighboring plates is the same as that of the preepisternum in the pterothorax. In
Trichoptera (Fig. 141A) the lateral cervical sclerite is so obviously serially homologous
with the preepisternum in the pterothorax, that Tindall (1965) had no hesitation in
identifying the two structures. In Sialis (Fig. 118A), again, the lateral cervical sclerite
is obviously a modified preepisternum. Mutuura (1961) also homologized the lateral
cervical sclerite in Mecoptera with the prothoracic episternum. In Odonata, which share
a number of plesiomorphic characters with Machilidae, the lateral cervical sclerite is of
preepisternal origin ( p. 386). In higher holometabolous insects, such as Diptera and
Lepidoptera (Monotrysia, Ditrysia), the preepisternal origin of the lateral cervical sclerite
has been generally obscured.
In all hemimetabolous orders and in Coleoptera the lateral cervical sclerite is of
presternal origin. In Lepisma (Fig. 37), as well as in Lepismachilis (Fig. 35), the pro-
thoracic presternum extends laterally anterior to the episternum. In the mesothorax of
Plecoptera (Fig. 48) the presternum occupies a corresponding position to those in
Lepisma and Lepismachilis; it is fragmented into median and paired lateral pieces, and
the latter is often called the interpleurite. In the neck region of Capnia (Fig. 46A) , the
lateral cervical sclerite and the presternum clearly correspond to the three presternal
sclerites in the mesothorax of Perla (Fig. 48). These correspondences in position of the
sclerites in question lead us to conclude that the lateral cervical sclerite in Capnia may
well be of presternal origin. Further, the fact that in Plecoptera the lateral cervical
sclerite articulates with the anepisternum and the preepisternum is fused with the basi-
sternum, excludes the possibility that the lateral cervical sclerite can be of pleural (pre-
episternal) derivation. -
Similarly, in the dermapterous genus Allostethus (Fig. 79A) the lateral cervical
sclerite clearly articulates with the reduced preepisternum; in Blattaria (Fig. 54A) the
lateral cervical sclerite articulates with the precoxal bridge, which is largely preepisternal;
in other related orders and in Coleoptera also, the lateral cervical sclerite articulates with
the preepisternal area. These facts suggest that the lateral cervical sclerite in these
orders is presternal in origin, as in Plecoptera. Fragmentation of the lateral cervical
sclerite into anterior and posterior pieces is common in lower hemimetabolous orders.
In most Hemiptera the lateral cervical sclerite has been lost.
In Megaloptera the preepisternum and the basisternum are firmly united, and
together they extend forward; an additional ventrolateral cervical sclerite of presternal
origin lies anterior to the preepisterno-basisternum (p. 285).
The above discussion on the origin of the lateral cervical sclerite leads us to conclude
that at least four distinct types of lateral cervical sclerites (undetached preepisternal,
detached preepisternal, presternal, preepisterno-presternal) occur in the Pterygota.
With regard to the origin of the dorsal cervical sclerite, Mutuura (1961) contended
that in Panorpa the dorsal cervical sclerite represents the antecosta, because a dorsal
longitudinal muscle [cv(d)-t 1] from the first phragma is attached to it. Alicata (1962b)
came independently to a very similar conclusion; he thought that the dorsal cervical
sclerite in orthopterous genera Dolichopoda and Tfoglophilus represents the acrotergite,
because the dorsal longitudinal muscle [cv(d)-t 1] is attached to it.
MATSUDA: THE INSECT THORAX 9
The muscle cv(d)-t 1 is serially homologous with t 14, which extends between the
phragmata. On the basis of associated musculature, therefore, these conclusions appear
to be valid. . However, the evolution of the muscles and the integument to which the
m u s c l e ~ are attached could have proceeded independently, both in phylogeny and
ontogeny, as Matsuda ( 1965) discussed fully.' Therefore, the dorsal cervical sclerite is
not necessarily the detached acrotergite of the phragma, as these authors believed. It
appears more probable that the dorsal cervical sclerite is purely a secondary sclerite
produced for the attachment of the muscle cv(d)-t 1. This idea of the secondary origin
of the sclerite explains better the reason why the dorsal cervical sclerite varies so much
both in shape and position in various insects. To be fully acceptable, the idea of the
acrotergal origin of the dorsal cervical sclerite requires more evidence, such as transitional
stages showing gradual detachment of the dorsal cervical sclerite from the pronotum in
phylogeny, and this gradual detachment should be comparable with the gradual detach-
ment of the preepisternum (lateral cervical sclerite) in the phylogeny of some holo-
metabolous insects.
The ventral cervical sclerite is merely a secondary sclerotization. It may be repre-
sented by one or more transverse sclerites between the lateral cervical sclerites, and
usually occurs in orders in which the dorsal cervical sclerite is present. In Hemipteroidea
and in' most holometabolous insects, no well-formed ventral cervical sclerite has
been found.
DuPorte ( 1960, 1962) proposed that the gula originated in the prognathous head by
incorporation of the ventral cervical sclerites that had become united with the postocciput
and the subgena. In support of his contention, DuPorte adduced the condition of the
neck region in Anisolabis, in which the anterior ventral cervical sclerite is definitively the
gula that is united with the submenturn, but is separated from the postocciput. DuPorte
believed that this condition represents a transitional stage toward the more solidified
gular area in Coleoptera. DuPorte (1962) admitted, however, that in certain larval
Coleoptera, as Snodgrass ( 1960) showed, the definitive gula is apparently formed by the
mesal growth of postgenae. Essentially in agreement with DuPorte, Akbar ( 1965)
admitted, in the definitive gula of some Coleoptera, that the sclerotized neck membrane
is included.
Theories: Verhoef! (1902-1904), as a corollary of his bold assumption that the
intersegmental areas of the thoracic segments represent independent segments, contended
that the neck region also represents an independent segment, the microthorax. This
theory is entirely inconsistent with embryology, which has consistently shown no evidence
that supports the presence of a segment between the head and the prothorax. Since the
theory has been repeatedly discredited over the years, it is no longer necessary to discuss
it in detail.
Crampton (1917a, 1926a) was right in negating the presence of a distinct segment
in this area and also in refuting the interpretation of Riley ( 1904) that the cervical area
is wholly labial in origin. But Crampton went too far when he denied, in his reinterpreta-
tion of Riley's work, the possible inclusion of the labial segment in the neck region of
termites (Holmgren 1909). It appears that to the mind of Crampton the neck region
was a specific area which does not include either the prothoracic or the labial element.
Essentially, therefore, Crampton's concept of the intersegmental area for the neck region
was equal to the microthorax of Verhoef!, although Crampton did not call the area a
segment. A definitive intersegmental area should belong to a segment that is either
anterior or posterior to the area in question, or should belong to both, as Crampton
( 1914b) thought.
Crampton homologized his (definitive) intersegmental pleural sclerite in the
posterior thoracic segments of Eosentomon and Japyx with the lateral cervical sclerite in
Capnia (Fig. 2). The interpleurite of Crampton was therefore partly based on the
See also p. 85 in this work.
10
MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA
FIG. 2. Ventral region of the prothorax of Capnia, the mesothorax of Eosentomon, and the
metathorax of Japyx; drawn as though spread out in one plane. Shaded areas represent inter-
segmental areas of Crampton (from Crampton).
contemplation of structures on Eosentomon and perhaps in other Protura, in which
fragmentation ofsclerites frequently occurs (Fig. 28).
Nevertheless, Crampton's homologization of the lateral cervical sclerite with the
interpleurite, which often can be a detached presternal sclerite (see p. 24), was essentially
correct. Such a homologization is valid, however, only for hemimetabolous insects and
Coleoptera. Crampton was, therefore, not justified in extending his homologization to
other orders.
Snodgrass' ( 1935) theory on the origin of the neck region states that this region
contains an invisible primary intersegmental line between the labial and the prothoracic
segments. It further states that the principal dorsal longitudinal muscle (op-t 3 or
cv( d) -t 1), which connects the back of the head and the antecosta of the mesonotum, and
the ventral longitudinal muscle between the posterior tentorium and the profurca (s 1 or
2) are fusion products of the muscles of the labial and prothoracic segments, which were
originally attached to this supposed primary intersegmental line (or to the antecosta).
Snodgrass' theory is correct is maintaining a composite nature of the neck region. The
muscles in question, however, are serially homologous with the segmental muscles in the
pterothorax and cannot be fusion products, although they may receive double
innervations.
Some earlier workers including Riley (1904) thought the neck region was labial in
ongm. Recently Wada (1966) came to a conclusion similar to Riley's. His study of
the results of artificially produced defects and other teratological materials in the embryos
of Tachycines indicated the presence of morphogenetical units, which Wada correlated
with segmental composition of the head, including the neck. His findings led him to
conclude that the largest part of the neck was of labial origin, the dorsal side of the neck
immediately behind the postocciput was maxiiiary, and only the very narrow hindmost
strip was prothoracic in origin. In this kind of experimentation, however, the defect-
producing operations are made at a certain time or times during the course of embryonic
development. After the operations, the regulative aspects of development would be
eliminated or at least disturbed. Consequently, the results may refer to artificially
produced mosaics of embryonic parts, and is not valid for the identification of a segment.
THE TERGUM
Division of the tergum: In Lepisma (Fig. 37) a tergum consists, from the front
backward, of the antenotum, the prescutum, the scutum, and the postnotum. In the
Pterygota a typical pterothoracic tergum consists of the acrotergite, the prescutum, the
scutum, the scutellum, and the postnotum. Besides the divisions of the scutum and
scutellum, which will be discussed later, misinterpretations have developed concerning
the homologies of the anterior and posterior parts of the tergum in Thysanura and the
Pterygota. The confusions arose from the fact that a well-formed mesothoracic post-
notum is absent in such lower pterygote orders as Blattaria, Mantodea, and Orthoptera,
and that the postnotum is often unrecognizable or difficult to recognize in immature
insects.
The fact that the postnotum is clearly present in Ephemeroptera and Plecoptera,
which exhibit so many primitive features, however, supports the idea that the presence of
the postnotum represents a primitive condition. The postnotum in these orders is prob-
ably homologous with the postnotum in Lepisma (Fig. 37) and Oncojapyx (Fig. 32),
which, according to Barlet (1951) and Barlet and Carpentier (1962), descends to meet
the dorsal posterior part of the pleuron as in pterygotes. Further, the postnotum in
Lepisma and Oncojapyx is dorsally hidden beneath the tergallobe as in pterygotes.
In the mesothorax of Blattaria and Orthoptera, to which Snodgrass (1927) attached
importance in postulating his theory (to be discussed below), the reduced postnotum is
hidden beneath the posterior area of the mesonotum (Figs. 54B, 70B), although
Snodgrass failed to recognize it. In some higher orders such as Diptera and Hemiptera
the postnotum has been greatly enlarged. Evidently, therefore, the evolution of the
mesothoracic postnotum has been either toward reduction, or toward enlargement.
When the postnotum is reduced laterally the postalar bridge is not formed.
Snodgrass (1909, 1927, 1935) believed that the postnotum is a secondary develop-
ment that occurred with acquisition of the wing. He ( 1927) attributed its origin to the
prolongation of the acrotergite that has developed along the a'ltecosta, or the true inter-
segmental ridge (Fig. 7 A). According to Snodgrass ( 1927, 1935), the postnotum
(Fig. 7C ). consists of the greatly enlarged acrotergite and the antecosta bearing the
phragma; the postnotum may contain a narrow strip, the posttergite, immediately behind
the antecosta; the phragma is a single walled structure that arises from the antecosta,
despite his earlier ( 1909) contention that the phragma arises from the posterior end of
the postnotum (the postphragma) and also from the antecosta (the prephragma) of the
succeeding segment. Weber (1924b) also emphasized the secondary nature of the post-
notum, but thought the phragma was primarily double layered as Snodgrass (1909)
thought. Weber attributed the development of the phragma to muscle action.
It should be pointed out, however, that in Ephemeroptera, Plecoptera, and some
other orders the second phragma is double walled, which clearly shows that both the
postnotum and the antecosta can produce the phragma, and this fact decisively disproves
the theory of Snodgrass (1927, 1935) on the origin of the postnotum. Clearly,
Snodgrass' theory was based on the preconception that the phragma arises only from the
antecosta; and this preconception apparently forced him to stretch his imagination and
postulate that the postnotum is an enlarged acrotergite. In any case, the formation of
PRESCUTOSCUTAL SUTU PH RAGMA PRESCUTUM GITE
ANTE COST
PARAPSIDAL
SUTURE
ANTEROLATERAL
SCUT AL SUTURE
TRANSSCUTAL
SUTURE
TERGAL FISSURE)
AXILLARY
CORD
LATERAL PARAPSIDAL
SUTURE
ARM

r---ANTERIOR NOTAL
WING PROCESS
AL
POSTMEDIAN
NOTAL WING
PROCESS
ERIOR
NOTAL WING
PROCESS
SCUTOSCUTELLAR SUTURE
RECURRENT SCUTOSCUTELLAR SUTURE
FIG. 3. Diagram of tergal sutures and structures in the pterothorax (original).
the double walled phragma, which frequently occurs in pterygotes, is difficult to explain
in terms of the theory of Snodgrass. If this theory is followed, one must assume that
the phragma invaginates from the antecosta, then turns sharply upward to become the
postnotum (anterior wall of the phragma), which is indeed difficult to conceive.
The acrotergite, which Snodgrass considered to be the forerunner of the postnotum,
is merely a narrow marginal strip anterior to the antecosta, and hence is homologous
with the antenotum in Lepisma. The acrotergite cannot develop into a large postnotum.
An intratergal phragma (pseudophragma) in Diplura, Thysanura, and Odonata is dis-
cussed in later sections (pp. 97, 388).
Tergal sutures of the pterothorax (Fig. 3): As a consequence of the acquisition of
wings, various sutures have developed on the tergum of wing-bearing segments. On the
tergum, therefore, two kinds of sutures occur: those inherited from the Thysanura-like
ancestor, and those developed after acquisition of wings.
The anteriormost suture in Lepisma is the antecosta that demarcates the antenotum
and gives rise to the endonotum internally. In the Pterygota the antecosta demarcates the
acrotergite and internally gives rise to the phragma. Obviously, therefore, the acrotergite
is homologous with the antenotum, and the endonotum is homologous with the phragma.
The histological difference between the endonotum and the phragma can be compared
with the tentorium of epidermal origin in Thysanura and the tentorium in the Pterygota,
which is produced by the ectodermal invagination.
The prescutoscutal suture in Lepisma (Fig. 37) is a nearly straight transverse suture
running across the tergum near its anterior margin and behind the antecostal suture.
The prescutum, demarcated posteriorly by this suture, is therefore a rather narrow trans-
verse plate. In the Pterygota the homologous suture is the anterior ventral notal ridge
of Snodgrass (1909) and the prescutoscutal suture of Snodgrass (1927, 1935), which
agrees in position with the prescutoscutal suture in Lepisma. The prescutoscutal suture
is present in Ephemeroptera, Plecoptera, Blattaria, Mantodea, Dermaptera, Orthoptera,
MATSUDA: THE THORAX 13
and some other orders. When this suture is retained laterally, it reaches the posterior
limit of the base of the prealar arm (Fig. 3).
Typically, the scutoscutellar suture in the Pterygota is an anteromesally produced
suture that arises laterally from the area anterior to the axillary cord on each side. It
occurs commonly in holometabolous insects and less frequently in hemimetabolous
insects. The fact that it is absent in Ephemeroptera and Plecoptera, as in Thysanura,
makes it doubtful that this suture has been inherited from the Thysanura-like ancestor,
despite the occurrence of a doubtful scutoscutellar suture in the pseudofetus of Dilta
(Bin Fig. 36A), which is discussed on p. 104.
Several other tergal sutures discussed below have evidently developed newly in the
pterothorax. While their functional significance is often unclear, some of these sutures,
especially the parapsidal and the lateral parapsidal sutures, recur frequently, regardless
of the phylogenetic relationships of the groups in which they occur. An extensive study
of these newly acquired sutures would yield a wealth of data for the study of convergence.
The anterolateral scuta! suture cuts the anterolateral angle of the scutum that bears
the anterior and the antemedian notal wing processes; this area is called the suralare. In
Pterondrcys (Fig. 46B), Stagmomantis (Fig. 62), and in many others this suture is con-
tinuous with the prescutoscutal suture. The suture is strongly ridged in Lepidoptera; in
the metathorax of some Homoptera and Hymenoptera, the corresponding area is mem-
branous and the suralare becomes an isolated sclerite.
The paired parapsidal sutures (Fig. 3) arise anteriorly from the prescutoscutal
suture. Often the sutures on the two sides become confluent posteromesally. Many
earlier workers, including Snodgrass (1909), Crampton (1914a), and some modern
workers, wrongly designated the area bounded medially by the parapsidal sutures as the
prescutum in Hymenoptera, Diptera, Hemiptera, Coleoptera, etc. In the mesothorax
of Ephemeroptera, Hymenoptera, Bittacus, and Diptera the prescutal area extends
posteriorly along the anterolateral margin of the greatly elongated mesonotum. In these
forms the parapsidal sutures have shifted their positions considerably caudad, but they
maintain the connections with the prescutoscutal suture, as in other orders.
The lateral parapsidal sutures in Plecoptera (Figs. 46B, 47A) and other orders arise
from the lateral of the prescutoscutal suture on each side, and extend posteriorly
towards the median longitudinal axis, where they may become confluent. Weber ( 1924b,
1933) mistook these sutures for the parapsidal sutures, and called the median area
bounded by them the prescutum in error. The lateral parapsidal suture always starts
from the anterior limit of the lateral scuta! margin or from the posterior (or lateral) end
of the prescutum delimited by the prescutoscutal suture. It is important to point out
that the anterior end of the anterolateral scuta! suture also usually extends to the point
where the lateral parapsidal suture starts (Fig. 3). Therefore, the three sutures (pre-
scutoscutal suture, lateral parapsidal suture, and anterolateral scuta! suture) typically
meet at one point along the anterior lateral margin of the tergum; this fact provides an
important landmark in determining the border between the prescutum and the scutum,
which is useful even when one or two of the three sutures are absent.
The transscutal suture connects the scuta! lateral margin immediately behind the
anterior (or antemedian) notal wing process on each side. The tergal fissure occurs in
this area in many lower forms of many orders. Most probably, therefore, the transscutal
suture was produced by sclerotization and ridge formation along the tergal fissure; this is
obvious, for instance, in Hymenoptera. In sawflies the membranous tergal fissure is
clearly present, but in many higher Hymenoptera this area is represented by an internally
ridged suture traversing the entire scutum of the mesothorax. A well-developed trans-
scuta! suture is also present in many Heteroptera, Diptera, Hymenoptera, and sporadically
in some other orders.
The posterolateral scutal suture is a marginal suture demarcating the posterolateral
angle of the scutum that bears the posterior notal wing process. It arises from the scutal
lateral margin anterior to the posterior notal wing process, and runs obliquely medic-
caudad. The degree of development of this suture varies considerably in different
insects; in some it extends as far as the scutoscutellar suture, and in others it extends
even beyond (e.g., Homoptera).
The recurrent scutoscutellar suture arises from or near the middle of the hind
margin of the scutellum, then diverges anteriorly on both sides in varying lengths. This
suture occurs in Ephemeroptera, Plecoptera, Orthoptera, and others; it lies proximal to
the posterolateral scuta! suture.
Modifications of the tergal surface: The most marked and frequent modification of
the tergum is the prolongation of the mesothoracic scutellum. The prolonged scutellum
either fits in special conformations of the metanotum, or simply lies above the metanotum
,and projects beyond. Such a secondary prolongation of the mesothoracic scutellum has
occurred independently at least in Dermaptera, Coleoptera, Diptera, Hymenoptera,
Hemiptera (Homoptera, Heteroptera), and perhaps in many other pterygote orders.
For accommodation of the forewing, modification of the scuta! area of the mesonotum
has occurred in certain orders, such as Hemiptera, Coleoptera, and Dermaptera. In
certain cases, the groove formed on the mesonotum for holding the wings may be
associated with the posterolateral scuta! suture. The loss of tergal sutures occurs in
short-winged and wingless forms.
The lateral margin of the pterothoracic tergum: With the acquisition of wings,
various articulatory processes have developed on the lateral margin of the pterothoracic
tergum. The most anterior of these is the prealar arm, which is often a lateral prolonga-
tion of the prescutum and extends behind the tegula; it is often called the prescutal arm.
The present work has shown, however, that the prealar arm can also be acrotergal in
origin in Blattaria, Mantodea, and some other orders, as LaGreca (1947b) pointed out
in Orthoptera. The prealar arm is often united with the dorsal or anterodorsal part of
the anepisternum, and together they form a prealar bridge. The prealar bridge is
usually absent in the metathorax, although it is present in some Neuroptera and perhaps
in some other lower orders. In Mantodea the prealar arm invaginates into the thoracic
cavity, and in some Orthoptera it is differentiated into an articular lobe. In most
Mecoptera the prealar arm is absent, but in Diptera ah arm, comparable in position and
shape with the prealar arm, is formed. In Hymenoptera also, the prealar arm is absent.
LaGreca (1947b) and his students carefully studied the scuta! lateral margin. In
several orders they discovered five points of articulation with the axillary sclerites
(Fig. 3), instead of two (Snodgrass 1935), or three (Weber 1933). These points of
articulation are projections with varying degrees of development. The anteriormost is
the processo notale anteriore (anterior notal process), and in some cases, the processo
notale media is differentiated immediately behind. These two processes together cor-
respond to the anterior notal wing process of Snodgrass. The processo notale media is
renamed the antemedian notal process, because of its position relative to three processes
that follow.
The membranous tergal fissure extends proximally, separating the antemedian notal
wing process from the median notal process which La Greca called processo b (hintere
Tergalhebel of Weber 1924b, 1933). Another small process, v of La Greca, lies
posterior to his b, which in this study is called the postmedian notal process. The
hindmost process is the long-recognized posterior notal wing process; the posterior notal
process is often detached from the scutum itself, and when this is so it. is usually called
the fourth axillary sclerite. Typically, the first four processes articulate with the first
axillary sclerite, and the last with the third axillary sclerite.
Of these notal wing processes, the median and the postmedian processes often are
pronounced in the higher orders. In Orthoptera, for example, the antemedian notal
process is well differentiated and lies clearly separated from the anterior notal process,
whereas in other lower hemimetabolous insects it is often difficult to distinguish from the
anterior notal process. The median and postmedian processes are conspicuously
developed only in a few orders such as Lepidoptera, Diptera, and Hymenoptera; and they
are inconspicuous or unrecognizable in Blattaria, Plecoptera, and Mantodea. All of
these facts suggests that primitively only the anterior and posterior notal processes were
MATSUDA: TiiE INSECT TiiORAX
15
c
An.
A
PROXIM. MED. PLATE JUGAL FOLD
FIG. 4. Diagram showing a primitive wing base (adapted from Snodgrass).
present and were articulated with the first and the third axillary sclerites, and that the
other processes differentiated later.
The tegula and the prealar sclerite: The tegula is usually a scalelike, setigerous lobe
overlapping the base of the wing. It has therefore intimate association with the humeral
plate. In Plecoptera and Dermaptera the tegula occurs both in the mesothorax and
metathorax; in other orders it is present only in the mesothorax, and in Coleoptera it is
absent even in the mesothorax. In Hymenoptera and Lepidoptera the tegula is greatly
enlarged and protects the base of the forewing; it is supported by the subtegula. In
Psocoptera the tegula is absent.
The small prealar sclerite often occurs, in association with the tegula. This sclerite
is presumably a detached prescutal sclerite. Usually muscle t-p 4 is attached to this
sclerite. The subtegula in Trichoptera and Lepidoptera is homologous with the prealar
sclerite in other orders.
The axillary sclerites: In Odonata the axillary sclerites are unique, except for the
first axillary sclerite, which can be homologized with that of the other pterygotes. The
second and third axillary sclerites in other orders are not separately differentiated in
Odonata, and the two sclerites are represented by a single, posterior articulatory plate
called the axillary plate. However, in another order, Ephemeroptera, in which also the
wings are not flexed at rest, all three axillary sclerites are differentiated and maintain the
same topographic relationships among themselves as they do in many other orders. An
uoique feature in Ephemeroptera, however, is that the median plate is not divided into
proximal and distal parts by the basal fold.
In lower neopterous orders each axillary sclerite has more or less fixed topographic
relationships with other axillary sclerites, the scuta! lateral margin, and with the wing
veins (Fig. 4).
The first axillary sclerite articulates with the anterior notal wing process and with
the scuta! lateral margin. Often the median and the postmedian processes also provide
16
points of connection with the first axillary sclerite. Laterally, this sclerite is associated
with the proximal margin of the second axillary sclerite, and connected with the proximal
end of the Sc vein. Its posterior margin is often connected with the proximal process
of the third axillary sclerite by a tendinous ligament.
Besides its association with the first axillary sclerite, a typical second axillary sclerite
is laterally joined to the proximal median plate and by its anterior margin to the base of
the R vein; posteriorly, it articulates with the third axillary sclerite. The second axillary
sclerite is sclerotized on the ventral surface of the wing base membrane, and rests on or
is connected by a ligament to the pleural wing process: it is therefore the pivotal sclerite.
The ventral second axillary sclerite is often associated, either by fusion or connection,
with the third axillary sclerite and with the subalare.
The third axillary sclerite has typically three processes or projections: anterior,
proximal, and distal. By its proximal projection it articulates with the posterior notal
wing process, and its proximal margin has a ligamentous connection with the posterior
margin of the first axillary sclerite. Its distal end connects with the A veins.
The proximal median plate lies distal to the second axillary sclerite. Typically, its
distal end is separated by the basal fold from the distal median plate. The basal fold
forms a prominent convex fold during flexion of the wings.
Judging from their stable connections with particular wing veins in lower pterygotes,
the first, second, and third axillary sclerites can be safely regarded as modified and
detached basal parts of the wing veins: the first axillary sclerite is the base of the Sc vein;
the second is that of the R vein; and the third is that of the A veins. The small humeral
plate on the proximal anterior margin of the wing represents the detached base of the C
vein; in Odonata this plate is large'. In Plecoptera (Fig. 47A) and Ephemeroptera
(Fig. 44A) the distal median plate is not differentiated, or poorly differentiated, and the
association of the proximal median plate with the M and Cu veins is definite and direct.
The proximal median plate is comparable with the axillary sclerites, because it represents
the basal part of the M-Cu veins.
The formation of the distal median plate is therefore clearly secondary, although it
is common in winged insects. The development of the distal median plate as a means of
resisting elevation of the fold during flexion of the wing must have occurred simul-
taneously with the formation of the basal fold; this fact accounts for the absence of the
distal median plate in mayflies: which never fold their wings. In Blattaria (Fig. 55 A)
other veins also have additional detached bases.
Among the Neoptera the most primitive relationship between axillary sclerites and
wing bases is seen in Plecoptera (Fig. 47A), which was studied by Onesto (1965). To
reconstruct a probable primitive wing base (Fig. 4), therefore, the wing bases in Plecop-
tera and Ephemeroptera were seriously taken into account. The diagram (Fig. 4) is a
modification of a diagram given by Snodgrass (19 3 5) . The distal median plate is not/
shown, and the tendinous connection extends from the third to the first axillary sclerites
in this diagram.
As Sharplin ( 1963c) showed in Lepidoptera, the sclerotized elements at the wing
base are unusually flexible. The fuchsinophil mesocuticle at the wing base is not a con-
tinuous layer, but is divided into cones and pegs (Fig. 5). This discontinuous meso-
cuticle is subject to distortions, projecting down through the endocuticle. The continuous
exocuticular layer and subdivided mesocuticle constitute the bending cuticle. Sharplin
distinguished two types of the bending cuticle: the mesocuticle projecting into the
endocuticle as a series of irregular pegs varying in length and diameter; and the large,
more regularly spaced cones of the mesocuticle reaching nearly to the inner margin of
the endocuticle.
A rubberlike protein (resilin) that constitutes the ligaments connecting structures at
the wing base was recognized by Weis-Fogh ( 1960). These ligaments evidently play an
important role in the flight mechanism, and have been observed from time to time by
insect morphologists since Voss (1905). LaGreca (1947b), Sharplin (1963a, b), and
Homologies of the wing base structures in Odonata with those in a paleodictypterous species, Ostrava nigra,
are discussed on p. 43.
NORMAL
EXOCUTICLE
. . : ..
B
MATSUDA; THE INSECT THOKAX
BENDING
EXOCUTICLE
ENDOCUTICLE
17
FIG. 5. A, diagrammatic vertical sections of cuticle (from Sharplin); B, diagram of the
action of bending cuticle (from Sharp lin).
Hatch ( 1966), among more recent workers, observed the ligamentous connections.
La Greca ( 1961) made an electron-microscopic study of the wing base in Leucophaea
maderae.
THE STERNUM
The endosternum, the sternal furca, and the spina: In the Apterygota the endo-
sternal skeletons or processes take various shes, such as plate, stalk, bridle, etc. A
series of studies by Carpentier and Barlet during the past two decades have elucidated
these endosternal structures in Diplura, Collembola, and Thysanura in sufficient detail;
these studies made the following analysis possible.
Endosternal processes are often represented by muscles. For instance, the postcoxal
endosternal bridges d and j are muscles in Campodea (Fig. 33) and Tetrodontophora
(Fig. 31A), kin Campodea (Fig. 33) and Tetrodontophora (Fig. 31A), and n in the
mesothorax of Tetrodontophora (Fig. 31A) are represented by muscles. These facts
parallel the cases of some head muscles that are homologous with endoskeletal structures
in Thysanura (Matsuda 1965).
As Carpentier (1946b, 1949), Barlet (1951, 1952a, b), Carpentier and Barlet
(1951), and Barlet and Carpentier (1962) attempted to show, the relative positions of
endosternal processes are fairly constant throughout the Apterygota. There are some
inconsistencies, however. For instance, e is a furcasternal structure in Ctenolepisma
(Fig. 38A), but it is attached to the anterior margin of the spinasternum in Tetrodonto-
phora (Fig. 31A) due to reduction of the sternum (in the latter); in Ctenolepisma
(Fig. 38A) d arises from b and in Tetrodontophora (Fig. 31A) it arises from g; in
Lepisma (Fig. 37) there are twice the number of h in the mesothorax; in Thysanura flies
on the anterolateral angle of the furcasternum and meets the descending katapleuron,
although it lies on the katapleuron in Collembola. It is therefore not surprising to find
some inconsistencies in, for instance, Carpentier's work (1949) regarding the homology
of the poststernum based on the attachment of processes in Tetrodontophora (p. 92).
As will be pointed out again later, overreliance on the endosternal processes as landmarks
must be avoided. However, the positions of certain processes appear to be useful in
determining associated external parts of the sternum; thus, p is always attached to the
dorsal margin of the katapleuron; a is always attached to the spinasternum and l is
attached to the anterior margin of the succeeding segment.
The base of the sternal furca in the Pterygota is probably homologous with the
endosternal process f in the Apterygota, since in the Pterygota tlie sternal furca arises
from an area that corresponds to the position off in the Apterygota.The sternal furcal
base usually has an external pit, and must therefore arise from ectodermal invagination.
The evolution of the endosternal f and the furca is thus comparable with the evolution of
the tentorium; the epidermal tentorium in the Apterygota has been replaced by the
tentorium produced by the ectodermal invagination in the Pterygota (Matsuda 1965).
In Nicoletia (Fig. 39B) and Ctenolepisma (Fig. 38A) a is attached to the posterior
margin of the spinasternum (the poststernite of Carpentier 1946b and of Barlet 1952a),
and l is attached to the anterior part of the presternum of the succeeding segment in
Ctenolepisma, and to the corresponding area (the border between presternum 1 and
presternum 2 of Bar!et 1952a) in Nicoletia (Fig. 40). In Oncojapyx (Fig. 34) also a
and l arise from the positions corresponding to those in Nicoletia and Ctenolepisma, viz.
a from the border between the spinasternum and the presternum (the intersternum of
Barlet and Carpentier, 1962), and l from the border between the poststernum and the
presternum (the presternum 2 of Denis, 1949) of the succeeding segment. In Campodea
(Fig. 33) too, a and l occur in the corresponding positions.
In regard to the spina (m of Carpentier and Barlet) in the Apterygota, Carpentier
(1946b) found in Cteno/episrna (Fig. 38A) that it is not formed exclusively from a, but
that l also contributes. In Lepisma (Fig. 37) the spina (m) is apparently associated
with a and /. In Carnpodea (Fig. 3 3) the spina ( m) is smaller and its location is similar
to that in Thysanura. In Japygidae Barlet and Carpentier ( 1962) found that the
strongly developed cuticular spina arises anteriorly from the spinasternum; Barlet (1965)
further showed that in Oncojapyx the spina is also connected with l.
These facts show that the location of the spina in Japygidae, at least its anterior
limit, is different from that in Thysanura and Campodea. This difference, however, has
probably resulted from the fact that in J apygidae the spina is so strongly invaginated that
even its cuticular part and the more anterior part of the sternum (the spinasternum)
have been affected. In this respect, the spina is more specialized in J apygidae than in
Thysanura and Campodea.
In the lower Pterygota also, the spinal invagination involves the external cuticule,
and in its location the spina in the lower Pterygota is more comparable with that of
Japygidae than with that of Thysanura. Besides the furcal base and the spina, the distal
part of the furca in the Pterygota is associated with the pleural arm; and this distal part
of the furca is probably homologous with g in Apterygota, since p' always connects, either
as a muscle' or as an endoskeletal bridge, g and the dorsal margin of the katapleuron,
the site of the pleural arm in most Pterygota. In Odonata the prefurca is the invagina-
tion from the basisternal lateral margin, and is probably homologous with h in the
Apterygota. Other parts of the endosternum in the Apterygota have been lost in the
Pterygota.
Division of the sternum in the Apterygota: In a generalized thysanurous genus
Nicoletia (Fig. 40) and in Oncojapyx (Fig. 34), each thoracic segment consists of five
subdivisions or apotomes. They are, from the front backwards, the presternum, the basi-
sternum, the furcasternum, the spinasternum, and the poststernum. Although there is
no problem with the four anterior apotomes as integral parts of the sternum, the question
has been raised whether the part here called the poststernum represents the frontmost or
the hindmost part of a segment. If the location of the spinal pit, which corresponds to
a in Japygidae, is taken as the intersegmental boundary, this apotome must belong to the
succeeding segment. This is, in fact, the opinion of Denis ( 1949, 1963), who called the
apotome the presternum I. However, the concept of the intersegmental location of the
spina by earlier workers including Snodgrass (1935), which Denis accepted, was based
on the observation of pterygotes, in which this apotome and the associated internal
process l have been lost. It is highly probable, therefore, that the poststernum in the
Apterygota represents the hindmost part of a segment in a more primitive sternum.
15
p is probably homologous with muscle p-s 1, which connects the furcal arm and the pleural arm in the
Pterygota.
MATSUDA: TilE INSECT TIIORAX 19
Barlet and Carpentier's ( 1962) proposal of the term the "intersternum" for this plate and
the for its dorsal counterpart has been unfortunate, since such terminologies
are confusing (see p. 94 and p. 99).
In Oncojapyx (Fig. 34) the Y ridge or line is present on the middle of the sternum.
Its lateral end articulates with the coxa near the endosternal tendon f, and posteriorly it is
continuous with the spina. In the mesothorax of Campodea (Fig. 33) also the lateral
end of the Y ridge reaches the base of f. In the prothorax of Campodea the Y line
apparently becomes aT-shaped line whose lateral ends connect the inner coxal margins.
In the prothorax of Eosentomon (Fig. 29) the anterior transverse line lies in the area
corresponding to the horizontal line in Campodea and the two lines are probably
homologous. In the metathorax of Eosentomon (Fig. 29) both the Y line and the
horizontal line connecting the anterior end of the Y are present. Among Thysanura only
theY line is present in the mesothorax of Ctenolepisma (Fig. 38A), and theY line and
the horizontal line are apparently present in Petrobius (Fig. 38B); in Nicoletia (Fig. 40)
only the horizontal line is present. The horizontal line is associated laterally with f in
Nicoletia (Fig. 40) and probably also in Petrobius (Fig. 38B). This horizontal line is
therefore the sternacosta of Weber ( 1933), which is associated with the furcal base.
The sternacosta may or may not form a condyle, which articulates with the coxa in the
Apterygota.
These facts suggest that both the Y line and the anterior line (the sternacosta) are
primary parts of the sternum, and that the absence of either of them is due probably to
loss. Barlet and Carpentier (1962) have suggested that the triangular space between
the branches of the Y line in Diplura is basisternal, in the belief that the undulation
formed by branches of the Y corresponds to the sternacosta. A close examination of the
facts mentioned above, however, shows that the two structures may have originated
independently. It therefore remains arbitrary to regard either the Y line or the sterna-
costa as demarcating the furcasternum from the basisternum. In the present work
Weber's definition of the sternacosta is accepted and it is used to mark the line of
boundary between the furcasternum and the basisternum. Snodgrass ( 1935) also used
the horizontal stern acosta as the line between the furcasternum and the basisternum.
Division of the sternum in the Pterygota: In the Pterygota the original division of
the sternum has become obscured because of fusion, loss, and modification of parts.
The presternum in pterygote insects is difficult to recognize, due to loss of the line
separating it from the basisternum. The true presternum homologous with the one
in the Apterygota is, however, present at least in some Plecoptera. In Perla (Fig. 48)
the median presternal area is continuous with the basisternum; the lateral isolated plate or
plates lying anterior to the pleuron (interpleurites) probably represent the true isolated
presternum. This interpretation is supported by the fact that in Thysanura (Figs. 35, 37,
38, 40) and Diplura (Fig. 34) the presternum extends laterally anterior to the pleuron.
Crampton (1909) also recognized the presternum in Nemoura and Pteronarcys.
Although the line demarcating the presternum from the basisternum appears to be
present in Nico/etia (Fig. 40) and Petrobius (Fig. 38B), this line has become lost in
Ctenolepisma (Fig. 38A) and Lepisma (Fig. 37). It may be said, therefore, that the
suture between the presternum and the basisternum was lost early in the evolution of the
Thysanura-Pterygota complex.
According to Snodgrass (1935), the marginal narrow presternum is separated from
the basisternum by the internally reinforced suture in some insects. However, the pre-
sternal ridge or suture of Snodgrass occurs in higher insects, such as Calliphora, and is
absent in lower insects. Therefore, this suture is not the primary border between the
presternum and the basisternum. Most probably the ridge is a secondarily produced
reinforcing ridge and the separated area is the prepectus (or hypopteron), which occurs
along the anterior margin of the pleuron and the sternum in higher insects. Weber's
( 1924b) assertion that the presternum in pterygotes is a plate secondarily separated from
the basisternum is also unfounded.
The separation of the basisternum from the furcasternum is usually obscurely
defined in the Pterygota. In some pterygotes, however, the sternacosta and the Y ridge
are present and are most probably homologous with those present in the Apterygota.
As Weber ( 1933) showed, in Plecoptera the Y ridge is connected anteriorly by the
horizontal sternacosta, which extends to the furcal base on each side; and the Y reaches
posteriorly as far as the spina. Grandi (1948) also showed theY ridge in the larva of
Perla. Among other pterygotes, the sternacosta and the Y ridge are found also in larval
Sialis (Weber 1928a). Yet, in many pterygotes only the horizontal line connecting the
furcal bases is present, and this line probably represents the sternacosta, which delimits
the furcasternum from the basisternum. In most adult holometabolous insects, in which
the furcal bases are approximated towards the median longitudinal axis, the sternacosta
is lost.
Delamare-Deboutteville ( 1952) defined the sternellum in the Blattopteroidea as the
median elongate plate that extends into the basisternal area and bears the furca on its
posterolateral angles. If Delamare-Deboutteville used the term sternellum for the
furcasternum as it is generally understood, this definition of the sternellum ( furca-
sternum) is unacceptable, since the area in question lies anterior to the furcal base. The
sternellum of Delamare-Deboutteville is purely a specialization in Blattopteroidea and it
cannot be homologized with the Y ridge, which lies posterior to the furcal base. Laterally
the basisternum meets, or is united with, the preepisternum in lower pterygotes, as in the
Apterygota.
Crampton ( 1909) showed paired plates behind the furcasternum ( furcisternum of
Crampton) and in front of the first spina in Nemoura, and called them the postfurcal
sclerites. Weber (1924b) thought that the postfurcasternite (postfurcal sclerite of
Crampton) behind the furcasternum in lower Ptergota arose from the membranous
region between two neighboring sternal areas, perhaps as a result of the muscle attach-
ment ( t-s 2) , and that the relationship between the postfurcal sternite and the spina that
follows is analogous to the relationship between the postnotum and the phragma. He
further thought that the spina is only a detached part of the postfurcasternum; Snodgrass
(1927) accepted the interpretation of Weber. For Weber and Snodgrass the post-
furcasternum was a secondary development just as, in their opinion, the postnotum was
secondary.
The location of the postfurcasternum and its association with the spina exactly
reflect the relation between the spinasternum and the spina in the Japygidae. It seems
highly probable, then, that the postfurcasternum in some insects is in fact the remnant
of the spinasternum, and not a secondarily formed area. In many higher pterygota the
postfurcasternum or the spinasternum has become greatly reduced or virtually lost.
Increased association of the spina with the spinasternum, in turn, must have been a
/
contributing factor in the reduction of this sternal part. At the same time, the spina
has lost its connection with the endosternal process I, which arises from the poststermim;
consequently, the poststernum has become functionally meaningless and altogether dis-
appeared along with I. As a result of the loss of the poststernum, further, the spina
has come to occupy the definitive intersegmental position in the Pterygota. It is
possible, however, that this poststernal plate has become indistinguishably fused with the
presternum of the succeeding segment in certain primitive insects, such as Perla
(Fig. 48).
In higher insects the reduction of the posterior parts of the sternum has involved
generally the spinasternum and the furcasternum. In the mesothorax of Orthoptera, for
instance, the spina has come to occupy a position between furcal bases, as a result of loss
of these posterior sternal areas.
In many higher orders, such as Homoptera, Diptera, Hymenoptera, Coleoptera,
Odonata, and others, even the mesothoracic spina has been lost, and the mesothoracic
furcasternum has greatly reduced; consequently, the anterior part of the metasternum has
come to occupy the area immediately behind the mesothoracic furcal bases. In these
forms, therefore, the ventral border between the mesothorax and metathorax has become
obliterated. This increased obliteration of the ventral segmental border is in
turn directly responsible for the formation of the synthorax as a functional unit, which
has occurred in varying degrees in various higher orders.
The idea of the invaginated sternum (cryptosternum) in holometabolous insects was
first enunciated by Weber ( 1928a), who showed a probable sequence of the median
longitudinal invagination of the sternum in holometabolous insects. Weber, starting with
the sternum in Plecoptera, which lacks such an invagination, was able to show that the
furcae also have migrated mesally, accompanying the median longitudinal invagination.
As Weber showed, the furcae lie on top of the posterior end of the invaginated sternum
in Neuroptera, Hymenoptera, and Lepidoptera. Kelsey's ( 1957) study, which traced
the developmental changes of thoracic parts in Corydalus, amply supported the concept
of the cryptosternum, or the invaginated sternum.
Daly's ( 1964) study on the postembryonic development of the thorax in A pis, how-
ever, showed that the median longitudinal ridge (lamella) in the basisternal area is
produced by a migration or proliferation of cells from a low midventral ridge; and this
suggested that the basisternal median longitudinal elevation is not formed by _invagina-
tion. Furthermore, there are holometabolous insects in which the pleurosternal suture
actually exists, as pointed out by Matsuda ( 1960b) for Gnophomyia (Diptera) and
Phassus (Lepidoptera). In these forms the median longitudinal suture is present, in
addition to the pleurosternal suture, and the leg bases are unusually widely separated.
This fact indicates clearly that the median longitudinal suture can be formed in-
dependently of the pleurosternal suture, and hence the suture does not represent the line
of union of the presterna as Ferris maintained. These apparent discrepancies regarding
the origin of the median longitudinal suture or the cryptosternum, however, can still be
accounted for, as discussed below.
In Sialis (Weber 1928b), Corydalus (Kelsey 1957), Apis (Daly 1964), and
Cicadidae (p. 258), in which the development of the sternum has been traced, leg bases
are invariably approximated towards the median longitudinal axis in the adult, thereby
contributing toward formation, or at least tdward the elevation, of the posterior part
of the median longitudinal ridge. This mechanical pressure, however, probably does not
extend too far anteriorly into the basisternal area of the greatly enlarged mesothorax such
as the one present in Apis, which Daly studied. Hence, the kind of developmental
mechanism for the production of the median longitudinal ridge on the basisternum Daly
(1964) found may often be at work. With the presumed invagination of the sternum,
the pleurosternal border (suture) has often become obliterated in holometabolous insects,
as will be discussed later (p. 37).
THE INTERSEGMENTAL RELATIONSHIP
In considering the relationship among thoracic segments, it should be remembered
that: ventrally, both the anterior and the posterior ends of a segment have undergone
reduction in varying degrees in various groups of the pterygota; and dorsally, both the
hind margin of the postnotum and the antecosta produce the phragma, anq both are
primary parts of a thoracic tergum.
In Fig. 6, A represents the condition in Thysanura in which no reduction of parts
has occurred. The inconspicuous endonotum, with which the phragma in other insects
is homologous, arises from the antecosta. Ventrally, the endosternal processes a and I
are present. "
In A, B, C, and D, the first phragma always arises from the antecosta of the meso-
notum. In the absence of the prothoracic postnotum, this is the only point where the
first phragma can be formed. Ventrally, the presternum has become reduced or lost in
B, C, and D.
In B, the second phragma is formed by the union of invaginations from both the
hind margin of the postnotum and from the metathoracic antecosta. This condition is
A
B
PRC. SC. + SCL.
PN. AN.
PHR.l-
PHR. 2 (EN.2)
(EN.!)
SP. 1
PHR. 1
SP. 1
POSTS.
PREST. BS. FS.
PRC. sc. SCL. PN. ACRT.
J
FS. SPS.
SCL.
__ __::.:::..._ ____ 2
'1
SP. 2

FS. SPS.
SCL. PN.
____ Z
FS.

FIG. 6. Diagram showing evolution of intersegmental relationships in insects (original).
ACRT, acrotergite; AN, antenotum; BS, basisternum; EN, endonotum; FS, furcasternum; PHR,
phragma; PN, postnotum; POSTS, poststernum; PRC, prescutum; PREST, presternum; SC,
scutum; SCL, scutellum; SP, spina; SPS, spinasternum; a and /, endosternal processes. See
text for explanation of A, B, C, D.
seen in Ephemeroptera, Plecoptera, and others. Ventrally, the second spina arises
wholly from the ectoderm of the spinasternum. In C, the second phragma arises from
the metanotal antecosta, the mesothoracic postnotum having become greatly reduced or
lost. This condition exists in Blattaria, lsoptera, Mantodea, and others. The dorsal
longitudinal muscle extends between the phragmata that arise from the antecosta of the
two neighboring segments. In this respect, Blattaria, Isoptera, and Mantodea resemble
Thysanura, but they differ from Thysanura in the great reduction of the postnotum. In
D, the second phragma is formed only from t!Ie postnotum. This condition exists in
higher Hymenoptera, Diptera, Hemiptera, and others, in which forewings are the prin-
cipal flight organs, and the postnotum is well developed in the mesothorax. Here the
metathoracic dorsal longitudinal muscle is attached anteriorly to the mesothoracic
phragma; hence, the muscle extends beyond the original segmental border. The single-
walled second phragma in Neuroptera and Mecoptera may also be the extension from the
mesothoracic postnotum.
In the orders where the hind wings are the principal flight organs, such as Orthop-
tera, Coleoptera, and Dermaptera, the mesopostnotum is greatly reduced or lost, and the
MATSUDA; THE INSECT THORAX
\ \
\:
''
':
"l
=
INTERSEGMENTAL
I FOLO
I
7
A
SECONDARY ,
---;- 1 -
' I
I :
I
MEMBRANE
SECONDARY /
SEGMENT/
PHRAGMA HRAGMA
B
c
23
FIG. 7. Snodgrass' diagrams showing intersegmental relationships. See text for explanation
of A, B, C.
second phragma is formed only from the metathoracic antecosta, and the third phragma
is formed from the metathoracic postnotum. In Trichoptera and Lepidoptera the meta-
postnatal third phragma is detached from the rest of the metanotum and is firmly united
with the anterior margin of the first abdominal tergite.
Ventrally, the primary intersegmental relationship has often been completely
obliterated, owing to loss of the spina, arrd to the reduction or loss of the anterior and
posterior parts of the sternum. In B and C, the reduction involves only the presternum.
In D, however, even the mesothoracic spina has become lost. As already noted (p. 20),
in some orders (Diptera, Coleoptera, Hymenoptera, Homoptera, and Odonata) even the
spinasternum and the furcasternum have become reduced or lost. The lost areas, both
on the anterior and the posterior ends of the sternum (shaded in B, C, D), are occupied
by the posterior and anterior ends of the neighboring sterna (the presternum and the
metasternum), and the union of the two thoracic segments are increasingly strengthened
(synthorax). Dorsally, such a firm union or fusion of segments occurs only rarely (e.g.,
Mallophaga, Anoplura).
Theories: According to Snodgrass ( 1927, 1935), in soft-bodied segmented animals,
such as annelids and larval insects, the segmented regions of the body are separated by
cuticular constrictions of the integument. Internally, the intersegmental grooves form
folds, and on these folds the fibers of the principal longitudinal bands of somatic muscles
are attached (Fig. 7 A). Snodgrass called such a system the primary segmentation.
With sclerotization of the body wall, however, membranous intersegmental areas
were left out to allow flexibility in the movement of body segments, and a definitive
sclerotized segment (Fig. 7B) in many arthropods is no longer coincidental with the
primary intersegmental folds, since it does not cover the posterior part of the original
segment and may extend anteriorly before the intersegmental groove, on which the
dorsal longitudinal muscle is attached. This new way of dividing the body is clearly
secondary, and constitutes a secondary segmentation.
6
Here the word intersegmental is used in the functional sense. Typically. the area belongs morpho1ogica11y to
the. segment that precedes the true intersegmental fold, as Snodgrass (1927) stated. .
24
In the thoracic segments, according to Snodgrass, the secondary segmentation
prevails. Snodgrass thought that the postnotal plate was actually an intersegmental plate
comprising two segmental elements (Fig. 7C), although the dorsal longitudinal muscles
are attached to the phragma that arises from the antecosta. Snodgrass also explained
the secondary ventral intersegmental relationships, by assuming the production of a plate
analogous to the dorsal postnotal plate.
As already concluded ( p. 11), Snodgrass' theory on the formation of the postnotum
is not supported by facts, and his idea of the secondary segmentation cannot be accepted.
The facts show clearly that the intersegmental relationship' is much simpler than
Snodgrass considered it to be. His complicated theory was clearly based on his pre-
conceptions that a phragma arises only from the antecosta and that the postnotum is a
secondary development.
Finally, it should be pointed out that in both Snodgrass' and Weber's diagrams
(1924b, 1933) of the hypothetical primitive intersegmental relationship (Fig. 7B), the
dorsal muscles are shown attached between the antecostae. Such dorsal musculature,
however, certainly is not typical in Thysanura. Dorsal muscles in Lepisma (Figs. 19-21)
and in Machilidae (Barlet 1950a, 1953, 1954, 1967) extend freely beyond segmental
borders. It is only in the Pterygota, with already well-developed phragmata, that the
distribution of dorsal and other muscles is largely confined within each segment.
Chadwick ( 1957) proposed the concept of the intersegmentallaterosternites for sites
of muscle attachments that lie at the lateral extremities of ventral intersegmental folds.
He inferred the presence of such an area from a comparison with the anterolateral area
of each abdominal sternite, which serves as an attachment point for apparently serially
homologous muscles. On the thoracic pleurosternal wall the primary intersegmental
line is obliterated. It is, therefore, difficult to know to which segment a sclerite on the
definitive intersegmental fold actually belongs. Often, however, these sclerites are prob-
ably the isolated presternum, as already discussed (p. 19).
These isolated plates have been recognized previously, for example, by Crampton
( 1926a), who called them the furcillae; but more often they have been called the inter-
segmental pleurites or the interpleurite. The term interpleurite is retained in this work,'
with the understanding that the plate may well be of the sternal (presternal) origin.
Barlet and Carpentier's (1962) and Barlet's (1967) concept of the intersegmental
relationship in the Apterygota is discussed on p. 94.
THE ORIGIN OF TIIE PLEURON
Since the embryological work on Hemiptera by Heymons ( 1899), the thoracic
pleuron in insects has generally been considered to owe its origin to the basal part of the
coxa or the subcoxa, which becomes separated and imbedded into the lateral wall of
the thorax between the tergum and the sternum. It has also been implicit in most works
that the same developmental mechanism as the one found in Hemiptera is at work to
produce the pleural wall throughout the Arthropoda. Recently, however, Snodgrass
(1952, 1958), Bekker (I 960), FUller (1963b), and Sharov (1966) have disputed this
theory of the subcoxal origin of the pleuron in the Arthropoda. It seems highly appro-
priate, therefore, to reexamine closely the pertinent facts.
Regarding the origin of the pleuron in the Crustacea, Snodgrass ( 1952) thought
that what Hansen (1925) considered to be the subcoxa in Anaspides (Fig. 8D) was
actually the laterotergite or the lateral subdivision of the tergite, because of the passage
of the body muscles through this area and their insertion on the coxa, as in other
arthropods. In various groups of the Isopoda, however, Gruner ( 1954) was able to
show varying degrees of incorporation of the coxa into the lateral wall of the thorax.
He showed also embryologically that in Porcellio the coxa becomes incorporated into the
'l'0iscussion on a similarly complex ventral secondary intersegmental relationship proposed by Snodgrass
is omitted.
BThe term proposed by Chadwick is confusing, since the Iaterosteroite has been in use for a part of
the pleuron.
COXOPODITE
PREISCHIOPODITE
ISCHIOPODI
A
PRECOXOPODITE
EPIPODITE
D
8
EPIPODITE
EXOPODITE
ENDOPODITE
25
Fm. 8. A-C, Basal part of right maxilliped of Anaspides tasmaniae Thompson, posterior view.
A, from Hansen; B, from Snodgrass; C, from Barlet and Carpentier; D, third left pereiopod
and adjoining part of tergum in Anaspides tasmaniae Thompson (from Snodgrass).
lateral wall and fuses with the tergum, and here the coxa is the definitive pleuron, or
epimeron in carcinology, and not the laterotergite. Snodgrass (1958) subsequently
showed that the corresponding plates in amphipods are freely suspended from the tergal
margins, and stated, "if these plates on the leg bases are truly coxal (as Gruner claims
them to be), there is no part of the skeleton in isopods and amp hi pods intervening
between the tergum and coxa that can be regarded as pleural." Snodgrass (1952, 1958)
thus denied the presence, in the Crustacea, of a pleuron homologizable with the subcoxa.
Carpentier and Barlet's ( 1959) detailed comparative study of the leg base of the
Malacostraca and the Apterygota, however, has shown that the lateral wall is divided into
two parts. They showed that, in the maxilliped of Anaspides (Fig. SA, B), what
Hansen (1925) and Snodgrass (1952) thought was the coxopodite (coxa) was actually a
segment dorsal to the true but reduced coxa; and Carpentier and Barlet called it the
precoxopodite (Fig. SC). What Gruner (1954) called the coxa, therefore, may also
be the precoxopodite.
The validity of Carpentier and Barlet's identification of the reduced coxopodite is
also seen, however, by comparing Fig. 9A with Fig. 39A. Both in Penaeus (Mala-
costraca) and Petrobius (Thysanura) the coxa has three points of articulation with the
precoxopodite (pleuron) and the sternum. The anterior articulation A in Penaeus
(Fig. 9A) corresponds to the trochantinocoxal articulation A' in Petrobius (Fig. 39A);
the posterior articulation B in Penaeus corresponds to the katapleurocoxal articulation C
in Petrobius; and the proximal coxal margin has the articulation C with the coxa that
corresponds to the base of the endosternal process f in Petrobius. Furthermore,
'Carpentier and Barlet (1959) thought A was the pleural apodeme, but it actually is the trochantinal condyle
as discussed on p. 106. '
FURCA
COXA
B
APPENDAGE
STERNUM
MEMBRANA VENTRALIS
FIG. 9. A, basal part of 3rd right pereiopod of Penaeus caramote Risso, seen from inside
(from Bar let and Carpentier). B and C, transverse section of a segment in embryo of
Sco/opendra; C represents a later stage (from Heymons).
Carpentier and Barlet ( 1959) showed in their figs. 4 and 5 most of the endosternal
skeletons in Penaeus (Fig. 9A), and they are apparently homologous with those in the
Apterygota. Based on these findings Carpentier and Barlet concluded that in these
malacostracans the precoxopodite is morphologically equivalent to the katapleuron in the
Apterygota; the laterotergite, with which the precoxopodite articulates, is therefore
homologous with the anapleuron in the Apterygota.
The next question is whether or not the precoxopodite in the Crustacea corresponds
to the subcoxa in insects. Carpentier and Bar!et's study now leads us to suspect that the
separation of the coxa and the precoxopodite can possibly occur during development.
Whatever the developmental origin of the precoxopodite may be, the area dorsal to the
MATSUDA: TI!B INSECT TI!ORAX
EUPLEURON
PROCQXA
.,.., '
' \
-o------"'-"EUCOXA
....
,..,
<=
TELOPODITE
8
><
>
<
w
c
i 1"'
<C =>
>< -
0 c
'-' c

/
=>
w
i
PROTOPLEURON
27
FIG. 10. Verhoeff's (1905) theory on the division of pleuron and coxa (from Verhoeff).
true coxa, i.e., the precoxopodite, corresponds by its location to the subcoxa in insects.
The difference is that in the Crustacea the precoxopodite is not imbedded onto the lateral
wall of the thorax in the way the subcoxa becomes imbedded in insects.
Regarding the origin of the pleuron in the Chilopoda, Heymons (1898) thought the
Pleuren, or B.asalglied, represented a fusion ( Verwachsung) of the coxa with the appen-
dicular segment lying dorsal to it. Further, in Scolopendra Heymons ( 1901) found
that both the tergum and the sternum were tripartite; the. tergum has a broad median
membrana dorsalis, and similarly the sternum has a broad membrana ventralis. The
appendage rudiment arises from the area between the lateral parts of the tergum and
the sternum (Fig. 98, C). A space is formed above the base of the appendage rudiment
and below the dorsally shifted lateral part of the tergum; this newly formed space is the
pleuron. The lateral part of the tripartite tergum shifts even further dorsally, and with
the median tergum forms the definitive tergum. Similarly, the lateral parts of the
sternum and the median sternum form the definitive sternum. The original tripartite
nature of the tergum and the sternum is still recognizable in the adult.
Verhoetf's (1905) study on the development of the coxopleuron in Lithobius essen-
tially agreed with the study of Heymons ( 1901) on Scolopendra. Verhoetf found, how-
ever, that the lateral appendicular bud eventually divides into the coxa and the pleuron.
According to Verhoeff, the coxa and the pleuron are formed from that part of his
pleuropodite that is dorsal to the telopodite (Fig. 10), of which the more dorsal mem-
branous area carrying the dorsal sclerites and the stigma (spiracle) is the eupleuron.
The definitive coxa (pleurocoxa) consists of four sclerites: the coxopleuron, the procoxa,
the metacoxa, and the eucoxa. Of these, Verhoeff equated the coxopleuron and the
procoxa to the subcoxa of Heymons. Thus, as Verhoeff emphatically stated, these sub-
coxal plates are, unlike the subcoxa in insects, not the plates that are separated off from
the coxa. In Hanseniella Tiegs ( 1940) found that the pleural sclerite lying dorsal to
the coxa was only incompletely separated from it, and he thought this was an indication of
the presence of a subcoxal segment.
For the Insecta, the subcoxal theory is now supported by four pieces of embryo-
logical evidence. Heymons ( 1899) first discovered in the embryo of Heteroptera that
the coxa becomes divided into two parts, and that its basal part, the subcoxa (Fig. 110),
eventually becomes a part of the pleuron. As in Scolopendra, the appendicular rudiment
lies between the rudiments of the tergum and the sternum. Hymons thought that only
the lower portion of the definitive pleuron immediately above the coxa in Naucoris and
Nepa (Fig. llH) corresponded to the embryonic subcoxa. Furthermore, in the young
nymph of Nepa (Fig. llH) Heymons showed a narrow pleuron beneath the paratergum.
Roonwal (1937) found in Locusta that, in the 100-hour stage, each leg consists of
subcoxa + coxa + trochanter, femur, tibia, tarsus, and terminal joint. By the 120-hour
stage, the basal joint divides into two segments, the proximal segment representing the
28
subcoxa. The entire subcoxa is converted into the pleuron and the basal . articular
sclerite of the thoracic legs. The tripartite sternum and two lateral tergal scierites are
present in Locusta, as in Heteroptera. Roonwal did not trace just how far the. subcoxa
takes part in the formation of the definitive pleuron in later stages of development. He
simply said, "It will thus be seen that the entire subcoxa is converted into the pleuron
and the basal articular sclerites of the thoracic legs." In Tachycines Ibrahim (1958)
noted the presence of the subcoxa." Furthermore, Bretfeld (1963), in three genera of
Collembola, described and illustrated the development of the two-segmented subcoxa.
11
Thus, the evidence in favor of the theory of the subcoxal origin of the pleuron continues
to grow, as far as the class Insecta is concerned.
The above reexamination of the more direct evidence clearly shows that the
definitive pleuron is different in its developmental process in different groups of the
Arthropoda, and that it is only in the Insecta that the pleuron or a part of the pleuron is
definitely known to be derived by separation from the dorsal part of the coxa (subcoxa),
which becomes imbedded. Therefore, it is to this specialized developmental process for
the production of the definitive pleuron in the Insecta that the theory of subcoxal origin
properly applies. Assuming that the pleural parts throughout the Crustacea-Chilopoda-
Insecta are homologous, as some scattered evidence indicates, the different developmental
processes producing them represent a case of mu!tiphased morphogenesis in evolution.
In the presumed evolutionary sequence through the Crustacea-Chilopoda-Insecta
the lateral wall of the thorax has probably become enlarged in the Chilopoda-Insecta.
This dorsoventrally enlarged thorax thus paved the way for eventual accommodation of
the powerful dorsoventral flight muscles in insects.
The subcoxa and the pleuron in insects: The extent to which the subcoxal element
is present on the definitive pleural wall must vary considerably in differept insects. As
already noted, Heymons' ( 1899) study on Heteroptera clearly showed that the subcoxa
lies mainly on the ventral part of the lateral wall of the thorax, and that the pleuron lies
lateral to it. Crampton (1909), however, pointed out that what Heymons called the
pleurite was actually the mesothoracic epimeron. He reasoned, further, that the meso-
thoracic episternum and the epimeron, which together form the subcoxa of Heymons in
N epa, were actually the mesothoracic episternum, the laterale, and perhaps also the
trochantin. Grandi (1950) also pointed out in her study of the nymphs of Plecoptera
that the epimeral area remains membranous while the episternal area is sclerotized, and
she suggested that the epimeral area might not be subcoxal in origin. In Heteroptera
and the nymphal Plecoptera the epimeral area occupies the dorsal part of the definitive
pleuron. It is logically to be expected, therefore, that the detached subcoxa is largely
incorporated into the ventrally located episternal area. In the pseudofoetus of Dilta
Barlet ( 1950b) observed that the anapleural area is membranous. He therefore assigned
a subcoxal origin only to the katapleuron, which is sclerotized.
All the above-mentioned facts clearly suggest that the definitive pleuron in the
Thysanura-Pterygota consists of the ventral subcoxal area and the dorsal pleural area,
which have become indistinguishable through the secondary sclerotization. Therefore,
Gouin's ( 1959) concept of the podopleuron ( = precoxopodite), which consists of the
anapleuron,
12
the katapleuron, and the trochantin, may be more properly defined as
consisting of the trochantin and the katapleuron alone, as Barlet ( 1950b) and Carpentier
and Barlet ( 1959) believed. It should be recalled, however, that in Collembola the
embryonic subcoxa is two-segmented. It is possible, therefore, that the relatively large
anapleural and katapleural plates in this order owe their origin to this two-segmented
subcoxa.
As Matsuda ( 1963a) pointed out, and as discussed again elsewhere (p. 68) in this
work, the subalare is tergal in origin. In the late embryos of Leucophaea surinamensis
10
Dr. G. Mickoleit of the University of Ttibingen informed me in private communication that he observed the
subcoxa during the course of Dr. Ibrahim's study.
UBretfeld did not trace the development of this peculiar subcoxa further. It remains to be seen if the claimed
subcoxa is really comparable with the subcoxa in other insects.
12Qouin's anapleuron does not include the preepisternum in Perla, as discussed on p, 40.
MATSUDA: TilE INSECT TiiDRAX 29
and in the early stage nymph of Locusta migratoria manilensis, Maki ( 1938) found that
the subalar-coxal muscle (t-cx 8) was dorsally attached to the lateral area of the tergum
or to the ventral surface of the wing base, and this area separates off as the subalare
during the final stage of nymphal development. Similar detachment of the subalar-coxal
muscle from the lateral tergal area has been observed, or can be surmised, in several
other orders (p. 68).
Roonwal ( 1937) showed, in the embryo of Locust a, that the sternum is divisible
into a median portion (the primary mediosternite) and two lateral portions (the primary
laterosternites), and he reported that the latter bore appendages at its lateral edge. He
asserts that his primary laterosternite is not the same as the laterosternite as understood
by morphologists." Since the ventropleurite presumably forms the ventral process for
articulation with the coxa in holometabolous insects, it is possible that the laterosternite
of Roonwal and the ventropleurite of the present work are the same. If this interpreta-
tion and the observation of Roonwal are correct, the ventropleurite, which is described as
a part of the katapleuron in this work, is actually sternal and not subcoxal in origin.
As will be discussed later, the katapleural ventropleurite lies between the inner coxal
base and the furcal base in the adult, and according to Weber (1928a) the furca arises
from the area that is both sternal and subcoxal in origin. On the basis of these observa-
tions in adult insects, therefore, the ventropleurite may be subcoxal in origin. It is
possible that in Locusta, where the subcoxa and sternum are indistinguishably fused in
the adult, and the primary border between the subcoxa and the sternum may be
obliterated even in the embryo. Roonwal's finding, therefore, needs to be supported by
similar studies on other insects, if the sternal origin of the ventropleurite is to be fully
accepted.
In higher holometabolous insects the pleuron does not arise from the subcoxa. For
instance, in Drosophila it has been known since the work of Chen ( 1929) that the thorax
and the appendages of adult Drosophila are represented, at the end of the larval period,
by six primordia, or imaginal discs; Geigy's (1931) experimental work indicated the
presence of imaginal cells of legs in the embryo of Drosophila. These embryonic and
larval anlagen of the imaginal legs and thorax, however, cannot be related to the
embryonic and nymphal subcoxa and coxa in hemimetabolous insects. Clearly, here the
anlagen replacement (or substitution) in ontogeny is involved in the production of
homologous adult structures."
Theories: In the past, many authors concerned with the morphology of the pleuron
of arthropods did not pay close attention to the difference in the developmental origin of
the pleuron in different groups of the Arthropoda, and they homologized the pleural
parts with the presumed subcoxal counterparts of other groups. Thus, Borner (1903,
1904), in his study of the comparative anatomy of various groups of arthropods, found
that the segment equivalent to the subcoxa of Heymons ( 1899) is present also in the
Crustacea and Chilopoda, but is not present, or is difficult to find, in the Chelicerata, the
Pantopoda, and the Progoneata; as a result, he agreed with Hansen ( 1893), who thought
there was a segment proximal to the coxa in the Crustacea and Insecta. Borner ( 1904)
proposed the concept of the basipodite, which consists of the subcoxa and the coxa.
Verhoeff (1905) criticized the concept of the basipodite, because the concept of the coxa
composing it fluctuates in different groups. Borner (1914) said that the subcoxa as an
independent sclerite was present in the Pantopoda and in many of the Malacostraca, and
in the Chelicerata and Hexapoda the subcoxal pleuron could split into smaller sclerites,
which are the epimeron, the episternum, and the trochantin. Thus, according to Borner,
nearly the whole lateral wall of the thorax is subcoxal in origin. Ewing ( 1928) applied
the subcoxal theory to the Arachnida.
"In insect morphology, the term laterosternite has been usually applied to the ventropleurite (e.g., Weber 1933).
"In Lepidoptera Kim (1959), Kuske et al. (1961), and Kuske (1963) found that the pupal leg, hence the
imaginal leg, is formed from the epidermal cells of the larval leg, no imaginal disc of the leg being present.
Amvng holometabolous insects themselves, therefore, the developmental process of the leg is not the same.
The theory of the subcoxal origin of the pleuron elaborated by Snodgrass and Weber
has had considerable influence upon later workers. According to Snodgrass ( 1927,
Fig. llA-F), the subcoxa was primitively a single plate surrounding the coxal base and
articulating with the latter by anterior and posterior hinges, from which later the anterior
trochantinal and posterior pleural articulations were derived; it was the ventral ring of the
subcoxa (trochantin) that bore the condyles that articulate with the coxa. With the
acquisition of the pleural ridge associated with the development of the wing, the dorsal or
anapleural ring split up into a few sclerites; the anterior sclerite formed the precoxal
bridge, and the posterior one formed the postcoxal bridge. Snodgrass implicitly applied
this theory of the subcoxal origin of the pleuron to other arthropods.
Weber ( 1928a) supplemented Snodgrass' theory by studying the ventral area of the
subcoxa and its relationship to the sternum in the insect thorax. He proposed that the
preepisternum (priicoxale Briicke) and the ventropleurite (Laterosternit) are the ventral
limit of the subcoxa that came in contact with the sternum. In the theory of Snodgrass
and Weber, no distinction was made between the primary pleuron and the subcoxal area.
It was implicit that the whole of the definitive pleuron was subcoxal in origin.
Ferris and his students applied the subcoxal theory in their series of studies of the
insect thorax. They also accepted implicitly the idea that the whole pleural wall was of
subcoxal origin. The basic new concepts they introduced were summarized in Ferris'
paper (1940b) entitled "The Myth of the Thoracic Sternites of Insects", in which it was
postulated that primitively the subcoxae met on the ventral median longitudinal axis of
the thorax, as seen in Neuroptera and many other holometabolous insects; this longi-
tudinal line of demarcation was called the discrimen. According to Ferris, then, the
exposed sternum in hemimetabolous insects has been produced by the retreat laterad of
the subcoxal element on each side. He also pointed out the usefulness of the paracoxal
suture (the pleural costa of Ferris) and the anapleural suture as landmarks in homolo-
gizing pleural parts.
Weber ( 1952) accepted Ferris' idea of the secondary nature of the sternum and
Carpentier's (194 7) idea of the katapleural origin of the furca, and proposed a more
elaborate theory of the production of the secondary sternum in phylogeny (Fig. 12).
Since, according to Weber ( 1952), the furca remains in the same relative position, viz.,
on the posterior inner margin of the katapleural ring (Fig. 12) during the retreat laterad
of the pleuron in evolution, the formation of the definitive sternum between the furcae is
dependent on the degrees of migration laterad of the katapleural ring on both sides.
Since the presence of the sternum as an integral part of the thorax is so evident in
any insect embryo, the idea of the secondary nature of the sternum in the theories of
Ferris and Weber was unfavorably criticized by Matsuda (1956a), Czihak (1956), and
Gouin (1959). Furthermore, as will be detailed later, more recent works on the Aptery-
gota by Carpentier and Barlet have shown that the furca does not necessarily lie on the
pleuron. Earlier, Hansen ( 1930) criticized Snodgrass' diagrams (Figs. llA-F) and
claimed that they were unrealistic, but the facts upon which he based his criticism were
mainly specialized features, and therefore largely not pertinent; in addition, he ignored
all embryological evidence.
Snodgrass ( 1952, 1958) criticized the subcoxal theory on which he had earlier
elaborated (1927, 1935). He attempted to show quite legitimately that the evidence in
favor of the subcoxal theory had been deduced mainly from a study of chilopods and
insects, and that mistaken significance had been attached to Heymons' ( 1899) finding of
the subcoxal origin of the pleuron in some insects, which led many authors including
Snodgrass himself ( 1927) to apply erroneously the concept of the subcoxal origin of the
pleuron to other arthropods. Snodgrass' ( 1952, 1958) conclusion was essentially the
!\arne as the one reached in this work.
In annelids the parapodium is a simple outgrowth of the pleural wall. Therefore,
Bekker ( 1923, 1924, 1960) thought that in arthropods also the same structural plan
must be maintained. Thus, what Heymons ( 1901) called the lateral tergum and the
TERGUM
SUBCOXA
TROCHANTIN
ANAPLEURITE
TROCHANTIN EUPLEURON
A
8 c D
ANAPLEURITE TROCHANTIN
\
EPISTERNUM PLEURAL SUTURE
\
EPIMERON
TROCHANTIN
E F
SUBCOXA
STERNUM
G
H
PLEURON
FIG. 11. A-F, diagram showing origin and evolution of pleural sclerites from a subcoxal
segment; a and b represent anterior and posterior coxal articulations (from Snodgrass); G,
right middle leg of embryo of Naucoris (from Heymons); H, last two thoracic segments and
abdomen in nymphal Nepa cinerea (from Heymons).
32
ANAPLEURON
MEMOIRS OF THE ENTOMOLOGICAL SOCIE1Y OF CANADA
KATAPLEURON
--H--.;
B
RCA
ANAPLEURON
EURON
I RCAL @1 ..
FIG. 12. Weber's (1952) theory on the origin of secondary sternum (from Weber).
lateral sternum in Scolopendra (Fig. 9B, C) are, according to Bekker, actually the
pleural bases of the leg. The definitive tergum and the sternum are formed secondarily
from the pleuron (tergite rudiment and sternite rudiment) by the median
true tergal and sternal parts. Therefore, what Heymons (1899) discovered in Heterop-
tera can best be explained as a process of formation of the coxal segment at the expense
of the pleuron, not as a separation of a basal part of the coxa, and this would be con-
sistent with the major evolutionary trends of the leg base in the Articulata.
The novelty of Bekker's theory is refreshing indeed. However, it leaves the
unanswered question of whether the kind of tripartite tergum and sternum that Heymons
( 1901) found in Scolopendra, in which the median membranous area is so broad, repre-
sents a primitive condition or not. Heymons attributed the broad membrana dorsalis to
the presence of the abundant yolk. In insect embryos Heymons (1895, 1899) and
Roonwal ( 1937) found that the tergum consists of two lateral plates that later become
fused. In insects, therefore, it would appear to be tripartite when the median area
between the two lateral plates is broad, and bipartite when the median area is very
narrow. Therefore, to regard the lateral tergum and the lateral sternite in the tripartite
tergum and sternum as the pleuron is quite arbitrary. If Bekker's theory is true most
insects have completely or nearly completely lost the primary tergum.
Matsuda ( 1963a) defended the subcoxal theory, pointing out the facts presented by
Snodgrass (1952), Gruner (1954), and Carpentier and Barlet (1959) which indicated
the presence of the basal part of the leg, or the part homologous with it, in the pleural
wall in the Chilopoda and Crustacea. Here again, however, the application of the sub-
coxal theory is not clear. He simply attempted to homologize the pleuron throughout
the Crustacea, Chilopoda, and Insecta, without giving a strict definition of the subcoxal
theory.
Fliller's ( 1963a, b) opposition to the subcoxal theory was directed mainly against
those applications of it by Ferris (1940b) and by Weber (1952) that proposed the
secondary nature of the sternum. Fiiller was able to show, in major groups of the
Chilopoda, the presence of a well-formed and distinct sternum that could hardly be
secondary. Fiiller observed that in the Chilopoda the presence of numerous small
sclerites on the pleural wall was a primitive character, and that the membranous pleural
wall was a specialization. Therefore, the sclerites on the pleural wall in Chilopoda are,
MATSUDA: TifE INSECT TifORAX 33
according to Fiiller ( 1963b), not homologous with the pleural sclerites in insects which
are of subcoxal origin.
It is clear from the foregoing review of various subcoxal theories that workers,
including the present writer, have tended to indulge in speculations, extending and
modifying the subcoxal theory, without really looking for reliable developmental facts in
support. The disputes have obviously stemmed from the preconception that one and the
same developmental mode of the pleuron prevails throughout the Arthropoda. Indeed,
the subcoxal theory, if it must be preserved, must be admitted to apply to hemimetabolous
insects and to some lower holometabolous insects, in which the embryonic subcoxa
develops continuously throughout all the postembryonic stages.
HOMOLOGIES OF THE PLEURAL PARTS
The main features that have emerged from the studies of the pleuron in Thysanura
and Diplura by Carpentier and Barlet are: the anapleuron extends as far ventrally as the
lateral margin of the basisternum; the katapleuron passes ventrally over the narrow space
between the furcal base and the inner coxal margin; the trochantin (s. str.) has two or
three points of articulation with the coxal margin and is part of the katapletiron; the
homologue of ~ t h e pleural arm (processus pleural), when present, demarcates the
anapleuron from the katapleuron. These features and a few others that will be pointed
out as occasions arise must be borne in mind before homologizations of the pleural parts
between the Apterygola and the Pterygota can be made.
Carpentier (1955) attempted to homologize the pleural parts in Ctenolepisma and
the last nymphal instar of Blatta (Fig. 13). Homologies of parts are obvious from the
designations of areas on the figure. A conspicuous difference is the presence, in Blatta,
of the pleural ridge bearing the pleural arm. The arm has a wide base, and its dorsal
limit, in the specimen examined, lies on the pleural ridge roughly where the dorsal margin
of the katapleuron meets it." In Blatta the dorsal margin of the katapleuron, or the
paracoxal suture, does not extend into the epimeron, although it does in Carpentier's
original figure, as reproduced in Fig. 13B.
In Ctenolepisma Carpentier (1955) did not refer to the line here labeled the
anapleural suture (Fig. 13A). This line represents the ventral limit of the sclerotized
part of the anapleuron, below which the anapleuron becomes membranous.' The line
roughly corresponds in position to the anapleural suture in most pterygotes, although in
Blattaria it is only an inconspicuous suture and absent in the last instar nymph of Blatta
orienta/is (Fig. 13B).
The proximal (ventral) end of the anapleuron in Ctenolepisma is broad and meets
nearly the whole lateral margin of the basisternum. In Blatta the proximal part of the
anapleuron, or the preepisternum, is reduced, although in some other lower pterygote
insects the preepisternum is broad proximally and meets the l"!tteral margin of the
basisternum.
The above discussion clearly shows that homologies of many pleural parts between
Ctenolepisma and Blatta can be established with reasonable certainty. A discussion on
some other primitive pleural structures in the Pterygota and their evolutionary fate
follows.
The primitive pleuron and its modifications in the Pterygota (Fig. 14) : The overall
evolutionary tendency of the pleural wall in the pterothorax has been toward increasing
sclerotization. This is evident from the fact that, in Thysanura, the pleural wall is not
so solid as in most of the Pterygota. Filller ( 1963a, b) found that, in the higher
Chilopoda, there is a tendency for the pleuron to be more membranous; in these related
groups the pleural wall is definitely softer than in most pterygotes.
The hardening of the pleural wall of the pterothorax has evidently been associated
with the acquisition of wings. Yet, in more primitive pterygote insects that have rela-
16In Fig. 13B the pleural arm was redrawn after actual examination of the last instar nymph of Blatta orienta/is.
1
6This was found in a reexamination of the specimen used by Prof. Carpentier.
PARACOXAL SUTURE ---T-.,......,.----...:--'-':;--:;:;/
PREEPISTERNUM ----r-
TROCHANTIN
h----4
BASISTERNUM ---
KATEPIST ERNUM
(ANTE COXAL E)
FURCASTLI\111UIII
ANTECOXALE
(KATEPISTERN
FURCA(f
A
PARACOXAL
SUTURE
B
A
COXAL PROCESS
PARACOXAL
SUTURE
ANEPIMERON
PLEURAL ARM

TROCHANTIN
A
8
FIG. 13. A, right mesothoracic pleuron in Ctenolepisma ciliata; B, same of Blatta orientalis
(from Carpentier).
tively weak flight ability, the sclerotization has not advanced so far as to obliterate the
original sclerites that presumably surrounded the coxal base on the pleural wall, as is
evident in the foregoing homologization of pleural parts between Ctenolepisma and
Blatta. It can be reasonably assumed, therefore, that the sutures separating the sclerites
were primitively membranous.
The internally ridged pleural suture was probably formed between the median
articulation of the trochantin B in Thysanura (Fig. 38A) and the dorsal margin of the
pleuron, as Carpentier (1955) and Matsuda ( 1963a) have suggested. The course of
this suture, from its pieurocoxal articulation to the dorsal pleural wing process, was
probably nearly vertical primitively, as is evidenced by the occurrence of a similar pleural
suture in some lower insects, such as Neuroptera. The most common specialization of
the pleural suture is its tendency to run obliquely forward. In some higher forms (e.g.,
Heteroptera) the suture has often become obliterated. It is well retained, however, in
wingless orders, such as Mallophaga, Grylloblattodea, and Aphaniptera.
MATSUDA: THE INSECT THORAX
SUBALARE
EPIST. LONGIT.
SUTURE
fBASALAR CLEFT)

- WING PROCESS
KATEPISTERNUM - (
----
BASISTERNUM
TRANSEPIMERAL SUTURE
__.L!=---+f___:::::::,..,._-------"7+--p ARACOXAL SUTURE
o---INTERPLEURITE
----J--,f--i"t-----KATEPIMERON
VENTROPLEURITE FURCASTERNUM
FIG. 14. Diagram showing sutures on pterothoracic pleuron (original).
35
The pleural arm arising from the internal pleural ridge is usually connected with the
sternal furca by a muscle, or at times both structures are firmly connected by a tendinous
structure. Externally, the point of origin of the pleural arm varies considerably in
different orders. In Dermaptera, Neuroptera, Plecoptera, and others it lies near the
middle of the ridge; the location of the pleurfil arm in these orders roughly corresponds
to the processus pleural of Carpentier and Barlet in Thysanura, which demarcates the
anapleuron from the katapleuron. In Blattaria, Mantodea, Phasmida, and Orthoptera,
the arm lies more ventrad, near the pleural condyle that articulates with the coxa. In
Coleoptera the apparent pleural arm, arising from near the dorsal margin of the pleuron,
may not be homologous with the pleural arm in other insects. In Hymenoptera and
Psocoptera the apparent pleural arm does not arise from the pleural ridge.
The paracoxal suture (Fig. 14) is the suture that separates the pleural wall into the
anapleural and katapleural rings. It is synonymous with the pleural costa (Ferris, etc.)
and the precoxal suture (Michener, Matsuda, etc.). It sometimes extends into the
epimeral area, and is primitively a complete suture, present both on the episternum and
on the epimeron, as seen in Thysanura (Fig. 13A) and some pterygotes, which are
discussed below. A more inclusive term, the paracoxal suture, is therefore proposed here
to replace the precoxal suture, which implies that the suture lies only anterior to the coxa,
and the pleural costa which is not specific in location.
Ferris was right in emphasizing the importance' of this suture as a landmark dividing
the pleuron into the dorsal anapleuron and the ventral katapleuron, although he and his
students did not always identify the suture correctly. In some pterygotes the paracoxal
suture crosses the pleural suture, at or near the point where it gives rise to the arm
internally.
17
Thus, in Blatta (Fig. 13B), Allostethus (Fig. 79C), Diapheromera,
Micropteryx (Fig. 147A), Merope (Fig. 122), Sialis (Fig. 120A), and Eucorethra
(Fig. 135) the paracoxal suture, after passing the pleural pit, runs caudad, thereby
producing a rather narrow katepimeron behind the pleural suture. It should be pointed
out here that the genera in which a complete paracoxal suture occurs are all primitive
pterygotes; and this fact indicates that the paracoxal suture is an archaic suture.
UThe point of invagination of the pleur-al arm can often be recognized as a pit.
The paracoxal suture is often retained only on the episternum in, many pterygotes,
such as Neuroptera, Trichoptera, some Lepidoptera, some Homoptera, some Plecoptera,
and others. In such cases the anapleural suture often meets distally the lateral end of
the paracoxal suture. In some Coleoptera (Fig. 870) and Zoraptera (Fig. 90C) the
paracoxal suture is retained on the epimeron.
In some Neuroptera (Fig. 120B), Hymenoptera (Fig. 161), Homoptera (Fig. 109A)
and others, the transepimeral suture (Fig. 14) is continuous from the paracoxal suture
on the episternum, and it gives a spurious impression of being the epimeral part of the
paracoxal suture. This suture is, however, definitely different from the latter, since its
course is horizontal and reaches the border between the lateral postnotum and the
epimeron. In some Diptera the transepimeral suture is continuous from the anapleural
suture.
The anapleural suture (Fig. 14) separates the episternal part of the anapleuron into
the dorsal anepisternum and the ventral preepisternum. This suture is very persistent
and well marked in holometabo!ous insects. Among hemimetabolous insects, this suture
may be inconspicuous, but it is found in Plecoptera, Blattaria, some Orthoptera
(Gryllidae), lsoptera, Embioptera, Mantodea, Phasmida, Derrnaptera, etc., as shown in
the discussion of each order. In Psocoptera, Thysanoptera, and Homoptera the ana-
pleural suture tends to lie more ventrad than in other orders. In spite of its prevalence,
DuPorte (1965) denies that the suture is primitive, and synonymizes it with his coxo-
sternal suture, which is discussed more fully later (p. 39).
As Ferris' earlier term for this anapleural suture, the pleural cleft, suggests, it is
often a membranous cleft in lower forms, but it may be an internally ridged suture in
higher forms. The anapleural suture arises proximally (ventrally) on the anterior
margin of the anapleuron, near and usually slightly behind the spiracle. As already
mentioned, its lateral end tends to merge with the paracoxal suture. In .Ctenolepisma
(Thysanura), as already discussed (p. 33), the ana pleural suture may be present.
In lower orders the basalare is a small detached anepisternal part immediately
anterior to the pleural wing process. However, the basalare is often represented by two
pieces of sclerites, as seen in Orthoptera, Lepidoptera, Diptera, and others. They are
called the anterior and the posterior, or the first and second basalaria; the presence of
this additional (anterior) basal are must be a specialized condition. The basalare is
sometimes not detached, and an internal apophysis arises from the .basa!are area as seen
in Neuroptera," or it fuses with the pleural wing process as in Coleoptera. In Odonata
a cap-shaped tendon occurs in a corresponding area, and it is connected to the humeral
plate by a ligament; it is not certain that this cap-shaped structure is homologous with
the basalare. In some orders such as Plecoptera, Trichoptera, some Lepidoptera, and
others, the prolonged prealar arm extends ventrally and attaches to the anterior dorsal
margin of the anepisternum, thereby forming a prealar bridge.
Often the area surrounding the basalare is membranized (incisura basalare of
La Greca, 1949a) and this membranized area sometimes extends posteriorly to form a
narrow cleft (e.g., in mantids). In the mesothorax of Dermaptera (Fig. 79A), the cleft is
apparently represented by a suture that meets the anapleural suture and the paracoxal
suture posteriorly. Matsuda ( 1960a, 1963a) called this suture in Dermaptera the basalar
suture, because it apparently corresponds to the basalar cleft in the metathorax of
Dermaptera. Since, however, this homology is somewhat uncertain, a new, more neutral
term the episternal longitudinal suture is proposed; the suture occurs also in Orthoptera
and Mantodea.
In Homoptera, in which the anapleural suture lies more ventrad, an additional
transverse suture (the transanepisternal suture) divides the anepisternum into dorsal and
ventral parts (Fig. 106B). The ventral part is the lateropleurite of Weber ( 1928b),
which differs, however, from the lateropleurite of Crampton (Fig. 15).
18Matsuda (1960a. 1963a) homologized this apophysis with a pleural process pc in Lepisma (Fig. 37), which is,
however, an anapleurokatapleural structure.
:n
The preepisternum forms the precoxal bridge by union with the basisternum. The
line of ljnion with the basisternum, when it is present, is genuinely the pleurosternal
suture, as can be seen in apterygotes and many hemimetabolous insects; the suture may
be represented by a membrane, as seen in Ephemeroptera, Blattaria, Isoptera, Gryllo-
blattodea, and some others. In some Plecoptera, the pleurosternal suture is seen as a
faint line (Fig. 48).
In holometabolous insects, however, the ventral limit of the preepisternum has been
controversial, because of its fusion with the basisternum, which invaginates on the median
longitudinal axis. Ferris (1939-1950) consistently maintained that the preepisternal
area had invaded ventrally from both sides to meet along the median longitudinal suture,
which he called the discrimen. It is for this reason, according to Ferris, that the sternum
is absent on the ventral surface of the thorax in holometabolous insects.
As already discussed (p. 21), however, there is no positive reason to believe that a
complete invagination of the basisternal area always occurs in holometabolous insects.
Despite the absence of a distinct pleurosternal suture, therefore, it is expected that often
the basisternal area is undefinably present on either side of the median longitudinal suture
( discrimen) .
In Merope (Fig. 124A), Tipula (Fig. 136A), and Limnephilus an oblique suture
on the median ventral area of the pterothorax appears to represent the pleurosternal
suture. The suture on each side, however, runs anteriorly toward the median longi-
tudinal axis; hence, the presumed basisternal area demarcated by them is narrower
anteriorly, instead of becoming wider anteriorly. When the basisternum is well exposed
in other insects, it is often wider anteriorly. Therefore, the identification of the suture
as the pleurosternal suture in these genera is somewhat uncertain.
In the prothorax of many holometabolous insects, in which the preepisternum is
represented by the lateral cervical sclerite, the association of the preepisternum with the
basisternum is absent. Even in the prothorax of hemimetabolous insects the typical
association of the preepisternum with the basisternum is often obliterated.
The katepisternum includes the trochantin. The antecoxale refers to the katepi-
sternal area between the paracoxal suture an9- the anterior margin of the trochantin, as
typically seen in Ctenolepisma (Fig. 13A) and Blattaria (Figs. 13B, 57B).
Primitively, the trochantin was probably bipartite, divided by an oblique line, of
which the anterior part was connected with the coxa, as Crampton and Rasey (1915)
maintained; the occurrence of this kind of trochantin in Lepismatidae, Dermaptera,
Isoptera, Blattaria, and others supports such an interpretation. In many other orders
the dividing line is lost.
In Ctenolepisma (Fig. 38A) the dorsal margin of the trochantin articulates with the
katapleuron; in the Pterygota this articulation has become lost, along with the loss of the
posterior articulation of the trochantin with the coxa that is present in Thysanura ( C in
Figs. 37; 38A, B; 39A).
In some lower pterygotes (e.g., Dermaptera, Blattaria) the distal margin of the
trochantin posteriorly almost reaches the pleural condyle for the coxal articulation. In
such cases, the distal margin of the trochantin may be confused with the paracoxal suture.
In Micropteryx (Fig. 147A), for instance, the paracoxal suture is bent sharply caudad
near the pleural suture and appears to be the distal margin of the trochantin. However,
its extension into the epimeral area helps identify it as the paracoxal suture. In the
prothorax of Allostethus (Fig. 79C) and nymphal Blatta (Fig. 13B) the two sutures
occur together; and it is clear that the paracoxal suture extends beyond the distal margin
of the trochantin.
In some insects (e.g., Eucorethra, Fig. 135; Sialis, Fig. 120A) the basal (distal')
part of the trochantin is fused with the antecoxale and the distal margin of the trochantin
is lost. When such a partial fusion of the trochantin with the rest of the katepisternum
occurs, the enlarged, definitive trochantin is called the eutrochantin. Often in higher
insects differentiation of the trochantin is lost, owing to increased sclerotization of the
katepisternum as a whole. The trochantin is absent also in Odonata and Ephemeroptera.
An isolated, undivided trochantin occurs in Heteroptera, some Homoptera, Coleoptera
(usually reduced), and in some Hymenoptera.
The ventropleurite, which may be sternal in origin (p. 29), is sometimes called the
laterosternite; it is the area between the furcal base and the inner margin of the coxal
base. Weber ( 1928a) first established the pleural origin of this area for the pterygotes.
In Thysanura (Figs. 38A, 40) and Diplura (Fig. 33) the area is a continuation of the
](_atapleural ring of the subcoxa. In other words, the katapleural ring extends ventra-
posteriorly the area between the endoskeletal process f, which corresponds to the furcal
base in the Pterygota, and the inner margin of the coxal base.
In Perla (Fig. 48) the ventropleurite is continuous with the anapleural pre-
episternum, but this condition must have resulted from the loss of the paracoxal suture.
In Acroneura and in Paragnetina, figured by DuPorte (1965), the paracoxal suture
(sternocostal suture of DuPorte) is continuous ventrally with the anterolateral angle of
the furcasternum as in Ctenolepisma (Fig. 13A), and the katepisternal antecoxale is
continuous with the ventropleurite area, indicating that the latter is a part of the kata-
pleuron. In Cicadidae (Fig. 109A) also the ventropleurite is obviously a katapleural
part. In Embioptera (Fig. 51) the ventropleurite can, upon superficial examination, be
either the anapleural or katapleural sclerite. In hemimetabolous insects, in which the
ventropleurite is present, the paracoxal suture is often incomplete, which obscures the
relationship of the ventropleurite with neighboring areas, including the katapleuron.
In holometabolous insects, the ventropleurite has presumably become the process for
ventral articulation with the coxa; and in some of the more primitive holometabolous
insects the paracoxal suture ventrally reaches the median longitudinal suture of the
invaginated sternum. In such forms, therefore, the association of the ventropleurite
with the katepisternum is clear. In Agulla (Fig. 118B) and Hepialus (Fig. 149A) the
ventropleurite (ventral process) is plainly continuous with the katepisternum. In
Limnephilus and Neuronia (Fig. 143A), in which the anapleural preepisternum is clearly
recognized by its line of union with the basisternum, the ventropleurite is clearly con-
tinuous with the katepisternum, and not with the preepisternum. These facts lead us to
conclude that the ventropleurite is in fact katapleural in origin.
Weber ( 1928a) proposed that the ventropleurite (his Laterosternit) is an anapleural
structure lying external to the trochantinal ring (katapleural ring) of the subcoxa, as can
be seen from his fig. 16b, c. This interpretation resulted from his belief that the ventro-
pleurite continuous from the undivided episternum (into the preepisternum and the
katepisternum) in the wingless female of Aphis jabae represents a primitive condition;
this interpretation ignores the improbability of finding a primitive structure in a highly
evolved group such as Aphis. His confusion resulted partly from his disregard of the
paracoxal suture. Matsuda (1960a) also postulated that the ventropleurite is part of the
anapleural ring, without paying sufficient attention to the paracoxal suture. This idea is
herewith retracted in favor of the katapleural derivation of the ventropleurite.
The epimeron, as already noted, is only rarely divided by the paracoxal suture into
the dorsal anepimeron and the ventral katepimeron. The dorsal margin of the epimeron
is often connected with the lateral extension of the postnotum to form a postalar bridge.
In Hemiptera and Coleoptera the dorsal part of the epimeron is modified for holding
wing veins.
The katepimeron demarcated by theparacoxal suture is small, as seen in Dermaptera
(Fig. 79A, C, D), Zoraptera (Fig. 90B), Coleoptera (Fig. 84), etc. In Odonata
(Fig. 171D) the postcoxal bridge is obviously formed by the katepimeron. The
epimeron is divided by the transepimeral suture into dorsal and ventral parts in some
Neuroptera, Hymenoptera, and Diptera.
The preceding discussion of the pleural parts leads the present author to revise the
proposal he made (1960a, 1963a) for the hypothetical primitive pleuron of the ptero-
thorax. In his new diagram (Fig. 14) the anapleural ring does not encircle the coxal
base ventrally, the ventropleurite is a part of the katapleural ring, and the transepimeral
PRESCUTUM
SURALARE:
PRETERGITE
INTERTERGITE
PARAPSIDE
PARASCUTELLUM
GITE
PTERALE
PREALAR BRIDGE MEDIOPHRAGMITE
TEGULA POSTERIOR BASALARE
NOTOPTERALE
ANTERIOR BASALARE POSTERIOR SUBALARE
EPISTERNUM ANTERIOR SUBALARE
ANEPISTERNUM PTEROPLEURON
HYPOPTERON HYPOEPIMERON
LATEROPLEURITE TROCHANTIN
LATEROSTERNITE
POSTCOXALE
CERVICAL STERNITE
PRESTERNITE BASISTERNUM VERACO POSTFURCASTERNITE
FIG. 15. Ground plan of thoracic parts (from Crampton).
suture is added. It should be noted that the transepimeral suture and the episternal
longitudinal suture are not primitive sutures, but they are the sutures that have developed
in the Pterygota. A few other sutures that rarely occur are not shown in Fig. 14.
Earlier homologizations of pleural parts: Crampton (1914a) proposed a ground
plan for a wing-bearing thoracic segment (Fig. 15). The pleural parts he recognized
agree fairly well with those recognized in this work. As Verhoeff (1902b) did, Crampton
apparently took Dermaptera into serious consideration. Primitively, according to
Crampton (1914b), the epimeron, the episternum, and the lateropleurite formed a single
continuous plate (eupleuron), which later split secondarily into three parts; the latero-
pleurite and the laterosternite together form a precoxal bridge, which connects the
eupleuron and the sternum. Crampton failed to recognize the paracoxal suture on the
epimeron; the transverse suture on the epimeron is the transepimeral suture in this work;
and oblique suture separating the episternum from the lateropleurite is the episternal
longitudinal suture.
DuPorte's (1965) concept of the coxosternum appears to have its source largely in
his misinterpretation of sutures, and in his lack of confidence in the anapleural suture as
a reliable landmark. Thus, in Paragnetina what DuPorte called the sternopleural suture
is, in fact, the anapleural suture. This is immediately clear from a comparison of his
fig. 3 with Fig. 48 in this work or with fig. 2 of the author's earlier work (1960a). It is
obviously this misidentification of the anapleural suture that led him to believe that the
suture was present only in holometabolous insects, and hence could not be a reliable
landmark in homologizing pleural parts.
Since the anapleural suture is the pleurosternal suture for DuPorte, the pre-
episternum is a sternal area; DuPorte called the preepisternum the coxosternum, and
thought that this area has probably been derived from the ventral region of the leg
segment. Thus, according to DuPorte, the leg is probably a sternal structure. The
sternum as thus conceived of is greatly expanded laterally in its anterior half. This idea
of the enlarged sternum has apparently misled him further to interpret the pani.coxal
suture in Paragnetina, Tibicen, and Sthenopis as the sternacosta.
Gouin (1959), in his study of the same order, Plecoptera (Perla), also interpreted
the preepisternum as the lateral part of the basisternum, and his podopleuron did not
include this area. He based his interpretation on an quite ambiguous premise that
muscles generally connect the neighboring segments (tergum, pleuron, sternum) within
a metamere. Thus, dvm 40 and ppm 55 of Wittig ( 1955) in Perla should be considered,
by their dorsal and ventral points of attachment, as a tergosternal muscle and a pleuro-
sternal muscle, respectively; and hence their ventral points of attachment should be the
lateral basisternum.
This interpretation harbors two misconceptions, viz. the muscles generally connect
different segments and the muscle insertions always determine the external areas, to which
the muscles are attached. The fallacy of this interpretation is immediately clear by
comparing the ventral part of the thorax in Perla (Fig. 48) with that of any Thysanura or
Diplura that have been studied (Figs. 34; 38A, B; 40). In none of these apterygotes is
the basisternum so much expanded laterally as Gouin and DuPorte ( 1965) thought it was
in Plecoptera. As Weber ( 1928a, b) clearly understood, the lateral basisternum of
Gouin and DuPorte is the ventral part of the subcoxa, and the muscles dvm 40 and
ppm 55 are t-p 5, 6 and p 3, respectively.
THE WING
The origin of wings: Since Crampton's ( 1916a) work, the theory of the paranotal
origin of wings, viz. the development of the wings from the lateral part of the tergum, has
been universally accepted. More recent studies dealing with the noninsect arthropods,
such as those on the Chilopoda by Bekker ( 1958a) and on oribatid pteromorphs (Acari)
by Woodring (1962), indicated that in these forms the tergum has the potential to
develop wings or winglike structures. Ross' ( 1964) genetical study dealing with the
prothoracic wing found in Blatella germanica also suggested the paranotal origin of the
true wing in the meso- and meta-thorax.
The question of how the paranotal lobes have become wings, however, has been
controversial. Lemche (1942) showed a type of reduced wing (his type 2) which lacked
the wing base structure in the mesothorax of Melanozosteria and others. This type of
wing is separated from the tergum only by a slit; Lemche thought that such a wing, being
comparable with the paranotum in shape and in having rounded impressions on the
surface, is definitely primitive. Further, he thought that this slit formation preceded the
differentiation of the wing base in evolution.
Lemche's type 2, however, appears to correspond to the micropterism produced in
the mesothorax of Loboptera, which was shown by Lefeuvre ( 1965). Five types of
wing development, which Lemche ( 1942) distinguished, are obviously the pterygopoly-
morphism which occurs in many pterygote orders, such as Blattaria, Orthoptera, Hemip-
tera, Diptera, etc., as summarized by La Greca ( 1954). If we follow Lemche's logic,
then, the wing arose many times in higher insects.
Based on the observation of wing deformities in Drosophila, Goldschmidt ( 1945)
thought that the entire wing evolved from the dorsal and ventral lobes of a polychaete
parapodium. Raw ( 1956) limited the origin of the wing to the dorsal lobe of the para-
podium. These theories appear to imply that insects are direct descendants from
polychaetes, since they assume, without giving sufficient anatomical details, direct
homologies of structures between the two remotely related groups.
The prothoracic wing that Ross ( 1964) studied was produced from the prothoracic
paranota in Blatella, and it had nothing to do with the pterygopolymorphism. She com-
pared the venations and associated tracheations of the pro-wing with those in the normal
wing, and postulated that if a dominant mutation, similar to that producing the pro-wing
41
but affecting all thoracic segments," occurred in such an insect, it is conceivable that a
pair of :rigidly extending wings with well-developed venation could be produced on each
of these segments. She went on to say that this potentiality makes it unnecessary to
postulate a series of gliding forms transitional between paranota and wings during the
evolution of insects. Indeed, there is no factual evidence that the paranota have ever
been of special functional importance associated with flight in early insects. As far as
can be seen from the anatomical viewpoint, the paranota were simply the preadapted -
so to speak- lateral swelling of thoracic segments (as seen in Chilopoda), from which
later wings arose, perhaps rather suddenly. In fact, winged insects were already present
during the lower Carboniferous time (Haupt 1941) and the Devonian time (Rodendorf
1961).
As this study shows, many pleural parts in Thysanura and in lower pterygotes are
homologizable. The pleural modifications required for the wing mechanism were,
besides a stronger sclerotization (Snodgrass 1958), mainly the formation of the pleural
ridge and the pleural wing process as a fulcrum, upon which the wing base could rotate.
Formation of the subalare and the basalare was also necessary; and elimination of a large
number of muscles was required. At the same time, some flight muscles had to be
produced newly through complex modifications of preexisting muscles and some leg
muscles had to develop a second function, viz. they had to become flight muscles while
retaining their functions as leg muscles for walking.
These structural and functional changes must have accompanied the production of
the functional wing. Therefore, Ross' ( 1964) result, although highly significant in con-
sidering the origin of wings, cannot be conclusive,::o unless sudden, simultaneous
changes in at least some of the above-mentioned structures in some apterygotes
are proven.
Types of wings and the phylogeny of winged insects: Two major types of wings
have been recognized since Martynov's ( 1924) work: one type shared by Ephemeroptera
and Odonata, in which wings are held vertically when at rest; and another type shared by
other pterygotes, in which wings are held folded on the back in their resting position.
The two groups bearing these different types of wings are commonly called the Palaeop-
tera and the Neoptera, respectively. Numerous theories have been put forward on
whether these groups constitute two distinct natural groups, and further whether these
two wing types are derivable from one to another.
Lemche ( 1940) showed that in all Pterygota except Odonata and Saltatoria, the
wing pads develop backwards. In Blattaria the wing pad grows backwards and its
posterolateral corner becomes the wing tip in the adult. In Odonata, as in the immature
Palaeodictyoptera, the wing pads grow essentially sidewards. They later become tilted
along the longitudinal axis of the body. In Saltatoria the early wing development
coincides with that of Blattaria; only in later stages of development the presumptive front
margin becomes directed upwards and inwards, thereby simulating the condition in
Odonata."
IBRoss inferred this possibility from a study on the recessive gene, balloon-wing (Cochran and Ross 1961) in
Blattella germanica, which produces swelling on the wings and the pronotum, indicating that a single
mutation can affect all segments.
SlDespite this inconclusiveness regarding the origin of wings, however, some recent workers have specu1ated on
the origin of flight and to some degree on the origin of wings, and in their arguments the gradual change of the
paranota into the wings was taken for granted. Alexander and Brown (1963) discussed the possible
selective advantage of nonfunctional wings (flaps) or the extended paranota, presumably connected with the
direct method of copulation that arose in the early Pterygota. Leston ( 1963), however, thought that the
paranota as an organ for attracting females would have led to a unisexual evolution of wings. He suggested,
therefore, that the paranota have been a species-recognition signal. Wigglesworth (1963a. b) suggested that
the paranotal planes first arose in small, aerial insects as a controlling device during takeoff. Hinton (1963)
thought that the paranotal lobes arose first in connection with altitude control in falling. Fowler (1964)
studied 'the selective advantage of the theoretical paranotal lobe from the viewpoint of aerodynamics, and
concluded that large insects (of perhaps one centimeter in length) with small legs were capable of the best
gliding performance if suitable altitude could be maintained.
msellier (1950) showed, in a grafting experiment in Gryllus, that the first turning of the wing is due to the
greater development of the hypodermis on its concave face before the molt. Thomas (1954a) also found in
Lamarckiana that wing buds (pads) alter their position between the third and fourth molts and again at the
final molt.
. 42 MEMOIRS OF TilE ENTOMOLOGICAL SOCIETY OF CANADA
FIG. 16. Diagram showing relationships of major groups of insects (from Lemche).
Lemche (1940) further pointed out the differences between Ephemeroptera and
Odonata in the articulating mechanism of the wing. Although his identifications of the
axillary sclerites was wrong, he was certainly right in pointing out the differences between
the two orders in major features of the axillary sclerites, and in emphasizing the overall
similarity between the wing base in Ephemeroptera and that in other winged insects ...
Chiefly based on the above-mentioned characteristics in structure and development,
Lemche proposed (Fig. 16) that the class Insecta can be divided into the Plagioptera,
which include Palaeodictyoptera, Megasecoptera, Protodonata, and Odonata, and the
Opisthoptera, which include all other pterygote orders except Ephemeroptera. He
regarded the Ephemeroptera as an offshoot that separated very early from the evo-
lutionary line of the Opisthoptera. Thus, Ephemeroptera and Odonata are not on the
same phylogenetic line. It is thus implicit in his theory that the wings of modern insects
other than Odonata cannot be directly derived from the type found in Palaeodictyoptera.
Lemche pointed out that the fossil record does not contradict this conception, since Proto-
blattoidea and Palaeodictyoptera are both represented by eight of the oldest fossil
wings known.
A closer scrutiny of pertinent facts, which have been ignored by recent workers,
reveals that Lemche's theory is well founded. Bekker (1954, 1956) and Bocharova-
Messner ( 1965) found that in mayflies the wing pad arises from the posterolateral angle
of the tergum. In Ecdyonurus Ide ( 1936) found that even the posterolateral angle of
the pronotum is produced during the nymphal development. In several old nymphs of
the Permian mayflies Kukalov:i (1968) has shown thatthe wing pad is directed postero-
laterally, instead of being fastened back over the thorax as seen in modern mayflies."'
Yet the posterolateral extension of the wing pad in these primitive mayflies is evident.
Thus the pattern of wing development in Ephemeroptera is essentially the same as in
Blattaria.
Among Odonata Bocharova-Messner ( 1959) found that in the representatives of
Anisoptera (Libellu/a, Aeshna, Fig. 17 A) the pleurotergal area is largely membranous,
and the wing rudiment is recognized as an isolated plate above the spiracle; during the
second nymphal stage it shifts its position dorsally, and later, during the fourth or fifth
nymphal stage, it comes to lie along the lateral margin of the tergum. In Zygoptera
z:snodgrass ( 1929b) also noted that "the small plates of the axilJary region in the Mayfly suggest the definite
sclerites of the flexor mechanism in insects that fold the wings horizontally over the back."
23
In the Permian mayflies the mesothorax and the rnetathorax are subequal in size. It would be at least
rnechanicalJy difficult for the forewing to be folded back on the dorsum during development when the hind
wing of similar size grows simultaneously from the metathorax. '
~
~
A
B
43
'
FIG. 17. Development of wings in Odonata. A, Libellula quadrirrzaculata; B, Agrion virgo
(from Bocharova-Messner).
(Agrion, Fig. 17B), however, the pleurotergal area is largely well sclerotized in the first
stage, and the membranous fold representing wing rudiments appears along the whole
lateral margin of the tergum in the second stage nymph. Both in Zygoptera and
Anisoptera, as is clear from the above description, the wing pad arises and extends
laterally; this fact clearly shows the difference in the mode of development of the wing
between Ephemeroptera and Odonata, and supports Lemche's idea.
Further, in the nymph of a fossil insect, Mischoptera douglassi (Megasecoptera),
Carpenter and Richardson ( 1968) have shown that wings join the thorax at nearly right
angles to the body before being directed obliquely backwards, and in this respect
Mischoptera is essentially the same as Odonata; the similarity in the position of wing pads
in the two groups in turn supports the inclusion of Megasecoptera into the Plagioptera
by Lemche (1940).
The differences in wing base structures between Ephemeroptera (p. 116) and
Odonata (p. 390) are obvious. In Ephemeroptera the axillary sclerites are, by their
relative positions, directly .homologizable with those in other winged orders, whereas the
wing base structures in Odonata, except possibly for the first axillary sclerite, can not be
homologous with those in other orders. Characteristic features of the wing base struc-
tures in Odonata, however, were probably derived from structures such as those seen in a
palaeodictyopterous species, Ostrava nigra (Fig. 18A). In this species the bases of all
wing veins are separated from each other and the association of the base of each vein
with the vein proper is quite clear. By comparing the wing base in Odonata with the
wing base in Ostrava, Kukalova (1960) suggested that the axillary plate in Odonata was
44
PRECOSTAL
FIELD
OSTORADIAL
FIELD
Sc
RtM
COSTAL PLATE
Sc ~ : . : . . . z . . ~ ~ ~ ~ r : :
RADIOANAL
PLATE
R
M
Cu
An
CUBITAL
PLATE
Cu
An
Fro. 18. A, wing base of Ostrava nigra; B, the same in Odonata (from Kukalov:i).
produced by fusion of the bases of R, M, Cu, and A veins; and this was also the con-
clusion Tannert (1958) reached in his study of Odonata. She further pointed out that
the greatly swollen basal part of the C vein (precostal field of Kukalova) probably gave
rise to the large costal (humeral) plate in Odonata.
It is much more difficult to derive the axillary sclerites in Ephemeroptera and other
orders from the wing base structure in Ostrava. In Ephemeroptera and other orders, or
the Opisthoptera of Lemche (1940), the bases of M and Cu veins were primitively the
proximal median plate (p. 16). In Ostrava the bases of these veins lie too proximal to
be the proximal median plate in Opisthoptera; the base of the A vein in Ostrava is very
.small, whereas the base of the A veins in Opisthoptera is the well-developed third axillary
sclerite. Furthermore, the large base of the C vein in Ostrava cannot be ancestral to the
humeral plate in Opisthoptera, which is always inconspicuous.
The position of the wings at rest, whether held vertically or folded back on the
dorsum, is a main characteristic distinguishing the Neoptera from the Palaeoptera. In
Odonata muscles t-s 10 and t-s 11 are the elevators of wings and they are responsible for
holding the wings vertically. In Ephemeroptera, as Grandi ( 1947b) showed, t-cx 4,
t-tr 2, and t-s 3 are responsible for holding the wings vertically. Clearly, the muscles
responsible for holding wings in the vertical position are not homologous between the
two orders.
Among fossil insects, Zalessky (1958) showed the wings folded on the dorsum in
Asthenohymen (Megasecoptera, Palaeoptera), and contended that the folded position of
wings arose polyphyletically in insects.
In higher orders, also, the position of wings can be variable. As Jeanne! ( 1949)
pointed out, many butterflies hold their wings vertically, although most moths hold them
on the dorsum. Schwanwitsch (1956) also noted that among Diptera a Psychoda holds
the costae downwards, and a Phlebotomus holds them upwards; in a Culex the wings are
held fiat over the abdomen, and in Tipula they are spread out.
As the preceding discussion clearly shows, the position of wings at rest fails to be
the absolute criterion for separating the Palaeoptera from the Neoptera, and for dis-
tinguishing one type of wing from another.
MATSUDA: THE INSECT TIIORAX 45
The absence or presence of the neala, which is also used as a character in separating
the Palaeoptera from the N eoptera, is also not a valid criterion, since the neala does not
develop in butterflies, termites, and others. The absence _of the jugum (neala) in
Rhopalocera can be accounted for by the vertically erected position of the wings at rest;
in this position the jugal fold simply would not develop. The claimed similarities in
wing venation between the Neoptera and Odonata (Forbes 1943; Hennig 1953) are,
according to Mackerras (1967), spurious. The thoracic musculature in Odonata is
unique (p. 399). Diirken (1907) and Schwanwitsch (1943, 1946) already pointed out
the basic differences in the thoracic musculature between Odonata and Ephemeroptera,
and Schwanwitsch called Odonata the Orthomyaria and other pterygotes including
Ephemeroptera the Chiastomyaria.
In addition to the above facts, the presence in Odonata of the intratergal apophysis,
the prefurca, the muscles associated with these structures (p. 60, 66) and the unique
head structures (see Matsuda 1965), etc. lead to the conclusion that Ephemeroptera and
Odonata are not closely related at all, and the differences between them are significant
enough for the separation of Odonata from the rest of the Pterygota, as Lemche (1940)
attempted to do. More discussion on the phylogenetic position of the Odonata is given
on p. 398.
As Grandi (1947a) showed and as Hennig (1953) referred to, the wing base struc-
tures in Odonata and Ephemeroptera are quite dissimilar. The alleged differences in
the wing base structures between Ephemeroptera and other Pterygota (Grandi 1947a)
are, however, untrue (p. 116). Hennig unfortunately accepted Grandi's (1947b)
concept of the pseudopteralia (p. 116), which proposed the uniqueness of the axillary
sclerites in Ephemeroptera, and her remark that the axillary sclerites in the Neoptera can
be derived from those in Odonata. Hennig was therefore led to believe that Odonata +
Neoptera (Metapterygota of Hennig) represent a sister group of Ephemeroptera. Such
a phylogenetic relationship is inconceivable, in the light of the preceding discussions.
Rodendorf (1961, 1962) described a winged fossil insect, Eopterum devonicum,
from the Upper Devonian beds. This is certainly the oldest winged fossil insect ever
discovered. Rodendorf erected a new order, Archaeoptera, for this species and assigned
it to the subclass Neoptera. He recognized the anal region which he considered to be a
characteristic of Neoptera. He further maintained that the wing structure of this
Devonian insect justified the belief that it is phylogenetically closest to the original forms
of all winged insects, both Neoptera and Palaeoptera.
Sha,rov ( 1966) pointed out that the wings in Eopterum were not folded on the
dorsum, but were spread somewhat obliquely sidewards. Sharov, therefore, contended
that this position of wings cannot be the characteristic of Neoptera, and he proposed a
new subclass, Archaeoptera, for Eopterum. The characteristics of Archaeoptera are:
ability of the wings to draw back to a tapered position in relation to the body, and the
absence of the jugal fold at the base of the wing. He further maintained that the
extremely simple wing venation of Eopterum could not be ancestral to those in Palaeop-
tera and Neoptera. Therefore, he pointed to the wing of a carboniferous family,
Paoliidae, with well-developed wing venation, as probably ancestral to both Palaeoptera
and Neoptera, and he assigned this family to the Archaeoptera because of the absence of
the anal fan and a similar resting position of the wings to that of Eopterum.
Thus in Sharov's theory, the position of the wings at rest was the primary charac-
teristic upon which his new subclass Archaeoptera was erected. The position of the
wings given in figs. 1 and 2 by Rodendorf and reproduced by Sharov, however, remind
us of a position which some moths or bugs might assume under some circumstances.
Moreover, as already discussed, the resting position of the wing cannot be a reliable
character in separating the subclasses of the Insecta. The absence of the neala occurs
both in the Plagioptera and the Opisthoptera. Therefore, this character is also insignifi-
cant in erecting a new subclass.
Earlier opinions on the origin of wings and its implication on the phylogeny of
insects were summarized by Crampton (1916a), Tillyard (1930), Jeanne! (1949), Wille
(1960), Sharov (1966), etc. For a reference or summary of palaeontological knowledge
of insects, works by Handlirsch (1906-08, 1930), Jeanne! (1949), Carpenter (1953),
and Rodendorf ( 1962) are useful.
THE COXA
The proximal end of the coxa is typically girdled by a submarginal basicostal suture
that internally forms a ridge known as the basicosta, and it externally demarcates a
marginal flange known as the basicoxite. The basicoxite forms a variable number of
sockets, and the condyles on the pleuron articulate with the coxa by fitting into these
sockets.
In Thysanura the pleuron has a maximum of three points of articulation with the
coxa: anterior, medial, and posterior (Fig. 38A). The ventral condyle for articulation
with the coxa in Diplura is represented by the thickened lateral end of the Y ridge. In
Thysanura and Plecoptera the sternacosta or the lateral end of the Y ridge does not form
the articulatory condyle. The anterior and median articulations with the coxa are
homologous in Thysanura and Pterygota, but the posterior articulation (with the coxa)
in Thysanura is absent in the Pterygota. With the development of the pleural ridge in
the Pterygota, according to Matsuda ( 1963a), the posterior condyle became lost and the
median condyle in Thysanura has become the definitive posterior pleural condyle that
articulates with the coxa in the Pterygota. This definitive posterior articulation occurs
most persistently in the winged insects.
In holometabolous insects, in which the leg bases are approximated toward the
median longitudinal axis, a ventral condyle is formed along the base of the furca. As
already discussed, this condyle is a transformed ventropleurite, which is katapleural in
origin and is therefore not homologous with the ventral condyle in some Apterygota.
This condyle is absent in lower hemimetabolous insects; in some nymphal Plecoptera and
in the nymph of Sialis, lateral ends of the Y ridge are associated with the furca, but they
do not participate in the ventral articulation with the coxa. These facts therefore
support the idea that the ventral condyle in adult holometabolous insects is a secondary
formation, although Ferris (1940b) thought it was primary.
The outer wall of the coxa in some insects is marked by a longitudinal suture, called
the coxal suture; it extends from the basicoxite to the anterior trochanteral articulation,
and is reinforced internally by a ridge. In the pterothoracic segments of some insects
the posterior margin of the coxa is enlarged to form the meron; the suture demarcating
this area is the meral suture, or the merocosta. According to Snodgrass ( 1927, 1935)
and Weber ( 1933), this suture is the posterior extension of the basicoxal suture. Larsen
( 1945a, b), however, found that it was formed secondarily. The meron is often greatly
enlarged, and the merocosta is, as Larsen maintained, an independent formation
developed for the accommodation of flight muscles. In Aphrophora Larsen showed
that the basicoxite is complete during the younger nymphal stages, and the meron is
formed at the final stage of the nymphal development.
POSITIONS OF THE THORACIC SPIRACLES
Normally 10 pairs of spiracles are recognized, two thoracic and eight abdominal.
There is no question about the segmental distribution of the metathoracic and the eight
abdominal spiracles, but the primitive position of the anteriormost spiracle, whether it is
prothoracic or mesothoracic, has been disputed. The issue arises from the fact that the
anterior spiracle lies often on the intersegmental area between the prothorax and the
mesothorax, or on the posterior part of the prothorax (Lepidoptera, Hymenoptera, etc.),
or on the anterior part of the mesothorax. This apparent inconsistency in position of
the anterior spiracle must be accounted for in terms of a shift from its primitive position,
both in phylogeny and ontogeny.
In Lepisma saccharina (Fig. 37) both the anterior and posterior thoracic spiracles
clearly lie within the segmental limit of the meso- and meta-thorax. A significant shift
47
of their positions during development in Thysanura is not known (Heymons 1897;
Woodland 1957). In Ephemeroptera the anterior thoracic spiracle lies clearly within the
mesothorax (Fig. 42B), both in the nymph (Diirken 1907) and in the adult (Knox 1935;
Matsuda 1956b). In the embryo of Blatta, Gryllus, and Forficula (Heymons 1895),
the thoracic spiracles lie within the segmental limit of the meso- and meta-thorax. In
Gry/lus and Forficula, according to Heymons (1895), the anterior spiracle migrates
cephalad towards the end of the embryonic development. Voss ( 1905) claimed that the
anterior spiracle was prothoracic in the adult of Gryllus, which must have been the result
in shifting during the postembryonic development. In Locusta (Roonwal 1937) the
thoracic spiracular invaginations in a 112-hour-old embryo belong to the meso- and
meta-thorax; they migrate forward, so that the anterior spiracle comes to lie on the
membrane between the pro- and meso-thorax, and the posterior one comes to lie on the
posterior margin of the mesothoracic pleuron. In Carausius (Lehmann 1926), also, the
anterior thoracic spiracle lies on the anterior part of the embryonic mesothorax and later
it migrates forward. In Odonata Ando's (1962) figures show that the thoracic spiracles
clearly lie within the embryonic meso- and meta-thorax.
The above facts concerning Thysanura and lower hemimetabolous insects lead us to
believe that the primitive position of the anterior thoracic spiracle is intrasegmental, and
that a definitive intersegmental position in the adult of some of these insects is due to a
shift in position during development.
In Neuroptera the anterior thoracic spiracle lies between the prothorax and the meso-
thorax, and at least in the larva of Myrmeleon (Sundermeier 1940) it lies on the
anterolateral part of the mesothorax. In the embryo of Hydrophilus (Heider 1889) the
anterior spiracle is mesothoracic, and in Chalastogastra (Weber 1923, 1926) the anterior
spiracle lies between the prothorax and the mesothorax. It is therefore clear that in
lower holometabolous insects including Coleoptera the position of the anterior spiracle
is essentially the same as in Hemimetabola.
In higher holometabolous insects, however, the anterior thoracic spiracle lies on the
prothorax. Thus, in Pteronidea (Shafiq 1954) and Pontanica (Ivanova-Kasas 1959)
among Hymenoptera, the anterior thoracic spiracle lies on the embryonic prothorax. In
the embryo of Pieris (Eastham 1930) the spiracular invaginations originate in front of
and above the paired appendage in the prothoracic, the metathoracic, and the first eight
abdominal segments. In three genera of fleas (Kessel 1939) the embryonic anterior
thoracic spiracle lies on the prothorax, and in the adult fleas (Snodgrass 1946) it lies
clearly on the area anterior to the sclerotized mesothorax. During the postembryonic
development of some Lepidoptera, as Weber (1927 b) showed, the anterior thoracic
spiracle clearly lies within the prothorax; it then comes to lie more posteriorly along the
wing base in the pupa. Yet in the adult the spiracle appears to lie on the prothorax.
Weber believed that the imaginal spiracle is, despite its shift, the same as the one present
in the larva. In larval Diptera, as Keilin ( 1924, 1944) showed, the thoracic
occur only on the prothorax and the metathorax.
These facts clearly show that in higher holometabolous insects the anterior spiracle
tends to occur on the prothorax, and this is especially true during earlier developmental
stages. Yet, the anterior spiracle in higher holometabolous insects must be homologous
with the anterior spiracle in Thysanura and lower orders which occurs on the mesothorax,
since it is highly improbable that in these holometabolous insects the prothoracic spiracle
suddenly appeared with the simultaneous disappearance of the mesothoracic spiracle.
Apparently, in phylogeny of insects, the anterior thoracic spiracle has shifted its position
anteriorly from its primitive position on the mesothorax; and in higher orders this
primitive position of the spiracle cannot be recognized even during earlier stages of
development.
The above evolutionary trend in the position of the anterior thoracic spiracle in the
Thysanura-Pterygota, however, does not apply to the entognathous Apterygota, more
spiracles being present in this group of insects. Thus, as Barlet and Carpentier (1962)
clearly showed, the thoracic spiracles in Diplura number four. In Oncojapyx (Fig. 32)
the first one is clearly prothoracic .. ; the second and fourth ones are intrasegmental; being
located above the anapleuron of the meso- and meta-thorax; the third one lies inter-
segmentally between the meso- and meta-thorax. These spiracles are difficult to homolo-
gize with those in Thysanura-Pterygota. In Sminthuridae (Collembola) a spiracle
occurs in the neck region; Davies (1927) thought it belonged to the prothorax and
Snodgrass thought it was labial in origin. In Eosentomon (Prell 1913; Fran9ois 1964)
the meso- and meta-thorax have a single spiracle that lies on the middle of the lateral
membranized tergal area. They appear to be homologous with two suprapleural spiracles
in Oncojapyx. Lehmann (1926) thought that the smaller number of spiracles in
Protura was due to reduction. Snodgrass (1935) pointed out that the positions of
spiracles in J apygidae correspond to the usual positions of the spiracles in the Chilopoda.
The structural details of the spiracle are not discussed in this study, as they will be
treated fully in the future in discussing the respiratory system. The most recent summary
of the physiology of the respiratory system was given by Miller ( 1964). This work
includes also a summary of morphology of the spiracle.
Maki's (1938) work shows variations in number and position of muscles attached to
the spiracle in the Pterygota. As Hassan's ( 1944) study shows, however, no muscle is
attached to the spiracle in Machi/is; in Lepisma Barlet showed that a muscle from the
endosternal area n is attached to the stalk supporting the spiracle and is probably an
occulsor. It seems certain, therefore, that in the Thysanura-Pterygota complex the
smaller numbers of muscles is primitive. The origin of the larger number of spiracular
muscles in the Pterygota remains unknown.
Theories: The conclusion regarding the primitive position of the anterior thoracic
spiracle and its secondary shift in the phylogeny of the Thysanura-Pterygota complex
agrees with those reached by Weber (1923, 1926, 1927b), Lehmann (1924, 1926), and
Hinton (1947), who expressed essentially the same opinions. Among earlier workers,
Boas (1899) and Snodgrass (1909) also presumed that the primitive position of the
anterior thoracic spiracle was on the mesothorax.
Burmeister (1832), Miiller-Biumenau (1884), Kolbe (1893), and Keilin (1924,
1944) thought that the primary positions of the spiracles were intersegmental. Keilin
based his contention on the fact that in the larval Diptera there are 11 segments posterior
to the head, while there are only 10 spiracles including prothoracic, metathoracic, and
eight abdominal ones. To recor>.cile these different numbers, Keilin thought that the
spiracles must have been primarily located intersegmentally, and this idea was accepted
by Richards and Davies (1957) and Miller (1964). As is clear from the foregoing
discussion, however, this idea of Keilin's is not supported by facts. Weber ( 1923, 1927b)
criticized the Schaltsegmenttheorie of Feuerborn (1922), in which the anterior spiracle
was considered to be an integral part of an extra segment between the prothorax and
the mesothorax.
2. THE MuscuLATURE
THORACIC MUSCULATURE IN LEPISMA SACCHARINA
(Figs. 19-21)
The thoracic muscles in Lepisma saccharina (Barlet 1953, 1954) are described
below to serve as a standard of reference for the muscles of the Pterygota. The numbers
of serially homologous muscles in parentheses are those used by Barlet. Homologies of
thoracic muscles between Lepisma saccharina and Lepismachilis Y-signata are given in
Table III (p. 112).
"'The embryonic prothoracic spiracle was also found in Blattel/a (Cholodkowsky 1891) and in Leptlnotarsa
(Wheeler 1889). In the embryo of the honey bee Nelson (1915) found a tracheal invagination on the
labial segment.
Tergal Muscles
(Fig. 19)
49
Muscle 1 (8, 14), a transsegmental muscle, extends between the prescutum and the
postmedian area or the posterior end of the succeeding tergum. The prothoracic muscle
1 of Barlet may also be homologous with 8 and 14, although it is listed below as the
homologue of muscle 9.
Muscle 2 (5, 10, 15), a transsegmental muscle, connects the anterior tergal area and
the posterolateral area of the succeeding tergum.
Muscle 3 ( 6, 9, 16), a long transsegmental muscle, arises from the median tergal
area of a segment and is inserted on the posterolateral area of the succeeding tergum
( 16), or on the median lateral area of the succeeding tergum ( 6, 9).
Muscle 4 (7, 11, 17) connects the median or anterior part of the tergum of a
segment and the antecosta of the succeeding segment.
Muscle 5 (4, 12, 18) is attached anteriorly near the prescutum (12, 18) or to the
upper part of the cervical membrane ( 4), and is inserted posteriorly on the antecosta.
Muscle 6 (13, 19) is attached anteriorly to the median or postmedian tergal part of
the meso- and meta-thorax, and posteriorly to the antecosta on the level of the endonotal
phragma ( 13), or to the tip of arched endonotal phragma ( 19). Barlet ( 1953) doubted
that they are homologous. No serial homologue occurs in the prothorax.
Muscle 7 (3) connects the median dorsal area of the pronotum and the postoccipital
sclerite 1. No homologue is present in the meso- and meta-thorax.
Muscle 8 (2) is attached anteriorly on the postoccipital sclerite 1, below the attach-
ment of muscle 7, and posteriorly to the mesothoracic antecosta.
Muscle 9 ( 1) arises from the middle of the mesonotum and attaches anteriorly to
the apodeme, which separates the postoccipital sclerite 1 from the hind end of the head.
Sternal Muscles
(Fig. 19)
Muscle 11 (28, 34, 42, 51) obliquely connects the endosternal m and n of the
succeeding segment ( 34, 42), or extends between the posterior tentorium and the pro-
thoracic endosternal stalk n ( 28) ; 51 is attached posteriorly to v of the first abdominal
endosternum.
Muscle 12 (30, 35, 43, 52) connects m and g or their equivalents (35, 43, 52), or
connects the posterior tentorium and g ( 30).
Muscle 13 (32, 36, 44) extends between m and b (36, 44), or between the labial
tendon below the posterior tentorium and the area in front of g (32). Since 32, 36, and
44 lie ventral to all other muscles they are here considered serially homologous.
Muscle 14 (37, 45) extends between d and b of the succeeding segment. No
homologue is present in the prothorax.
Muscle 15 (31, 38, 49, 53?) extends between the base of d and g of the succeeding
segment ( 3 8, 49), or extends between the posterior tentorium and g of the prothorax
(31); 53 attaches anterior ton of the metendosternum and may not be homologous.
Muscle 16 (29, 39, 47, 57) extends between nand d of the succeeding segment (39,
47, 57), or between the collar tentorium and g of the prothorax (29).
Muscle 17 (75, 40, 48, 58?) extends between n and g of the succeeding segment
( 40, 48), 75 is a prothoracic muscle that extends from the postoccipital sclerite 2 to g
of the prothorax; 58 arises anteriorly from d and may not be homologous.
Muscle 18 (41, 46, 56) connects g and g of adjacent segments. No homologue is
present in the prothorax. Barlet homologized 56 with his 37.
Muscle 19 (33), a prothoracic intraendosternal muscle, connects g and m (not
shown in Fig. 19).
Muscle 20 (50, 55?) extends between the mesothoracic b and the metathoracic b
(50); it is absent in the anterior segments. 55 arises anteriorly from near the base of n,
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Muscle 21 (62), a prothoracic muscle, extends between the anterior angle of the
furcasternum and /.
Pleurosternal Muscles
(Figs. 19, 20)
Muscle 25 (59, 60, 61), a transsegmental pleurosternal muscle, extends between the
posterior half of the anapleural apophysis and the endosternal b of the succeeding
segment.
Muscle 26 (81, 88, 95) extends between the lateral extremity of the tigella n and
the dorsal anapleural area. It is a spiracular muscle and is absent in the prothorax.
Muscle 27 (201), a prothoracic muscle, extends from the anapleural tendon p to
the endosternal g.
Sternocoxal Muscles
(Fig. 20)
Muscle 31 (106, 112, 118) connects m or its equivalent and the coxal process of the
succeeding segment in the mesothorax and meta thorax ( 106, 112), or m and the pre-
sumed coxal area of the first abdominal segment ( 118).
Muscle 32 (100, 107, 113) penetrates deeply into the coxal cavity and is inserted
along the groove between the anterior and posterior faces of the coxa. The anterior
point of attachment is on the collar tentorium ( 100) or m ( 107, 113).
Muscle 33 (1 02, 108, 114) extends between the median trochantinocoxal articula-
tion and the external side of g.
Muscle 34 (103, 109, 115) arises from the basisternum, passes through f and e,
and is inserted on the coxal margin behind the coxal process.
Muscle 35 (104, 110, 116) connects the posterodorsal coxal margin and the endo-
sternal area beneath b.
Muscle 3 6 ( 105, 111, 117) is attached to the posterior coxal margin and m of the
endosternum.
Muscle 37 (101), a prothoracic muscle, extends between the area behind the
posterior trochantine-coxal articulation ( C of Barlet) and the base of f.
Muscle 38 (99) extends between the prothoracic coxal process and the posterolateral
angle of the collar tentorium on the opposite side. It crosses the counterpart coming
from the coxal process on the opposite side.
Tergosternal Muscles
(Fig. 19)
Muscle 41 ( 68, 80, 87, 94) connects the endonotum n ( 80, 87, 94), or the post-
occipital sclerite 2 with a structure (feuillet posterieur de Ia mentonniere of Barlet)
beneath the labium.
Muscle 42 (72, 78, 79, 86, 93) connects the endosternal area behind g and the
anterolateral area of the tergum of the succeeding segment (78, 79, 86, 93), or the collar
tentorium and the anterior lateral area of the pronotum (72).
Muscle 43 (73, 82, 89, 96), a transsegmental muscle, connects m and the antero-
lateral area of the tergum of the second succeeding segment (82, 89, 96), or the posterior
margin of the anterior tentorium and the lateral margin of the mesonotum behind the
middle (73).
Muscle 44 (74, 83, 90), a transsegmental muscle, has the anterior point of attach-
ment on m (83, 90) or the anterior tentorium (74), and its posterior point of attachment
on the antenotum of the second succeeding segment.
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MATSUDA: TilE INSECT THORAX
53
Muscle 45 (76, 84, 91) connects the median lateral area of the tergum and g (84,
91); in the prothorax it lies parallel to the endosternal tigella k (76).
Muscle 46 (77, 85, 92) connects the endonotal phragma and g.
Muscle 47 (69, 70) arises dorsally from the cervical membrane (69) or from the
area anterior to the pronotum (70), and is attached to a tendinous lamella associated
with the labial gland.
Muscle 48 (71), a prothoracic muscle, connects the distal end of the tigella k and
the small process on the posterior margin of the presternum ( sclerite Y of Barlet). The
ventral point of attachment is very close to that of muscle 87 ( 172 of Barlet).
Muscle 49 (97) extends between the metendosternal n and the antecosta of the
second abdominal tergum.
Trochanteral Muscles
(Figs. 20, 21)
Muscle 50 (119, 122, 125) connects the interior angle of the trochantin and the
trochanter in all thoracic segments.
Muscle 51 ( 120, 123, 126) shares the same point of attachment on the trochanter
with muscle 50. Its other point of attachment is f of the endosternum (not shown for
the prothorax and metathorax in Fig. 20).
Muscle 52 ( 121, 124, 127) is a powerful muscle which comes from the middle of
the tergum and is inserted on the trochanter.
Muscle 53 (193, 198) connects the tip of the endokatapleural apophysis and the
upper angle of the trochanter. It occurs only in the mesothorax and metathorax.
Muscle 54 (189, 194, 199) connects either the katapleural part of the endopleuron
(189 in prothorax) or the katapleural margin (194, 199) and the upper angle of the
trochanter.
Muscle 55 ( 190, 195, 200) connects the anapleural tendon p and the upper angle of
the trochanter, as muscle 54. Large part of this muscle is confluent with muscle 53.
Tergocoxal Muscles
(Fig. 20)
Muscle 61 ( 129, 134, 139, 144) originates from the anterior area of the tergum and
is inserted on the coxal process in all thoracic segments (129, 134, 139). Barlet (1954)
homologized 144 in the first abdominal segment with 129, 134, and 139.
Muscle 62 ( 130, 135, 140) originates from the tergum posterior to the origin of
muscle 61 and is inserted ventrally on the median point of the trochantinal articulation
with the coxa (B of Barlet).
Muscle 63 ( 131, 136, 141, 145) originates from the posterior area of the tergum
and is inserted on the dorsal posterior margin of the coxa (131, 136, 141); 145 is prob-
ably the serial homologue in the first abdominal segment.
Muscle 64 (132, 137, 142) originates from the tergum anterior to the origin of
muscle 63, and its insertion is on the posterior internal margin of the coxa.
Muscle 65 ( 133, 138, 143) originates on the tergum posterior to the origin of
muscle 63, and its ventral point of attachment is on the posterior margin of the coxa,
deeper beyond that of 64.
Muscle 66 ( 128), a prothoracic muscle, is attached anteriorly on the postoccipital
sclerite 2, and its posterior point of attachment is on the groove that corresponds
internally to the distal edge of the coxa.
Tergotrochantinal Muscles
(Fig. 20)
Muscle 71 ( 146, 149, 153) arises from the postoccipital sclerite 2 (146) or from
the endonotal phragma ( 149, 153) and is inserted ventrally on the interior angle of the
apex of the trochantin.
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Muscle 72 (150, 154, 157?) originates from the median area of the meso- and
meta-ndtum and is inserted ventrally on an area near the point of attachment of 71.
This muscle is absent in the prothorax.
Muscle 73 (147, 151, 155, 157?) originates from the tergum posterior to the origin
of 72, and is inserted on the tendon from the interior angle of the trochantin.
Muscle 74 (148, 152, 156) arises from the posterior area of the tergum and is
inserted on the posterior margin of the trochantin in all thoracic segments.
Tergopleural Muscles
(Fig. 21)
Muscle 81 (158, 159, 165) arises from the middle of the tergum and is inserted on
the anterior margin of the katapleuron (the paracoxal suture) near its ventral end (the
meso- and meta-thorax) or at its middle (the prothorax).
Muscle 82 (160, 166) connects the endonotal phragma and the paracoxal ridge
above the point of attachment of 81 in the meso- and meta-thorax. This muscle is
absent in the prothorax.
Muscle 83 (161, 167) originates from near the prescutal area on the meso- and
meta-notum and is inserted on the paracoxal ridge above the point of insertion of
muscle 82. This muscle is absent in the prothorax.
Muscle 84 (162, 168) lies posterior and parallel to muscle 83 in the meso- and
meta-thorax. This muscle is absent in the prothorax.
Muscle 85 (163, 169) arises from the anterior area of the meso- and meta-notum
and is inserted on the endopleural process (katapleural). It is absent in the prothorax.
Muscle 86 (164, 170) is external to all tergokatapleural muscles. It is inserted on
the endopleural process in the meso- and meta-thorax, and is absent in the prothorax.
Muscle 87 (172, 177, 181) originates on the tip of the stalk k in the prothorax
( 172) or on the anterior area of the meso- and meta-notum ( 177, 181); it is inserted on
the anterior margin of the sclerotized anapleuron near its distal end in the prothorax, or
at its middle in the meso- and meta-thorax.
Muscle 88 (173?, 178, 182) arises from the. anterior region of the tergum anterior
to the origin of 87 ( 178, 182) and is inserted on the anapleural margin above the point
of insertion of muscle 87; 173 originates from the pronotum posterior to the point of
origin of 87 and may not be serially homologous.
Muscle 89 (174), a prothoracic muscle, arises from the neck membrane and is
inserted on the anapleuron. This muscle may be serially homologous with muscle 82 of
the meso- and meta-thorax, which, however, is inserted on the katapleuron.
Muscle 90 ( 175), a prothoracic muscle, connects the postoccipital sclerite 2 and the
anapleural tendon.
Muscle 91 (179, 183, 186?) arises from the midlateral part of the tergum and is
inserted on the anterior edge of the dorsal (superieur) anapleural apophysis. It is not
present in the prothorax.
Muscle 92 (176, 180, 185, 187?) originates from the posterolateral area of the
tergum (176 in the prothorax), or from the posterolateral angle of the tergum, where the
postnotal membranous infolding begins ( 180, 185) ; it is inserted on the dorsal anapleural
anophysis.
Muscle 93 ( 184), a metathoracic muscle, lies posterior to muscle 91 and is inserted
on the dorsal anapleural apophysis.
Pleurocoxal Muscles
(Fig. 21)
Muscle 101 (191, 196) originates on the upper margin of the katapleuron near the
endopleural apophysis and is inserted on the anterior margin of the coxa. It is absent
in the prothorax.
Muscle 102 (188, 192, 197) connects the katapleuron near the endopleural
apophysis and the posterior ventral surface of the coxa.
Other Muscles
(Fig. 21)
Muscle 111 (202, 203) is present only in the meso- and meta-thorax. It connects d
and cross-figures (see Fig. 21) in the posterior membranous folded areas of the meso-
and meta-thorax. The dorsal attachment corresponds to the area where a spiracle occurs
in Campodea (Fig. 33).
THORACIC MUSCULATURE IN THE PTERYGOTA
(Figs. 22-26)
The homologies of muscles of pterygotes referred to in the following discussions are
listed in Tables II to XXIV. Prothoracic tergopleural.muscles are omitted, since they
are often too difficult to compare with tergopleural muscles in the pterothorax. Larval
muscles listed are mainly those homologizable with muscles in the adult of the Apterygota
and of lower pterygote orders. Some muscles, whose derivations are described as
unclear, are neogenetic muscles (p. 81).
In designating the muscles, the anterior or dorsal points of attachment always come
first and are followed by the posterior and ventral attachments; for instance, op-t
( occipitotergal), cv-p ( cervicopleural), t-p (tergopleural), t-s ( tergosternal), t-cx ( tergo-
coxal), etc. This system is consistently followed throughout the work.
Voss' (1905) designations of muscles by symbols such as dv, bm, zm, etc. have had
some currency in the past, especially among German workers, but this system is no
longer useful, because many more thoracic muscles are recognized now than were known
at Voss' time. Similarly, the number of thoracic muscles recognized by Keler (1963) is
hardly enough for describing the thoracic musculature in insects.
Cervical Muscles
(Fig. 22A, B)
cv(d)-t 1 arises posteriorly from the middle of the first phragma in most species and
is inserted either on the dorsal cervical sclerite (e.g., Periplaneta), or on the tendon from
the dorsal cervical sclerite (e.g., Stagmomantis), or on the dorsal cervical membrane.
This muscle must be serially homologous with t 14 in the pterothorax (median dorsal
longitudinal muscle), and is probably homologous with muscle 5 in Lepisma (Fig. 19).
op-t 1 arises from the middle of the anterior margin of the pronotum or the dorsal
cervical sclerite (e.g., Periplaneta) and is inserted either on the dorsolateral cervical
membrane near the posterior end of the head, or on the dorsolateral angle of the post-
occipital ridge (e.g., Siphlonurus). 27 in Lepidocampa (Diplura) agrees in position
with this muscle.
op-t 2 arises from the median or posterior part of the pronotum and is inserted on
the dorsolateral angle of the postocciput (e.g., Dissosteira), or on the apodeme which is
on the dorsolateral portion of the cervical membrane near the posterior end of the head.
This muscle is probably homologous with 7 of Lepisma (Fig. 19), which also occurs only
in the cervico-prothorax.
op-t 3 arises posteriorly from the lateral area of the first phragma, and is inserted
either on the dorsolateral angle of the postocciput (e.g., Siphlonurus, Dissosteira) or on
the apodeme on the dorsal cervical membrane near the postoccipital ridge in some insects.
This muscle is probably homologous with 8 in Lepisma (Fig. 19); in position, op-t 3 also
agrees with the lateral part of the greatly enlarged t 14 in higher insects.
op-t 4 is 3.40 in larval Limnephilus which stretches between the middorsal region of
the anterior part of the mesonotum and the posterior region of the head capsule. In
57
A
0
/
';;)ZJi("'

/ s-cx(cv) 1 dv-cx 3
cv-cx l(x)
B
FIG. 22. A and B, cervical muscles in Pterygota (original).
position this muscle agrees with muscle 9 in Lepisma (Fig. 19) and may be homologous
with it; L.D. 1 in the pseudofoetus of Dilta (Barlet 1950a) is also probably homologous
(not figured).
op-cv 1 connects the dorsolateral angle of the postocciput with the anterior end of
the lateral cervical sclerite. In Corydalus op-cv 1 is inserted on the ventrolateral surface
of the jugulare. In some Orthoptera, as Alicata (1962b) showed, the anterior (dorsal)
point of attachment can be more ventral, and the muscle itself is shorter and sub-
horizontal in position. This muscle may be homologous with 41 in Lepisma (Fig. 19).
58 MEMOIRS OF 1HE ENTOMOLOGICAL SOCIETY OF CANADA
op-cv 2, 3 arise from the dorsolateral angle or sometimes from the ventrolateral part
( op-cv 3) of the postocciput, and is inserted on the middle and posterior areas of the
lateral cervical sclerite. op-cv 2, 3 might have been derived as a result of secondary
split of an enlarged op-cv 1. No such muscle is present in Thysanura.
op-cv(v) 4 (58a in Forficula) connects the submental corner of the head with the
posterior ventral cervical sclerite. The derivation of this muscle is unclear (not figured).
op-p 1 arises from the dorsal or lateral part of the postocciput (0 lm 9 in Perla) and
is inserted on the anterior angle of the prothoracic episternum. 0 idvm 1 in Dixippus,
which connects the lateral cervical membrane and the coxosternal plate (of Jeziorski,
1918), may represent this muscle. This muscle is probably serially homologous with
t-p 3 in the pterothorax, and is also probably homologous with 89 in Lepisma (Fig. 21).
op-p 2 (12 in Odontotermes) connects the dorsolateral portion of the posterior end
of the head and the pleural ridge above the pleural arm. This muscle may be the serial
homologue of t-p 4.
op-p 3 (DVL 2 in Stenopsocus) connects the dorsal part of the postoccipital ridge
with the ventral end of the preepisternum (precoxale) of the prothorax. This muscle
may be serially homologous with t-ti 1. in the pterothora"{, which is homologous with 71 in
Lepisma (Fig. 20).
op-s 1 (12 in Siphlonurus) connects the dorsal postoccipital ridge and the antero-
lateral margin of the basisternum. The derivation of this muscle remains unclear.
This muscle may be the same as op-p 3.
op-s 2 (67 in Chauliodes) connects the profurcal arm and the dorsolateral portion
of the postocciput. This muscle is present also in Ephemeroptera, Mecoptera, Hymenop-
tera, Trichoptera, and Lepidoptera. op-s 2 may be homologous with 55 in Lepismachilis,
which extends between the prothoracic endosternal tendon c and the postgenal sclerite.
t-s( cv) 1 arises dorsally from the dorsal cervical sclerite or its equivalent, or from
the dorsolateral cervical membrane, or from the anterolateral part of the pronotum. It
is inserted ventrally on the posterior tentorial bridge (e.g., Periplaneta) or on the ventral
occipital ridge (e.g., Sialis), or on the anterior end or area of the lateral cervical sclerite
(Orthoptera), or on the dorsal surface of the apodeme of the lateral postgular sclerite
(Corydalus). This muscle is 7 of Alicata (1962b) in Orthoptera.
The serial homologue in the posterior segments is also t-s 1, which connects the
phragma or the anterolateral angle of a segment and the furca of the preceding segment.
This is the derivative of 42 or 46 or both in Lepisma (Fig. 19).
t-s( cv) 9 arises from the lateral antecosta of the mesothorax or the first phragma
and is inserted on the tentorium (e.g. Sialis), or on the ventral process of the postoccipital
ridge (Myrmeleon). This muscle occurs in Neuroptera, Coleoptera, Trichoptera,
Lepidoptera, Hymenoptera, Homoptera and in the larval pterothorax of Hexagenia.
67 in Corydalus, which extends between the intersegmental dorsite and the posterolateral
dorsal portion of the jugulare, may be a modified t-s(cv) 9. t-s(cv) 9 is most probably
homologous with muscle 44 in Lepisma (Fig. 19).
t-cv 1 arises dorsally from the anterior area of the pronotum and is inserted on the
posterior end of the lateral cervical sclerite. This muscle runs subvertically and occurs
in many orders. In Corydalus, t-cv 1 is probably represented by muscle 66, which arises
from the midlateral portion of the pronotum and is inserted on the posterolateral portion
of the jugulare.
t-cv 1 is probably the derivative of 48 in Lepisma (Fig. 19), which is inserted on
the posterior margin of the presternum in the prothorax. Since 87 (Fig. 21) and 48
(Fig. 19) in Lepisma might have been produced by a secondary split of originally a
single muscle, and since 87 can be homologous with t-p 5, t-cv 1 can also be a serial
liomologue of t-p 5.
t-cv 2, 3 arise posterodorsally from the medial area of the pronotum posterior to the
origin of t-cv 1, and is inserted on the medial area of the lateral cervical sclerite or, when
MATSUDA: TilE INSECT TiiORAX 59
the latter consists of two parts, on the posterior part of the anterior lateral cervical
sclerite" ( t-cv 2) and on the anterior end of the anterior lateral cervical sclerite ( t-cv 3).
t-cv 2 may consist of two or more bundles. These muscles are probably homologous
with 50 and 54 in Lepisrnachilis.
cv-s 1 connects the prothoracic furca and the posterior part of the posterior lateral
cervical sclerite. This muscle occurs in many orders, but its derivation is unclear.
cv-s 2(x) (54 in Dissosteira) connects the anterior lateral cervical sclerite and the
prothoracic furca on the opposite side. The derivation of this muscle is unclear.
cv-s 3 (73 in Corydalus) connects the anterior margin of the basisternum and the
lateral face of the jugulare. Its derivation is unclear (not figured).
cv-s 4 is 3 in Siphlonurus and is a common muscle in Hymenoptera; it connects the
prothoracic furca with the anterior end of the lateral cervical sclerite (Siphlonurus), or
with the anterior end of the cervicopleuron (Hymenoptera). In Hymenoptera this
muscle may be a modified cv-s 1. In the nymph of H exagenia 1 VIm 2 ( cv-s 4) is a
furcotentorial muscle. cv-s 4 in Ephemeroptera, therefore, can be a derivative of s 1.
In Lepisrnachilis 28 agrees in position with the nymphal muscle 1 Vlm 2 in Hexagenia.
cv-cx 1 (x) connects the trochantin or the anterior coxal margin with the anterior
lateral cervical sclerite on the opposite side. This muscle occurs in Blattaria, Mantodea,
Orthoptera, Psocoptera, Hymenoptera, Mallophaga, and Mecoptera (not crucial in
Panorpa and Boreus). It is probably homologous with 38 in Lepisrna (Fig. 20),
although in the latter the posterior point of attachment is on the coxal process.
cv-cx 2(x) ( 49 in Grylloblatta) connects the anterior coxal margin and the posterior
lateral cervical sclerite on the opposite side. This muscle is probably a modified
cv-cx 1 ( x) (not figured) .
cv-cx 3 (87 in Corydalus) connects the dorsal portion of the jugular apodeme and
the apodeme arising from the membrane at the anterior midportion of the coxal base.
This muscle may also be a modified cv-cx 1 (x).
cv-p 1 (64 in Forficula and 14 in Anisolabis and Labidura) connects the epimeron
of the prothorax and the anterior end of the lateral cervical sclerite. This muscle may be
a derivation of t-s ( cv) 9 (not figured).
cv-cv 1 (27 in Papilio) connects the extremity of the posterior ventral cervical
sclerite and the posteroventral surface of the lateral cervical sclerite. This muscle may
be a modified cv-s 3 (not figured) .
s 1, 2 arise from the profurca and are usually inserted on the tentorium anteriorly,
but sometimes on the ventral postoccipital ridge (Siphlonurus) or on the postgula
( Corydalus), etc. The one attached more medially is s 1, and the one which lies lateral
to s 1 is designated ass 2. These muscles are homologous with two of 12-16 in Lepisrna
(Fig. 19).
s 3 (72 in Megacrania) connects the anterior end of the prothoracic spinasternum
and the tentorium. This muscle is, therefore, probably serially homologous with s 15 in
the pterothorax, and is homologous with 11 in Lepisrna (Fig. 19).
s-ex( cv) 1 connects the tentorial bridge and the basicosta of the front leg in Lepidop-
tera. In Mallophaga s-ex 7 occurs in a corresponding position of the meso- and meta-
thorax, perhaps by convergence. 36 in Lepidocarnpa and 81 in Lepisrnachilis (Fig. 35)
arise from the labial spina and are inserted on the prothoracic coxal margin, but they are
probably not homologous.with s-cx(cv) 1 in Lepidoptera.
p-s( cv) 7 in the cervical region is represented by a muscle, which connects the
interpleurite between the prothorax and the mesothorax with the tentorium in larval
Myrmeleon (0 vlm 3). 62 in adult Corydalus, which connects the ventral margin of the
intersegmental pleurite between the prothorax and the mesothorax with the lateral margin
of the postgula, also probably represents this muscle. s 3 and p-s(cv) 7 may be the
same, and p-s( cv) 7 is probably homologous with 11 in the cervical area of Lepisrna
(Fig. 19).
Tergal Muscles
(Fig. 23A)
t II is a slender muscle connecting the lateral sides of the first and second phragmata
in Chauliodes (104) and larval Corydalus ( 117). In adult Corydalus 115 connects the
lateral base of the prescutum and is inserted on the anterior projection of the latero-
phragma and it may also be t 11. This muscle lies lateral to t 14 and itsposterior point
of attachment is distinctly separated from t 14.
This muscle may be a derivative of 23 in Lepismachilis; in Collembola also, as Maki
(1938) showed, a lateral dorsal longitudinal muscle is present.
t I2 is the lateralmost oblique dorsal muscle that extends between the ventral end of
the phragma and the posterolateral area of the tergum. This muscle may often be
difficult to distinguish from t 13, when either t 12 or t 13 alone is present. It is usually
considered as an indirect flight muscle and is the antagonist of t 14 functionally; it is
strongly developed in Homoptera. This muscle is probably a derivative of 7, 13 or 15,
24 in Lepismachilis.
t I3 is another posterior oblique dorsal muscle and is usually the antagonist of t 14
functionally. It lies dorsal to t 12 and in Neuroptera its anterior point of attachment is
on the internal ridge of the scutoscutellar suture. This muscle is probably homologous
with 6 in Lepisma (Fig. 19).
t I4 is the important indirect flight muscle that extends between the phragmata in
the pterothorax. In Blattaria it is single and relatively weakly developed, and such a
state probably represents a primitive condition in the Pterygota. In strong fliers t 14 is
represented by several bundles of muscle fibers. This muscle is often only weakly
developed or absent in wingless forms, such as Phasmida. It is, however, present in
wingless Grylloblatta and is strongly developed in Siphonaptera. Its mode of formation
during the postembryonic development varies in higher insects (p. 86).
A probable homologue in Lepisma is 5, and possibly 4 in Lepisma is also a homo-
logue (Fig. 19). In Ctenolepisma (Tiegs 1955) there are three dorsal longitudinal
muscles that correspond in position to t 14.
t 15 ( 65b in Perla) is a small muscle which runs at the base of the scutoscutellar
ridge between the scutum and scutellum. The derivation of this muscle is unknown
(not figured).
t 16 is an unpaired transverse muscle in Lamarckiana (Orthoptera). It passes
transversely across the median lobe of the definitive scutellum, and is attached to it at
either side. This is present in the mesothorax but sometimes absent in the metathorax,
and it regulates the curvature of the tergum. The derivation of this muscle is unclear.
t I7 is an intersegmental muscle 1 dlm in the nymph of Ephemerella and adult
Centroptilum (Ephemeroptera); it connects the second phragma and the anterior margin
of the second abdominal tergum. This muscle is probably homologous with 1 in the
metathorax of Lepisma (Fig. 19) (not figured).
t I8, a transsegmental muscle in Odonata (2 of Odonata), connects the first and
second tergal apophyses. This muscle is probably a serial homologue of 9 in Lepis-
machilis, which connects meso- and meta-thoracic pseudophragmata (not figured).
t I9, a transsegmental muscle in Odonata ( 25 and 45 of Odonata), connects the
tergal apophyses 3 and 4 with the anterolateral areas of the tergum of succeeding
segments. This muscle is probably homologous with 11, 12, 17, and 23 in Lepis-
machilis, which connect the pseudophragmata with the anterolateral margins of the
tergum of the succeeding segments (not figured).
Tergopleural Muscles
(Fig. 23A, B)
Many of the tergopleural (t-p) muscles connect the lateral area of the tergum and
the dorsal area of the pleuron; they have probably been produced newly in the Pterygota,
s 19 )
2abd. seg.
1 abd. sea.
B
FIG. 23. Muscles in pterothorax. A, tergal, tergopleural, pleural, and sternal muscles; B,
tergopleural and pleurosternal muscles (original).
62
with increased connection between the tergum and pleuron (which occurred with. acquisi-
tion of wings). The derivation of many of them is, therefore, unclear.
Some of the tergopleural muscles are inserted ventrally on the interpleurite. Since
the interpleurite may, as already discussed (p. 19), represent the isolated lateral part of
the presternum, these muscles may be classified as pleurosternal, upon elucidation of the
origin of the interpleurite in the future.
t-p 1 (84 in Corydalus) connects the posterolateral area of the pronotum and the
dorsal extension of the interpleurite between the pronotum and the mesonotum. In
larval Trichoptera 5.31 of Tindall (1963) is also t-p 1.
t-p 2 connects the posterolateral portion of the pronotum and the small interpleurite
(more dorsal) between the pro- and meso-thorax (85 in Corydalus), or the interpleural
membrane just in front of the mesepisternum (58 in Dissosteira). This muscle lies
posterior to t-p 1 in Corydalus.
t-p 1 or t-p 2 is probably homologous with 111 in Lepisma (Fig. 21) and with 66
and 77 in Lepismachilis.
t-p 3 is inserted posteriorly on the anterior dorsal margin of the episternum or on the
basalare. The dorsal point of attachment is the first prealar sclerite near the phragma
(Chauliodes), or the phragma (Blattaria, Isoptera), or the anterolateral end of the
scutum or the prescutum (Grylloblatta) etc. In Siphlonurus 44 may represent this
muscle.
t-p 3 is one of the persistent t-p muscles in the Pterygota, but its derivation is
unclear. As already noted, this muscle is serially homologous with op-p 1, which, in
turn, is homologous with a prothoracic muscle 89 in Lepisma (Fig. 21). In Lepisma,
however, there is no exact serial homologue of 89 in the posterior thoracic segments.
t-p 4 connects the subtegula (Trichoptera, Lepidoptera), or the 1aterophragma
(Coleoptera), or a detached muscle disc (Dermaptera), or the prealar sclerite (Psocop-
tera, Neuroptera) with the pleural ridge just below the pleural wing process or with the
pleural arm. This muscle occurs in many orders. Its derivation remains unclear.
t-p 5, 6 are important flight muscles extending dorsally from the anterior part of the
tergum to the ventral part of the preepisternum, or to the lateral part of the basisternum.
Primitively, these muscles were represented by a single muscle; when represented by two
or more bundles, the anterior one is designated as t-p 5, and the posterior one as t-p 6.
These muscles are absent in weak fliers or non-fliers such as Dixippus, Anisolabis,
etc. In Gryllus the only muscle that belongs to the category of t-p 5, 6 is a mesothoracic
tergo-preepisternal muscle which represents t-p 5. In Dissosteira, two bundles of dorso-
ventral muscles in both segments of the pterothorax represent t-p 5 and t-p 6, respec-
tively. In higher orders t-p 5 and t-p 6 are often strongly developed.
In immature stages of lower hemimetabolous insects (e.g., Hexagenia, Knox 1935;
Perla, Wittig 1955) t-p 5, 6 are represented by a single, slender and weak muscle.
During the final stage of nymphal development it becomes greatly enlarged and splits
into two or more muscles. The multiplied bundles of muscle fibers can now be identified
as t-p 5 and t-p 6. At the same time, they tend to become more vertical in position, and
the dorsal points of their attachment (especially of t-p 6) occupies a more posterior
position on the tergum. Thus, it is evident that at least t-p 6 is a newly acquired muscle
in the Pterygota. In higher holometabolous orders, the rudiments of t-p 5, 6 are often
merely a group of myoblasts in the earlier stages of postembryonic development. The
powerful t-p 5, 6 in the adult are produced through various developmental processes
during later stages of development, as shown in discussing the muscle development in
each order.
t-p 5 in the Pterygota may be homologized with 87 in Lepisma (Fig. 21), which
arises from the anterior part of the tergum and is inserted on the ventral part of the
anapleuron.
t-p 7 connects the anterolateral scutal area or the prescutal area with the lower part
of the basalare. This muscle occurs in Plecoptera, Mantodea, Phasmida, Diptera,
63
Trichoptera, and Lepidoptera. t-p 7 may be difficult to distinguish from t-p 3 in some
cases. :Thus, in Mecoptera t-p 3 is present and t-p 7 is absent, whereas in Diptera
apparently t-p 7 is present and t-p 3 is absent. In these orders, therefore, t-p 7 may
appear to be a modified t-p 3, or vice versa. In Trichoptera and Lepidoptera, however,
t-p 3 and t-p 7 occur together, and it is clear that t-p 7 should be recognized as a muscle
distinct from t-p 3. In Plecoptera, as Wittig ( 1955) showed, t-p 3 and t-p 7 are
produced by a secondary split of a muscle, which may be considered as t-p 3.
t-p 8 connects the anterior margin of the basalare or of the episternum with the
lateral scuta] area anterior to the anterior notal wing process. t-p 8 occurs in
Ephemeroptera, Psocoptera, Phasmida, Coleoptera, Homoptera, Neuroptera, Mecoptera,
and Diptera. This muscle may be a modified t-p 7, since when t-p 7 is present t-p 8 is
usually absent, and vice versa.
t-p 9 ( 45 in Tipula) arises from the apodeme on the lower part of the basalare and
extends deeply into the scutum dorsal to the anterior notal wing process. In Tabanus
32, which is also t-p 9, is dorsally attached to a hooklike process by a long tendon.
Apparently this muscle is peculiar to Diptera, and may have been derived from t-p 8
through a secondary split. In Tipula both t-p 9 ( 45 of Mickoleit, 1962) and t-p 8 ( 46
of Mickoleit, 1962) occur together.
t-p 10 usually connects the pleural arm with the anterior notal wing process in many
orders, including Ephemeroptera and Blattaria. In Stenopsocus, Trichoptera, Lepidop-
tera, Mecoptera and Diptera t-p 10 is dorsally attached to the first axillary sclerite, and
this point of attachment is probably secondary; in Diptera it is represented by two
muscles. According to Mickoleit (1966), the presence of this muscle in Trichoptera,
Lepidoptera, Mecoptera and Diptera is a synapomorphic character of the Mecopteroidea;
in the sister group, Neuropteroidea, it is absent. Mickoleit further thought that this
muscle was derived from t-p 12 (hinterer Tergopleuralmuskel of Mickoleit) by
shift in the point of dorsal attachment. In Panorpa, however, both t-p 10 and t-p 12 are
present. This muscle may possibly be the derivative of 65a, 74 or 116, 121 in Lepisma-
chilis, which connects the dorsal face of the endopleural process and the tergum,
roughly at the level of paranotal duplication.
t-p 11 connects the anterior (or dorsal) part of the pleural ridge (144 in Forficula)
or the anterior part of the pleural wing process (W 3 in Stenopsocus) with the first
axillary sclerite. This muscle may be a more dorsally located t-p 10, although in
Psocoptera both t-p 10 and t-p 11 occur together.
t-p 12 usually arises from the pleural arm and is inserted on the lateral margin of
the scutum between the anterior notal wing process and the posterior notal wing process.
This muscle occurs in many orders. It may be confused with t-p 15 which is inserted on
the posterior notal process, but they are distinct, the two muscles being present together
in some orders. t-p 12 may possibly be the derivative of 116, 121 in Lepismachilis.
t-p 13 connects the dorsal episternum ( anepisternum) and the third axillary sclerite.
In Siphlonurus this muscle (51) is inserted on the second axillary sclerite; it is one of the
flexors of the wing in many insects. The derivation of this muscle is unclear.
t-p 14 connects the dorsal part of the pleural ridge near the pleural wing process, or
the pleural wing process itself, with the third axillary sclerite. In Forficula t-p 14
(112, 146) arises from the pleural arm, and in Psocus it (28, 51) arises from the middle
of the pleural ridge. The attachment of this muscle on the pleural ridge or arm makes
this muscle distinct from t-p 13, which arises from the dorsal part of the anepisternum.
The derivation of t-p 14 is unclear.
t-p 15 connects the middle of the pleural ridge and the posterior notal wing process.
In M egacrania t-p 15 ( 115, 154) connects the dorsal portion of the pleural ridge with
the fourth axillary sclerite, which is the detached posterior notal wing process. This
muscle occurs also in Ephemeroptera, Plecoptera, Blattaria, Phasmida, Diptera and some
other orders. In Ephemeroptera this muscle is possibly a modified t-p 14, since when
t-p 14 is present t-p 15 is absent, and vice versa.
64 MEMOIRS OF THE ENTOMOLOGICAl_ SOCIETY OF CANADA
t-p 16 connects the posterior end of the subalare and the posterior dorsal part of the
epimeron. In Gryllus t-p 16 (pm 7) arises dorsally from a small sclerite posterior to the
subalare proper. This muscle is present in lower groups of the Pterygota. In larva(
Dytiscus II lOg of Speyer (1922) is the tergal remotor of the pleuron that arises from
the posterolateral area of the pleuron ( epimeron). It is probable that this tergal
remotor of the pleuron becomes t-p 16, with detachment of the subalare from the tergum.
In the nymph of Perla, according to Wittig ( 1955), this muscle is inserted on the posterior
membrane near the area where the subalare (Fliigelscheide) is formed later.
In a machilid, Pedetontus, as Maki (1938) showed, the posterior tergopleural
muscles ( 10, 11 of Maki) are present in the prothorax, but it is not certain whether
t-p 16 is the derivative of these muscles. In Lepismachilis 115 and 120 agree approxi-
mately with t-p 16 in position.
t-p 17 connects the phragma or the anterolateral angle of the tergum and the inter-
pleurite below it. This muscle is present in Neuroptera and Plecoptera; in Embioptera
the apparent homologues (37, 67) arise from the prealar sclerite. It is possibly the
derivative of 41 in Lepisma (Fig. 19).
t-p 18 (145 in Megacrania) connects the anterior notal wing process with the
posterior margin of the basalare. This muscle may be a modified t-p 10.
t-p 19 (pm 16 in Panorpa and 35, 65 in Neopanorpa) connects the pleural ridge or
the pleural arm with the subalare. In Diptera this muscle is absent, although present in
Hymenoptera. The derivation of t-p 19 is unclear.
t-p 20 is a muscle in Thysanoptera (M.p!.-terg.condyl. in Phloeothrips and Thrips)
that connects the acrotergite with the pleural wing process. Similar muscles are 53 and
54 in the nymph of Pseudococcus (Makel1942), which connect the mesothoracic pleural
arm and the posterior pronotum. This muscle may be a derivative of t-p 4 (not figured).
t-p 21 (34 and 56 in Odonata, M. gubernator ana/is 2 of Tannert 1958) extends
between the interpleural ridge near the metathoracic spiracle and the ventral posterior
margin of the axillary plate (Fig. 170A). The derivation of this muscle remains unclear.
t-p 22 (21 and 43 in Odonata, M. gubernator costae of Tannert, 1958) originates
from the preepisternal sclerite and is inserted on a small cap-tendon arising from the
dorsal intersegmental ridge between the mesothorax and metathorax; the cap-tendon, in
turn, is inserted into the humeral plate (Fig. 169D). The derivation of this muscle is
unknown.
t-p 23 (22 and 44 in Odonata, M. depressor anterior of Tannert, 1958) arises from
the preepisternal apodeme and is inserted on the anterior cap-tendon, which is in turn
connected with the medial area of the humeral plate (Fig. 169D). Maki (1938) thought
that t-p 22 and t-p 23 were sternobasalar muscles, but they should be regarded as tergo-
pleural. In position this muscle is similar to p 3 in other insects, but its exact dorsal
point of attachment is tergal. Furthermore, t-p 23 is present from the first nymphal
stage on, while p 3 is formed during the pupal stage.
t-p 24 (32 and 54 in Odonata, M. depressor medianus of Tannert, 1958) arises from
the ventral portion of the epimeron, and is inserted on the posterior cap-tendon (or
FZUgelgelenkfortsatz der R + M Ader of Tannert, Fig. 172B). Despite its superficial
similarity to t-p 19, this muscle is probably not homologous with t-p 19, since the
posterior cap-tendon in Odonata is probably not homologous with the subalare, to which
t-p 19 is attached. The derivation of this muscle is unclear.
t-p 25 (33 and 55 in Odonata, M. depressor posterior of Tannert) arises from the
ventral portion of the epimeron and is inserted on the ventral posterior margin of the
l!J(illary plate (Fig. 170A). The derivation of this muscle is unknown.
t-p 26 (31 and 53 in Odonata, M. gubernator ana/is la, b of Tannert, 1958) arises
from the posterior surface of the pleural wing process and is inserted on the proximal
part of the A vein in the axillary plate. The derivation of this muscle is unknown (not
figured).
MATSUDA: ffiE INSECT THORAX
65
t-p 27 is .a larval muscle ( 139 in Corydalus) which passes anteroventrally from its
origin on the midlateral portion of the scutum to an insertion on the anterodorsal margin
of the invaginated episternum. This muscle is a relatively robust one that becomes lost
in the pupa and adult. In position, t-p 27 agrees roughly with 87 and 88 in Lepisma
(Fig. 21) (not figured).
t-p 28 is a larval muscle ( 140 in Corydalus) which passes ventrally from the antero-
lateral portion of the scutum to an insertion on the invaginated episternum laterad of the
insertion of t-p 27 (139). In larval Limnephilus mesothoracic 7.13-7.16 also represent
the t-p 27, 28 group. t-p 28, together with t-p 27, are probably homologous either with
87 or 88, or with both (not figured).
t-p 29 is a larval muscle ( 147 in Corydalus) which passes ventrally from its origin
on the posterolateral margin of the pronotum to an insertion on the anterior margin of
the episternum (the precoxal bridge of Kelsey, 1957). The derivation of this muscle is
unclear (not figured) .
t-p 30 is a larval muscle (5.34 in Trichoptera, and plom 1 in Psychoda), which
connects the anterior (or median) lateral part of the tergum and the posterior inter-
segmental pleurite. In larval Thymalus (Coleoptera) Breed ( 1903) described a similar
muscle which extends obliquely across the whole metathorax.
In Ctenolepisma the second oblique muscle of Tiegs (1955) passes from the spina
to the preceding tergum, and is similar to this muscle in position. In Lepidocampa
Maki's (1938) 57 extends from the anterior dorsolateral region of the mesonotum to the
posterolateral basal wall of the middle coxa; in position this muscle is, therefore, similar
to the above larval muscles in the Pterygota. In nymphal Ephemeroptera an oblique
intersegmental dorsoventral muscle (idvm) in the abdomen (Diirken 1907, 1923) is
most probably a serial homologue (not figured).
Tergosternal Muscles
(Fig. 26)
t-s 1 arises from the phragma and is inserted ventrally on the furca of the preceding
segment in many orders. It is clearly homologous with 46 in Lepisma (Fig. 19).
t-s 2 (75 in Corydalus) originates on the midlateral portion of the mesothoracic
prescutum, laterad of the phragma, and is inserted on the posterodorsal surface of the
prothoracic furca. In the pterothorax of Corydalus the homologues are retained only
during the larval stage. 36 in Stagmomantis, which is inserted on the posterolateral
region of the prosternum, may also represent t-s 2.
t-s 2 is probably homologous with 42 in Lepisma (Fig. 19). As discussed later,
p-s 12 in Ephemeroptera may also be homologous with 42 in Lepisma and, hence,
with t-s 2.
t-s 3 is a direct flight muscle which connects the tip of the furca and the first or
second axillary sclerite in Ephemeroptera (Fig. 44B). In larval Corydalus muscle 134,
which probably represents t-s 3, arises dorsally from the midlateral anterior portion of
the scutum and is inserted on the anterior dorsal half of the furca. II 11a in larval
Dytiscus (Speyer 1922) also represents t-s 3. Among the adult Pterygota, this archaic
muscle has been retained only in Ephemeroptera. t-s 3 is probably homologous with 45
in Lepisma (Fig. 19).
t-s 4 is a subalar-furcal arm muscle in Ephemeroptera (Fig. 44B). In Siphlonurus
39 ( t-s 4) arises from the. subalar apodeme and is inserted on the tip of the mesofurca.
Homologous muscles occur also in the metathorax of Siphlonurus and possibly' in
Euthomostethus (Hymenoptera); its derivation is unclear.
t-s 5 is a large muscle in Ephemeroptera, which arises from the subalare and is
inserted on the furcasternum behind the middle coxa (Fig. 44B). This muscle is present
also in the metathorax of Siphlonurus, and may be a derivative of the subalar-coxal
(meral) muscle (t-cx 8).
t-s 6 (Ill Dvm 4 in Hexagenia) connects the scutellum and the 'furca. t-s 6 may
possibly be a displaced t-s 1 that has changed its dorsal point of attachment.
t-s 7 (24, 50 in Ecdyonurus) originates from the anterolateral portion of the tergum
and is inserted on the anterolateral corner of the sternum; this muscle may be present
also in Phasmida. t-s 7 may be homologous with 48 and 68 in Lepidocampa (Mald
1938) which arises from the anterior transverse ridge of the tergum and is attached to
the lateral end of the presternum; t-s 7 may also be homologous with 48 in Lepisma
(Fig. 19), although the latter is a prothoracic muscle.
t-s 8 (24 in Perla) connects the mesothoracic antecosta and the prothoracic spina.
13 of Mantis, which connects the first phragma and an area near the prothoracic spina,
also represents t-s 8. t-s 8 may be homologous with 41 in Lepisma (Fig. 19).
t-s 9 occurs more often in the cervico-prothoracic region. In nymphal Hexagenia
II Dim 2 connects the second phragma and the profurca and is obviously a serial homo-
logue of t-s( cv) 9. In the pterothorax of adult pterygotes this muscle has apparently
been lost. This muscle appears to be the derivative of 44 in Lepisma (Fig. 19) (not
figured).
t-s IO (23, 46 in Odonata) extends between the anterolateral part of the scutum near
the tergal apophysis (Fig. 169B) and the internal basisternal skeleton (the prefurca). In
the prothorax of Corydalus muscle 79, which passes posteroventrally from its origin on
the anterolateral portion of the pronotum to an insertion on the posterolateral portion of
the basisternum, may possibly represent this muscle. This muscle has often been con-
fused with t-p 5 or 6 (Berlese 1909b; Tillyard 1917; Chao 1953, etc.); but it is a
genuinely tergosternal muscle from the first nymphal stage on, whereas t-p 5 (and 6) is
primarily a tergopleural muscle. t-s 10 is probably homologous with 64 in Lepisma-
chilis, which connects the pseudophragma and the endosternal tendon h.
t-s II (23', 46' in Odonata) is a fine elevator muscle of the wing. It arises from the
prefurca and is inserted on a notal wing process (Ventralausliiufer des Scutum of
Tannert, 1958). The derivation of this muscle remains unclear (not figured).
Tergotrochantinal (coxal) Muscles
(Fig. 24A)
t-ti(cx) I arises from the anterior edge of the tergum or the phragma, and is
inserted on the trochantin near the point of trochantinocoxal articulation. This muscle
occurs in Blattaria, Orthoptera, Isoptera, Dermaptera, and Coleoptera and is most prob-
ably homologous with 71 in Lepisma (Fig. 20).
t-ti(cx) 2 arises usually from the anterior portion of the tergum behind t-ti 1, and is
inserted on the apodeme from the anterior margin of the trochantin. It may be inserted
on the anterior coxal margin in the pro thorax of some forms (e.g., 79 in F orficula).
t-ti(cx) 2 is probably homologous with 72 in Lepisma (Fig. 20).
t-ti(cx) 3 arises, in Periplaneta (120, 163) and Mantis (47, 75), from the middle of
the tergum, and is inserted on the apodeme from the trochantinal edge on which t-ti 2 is
also inserted. It is difficult to distinguish t-ti 2 from t-ti 3 or vice versa, when only one
of these muscles is present. t-ti(cx) 3 may be homologous with 73 in Lepisma (Fig. 20),
although the ventral point of attachment of the latter does not agree with the typical
point of attachment of t-ti 3 in the Pterygota, which lies more dorsad.
t-cx 4 is a direct flight muscle in Ephemeroptera (Fig. 44B) and Odonata. In
Siphlonurus (50, 78) t-cx 4 connects the second axillary sclerite and the coxal process,
and in Ecdyonurus it is inserted on the first axillary sclerite. In Odonata there are two
muscles that connect the disc internal to the coxal condyle with the first axillary sclerite
and with the axillary plate (second and third axillary sclerites). Since in Odonata they
arise ventrally from the same disc, and since their dorsal points of attachment are very
close, they are designated as t-cx 4 and t-cx 4', respectively. t-cx 4 appears to be homo-
logous with 62 in Lepisma (Fig. 20).
67
0
B
FIG. 24. Muscles in pterothorax. A, tergocoxal muscles; B, pleurocoxal and sternocoxal
muscles (original).
t-cx 5 ( 62, 87, and 117 in Grylloblatta) is a tergocoxal muscle which originates "from
the anterior half of the tergum. Running obliquely caudad from the origin, it is ipserted
on the posterior part of the coxal base. This muscle occurs also in the metathorax of
Gryllus and in the prothorax of Blattaria and Neuroptera. t-cx 5 is probably homo-
logous with 61 in Lepisma (Fig. 20), which, however, is inserted on the coxal process.
The muscle has probably shifted its attachment onto the posterior coxal margin, with
reduction of the coxal process in the Pterygota.
t-cx 6, 7 are posterior tergocoxal (meral) muscles which have often been described
as tergal remotors. The primitive number of these muscles is rather difficult to deter-
mine. In holometabolous insects including Neuroptera, Mecoptera, Diptera, Trichoptera
and Lepidoptera, Larsen (1948) and Mickoleit (1966) recognized only two posterior
tergocoxal muscles. In Ephemeroptera, Corydalus and Chauliodes, etc., however, there
are three such muscles, and the presence of an apparently extra muscle in these insects
contradicts the idea that primitively there were two posterior tergocoxal muscles.
In these lower forms, however, two of the three definitive muscles can be counted as
representing a single muscle that has secondarily split into two. For example, 159a and
159b in Corydalus lie so close together in their dorsal and ventral points of attachment
that they could well be counted as representing a single muscle. In fact, this is
apparently the way Mickoleit ( 1966) counted the number of muscles; he recognized only
two posterior tergocoxal muscles in Coryda/us. Provisionally, therefore, the primitive
number of the posterior tergocoxal muscles is considered as two.
In holometabolous insects, as Larsen (1948) and Mickoleit (1966) have shown, the
two posterior tergocoxal (meral) muscles are the inner tergal remotor and the outer
tergal remotor. The latter often lies lateral to the former (Fig. 25); these muscles are
designated as t-cx 6 and t-cx 7, respectively.
Since, according to Larsen (1948) and Mickoleit (1966), t-cx 7 has increased its
function as a flight muscle, it has increased its size and probably has split into two bundles
of muscle fibers (as discussed above) ; the muscle is retained in all holometabolous orders
they have examined. t-cx 6, on the other hand, has undergone progressive reduction and
has been completely lost in Mecoptera, Diptera, Trichoptera, and Lepidoptera.'" Pre-
sumably, a parallel loss of t-cx 6 has occurred in other higher orders. When only one
of the two posterior tergocoxal muscles is present, therefore, the muscle may often
represent t-cx 7.
The origin of t-cx 6 and 7 may be traced to 63, 64, and 65 in Lepisma (Fig. 20) or
to 103, 104, 107, 108 in Lepismachilis.
t-cx 8 is a subalar-coxal (meral) muscle which occurs in many insects, and is a
flight muscle. Based on the evidence given below, this muscle is treated as a tergocoxal
muscle. Maki (1938) has shown that this muscle, which later becomes the subalar
coxal muscle, is attached dorsally on the lateral margin of the tergum in the late embryo
of Leucophaea (Blattaria), or on the ventral base of the rudimentary wing in the early
nymphal stages of Locusta (Orthoptera). At the last molting, the area where the
muscle is attached becomes the isolated subalare by membranization of the surrounding
area, and thus, in the adult, the muscle is the subalar-coxal muscle. Storch and
Chadwick ( 1967) also found that in Periplaneta this muscle is a tergal remotor of the
coxa during the late embryonic and nymphal stages. In Dysdercus (Heteroptera)
Khanna ( 1963c) found that during the fourth nymphal stage a tergocoxal muscle splits
in two bands; with formation of the subalare during the final molt, the anterior band
shifts its dorsal attachment onto this subalare. In Thy malus (Coleoptera) Breed ( 1903)
found that the eventual subalar-coxal muscle is a two bundled tergocoxal muscle to begin
with; during the pupal stage it becomes fused into a single bundled muscle, which
shifts its dorsal point of attachment to the subalare (posterior cupula of Breed).
"'Mickoleit regarded the-loss of t-cx 6 in these orders as a characteristic (synapomorphic) of the Panorpoidea.
BASALARE
p3-------.
p-ex 5 -==tf;
p-ex 3
s-ex 5 -------
PLEURAL WING PROCESS
;;....;:------. s-ex 2

'--------- s-ex 6
FIG. 25. Muscles attached to coxal base in Dissosteira carolina (from Snodgrass).
69
Other facts also favor the idea of the tergal origin of this muscle. Maki ( 1938)
thought that in Labia (Dermaptera) 43 (subalar-coxal muscle) might be homologous
with the external tergal remotor, 42 in the wingless Aniso/abis. In Forficula (Dermap-
tera) Kleinow ( 1966) showed that his muscle 122 in the mesothorax arose dorsally from
the posterior end of the jugal field of the elytron, where the third axillary sclerite has
become indistinguishably fused. In other words, when the small subalare is not well
differentiated, the origin of t-cx 8 ( 122 of Kleinow) lies more dorsad. II dvm 2a in
wingless Dixippus (Phasmida), which arises from the posterolateral area of the tergum,
is obviously homologous with 127 (t-cx 8) in winged Megacrania.
All of the above facts suggest that t-cx 8 arise from the tergum, when the wing is
absent or weakly developed. If this tendency is generally true, the number of posterior
tergocoxal muscles in the prothorax, which lacks the subalare, should tend to equal the
number of posterior tergocoxal muscles plus the subalar-coxal muscle in the pterothorax
of the same species. This tendency is true in lower orders. For instance, in Odonto-
termes (Table VIII) there are three prothoracic posterior tergocoxal muscles, and in the
pterothorax there are two posterior tergocoxal muscles and a subalar-coxal muscle.
The same relationship is seen in Plecoptera, Dermaptera, Phasmida, and Orthoptera
(Dissosteira and Xyphidion). In the prothorax of Blattaria, Isoptera, and Mantodea
(Table VII) the posterior tergocoxal muscles are split in their origin, and this makes it
difficult to obtain an accurate count. However, if the muscle counts suggested in
Table VII are correct, the number of the posterior prothoracic muscles is roughly equal
to the number of the pterothoracic muscles under discussion. An exception appears to
be in Psocus, where the prothoracic posterior tergocoxal muscle is represented by a
single muscle (14). Yet, this muscle is very thick due probably to a secondary fusion
of what were originally two or more muscles. t-cx 8 may be homologous with 62 in
Lepisma (Fig. 20).
Pleural Muscles
(Fig. 23A)
Under the pleural muscles, only those which occur within the pleuron are described.
p 1 usually bridges the anapleural cleft; it is short but can be broad. In Phasmida
this muscle is represented by a row of short vertical muscle fibers that connect the
anepisternal area and the dorsal margin of the definitive basisternum (actual pre-
episternum). p 1 is absent in the larva of Perla (Wittig 1955), but is present in the
larva of Corydalus (Kelsey 1957). The derivation of this muscle is unclear.
p 2 connects the posterior face of the intersegmental pleurite and the anterior face
of the basalare. p 2 occurs in Neuroptera, Trichoptera, Lepidoptera, Mecoptera,
Diptera, Thysanoptera, Homoptera, Embioptera, Hymenoptera, Phasmida, and Plecop-.
tera. This muscle may be homologous with 26 in Lepisma (Fig. 19) which, however, is
a muscle for the spiracle.
p 3 arises dorsally from the anterior part of the anepisternum or the basalare and is
inserted on the ventral part of the preepisternum. It has often been described as a
pleurosternal muscle, because of its insertion on the definitive sternum. However, this
muscle is primarily a pleural muscle. It lies external to the tergopleural muscle ( t-p 5,
6), when the latter is present at the same time.
. p 3 in the nymph of Perla (Wittig 1955) becomes greatly enlarged during the final
molt into the adult; it is also very weak in the nymph of Orthoptera (Wiesend 1957).
Breed ( 1903) found that in the larva of Thymalus it is represented by three or four short
fibers lying immediately above the larval leg, and they probably have connection with the
leg movement. These fibers elongate very rapidly in the pupa. By the time the imaginal
state is attained, the muscle has increased still more in size, and its fibers are so fused as to
show but two parts, which are separated at the ventral end only. The imaginal muscle
thus formed extends from the basalare (large cupula of Breed) backward and downward
to the middle of the lateral expanse of the definitive metasternum, and then, the muscle is
p 3. In the pupa the connection with the leg base becomes lost, and Larsen's ( 1966)
observation that p-cx 2 is absent in the adult Coleoptera supports the above observation
by Breed (1903). Tiegs (1955) also noted that in Acrididae p 3 is formed after it has
lost its association with the leg during development.
In the absence of p 3 in Thysanura and in the light of its peculiar development, it is
clear that p 3 is purely a new muscle acquired in the Pterygota.
p 4 is a pleural muscle in Dermaptera which connects the anterior part of the pleural
ridge and the anapleural cleft. The derivation of this muscle is unclear.
p 5 ( 42 in Siphlonurus) connects the pleural wing process and the basalare.
muscle ism. pleurocondilo scutale of Grandi (1947b, Fig. 44B) and may be t-p 4.
derivation of this muscle is unclear.
Pleuro!lternal Muscle
(Fig. 23B)
This
The
Some of the pleurosternal muscles in the Pterygota are homologous with endosternal
muscles in Thysanura. In Thysanura the posterior dorsal extension of the endosternum
reaches the definitive pleural area internally and muscles are attached to this area
(Fig. 19); in the Pterygota the homologues of some of these muscles are attached to the
interpleural area, which may be presternal in origin. Therefore, these muscles can be
sternal muscles.
p-s 1 connects the pleural arm with the furcal arm. It occurs in many orders, and
may sometimes be represented by a ligamentous strap. In a machilid genus, Petrobius,
Carpentier (1946b) showed a pleural process (p) that connects the pleuron with the
endosternal g (Fig. 39), which appears to be the same as the ligamentous connection
between the pleural arm and the furca in the Pterygota. In Lepisma a prothoracic
muscle 27 (Fig. 20) may represent p-s 1, although this muscle connects the anapleural
tendon and the endosternal process g.
p ~ s 2 connects the spina and the anterior margin of the episternum, or basalare, of
the succeeding segment. This muscle is present in lower orders such as Blattaria,
Isoptera, Gryllob!attodea, Mantodea, Orthoptera, Psocoptera, Phasmida, Embioptera,
Plecoptera, and Neuroptera. The derivation of this muscle is not clear.
p-s 3 (59 in Dissosteira and II pm 17 in Myrmeleon) connects the furca and the
anterior margin of the episternum of the succeeding segment. This muscle occurs also in
Psocoptera, Coleoptera, Hymenoptera, and Lepidoptera. p-s 3 is not a modified p-s 2,
since p-s 2 and p-s 3 occur together (Psocoptera and Orthoptera). This muscle may be
a derivative of p-s 12.
p-s 4(x) is a crucial muscle in Periplaneta and some other cockroaches and
Mantodea (Chadwick 1957); it connects the profurca and the mesepisternum on the
opposite side. No such muscle has been found in the Apterygota (not figured).
p-s 5 is 13, 24, and 31 of Chadwick (1957) in Blattaria, Mantodea, and Isoptera;
it connects the postcoxal membrane with the furca. It is often ligamentous. In larval
Corydalus muscle 177 is p-s 5, as it connects the intersegmental pleurite and the furca of
the preceding segment. p-s 5 occurs also in Psocoptera, Homoptera, Heteroptera, and
Mallophaga. This muscle is similar to p-s 3. The derivation of this muscle remains
unclear.
p-s 6 is 4, 17, 28 of Chadwick (1957) in Blattaria and Isoptera, and 83 in Stagmo-
mantis; it connects the spina and the intersegmental pleurite lying just dorsal to it. p-s 6
occurs also in Orthoptera, Dermaptera, Neuroptera, Mecoptera, and Trichoptera. This
muscle may be homologous with similar muscles in Collembola (36 and 61 in Folsomia
and 46 and 67 in N eanura). This muscle may be confused with p-s 2.
p-s 7 is 1sps-2ils, 2sps-3ils, 3sps-1ils of Chadwick ( 1959b); it connects the spina and
the intersegmental pleurite of the succeeding segment. This muscle occurs in immature
forms of Coleoptera (II, III, 4d of Speyer, 1922), Neuroptera (Ovlm 3 of Korn, 1943;
2sps-1ig. 3 of Chadwick, 1959b), and possibly of Odonata (68 of Maloeuf, 1935a).
Among adult insects, it may be present in Dermaptera (29 in Anisolabis and Labia).
p-s 7 is probably a derivative of 11 in Lepisma (Fjg. 19).
p-s 8 is 2sps-1ils and 3sps-2ils of Chadwick ( 1959b), which connects the second
and third spinae and the respective preceding interpleurites. p-s 8 occurs in larval
Coleoptera and larval Neuroptera.
No homologue is present in Lepisma, but 31 or 32 in Lepismachilis, 47 and 67 in
Lepidocampa may be homologous. As Chadwick (1959b) said, this muscle must repre-
sent a primitive part of the hexapod muscular pattern that began to disappear at a
relatively early date.
p-s 9 (184 in Corydalus) arises from the second intersegmental pleurite and is
inserted on the metafurca. 35 in Mantis and 44 in Stagmomantis have their anterior
points of attachment on the prosternum lateral to the first spina and are inserted on the
mesothoracic furca, and they appear to be homologous with 184 in Corydalus. In
Anisolabis the apparent homologue 57 connects the anterolateral angle of the meta-
sternum (presternum) and the metafurca.
In larval C orydalus 123 and 179 are p-s 9, and 120 ( p-s 8) and 123 are present at
the same time. Therefore, p-s 9 is not the same as p-s 8. This muscle is homologous
with 17 in Lepisma (Fig. 19).
p-s 10 ( 122 in Corydalus) passes from the prothoracic furcasternum to the anterior
face of the second trl:!Ilsverse intersegmental pleurite-spinal tendon. II, 1a vim in
Mallophaga and M. mesfurc.-abdst. in Thysanoptera, which connect the mesofurca and
the first and( or) second abdominal segment, may represent p-s 10. No such muscle is
present in the Apterygota, and hence its derivation is unclear.
p-s 11 (unnumbered muscle in Siphlonurus, 80?) connects the paracoxal ridge and
the furca. No such muscle is known in the Apterygota.
p-s 12 is two paired muscles in Ephemeroptera which originate on the mesothoracic
basalare; one pair is inserted on the profurca, and another pair is inserted commonly on
the posterolateral area of the prothoracic sternum, behind the furca (Fig. 44B). They
act as direct flight muscles of the forewing. .
Possible homologues of p-s 12 are two paired muscles 42 in Lepisma (Fig. 19)
which, however, are attached to the anterolateral area of the mesonotum above the dorsal
anapleural sclerite.
p-s 13 is 189 in Periplaneta and 111 b in Grylloblatta; it connects the second spina
and the anterolateral part of the second abdominal sternum. Chadwick ( 1959b)
thought that this muscle results from the fusion of 2sps-3ils (p-s 7) and 3 ils-Iils which
occurs in the larva of Corydalus. Similarly, according to Chadwick ( 1957), 2sps-llils
can be formed by incorporation of another ils-ils.
p-s 14 (142 in larval Corydalus) connects the furca with the pleural ridge. It is
distinct from 151 which is p-s 1 in the larva. No such muscle is present in the Aptery-
gota (not figured) .
Pleurotrochantinal (coxal) Muscles
(Fig. 24B)
Of the nine pleurotrochantinal muscles, five anterior pleurotrochantinal muscles
(p-ti 1-5) that extend from the episternum to the coxal base are sometimes difficult to
identify. The best criterion for the identification of them appears to be the points of
their ventral attachment, relative to one another, on the coxa or on the trochantin. On
this basis, the episterno-coxal (or trochantin a!) muscles can be classified into two groups:
the anterior three muscles (p-ti 1, 2, 3) that are inserted on the trochantin or the coxa
immediately behind the point of attachment of a tergocoxal muscle t-cx 3, and two other
muscles (p-cx 4, 5) that are inserted on the coxal margin (and never on the trochantin)
lateral to the first three muscles.
p-ti(cx) 1, 2, 3 are often difficult to distinguish. It is especially difficult when only
one or two of these muscles are present. Even when all three muscles are present,
identification of them is sometimes difficult, because of the crowded points of origins and
insertions. In some insects, such as Stagmomantis and Dissosteira (Fig. 25), the
anteriormost pleurotrochantinal muscle originates from the lower anterior margin of the
episternum, and in Psocus muscle e arises obviously from the preepisternal area; in such
cases they can safely be identified as p-t 1.
In the absence of corresponding muscles in Thysanura, it is probable that these
muscles have arisen secondarily as flight muscles, with the acquisition of the wing. At
least in Acrididae, a basalar coxal muscle (p-cx 2 ?) arises from the pleurotrochanteral
muscle by shift of the ventral point of attachment during development (Tiegs 1955).
Furthermore, the development of p 3 in Thymalus (discussed,,on p. 70) is highly sug-
gestive. If the posterior part of the ventrally divided (into two) p 3 in Thy malus
becomes attached to the coxal margin during the final stage of development, it would
become a basalar-coxal muscle (p-ti 2 or 3). p-ti 1 may be of a quite different origin,
which cannot be explained at present.
p-cx 4, 5 are usually the abductors of the leg. p-cx 4 arises usually from the dorsal
area of the anepisternum immediately anterior to the pleural wing process. The origin
of p-cx 5 is on the paracoxal suture (formed internally by the paracoxal ridge), when it
is present; or it may arise from the lower margin of the pleural arm. Both muscles are
inserted directly on the coxal margin anterior to the pleurocoxal articulation.
p-cx 5 is obviously homologous with 101 in Lepisma (Fig. 21). Barlet (1953,
1954) did not describe a dorsal pleurocoxal muscle homologizable with p-cx 4. In the
same species, however, Maki described a pleurocoxal muscle in each thoracic segment
(29, 61, 95), which arises from the dorsal portion of the pleuron and is inserted on the
lateral basal wall of the coxa. It is impossible, however, to discern the exact positions
of these muscles from his smudgy figure. If such muscles really exist in Lepisma, p-cx 4
is homologous with them.
p ~ c x 6 (91, 129, 166 in Megacrania) arises from the pleural arm, and is inserted on
the basal ventral margin of the coxa. 102 in Lepisma (Fig. 21) roughly agrees in
position with this muscle.
p-cx 7(x) (M. postpleuro-coxa/is transversus in Sialis) extends from the lower part
of the postpleural (interpleural) sclerite, and is inserted on the pleural articulation with
the coxa on the opposite side. It is probably homologous with 97 in Periplaneta, which
arises from the internal edge of the mesepisternum and goes to the coxopleural articula-
tion of the coxa on the opposite side. In the larva of Dytiscus the homologue (I 15)
arises posteriorly from the intersegmental pleurite (Furcilla of Speyer, 1922). Accord-
ing to Chadwick (1957), the homologue of p-cx 7(x) in larval Coryda/us arises from the
furcal arm. The derivation of this muscle is unclear (not figured) .
p-cx 8 is muscle 15 in Calobata, 15 in Orthellia, and muscle H in Stenopsocus; it
connects the anterior portion of the mesothoracic episternal region behind the first
spiracle (Diptera) or the metepisternum (Psocoptera) with the posterior basal rim of the
prothoracic coxa. This muscle must act as a remotor of the coxa. Unlike p-cx 7(x),
p-cx 8 is not a crucial muscle, and agrees in position approximately with 135 in
Lepismachi/is.
p-cx 9 arises from the epimeral area, and is inserted on the coxal margin behind the
pleurocoxal articulation. This muscle occurs commonly in the prothorax of Trichoptera
and Lepidoptera. Among immature insects, 162 in Corydalus, III pm 2 in Hexagenia,
and 15 in Plathemis are p-cx 9. p-cx 9 is probably homologous with coxoepimeral
muscle 136 or 137 in Lepismachilis.
Sternal Muscles
(Fig. 23A)
s 11 is the spina-spinal muscle that connects the spina of one segment with that of
the succeeding segment. The occurrence of this muscle is limited to lower orders such
as Blattaria, Mantodea, Psocoptera, Neuroptera, and Orthoptera. There is no exact
homologue of s 11 in Lepisma, although Chadwick ( 1959b) thought that 13 in Lepisma
(36, 44, 54 of Barlet) was homologous. However, 13 in Lepisma is a spina-furcal
muscle. s 11 may be homologous with 38 or 40 in Lepismachilis.
s 12 is a spina-furcal muscle that extends between the spina and the furca of the
succeeding segment. This muscle occurs commonly in lower orders. s 12 is most
probably homologous with 12 in Lepisma (Fig. 19).
s 13 is a furcofurcal muscle that connects furcae of two adjacent segments. It is
represented by three muscles or three bundles of muscle fibers in Corydalus, or in
Blattaria by two or three bundles (Chadwick 1957). In many other orders s 13 is
represented by a single muscle. In Hymenoptera it is well developed between the meso-
furca and profurca, and is a retractor of the propectus (cervicopleuron). In Lepisma
homologous muscles are 15 and 18, and possibly also 14 (Fig. 19).
s 14 is a muscle which extends from the furca to the spina immediately behind it.
Its occurrence is more common in the prothorax than in other segments. This muscle is
homologous with 19 in the prothorax of Lepisma (Fig. 19).
s 15 is a transsegmental furca-spinal muscle that stretches between the furca and the
spina of the succeeding segment. The profurca-second spinal muscle is more common
than the serial homologue in posterior segments. s 15 has been found in Embioptera,
Blattaria, Isoptera, Psocoptera, Mallophaga, Thysanoptera, Dermaptera, Orthoptera, and
Grylloblattodea. The derivation of this muscle is not too clear, but 14 of Lepisma
(Fig. 19) may be its homologue, as Chadwick (1959b) suggested. Another possible
homologue is 20 in Lepisma (Fig. 19).
s 16 connects the furcal arm on the two sides of the same furca; it has been found
in Phasmida, Blattaria, Mecoptera, Mallophaga, Heteroptera, and Ephemeroptera.
Chadwick (1959b) believed this muscle to be the derivative of fu-sps muscles (s 12); he
thought that s 16 was formed as a result of loss of the spina on which it was primitively
attached. In Neopanorpa s 16 is present in all segments and this condition is correlated
with the absence of the spina. Daly ( 1964) observed in A pis that during t h ~ post-
embryonic developments 12 becomes s 16; apparently, therefore, Chadwick's suggestion
was correct (not figured).
s 17 (9 in the prothorax of Oligotoma) connects the anteromedian portion of the
furcasternum (sternellum of Maki, 1938) and the furcal arm. It is probably homo-
logous with 21 of Lepisma (Fig. 19), which occurs also only in the prothorax (not
figured).
s 18 ( 110 in C hauliodes) arises from the median base of the furcal arm. It divides
into two branches, one extends forward and the other backwards, and joins in the
formation of a common ventral muscular net. The presence of the homologous muscle
in other insects needs to be studied. No such muscle is known in the Apterygota (not
figured).
s 19 (II Vim 1 in nymphal Hexagenia) connects the profurca and the anterior
margin of the first abdominal segment. In larval Myrmeleon the apparent homologue
(I VIm 1) reaches the sternum of the second abdominal segment; in nymphal Perla the
"atypische ventrale Liingsmuskeln" of Wittig ( 1955) also probably represent this muscle,
although they arise from the tentorium. Among adult insects it occurs only in Odonata.
No such muscle is present in the Apterygota.
s 20 ( 193 in Periplaneta) connects the metafurca and the boundary between the first
and second abdominal segments. In the larva of Thy malus, according to Breed ( 1903),
this is a muscle of the first abdominal segment, and it becomes attached to the forming
metafurca during the pupal stage. This muscle has been found also in Thysanoptera,
Homoptera, Mecoptera, Lepidoptera, etc. The derivation of this muscle remains
unclear. This muscle may be serially homologous with p-s 3.
s 21 (66 in Odonataj connects the metathoracic poststernum and the anterior
process of the second abdominal sternum. The derivation of this muscle is unknown
(not figured) .
s 22 ( 68 in Odonata) connects the posterior end of the metafurcal invagination and
the membrane posterior to the isolated metathoracic poststernum. In the nymph of
Plathemis (68 of Maloeuf, 1935a) this muscle appears to be p-s 7.
s-stg in Heteroptera is the muscle of the metathoracic scent gland. This muscle is
probably a derivative of a sternal muscle.
Sternocoxal Muscles
(Fig. 24B)
s-ex 1 usually connects the spina and the anterior coxal margin of the succeeding
segment. It may be inserted near the pleurocoxal articulation (e.g., Periplaneta), or on
the trochantin near the trochantine-coxal articulation (e.g., Anisolabis and Labia).
s-ex 1 has been retained only in lower orders such as Blattaria, Isoptera, Orthoptera,
Psocoptera, Dermaptera, Grylloblattodea, and Neuroptera. This muscle is homologous
with 31 in Lepisma which connects the spinal area of endosternum (m) and the coxal
process of the succeeding segment (Fig. 20). The point of attachment in Periplaneta is
therefore similar to that of Lepisma.
s-ex 2 connects the tip of the furca and the ridge limiting the meron in Periplaneta
(133, 170). In a termite (fig. 7 of Mickoleit, 1965) the muscle is apparently inserted
on a ridge formed below the ventral end of the pleural ridge. In Gryllus II, III bm 5
connects the tip of the furca and an inwardly directed process on the posterior coxal
margin, immediately behind the pleurocoxal articulation. At least in lower forms, there-
fore, this muscle is a sternocoxal muscle.
In some forms, however, the area of its attachment on the coxal margin has become
indistinguishably continuous with the pleural ridge. In Diptera, for instance, the ventral
end of the pleural ridge is modified into a well-developed process, and s-ex 2 is attached
to it, and thus the muscle appears to be a pleurocoxal muscle.
The position of s-ex 2 in lower pterygotes agrees with 33 in Lepisma (Fig. 20), and
hence they are probably homologous.
s-ex 3 is 3 of Mickoleit ( 1965); it usually arises from the posterior margin of the
furca and is inserted on the posterolateral area of the coxal margin, or on the meron
posterior to the attachment of s-ex 2. In the prothorax of nymphal H exagenia s-ex 3 is
not differentiated from s-ex 2, although they become distinct in the adult. In the pro-
thorax of Lepisma (Fig. 20) 37 lies posterior to 33 (s-ex 2), and in position it agrees
with s-ex 3. Hence, s-ex 3 is probably homologous with 37 in Lepisma.
s-ex 4 connects the spina and the posterior coxal margin or the meron of the
preceding segment. With the loss of the spina, this muscle may become a transverse
muscle that connects the posterolateral area of the coxal margin on both sides (58 in
Siphlonurus). This muscle has been found in Blattaria, Isoptera, Mantodea, Orthoptera,
Grylloblattodea, Neuroptera, Psocoptera, Trichoptera, Lepidoptera, and Hymenoptera.
In higher Hymenoptera this muscle arises posteriorly from the anteromesial area of the
mesosternum. s-ex 4 is probably homologous with 36 in Lepisma (Fig. 20), which
connects the endosternal area m and the posterolateral area of the coxal margin behind
the coxal process.
s-ex 5 usually arises from the base of the furca, or the basisternal area immediately
anterior to the furca, and is inserted on the anterior coxal margin between the pleurocoxal
articulation and the trochantino-coxal articulation, as is the case with II, III bm 57 in
Perla; in Dissosteira s-ex 5 is 121, which arises partly from the sternellar (furcasternal)
lobe (Fig. 25). This muscle occurs commonly in many orders and is probably homo-
logous with 34 in Lepisma (Fig. 20), which connects the basisternum and the base of
the coxal process.
s-ex 6 is 2 of Mickoleit ( 1965); it arises from the posterior margin of the furca and
is inserted on the inner (mesial) rim of the coxa. Mickoleit (1965) found this muscle
in many groups of Neuropteroidea. s-ex 6 may be homologous with 35 in Lepisma
(Fig. 20), which connects the endosternal area beneath b and the posterolateral coxal
margin behind the coxal process.
s-ex 7 is a characteristic muscle in Mallophaga, which connects the mesofurca and
the meta thoracic coxal margin. It is possibly a modified s-ex 1 (not figured).
s-ex 8(x) (Ibm 1 in Myrme/eon) connects the basisternal invagination and the basal
margin of the coxa on the opposite side, anterior to the pleurocoxal articulation. Larval
muscle 171 in Corydalus is also s-ex 8 (x). 58 in Siph/onurus, which transversely
connects the posterolateral coxal margin on both sides, may be the derivative of s-ex 8 (x),
although it may also be a modified s-ex 4 (not figured). The derivation of this muscle
is unclear.
s-ex 9 (ISm 1 in Hexagenia) arises from the furca and is inserted on the internal
ridge of the coxa. The derivation of this muscle is unclear (not figured).
Trochanteral Muscles
(Fig. 26)
t-tr 1 arises from the scutum and is inserted on the trochanter. It is usually des-
cribed as the depressor of the trochanter-femur in many orders. In Corydalus 165
(t-tr 1) is an outgrowth of 175 (p-tr 1 or 2) in the larva. This muscle is obviously
homologous with 52 in Lepisma (Fig. 20).
t-tr 2 is M .. troeantero-aseellare of Grandi (1947b) in Ephemeroptera which is
inserted on the first axillary sclerite, very close to the point of attachment of t-cx 4
(Fig. 44B). In some Ephemeroptera this muscle is absent, in some it is represented only
by a few fibers and fused with t-cx 4, and in some the two muscles are quite distinct.
The derivation of t-tr 2 is unclear, although it may be a detached part of t-cx 4.
p-tr 1 arises from the middle of the episternum or from the pleural arm, and is
inserted on the trochanter. This muscle is probably the derivative of 53 or 54 in Lepisma
(Fig. 21 ).
76
FIG. 26. Tergosternal and trochanteral muscles in pterothorax (original).
p-tr 2 arises from the dorsal part of the episternum or the basalare and is inserted on
the trochanter. This muscle may be confluent with t-tr 1 for the most part (e.g.,
Stenopsocus). p-tr 2 is probably the derivative of 55 in Lepisma (Fig. 21).
p-tr 3 arises from the epimeron and is inserted on the trochanter. This muscle has
been found in Siphlonurus and Odonata, and appears to be homologous with 129 and 132
in Lepismachilis, which are epimerotrochanteral muscles in the meso- and meta-thorax.
If this homologization is valid, p-tr 3 must represent an archaic thoracic muscle in the
Insecta which has become lost even in Lepisma.
p-tr 4 is the levator of the leg in Mecoptera and Diptera, and connects the ventral
pleural arm (the apodeme at the ventral end of the pleural ridge) and a trochanteral
apodeme. It is actually homologous with a coxotrochanteral muscle in insects in which
the ventral end of the pleural ridge is not conspicuously modified (not figured).
s-tr 1 connects the furca and the trochanter in many orders. It is homologous with
51 in Lepisma (Fig. 20).
MAJOR EVOLUTIONARY FEATURES OF THORACIC MUSCULATURE
As the foregoing discussion shows, muscles of certain categories are more abundant
in the Apterygota than in the Pterygota; for muscles of certain other categories, however,
the reverse is true. Such increase and decrease in number of muscles during evolution
must be accounted for in terms of functional significance and the underlying develop-
mental processes involved.
Evolution of thoracic muscles in adult insects: Despite the drastic modification of
the neck region in the Pterygota, which has resulted in the formation of at least four
major types of lateral cervical sclerites, many muscles occurring in this region in the
Pterygota are probably homologous with similar muscles in Thysanura and Diplura
(Table 1). However, some cervical muscles (e.g., op-cv 2, 3; cv-s 1, 2(x); cv-s 3)
have apparently been acquired newly in the Pterygota, probably by enlargement and
secondary splitting of the precursors. In any case, there has been no conspicuous
tendency for the cervical muscles to become reduced in number, as the insects became
TABLE I
Comparison o{ muscles of the Apterygota and Pterygota. The preceding homologizations of mus-
cles in the Pterygota with those in the Apterygota, especially with those in Thysanura, are tab-
ulated below. The numbers in the column of the Apterygota are those of Lepisma saccharina
Pterygota
cv(d)-t 1
op-t I
op-t 2
op-t 3
op-t 4
op-cv 1
op-cv 2, 3
op-cv(v) 4
op-p 1
Op-p 2
op-p 3
op-s 1
op-s 2
t-s(cv) 1
t-s(cv) 9
t-cv 1
t-cv 2, 3
cv-s 1
cv-s 2(x)
cv-s 3
cv-s 4
cv-cx 1(x)
cv-cx 2(x)
cv-cx 3
cv-p 1
cv-cv 1
s 1, 2
s3
s-cx(cv) 1
p-s(cv) 7
t11
t 12
t 13
t 14
t 15
t 16
t 17
t 18
t 19
t-p 1
t-p 2
t-p 3
t-p 4
t-p 5
t-p 6
t-p 7
t-p 8
t-p 9
t-p 10
t-p 11
t-p 12
t-p 13
t-p 14
t-p 15
t-p 16
t-p 17
(pp. 48-56) unless otherwise noted.
Apterygota
Cervicoprothoracic muscles
5
27 in Lepidocampa
7
8
9
41?
?
?
89
?
71?
71?
55 in Lepismachilis?
42 or 46
44
48?
50 and 54 in Lepismachilis
?
?
?
28 in Lepismachilis
38
38
38?
44
?
12-16
11
36 in Lepidocampa and 81 in Lepismachilis??
11
Tergal muscles
23 in Lepismachilis
7, 13, or 15, 24 in Lepismachilis
6
4, 5
?
?
1
9 in Lepismachilis
11, 12, 17, 23 in Lepismachilis
Tergopleural muscles
111 in Lepisma? 66, 77 in Lepismachilis
111 in Lepisma? 66, 67 in Lepismachilis
89
New formation in Pterygota
87?
116, 121 in Lepismachilis?? or new formation in Pterygota
116, 121 in Lepismachilis?? or new formation in Pterygota
115, 120 in Lepismachilis? or new formation in
Pterygota
41? or new formation in Pterygota
TABLE I (Continued)
Pterygota Apterygota
t-p 18
t-p 19
t-p 20
t-p 21
t-p 22
t-p 23
t-p 24
t-p 25
t-p 26
t-p 27
t-p 28
t-p 29
t-p 30
t-s 1
t-s 2
t-s 3
t-s 4
t-s 5
t-s 6
t-s 7
t-s 8
t-s 9
t-s 10
t-s 11
t-ti(cx) 1
t-ti(cx) 2
t-ti(cx) 3
t-cx 4
t-cx 5
t-cx 6
t-cx 7
t-cx 8
p 1
p2
p 3
p4
p5
p-s 1
p-s 2
p-s 3
p-s 4(x)
p-s 5
p-s 6
p-s 7
p-s 8
p-s 9
p-s 10
p-s 11
p-s 12
p-s 13
p-s14
p-ti(cx) 1
p-ti(cx) 2
p-ti(cx) 3
p-cx4
87, 88
87, 88
Second oblong muscle in Ctenolepisma?, or new
formation in Pterygota
Tergosternal muscles
46
42?
45
'
?
46?
48, 68 in Lepidocampa?
41?
44
64 in Lepismachilis
?
Tergotrochantinal (coxal) muscles
71
72
73?
62
61
One of 63, 64, 65 in Lepisma, and one of 103, 104, 107,
108 in Lepismachilis
One of 63, 64, 65 in Lepisma, and one of 103, 104, 107,
108 in Lepismachilis
62?
Pleural muscles
26?
?
Pleurosternal muscles
27?
?
42??
?
?
38, 61 in Folsomia?
11
31 or 32 in Lepismachilis ?
17
?
?
42?
?
?
Pleurotrochantinal (coxal) muscles
?
29, 61, 95 in Lepisma (Maki 1938)?
TABLE I (Continued)
Pterygota Apterygota
p-cx5 101
p-ex 6 102
p-ex 7(x) ?
p-ex 8 135 in Lepismachilis?
p-ex 9 136, 137 in Lepismachilis
Sternal muscles
s 11 38 or 40 in Lepismachilis?
s 12 12
s 13 15, 18
s 14 19
s 15 14? or 20?
s 16 12
s 17 21
s 18 ?
s 19 ?
s 20 ?
s 21 ?
s 22 ?
s-stg. ?
Sternocoxal muscles
s-ex 1 31
s-ex 2 33
s-ex 3 37
s-ex 4 36
s-ex 5 34
s-ex 6 35?
s--ex 7 31?
s-ex 8(x) ?
s-ex 9 ?
Trochanteral muscles
t-tr 1 52
t-tr 2 62??
p-tr 1 53 or 54
p-tr 2 55
p-tr 3 129, 132 in Lepismachilis
p-tr 4 ?
s-tr 1 51
winged. This is obvious in a comparison of the cervical muscles between lower ptery-
gotes and apterygotes.
In Thysanura some tergal muscles are transsegmental, clearly crossing the segmental
border to be attached on the tergum of the next segment. Apparently these muscles
cannot exist in the Pterygota, because of the greater development of the phragma. In
Lepisma muscles 1, 2, 3 are such transsegmental muscles (Fig. 19); they have been com-
pletely lost in the Pterygota. However, t 18 and t 19 in Odonata are transsegmental
muscles, which are probably homologous with similar muscles in Lepismachilis. Other
tergal muscles in Le pisma ( 4, 5, 6) inherited by the Pterygota, have become important
fiight muscles.
Most of the tergoleural muscles in the Pterygota connect the dorsal part of the
pleuron and the lateral area of the tergum; and most of them cannot be homologized
with tergopleural muscles in Thysanura. It is an obvious hypothesis that most of these
tergopleural muscles in the Pterygota are new muscles that arose in connection with
acquisition of wings. Of these tergopleural muscles the postembryonic development
process of t-p 5 and t-p 6 hasbeen well investigated, and t-p 6, at least, is an entirely new
muscle that has been added to the thoracic musculature of the Pterygota (p. 62). The
same is apparently true of some other tergopleural muscles. For instance, in Perla
(Wittig 1955) t-p 3 and t-p 7 are produced by the secondary division of a muscle, which
may be considered to be t-p 3; in Baetis (Bocharova-Messner 1965) p 5 and t-p 10 are
indistinguishable in earlier stages of development, and only during the 14th nymphal
stage do these muscles become differentiated from a single muscle. In the mesothorax
of Diptera some individual tergopleural muscles are represented by more than one bundle
of muscle fibers each, obviously as a result of a secondary enlargement and subsequent
splitting of originally single muscles; t-p 9 in Diptera has probably been produced through
such secondary processes.
Some of the tergopleural muscles in Thysanura have apparently become lost in the
Pterygota. For instance, some of the 81-87 group of muscles in Lepisma and many of
the posterior tergopleural (perhaps partly tergosternal) muscles present in Lepismachilis
are no longer found in the adult Pterygota.
Tergosternal muscles are abundant in kinds in the Apterygota, including Diplura,
Collembola, and Thysanura. Among adult pterygotes, only Ephemeroptera have retained
relatively large numbers of tergosternal muscles (t-s 2-7, 9), although probably not all of
these muscles are homologous with those in Apterygota. Odonata also have two peculiar
tergosternal muscles (t-s 10, 11), and one of them (t-s 10) is probably homologous with
a muscle in Lepismachilis (p. 78). In other pterygotes the only persistent tergosternal
muscle is t-s I, which is also present in Lepisma. It can be said that tergosternal muscles
have had an overall evolutionary tendency to bc;:come progressively reduced in number.
All tergotrochantinal (coxal) muscles in the Pterygota can, with varying degrees of
certainty, be homologized with those present in Thysanura. This fact represents a
highly conservative aspect of the evolution of the thoracic musculature in the class
Insecta. It is not surprising, therefore, that t-cx 4 occurs both in Ephemeroptera and
Odonata which are not closely related (p. 45). Most probably, the origin of tergocoxal
muscles can be traced back into more primitive groups of Arthropoda. Among major
pterygote orders, a significant decrease in number of tergocoxal muscles has occurred in
Hymenoptera.
Although the original function of the tergocoxal muscles was undoubtedly the move-
ment of legs, they have undergone functional changes to become flight muscles, in varying
degrees in various groups of winged insects. Thus, in Orthoptera, as Wilson ( 1962)
has experimentally shown, t-ti 2 is an elevator of the wing and promotor of the coxa;
t-cx 6 and t-cx 7 are elevators of the wing and the remotors of the coxa; t-cx 8 is a
supinator depressor of the wing and remotor of the coxa. In Dermaptera, according to
Kleinow (1966), t-cx 6 or 7 moves both the legs and wings. In Blattaria also leg
muscles are bifunctional. In many Coleoptera anterior tergocoxal muscles have lost
their original function as leg muscles and they are actually flight muscles. This func-
tional change in Coleoptera has accompanied increased incorporation of the leg base into
the thoracic wall, and the incorporation of the leg base in turn has resulted in the change
of points of origin and insertion of these muscles. The tergocoxal muscles in Coleoptera
are highly effective as depressors of the tergal margin, and hence they are elevators
of wings.
None of the pleural muscles can be homologized with muscles in the Pterygota with
certainty. As studies on the development of p 3 (p. 70) suggest, muscles occurring
within the boundary of the pleuron are probably new formations in the Pterygota. Tiegs
( 1955) has already suggested such an origin of the pleural muscles.
Some of the pleurosternal muscles in the Pterygota have been inherited from the
Thysanura-like ancestor, and some of them have apparently arisen newly in the Ptery-
gota. The author (1963a, b) attempted to derive the pleural muscles (p-s, p-cx) from
tergopleural muscles in Lepisma by shift in their dorsal points of attachment from the
tergum to the dorsal part of the episternum. This idea is here retracted, since it is not
supported by sufficient facts.
The tendency for the muscles arising from or occurring within the pleuron to
decrease in number in higher Pterygota is generally correlated with increased sclerotiza-
MATSUDA: THE INSECT THORAX Rl
tion of the pleural wall. The hardened skeletal wall in these insects simply would not
serve ~ a flexible and functional surface for the action of small muscles.
Among sternal muscles, those most frequently present (s 11-16) are probably
homologous with those present in Lepisma. Although the origin of some muscles,
especially those in Odonata (s 21, 22), remains unclear, there is at least no positive
evidence which indicates an addition of new sternal muscles in the Pterygota. Further,
there is also a marked tendency for the sternal muscles to be fewer in higher Pterygota.
These facts indicate that the overall evolutionary tendency of the sternal muscles has
been toward reduction in number.
Similarly, most of the sternocoxal muscles and the trochanteral muscles are homo-
logous with those present in Thysanura. Here also, the overall evolutionary tendency
has been toward reduction in number.
The above discussion shows that reduction in number of muscles has been a pre-
dominant evolutionary trend of the adult thoracic musculature in the Insecta. At the
same time, however, some muscles have been added to the thoracic musculature; and in
their turn these secondary muscles have decreased in number in the higher pterygotes.
Clearly, then, the thoracic musculature includes two kinds of muscles of different
evolutionary origins; viz. muscles that have been inherited from the wingless ancestor, and
those which have arisen de novo in the Pterygota. The former may be called palaeo-
genetic muscles, and the latter neogenetic muscles. Most t-p muscles, all p muscles,
many p-cx and p-s muscles are neogenetic.
Gross developmental patterns of the thoracic muscleS'":
( 1) In hemimetabolous insects caenogenetic muscles'" are absent or few in
number. In Gryllus the nymphal and adult muscles are nearly the same; only one
muscle is lost during development. In Acrididae several muscles are lost during the
last nymphal stage and after the adult stage is reached, but these muscles are homologous
with adult muscles in other insects; no new muscle is formed during development.
Hence, in Orthoptera there is no evidence that caenogenetic muscles occur. In Perla
and Blattaria also, all or nearly all degenerating muscles are homologous with adult
muscles in other insects. In Homoptera some degenerating muscles are not homologous
with adult muscles, but this is due partly to the inadequacy of the original descriptions of
muscles studied. In Heteroptera there is no evidence for the presence of the caeno-
genetic muscles; all degenerating muscles can be homologized with adult muscles. In
Odonata three degenerating nymphal muscles cannot be homologized with adult muscles
and they appear to be caenogenetic. Considering the obscure phylogenetic position of
this order, however, these muscles may be found in some apterygotes in the future.
(2) Caenogenetic muscles tend to increase in number in higher holometabolous
insects. In Corydalus only two muscles are caenogenetic; and in a more derived genus
of the same order, Myrmeleon, the caenogenetic trend is obviously more pronounced.
In Dytiscus 13 degenerating muscles cannot be homologized with adult musCles and they
appear to be caenogenetic. In Trichoptera and Lepidoptera also the caenogenetic trend
of the thoracic muscles is pronounced. In Psychoda most of the larval muscles cannot
be homologized with adult muscles, and hence many of these unidentifiable muscles are
presumably caenogenetic. In Apis two larval muscles are clearly caenogenetic.
(3) The time of appearance and degeneration of muscles during postembryonic
development varies in different groups of insects (heterochrony). Muscles start their
development at different times in different insects. This shifting in time of appearance,
which may be termed heterochrony, is evident, for instance, in the production of t-p
2'8For more information about the muscle development in each order, refer to p. 122 (Ephemeroptera), p. 132
(Plecoptera), p. !53 (Blattaria), p. 186 (Orthoptera), p. 216 (Coleoptera), p. 262 (Homoptera), p. 282
(Heteroptera), p. 292 (Neuroptera), p. 324 (Diptera), p. 332 (Trichoptera), p. 354 (Lepidoptera), p. 372
(Hymenoptera), p. 399 (Odonata).
27
Caenogenetic muscles are muscles that have been developed exclusively in relation to the immature
stages and a r ~ not present in. the adult .insect. In this study caenogenetic muscles are identified as such when
the degenerating muscles of tmmature l!lSects cannot. be hom?logized, by morphological correspondence, with
the known adult muscles. Caenogenetic muscles, 1n evolut10n, are not related to changes in the adult
musculature.
muscles. In lower insects (Blattaria, Orthoptera, Plecoptera, etc.), t-p muscl!es (except
t-p 5, 6) are already clearly present in early stage nymphs, whereas in A pis these 'muscles
are formed during the pharate pupal stages. Indirect flight muscles (t 13, t 14, t-p 5,
t -p 6) start their development at different developmental stages in different insects (see
p. 86).
Heterochrony applies also to the time of degeneration of muscles. For instance, in
Orthoptera some t-p muscles degenerate after the adult stage is reached; whereas in many
other insects the homologous muscles degenerate before the adult stage is reached. The
postimaginal degeneration of muscles occurs also in Apis, Gerris, Coleoptera, lsoptera,
Diptera, and probably in others. The degeneration of the flight muscles in some insects
is known to be related to certain aspects of behavior, such as nuptial flights, gallery
construction in wood, etc.
Conservative aspects of the thoracic musculature in immature insects: Apparently
in association with their structural similarity to the adult Apterygota, immature ptery-
gotes tend to preserve muscles present in Apterygota but absent in the: adult Pterygota.
Such muscles include op-t 4 (9 in Lepisma), which is present in larval Trichoptera
28
;
t-p 27 (87 or 88 in Lepisma), which is present in larval Corydalus; t-p 28 (87 or 88),
which is present in larval Corydalus and larval Limnephilus; t-p 30 (a muscle in Cteno-
lepisma and 57 in Lepidocampa), which is present in the abdomen of nymphal
Ephemeroptera and may be present in the thorax of larval Trichoptera and Diptera; t-s 9
( 44 in Lepisma), which is present in nymphal H exagenia; p-s 7 ( 11 in Lepisma), which
is present in larval Dytiscus and larval Neuroptera; p-s (cv) 7 (11 in Lepisma), which is
present in larval Myrmeleon; t 17 ( 1 in Lepisma), which is present in nymphal
Ephemeroptera; p-s 8 (31 and 32 in Lepismachilis), which is present in larval Corydalus.
All these muscles are palaeogenetic muscles.
There is also a tendency for nymphal and larval insects to preserve muscles present
in the adults of related but generally less derived groups.
In the nymph of Psylla, degenerating t-cx 6 or 7 (meso- and meta-thorax) and p-s 1
(metathorax) is present in the adult of Aphis which is generally more primitive struc-
turally. t-cx 6 or 7 and t 12 or 13 (metathorax), which degenerate in Psylla, are present
in the adults of Psocoptera, to which the sternorrhynchous Homoptera are closely related
(Crampton 1921). Another degenerating muscle in Psylla, s-ex 5, occurs in the adults
of other orders.
Among Heteroptera, all or nearly all degenerating muscles in Ranatra, Gerris,
Notonecta, and Dysdercus are homologous with muscles in the adults of other groups of
Heteroptera, no caenogenetic trend being evident. Of the degenerating muscles in these
genera, p-s 9 occurs in the adult of lower hemimetabolous orders, but not in adult
Heteroptera. Comparing the adult muscles in Heteroptera and those of auchenorrhyn-
chous Homoptera, it is evident that some muscles of auchenorrhynchous Homoptera are
absent in Heteroptera, and nearly all muscles of Heteroptera are homologous with those
present in auchenorrhynchous Homoptera. These facts suggest that those muscles lost
during the presumed phylogeny of the Homoptera-Heteroptera might be found in the
nymphs of less derived Heteroptera in the future.
Among Orthoptera, degenerating muscles t-p 3, t-p 4, t-p 16, p-s 6, and s-ex 1 in
Locusta are present in the adults of Tettigoniidae and Gryllidae, which are generally
more primitive; other degenerating muscles, t-p 8 or 9, t-p 11, t-p 15, and t-p 17, are
present in other orders. A similar tendency is noted also in the development of thoracic
muscles in Acanthacris, Phymateus, etc.
In Periplaneta (Blattaria) and Perla (Plecoptera) relatively few muscles degenerate
and only one species of Plecoptera has thus far been studied. The tendency noted for
Homoptera, Heteroptera and Orthoptera is therefore not evident. In H exagenia
2B'The occurrence of this muscle in larval Trichoptera may be by convergence.
(Ephemeroptera) some degenerating muscles are homologous with muscles present in the
adult of the Apterygota (already noted).
Among Neuroptera, most of the degenerating larval muscles in Corydalus, whether
palaeogenetic or neogenetic, are homologous with muscles present in the adults of the
Apterygota and of other pterygote orders. Some degenerating noncaenogenetic muscles
(e.g., s 12, s 14, p-s 6, p-s 9) in Myrmeleon are present in the adult of Corydalus.
In the mesothorax of Dytiscus (Coleoptera) some degenerating palaeogenetic and
neogenetic muscles (p-s 3, t-p 3) are present in the adults of other Coleoptera, and 19
degenerating larval muscles are present in the adults of other, mainly lower pterygote
orders.
Although the caenogenetic trend is pronounced in Diptera, Trichoptera, and
Lepidoptera, some degenerating larval muscles in these orders are homologous with
muscles present in the adults of other groups. In Apis 12 degenerating pupal muscles
are homologous with adult muscles in less derived genera, Sirex and Xyela of the same
order.
Relationships between ontogeny and phylogeny in muscle development: The
tendency for immature insects to preserve the muscles (neogenetic and palaeogenetic
muscles) present in the adults of generally more primitive and related groups of in&ects
supports Haeckel's (1866) theory of recapitulation or the "biogenetic law," which stated
"ontogeny is a short recapitulation of phylogeny and phylogeny is the mechanical cause
of ontogeny." What is important here is the first part of the law"" that ontogeny
recapitulates phylogeny, and the relation between ontogeny and phylogeny in the muscle
development can be explained by a simplified diagram shown in Fig. 27. The three
developmental patterns already discussed must be remembered in analyzing the
relationship.
In a primitive apterygote species A (Fig. 27), all muscles formed in the embryo
become adult muscles and no new muscles are added during postembryonic develop-
ment.30 The development of thoracic muscles in the Apterygota is not sufficiently known
to allow an analysis of the relationship between ontogeny and phylogeny in muscle
development within the Apterygota. '
In a primitive hemimetabolous species B, all palaeogenetic muscles present during
the immature stage are those inherited from A, and they are equal in number in A and B.
A few neogenetic muscles are produced in B, but for the palaeogenetic muscles the
recapitulatory process in development is clear. A few of these palaeogenetic muscles
degenerate later.
In a more derived hemimetabolous species C, all muscles present during the nymphal
stage are homologous with all muscles present in the adult of species B. The fact that
some muscles are palaeogenetic and some others are neogenetic in origin does not obscure
the recapitulatory process, since both kinds of muscles have been inherited from B.
In a primitive holometabolous species D, a few caenogenetic mus.cles occur during
the larval stage, in addition to palaeogenetic and neogenetic muscles. Caenogenetic
muscles and a relatively small number of palaeogenetic and neogenetic muscles
degenerate during development."
In the larva of a more derived holometabolous insect E, more caenogenetic muscles
occur, but all palaeogenetic and neogenetic muscles in the adult of species D are present
in the larva of species E, and the recapitulatory process of muscle development is clear.
_Beer (1958) rejected the l?_iogenetic law on two major as is evident from his statement (p. 125),
It ts these early-developed characters ( caenogeneuc characters) are not exceptions in phylogeny,
together wxth the fact that phylogeny is the result of ontogeny instead of being its cause, that we reject
Haeckel:s. theory of The first part of the biogenetic law that ontogeny recapitulates phylogeny
as emp1ncal _fact remams intact in de Beer's rejection of the theory. Caenogenesis is indeed a universal
phenomenon m insects, but what the author is concerned with here, in testing the first part of
Haeckel's law, IS the palaeogenetic and neogenetic muscles. For more discussions on the biogenetic
see Sewertzov (1931), Kryzanowsky (1939), Remane (1952, 1960), and Rensch (1959). Among entomologists
Snodgrass (1963) and Hennig (1966) admitted the applicability of the first part of the biogenetic law. '
3
0'fhis hypothesis is based on the fact that in Orchesella (Fig. 30) all embryonic muscles become adult muscles.
31
This agrees with Hinton's (1948) postulate that in primitive holometabolous insects more muscles persist
into the adult stage than in specialized holometabolous insects.
84
A
B
c
D
E
F
IMMATURE STAGE
ADULT
II
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FIG. 27. Diagram showing relationship between ontogeny and phylogeny of thoracic muscles
in insects (original). Black bars are palaeogenetic muscles, striped bars neogenetic muscles,
white bars caenogenetic muscles, thin bars degenerating muscles. A-F are stages of evolution
(see text).
During the final molt into the adult, however, some neogenetic muscles may increase
their numbers by a secondary splitting (with a consequent production of muscles with
new designations).
The recapitulatory process of muscle development in an even more derived holo-
metabolous insect F is essentially the same as that seen in the muscle development in
species E, but the reduction in number of muscles is more pronounced than in E.
In the larvae of holometabolous insects the production of caenogenetic muscles tend
to obscure the recapitulaJ:ory tendency in muscle development. Nevertheless, by dis-
tinguishing the caenogenetic muscles from palaeogenetic and neogenetic muscles, the
recapitulatory tendency in the development of the muscles of the last two categories can
still be discerned.
As already noted, the time of emergence and of degeneration (heterochrony) of
muscles varies in var-ious insects. Therefore, examination of muscle development at a
certain time or times during development often does not reveal the whole recapitulatory
tendency. Also, production of new muscles (see below) during later periods of develop-
ment may obscure the recapitulatory tendency.
Underlying developmental mechanisms for production of new muscles: Enlarge-
ment and secondary splitting of an originally single muscle and shifting of the point of
attachment are major processes by which muscles with new designations are produced;
the development of muscles t-p 5, 6 .and p 3 (p. 62 and p. 70) provide good examples.
As Wittig ( 1955) showed in Perla (p. 132), however, differentiation or sclerotization of
insertion points is also a necessary prerequisite for the eventual establishment of adult
muscles.
The shift in points of attachment is, after all, attributable to the difference in the
embryonic and postembryonic developmental potentials between the musculature and the
exoskeleton to which muscles are attached. For example, Daly ( 1964) found that
during the postembryonic development of the thorax in Apis, the imaginal musculature
develops prior to the production of the skeletal mould. According to Daly, most muscles
of the bee do not grow between final sites of attachment. In reality, the myoblasts
develop appressed to the undifferentiated body wall, or they are supported by larval
muscles which are already suspended within the body cavity. Daly thought attachment
of some kind, perhaps not always involving tonofibrillae, are established by the incipient
muscles, and the subsequent growth of epidermis carries the muscles to near their final
positions. Daly ( 1964) also found that in A pis the form of the thoracic skeleton is
virtually complete before the direct and indirect flight muscles enlarge their areas of
attachment. In Simulium (Hinton 1959a, 1961), in which the imaginal indirect flight
muscles are already attached to the epidermis in the second larval stage, changes in the
position and orientation of the indirect flight muscles are also brought about by dif-
ferences in the epidermal growth rate of different parts of the mesothorax. All of these
facts lead to the conclusion that differences in the growth rate of epidermal cells and
differences in time of connection between muscles and the epidermal cells in various
insects result in the production of homologous muscles with new points of attachment,
and hence result in the production of muscles with 'new designations.
Although the evidence is only circumstantial, the development of some external
structures appears to be dependent on muscle attachment. For instance, muscles are
scarce in the embryonic pronotum of Orchesella (Fig. 30A), and this embryonic pro-
notum becomes almost completely lost in the adult (Fig. 30B). All muscles attached to
the anterior end of the embryonic mesonotum come to occupy the anterior end of the
thorax which is actually devoid of the pronotum." Similarly, shifting in points of attach-
ment of certain dorsoventral muscles in Blattaria is probably due to reduction of the pre-
episternum in this order.
As the discussion shows, the underlying developmental mechanism for the
shift of muscle attachment is complex. It may be said that the development of
ectodermal parts is often dependent on muscles after the connection of the two systems
is established, but not before. However, some external structures appear to differentiate
independently of the musculature. For instance, elaborate modifications of the pronotum
in Membracidae do not seem dependent on the influence of muscle development.
Relationships between muscles and internal skeleton: Weber (1933), Maloeuf
(1935b), and Chadwick (1957) attributed the development of the internal thoracic
skeletons (the phragma, furca, etc.) to the action of muscles that are attached to them.
Daly ( 1964) found in A pis that an epidermal bridge is formed in exactly the position
occupied by a muscle that has degenerated. According to Daly ( 1964), therefore, the
epidermal bridge (ligament) replaces, but is not derived from, a former muscle.
Whatever the developmental mechanism may be, some muscles are replaced by
ligamentous structures which must be epidermal in For instance, in many insects,
32ffence, the points of attachment of muscles in the adult do not identify the external parts. For further
discussion on the reliability of musculature in homologizing external structures, see Remane (1963), Matsuda
(1965),.and Gouin (1968).
muscle p-s 1 is often replaced, partly or completely, by a ligamentous bridge that
connects the pleural arm and the furcal arm; t-p 4 in Lepidoptera is often replaced by the
tergopleural process. In the Apterygota various endosternal processes are sometimes
represented by muscles.
In the Pterygota, formation of an internal apodeme or apophysis usually involves
cuticular invagination, and muscles become directly connected with the cuticle (through
tonofibrillae). However, in Machilidae, as Barth ( 1963) showed, various internal
tendons associated with muscles are formed by the thickening of the basement membrane,
through which the tonofibrillae pass. The endosternum which Carpentier and Barlet
found in various apterygotes must be of the same histological constitution. Thus, homo-
logous muscles may be inserted on the cuticle in the Pterygota, and on the epidermis in
Apterygota and larval Pterygota.
33
Development of flight muscles: Flight muscles (mainly t 14, t-p 5, 6) are known to
be produced through various developmental processes. Hinton ( 1959b) classified these
processes as follows:
I. Nuclei of rudimentary fiber divide and there is no incorporation of free
myoblasts.
A. Fibril formation is precocious - Blattella germanica (Blattaria), Acrido-
peza reticulata ( Orthoptera), Cyclochila australasiae (Homoptera),
Odonata.
B. Fibril formation begins at about the time that nuclear division ceases -
Diptera-N ematocera.
II. Free myoblasts are incorporated into the muscle rudiment and the muscle
nuclei do not divide - Erythroneura ix (Homoptera).
III. Free myoblasts are incorporated into the muscle rudiment and the muscle
nuclei also divide- Bathylus albicinctus (Homoptera).
N. All myoblasts of the indirect flight muscles are contained within the larval
muscle - Neuroptera, Coleoptera, some Hymenoptera, and Lepidoptera.
V. Free myoblasts adhere to or penetrate degenerating larval muscles - Ptero-
malidae (Hymenoptera), Diptera-Cyclorrhapha including Drosophila.
The above facts and others that are mentioned for each order in part II of this work
show that the derivation of flight muscles varies considerably in various groups;
here the substitution of rudiments in ontogeny is involved in producing homologous
muscles. The fact that myoblasts are incorporated into the muscle rudiment in some
Homoptera shows that the substitution has not occurred exclusively in holometabolous
insects. The flight muscles start their development at different times in different insects.
This shifting along the time scale (heterochrony), as already noted, must have been an
underlying factor in the evolution of the thoracic muscles, including flight muscles.
Histological and physiological evolution of flight muscles: Based chiefly on dif-
ferences in the relative positions of the nuclei, and in sizes and shapes of the contractile
organelles, three histological types of muscles are usually recognized. The normal non-
flight muscles are usually called tubular muscles. They consist of the central core
containing the nuclei, and myofibrils radiating in a row from this central core. The
contractile organelles (fibrils) exist as thin lamellae, about 0.5 p. or less in diameter.
The other two types are the close-packed type and the fibrillar type, in which nuclei are
just beneath the sarcolemma, or scattered irregularly among myofibrils, or both. The
fibers of the close-packed type muscle have myofibrils, the diameter of which range from
0.5 to 1.5 p.. The myofibrils in the fibrillar type muscle are large, each being 1.5 p. or
greater in diameter. Tiegs (1955) recognized in Cicadidae a rather unusual type of
flight muscle with fibers arranged in small bundles in a nucleated sheath, and fibrils in
33This parallels the evolution of the tentorium in the Thysanura-Pterygota complex (Matsuda 1965). Barth
(1?63), similarity of the dermal aoodeme in Machilidae and larval Diptera (Perez 1910),
sa1d that th1s stmtlanty supports Handlirsch's (1930) untenable hypothesis that Thysanura represent a
neotenous state of the Pterygota. The attachment of muscles to the epidermal apodeme in larval insects is
better viewed as recapitulating, in a general manner, the ancestral state of muscle attachment.
87
complex lamellae; but the dimensions of these fibrils are not significantly different from
those in the close-packed type of muscle.
The trunk and leg muscles are usually of the tubular type, and in Blattella, which
is a weak flier and is generally primitive structurally, a suspected flight muscle (t-p 5?) is
also of the tubular type. It appears certain, therefore, that the fibrillar and close-packed
types of flight muscles in other insects are specializations. The Odonata are unique in
that all flight muscles are of the tubular type, but they have relatively large sarcosomes
which provide enough energy for flight.
In ordinary flight muscles, the relationship between muscular contraction and
motor-nerve impulse is (:1; in other words, they are synchronous. Boettiger (1957),
however, established that in the flight muscles of the fibrillar type in Diptera, Hymenop-
tera, Coleoptera, and some Hemiptera, a single nerve impulse causes more than one
muscle contraction. Pringle (1957a, b) suggested that the rate of beat in such instances
is determined not by the nerve impulses, but by the load - in" terms of the wing area -
imposed on a natural rhythm that is produced within the flight muscles themselves; he
called this the myogenic rhythm. Of the two physiological types, the 1: 1 muscles are
histologically less different from normal trunk muscles. Pringle (1957a) further sug-
gested that the asynchronous relationship between the nerve impulse and the muscular
contraction has a definite biological advantage and has developed in accordance with the
requirement for increasingly higher frequency of wing beat in higher insects. Further-
more, as Pringle pointed out, these histological and physiological specializations have
evidently occurred independently several times in evolution, since the orders in which
such specializations occur have not been derived from a common stock.
Recent studies using the electron microscope have revealed that the fibril is itself
composed of subunits. Huxley and Hansen (see Huxley, 1958, for summary) proposed
that a muscle fibril contracts by the process of two sets of filaments sliding past each
other. The energy required for the sliding process is provided by ATP, and the utiliza-
tion of this substance within the fibrils during contraction requires the presence of calcium
ions. Recent electron-microscopic studies have revealed further details of the synapses
between nerve and muscle, the oxygen transport system, structural differences in the
fibrils of asynchronous and synchronous muscles, etc. Smith ( 1965) has summarized
these more recent discoveries.
II. SPECIAL DISCUSSION
THE PROTURA
The peculiar interpretations and terminologies used by Berlese ( 1909b) and Prell
( 1913) were corrected by Franc;:ois ( 1964), and unless otherwise noted the following
discussion is based on his study of Eosentomon transitorium and Acerentomon
propinquum.
The pronotum and the neck region: The pronotum (Fig. 28) is greatly reduced and
its anterolateral margin is strongly sclerotized. In both Eosentomon and Acerentomon
the dorsal area between the head and the reduced pronotum is largely membranous,
although Prell ( 1913) described a series of sclerites in this area in Eosentomon
germanic urn.
Paclt (1956) thought that the neck region was prothoracic in origin, as in Collem-
bola. Based on the musculature, Franc;:ois ( 1964) believed that the head capsule con-
tained some prothoracic elements, and hence the dorsolateral part of the neck region is
entirely prothoracic in origin.
The propleuron: In both Eosentomon (Fig. 28) and Acerentomon clear-cut con-
centric supracoxal arcs are absent; neither the precoxal nor the postcoxal bridge is
present. The posterior sclerite above the coxa is, probably katapleural in origin, because
it carries a condyle that articulates with the coxa; its posterior position is probably due
to a secondary displacement of the inner coxal base, which lies obliquely (Fig. 29).
This posterior, more ventral point of articulation is apparently serially homologous with
the dorsal, pleurocoxal articulation in the succeeding thoracic segments. A well-defined
trochantin is absent. The two anterior pleural sclerites are probably anapleural sclerites;
the posterior sclerite must be, as already noted, a katapleural sclerite.
The prosternum: The prothoracic presternum in Eosentomon (Fig. 29) is frag-
mented into three lobes, and in Acerentomon it is a simple lobe. In both, the presternum
is separated from the basisternum by a transverse suture; and the sternacosta provides,
on each end, a condyle that articulates with the coxa. The sternacosta demarcates the
furcasternum posteriorly in Eosentomon. A poorly developed Y ridge in Eosentomon
continues posteriorly to the spina; the Y ridge is absent in Acerentomon. In Eosentomon
the spinasternum appears to be represented by a semimembranous area along the
posterolateral margin of the furcasternum; in Acerentomon the spina is apparently absent
and the posterior margin of the presternum is represented by the apodemal ridge.
The meso- and meta-notum: The anterior margin of the tergum in both segments is
provided with a thickened antecosta, although no clear development of the acrotergite
occurs."' In Acerentomon the transverse suture behind the antecosta may represent the
pseudoprescutoscutal suture (p. 104), although Franc;:ois did not refer to it. In
Eosentomon a median longitudinal ridge is present, whereas in Acerentomon this ridge
is reduced. A transverse ridge (apodeme transverse of Franc;:ois) also occurs in the,
posterior area of the tergum in Eosentomon (Fig. 28), and in Acerentomon this line
(ridge) appears to correspond to the posterior margin of the sclerotized part of the
tergum. The line extends anterolaterally to the anterior margin of the tergum (Fig. 28);
the lateral tergal area demarcated by this line may be homologous with the paranotum in
Diplura, although in Protura the spiracle lies within this area. The hindmost part of the
tergum demarcated by the transverse ridge may be called the postnotum; however, it
lacks the lateral connectio.n with the pleuron. A subtriangular plate, lateral to the
antecosta in Eosent01non (Fig. 28), was called the Iaterotergite by Franc;:ois, because it
appeared to be a detached part of the tergum. It is possible, however, that this plate
represents the interpleurite.
did not study the endoskeleton, so we. do not know if the endonotal phragma is present or not, and,
tf present, where. The homology of the antenor part of the meso- and meta-notum is therefore unclear
Tuxen (1931) called the greatly reduced area anterior to the ridge the "precosta.'" ' '
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The meso- and meta-pleuron: Two concentric supracoxal arcs representing the
ana- and the kata-pJeuron are present both in Eosentomon (Fig. 28) and in Acereniomon.
The dorsal pleural articulation with the coxa probably corresponds to 8 in Thysanura
(Fig, 3 SA), and to the lateral (dorsal) pleurocoxal articulation in the Pterygota. A
well-differentiated trochantin is absent, and the trochantino-coxal articulation is replaced
or represented by the articulation of the anterior end of the katapleuron with the coxa.
In Eosentomon the anapleuron consists of three sclerites; the median sclerite has a ventral
socket that articulates with the katapleuron, and its dorsal extremity carries a condyle
which articulates with the tergum. As in other Apterygota, there is no well-formed
pleural ridge, although Prell (1913) thought the median sclerite of the anapleuron was the
pleural ridge (Pleuralleiste).
The meso- and meta-sternum: As in the tergum, the anterior margin of the sternum
is strengthened by the apodemal ridge, The presternum is distinctly separated from the
definitive basisternum by a transverse suture (Fig. 29), In Eosentomon the median
longitudinal ridge of the mesosterna! furcasternum continues posteriorly to the spina,
and anteriorly to the posterior margin of the definitive basisternum. The sternacosta
and the ventral articulation are absent in the mesothorax of Eosentomon. Since the
sternacosta, which is present in the metasternum, is a landmark that separates the basi-
sternum from the furcasternum, the definitive furcasternum in the mesothorax contains
a basisternal area anteriorly.
In the metasternum of Eosentomon, however, Fran9ois ( 1964) says that a distinct
Y ridge provides a condyle that articulates with the coxa; the sternacosta connects the
anterior ends of the Y ridge. In the mesosternum and the metasternum of Acerentomon
the lateral ends of the medially obliterated sternacosta articulate with the coxa. In
Acerentomon neither the median longitudinal ridge nor the Y ridge is present. In
Eosentomon Fran9ois called the hindmost division of the metasternum behind the spina
the posterior spinasternum. However, this area may be equivalent to the poststernum.
The adult musculature: 8erlese ( 1909a) studied the thoracic muscles of Acerento-
mon doderoi and Eosentomon transitorium, but his illustrations and descriptions are too
sketchy to be restudied here.
THE COLLEMBOLA
The tergum and the neck region: In more generalized Collembola the pronotum is
clearly retained. In Entomobryomorpha earlier authors (e,g,, Willem 1900; Lameere
1935; Imms 1951) thought that the pronotum was greatly reduced or undeveloped, and
the anterior tergum is the mesonotum, Carpentier (1946b, 1947), however, thought
that the so-called mesonotum was a fused promesonotum, because certain dorsoventral
muscles of the prothoracic leg arise dorsally from it. Janin ( 1947) observed a similar
arrangement of muscles in Tomocerus, and agreed with Carpentier. Denis (1949) and
Paclt ( 1956) also accepted Carpentier's interpretation.
8retfeld (1963) showed that the same muscles are present both in the embryo and
the adult of Orchesella, In the embryo (Fig. 30A), the border between the pronotum
and the mesonotum is marked by an internal structure (phragma?), to which muscles
I ism 1-4 and II dlm and II dlm 1 are attached. In the adult (Fig. 308) all of these
muscles have evidently shifted their dorsal points of attachment forward, accompanying
the reduction of the pronotum, In the adult of Entomobryomorpha the only part
assignable to the pronotum is the neck region, and the alleged promesonotum of
Carpentier ( 1946a, 1947) is actually the mesonotum, as Bretfeld ( 1963) concluded and
as the earlier workers believed. The mere presence of the prothoracic muscles in the
mesonotal area of the adult, therefore, does not justify the interpretation that the meso-
notum is a fusion product."" The same criticism applies also to the interpretation by
Denis (1964), who thought that the true border between the pro- and meso-notum lies
posterior to ism 3, 4 and at the level of dvm 1a, b (Fig. 308).
:lliSee also p. 85.
VENTRAL
COXAL
ARTICULATION
COXA------
BASISTERNUM
STERNA COSTA
COXA
POSTSTERNUM?
MATSUDA: THE INSECT TIIORAX
PRESTERNUM
STERNACOSTA
ERNUM
SPINA
PRESTERNUM
Y RIDGE
FURCASTERNUM
SPINASTERNUM?
FIG. 29. Ventral view of thorax in Eosentomon transitorium (from Fran!,:ois).
91
In the ventral part of the neck region in Neanura, Maki (1938) recognized the
spina, which is probably homologous with the labial spina in Diplura. Muscle 3;l of
Maki in Neanura, which is inserted on this spina, is probably homologous with muscle 36
of Maki in Lepidocampa, which is inserted on the labial spina.
The pleuron: Using the endosternal process f as a landmark, the katapleuron in
Tetrodontophora (Fig. 31A) occupies a relatively large area along the inner margin of
the coxa. The anapleuron is present above the katapleuron, and its border with the
basisternum is obliterated because of the loss of h as a landmark. The endosternal
0
reaches the dorsolateral margin of the anapleuron. In Tomocerus (Fig. 31B), as
Carpentier ( 1949) showed, the anapleuron and the katapleuron extend farther toward
the median longitudinal ridge than in Tetrodontophora.
Hansen (1930) thought that in Tetrodontophora the trochantin was present. How-
ever, Carpentier (1947) thought that the trochantin referred to by Hansen was actually
the katapleuron. As Crampton ( 1926a) pointed out, in an unidentified collembolan
species, the apparent prothoracic trochantin is actually a thickening of the coxal margin.
The alleged trochantin in Tomocerus (Fig. 31B) also seems to be a thickening of the
coxal margin. Maki (1938) described muscles 44 and 67 in Folsomia as tergal pro-
motors of the coxa that are inserted directly on the coxal margin. All these facts lead
one to conclude that the trochantin is probably absent in Collembola ..
The sternum: The most pronounced characteristic of the sternum in Collembola is
the presence of the median longitudinal ridge (linea ventralis of Carpentier, 1947),
which divides the sternal surface into two lateral halves (Fig. 31A, B); Carpentier
( 1947) compared this condition with the sternum in neuropteroid insects, in which also
the median longitudinal elevation occurs. In Collembola, however, the leg bases are not
approximated along this ridge, as they are in neuropteroid insects.
Carpentier (1949) compared the endosternum in Tetrodontophora (Fig. 31A) with
that in Petrobius (Fig. 39D), and interpreted the posterior transverse stalk as a fusion of
b and m. However, bm in Petrobfus is probably m, as discussed on p. 110. Hence; in
Tetrodontophora the corresponding structure (partly muscular in. the mesothorax, entirely
so in the metathorax) is probably m.
Carpentier ( 1949) thought that the anterior transverse stalk in Tetrodontophora
(Fig. 31A) is q, which in Nico/etia (Fig. 39B) remains unconnected medially as an
extension of g. This interpretation is justified since q lies anteromedial to g, and e and f
arise from the base of g, as in Nicoletia. In Tomocerus (Fig. 31B) q, a muscular
formation, lies obliquely, and is conrrected with the median longitudinal ridge of the
basisternum. Both in Tetrodontophora and in Tomocerus, e arises from the anterior
part of the poststernum or from the spinapoststernum, and in Thysanura it is furcasternal.
The homologies of the lateral parts of the endosternum are easier to determine than
those of the anterior parts. As In Lepismatidae, i, ;, and k arise from g, although the
last two are represented by muscles in Tetrodontophora. In both Tetrodontophora and
Tomocerus p is attached to the dorsal margin of the katapleuron, as in Thysanura. A
stalk o, which is absent in Thysanura, extends from g to the anapleuron in Tetrodonto-
phora. In the meso- and meta-thorax of Tomocerus, a long two-part tigel!a lies on the
thickened anterior coxal margin (trochantin of Carpentier).
In both Tetrodontophora and Tomocerus Carpentier ( 1949) recognized four sub-
divisions of the sternum. According to Carpentier, the narrow presternum is demarcated
posteriorly by a transverse line. In the mesosternum of Tomocerus (Fig. 31B) the
basisternum is separated from the furcastemum by a transverse suture, which is
presumably the sternacosta; it reaches laterally the endostemal f. In Tetrodontophora
t h ~ definitive basisternum is demarcated by the mesal extension of the paracoxal suture.
Therefore, the definitive basisternum in Tetrodontophora is not homologous with the
basisternum in Tomocerus. Carpentier (1949), probably in error, called the proximal
katapleural area lateral to f in Tetrodontophora the furcasternum. In the absence of a
and I as landmarks the hindmost plate is difficult to interpret. However, since m is
d lm
Lb dvm--"
ism (3,4)--'
ism (1,2)
Lb vim 2
Lb
Lb ldvm
I ism 1
93
I
11
dim 1 dim 2 ism 3 dvm la,b
dvm 2
A
ldvm 1
B
Fro. 30. Thoracic muscles in Orchesella cincta (from Bretfeld). A, embryo; B, adult.
evidently associated with it, the plate may be partly the poststernum. Both in Tetro-
dontophora and in Tomocerus the hindmost plate is, therefore, regarded as a fusion of
the spinasternum and the poststernum.
The adult musculature: Maki (1938) studied the thoracic muscles of Neanura
dubiosa (Poduridae) and Folsomia sp. (Entomobryidae), and Bretfeld (1963) studied
the same in Orchese/la cincta (Entomobryidae) and Neanura muscorum (Poduridae).
Muscles in the first two species are shown in Table II. As in Diplura, no muscles are
attached to the pleuron. Tergosternal muscles and sternal muscles are relatively
abundant. Apparently no muscles are lost during the postembryonic development of
Orchese/la cincta (Fig. 30A, B).
THE DIPLURA
The neck region: In Oncojapyx (Fig. 34) and Campodea a postcephalic spina is
present. In Lepidocampa Maki ( 1938) also showed the spina in the corresponding
area. It appears highly probable that this postcephalic spina is the spina of the labial
segment, which is serially homologous with the spina in thoracic segments. The inter-
sternum of Barlet and Carpentier ( 1962), on which the spina occurs, is the poststernum
of the labial segment. In Oncojapyx (Fig. 34) and Campodea the ventral neck region
posterior to the poststernum must be the prothoracic area.
The pronotum: In Oncojapyx the medial apodeme on the anterior margin of the
pronotum is continuous with the apodeme on the posterior part of the head capsule, and
it functions as a buffer stop. In lapyx (Denis 1949) the anterior part of the pronotum
is demarcated by a transverse area, which Denis ( 1949) called the pretergite.
The meso- and meta-notum and the intersegmental relationship: In Oncojapyx
(Fig. 32) the mesonotum, the metanotum, and the first abdominal tergite are each divided
by sutures into three areas, which Barlet aJ:ld Carpentier ( 1962) called the internotum,
the notum, and the paranotum. A phragma arises from the suture between the inter-
notum and the notum. Controversy has existed over the homology of the internotum
of Barlet and Carpentier.
Snodgrass ( 1927) interpreted the anterior transverse suture in ]a pyx as the ante-
costa, or the true intersegmental suture, and the area anterior to the suture as the precosta
(acrotergite). Weber (1933) called the same area in Japyx the postnotum, and thought
it was annexed to the succeeding segment. Barlet and Carpentier ( 1962) considered
the area to be an intersegmental area, which does not belong to either of the two neigh-
boring segments. They also thought that the postnotum was represented by a narrow,
poorly sclerotized area, which is distinct from their internotum and is continuous with the
pleuron (postalar bridge). Further, according to Barlet and Carpentier, the dorsal
apotome (internotum), its ventral counterpart (intersternum), and the intervening
pleuron form a complete intersegmental ring.
Denis ( 1963) was right in criticizing Barlet and Carpentier's concept of the inter-
segment as a morphological entity. Denis interpreted the internotum of Barlet and
Carpentier as a detached part of the preceding segment that has become united with the
succeeding segment, since, as Snodgrass has maintained, the dorsal resegmentation (the
secondary segmentation) cannot be the same as the ventral resegmentation. According
to Denis, the dorsal apotome belongs to the preceding segment, and the corresponding
ventral apotome (intersternite of Barlet and Carpentier) belongs to the succeeding
segment. However, Snodgrass' concept of the intersegmental relationships, which Denis
accepted unconditionally, is artificial and ill founded (p. 24), and Denis' interpretation
is not acceptable.
The internotum of Barlet and Carpentier ( 1962) must be the acrotergite, if the line
that demarcates this area corresponds to the antecosta in the Pterygota. However, since
the supposed acrotergite in Diplura is very large compared with the very narrow ante-
notum (acrotergite) in Lepisma (Fig. 37), the line probably does not correspond to the
antecosta in Lepisma. As will be discussed later (p. 104), the line (suture) is the
SPINA-
POSTSTERN
KATAP
q
KATAPLEURON
PARA.COXAL
SUTURE
SPINA-
POSTSTERNUI
A
SPINA-POSTSTERNUM
ANAPLEURON
TROCHANTIN?
EURON
PRESTERNUM
ARACOXAL SUTURE
95
FIG. 31. A, ventral view of thoracic endoskeleton in Tetrodontophora gigas (from Carpentier);
B, the same in Tomocerus plumbeus (from Carpentier).
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TROCHANTIN
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FIG. 33. Internal view of thorax in Campodea sp. (from Carpentier and Barlet).
97
pseudoprescutoscutal suture, and the area anterior to this suture is the pseudoprescutum.
In any case, the pseudoprescutum and the area posterior to the pseudoprescutoscutal
suture should belong to the same segment. The phragma arising from the pseudo-
prescutoscutal suture in Oncojapyx is intranatal and is most probably homologous with
the pseudophragma in Lepismachilis (p. 104).
The presternite 1 of Denis, or the intersternite of Barlet and Carpentier, is the post-
sternum of the preceding segment. The ventral intersegmental relationship is treated
more fully in discussing the sternum later ( p. 97).
The pleuron: In the mesothorax of Campodea (Fig. 33), as Carpentier and Barlet
( 1951) showed, the anapleuron is identified by its association with the endosternal h and
with the basisternum, and by the presence of the muscle a-n on its dorsal margin, which
is present also in the corresponding area of Ctenolepisma (Fig. 38A). In Oncojapyx
(Fig. 34) the anapleuron is identified externally by its association with the branch of
the Y ridge and the basisternum. In this genus, however, the anapleuron is divided into
anterior (or ventral) and posterior (more dorsal) plates (Fig. 32).
The katapleuron in Campodea (Fig. 33) is identified by its contact with f, and by
the attachment of p on its dorsal margin. The katapleuron is narrowed caudad and is
apparently articulated with the posterior end of the trochantin. In Oncojapyx (Fig. 32)
the katapleuron becomes membranous posteriorly, and the posterior end of the trochantin
is attached to the elevated posterior anapleural sclerite. The relativly narrow kata-
pleural area encircles the inner coxal margin in both Oncojapyx and Campodea.
In Campodea (Fig. 33) the anterior articulation of the trochantin with the coxa is
apparently present. If the shaded area above the coxal cavity and below the trochantin
in Campod'ea represents the outwardly exposed coxal rim, which is comparable with the
coxal apodeme or pro<;ess in Ctenolepisma (Fig. 38A) and Lepisma (Fig. 37), the
posterior end of the trochantin appears to have a posterior point of articulation with the
coxa. This apparent posterior trochantinal articulation with the coxa cannot be seen in
the figures of Oncojapyx and is not mentioned in the description by Barlet and Carpentier
( 1962).
The sternum: In Campodea (Fig. 33) the endosternal structures, as Carpentier and
Barlet (1951) showed, closely resemble those in Ctenolepisma (Fig. 38A) and other
machilids. The endosternal processes j, k, and dare represented by muscles; p connects
98
POSTSTERNUM
SPINA
PRESTERNUM
FURCASTERNUM
- POSTSTERNUM
SPINASTERNUM

SPIRACLE
BASISTERNUM ANAPLEURON
FURCASTERNUM
KATAPLEURON
SPINASTERNUM
POSTSTERNUM
FIG. 34. Ventral view of thorax in Oncojapyx basilewskyi (from Barlet and Carpentier).
MATSUDA; THE INSECT THORAX 99
the dorsal mlj.rgins of the katapleuron, and f and h are associated with the katapleuron
and the anapleuron, as in Ctenolepisma.
In Oncojapyx (Fig. 34) the lateral end of the branches of the Y ridge reaches the
base of the endosternal tendon /, which lies on the anterolateral angle of the furcasternum.
The lateral end of the branch also forms the condyle that articulates with the inner
margin of the coxa. In the prothorax of Campodea a T ridge is present, instead of the
Y ridge; Barlet and Carpentier (1962) thought that the anterior horizontal line of this T
ridge had resulted from the approximation mesad of the branches of the Y ridge. They
also thought that the arms of the Y in the meso- and meta-thorax of Diplura are homo-
logous with the sternacosta, and therefore the median triangular space delimited by these
arms is basisternal. This interpretation is probably invalid, since in some insects, in-
cluding Protura (Fig. 29), the sternacosta and the Y ridge occur together. Both in
Oncojapyx and Campodea the meso- and meta-thoracic sternacosta is absent; it may be
represented by the anterior line of the T in the prothorax of Campodea. The posterior
part of the median triangular space in Oncojapyx (Fig. 34) is therefore regarded as
furcasternal. The posterior end of the Y ridge is continuous posteriorly with the well-
developed spinal m (Fig. 34).
The furcasternum is flanked on either side by a medially reduced plate, the spina-
sternum.'" The spinasternum, as Barlet and Carpentier maintained, is thus segmental
and not intersegmental. In Oncojapyx the spina, as Barlet ( 1965) showed, is a cuticular
invagination; in Diplura the spina is present also in the metathorax.
In Oncojapyx (Fig. 34) two transverse plates lie behind the spinasternum; Barlet
and Carpentier (1962) called them the intersternum and the presternum. According to
these authors, the intersternite is a secondarily expanded part below the endosterna a
and !, which does not belong either to the preceding or to the following segment, and it
corresponds to the presternum 1 of Denis ( 1949). Their presternum corresponds to the
presternum 2 of Denis (1949).
Denis (1963) was opposed to this interpretation on the grounds that Barlet and
Carpentier were not consistent in regarding the spina as the landmark for intersegmental
demarcation, and that the endoskeletal apodemes a and l were probably derived from a
longitudinal muscle occurring in that area and these muscles belonged to the succeeding
segment. Hence, the anterior plate is the one that had been secondarily constricted off
(Apotom in the sense of Enderlein, 1907) from the presternum of the following segment,
as Denis (1949) had maintained earlier.
As already fully discussed (p. 18), it is l, not the spina, that demarcates the seg-
mental border in the Apterygota; hence the presterna 1 and 2 are not the secondarily
divided parts of the originally single presternum. The two plates are the poststernum
(presternum l of Denis 1949) and the presternum (presternum 2 of Denis 1949) of the
succeeding segment.
The adult musculature (Table II): The thoracic musculature in Lepidocampa
weberi, described by Maki (1938), shows that most muscles are homologizable with
those occurring in Thysanura. Characteristically, the spina-coxal muscles (s-ex 1,
s-ex 4) are present in all segments.
THE THYSANURA
Carpentier (1946b, 1949, 1955), Barlet (1946, 1947, 1949, 1950a, b, 1951,
1952a, b, 1953, 1954, 1967), and Carpentier and Barlet (1951, 1956, 1959) have studied
the thoracic anatomy of Thysanura in detail. The following discussion refers to the
meso- or meta-thorax, unless otherwise noted.
The neck region: In the membranous neck region in Lepisma (Barlet 1951) two
dorsal plates occur, which Barlet ( 1951) called sclerites postoccipitaux (Fig. 37). In
Lepismachilis (Fig. 35), as already discussed (p. 7), a structure comparable with the
lateral cervical sclerite in most holometabolous insects is present.
38
According to Barlet and Carpentier (1962), the spinal pit is flanked by this spinasternum. This association
is not evident in Fig. 34.
100
Lepidocampa
(Maki 1938)
25
26
27
44, 62
45, 63?
64
28
29
46, 66
47, 67
48, 68
31, 49
32, 50, 70
33, 51, 71,
34, 52, 72
69
30
35, 53, 73
37, 55, 75
38
39, 57, 77,
40
42
58, 78
56, 76,
60, 80
36, 54, 74
41, 59, 79
43, 61, 81
65
TABLE II
Thoracic in Diplura, Collembola, and Thysanura*
Folsomia
(Maki 1938)
16, 31, 54
17, 32, 55
30, 53
34, 57
19, 35, 58
36, 59
37, 60
38, 61
18
39, 40
62, 63
21, 41, 64
20, 42, 65
22, 43, 66
28, 51, 74
25, 48, 71
44, 67
23? 46, 69
24, 47, 70
26, 27, 49
50, 72, 73
45, 68
29, 52, 75
33, 56, 69
Neanura
(Maki 1938)
Thysanura
(Barlet 1952, 1953, 1967)
Dorsal musdes
15, 22, 40, 61
16, 23, 41, 62
Sternal muscles
24
43, 64
46, 67
25, 44, 65
26, 45, 66
19
9 (L.)
2 (Lm.)
4 (L.)
6 (L.)
15, 24 (Lm.)
5 (L.)
8 (L.)
2 (L.)
26? 29, 39 (Lm.)
11 (L.)
40 (Lm.)
32 (Lm.)
48 (Lm.)
25 (Lm.)
Teraosternal muscles
27? 28, 47, 68 41 (L.)
48, 49
69, 70
20, 30, 31?
29, so, 51, 71, 72
43 (L.)
57, 58, 69
72, 78 (Lm.)
66, 75, 76 (Lm.)
56, 69, 78 (Lm.)
44 (L.)
64 (Lm.)
42 (L.)?
Teraocoxal muscles
32, 52, 73 71 or 72 (L.)
64 (L.)
38, 59, 80
36, 57, 78
35, 56, 77
53, 74
61 (L.)
98 (Lm.)
63 or 64 (L.)
65 (L.)
62 (L.)
73 (L.)
Sternocoxal muscles
33, 54, 75 31 (L.)
. 37, 58, 79 36 (L.)
34 (L.)?
34
55, 76
39, 60, 81 51 (L.)
Other muscles
42, 63 33, 56
Remarks
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
*The muscles in Thysanura were arbitrarily chosen either from Lepisma (L.) or from Lepismachilis (Lm.) for reference
in homologizing the muscles in Collemboia and Diplura.
MATSUDA: TilE INSECT THORAX 101
REMARKS (Table II)
(1) 25 in Lepidocampa extends between the dorsolateral part of the head capsule
and the posterior part of the mesonotum near the median longitudinal line.
(2) 27 in Lepidocampa extends between the dorsolateral part of the head and the
median anterior part of the pronotum. This muscle is similar in position to op-t 1 in
Pterygota.
(3) 63 in Lepidocampa may not be serially homologous with 45, as the anterior
attachment is on the anterior end of the metanotum.
( 4) 22 and 40 in N eanura together correspond to 31 in F olsomia.
(5) 23 and 41 in Neanura together correspond to 32 in Folsomia.
( 6) 46 and 66 in Diplura extend between the anterolateral ends of the presterna
of adjacent segments; in Collembola and Thysanura they connect endosternites of
adjacent segments.
(7) p-s 8 in the Pterygota may be homologous with these muscles.
(8) 36 and 59 extend from the anterolateral part of the sternum to the median
lateral portion of the sternum inside the leg base.
( 9) 3 7 and 60 extend between the posterolateral and the median lateral portions
of the sternum.
(10) These muscles extend between the anterolateral and the median anterior ends
of the sternum; they may be homologous with p-s 6 in Pterygota.
( 11) These muscles extend between the middle of the lateral margin of the
sternum and the posteromedian portion of the sternum.
( 12) These muscles extend between the posterolateral end of the sternum and the
lateral margin of the sternum where 25, 44, and 65 are attached.
(13) In N eanura 19 extends between the labial appendage and the lateroventral
part of the intersegmental ridge; in Folsomia 18 arises anteriorly from the tentorial plate
and is inserted posteriorly on the ventrolateral part of the anterior end of the mesothorax.
It is not certain whether the two muscles are homologous. In Lepismachilis 25 connects
the tentorial lobe with the proendosternal process n.
( 14) Muscles in Lepidocampa connect the anterolateral angle of the tergum with
the lateral margin of the presternum. t-s 7 in the Pterygota appears to be homologous
with these muscles. 27 in N eanura is inserted on the median anterior end of the
prosternum.
(15) Muscles in Lepidocampa are inserted ventrally on the lateral part of the pre-
sternum; those in Collembola, on the ventrolateral part of the segment (sternum?); in
Lepismachilis they are inserted on the dorsolateral part of the endosternum.
(16) In Neanura muscles arise from the anterolateral area of the tergum, while in
others they arise from the posterolateral area of the tergum. The muscles are inserted
on the posteromedial or posterolateral portion of the sternum, or on the endosternal n
(Lepismachilis).
( 17) These muscles connect the ventrolateral portion of the anterior margin of
the segment with the anterior margin of the tergum of the following segment.
(18) t-s 10 in Odonata is probably homologous.
(19) 30 in Lepidocampa arises from the anterior area of the pronotum and
extends anteriorly into the head beyond the labial spina (anterior end of the prothoracic
sternum of Maki).
(20) 38 arises from the anterior portion of the mesonotum and is inserted on the
posterior dorsal coxal margin in the prothorax.
(21) 40 arises from the posterolateral part of the mesonotum and is inserted on
the posterolateral coxal margin in the prothorax.
(22) 35, 56, and 77 in Neanura arise from the posterior end of the tergurn.
(23) 23 in Folsomia arises from the tentorium, while 36 in Lepidocampa and 33
in N eanura arise from the median anterior portion of the sternum, where the labial spina
occurs.
(24) Muscles in Folsomia connect the lateral portion of the sternum inside the leg
base with the anterior basal rim of the coxa.
(25) These muscles connect the lateral portion of the sternum inside the leg base
with the posterior basal rim of the coxa.
( 26) 34 arises from the ventrolateral part of the posterior end of the prothorax
and is inserted on the anterior basal rim of the prothoracic coxa.
(27) These muscles arise on the ventrolateral portion of the anterior end of the
segment (the anterolateral portion of the sternum?) and are attached to the anterior
basal rim of the coxa.
(28) These muscles are dorsal transverse muscles that are inserted on the dorsal
vessel.
POSTG AL PROCESS
(GENAL APOPH'I'SIS)--....,
POSTLABIAL
SCLERITE
PREEPISTERNUM
(Sci. anapl. super.) -H.!S!--::-":JV
AN APLEURON - - + r ; . ~ C
SPIRACLE
bm?
Fm. 35. Ventral view of thoracic endoskeletons in Lepismachilis Y-signata (from Barlet).
PSEUDOPRESC
A
PARANOTUM
PSEUDOPHRAGMA
B
PSEUDOPHRAGMA
( ENDONOTUM)
B
FIG. 36. A, lateral view of thorax in pseudofetus of Di/ta hibernica (from Bar!et); B,
muscles associated with tergal sutures and pseudophragmata in pseudofetus of Dilta hibernica
(from Barlet).
The tergum: In Lepisma (Fig. 37) the tergum consists of the narrow antenotum,
which is demarcated by the antecosta; the prescutum, which is demarcated by the pre-
scutoscutal suture; the scutum; and the postnotum, which lies hidden beneath the tergal
lobe of the preceding segment and is continuous with the pleuron. As already discussed
(p. 12), all these subdiv.isions of the tergum in Lepisma are homologizable with those
present in the tergum of the Pterygota. In a primitive lepismatid, Nicoletia, Barlet
( 1952a) did not recognize the presence of a prescutum comparable with that in
Lepisma. However, what he called the antenotum (pretergite) in Nicoletia is com-
parable with the pseudoprescutum in Diplura and Machilidae, as discussed below.
Compared with the antenotum in Lepisma (Fig. 37), the alleged antenotum in the
pseudofetus in Dilta (labeled as pseudoprescutum in Fig. 36B) is very large. Most
probably, therefore, they are not homologous. Instead, the area in question in Dilta is
probably homologous with the anterior tergal area or the pseudoprescutum in Oncojapyx
(Fig. 32), since both areas occupy comparably large anterior areas of the tergum, and the
phragma (Diplura) and the endonotum (Dilta) arises from a transverse suture that
demarcates the area under consideration. Although the internal phragma is absent, the
anteriormost division of the tergum in Nicoletia (Fig. 40) is, according to Barlet
( 1952b), broad proximally and is not homologous with the antenotum in Lepisma.
Therefore, this area in Nicoletia is probably homologous with the anterior tergal area in
Oncojapyx and Dilta.
The phragma in Diplura and the endonotum in Dilta, which arise from the suture
that demarcates the homologous anterior part of the tergum, are therefore homologous.
Although in Lepismachilis (Barlet 1967) a comparable suture is absent, the intratergal
pseudophragma in this genus is, as Barlet (1967) pointed out, probably homologous with
the endonotum in Dilta, and hence with the phragma in Oncojapyx.
These three homologous internal structures with their three different names should
therefore be designated by a single term, the pseudophragma, and they should be dis-
tinguished from the endonotum in Lepisma, with which the phragma in the Pterygota is
homologous. Further, the transverse suture crossing the pseudophragma in Japygidae
and Dilta, and the transverse suture in Nicoletia, are not homologous either with the
antecosta or with the prescutoscutal suture, which lacks an internal phragma. An
appropriate new term for this suture is therefore the pseudoprescutoscutal suture; and
the anteriormost tergal area demarcated by the pseudoprescutoscutal suture is the
pseudoprescutum.
It may be argued that the prescutoscutal suture in Lepisma and in the Pterygota is a
modified pseudoprescutoscutal suture that has lost the pseudophragma, and this interpreta-
tion may be valid especially for Nicoletia. However, for the present, these two sutures
are considered to be distinct, until more information becomes available.
Barlet ( 1949, l950a) attempted to show that in the pseudofetus of Dilta (Fig. 36A)
a scutoscutellar suture (suture B of Barlet) divides the tergal surface behind the
antenotum into the scutum and the scutellum. Because of the apparent similarity
between suture B and the scutoscutellar suture in the Pterygota, Barlet ( 1950a) alluded
to an unacceptable theory of Handlirsch ( 1930) ,"' and suggested a possible homology
between the two sutures. Most probably, however, suture B merely represents a
secondary suture formed as a result of attachment of certain dorsoventral muscles in this.
area (Fig. 36B). No dorsoventral muscle is attached to the scutoscutellar suture in the
Pterygota. Moreover, the absence of the scutoscutellar suture in Ephemeroptera and
Plecoptera, which retain so many primitive features in the thorax, makes it further im-
probable that the suture in question is homologous with the scutoscutellar suture in the
Pterygota.
The pleuron: In Thysanura there is no pleural ridge that separates the pleuron into
the anterior episternum and the posterior epimeron. The structure homologous with the
pleural arm in the Pterygota is what Carpentier (1946b) and Barlet (1950b) called the
processus pleural in some machilids (Fig. 39A). Barlet (1950b) found, in the pseudo-
fetus of Dilta, that the pleural process arises from the dorsal margin of the katapleuron
and has an external orifice. He therefore thought that the external point of invagination
of this structure demarcates the border between the anapleuron and the katapleuron. In
Petrobius (Fig. 39A) and Lepismachilis (Barlet 1967) the well-developed pleural process
(p) is continuous with the endosternal g. According to Barlet (1950b), the pleural
process (arm) occurs only in the meso- and meta-thorax of jumping apterygotes such
as machilids.
In Lepisma the endopleural processes, labeled p and c (Fig. 37), are separate endo-
pleurites in the prothorax, and they are fused to form pc in the mesothorax and the
metathorax. These structures are therefore not homologous with the pleural process in
Machilidae; in Ctenolepisma (Fig. 38A), a comparable pleural process or processes
are absent.
3
7
Accordjng to Handlirscb (1930), Thysanura represent neotenous wingless pterygotes.
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The anapleuron is more or less reduced in Machilidae, but probably not as. much as
Carpentier (1946b) and Barlet (1950b) believed. Using the pleural process (pleural
arm) or the point of attachment of the endosternal process, p, as a landmark that
separates the katapleuron from the anapleuron, the mesothoracic anapleuron in Petrobius
(Fig. 39A) must be the area dorsal to the pleural process. The area labeled cp (cata-
pleuron) by Carpentier ( 1946b) is largely anapleuron. The anterior triangular area
demarcated by a ridge is the trochantin.
38
The posterior part of the anapleuron is not
distinctly separated from the katapleuron. The propleuron of Petrobius also can be
reinterpreted, as suggested in Fig. 38B. What Carpentier labeled cp ( catapleuron) and
so (sc/erite oblique) are probably ana pleural. The trochantin has no clear articulation
with the coxa in the prothorax.
If what Barlet ( 1950b) called apodeme pleural in the mesothorax of adult Dilta was
meant to be the homologue of the internal pleural ridge in the Pterygota, its anterior
position, as Barlet himself admitted, is quite abnormal. It seems more reasonable to
interpret this area as homologous with the katepisternum and the trochantin in Petrobius
(anterior part of the katapleuron in Fig. 39A), and the point of articulation with the
coxal margin (trochantin of Barlet) is then homologous with the anterior point of
articulation, A, in Lepismatidae (Fig. 38A) and in Petrobius (Fig. 39A).
In Lepismatidae the pleural process as a landmark is absent. However, the division
of the pleural wall into the anapleuron and the katapleuron can be easily recognized,
since the two pleural arcs are more clearly separated from one another (by the paracoxal
suture) than in Machilidae. In Ctenolepisma (Fig. 38A) the precursor of the anapleural
suture separates the membranous ventral part of the anapleuron, which corresponds to
the preepisternum in the Pterygota. In all three genera including Ctenolepisma
(Carpentier 1946b), Lepisma (Barlet 1951), and Nicoletia (Barlet 1952a), the ana-
pleural ring ventrally reaches the lateral margin of the basisternum, and the endosternal
tendon, h, lies on the border between the two areas.
The interpretation of the katapleural parts in Lepismatidae is somewhat difficult,
because of the presence of two different structures that have been called the trochantin.
In the mesothorax of Petrobius (Fig. 39A) the katapleuron extends posteriorly and
provides a condyle C that articulates with the posterior end of the marginal area of the
coxa, which Carpentier ( 1946b) called the trochantin. In the prothorax of the same
species (Fig. 3 8B) , the corresponding coxal marginal area is thicker. A conspicuous
structure, which Carpentier called the coxal apophysis (apophyse coxale), lies im-
mediately posterior to this area.
A comparison of these structures with the corresponding areas in Lepisma (Fig. 37)
and Ctenolepisma (Fig. 38A) shows that in these genera the coxal apophysis (processus
coxal of Carpentier, 1955) and the marginal thickening of the coxa (the trochantin of
Carpentier in Petrobius) are obviously turned into the coxal cavity. Actually the
position of the trochantin in Lepismatidae agrees with those of the true trochantin in
Machilidae that lies above the thickened, turned out coxal margin.
Barlet ( 1951) showed three points of articulation of the trochantin with the coxa
in Lepisma (Fig. 37). In Nicoletia, according to Barlet (1952a), the median and the
posterior (Band C in Lepisma) articulations are clearly present; in Lepisma the posterior
condyle C articulates with the spinous coxal apophysis (Fig. 37). .
In Lepismatidae the katapleuron is distinct from the anapleuron, and there is a
considerable space above the dorsal margin of the trochantin. This space corresponds to
the antecoxale in the Pterygota. In Ctenolepisma (Fig. 38A) the katapleuron has an
articulation with the dorsal margin of the trochantin.
In Lepismatidae the katapleural ring proximally occupies the space between the
base of f (homologue of the furca in the Pterygota) and the inner coxal margin. In
Ctenolepisma f may appear to be captured by the katapleuron, as Carpentier (1947)
38Carpentier did not name this area, but he named the narrow anterior part along the coxal margin the
trochantin. This area is a part of the coxa, or one of the two kinds of the trochantin to be discussed shortly.
PRESTERNUM -+---
BASISTERNUM --+---
KATAPLEURON
FURCASTERNUM
THICKENED
COXAL MARGIN
PRESTERNUM ?
ARACOXAL SUTURE
AN EPISTERNUM
TROCHANTIN
-------SPINASTERNUM
A
TROCHANTIN
B
COXAL PROCESS
(APOPHYSIS)
FIG. 38. A, mesothoracic pleuron and sternum in Ctenolepisma sp. (from Carpentier and
Bar let); B, prothoracic pleuron and sternum in Petrobius balticus (from Carpentier).
noted in Tomocerus." However, f is on the anterolateral angle of the furcasternum, and
meets the katapleural extension.
The sternum: The endosternal tendons (stalk, bridge, plate) occurring in various
genera of Thysanura are as follows:
a connects the middle of the posterior margin of the spinasternum (poststernite of
Carpentier, 1946b) and the endosternal plate above.
b is the median unpaired endosternal plate above the ganglion, and refers to the
area anterior to the point where a arises (Fig. 39C).
c is a furcal projection behind g in the prothorax of Petrobius (Fig. 39C). Barlet
used c also for the katapleural tendon in Lepisma (Fig. 37).
d is a postcoxal bridge from the anterolateral angle of b that reaches the lateral end
of the spinasternum (poststernite of Barlet, 1952a, b). It is double in Nicoletia
(Fig. 39B).
e lies on the lateral margin of the furcasternum behind f.
f lies on the anterolateral angle of the furcasternum, and supports g. It meets the
posteriorly extending katapleuron. In the prothorax of Petrobius (Fig. 38B), e and f
are probably indistinguishably fused (Carpentier 1946b), or f has become lost. It is
double in the mesothorax of Nicoletia (Fig. 39B).
g is a common plate from which f, h, i, j, k, and p arise. In the prothorax of
Petrobius (Fig. 39C) g is greatly reduced, and gives rise to only h, p, and k.
h connects g and the border area between the basisternum and the descending ana-
pleuron. It is double in the meso thorax of Lepisma (Fig. 3 7).
i is a bridge that connects g and the posterior coxal margin near d.
j is a narrow bridge that connects g and the median trochantine-coxal articulation in
Ctenolepisma (Fig. 38A). In Lepisma muscle 33 (Fig. 20) is the corresponding
structure.
k is a long stalk from g to the tergum (Fig. 3 SA).
1 is dependent on the poststernum and the presternum of the succeeding segment,
and dorsally supports the endosternal plate m. l is absent in the metathorax of Nicoletia
and perhaps in others. l demarcates the intersegmental border.
m is the posterior part of the unpaired median endosternal plate.
n is a posterior lateral stalk from m that reaches near the spiracle.
p arises from g, and reaches the dorsal margin of the katapleuron. It is present in
the prothorax of Nicoletia and Petrobius, and in all segments of Lepismachilis.
q is an expansion mesad of g, present in the prothorax of Nicoletia (Fig. 39B).
t is a small pillar that connects b and the point of divergence of the branch of Y on
the sternum in Ctenolepisma (Fig. 38A).
v is, according to Barlet (1967), a vertical tigella that arises from m in the pro-
thorax of Lepismachilis; v was used also for an endosternal part of the first abdominal
segment in Lepisma (Fig. 37).
v', v'' are other vertical tigellas, which arise from m in Lepismachilis and lie posterior
to the base of n.
w is a triangular area formed between v' and v" in Lepismachilis.
In the mesothorax of Petrobius (Fig. 39D) b and m are free, being medially dis-
continuous. In the metathorax of the same species, apparently b and m are also free, as
in the mesothorax. Carpentier (1949), however, interpreted the posterior transverse
stalk in the metasternum as a fusion of b and m, because it is attached ventrally to the
poststernum and the presternum of the succeeding segment, and called the anterior
transverse stalk q (not present in the mesothorax). If the ventral points of attachment
of the tendons are the infallible and only landmarks, Carpentier's interpretations are
justified. However, if the positions of these stalks and the neighboring tendons relative
to d are used as landmarks, the anterior stalk (q of Carpentier) in the metathorax is b,
30
It was this appearance, remarked by Carpentier (1947), that misled Weber (1952) to formulate an unaccep-
table theory (p. 30).
109
~ - - - KATAPLEURON
PLEURAL
PRO C ES:S---'t''/
TROCHANTIN
n
i+d
k
Fro. 39. A, endoskeleton of mesothoracic pleurostemum in Petrobius balticus (from
Carpentier); B, prothoracic endostemum in Nicoletia neotropicalis (from Barlet); C, pro-
thoracic endostemum in Petrobius balticus (from Carpentier); D, meso- and meta-thoracic
endostemum in Petrobius baltic us (from Carpentier).
110
PSEUOOPRESCU
BASIST
STERNACOSTA
FURCASTERNUM
l<ATAPLEURON
PRESTERNUM
ANAPLEURON
STERNACOSTA
SPINASTERNUM

ERNUM
POSTSTERNUM
PRESTERNUM
SPIRACLE
ANAPLEURON
RNA COSTA
SPINASTERNUM
FIG. 40. Thoracic sternum in Nicoletia neotropicalis (from Barlet).
because it is laterally continuous with d and p, just as b is in the mesothorax; and the
posterior transverse stalk should be m (bm of Carpentier).
In Lepismachilis (Fig. 35) Barlet (1967) showed a tubercle on the middle of the
anterior margin of the prothoracic sternum, and called it the spina. Maki (1938) also
showed such a spina on the corresponding area in Pedetontus. The coxal muscle that is
attached to this tubercle (81 in Lepismachilis, Fig. 35) is apparently s-ex 1. If this
structure is the labial spina homologous with the one in Diplura, as the associated muscle
suggests, its more posterior position must be due to shifting. In Nicoletia the labial spina
is absent, but the anteriormost plate (presternum 1 of Barlet, 1952a) may represent the
labial sternum, comparable with that in Oncojapyx (Fig. 34).
In Nicoletia and Ctenolepisma the basisternum is not evaginated, and the division of
the sternum in the posterior thoracic segments is more clearly recognized than in other
genera. In Nicoletia (Fig. 40) the presternum is clearly separated from the basisternum
by a transverse line. A line reaching h in Petrobius (Fig. 38B) may not be homologous
with the transverse line under consideration. In Lepisma (Fig. 37) and Ctenolepisma
(Fig. 38A) this line of demarcation is lost. It should be noted that the presternum
extends laterally and lies anterior to the pleuron in all genera.
The basisternum meets the descending anapleuron laterally, and the endosternal h
lies on the border line between the two areas. In Nicoletia a transverse line, which
represents the sternacosta, connects f on each side in the pro- and meso-thorax, and it
separates the basisternum from the furcasternum. The sternacosta in Nicoletia does not
form a distal condyle that articulates with the coxa; it may be present also in the pro-
thorax of Petrobius (Fig. 38B). In other genera the border between the furcasternum
and the basisternum is obliterated, the stern acosta being absent. The desclerotized Y
line, which may be represented by a ridge in other insects, is recognized in Ctenolepisma
(Fig. 38A).
In Nicoletia (Fig. 40) the spinasternum (poststernum of Barlet, 1952a) is clearly
separated from the poststernum by a complete transverse line; in contrast to Oncojapyx
(Fig. 34), the spinasternum is not lost medially. The endosternal a is associated with the
spinasternum, and I is associated with the poststernum (presternite 1 of Barlet, 1952a).
In Ctenolepisma (Fig. 38A) the same division of the posterior part of the sternum occurs.
Here, however, one of the double /'s is attached to the presternum of the succeeding
segment, and another is on the poststernum. In Lepismachilis, according to Barlet
( 1967), the metathoracic poststernum (intersternum of Barlet) is associated with a and
/, and the metathoracic spina is present, although they are not clearly shown in Fig. 35.
It is evident from the foregoing discussion that the primitive five divisions of the
sternum are retained in Nicoletia; in other genera studied the divisions are obliterated in
varying degrees in different parts of the sternum.
The adult musculature: Malci (1938) studied the thoracic musculature of
Pedetontus sp. and Lepisma saccharina; and Argilas ( 1941) studied the same in Dilta
littoralis. Since Barlet's (1953, 1954, 1967) studies on Lepisma saccharina (Figs. 19,
20, 21) and Lepismachilis Y-signata are obviously more reliable than the previous works,
the muscles in the last two genera are homologized in Table III. The two species
represent Lepismatidae and Machilidae.
The thoracic musculature in Lepismachilis differs conspicuously from that in
Lepisma in the possession of some torsion muscles (34-37, 42-47). As in Lepisma,
muscles strictly within the pleuron are absent in Lepismachilis.
THE EPHEMEROPTERA
The following discussion on external structures refers mainly to Hexagenia recurvata
(Knox 1935) and Siphlonurus columbianus (Matsuda 1956b).
The neck-region: The cervical sclerite in Siphlonurus (Fig. 41A) is represented by
the lateral cervical sclerite, which connects the ventral lateral angle of the postoccipital
ridge and the anterior part of the episternum. Because of the short neck membrane, the
undivided lateral cervical sclerite lies almost vertically. In H exagenia also, only the
lateral cervical sclerites occur. The lateral cervical sclerite is probably presternal in
origin, since it lies anterior to the presumed preepisternal area (p. 114 and Fig. 41B).
The pronotum: The pronotum is a simple plate hanging over the pleuron on each
side; it is connected with the pleuron by membranous infolding. In Siphlonurus an
elongate, heavily sclerotized area is present on either side of the median longitudinal axis.
Ide (1936) described the posterolateral expansion of the pronotum in the nymph of
Ecdyonurus.
The propleuron: In Siphlonurus (Fig. 41A) the propleuron is divided into the
anterior episternum and the posterior epimeron; in Hexagenia a distinct pleural suture is
absent. In Siphlonurus the narrow trochantin is present and has an anterior articulation
with the coxa (Fig. 41B). Matsuda (1956b) called this plate the katepisternal plate,
but he now believes it is the trochantin that is distally fused with the katepisternum; its
TABLE III
Thoracic musculature in Thysanura
Lepisma Lepisma
(Barlet 1952, Lepismachilis (Barlet 1952, Lepismachilis
1953) (Bar!et 1967) Remarks 1953) (Barlet 1967) Remarks
1
2
3
4
5
6
7
8
9
11
12
13
14
15
16
17
18
19
20
21
25
26
27
31
32
33
34
35
36
37
38
41
42
43
44
45
46
47
48
Ter!l,al muscles
18
19? 1
8, 21 2
5, 14, 15, 24 3
6
7, 13, 20
4
2, 3
1 4
10, 16 5
9, 22 6
11,12,17,23 7
Sternal muscles
25, 30, 40
26?, 29, 39 8
28, 33a,b, 41
32, 55? 9
27 10
31, 38 11
48 12
34-37, 42-47 13
Pleurosternal muscles
60? 14
Sternocoxal muscles
81, 86, 88 15
84
85, 87
82 16
83 17
Tergosternal muscles
68
49, 52, 61, 62 18
63, 70, 71
51, 53?, 59
65b
79, 80
49
50, 54
56, 69, 78
57, 58, 72
64
65a, 74
66, 75, 76
73
111 67, 77
50
51 126
52 123
53 127, 128, 130, 131
54
55 124
125
129, 132
Ter!l,ocoxal muscles
61 98
62 97, 102, 106
63 99, 103, 107
64 104, 108
65 100?
66
95, 101, 105
Tergotrochantinal muscles
71 93, 96?
72 89, 91, 94
73 90, 92
74
TerJtopleural muscles
81
82 109, 118
83 113, 118?
84 114, 119
85
86
87
88
89
90
91 117?, 122?
92
93
111
91 112?
110, 115, 120
116, 121
Pleurocoxal muscles
101
102
133, 134
135
136, 137
19
20
21
22
23
24
25
26
27
28
29
30
31
31
32
33
34
35
36
37
38
39
REMARKS (Table III)
The following remarks refer to the muscles in Lepismachilis in Table III.
( 1) 19 is attached posteriorly to the pseudophragma of the first abdominal
segment.
(2) 8 and 21 are inserted posteriorly on the pseudophragma of the succeeding
segment.
(3) 5 arises anteriorly from the occipital sclerite, and 14, 15, and 24 from the
middle of the tergum.
( 4) 1 extends between the occipital groove and the middle of the metanotum.
(5) These muscles extend between the pseudophragma and the middle of the
tergum of the succeeding segment.
(6) These muscles extend between the pseudophragmata of two neighboring
segments. t 18 in Odonata may be homologous with these muscles.
(7) These muscles arise from the pseudophragma and are inserted on the ridged
anterior margin of the succeeding segment. t 19 in Odonata is probably homologous
with these muscles.
(8) The anterior point of attachment of 26 is more lateral than the others.
(9) The anterior point of attachment of 55 is on the postgenal sclerite, and its
posterior attachment is endosternal c. op-s 2 in the Pterygota is probably homologous
with 55.
(10) 27 extends between the tentorial lobe and the furcal area of the prothoracic
endosternum, and lies close to 26.
( 11) 31 extends between the lateral palette (palette laterale, roughly n) and the
base of the mesothoracic endosternum, and 38 between v" and v' of the meso- and
meta-thoracic endosterna.
(12) 48 connects the median longitudinal ridge of the basisternum with the endo-
sternal process p of the metathorax. This muscle was also described in Petrobius
(Fig. 39D).
(13) These muscles are twisted muscles. 34-37 connect the prothoracic spina or
m with mesothoracic n posteriorly; and 42-47 stretch between the proximal region of the
mesothoracic endosternum and the anterior surface of n of the metathoracic endosternum.
(14) 60 connects the membranous lateral expansion n' (arising from v) and the
lateral area of the mesonotum immediately above the dorsal margin of the pleuron.
This muscle was described as a tergopleural muscle.
( 15) 81 arises anteriorly from the middle of tqe prothoracic presternum (spina
postcephalique of Barlet).
(16) 82 is a prothoracic muscle that extends from the endosternal area c to the
tip of the trochantin.
(17) 83 is a prothoracic muscle that extends from near f to the tip of the
trochantin.
( 18) These muscles are inserted posteriorly on or near the pseudophragma of the
succeeding segment.
(19) Both muscles arise from the anterior part of the pronotum. 50 is inserted
on the postlabial sclerite, and 54 on the postgenal apophysis. t-cv 2, 3 in Pterygota are
probably homologous with these muscles.
(20) 56 and 69 extend between the mesonotal pseudophragma and v' of the pro-
thoracic endosternum. .They may be serially homologous with 50 and 54.
(21) These muscles connect the middle of the mesonotum and the lateral palette
(n?) of the endosternum of the preceding segment.
(22) 64 is a mesothoracic muscle that connects the basisternum near h with the
median lateral area of the mesonotum; t-s 10 in Odonata is probably homologous with
this muscle.
(23) These muscles may be tergopleural muscles. Barlet ( 1967) was uncertain
about their dorsal points of origin.
114 MEMOIRS OF THE ENTOMO.LOGICAL SOCIETY OF CANADA
(24) 66 and 75 in the meso- and meta-thorax connect the posterior part of the
tergum and the endosternal n of the same segment. t-p 1, 2 in the Pterygota are
probably homologous with these muscles.
(25) 73 is a metathoracic muscle that connects the presternum and the anterior
part of the tergum. This muscle may be homologous with 48 in the prothorax of
Lepisrna.
( 26) 125 connects the lateral palette of the prothoracic endosternum and the
trochanter.
(27) These muscles are epimero-trochanteral muscles. p-tr 3 in the Pterygota is
homologous with these muscles.
(28) 100 is inserted on the tip of the coxal process.
(29) These muscles may possibly be homologous with 62 in Lepisrna, but they
are iPserted on the coxal margin anterior to the pleural process.
(30) 96 is not inserted on the tip of the trochantin.
( 31) 118 arises from the anterior ridge of the metanotum and appears to be
serially homologous with 109 in Barlet's figure. Barlet homologized this muscle with ~ 1 1 3 .
(32) These muscles are intraparanoto-anapleural muscles. They may not be
homologous with 91 in Lepisrna.
(33) 111 is an oblique muscle that connects the occipital sclerite with the
posterior end of a large, dorsal anapleural sclerite in the prothorax.
(34) 112 connects the pronotum and the dorsal anapleural sclerite as 111; it may
be homologous with 91 in Lepisma.
( 35) These muscles are posterior katapleuro-notal muscles.
(36) These muscles extend between the tergum and the pleural processes. Barlet
( 1967) was uncertain about their dorsal points of attachment.
(37) These muscles are anapleuro-katapleuro-coxal muscles.
(38) 135 is an anapleuro-coxal muscle that lies more posteriorly. p-cx 8 in the
Pterygota is probably homologous with this muscle.
(39) These muscles are epimero-coxal muscles that are inserted on the posterior
wall of the coxa. p-cx 9 in the Pterygota is probably homologous.
proximal end articulates with the coxa. The plate has a muscle that originates on the
pronotum (t-ti 2). In no other species of the mayfly has the trochantin been found.
The prosternum: In Siph/onurus (Fig. 4JB) the sclerotized part of the sternum is
represented by the basisternum, which is demarcated posteriorly by the sternacosta that
connects the furcal pits. The definitive basisternum has an anterolateral extension on
each side. This lateral expanded area probably represents the preepisternum that has
become indistinguishably fused with the basisternum, since it lies directly anterior to the
katepisterno-trochantin. In Hexagenia a transverse line demarcates the marginal strip
along the anterior margin of the expanded area. Knox ( 1935) called this area the pre-
sternum, but the area is probably the secondarily formed area, comparable with the
prepectus in other insects. The furcal pits bear conspicuous furcae internally. In neither
genera does spina occur in the large posterior membranous sternal area. In Siph/onurus
the sclerotized anterior limit of the mesopleuron continuous from the prealar arm
roughly demarcates the posterior limit of the membranous prothoracic sternum.
The mesonotum: In Ephemeroptera the mesonotum is strongly elongated to accom-
modate huge dorsal longitudinal muscles ( t 14). It consists of the acrotergite ( antecosta
of Knox), prescutum, scutoscutellum, and postnotum. In Hexagenia the acrotergite
bears two small lobes that project upward and serve as the point of attachment of the
dorsal longitudinal muscles of the prothorax. The paired first phragma appears to be an
extension from the anterior margin of the acrotergite, judging from the figures given by
STERNACOSTA
B
PARACOXAL SUTURE
KATEPISTERN
FURCA
STERNACOSTA
KATEPISTERN
c
115
BASISTERNUM
TROCHANTINOCOXAL
ARTICULATION
fURCASTERNUM
PREEPISTERNUM
BASISTERNUM
COXA
FURCASTERNUM
+MET A STERNUM
BASISTERNUM
FURCAL PIT
FURCA
FIG. 41. Thoracic structures in Siphlonurus columbianus (from Matsuda). A, lateral view
of prothorax; B, ventral view of prothorax; C, ventral view of pterothorax.
Knox (1935) and M a t ~ u d a (1956b; Fig. 42C) and from observation of this area in a
mayfly species (Hexagenia?). The prescutoscutal suture runs marginally in the anterior
third of the mesonotum both in Hexagenia and in Siphlonurus. The prealar arm or
bridge, arising from the prescutum, extends to the anterior limit of the pleuron
(Fig. 42B). In Hexagenia, according to Knox (1935), the prescutum articulates
posteriorly with the basalare, which in tum supports the humeral plate. Grandi (1947b)
found, however, that in Ephemeroptera the basalare articulates with the tegula, which
articulates with the humeral plate, as in Siph/onurus (Matsuda 195 6b).
The posterolateral scuta! suture cuts the posterior notal wing process on the lateral
margin, and proximally reaches the recurrent scutoscutellar suture (Fig. 42C). This
suture has been erroneously called the scutoscutellar suture by previous workers, including
Knox (1935), Snodgrass (1935), Grandi (1947b), and Matsuda (1956b). In
Ephemeroptera the scutoscutellar suture is absent, and hence, the division of the tergum
into the scutum and scutellum is absent. The scutoscutellar suture is also absent in
Thysanura and Plecoptera, and hence its absence probably represents a primitive
condition.
In Hexagenia (Fig. 42A) the anterolateral scuta! suture demarcates the narrow
marginal strip below the prescutoscutal suture, and extends posterodorsally along the
tergal fissure, although it nearly meets the prescutoscutal suture in front of the anterior
notal wing process. The lateral parapsidal suture also approaches the area where the
prescutoscutal suture and the anterolateral scuta! suture nearly meet. The round area
shown by a broken line in Fig. 42A represents the area where these three sutures
converge. In Hexagenia the anterolateral scutal suture and the lateral parapsidal suture
secondarily extend forward and backward. In Siphlonurus (Fig. 42B) these three
sutures are apparently merged into a single line, and the recurrent scutoscutellar suture
is also anteriorly contiguous to the lateral parapsidal suture. The presence of the lateral
parapsidal suture in many mayflies is evident in the drawings by Grandi (1942, 1947b).
The postnotum is represented by a transverse sclerite lying beneath the posterior end
of the scutoscutellum (Fig. 42A, B). It extends downward towards the membranous
epimeral area in Siphlonurus. The anterior wall of the second phragma is continuous
posteriorly from the postnoturn, and it is united with the posterior wall of the second
phragrna corning from the metanotum (Fig. 43A).
The axillary sclerites of the forewing: In Siphlonurus (Fig. 42C) the first axillary
sclerite is connected with the anterior notal wing process; the second axillary sclerite lies
posterolateral to the first. The narrow third axillary sclerite lies behind the second
axillary sclerite, and is articulated with the detached posterior notal wing process. Thus
in Siphlonurus there is nothing unusual regarding the axillary sclerites, and their arrange-
ment is comparable with those in other orders.
In a series of species of Epherneroptera Grandi (1947b) erroneously interpreted the
base of the median plate as the third axillary sclerite. In Ecdyonurus (Fig. 44A) Grandi
studied the third axillary sclerite (sclerificazione semilunare of Grandi) which is fused
with the scutallateral margin, and this third axillary area has the same position relative to
the second axillary sclerite and the posterior notal wing process, as the third axillary
sclerite in Siphlonurus. The median plate (third axillary sclerite of Grandi) lies parallel
to the lateral (distal) margin of the second axillary sclerite, and it is undivided (into
proximal and distal plates); this is evidently associated with a vertical position of the
wing at rest. In other Ephemeroptera that Grandi (1947b) studied, the axillary
sclerites have evidently undergone further modifications and reductions, and the third
axillary sclerite has become lost in many mayflies. Grandi also noted the presence of a
cephalocaudal inflexion between the scuta! lateral margin and the first axillary sclerite
(inflessione ascellare of Grandi), along which the forewing is flexed (Fig. 44A).
The apparent peculiarity of the wing base that resulted from the misinterpretations
led Grandi to propose the concept of the pseudopteralia, which she thought was peculiar
to Ephemeroptera. Bekker ( 1954) also studied the wing base of a Siphlonurus sp. His
identifications of axillary sclerites are, however, incorrect. What he considered to be
the first axillary sclerite is most probably the second. His second axillary sclerite is
obviously the base of the median. plate and so on.
The mesopleuron: In Siphlonurus (Fig. 42B) the pleural suture is distinct. The
apparent paracoxal suture on the episternum is well marked, and extends into the
epimeron after reaching the pleural suture. The ventral end of this suture is far in front
of the coxa, which is atypical if this suture is really homologous with the paracoxal
suture in other insects. In Hexagenia the anterior margin of what Knox (1935) called
117
PRESCUTOSCUTAL SUTURE
ACROTER'GITE
PHRAGMA
ANTERIOR NOTAL WING PROCESS
PRESCUTOSCUTAL +
ANTEROLATERAL SCUTAL +
LATERAL PARAPSIDAL+
RECURRENT SCUTOSCUTELLAR
suTURES
ACROTERGITE

ANEPISTERNUM +
PREEPI
PRESCUTOSCUTAL
A
B
SUTURE-------H

------- _/ -----,
LATERAL

SCUT AL SUTURE J
RECURRENT SCUTOSCUTELLAR
SUTURE-------'""---\
!1------POSTNOTUM
ANTECOSTA
BASAL ARE
TEGULA
HUMERAL PLATE
SCLERITE
'\. Z'ND AXILLARY SCLERITE
3'RD AXILLARY SCLERITE
NOTAL
WING PROCESS
MEDIAN LONGITUDINAL
SUTURE
FIG. 42. A, lateral view of mesothorax in Hexagenia recurvata (from Knox); B, lateral view
of mesothorax in Siph/onurus columbianus (from Matsuda); C, dorsal view of mesothorax in
Siph/onurus columbianus (from Matsuda).
the pJeurotrochantin probably represents the paracoxal suture; this suture has the typical
curvature of the paracoxal suture ventroposteriorly, though it has become lost in the
epimeral area. The anapleural cleft is absent in both genera. Hence, the anapleural
plate is a single continuous plate that is ventrally separated from the basistemum by a
narrow membranous area (pleurosternal cleft) in Siphlonurus (Figs. 41C, 42B).
The katepisternal plate is separated from the coxal margin by the intervening
membranous area, and the trochantin is absent both in Siphlonurus and in Hexagenia.
In H exagenia the episternal sclerotization extends between the basalare and the
spiracle, and the corresponding area in Siphlonurus is membranous. The mesothoracic
spiracle lies clearly within the segment, and lies more ventrad than in other orders. The
basal are is connected with the tegula in many species of Ephemeroptera (Grandi 194 7 b).
The epimeral area in Siphlonurus is sclerotized along the paracoxal suture. In
H exagenia a transverse epimeral sclerotization above the coxal base forms a postalar
bridge by fusion with the postnotum. The subalare in Ephemeroptera is a large sclerite
bearing an apodeme, or apodemes, for the attachment of muscles.
' The mesosternum: Both in Siphlonurus (Fig. 41C) and Hexagenia the sternacosta
connecting the furcal pits separates the basisternum from the furcasternum. The large
furcasternal area is simple and its posterolateral angles are met by the metapleural para-
coxal ridges. This broad, definitive furcasternal area may therefore be partly metasternaL
The mesothoracic spina is absent in Ephemeroptera.
The metanotum: The prescutum and the antecosta are not differentiated on the
anterior margin in Siphlonurus and Hexagenia. In Siphlonurus the paired posterior wall
of the second phragma develops directly from the anterior margin of the metanotum; it
(Fig. 43A) extends sharply downwards and is united with the phragma from the meso-
thoracic postnotum. As in the meso thorax, the scutoscutellar suture is absent in the
metanotum. The oblique posterolateral scuta! suture that cuts the posterior notal wing
process occurs in the metanotum of Siphlonurus. The postnotum is clearly present
behind the metathoracic scutellum. As in the mesonotum, the postnotum extends
laterally to form a postalar bridge in Siphlonurus and Hexagenia.
The metap/euron (Fig. 43B): The anapleural area in Siph/onurus is strongly
reduced and membranous, and the metathoracic spiracle clearly lies intrasegmentally.
In H exagenia the ana pleural sclerotization extends forward to encircle the spiracle. The
pleural suture is distinct in Siphlonurus, and it is obliterated in Hexagenia. In Siphlo-
nurus (Fig. 43B) the apparent paracoxal suture is recognized only on the episternal area
and it represents the anterior ventrolateral border of the sclerotized metepisternum. In
neither genera is the trochantin present in the membranous area between the katepi-
sternum and the coxal base. The basalare is unrecognized in both, whereas the subalare/
~ ~ ~ /
The metasternum: In Siphlonurus (Fig. 41C) tl:le metasternum is a single sclero-
tized plate, the sternacosta connecting the furcal pits being absent. The posterior end
of the sternal plate is recognized by the presence of a transverse suture that separates the
abdomen in Siphlonurus; the suture is lost in Hexagenia.
The anatomical basis of flight: The special characteristic of the flight mechanism in
mayflies lies- in the greater numbers of direct flight muscles. Besides the indirect flight
muscles (t 12, t 14, t-p 5, 6, t-tr 1 ), which are well developed and probably function in
a similar manner to those of other insects, Grandi (194 7 b) recognized the following
mesothoracic muscles that are concerned with flight (Fig. 44B): two m. prosterno-
basalare (p-s 12), m. episterno-scutale (t-p 10), m. pleurocondilo-scutale (p 5), m.
coxo-ascelare (t-cx 4), m. trochantero-ascellare (t-tr 2), m. furco-ascellare (t-s 3),
m. sterno-subalare (t-s 5), m. /urco-subalare (t-s 4), m. coxo-subalare (t-cx 8), m.
pleurocondilo-scutellare (t-p 15), Grandi explained the functions of these muscles as
follows:
PLEURAL WING
PROCESS
PARACOXAL
SUTURE,
A
8
POSTERIOR WALL
of 2'ND PHRAGMA
119
UTELLUM
EURAL SUTURE
FIG. 43. Thoracic structures in Siph/onurus columbianus (from Matsuda). A, dorsal border
between mesothorax and metathorax; B, lateral view of metathorax.
Since the basalare is connected with the tegula, and the tegula with the humeral
plate, contraction of the paired basalar-prosternal muscle (p-s 12) causes extension of
the costal region of the wing.
t-cx 4 and t-tr 2 are inserted dorsally very close to each other on the first axillary
sclerite; in 0/igoneuriella and Caenis t-cx 4 is attached to a small supplementary plate
anterior to the first axillary sclerite. These muscles are mainly responsible, when at
rest, for holding the wing vertically along the axillary inflexion.
t-s 3 also varies in the dorsal point of its attachment. Thus, in Ecdyonurus it is
dorsally attached to the point of articulation of the second axillary sclerite with the third
axillary sclerite. As the second and third axillary sclerites of Ecdyonurus lie per-
pendicular to the dorsal surface of the body, the contraction of this muscle keeps the
wing vertical when at rest. In Oligoneuriella neither t-tr 2 nor t-s 3 is present, and the
wings are not raised vertically. In Caenis the t-s 3 is attached to the anterior notal wing
process without having a direct relation to the second axillary sclerite; the contraction of
this muscle therefore causes only a slight flexion of the wing.
Bekker ( 1954) emphasized the importance of the subalar muscles, which he thought
were responsible for lowering the wings. Bocharova-Messner (1965) thought that in
Baetis the pleural arm-prealar bridge muscle (t-p 10), pleural ridge-basalar muscle (p 5),
and a pleurocoxal muscle (p-cx 5) were directly concerned with the movement of wings.
These explanations of the flight mechanism did not refer to Grandi ( 1947b).
According to Edmunds and Traver (1954), some of the concave wing veins have
weakened spots called bullae, which allow the concave veins to bend; consequently during
the upstroke, these concave veins fail to support the distal half of the wing. On the
upstroke, the distal half of the wing then bends downward under the presence of the
opposing air. The opposing air slips away in such a manner that itoffers lessened
resistance to the upstroke of the wing. In such a flight the prosterno-basalar muscles
( p-s 12) most likely help keep the anterior margin of the wing rigid.
The adult musculature: The thoracic musculature of Ephemeroptera was studied by
Diirken ( 1907) in Centroptilum luteolum and Ephemerella ignitq, by Knox (1935) in
Hexagenia recurvata, by Maki (1938) in Ecdyonurus hyalinus, by Grandi (1947b) on
direct flight muscles in various species, by Bekker ( 1954) on some muscles in Siphlo-
nurus, by Matsuda ( 1956b) in Siphlonurus columbianus, and by Bocharova-Messner
( 1965) on some muscles in Baetis. Maki homologized muscles in the species studied by
Diirken, by Knox, and by himself. In Table IV homologies of muscles in Siphlonurus
(Matsuda 1956b), Ecdyonurus (Maki 1938), and Hexagenia (Knox 1935) are given.
The sternocoxal muscles in Ecdyonurus are not included.
The presence of large numbers of tergosternal muscles is definitely a primitive
feature of the thoracic musculature in Ephemeroptera, because such muscles are also
abundant in the Apterygota. Of these, t-s 3 and t-s 7 are probably the muscles inherited
from the wingless ancestor. t-s 4, t-s 5, and t-s 6 are present only in Ephemeroptera,
and may have been secondarily acquired. Other muscles peculiar to Ephemeroptera
include t 17, p 5,s-cx 8?, s-ex 9, p-s 11, and p-s 12; t-cx 4 is shared only by Ephemerop-
tera and Odonata.
t 14, t-p 5, 6, and t-cx 8, which are important flight muscles in other pterygotes
(except for Odonata), are also well developed in Ephemeroptera. t-p 13 and t-p 14,
which are flexors of the wing in other insects, are present in primitive genera, Siphlonurus
and Ecdyonurus; but they are absent in Hexagenia and many other genera that Grandi
(l947b) studied. In those genera without t-p 13 and t-p 14 the third axillary sclerite is
lost and t-p 15 is usually present (in Siphlonurus and Ecdyonurus t-p 15 is absent).
Peculiarly, flight muscles p 3, p-cx 2, and p-cx 3 in other insects are absent in
Ephemeroptera.
The above features of the thoracic musculature in Ephemeroptera are quite distinct
from the thoracic musculature in Odonata (p. 399), as Diirken (1907) noticed a long
time ago.
INFLEXION
ANTERIOR
WING
PROCESS
3'RD AXILLARY
SCLERITE - - - - 4 ~ G Z ~
A
t-p 15
B
r
I
;
J
I
.J
121
2'ND AXILLARY
SCLERITE
t-s 5
Fro. 44. A, axillary region of forewing in Ecdyanurus fiuminum (from Grandi); B, flight
muscles of forewing in Ephemera danica (from Grandi).
Gross aspects of postembryonic muscle development: Knox ( 1935) described
nymphal thoracic muscles in Hexagenia recurvata. She did not trace the development
of each individual muscle. However, the following developmental pictures were found,
by comparing the accounts of the nymphal and adult muscles (although some of the
pleural muscles in the nymph could not be identified from the figures and descriptions
given). One characteristic feature of the development of thoracic muscles appears to be
that no caenogenetic trend is really evident"; most muscles that become lost in the adult
can be homologized with muscles present in the adults of other insects, and some other
nymphal muscles are transformed into the adult muscles, as shown below.
Nymphal muscles that become lost in the adult: I Pm 4 (p ?) , Im (t-s 1), II Pm 1
(t-p 3 or 4?), II Dlm 2 (t-s 9), II Vlm 1 (s 19), II Pm 11 (t-p 17), II Sm 3 (s-ex 6?),
II Sm 4 (p-s 5), III Sm 2 (s-ex 2 or 4), III pm 2 (p-cx 9).
Nymphal muscles that develop into adult muscles: by split: I Dvm 4 (nymphal
t-cx 7) ~ I Dvm 4, 5 (t-cx 7); Cv 1 (nymphal t-s(cv) 1) ~ Cv 1-5 (t-s(cv) 1 and
t-cv 2, 3); ISm 1 (nymphal furcacoxal muscles) ~ I Sm 1, 2, 3 (s-ex 2, 3, 6); II Dvm 4
(nymphal t-cx 6, 7) ~ II Dvm 4, 5 (t-cx 6, 7); by differentiation of external parts:
II Pm 6 (nymphal scutofurcal muscle) ~ I I Pm 10 (t-s 3), II Pm 10 (nymphal wing
pad- pleural ridge muscle) ~ I I Pm 6 (t-p 15); by shift of attachment: I VIm 2
(nymphal s 1) ~ I VIm 2 (cv-s 4); by unknown process: I Pm 2 (p-tr 2), II Pm 4
(t-cx 4), II Pm 5 (p 5), II Pm 11 (t-p 10), II Pm 1, 2 (p-s 12).
In Baetis Bocharova-Messner (1965) found that muscles 16 (p 5) and 17 (t-p 10)
were indistinguishable in earlier stages of development, and that only in the 14th
nymphal stage did these muscles become differentiated from the originally single muscle.
The same developmental mechanism is probably at work in Hexagenia, although this is
not possible to discern from Knox's ( 1935) study.
Bocharova-Messner (1965) also found that t-p 5, 6 (5 of Bocharova-Messner) is
absent in the first stage nymph. Thus,"the development of large t-p 5, 6 takes place in
later stages of development, as in other insects. Furthermore, according to Bocharova-
Messner, t-s 4, 5 (11, 14 of Bochamva-Messner) are already present in the first stage
nymph as the subalar-sternal muscles.
THE PLECOPTERA
The neck region: In relatively primitive Plecoptera, Pteronarcys (Fig. 45A) and
Capnia, the lateral cervical sclerite is a simple, undivided plate. Its posterior point of
articulation is with the area that corresponds, by all criteria, to the anepisternum. This
posterior point of connection in Capnia thus differs from that in Blattaria (Fig. 54A) and
related orders, in which it occurs in the area that corresponds roughly to the pre"
episternum. In Plecoptera, as discussed later, the preepisternum has probably become
fused with the basisternum as in Ephemeroptera. The presternal origin of the lateral
cervical sclerite was already discussed (p. 8).
The lateral cervical sclerite appears to be divided into the anterior and posterior
sclerites in some other Plecoptera such as Isopteryx and Chloroperla (Grandi 1950),
Perla (Wittig 1955), etc. Since the lateral cervical sclerite is undivided in relatively
primitive genera, Capnia and Pteronarcys, its divided condition in Plecoptera probably
represents a derived condition.
The dorsal cervical sclerite is present in Capnia (Crampton 1926a). The ventral
cervical sclerite appears to be represented by a single transverse plate lying anterior to
the presternum (Fig. 46A).
The pronotum: The pronotum in adult Plecoptera is a simple, broad plate. Its
anterior margin is often more or less carinated, and a transverse impression is present
near the posterior margin.
40lt is possible that some unidentified nymphal muscles are caenogenetic.
TABLE IV
Thoracic musculature in Ephemeroptera
Siphlonurus Hexagenia Ecdyonurus Muscle
(Matsuda 1956b) (Knox 1935) (Maki 1938) designations Remarks
I Dim l cv(d)-t 1
4 op-t 1
5 3 op-t 2
6 I Dlm 2 1 op-t 3
2 5 op-cv 3
11 I Pm 3 9 op-p 1
12 ISm 4 op-s 1
26, 27 op-s 2
1 Cv 1-5 6, 7 t-s(cv) 1 1
and t-cv 2, 3
3 I Vlm 2 cv-s4
I VIm 1 4 s 1 or 2
8, 9 (I) t 12
31 II Dm 21
64 (III)
10 I Dm t13
7(!) 2 t 13? 2
6 I Dim 1 1 t 14
30 II Dim 20
63 III Dim 47
13 (I) 10 t-p 3
44 t-p 3 4
43 (I I) II Dvm 1 25 t-p 5, 6
71 (III) III Dvm 1 51
45 (II) t-p 8 5
II Pm 11 t-p 10 6
51 (II)
76 (III)
t-p 13
32 (II)
57 (III)
t-p 14 7
II Pm 6 t-p 15 8
III Pm 6
29 (I I) t-p 8? 9
54 (III)
Ill Pm 2 t-p 8? 10
72 (III) t-p 8? 11
8 (HI) t-s 2 12
52 (I I) II Pm 10 28 t-s 3 13
77 (III) III Pm 4 53
39 (II) II Pm 8 27 t-s 4 14
70 (Ill)
38 (I I) II Pm 7 26 t-s 5 15
52 (III)
III Dvm 4 t-s 6
24 (II) t-s 7
50 (III)
14 (I) 11 t-ti 2
18 I Dvm 1 16, 17 t-ti 3
32 II Dvm 2 36
65 III Dvm 2 59
TABLE IV (Continued)
Siphionurus Hexagenia Ecdyonurus Muscle
(Matsuda 1956b) (Knox 1935) (Maki 1938) designations Remarks
50 (I I) II Pro 4 31 t-ex 4 16
35 (II)
67 (III)
II Dvm 5 t-ex 6
15, 16, 17 I Dvm 3, 4, 5 12, 13, 14 t-cx 7 17
33, 34 II Dvm 3, 4 39, 40
66 I!IDvm3 62, 63
37 (II) II Pm 9 41 t-cx 8
69 (III) III Pm 5
42 (II) II Pm 5 30 p 5 or t-p 4 18
III Pm 3 55 (III)
80? (Ill) p-s 11
40, 41 (I-II) II Pm 1, 2 33, 34 (II) p-s 12
19
19(I) I Pm 1 18 (I) p-ex 4
36 (II) II Pm 12 43 (I I) p-cx 5
68 (III) III Pm 8 65 (Ill)
ISm 1 s-ex 2
55 (II) 35(!1)
Ill Sm 1 58 (III)
ISm 2 s-ex 3
54 (I l)
Ill Sm 2?
25 (I) s-ex 5
56 (II)
III Sm 4
ISm 3 s-ex 6
58 (I I) s-ex 8? or s-ex 4? 20
II Sm 1 s-ex 9
57 (I-II) II vim 22 (I-II) s 13
48 (II-III)
59 (I I) II Sm 2 23 (II) s 16 21
81 (III) III Sm 3 49 (III)
20
I Dvm 2 t-tr 1
48 II Dvm 6 44 (II)
74 III Dvm 5 66 (III)
47 (II) II Pm 13 45 p-tr 1
I Pm 2 19 (I) p-tr 2
II Pm 3
21 (I) p-tr 3
REMARKS (Table IV)
( 1) Cv 1 in Hexagenia is t-s( cv) 1 that arises from the dorsal cervical membrane;
it is inserted on the anterior half of the lateral cervical membrane. All others arise from
the pronotum and they may be considered as t-cv 2, 3; though they are inserted on the
anterior part of the lateral cervical sclerite.
(2) 7 in Siphlonurus is an oblique muscle that extends between the mesothoracic
antecosta and the pronotum near its median longitudinal axis; 2 in Ecdyonurus stretches
between the dorsolateral region of the pronotum, immediately behind the faint transverse
tergal ridge and the lateral anterior end of the mesonotum.
( 3) These muscles are prothoracic muscles that connect the lateral pronotum and
the dorsal part of the propleuron.
( 4) 44 in Siphlonurus connects the anterior margin of the sci erotized episternum
and the prescutum near its anterior end.
(5) 45 in Siphlonurus may be t-p 10, as II Pm 11 in Hexagenia.
(6) II pm 11 is 3 of Grandi (1947b).
(7) 32 and 57 in Ecdyonurus are small muscles that connect the pleural wing
process and the third axillary sclerite.
(8) These muscles are 8 of Grandi (1947b).
(9) 29 and 54 in Ecdyonurus connect the episternum (apparently paracoxal
suture) and the lateral scuta! margin above the basalare. They may be homologous with
45 in Siphlonurus.
( 10) III pm 2 in H exagenia is a small muscle that extends between the pleuron
anterior to the pleurocoxal articulation and the scuta! lateral margin anterior to the
anterior notal wing process. It may be homologous with 72 in Siphlonurus.
( 11) 72 in Siphlonurus is a small muscle arising from the dorsal membranous
part of the pleuron, and it is inserted on the scuta! lateral margin anterior to the anterior
notal wing process.
( 12) 8 in Ecdyonurus may bet-s 1.
(16) These muscles are 5 of Grandi (1947b). They are inserted dorsally on
the first axillary sclerite, or on the axillary inflexion formed between the scuta! lateral
margin (anterior notal wing process?) and the first axillary sclerite, or in H exagenia on
the basalare near the articulation with the tergum.
( 17) One or two of the prothoracic muscles must be serially homologous with
t-cx 8.
(18) These muscles are 4 of Grandi (1947b) which, however, may be t-p 4. 42
in Siphlonurus connects the pleural wing process with the point where the basalare
articulates with the tegula.
(19) These muscles are 1 and 2 of Grandi (1947b). In Siphlonurus 40 and 41
are inserted on the profurca; in Ecdyonurus the ventral points o( attachment are on the
anterolateral area of the mesosternum; in Hexagenia on the posterior prosternum (furcilla
of Knox) behind the furca; in the species of Grandi (1947b) studied they are attached
on the small sclerotized prominences in the posterolateral area of the prosternum.
(20) 58 in Siphlonurus is a transverse muscle that connects the outer posterior
wall of the coxa on the two sides.
( 21) These muscles connect the furcal base on the two sides.
Besides the muscles listed in the Table IV, 6 of Grandi (1947b) is the second
tergotrochanteral muscle ( t-tr 2) that is present only in Ephemeroptera; it is dorsally
attached to the first axillary sclerite, very close to the point of attachment of t-cx 4.
According to Grandi, this muscle is absent in some species, and it is represented by a few
fibers and fused with t-cx 4 in some species. In both the meso- and meta-thorax of
Centroptilurn and Ephernere/la (Diirken 1907), the sternotrochanteral muscle (s-tr 1) is
present. In Centroptilurn 55 of Diirken is s 20. Furthermore, i dim of Diirken in
Centroptilurn is t 17, which has not been found in other winged insects.
The propleuron: In the propleuron of Pteronarcys dorsata (Fig. 45A) the pleural
suture is bent strongly forward in its dorsal half. The pleural arm arises internally from
the pit formed along the dorsal half of the pleural suture. The paracoxal suture is
probably represented by the dorsal margin of the katepisternum that meets the middle
of the pleural suture. In Pteronarcys californicus, as Snodgrass (1909) has shown, a
complete paracoxal suture extends into the epimeron. In Allocapnia (Fig. 45B) the
apparent paracoxal suture reaches the dorsal (anterior) end of the pleural suture, but
KATEPISTERNUM
PLEURAL ARM
AN EPISTERNUM
+ PREEPISTERNUM
PARACOXAL
SUTURE
A
PLEURAL SUTURE
ANEPISTERNUM
- EPIMERON
PARACOXAL SUTURE
tROCHANTIN
LATERAL -COXA
CERVICAL
SCLERITE
POSiCOXAL
BRIDGE
B
FIG. 45. A, lateral view of prothorax and cervical sclerite in Pteronarcys dorsata (original);
B, the same in Allocapnia granulata (original).
this is most probably produced by sclerotization of the shaded area that remains mem-
branous in Pteronarcys; hence it may not be the true paracoxal suture. In Capnia
(Crampton 1926a) a similar suture is apparently present. In Allocapnia (Fig. 45B),
Capnia and Eusthenia, studied by Crampton (1926a), the sclerotized postcoxal bridge
encircles the posterolateral coxal margin and extends medially almost to the furcal base.
The prosternum: In the prosternum the presternum is often clearly recognized as a
small, isolated, anterior plate (Fig. 46A), although it may be fused with the basisternum
as in Eusthenia (Crampton 1926a). Since the preepisternum tends to become fused
with the basisternum in the pterothorax, the expanded anterolateral angle of the definitive
basisternum (Fig. 46A) is probably of preepisternal origin. The furcal pits are distinct,
and the furcasternum is demarcated from the basisternum by the horizontal sternacosta
that connects the furcal bases on the two sides.
The spinasternum in Perla (Fig. 48) is a small triangular plate bearing the spina, and
in Allocapnia and Capnia (Fig. 46A) it is a narrow, transverse plate. Martin ( 1916)
and Crampton (1926a) have called the transverse plate behind the furcasternum in
Capnia the postfurcasternum. If this plate represents a primary subdivision of the
sternum homologous with that in Thysanura, the postfurcasternum of Crampton is the
127
LATERAL
CERVICAL
SCLERITE
AN EPISTERNUM
POSTCOXAL
BRIDGE
POST STERNUM
ACROTERGITE
A
VENTRAL CERVICAL
SCLERITE
PRESTERNUM
PREEPISTERNUM
SPINASTERNUM
PHRAGMA
ANTE COSTA
SUTURE
POSTEROLATERAL
SCUTAL
AXILLARY
CORD
RECURRENTSCUTOSCUTELLAR B
SUTURE
PREALAR ARM
<< ANTEROLATERAL SCUTAL
u SUTURE
-SCUTUM
FISSURE
:{.
POSTERIOR
NOTAL PROCESS
FIG. 46. A, ventral view of prothorax in Capnia sp. (from Crampton); B, dorsal view of
mesonotum in Pteronarcys dorsata (original).
spinasternum in Thysanura, and the hindmost plate bearing the spina is the poststen'lum;
this interpretation assumes a certain degree of shift caudad of the spina. n
The meso- and meta-no tum: The meso- and meta-notum are similar in Plecoptera;
the following note refers to the mesonotum, unless otherwise noted.
A characteristic feature of the mesonotum is its well-developed anterior region. In
Pteronarcys (Fig. 46B) the weakly developed phragma arises from beneath the antecostal
suture. The narrow acrotergite is represented by the area anterior to the antecostal
suture. The prealar arm is well developed and reaches ventrally the episternum in
Perla (Fig. 47B) and Pteronarcys, but not in Nemoura (Onesto 1965). In Plecoptera
the arm is clearly the lateral extension of the prescutum.
The prescutoscutal suture in Pteronarcys (Fig. 46B) and Perla (Fig. 47 A) runs
obliquely forward, and is lost medially; in the metanotum of N emoura (Ones to 1965),
however, the prescutoscutal suture is horizontal, demarcating a complete prescutum
anteriorly. In Pteronarcys (Fig. 46B) the prescutoscutal suture is continuous from the
anterolateral scuta! suture, as is often the case in other insects. The lateral parapsidal
suture arises laterally from the point of union of the anterolateral scuta! suture and the
prescutoscutal suture; Grandi (1948) erroneously called the area anterior to this suture
the prescutum. The paired parapsidal sutures arise from the antecosta! suture in
Pteronarcys (Fig. 46B).
The tergal fissure is represented by a suture in Pteronarcys and Perla (Fig. 47 A).
Often, however, this suture appears to be absent in Plecoptera. The scutoscutellar
suture is absent in Plecoptera, as in Thysanura and Ephemeroptera. The recurrent
scutoscutellar suture arising from the posterior margin of the scutoscutellum is present
(Figs. 46B, 47 A). The posterolateral scuta! suture is only weakly developed from the
base of the posterior notal wing process (Figs. 46B, 47A).
The mesonota! postnotum has well-developed, paired invaginations arising from its
posterior margin, which form the anterior walls of the second phragma. In Pteronarcys
the laterally extended part of the postnoturn unites with the posterolateral angle of the
rnesepimeron along a suture, whereas in Nemoura (Onesto 1965) it does not reach the
epimeron (Fig. 47B).
The bilobed posterior wall of the second phragma arises from the anterior margin
of the metanotum. In the metanotum of Pteronarcys the antecosta cannot be observed.
The third phragma is so weakly developed that its double-walled constitution is less
evident than in the second phragma.
The mesoscutal lateral margin and forewing base structures: In Perla (Fig. 47A)
the anterior notal wing process, as Onesto ( 1965) has shown, lies on the distal end of the
oblique anterolateral scuta! area. The antemedian notal wing process is also clear. The
emargination posterior to the antemedian notal wing process is shallow, and the oblique .
tergal fissure starts from this emargination. The median notal wing process is recog-
nizable in Perla, but not in N emoura. In many Plecoptera the scuta! lateral margin
from the median notal wing process to the base of the posterior notal wing process is
nearly straight, and lies in opposition to the long proximal margin of the first axillary
sclerite, as seen in the figures by Grandi ( 1950). Grandi's posterior notal wing process
is actually the first axillary sclerite, as Onesto ( 1965) has pointed out. The postmedian
notal process is not differentiated in Plecoptera; the posterior notal wing process is large.
In the metanotum the posterior notal wing process is often detached to be the fourth
axillary sclerite. The tegula, as Crampton (1918a) showed, occurs both in the meso-
thorax and the metathorax in Plecoptera.
Onesto's work ( 1965) shows that in Perla (Fig. 47A) the anterior end of the first
axillary sclerite is either articulated or connected with the anterior notal wing process,
with the basisubcostale, with the basiradiale and, by the tendinous ligament, with the
proximal expansion (process) of the third axillary sclerite across the parascutal mem-
4J.The shift in position of the spina occurred in many orders (p. 20).
129
PRESCUTOSCUTAL TEGULA
ANTEROLATERAL
SCUTAL SUTURE
LATERAL
PARAPSIDAL
SUTURE
TERGAL FISSURE
SUTURE
POSTEROLATERAL:.----!----!--i--1
SCUTAL SUTURE
RECURRENT
POSTNOTUM
PREALAR ARM-
EPIMERON
A
B
PREALAR ARM
BAStSUBCOSTALE
BASIRADIALE
t'ND AXILLARY
SCLERITE
PROXIMAL
MEDIAN PLATE
3'RD AXILLARY
SCLERITE
l'ST AXILLARY SCLERITE
2'ND
AXILLARY
SCLERITE
V ~ ~ : ; f 4 ~ : 3'RD
AXILLARY
SCLERITE
SUBALARE
FIG. 47. Forewing base in Perla maxima (from Onesto). A, dorsal view; B, lateral view.
branous area. The posterior half of the proximal margin of the first axillary sclerite is
closely united with the scuta] margin by very short ligaments.
Besides the articulation with the first axillary sclerite, the dorsal second axillary
sclerite is connected with the proximal median plate, with the proximal process of the
third axillary sclerite, and anteriorly with the basiradiale. The ventral second axillary
sclerite (Fig. 4 7B) is connected with the pleural wing process by a ligament. Its
associations with the ventral third axillary sclerite and with the subalare are evident.
Besides the ligamentous connection with the first axillary sclerite, the dorsal third
axillary sclerite is articulated proximally with the posterior notal wing process and is
fused with the proximal median lobe. The ventral third axillary sclerite is associated
with the subalare and with the ventral second axillary sclerite.
The sclerotized humeral plate (or basicostale) is isolated ventrally, and is connected
with the basalare (Fig. 47B).
The proximal median plate is distally separated by a membrane from the base of the
M-Cu vein complex, which indicates that the distal median plate, if formed, represents
the base of the M-Cu vein. In Plecoptera, as in Ephemeroptera, a distinct distal median
plate is not formed. In Perla a short suture appears to represent an incomplete separation
of the distal median plate from the base of the M-cu vein (Fig. 47A); in the metathorax
the separation of the plate is more advanced. The wing is folded on the narrow mem-
brane along the distal margin of the proximal median plate. The absence of the well-
differentiated distal median plate represents a highly primitive condition of the wing base.
The association of the base of the R-vein with the second axillary sclerite is clear and
needs no explanation (Fig. 47 A). The basisubcostale is ventrally united, by a tendinous
ligament, with the pleural wing process (Fig. 47B). The small basijugale in the hind
wing base has an association with the third axillary sclerite, as in Blatta (Fig. 56).
Holdsworth's ( 1942) study indicated that during postembryonic development the
first axillary sclerite and part of the third axillary sclerite are formed from the tergum,
not from the wing tissue.
The meso- and meta-p/euron: The pleuron is clearly separated by the pleural suture
into the anterior episternum and the posterior epimeron. On the episternum of Perla
(Fig. 48) an oblique suture arising from the pit on the pleural suture probably represents
an incomplete paracoxal suture that has become lost proximally. Such an apparent para-
coxal suture is present in Chloroperla and Isopteryx, as was shown by Grandi (1950).
The anepisternum has an incompletely separated basalare (Fig. 47B). The sclerotized
katepisternal area is often represented by a narrow katepisternotrochantin, and by the
area between the furcal pit and the inner basal margin of the coxa, which is the
ventropleurite.
The anapleural suture (Fig. 48) extends posteriorly to the base of the trochantin as
in other insects. The preepisternum is united with the lateral margin of the basisternum.
The line of union (the pleurosternal suture) is clearly recognized by different dgrees of
sclerotization between the basisternum and the preepisternum in Pteronarcys observed in
this study, although in Allocrania (observed) they are indistinguishably fused. In Perla
(Fig. 48) Weber (1933) showed the pleurosternal suture.
The more or less obliterated pleurosternal suture led DuPorte ( 1965) to regard the
anapleural suture as the sternopleural suture (see p. 39). If this interpretation is
followed, the basisternum extends laterally above the trochantin. No such relation of
the two areas occur in lower insects. As already discussed ( p. 40), Gouin's (1959)
interpretation that the preepisternum represents the lateral basisternum is also invalid.
The epimeron is more or less reduced, and in some Plecoptera it is united with the
lateral extension of the postnotum to form a postalar bridge. The small subalare is
present in the membrane above the epimeron.
The meso- and meta-sternum: In Perla (Fig. 48) the mesosternum consists, from
the front to the back, of the presternum, the basisternum and the definitive furcasternum
bearing the spina at its. posterior end. The presternum is often clearly separated from
MATSUDA: TiiE INSECT TiiORAX
FURCAL PIT
131
SPIRACLE
PARACOXAL
SUTURE
SUBALARE
POSTNOTUM
KATEPISTERNUM
FIG. 48. Ventral view of thorax in a plecopteron (Perla) (from Weber).
the basisternum, although in the metathorax of Perla (Fig. 48) it is apparently com-
pletely fused medially with the basisternum. In Perla a two-part sclerite lies immediately
in front of the preepisternum, and this is what is usually called the interpleurite; however,
it is probably a detached part of the presternum, since the presternum in Thysanura
and Diplura extends laterally anterior to the episternum.
In Perla (Fig. 48) the basisternum is separated from the furcasternum by the
horizontal sternacosta that connects the furcal bases. The furcal bases are also con-
nected by a posteriorly converging Y-ridge, whichreaches the spina at its posterior end.
The Y-ridge and its relation to the spina is thus remarkably similar to that in Diplura
(Fig. 34); the difference in Plecoptera is in the absence of any direct association of the
Y-ridge with the coxal margin. Compared with the mesosternum in Oncojapyx
(Fig. 34), it is evident, further, that in Perla the spinasternum and the poststernum have
become obliterated, due to membranization. In Capniidae, as Hanson ( 1946) and
Kawai ( 1967) have shown, the thoracic sternal sclerites provide generic characters.
The adult musculature: Maki (1938), Grandi (1948), and Wittig (1955) have
studied the thoracic muscles in Neoperla formosana, Perla marginata, and Perla
abdomina/is, respectively. Homologies of muscles in the first and the last species, except
forthe s-ex muscles in Neoperla, are shown in Table V.
Despite the retention of many primitive features in the external structures, the
musculature does not exhibit notably primitive features. As in some strong-flying
insects, t 14 (a dorsal longitudinal muscle) and t-p 5, 6 (dorsoventral flight muscles) are
well developed. Unlike Ephemeroptera, however, p 3 is present, and in this respect
Plecoptera is more similar to typical pterygotes with strong power of flight.
Gross aspects of postembryonic muscle development: Wittig (1955) summarized
the postembryonic development of thoracic muscles in Perla marginata as follows: The
few degenerating nymphal muscles are homologizable with the adult muscles in other
insects; other nymphal muscles are transformed into the adult muscles. Thus, there is
no caenogenetic trend.
Muscles with imaginifugal development.
Reduction (loss): avlm (s 19), I dlm m (t 11? or part of op-t 3)
Diminution: I ism 22 ( t-s 1), II ism 44 ( t-s 1), II tpm 48 ( t-p 13)
Muscles with imaginipetal development.
New formation: II im 65a (p 1)
Enlargement: II dim 35 (t 14), II dlm 36 (t 12), II dvm 40 (t-p 5, 6), II epm 53
(t-cx 8), II ppm 55 (p 3)
Split: I tpm 25 into I tpm 25a and I tpm 25b (prothoracic tergo-pleural muscles);
II tpm 46 into II tpm 46a (t-p 3) and II tpm 46b (t-p 7); ppm 54 into II ppm 54a (p 2)
and II ppm 54b (p 2)
Differentiation of external insertion points: II dlril 37 (t 13), II tpm 46 (t-p 3),
II tpm 47 (t-p 4), II tpm 48 (t-p 13), II tpm 49 (t-p 15), II epm 50 (p-tr 2), II cpm 51
(p-cx 2 or 3).
Sclerotization of insertion-points: 0 dim 2 (op-t 1); I dim 10 (op-t 3), II cpm 53
(t-cx 8), II ppm 56 (t-p 16), II stm 62, 63 (spiracle muscles)
Shift of insertion points: I ism 22 (t-s 1), I ism 23 (t-p 17), II tpm 46a, b
(t-p 3, t-p 7)
THE EMBIOPTERA
In association with their movements under a web formed subterraneously or under
bark, components of each thoracic segment have undergone considerable elongation and
fragmentation. Upon superficial examination, the thorax appears to be composed of
more than three segments. This peculiarity led Verhoeff (1904) to believe that there
were six segments in the thorax of embiids and other insects. However, Crampton
( 1918a) pointed out that the thoracic structures in embiids have certain remarkable
similarities to those in Plecoptera, and that Verhoeff's theory was utterly ill-founded.
The neck region: In Eosembia and Clothoda (Fig. 49) examined in this work, the
large lateral cervical sclerite consists of anterior and posterior plates. The posterior
point of articulation is obviously on the anepisternum, as in Plecoptera. The presternal
origin of the lateral cervical sclerite is obvious, since it lies anterior to the reduced
preepisternum. In Clothoda two ventral cervical sclerites are present; the anterior is
paired (Fig. 51). In Eosembia (Fig. 50) the dorsal cervical sclerite is represented by a
small median plate, and in Clothoda by three small sclerites arranged along the anterior
margin of the pronotum.
The pronotum: In Clothoda and Eosembia (Fig. 50) the pronotum is a posteriorly
swollen, subquadrangular plate. A median longitudinal impression follows a transverse
depression on the anterior third of the pronotum. The anterolateral area of the pro-
notum on each side has become detached to form an elongate plate, as will be discussed
shortly.
The transverse plate behind the posterior margin of the definitive pronotum is the
tergum of the stenothorax of Verhoeff ( 1904). This plate is a mere secondary sclerotiza-
tion, comparable to the dorsal cervical sclerite, and should be part of the pronotum,
because it lies anterior to the mesothoracic acrotergite.
The propleuron: Using the posterior point of articulation of the lateral cervical
sclerite with the anepisternum as a fixed landmark (both in Plecoptera and Embioptera
Figs. 45A, B, 49), it is immediately noted that in Embioptera there is an extra plate
TABLE V
Thoracic musculature in Plecoptera and Embioptera
Perla Neoperla Oligotoma Muscle
(Wittig 1955) (Maki 1938) (Maki 1938) designations Remarks
I dlm 12 2 cv(d)-t 1
0 dlm 2 4 5 op-t 1
0 dlm 1 op-t 2
I dim 10, 11 1, 2 1 op-t 3
0 lm 7 7 11 op-cv 1, 2 1
0 lm 8 8 12 op-cv 3
0 lm 9 op-p 1
0 Jm 5 9, 10 t-s(cv) 1
15 t-cv 1
0 lm 6 13, 14 t-cv 2
16 t-cv? 2
0 vim 4 6 8 cv-s 4
0 vim 3 5 6 s 1
7 s 2
3(1) 4 t 12
II dim 36 28 33
III dim 36 61 64
3 t 13
II dim 37 32
III dim 37 63
I dim 10, 11, 12? 1, 2 1 t 14 3
II dim 35 27 31
III dim 35 60 62
II im 65b t 15
III im 65b
I tpm 25, 26, 27 12, 13 19 t-p? 4
II tpm 46a 35 40 t-p 3
III tpm 46a 68 70
II tpm 47
III tpm 47
t-p 4
II dvm 40 a,b,c, 33 38 t-p 5, 6
III dvm 40 a,b,c, 66 68
II tpm 46b 36 t-p 7
III tpm 46b 69
II tpm 48 38 42 t-p 13
III tpm 48 71 72
II tpm 49 37 41 t-p 15 5
III tpm 49 70 71
II ppm 56 39 43 t-p 16
III ppm 56 72 73
I ism 23 t-p 17 6
II ism 45 32 37?
III ism 45 65 67?
I ism 22 11 17 t-s 1
II ism 44 34 39
III ism 44 67 69
I ism 24 t-s 8
I dvm 15 15, 16 20, 21 t-ti(cx) 2
II dvm 41 45 48
III dvm 41 78 78
Idvm16,17 24 27 t-ti(cx) 3
46
79
TABLE V (Continued)
Perla Neoperla Oligotoma Muscle
(Wittig 1955) (Maki 1938) (Maki 1938) designations Remarks
I dvm 19 19 23 t-cx 6
II dvm 43 48 so
Ill dvm 43 81 80
I dvm 20 20 24 t-cx 7
II dvm 43 49 51
Ill dvm 43 82 81
I dvm 211 21? t-cx 8
II cpm 53 50 52
III cpm 53 83 82
II im 6Sa 44 p 1
III im 65a 77
II ppm 54a, b 40 45 p2
II I ppm 54a, b 73 75
II ppm 55 41 46 p3
III ppm 55 74 76
I zm 34 14 p-s 1
II zm 61 42, 43 47
Ill zm 61 75, 76 77
3! (II) 44 p-s 2
64(11!) 74
II cpm 51 54 57 p-cx 2 or 3
III cpm 51 87 87
I cpm 28 28 p-cx5
II cpm 52 53 56
Ill cpm 52 86 86
II vlm 39 30 36 s 12
III vlm 39 63 66
I vlm 13 29 34 s13
II vlm 38 62 65
I vlm 14 10 (I) s 14
35 (II) s 15
9(1) s 17
avlm s 19 7
III vlm 64 s 20
Ibm 32 25? s-ex 2 8
II bm 59 53?
Ill bm 59 83? 84?
Ibm 33 s-ex 3
II bm 60
Ill bm 60
Ibm 30 22 s-ex 5
II bm 57 49
III bm 57 79
26 s-ex 6
55
85
I dvm 18 25 29 t-tr 1
II dvm 42 55 58
Ill dvm 42 88 88
I 29 p-tr 1
30(!) p-tr 2
II cpm 50 56 59
III cpm SO 89 89
Ibm 31 26 s-tr 1
II bm 58 57 60
III bm 58 90 90
REMARKS (Table V)
(1) 11 in 0/igotoma is op-cv 2.
(2) 16 in Oligotoma is a very small muscle extending from the middle of the
pronotal lateral margin to the lateral portion of the posterior lateral cervical sclerite.
(3) I dim 12 in Perla may represent t 12.
( 4) These muscles are prothoracic tergopleural muscles; homologizations of them
are not attempted.
(5) All muscles in Plecoptera and Embioptera are thick vertical muscles.
(6) In Oligotoma the dorsal points of attachment of 37 and 67 are on the prealar
sclerite; in Plecoptera the homologues are on the lateral part of the antecosta (Perla)
or on the anterolateral corner of the tergum (Neoperla).
(7) avlm in Perla is lost in the adult.
( 8) Muscles in Oligotoma are difficult to identify.
(Halspleuren of Verhoeff 1904) just anterior to the anepisternum and along the lateral
margin of the definitive pronotum. This plate represents the detached anterolateral part
of the pronotum, because in Plecoptera the pronotum is not narrower in the anterior half
as in Embioptera, but often wider than long. It is probable that in Embioptera the
pronotum of a similar shape has become prolonged, and that the wide anterolateral area
has become secondarily detached to lie anterior to the anepisternum. As a consequence
of this separation, the definitive pronotum (pronotum minus its anterolateral angle) in
Embioptera is anteriorly narrowed, or posteriorly swollen (Fig. 50).
In Clothoda (Fig. 49) the propleuron posterior to the detached anterolateral pro-
notum is similar to the propleuron in Plecoptera. The small plate proximal to the
anepisternum is the preepisternum, which is absent in Eosembia. The membranous cleft
anterior to the trochantin must correspond to the area where the paracoxal suture would
occur, if sclerotized.
The prosternum: The prosternum has become greatly prolonged in Embioptera.
The small presternum (Fig. 51) is clearly separated from the anterior margin of the
basisternum as in Plecoptera. The broad basisternum is separated from the furcasternum
by the transverse sternacosta. The lateral longitudinal line on the basisternum is
probably purely secondary and is not the pleurosternal suture, since the pleurosternal
suture is absent in the prothorax of Plecoptera and the reduced preepisternum is present
separately (Fig. 49). The furca arises from each lateral end of the sternacosta. The
large plate posterior to the sternacosta is a fusion product of the furcasternum and the
spinasternum. As already observed, in Capnia (Fig. 46A) the sclerotized spinasternum
is present behind the furcasternum. Apparently in Embioptera this sclerotization is
enlarged to bear the spina at its posterior end. In Eosembia the spinal invagination
is absent.
The mesonotum: The anterior part of the mesonotum (Fig. 50) is similar to that of
The antecosta clearly separates the narrow acrotergite anteriorly. The first
phragma is only weakly developed beneath the antecosta. The prescutoscutal suture,
anterolateral scuta! suture, lateral parapsidal suture, and posterolateral scuta! suture have
the same relative positions as in Plecoptera (Figs. 46B, 47A). In Embia, as Crampton
( 1918a) showed, the latt<ral parapsidal suture is a complete transverse suture without any
median interruption. The prealar arm does not extend ventrally as in some Plecoptera.
The tegula is large and lies along the suralare. The broad area posterior to the antecosta
is the scutoscutellum. As in Plecoptera, the scutoscutellar suture is absent. The meso-
thoracic postnotum (the tergum of the cryptothorax of Verhoeff, 1904) is large and'
continuous with the metathoracic acrotergite, without forming the anterior wall of the
second phragma. The postnotum also lacks the lateral extension which, in Plecoptera,
forms the postalar bridge.
PRONOTUM PREEPISTERNUM
LATERAL CERVICAL SCLERITE
PLEURAL
SUTURE
KATEPISTERNUM
COXA
TROCHANTIN
Fro. 49. Lateral view of prothorax and cervical sclerites in Clothoda nobilis (original).
The anterolateral scuta! suture demarcates the narrow suralar plate, which bears the
anterior and antemedian notal wing processes. A small membranous emargination
follows the antemedian process. A characteristic feature of the lateral @argin of the
mesonotum in Embioptera is the fusion of the first axillary sclerite with the posterior
notal wing process (Fig. 50). The posterolateral scuta! area is demarcated by the
posterolateral scuta! suture. The tegula is present in the metathorax, as in Plecoptera.
The mesopleuron: Both in Antipaluria (Matsuda 1960a) and in Clothoda (Fig. 51)
the anapleural cleft extends posteriorly to the base of the trochantin, as in Plecoptera.
In Clothoda the incomplete paracoxal suture is apparently present on the episternal
surface anterior to the trochantin. In the anepisternal area the basalare is a small, free
plate connected to the humeral plate of the wing, as in Plecoptera. At least in Clothoda
and Eosembia a secondary ridge near the dorsal margin of the anepisternum separates a
marginal area, which is comparable with the prepectus in other orders. The pre-
episternum is the laterai swollen area of the definitive basisternum. The original line of
fusion between the basisternum and the preepisternum (pleurosternal suture) is lost in
Embioptera (shown by an imaginary broken line). In the wingless form of Antipaluria,
as Matsuda (1960a) showed, however, the preepisternal plate is separated from the basi-
sternum by the sternopleural cleft. The trochantin is a single-lobed, slender plate that
articulates with the anterolateral coxal margin. The ventropleurite is a triangular area
lateral.to .the furcal base, as in Perla (Fig. 48). The subalare is a small plate that is
associated with the third axillary sclerite.
The mesosternum: In Clothoda (Fig. 51) the original division of the anterior part
of the sternum i n t ~ the presternum and the basisternum is indicated by the incomplete
transverse line that separates them. In Antipaluria and Eosembia this suture is lost.
Unlike the prothorax and like the mesosternum of Plecoptera, the furcal pits are not
connected by the sternacosta. A transverse plate bearing the spina at its posterior end
may be regarded as the spinasterhum; it is separated from the fucasternum by a
transverse suture.
The metanotum: In Eosernbia (Fig. 50) the second phragma is produced by
invagination from the metathoracic antecosta, and it is continuous with the mesothoracic
postnotum. There is no other notew6rthy difference between the mesonotum and the
metanotum.
PRONOTUM,
ANTE COSTA
PREALAR ARM
LATERAL
PARAPSIDAL
SUTURE
SURALARE
SCUTUM
ANTEROLATERAL
SCUTAL SUTURE -
l'ST AXILLARY
SCLERITE
POSTEROLATERAL
SCUTAL SUTURE
DORSAL CERVICAL
SCLERITE
T ~
I
PRO NO TUM
ACROTERGITE
PRESCUTOSCUTAL
SUTURE
TEGULA
ANTEROLATERAL
SCUTAL SUTURE
TERGAL
FISSURE
-----POSTEROLATERAL
SCUTAL SUTURE
SCUTUM
POSTERIOR NOTAL
PROCESS
POSTNOTUM
Fro. 50. Dorsal view of thorax in Eosembia sp. (original).
The metapleuron and the metasternum: In the embiids examined in this study, the
metapleuron has no noteworthy difference from the mesopleuron. The furcal pits in
the metasternum are connected by a transverse ridge.
The adult musculature: The only reliable study of the thoracic musculature in
Embioptera is on 0/igotoma jormosana by Maki (1938), which is shown in Table V.
As the great similarities in external structures between this order and Plecoptera
suggest, there is also a striking similarity in the thoracic musculature between the two
orders, as seen from Table V. In Embioptera, however, more sternal muscles have been
retained than in Plecoptera. This condition is clearly associated with locomotion in
their peculiar habitats.
BASAL ARE
ANAPLEURAL CLEFT
PRE EPISTERNUM
KATEPISTERNUM
FURCAL
FURCASTERNUM
PREEPISTERNUM ?
N
ECTUS
PLEURAL WING
PROCESS
AN EPISTERNUM
BASISTERNUM
TROCHANTIN
SPINASTERNUM
PRESTERNUM
FIG. 51. Ventral view of thorax in Clothoda nobilis (original).
139
THE PHASMIDA
The neck region: In Anisomorpha (Fig. 52A, B) and Pseudophasma the lateral
cervical sclerite consists of a plate in two parts. The anterior part articulates with the
postoccipital ridge, and the posterior part with the preepisternal area. In Dixippus
(Jeziorski 1918) the lateral cervical sclerite is not divided into two. Neither the dorsal
nor the ventral cervical sclerite is known to occur in Phasmida. The fact that the lateral
cervical sclerite articulates with the preepisternum suggests that the lateral cervical sclerite
is of presternal origin.
The pronoturn: The pronotum is a simple plate with a median longitudinal line,
which is crossed by a median or antemedian transverse internal ridge. Another trans-
verse ridge behind the anterior margin of the pronotum becomes continuous with the
similarly ridged lateral margin of the pronotum. Often the anterolateral corners of the
pronotum are notched to expose the external openings of the thoracic repellent glands
(Fig. 52A), which Gangrade ( 1964) studied histologically. The first thoracic spiracle
lies on the membrane below the posterolateral angle of the pronotum.
The propleuron: At its ventral end, the pleural ridge forms a well-developed
process, which articulates with the coxa (Fig. 52B); no distinct pleural arm is formed on
the pleural ridge. The episternal area is generalized, and similar to that in Plecoptera.
In A nisornorpha the paracoxal suture is probably represented either by the internally
ridged posterior margin of the episternum or by a suture dorsal to it (Fig. 52B).
DuPorte (1965) also noted the probable paracoxal suture in Diapheromera. In Aniso-
rnorpha a broken vertical line from the anterior dorsal part of the episternum may
represent the anapleural suture. The preepisternal arm extends ventrally to meet the
anterolateral angle of the sternum; the narrow trochantin is continuous from or
articulated with the internally ridged posterior margin of the sclerotized episternum, and
articulates peculiarly with the inner margin of the coxa. The narrow epimeron is present
in Anisomorpha.
The prosternum: In Phasmida the prosternum consists of two plates, the anterior
and the posterior, as Jeziorski (1918) and Chopard (1949) maintained. While the
anterior plate must be largely basisternal in origin, homology of the posterior plate is
quite unclear. Since t-s 7 (or 8) is inserted on the middle of the lateral margin of this
plate, and since its posterior margin lies caudal to the anterior margin of the mesonotum
in Megacrania (Maki 1935) and Dixippus (Jeziorski 1918), it is quite possible that this
plate is formed by fusion of the posterior part of the prosternum and the anterior part of
the mesosternum. It is also possible that this posterior plate corresponds to the posterior
prosternal plate in Embioptera (Fig. 51), which does not include the mesosternum.
The furca and the spina are absent in the prothorax of Phasmida.
The rnetanotum: In winged Pseudophasma (Fig. 53A) the first phragma on each
side arises from the antecosta. The prescutum and the lateral prealar arm are behind
the antecosta. Hence, the prealar arm is probably prescutal in origin, as in Plecoptera.
The oblique prescutoscutal suture is recognized only laterally, as in Plecoptera and
Embioptera. The anterolateral scuta! suture, the parapsidal suture, and the tergal fissure
lie in their typical positions, although the association of the anterior end of the parapsidal
suture with the prescutoscutal suture has become obliterated. On the posterior half of
the metanotum, the posterolateral scuta! suture is distinct, and the recurrent scuto-
scutellar suture is obliterated anteriorly.
On the scutallateral margin (Fig. 53 A) the anterior, the antemedian, and the median
notal wing processes are clearly recognized, and the posterior notal wing process is
detached. Characteristically, the proximal margin of the first axillary sclerite is firmly
attached to the scuta! lateral margin in Pseudophasma, Necroscia (Gangrade 1966) and
Orxinus (Ragge 1955), as in Plecoptera.
The metathoracic postnotum is fused with the first abdominal tergite; such a fusion
occurs during the embryonic stage (Heymons 1897 b). An inconspicuous phragma
LATERAL CERVICAL
SCLERITE----------""""""'
BASI STERNUM
..
. . .
+ FURCASTERNUM -----+--
A
REPELLENT GLAND
B
Fm. 52. Prothorax of Anisomorpha sp. (original).
~ C O X A
TROCHANTIN
AN EPISTERNUM
A, ventral view; B, lateral view.
between the ,postnotum and the first abdominal tergite, which Maki described in
Megacfania, is a mere thickening in Pseudophasma examined (Fig. 53 A). In wingless
phasmids the metanotum is a simple plate without subdivision by sutures.
The metap/euron: In winged Pseudophasma (Fig. 53B) the horizontal pleural ridge
can be traced as far as the anterior limit of the metapleuron. The metepimeron is com-
pletely lost, and a narrow subalare is recognized along the pleural ridge. The pleural
arm lies at the level posterior (ventral) to the middle of the pleural ridge. The dorsal
end of the episternum has a transverse membranous cleft, which separates off two dorsal
basalaria; the large anterior basalare is comparable with that in Mantodea, and the
smaller posterior basalare is comparable with the basalare in Blattaria. The condyle at
the base of the Sc vein fits in a notch formed between the pleural wing process and the
dorsal margin of the posterior basalare (x in Fig. 53B). The humeral plate articulates
with the anterior dorsal angle of the posterior basalare.
The long membranous cleft that separates the episternum and the definitive meta-
sternum in Pseudophasma is the anapleural cleft, because it arises anteriorly from the
area ventral to the spiracle and the anterior basalare, and reaches the base of the
trochantin posteriorly. The preepisternum ventral to the anapleural cleft must have
been indistinguishably fused (shown by a broken imaginary line) with the sternum.
The paracoxal suture is absent. An imaginary paracoxal suture that has become lost is
shown by a broken line reaching the level of the pleural arm (Fig. 53B). The point of
articulation of the trochantin with the coxa, as in the prothorax, is more ventral
(proximal) than in most other orders.
In the metapleuron of wingless phasmids the basalar sclerites are not differentiated.
Littig (1942) showed in Anisomorpha that for its entire length the episternum is divided
horizontally into two areas. The ventral one was found, by actual examination, to be
merely a desclerotized area. No distinct pleurosternal suture has been found in wingless
phasmids. In Dixippus Jeziorski (1918) showed an additional trochantinal piece lying
between the pleurocoxal articulation and the anterior trochantinocoxal articulation.
The metasternum (Fig. 53B) : The furcal pits with poorly developed internal
furcae are rather widely separated from each other. The sulitriangular space from the
posterior margin of the mesosternum reaches (anteriorly) the level of the furcal pits or
beyond. This posterior area is the definitive furcasternum that has incorporated the
first abdominal sternum by fusion during the embryonic stage (Heymons 1897 b).
The mesonotum: In Pseudophasma, in which the forewing is brachypterous and is
located on the posterolateral angle of the mesonotum, a typical division of the mesonotum
by sutures is absent. The mesonotum in winged Phasmida is therefore a simple plate
and is separated from the mesopleuron by the membranous infolding.
The mesopleuron and mesosternum: The mesopleuron in Pseudophasma is essen-
tially the same as in the metapleuron of wingless forms. The separation of dorsal parts
of the episternum into the basalaria is absent. The definitive mesosternum is laterally
preepisternal in origin. The vestigial furcal pits are continuous with the vestigial furcae
internally. The small, definitive posterior plate of the mesosternum (presternum of
Maki 1938) may be a composite plate, similar to the corresponding definitive posterior
plate of the presternum. The mesosterna! spina is absent.
Phylogenetic relationships: The thorax in Phasmida is similar to that in Plecoptera
in having the prealar arin of prescutal origin, the first axillary sclerite that is closely
attached to the scuta! lateral margin, the short and incomplete prescutal suture and the
recurrent scutoscutellar suture and in lacking the scutoscutellar suture. The thoracic
musculature is also similar in the two orders. Of these characters, at least the shape and
position of the first axillary sclerite can be regarded as synapomorphic and support
Crampton's ( 1916b) contention that the origin of the Phasmida may be traced back to
the perloid insects.
142
PHRAGMA ---------..1_
ANTECOSTA
TEGULA
PARAPSIDAL
SUTURE
l'ST AXILLARY
SCLERITE
TERGAL FISSURE
RECURRENT
SCUTOSCUTELLAR
SUTURE
1 ABO. TERG.
BASALARE------------
PLEURAL WING
PROCESS------
A
AN EPISTERNUM -------'-o-. .!\\. ;..._ __
,'
SUB ALAR E -----------:il\1
:, \
PLEURAL SUTURE ---------m\
FURCASTERNUM
' \
' '
PRE SCUTUM
PRESCUTOSCUTAL
SUTURE
ANTEMEO.IAN NOTAL
MEDIAN NOTAL
PROCESS
DETACHED
POSTERIOR NOTAL
PROCESS
POSTEROLATERAL
SCUIAL SUTURE
POSTNOTUM
------- ANTERIOR BASALARE
- <ANEPISTERNUM)
SPIRACLE
, ANAPLEURAL
: ~ I CLEFT
L__t__ IMAGINARY PLEUROSTERNAL
SUTURE
-;--+------_FUSED
PREEPISTERNUM
IMAGINARY
PARACOXAL
SUTURE
FURCAL PIT
FIG. 53. Metathorax of Pseudophasma menius (original). A, dorsal view; B, lateral view.
143
IUs also interesting to point out that in Embioptera, which are specialized Plecop-
tera"' as far as the thoracic structures studied are concerned, prolongation of the body and
fragmentation of thoracic sclerites have occurred, and this character may be regarded as
synapomorphic in the two orders. Assuming, then, that Phasmida and Embioptera are
apomorphic sister groups of Plecoptera, the two orders have become differentiated
through autapomorphic changes peculiar to each. For instance, in Phasmida the antero-
lateral angle of the pronotum, which is an isolated plate in Embioptera, has been modified
to be the opening of the repellent gland in Phasmida. The mode of fragmentation of
dorsal and ventral thoracic sclerites is also obviously different in the two orders.
The adult musculature: Jeziorski (1918) and Maki (1935) studied the thoracic
musculature of wingless Dixippus morosus and winged Megacrania tsudai, respectively.
Homologies of the muscles in these forms are shown in Table VI. The definitive
posterior plates of the prosternum and mesosternum in Megacrania were identified as the
presternum by Maki (1935). Therefore, his descriptions of the muscles attached to
these areas could not be followed literally. The actual positions of the muscles in Maki's
figures, rather than his descriptions, were followed where discrepancies existed.
In wingless Dixippus and winged Megacrania, t 14 (dorsal longitudinal muscle) is
absent in the pterothorax, whereas in wingless Gryl/oblatta, t 14 is well retained in all
thoracic segments. Other important indirect flight muscles, t-p 5 and 6, are also absent
in Phasmida. A remarkable fact is the presence of p 3, which is present in Plecoptera
and in some other rather strong fliers and is absent in Blattaria, Isoptera, and Mantodea.
A comparison with Table V shows an overall similarity in the musculature between this
order and Plecoptera and Embioptera. The presence of p-cx 6 and t-s 8 represents a
primitive feature in the thoracic musculature of this order.
In a comparison of winged Megacrania with wingless Dixippus, the most conspicuous
differences are the absence of t-p muscles and the scarcity of pleural muscles in Dixippus;
they are the muscles directly or indirectly concerned with flight.
TIIE BLAITARIA
The neck region: The dorsal cervical sclerite in Blattella (Matsuda 1956a) and
Periplaneta (Crampton 1927; Snodgrass 1952) is a horseshoe-shaped plate, which
articulates anteriorly with the middle of the dorsal margin of the occipital ridge. The
lateral cervical sclerite (Fig. 54A) consists of the anterior part articulated with the post-
occiput, and the posterior part, which is folded and connected with the preepisternal area.
The articulation of the latter with the preepisternal area precludes the possibility that the
lateral cervical sclerite is of preepisternal origin; and hence the sclerite is of presternal
ongm. Two transverse ventral cervical sclerites are present on the neck membrane
between the lateral cervical sclerites.
The prothorax: The pronotum has a well-developed marginal paranotal area, which
does not overgrow the pleuron.
The narrow, oblique, anterior episternal plate (Fig. 54A) was called the precoxale
by Crampton ( 1927) and the preepisternum by Matsuda ( 1956a). This plate is actually
the anepisternum plus the preepisternum, judging from its elongate shape and position.
The crooked pleural suture bears the well-developed pleural arm internally, and the
narrow epimeron is separated off by this pleural suture. The episternum posterior to
the anepisterno-preepisternal plate (Fig. 54A) is difficult to interpret, owing to reduction
and to the absence of such landmarks as the anapleural cleft (suture) and the paracoxal
suture. The episternal area along the pleural suture, however, must be partly the
anepisternum. The two-part trochantin is present.
In Blatel/a (Fig. 54A) the sclerotized sternal plate is largely the basisternum, if the
bases of furcae are taken as landmarks separating the basisternum from the furcasternum.
In Periplaneta, as Crampton (1927) showed, a short, transverse furcasternal plate is
4.!lSynapomorphic characters of Plecoptera and Embioptera are the reduction of the paramere, the modification
of the tenth tergum to form a copulatory organ, and the reduction of the ovipositor.
144
TABLE VI
Thoracic musculature in Phasmida and Grylloblattodea
Megacrania Dixippus Grylloblatta Muscle
(Maki 1935) (Jeziorski 1918) (Walker 1938) designations Remarks
65 0 idim 2 56? cv(d)-t 1
1
71 i Fam c, d 51 op-t 1 2
70 0 dim 1 52 op-t 2 3
45 op-t 3
0 idvm 1, 2 op-p 1
76, 77 0 dvm 2b, c 46 op--cv 2, 3
0 ism 2, 0 dvm 2a t-s(cv) 1 4
62 0 ism 48 t-cv 1
47 t-cv 2
73 0 vlm 3 cv-s 1 5
49 cv-cx 2(x)
72 0 vim 4, 5 50 s 1, 2
67, 69 I dim 3b t 12
94,96 II dlm 4
134, 136 Ia dlm 4, 5
66, 68 I dim 3a 56? t 13
93, 95 II dim 1, 2, 3 83
133, 135 Ia dlm 1, 2, 3 113
65 0 idim 2 45, 56? t 14
82
112
133-136 (II) t?
143 (III) t-p 7
144 (III) t-p 8
76 (II)
107 (Ill)
t-p 9 or 4
105 (II) 77? t-p 12 6
146 (III) 108?
114 (II) t-p 13
153 (III)
115 (II) t-p 15
154 (III)
106 (II, pars) t-p 16 7
147 (III, pars)
145 (Ill) t-p 18
74, 75 (I) t-p? 8
104, 106 (II)
14 7, 237 (III)
53, 54 (I) t-p? 9
101 (HI) I ism 58 t-s 1 10
100 (II-III) II ism 85
141 (III) ? III ism ? 115
102 (Hf) I ism 2 t-s 7 or 8
103, 142 (II-III)
II ism 2
81 I dvm 6 60 t-ti 2 or 3
116 II dvm 6 86
155 116
62 t--cx 5
87
117
TABLE VI (Continued)
(Maki 1935) (] eziorski 1918) Walker 1938) designations Remarks
88 (I) I ldvm 1 63 t-ti 2 or 3? 11
83 I dvm 3, 4 t-cx 6
117 II dvm 3, 4
!56 III dvm 3, 4
84 I dvm 2 t-cx 7
127 II dvm 2
!57 III dvm 2
85 I !dvm 2a t-cx 8 12
127 II ldvm 2a
164 III ldvm 2a
107, 109 (II) Muske! be lag p 1
148 (II) p 2
149 (II) p 2?
110(Il) 78 p3
150 (III) 109
I Zm 55 p-s 1
111(II) II Zm 79
!52 (III) III Zm 110
108 (I-II) I ildvm 75 p-s 2 13
II i!dvm 106 (I I-Ill)
139 (II-III) II vlm p-s 9
111b (II-2A) p-s 13 14
I pm 5 p-ti(cx) 1
122 (II)
Ill pm 5
89 (?) p-ti(cx) 2
123
160
82 (?) 61 (I) ? p-ti(cx) 3
125, 126 Illdvm 1
162, 163 III !dvm I
91 (II)
121 (III)
p-cx 4
124 (II) p-cx 5
!61 (III)
91 I ldvm 2 p-cx 6
129 II ldvm 2
!66 III ldvm 2
80 (I-II) s 12
138 (IT-III) 111a
57 (I-ll) s 13
84 (II-Ill)
98 (I) I vlm 59 s 14
81 (I-II) s 15
99 (II) s 16
140 (III)
114 (Ill-2A) s 20
88 (I-II) s-ex 1
118 (II-III)
92 (II) s-ex 2
122 (III)
87 Ibm 2
s-ex 3
121 II bm 2
!59 III bm 2
TABLE VI (Continued)
(Maki 1935) (Jeziorski 1918) (Walker 1938) designations Remarks
90 (I I) s-ex 4
120 (III)
86 Ibm 1 64 s--ex 5
119, 120 II bm 1 89
158 III bm 1 119
90 Ibm 3 65 s-ex 6 15
128 ? II bm 3 93
165 ? III bm 3 123
I dvm 5 67a t-tr 1
130 II dvm 5 98a
167 III dvm 5 128a
67b (I) p-tr 1
92 I ldvm 3 p-tr 2
131 II ldvm 3 98b
III ldvm 3 128b
168 III bm 4 s-tr 1
REMARKS (Table VI)
(1) 56 in Gryl/oblatta may represent t 13.
(2) 71 and i Fam c, din Phasmida are inserted on the hind margin of the head
more ventrad than in other insects; 51 in Gryl/oblatta is inserted on the neck membrane.
( 3 ) 70 and 0 dlm 1 in Phasmida are inserted on the dorsal neck membrane
behind the posterior end of the head.
( 4) 0 ism 2 in Dixippus is not inserted on the tentorium, but on the anterior part
of the lateral cervical sclerite. Some other cervical muscles in Phasmida also have
shifted their anterior points of attachment.
( 5) 73 in M egacrania is inserted posteriorly on the posterior end of the anterior
prosternal plate.
(6) Muscles in Gryl/oblatta are difficult to homologize, in the absence of such
landmarks as the anterior and posterior notal wing processes.
(7) The posterior parts of 106 and 147 in Megacrania are attached to the
subalare; hence they are t-p 16.
( 8) Each one of these muscles in M egacrania consists of a row of short and
vertical fibers that connect the lateral area of the tergum and the dorsal margin of the
pleuron. These muscles are dvm Belag of Jeziorski (1918).
(9) 53 and 54 in Grylloblatta are prothoracic tergopleural muscles.
(10) Metathoracic muscles in Megacrania and Dixippus arise from the anterior
lateral margin of the first abdominal tergite, and are inserted on the metasternal furca.
( 11) The dorsal origins of these muscles lie anterior to that of the preceding
muscle ( t-ti 2 or 3), and they are abductors of the coxa.
(12) t-cx 8 in wingless Dixippus, in the absence of the subalare, arises from the
posterolateral margin of the tergum.
(13) In Megacrania 108 may be serially homologous with 148 (p 2).
(14) 111 b in Grylloblatta arises from the mesothoracic spina and is inserted on the
anterolateral angle of the second abdominal segment. Walker ( 1938) expressed some
doubt about the exact pojnt of insertion of this muscle.
(15) The points of attachment of 128 and 165 on the coxal margin in Megacrania
are the same as those of 121 and 159, which are s-ex 3.
Besides the muscles homologized in Table VI, Maki (1935) described spiracle
muscles (78-80, 112, 113), heart muscles (97, 137), thoracic gland muscles (63, 64),
and an ovary muscle. ( 132).
LATERAL CERVICAL
. SCLERITE
KATEPISTERNUM +
AN EPISTERNUM
PREALAR

SCUTOSCUTELLAR
SUTURE--t----_j

----
c
VENTRAL
CERVICAL SCLERITE
EPIMERON
RCA
TROCHANTIN
l'ST PHRAGMA
:/
POSTERIOR
NOTAL
PROCESS
147
FIG. 54. A, ventral view of prothorax and lateral cervical sclerite in Blattella germanica
(from Matsuda); B, connection of mesonotum and metanotum in Arenivaga aegyptica
(original); C, dorsal view of pterothorax in Arenivaga aegyptica (original).
present between the bases of the furcae. The prothoracic spina arises from the small
sci erotized median plate lying in the membranous area behind the furcae (Fig. 57A, B).
The meso- and meta-notum: The meso- and meta-notum are nearly equal in size
and similar in shape in most cockroaches. In the mesonotum of Arenivaga (Fig. 54C)
the weakly developed phragma extends internally from shallow, paired depressions
beneath the antecostal suture. The second phragma in Arenivaga (Fig. 54B) consists
of the anterior wall arising from the greatly reduced mesothoracic postnotum (hidden
beneath the scutellar evagination) and the narrow posterior wall arising from the meta-
thoracic antecosta. The third phragma is also a narrow internal inflexion which is
presumably of postnatal origin.
The mesothoracic postnotum forming the second phragma has not been correctly
recognized by previous authors. According to Crampton (1927), the postnoturri is a
small plate lying on each side of the secondary reduplication of the &cutellum in Peri-
planeta. Matsuda ( 1956a) labeled the posterior evagination (reduplication) of the
scutellum, the postnotum. Snodgrass (1952) did not even recognize the presence of the
postnotum in In Blatella, as the figures given by Snodgrass (1909) and
Matsuda ( 1956a) show, the postnatal phragma is horseshoe-shaped; and in Periplaneta
Snodgrass ( 1952) called it the connective sclerite. It is now clear that this connective
sclerite is nothing but the reduced anterior wall of the second phragma.
In some forms the anterior area of the tergum behind the antecosta is demarcated
by a transverse suture, which probably represents the prescutoscutal suture. This suture
extends almost to the anterolateral margin of the tergum in Periplaneta (Crampton
1927; Snodgrass 1952), or it exists as only a short median transverse suture in Arenivaga
(Fig. 54C), or it is completely absent in Blat ella (Matsuda 1965a). Crampton (1927)
and Matsuda ( 1956a) recognized a small triangular plate lying anterior to the anterior
notal wing process. This plate is either detached from the anterolateral angle of the
tergum or continuous with it (Fig. 56); Crampton regarded this plate as the incompletely
developed prealar arm. This presumed prealar arm has, however, lost direct connection
with the prescutoscutal suture, and the structure is clearly an extension of the acrotergite,
as in Orthoptera.
The scutum and the scutellum are separated by the inverted V-shaped, internally
ridged scutoscuteJ!ar suture in many cockroaches. The anterolateral scuta! suture that
separates the anterior notal wing process is also present.
The mesoscutal lateral margin and forewing base structures: Onesto (1959a) des-
cribed the axillary sclerites and the other structures associated with wing opening and
closing mechanisms in Blatta orienta/is. This species exhibits the sexual pterygo-
dimorphism, the female being micropterous. The following description therefore relates
only to the male of Blatta orienta/is.
Besides the anterior, antemedian, and posterior notal wing processes, the small
median process is present on the scuta! lateral margin (Fig. 55 A). The postmedian
process is absent. The median processes are also clearly seen on the metanotal lateral
margin (Fig. 56).
The neck region of the first axillary sclerite (Fig. 55A) articulates with the anterior
notal process and the base of the Sc vein, as in other insects. It articulates laterally with
the second axillary sclerite, and it is proximally connected with the scuta! lateral margin
at the anterior, antemedian, and median notal processes.
The dorsal sclerotization of the second axillary sclerite (Fig. 55A), besides its
articulation with the first axillary sclerite, articulates with the proximal median plate and
with the basiradiale, .. it is also connected posteriorly with the third axillary sclerite. The
ventral sclerotization of the second axillary sclerite (Fig. 55B) articulates with the
pleural wing process, and has close association with the ventral sclerotization of the third
axillary sclerite and with the subalare.
Onesto (1959a). did not describe it, the plate anterior to the proximal median plate probably repre-
sents the base of the R vein that has become fused with the distal median plate. This presumed basiradiale
forms a small process that fits in the anterior end of the second axillary sclerite, as in other insects.
149
TEGULA
BASI COST ALE
BASIRADI
ANTERIOR
NOTAL PROCESS
ANTEMEDIAN
NOTAL PROCESS---+---
TERGAL
MEDIAN NOTAL
PROCESS
l'ST AXILLARY
SCLERITE
POSTERIOR
NOTAL PROCESS
A
8
BASISUBCOSTALE
DISTAL MEDIAN
PLATE
'-----+--- PROXIMAL
MEDIAN PLATE
BASANALE
3'RD AXILLARY
SCLERITE
BASIJUGALE
FIG. 55. Forewing base in Blatta orienta/is (from Onesto). A, dorsal view; B, lateral view.
3'RO AXILLARY
SCLERITE
POSTERIOR
NOTAL PROCESS
)<
CORD BASIJUGALE
SASICOSTALE
BASISUBCOSTALE
2'NO AXILLARY
SCLERITE
FIG. 56. Dorsal view of hind wing base in Blatta orienta/is (from Onesto).
The dorsal sclerotization of the third axillary sclerite (Fig. 55A) articulates with the
posterior notal wing process, and its proximal anterior process articulates with the dorsal
posterior end of the second axillary sclerite, and has the tendinous connection with the
first axillary sclerite; its distal anterior expansion continues to the posteromedial angle
of the proximal median plate. Posterodistally, it is narrowly continuous with the
basanale. The ventral third axillary sclerite (Fig. 55B) articulates with the ventral
second axillary sclerite and with the subalare.
Both the proximal and distal median plates are well defined. The distal median
plate, which contains the basiradiale proximally, is also united with the basal part
of the Sc vein (basisubcostale) and with the R+M vein complex via a narro.v,sclerotized
bridge. A faint line on the distal median plate allows movement of the forewing.
Onesto thought that the distal median plate was derived from the base of the M vein.
The basicostale (Fig. 55A) is the humeral plate of Snodgrass (1935), which repre-
sents the base of the costa; it is associated medially with the tegula, and posteriorly with
the basisubcostale.
The basisubcostale (Fig. 55A) has dorsal and ventral sclerotization. In addition
to its association with the tegula and the basicostale, the dorsal sclerotization
connects proximally with the neck of the first axillary sclerite, posteriorly with the basi-
radiale, and distally with the proximal end of the Sc vein. The ventral sclerotization
(Fig. 55B) connects anteriorly with the basicostale, proximally with the basalare by a
robust tendinous ligament, and laterally with the bases of the Sc and R veins. Onesto
thought that the R vein participates in the formation of the ventral basisubcostale.
The ventral surface of the R vein base (Fig. 55B) is swollen and its posterior margin
lies above the surface of the wing and projects caudally.
The basanale (Fig. 55A) lies between the third axillary sclerite and the A veins. It
connects with the A vein laterally and with the distal median plate and the third axillary
sclerite by a narrow bridge.
The basijugale is a vast area of sclerotization located on the dorsal side of the
forewing, occupying the area between the posterior notal wing process ,and the anal field
(Fig. 55 A). Anteriorly, it is associated with the third axillary sclerite and the basanale.
Characteristically, as noted in Fig. 55A, the bases of the Sc vein and the A vein are
isolated by membrane in the forewing of Blatta. In the hind wing (Fig. 56) the separa-
tion of the bases of the C and Sc veins is absent.
151
The meso- and meta-pleuron: In the nymph of Blatta (Fig. 13B) the paracoxal
suture extends into the epimeron. However, in most adult cockroaches this suture is
obliterated posteriorly, and cannot be recognized on the epimeron.
The anapleural suture, which is often indistinct or completely obliterated, is clearly
present in Arenivaga (Fig. 57A) and Ischnoptera (Crampton and Rasey 1915). The
small basalare is present in the membranous basalar cleft (incisura basalare of La Greca
1949a). According to Onesto ( 1959a), the basalare lies isolated in the winged male,
but is fused with the neighboring sclerotized area in the wingless female.
The proximal part of the preepisternum lies along the lateral margin of the basi-
sternum, and is always relatively short in cockroaches. In many forms it is posteriorly
demarcated by the paracoxal suture and the antecoxale (Fig. 57B) covers the anterior
part of the large trochantin, when the insect is at rest. In the mesothorax of Arenivaga
(Fig. 57A) the paracoxal suture is absent and hence the antecoxale is not formed; the
posterior area of the relatively large definitive preepisternum covers the anterior part of
the trochantin. In many cockroaches the antecoxale has a definite functional advantage;
the membrane folded beneath it gives flexibility in the movements of the pleuron and the
leg base.
As in some Thysanura and in some other hemimetabolous insects, the large
trochantin is divided into two parts by an oblique suture. Its distal end meets the pleural
suture near the pleurocoxal articulation. The narrow subalare lies along the somewhat
reduced epimeron.
The meso- and meta-sternum: The sternal area of the pterothorax (Fig. 57 A, B) is
often considerably membranized. The presternum is absent. The large sclerotized
plate is the basisternum; the paired longitudinal sutures (sternellum, see p. 20) often
develops on the middle of the basisternum. The furcal pits lie close to the median longi-
tudinal axis. The sclerotized furcasternum is greatly reduced and no sternacosta is
present; the second spina is present.
The anatomical basis of flight: In cockroaches only one direct flight muscle (t-p 14)
is inserted on the third axillary sclerite. Contraction of this muscle as flexor is directly
responsible for opening and folding of the wing; a highly complex connection of the third
axillary sclerite with neighboring sclerites accounts for this movement. Muscles t-p 6
and p 3 are absent and t 14 is weakly developed. Cockroaches are therefore weak fliers.
The contraction of relatively large numbers of the tergocoxal (trochantinal) muscles and
the tergotrochanteral muscle is probably responsible for the elevation of wings; the leg
muscles inserted dorsally on the subalare (t-cx 8), the basalare and the anepisternum
(p-ti 2, 3) probably directly lower wings, as Chadwick ( 1953) thought regarding the
flight mechanism in insects lacking a well developed t 14. Ewer and Nayler (1967)
thought that p-ti 2, 3 depress the anterior margin of the wing and cause the wing
to extend.
In a comparison of the pterothoracic musculature between the winged male and the
wingless female of Deropeltis Ewer and Nayler ( 1967) only t-p 14, t-p 3, and
t-ti 1 (or t-p 5) are absent in the wingless female, and they 'discussed the bifunctional
aspects of leg muscles by studying their relative development iri\ both sexes. Ewer and
Nayler (1967) further proposed a hypothetical relation the sequence of
muscular contraction in walking and their implication in possible\wing movement. In
various cockroaches Kramer (1958) found that flight ability is correlated with the
presence of cytochrome ?ontent, which gives a pink color to the muscles.
The adult musculature: Thoracic muscles were studied by Carbonell (1947) in
Periplaneta americana, by Chadwick (1957) in various cockroaches (ventral muscles),
by Ewer and Nayler (1967) in Deropeltis erythrocephala (pterothoracic muscles), and
by Storch (1968) in Nauphoeta cinerea. In Table VII (Chadw.) refers to the muscles
found by Chadwick.
Besides some obviously primitive muscles that are retained, the thoracic musculature
in Blattaria is characteristic in the absence of t-p 6 and p 3, which are strongly developed
SPIRACLE TEGULA
ANEPISTERNUM PLEURAL
SUTURE
SPINA
BASISTERNUM
PREEPISTERNUM
+ ANTECOXALE
TROCHANTIN
FURCA
SPINA
PARACOXAL
SUTURE
BASI STERNUM
--EPIMERON
PREEPISTERNUM
TROCHANTIN
-+ ANTECOXALE
FURCA
A
ANEPISTERNUM
BASALARE
SPINA
EPIMERON
BASI STERN
ANTECOXALE
PLEURAL
TROCHANTIN
ANEPISTERNUM
ANTECOXALf
F

BASISTERUM
B
Fro. 57. A, ventral view of pterothorax in Arenivaga aegyptica (original); B, ventral view
of pterothorax in Blattella germanica (from Matsuda).
in other orders; their absence is also correlated with considerable reduction of the
preepisternurri in this order. In the possession of comparatively large numbers of
muscles arising from and within the pleuron Blattaria is more specialized than
Ephemeroptera.
Gross aspects of muscle development: Storch and Chadwick (1967) traced the
development of cervicothoracic muscles in Periplaneta americana throughout embryonic,
nymphal and adult stages. Nearly all nymphal muscles are formed at late stages of the
embryonic development. A comparison of the nymphal (first stage) and adult muscles
is given below. Note the fact that all or nearly all nymphal muscles can be homologized
with adult muscles in Periplaneta americana, or with the adult muscles in other insects.
Muscles that become lost in the adult: 1 Ph-POR (op-t 3), 1-3 Ph 1-3 LMemb
(t-p 17), Pl1-Cx No.3, 4 (p-cx 6), 2 Sp-S II (p-s 13?), Eps 2, 3-Tn2, 3 No.1 (p-ti 1),
Fu 2-(3 Sp) (s 13?) L Cer-Cxlx (cv-cx 1(x)).
Decrease in number of muscles due to loss or fusion:
1st stage nymphs Adult
1 Ph-DMemb [cv(d)-t 1]
T 1-POR (op-t 2)
T 1-Tn 1 (prothoracic t-ti muscles)
T 1-PIR 1 (prothoracic t-p)
T 2-2 Ph (t 12, 13)
2 Ph-Eps 3 (t-p 3)
P!R 2, 3-Cx 2, 3 (p-cx 4, 5)
Increase in number of muscles probably due to split:
2
2
5
5
4
2
3
1
1
4
4
3
1
2
1st stage nymphs Adult
Tl-Cxl No. 3-7 (t-cx 6, 7, 8)
PIR 1-Tr 1 (p-tr 1 or 3)
Eps 2, 3-Tn 2, 3 (p-ti 2, 3)
6
2
3
Sclerotization or modification of external parts where muscles are attached:
Tl-1Ph (t 12,13), T2,3-Cx 2,3 No.4 (t-cx 8), Eps 2,3-Tr 2,3 (p-tr 2) T 2,3-PlR 2,3
(t-p 14).
A muscle or a ligament with some muscle fibers in the nymph, and wholly liga-
mentous in the adult:
Fu-PlR (p-s 1) in the prothorax, Fu 1,2,3-LMemb 1,2,3 (p-s 5), fu3-fu3 (s 16).
In Nauphoeta (Storch 1968) the following ventral muscles are present only during
the nymphal stage: L Cer-Cx IX (cv-cx 1 (x)), SA 1-Eps 2X(p-s 4(x)), Cx 1-Eps
2X(p-cx 7(x) ), SA 1-2 Sp (s 15), (3 SP)-Ventr. diaph.
The histological differentiation during development: Tiegs ( 1955) traced the histo-
logical differentiation of the tergosternal muscle (t-p 5?, 121 of Carbonnel) and a
tergocoxal muscle (t-ti 1) during the postembryonic development of Blatt ella germanica,
as follows.
The tergocoxal muscle (t-ti 1) consists of about 60 fibers at the first nymphal stage.
In the large fibers the pattern of marginal lamellate sarcostyles (compound fibrils) is
already evident. The initial growth of the muscle is due almost entirely to the enlarge-
ment of fibers. Within the fibers the lamellae increase in number and in width, and th.e
nuclei begin to move from the axial sarcoplasm into the outer zone of lamellae. Later a
cleavage of fiber upsets the sarcostyle pattern of fibers. The fiber cleavage proceeds
more actively in Periplaneta americana.
The tergosternal muscle (t-p 5?) in Blattella is not recognized in dissections until
late in the nymphal period, the rudiment of this muscle being a mere delicate wisp in
early nymphs. As the nymph grows, the muscle rudiment, plainly a rudimentary inuscle
fiber, enlarges, its fibrils increasing in number and displaying more distinct cross striation.
Within the sarcolemma the fibrils now become grouped into separate columns, which in
the later nymph are recognizable as daughter fibers with a marginal zone of lamellae, and
enclosing an axial core of sarcoplasm. The entire muscle, comprised of some 12 fibers,
is therefore derived from a single rudimentary fiber.
According to Tiegs (1955), most of the muscles, whether or not they are flight
muscles, are composed of fibers which are the radial lamellar type. In Periplaneta
americana Smit (1958) showed histological differences between the fast and the slow
contrasting coxal muscles under different fixatives.
THE ISOPTERA
The neck region: In shape and location, the lateral cervical sclerite in typical
termites (Fig. 58A) closely resembles that in Blattaria (Fig. 54A); its elongate anterior
part articulates with the occipital ridge, and the posterior part with the anapleuron. The
vestigial dorsal cervical sclerite is present in Archotermopsis (Grasse 1949), although it is
usually absent in Isoptera. The ventral cervical sclerite is also present in some termites.
The pronotum: The pronotum in Isoptera varies greatly in shape. It may be
flattened and shieldlike, or lateraliy lobed or often saddle-shaped. It is also typically
contiguous to the propleuron. In Arrhinotermes the pronotum is known to possess
winglike extension during postembryonic developmental stages ( cf. Fuller 1924).
The propleuron: The propleuron in typical termites (Fig. 58A) is generally similar
to the propleuron in cockroaches. The anterior oblique plate (precoxale of some
authors) is probably a fusion product of the anepisternum and preepisternum as in
Blattaria, although in Leucotermes (Crampton 1926a) the anapleural suture, which
divides the plate into the two original components, is apparently present. The dorsal end
of the anapleural plate ( anepisternal area) is attached dorsally to the anterior rim of the
pronotum (Fig. 58B). Thus, the dorsal end of the anapleuron on both sides, together
with the pronotum, form an almost entire and quite rigid collar band, to which the neck
membrane is attached. According to Fuller ( 1924), this condition is present throughout
the whole series of termite species he examined.
The katapleuron (Fig. 58A) is represented by two plates, the trochantin and the
lateral plate that is continuous with the ventral area of the pleural suture; the latter forms
the pleural process, which articulates with the coxa. The narrow membranous area
anterior to these sclerites roughly corresponds in position to the paracoxal cleft (suture).
The epimeron is an undivided lobe.
The prosternum: The presternum is largely membranous. In Syntermes, as
Crampton (1926a) showed, the basisternum is a triangular plate, and in Leucotermes
(Crampton 1926a) it is represented by paired elongate plates. The furcasternum in
these forms is represented by a small sclerite bearing the furcae internally. The pro-
thoracic spina was shown to be present by Crampton ( 1926a) on the membranous area
behind the furcal base in Syntermes.
The mesonotum: In typical termites (Fig. 59A) the antecosta demarcates the
laterally preserved acrotergite. The first phragma arises medially from the antecosta.
Evidently the prealar arm represents the modified lateral end of the acrotergite, as in
Blattaria. The horizontal prescutoscutal suture demarcates anteriorly the narrow pre-
scutum (antenatal ridge of Fuller, 1924). The anterolateral scutal suture demarcates
the anterior notal wing process, and is anteromesally continuous with the prescutoscutal
suture. Unlike Blattaria, however, the scutoscutellar suture is absent. Instead, in
certain termites (Cryptotermes, Calotermes, and Holotermes) the mesonotum is provided
with a median longitudinal ridge that arises from the prescutoscutal suture, although such
ANEPISTERNUM &
PREEPISTERNUM
ANTERIOR REDUPLICATION
of PRONOTUM
EPIMERON
PLEURAL
PROCESS
B
LATERAL
CERVICAL
SCLERITE

, SUTURE
KATEPISTERNUM
ANEPISTERNUM
AN EPISTERNUM &
PREEPISTERNUM
PARACOXAL
SUTURE
155
FIG. 58. Thoracic structures in a typical termite (from Fuller). A, ventral view of pro-
thorax and neck region; B, connection of propleuron and pronotum; C, lateral view of
propleuron.
a ridge is absent in Mastotermes (Fuller 1924). Typically, the transverse internal ridge
divides the mesonotum into the anterior and posterior halves. Fuller ( 1924) thought
erroneously that the anterior half was the scutum and the posterior half the scutellum.
This suture cannot be the scutoscutellar suture, which is absent in Isoptera. The position
of the transscutal suture in Isoptera is more caudal than the medially confluent terga1
fissure in Hymenoptera, Heteroptera, etc.
In typical termites the lateral margin of the mesonotum carries the anterior and
posterior notal wing processes. Other notal processes are not discernible, at least in a
Chilean species of Isoptera (unidentified) examined in this study. The emargination
behind the anterior notal process is inconspicuous; in the Chilean species a well-developed
ANTE COSTA
PRESCUTOSCUTAL
SUTURE
POSTERIOR
NOTAL
PROCESS
i
)
A
PLEURAL WING PROCESS --------.
ANEPISTERNUM &
KETEPISTERNUM -------1---
ANAPLEURAL
CLEFT--------
PREEPiSTERNUM -------+
B
ANTERIOR
3'RD AXILLARY
SCLERITE
AXILLARY
CORD
TRANSVERSE
RIDGE
---EPIMERON
------TROCHANTIN
FIG. 59. Mesothorax in a typical termite (from Fuller). A, dorsal view of mesonotum;
B, lateral view of mesopleuron.
longitudinal ridge follows this emargination along the lateral margin. The large, less
sclerotized posterior notal wing process forms a stump on which the folded wing lies.
The first and third axillary sclerites maintain their typical topographic relationships with
the anterior and posterior notal wing processes. The postnotum is absent.
The metanotuni: The anterior area of the metanotum is essentially the same as in
the mesonoturn. The mesonotum and the metanotum are united by a connecting mem-
brane. The postnotum is absent also in the metanotum.
The meso- and meta-pleuron: In a typical mesopleuron of Isoptera (Fig. 59B) the
anapleural cleft is distinctly present and continuous with the membrane at the base of the
trochantin. In this respect, the mesopleuron in Isoptera is conspicuously different from
the pterothoracic pleuron of Blattaria, in which the anapleural suture is often indistinct.
The pleural arm arises from the pleural process (Holmgren 1909; Fuller 1924; Ahrens
1930) that articulates with the coxa.
The paracoxal suture is not evident on the pterothoracic pleuron of Isoptera. Fuller
( 1924), however, suggested that the episternal depression v (Fig. 59B), which is the
muscle disc, represents the dividing line of the episternum into the anepisternum and the
katepisternum. Matsuda ( 1960a) thought that the muscle disc represented the point of
origin of muscle p-cx 5, which originates from the paracoxal suture in some insects.
Matsuda also pointed out the presence of a presumed paracoxal suture crossing the
pleural suture in a termite (La Greca 1949a) and in Zootermopsis (Grasse 1949); these
sutures, which extend dorsad on the epimeron are, however, probably not the true
paracoxal sutures.
In Isoptera the basalar cleft enclosing the basalare is distinct. It may even reach the
base of the coxa, as was found in Macrotermes (Fuller 1924). The preepisternum is
small as in Blattaria, and faces ventrally the lateral margin of the basisternum. The
trochantin is divided into two parts by an oblique suture on the surface. The subalare is
present. The metapleuron is similar to the mesopleuron, and the discussion on the meso-
pleuron generally applies to it.
The meso- and meta-sternum: In the mesosternum of typical termites (Fuller 1924)
the presternum is absent. The broad basisternum is provided with a median longitudinal
area, the sternellum of Delamare-Deboutteville ( 1952). The furcal pits are near the
median longitudinal axis of the sternum. In the Chilean species examined, the second
spina is continuous from the middle of the base of the furcasternum. In the metathorax
of typical termites (Fuller 1924) the median sternellum is not differentiated.
The adult musculature: Fuller (1924) studied the thoracic muscles of termites, and
Maki (1938) worked on the same in Odontotermes jormosanus. Table VI shows the
muscles in Odontotermes. The thoracic musculature in lsoptera is similar to that in
Blattaria, although fewer muscles have been recognized in Odontotermes than in Blattaria.
Conspicuously, the dorsal longitudinal muscle ( t 14) and p-ti 2, 3 are absent in Isopter a.
In this order t-cx 8 alone may be responsible for lowering wings.
THE MANTODEA
The neck region: The neck region in Mantodea is characterized by its suspended
condition, viz. it lies outside and below the prothorax. In Stagmomantis (Levereault
1936) and Mantis (LaGreca and Raucci 1949) the neck is provided dorsally with paired,
slender cervical sclerites and a lightly sclerotized anterior plate on each side of these
sclerites. As in Blattaria, the median slender sclerites lie beneath the pronotum, and
their extensions provide the points of attachment for the longitudinal muscles.
In Mantis each lateral cervical sclerite (Fig. 60A) is represented by four sclerites.
In Stagmomantis the small, first plate is, however, indistinguishable from the second
sclerite. The third sclerite in Mantis (second of Stagmomantis) articulates with the
posterior margin of the second, and bends horizontally inward. The fourth sclerite
articulates with the prothoracic episternum. In both genera two transverse ventral
cervical sclerites are present between the paired lateral cervical sclerites.
The pronotum: The prolonged pronotum is arched over the sternum. The inner
surface of the pronotum is provided with a transverse ridge at the level of the coxa
(Fig. 60C); the ridge separates the pronotum into the anterior prozona and the posterior
metazona. This transverse ridge cannot be homologized with any suture on the ptero-
thoracic terga, although Chopard (1949) thought that the ridge separated the prescutum
anteriorly. Another characteristic of the pronotum in Mantodea is a peculiarly folded
lateral edge of the pronotum (Fig. 61).
The propleuron: Despite the fact that the lateral pronotal area extends ventrally, the
propleural parts are well retained beneath the pronotum. In Stagmomantis (Fig. 60C)
158
VENTRAL
CERVICAL
SCLERITE
A
ANAPLEURAL
RIDGE?
PREEPISTERNUM
BASISTERNUM ------'
PARACOXAL
RIDGE
c
RCASTERNUM
SPIRACLE
B
ARM
TEPIMERON
PLEURAL CONDYLE
UM
FIG. 60. Prothorax in Stagmomantis carolinus (from Levereault). A, ventral view of neck
region; B, ventral view of posterior portion of prothorax; C, internal surface of propleuron.
the pleural ridge extends dorsally to the pronotum near the transversal ridge, and its
ventral end forms three knobs, including the pleural process (condyle), the pleural arm,
and a small process, which perhaps represents the katepimeron.
The paracoxal suture is perhaps represented by the ridge that is continuous from the
pleural ridge near its ventral end, at the level where the pleural arm occurs; in Mantis the
suture is apparently continuous with the pleural arm. The broad episternum is divided
into two parts by an oblique ridge that may represent the ridged anapleural suture. The
reduced katepisternal area is represented anteriorly by the nearly vertical trochantin and
the area beneath the paracoxal suture.
The prosternum: In the prothorax of Stagmomantis (Fig. 61) the anterior basi-
sternum meets the preepisternum along the pleurosternal suture; the posterior basisternum
between the coxae and anterior to the furcal pits is surrounded by a membranous area
which allows a considerable flexibility in the movement of the front leg. In the absence
of the sternacosta, the furcal pits represent the anterior limit of the furcasternum.
Aecording to Levereault ( 19 3 6) , the border between the furcasternum and the spina-
sternum in Stagmomantis lies at the level of the submarginal notches (Fig. 60B). In the
Cameroon mantid, according to Levereault, a transverse line between the corresponding
notches separates the furcasternum from the spinasternum; Levereault thought that in
Stagmomantis the two sternal areas had become indistinguishably fused by loss of this
suture.
159
PRE EPISTERNUM
FIG. 61. Ventral view of anterior portion of prosternum m :;tagmomanns caroanus \HOm
Levereault).
The mesonotum: The anterior area of the mesonotum (Fig. 62) is somewhat similar
to that of lsoptera (Fig. 59 A). As in Isoptera, the prescutoscutal suture is laterally
continuous with the anterolateral scuta! suture. The antecostal suture is medially lost,
and the poorly developed phragma arising from the antecosta on each side lies wide apart.
The anterior margin of the mesonotum is medially represented by the prescutum, and the
acrotergite is retained only laterally. The parapsidal suture reaches anteriorly the pre-
scutoscutal suture. A conspicuous process at the anterolateral angle of the mesonotum is
the prealar arm of acrotergal origin, as in Blattaria and Isoptera. It is a hollow process
with a wide opening at the base, and the whole process is invaginated into the thoracic
cavity. Snodgrass (1909) thought that the process was a phragma, because of its
invaginated state; but its position leads us to believe that it is a specialized prealar arm
with unknown function. Paired, median longitudinal sutures behind the parapsidal
sutures converge posteriorly (Fig. 64A); their rounded posterior ends may represent the
recurrent scutoscutellar suture. A well-formed postnotum is absent, as in Isoptera and
Blattaria.
The mesoscutallateral margin and forewing base structures: The mesoscutallateral
margin, as Onesto (1960) showed in Ameles (Fig. 63A), is similar to that of Isoptera in
having rather inconspicuous notal wing processes and a greatly swollen posterior area
ACROTERGITE
PRESCUTOSCUT AL
SUTURE
PHRAGMA
PRESCUTUM
PARAPSIDAL
SUTURE
FIG. 62. Dorsal view of anterior part of mesonotum in Stagmomantis carolinus (original).
MEDIAN
NOTAL PRO,CESSt'
POSTMEDIAN
NOTAL PROCESS
PLEURAL
WING PROCESS
SUBALARE
A
B
2'ND AXILLARY SCLERITE
DISTAL MEDIAN PLATE
3'RD AXILLARY PLATE
r----2'ND AXILLARY
SCLERITE
3' RD AXILLARY
SCLERITE
PLtURAL SUTURE
FIG. 63. Forewing base of Ame/es decolor (from Onesto). A, dorsal view; B, lateral view.
bearing the posterior notal wing process. As in Blattaria, the jugal area of the forewing
is broadly attached to the scuta! lateral margin behind the posterior notal wing process.
Onesto ( 1960) studied the wing articulation in the macropterous male of Ameles
decolor (Fig. 63A, B), as cited below. The degree of fusion of the sclerites involved
with wing articulation is more pronounced in the micropterous female.
The neck of the elongate first axillary sclerite connects with the basisubcostale and
with the anterior notal wing process; its posterior proximal margin articulates with the
scuta! lateral margin; its anterior distal margin connects with the second axillary sclerite;
and its posterior distal margin and the third axillary sclerite are sustained by the tendinous
parascutal membrane.
The dorsal second axillary sclerite is anteriorly connected with the proximal base of
the definitive distal median plate, which probably represents the basiradiale," and
proximally with the first axillary sclerite; distally it is associated with the proximal median
plate and separated from the latter by a narrow membrane (incisura submediana). As
"'0nesto (1960) found the second axillary sclerite was anteriorly connected with the base of Sc+R vein.
PRESCUTUM
PARAPSIDAL
SUTURE
SCUT UM--------\j[-
PARAPSIDAL SUTURE
POSTEROLATERAL
SCUTAL SUTURE
RECURRENT
SCUTOSCUTELLAR SUTU
161

BASISTERNUM
A
B
FIG. 64. Pterothorax in Stagmomantis carolinus (from Levereault). A, dorsal view of
pterothorax; B, dorsal view of inner pterothoracic sternum.
in Blatta the ventral second axillary sclerite (Fig. 63B) has associations with the pleural
wing process and with the ventral third axillary sclerite.
The proximal process of the dorsal thiq:l axillary sclerite (Fig. 63A) articulates
anteriorly with the dorsal posterior process of the second axillary sclerite, with the
posterior end of the proximal median plate and also with the first axillary sclerite by a
tendinous membrane. Its proximal process is articulated with the posterior notal wing
process. The posterior distal part of the definitive third dorsal axillary sclerite may
correspond to the basanale of Blatta (Fig. 55 A). The ventral third axillary sclerite
(Fig. 63B) is robust and elongate, and it articulates with the ventral second axillary
sclerite and with the subalare.
The definitive distal median plate, which is proximally the basiradiale, is distally
fused with the M+Cu vein, and is connected with the basisubcostale and the second
axillary sclerite. It is also distally connected with the base of the A veins complex.
In the male the humeral plate (basicostale) is present only ventrally (Fig. 63B) and
is articulated with the basalare, with the C vein, and distally with the basisubcostale. In
the female the humeral plate develops both dorsally and ventrally; dorsally, the humeral
plate articulates with the tegula, and posteriorly with the basisubcostale; and the ventral
humeral plate articulates as in the male.
The basisubcostale is present both dorsally and ventrally. The ventral basisub-
costale is represented by a small triangular plate that articulates anteriorly with the
humeral plate; proximally it articulates with the basalare, and also partly with the pleural
wing process. The base of the hind wing is very similar to the base of the forewing.
The metanotum: In Stagmomantis (Fig. 64A) the metanotum is similar in shape
and size to the mesonotum. The recurrent scutoscutellar suture is apparently con-
tinuous with the posterolateral scutal suture.
The mesopleuron: The elongated mesopleuron (Fig. 65A-c) is divided by a nearly
horizontal pleural suture into the anteroventral episternum and the posterodorsal
PREEPISTERNUM PLEUROSTERNALSUTURE
A
PLEUROSTERNAL SUTURE(CLEFT) f\
( _ __., \ ___ ANEPISTERNUM
BASI STERNUM----- ( - t--+EPISTERNAL
----,..__ ( LONGITUDINAL S.UTURE
TROCHANTIN \0l fl
FURCA
EPISTERNAL
LONGITUDINAL SUTURE
AN EPISTERNUM
TROCHANTIN
B
c
ANTERIOR
BASALARE
PLEURAL SUTURE
FIG. 65. A, lateral view of pterothorax in Mantis religiosa (from La Greca and Raucci);
B, ventral view of mesothorax in wingless Geomantis larvoides (from La Greca); C, ventral
view of mesothorax in Mantis religiosa (from LaGreca and Raucci).
epimeron. - The pleural arm arises from near the point of pleurocoxal articulation, as in
the prothorax ..
The paracoxal suture is absent. The presence or absence of the anapleural suture
(or cleft) is somewhat difficult to prove. One interpretation (Fig. 65B), which is pro-
visionally accepted here, is that the anapleural cleft changed its course caudally, accom-
panying the prolongation caudad of the an episternum. This ana pleural cleft still reaches,
as in Dermaptera (Fig. 79A), the anterior end of the trochantin where the episternal
longitudinal suture reaches. The preepisternum, which is demarcated posteriorly by this
anapleural cleft, is present in some mantids, such as Geomantis (Fig. 65B). This
interpretation was proposed by Matsuda ( 1960a).
Another interpretation is that the anapleural cleft is absent, since the proximal end
of the apparent anapleural cleft, as identified above, is abnormal. If this is so, the
elongated anepisternum in Fig. 65A-C must be a fusion product of the anepisternum and
the preepisternum, and the elongate plate along the posterior lateral margin of the
sternum (preepisternum) must be the ventropleurite. Since the ventropleurite is absent
in related orders and since the plate lies somewhat too anteriorly to be a katapleural
ventropleurite, this second interpretation is not highly acceptable.
The anterodorsal anepisternal area is usually separated off by the membrane that
surrounds it. The production of this additional basal are (the anterior basal are of
La Greca, 1949a) is evidently a specialization in this order; a corresponding plate is
absent in Blattaria and Isoptera. A longitudinal suture that extends anteriorly from the
anterior end of the trochantin (Fig. 65A-C) is apparently homologous with the episternal
longitudinal suture present in Dermaptera (Fig. 79A), although in Mantodea it may not
reach the membrane surrounding the basalare.
The katepisternal area is represented by a large, bipartite trochantin; its oblique,
distal margin reaches the pleurocoxal articulation. The original katepisternal area distal
to the trochantin has, with the loss of the paracoxal suture, become completely fused with
the anepistemum. The epimeron is reduced, and the subalare lies above the dorsal
margin of the epimeron.
The metapleuron: The metapleuron is similar to the mesopleuron. The anapleural
cleft identified in the mesopleuron is, however, often absent (e.g., Mantis, Fig. 65A;
Stagmomantis, Levereault 1936; Hierodula, Matsuda 1960a).
The mesosternum: The presternum is absent. The anterior half of the lateral
margin of the basisternum faces the proximal margin of the caudally prolonged
anepistemum (Fig. 65B, C). Often the posterior half of the definitive basisternum is
laterally swollen, due to incorporation of the preepisternum (Fig. 65C). The basisternal
surface is often provided medially with a V-shaped, internally ridged (Fig. 64B) suture;
the ridge is posteriorly continuous with the furcal base. The mesothoracic spina is well
retained in Stagmomantis.
The metasternum: The metasternum (Fig. 64B) is similar to the mesosternum,
although the distinct spina is absent
The adult musculature: Maki (1938) studied the thoracic musculature of Hierodula
patel/ifera, and Levereault (1938) and La Greca and Raucci (1949) studied it in
Stagmomantis carolinus and Mantis religiosa. Since La Greca and Raucci homologized
the muscles in Hierodula and Mantis, only the muscles of Stagmomantis and Mantis are
shown in Table VII. The musculature in both genera is remarkably similar. As in
Isopter a, t 14 and p 3 are absent, and the flight ability of mantids is restricted.
The postembryonic development and histology of muscles: According to Tiegs
(1955), the development of leg muscles in Orthodera ministralis proceeds first by the
usual fiber enlargement, then fiber cleavage ensues after the nymph has grown 2 em long.
In cross-section t-tr 1 has a somewhat irregular radial pattern of fibrils.
TABLE VII
Thoracic musculature in Blattaria, Isoptera, and Mantodea
Mantis Stagmomantis
Periplaneta Odontotermes (LaGreca & (Levereault Muscle
(Carbonell 1947) (Maki 1938) Raucci 1949) 1938) designations Remarks
59 1 2 5 cv(d)-t 1
52
5.
7b op-t 1
53 4 6 1c op-t 2
1 7a op-t 3
57 7 op-cv 1 or 2
58 8 8 1a, b op-cv 2 or 3
56 op-p 1
12 op-p 2
54 11 6 t-s(cv) 1
10 2 t-cv 1
63, 64 9 9 3 t-cv 2
69 t-cv? 1
84 cv-s 1
lcv-cxlx (Chadw.) 22 cv-cx 1(x)
4 cv-cx 3
55 5 7 8 s 1
62 2 3 -37 t 12
112, 113 26, 27 29, 32 52, 53
155, 156 56, 57 66 90
60, 61 3 4 35 t 13
111 25 30, 31 54
154 55 64, 65 89
59 1 1, 2 5, 7a t 14
110
153
56 t-p 3
108, 109 34 39 47
150 63 69 85
121 (II)? 32 36 48 t-p 6 or 5 2
108? 40 (II) 46 t-p 7
65, 66 (I)?
116 (II)
t-p 10
159 (III)
41 (I I) 51 (II) t-p 12
70 (III) 88 (III)
115(II) 36 43 49 t-p 14
158 (III) 65 72 86
114 (II)? 35 42 50 t-p 15
157 (III)? 64 71 87
67, 68 (I) 13 14, 15 20, 21 t-p (prothoracic)
54 (0) t-s 1
96 (HI) 10
125 (II-III) 33 38 55
190 (III-IA) 62 68 91
12 (I-II) 36 t-s 2
13 (HI) 38 t-s 8
121 (II) 32 t-ti 1? 3
or t-p 5?
70? 14 t-ti(cx) 1
118 38
16.1 67
TABLE VII (Continued)
Mantis Stagmomantis
Periplaneta Odontotermes (LaGreca & (Levereault Muscle
(Carbonell 194 7) (Maki 1938) Raucci 1949) 1938) designations Remarks
71? 18 t-ti(cx) 2 4
119 46 62a
162 74 95a
72? 73? 21 (1)? 17 9 t-ti(cx) 3 4
120 47 62b
163 75 9Sb
74 (I) t-cx 5
129 (II) 43 49 t-cx 6
174 (III) 72
130, 131 (II) 42 48 65 t-cx 7 5
175, 176 (III) 71 76 97
128 (II) 44 60 70 t-cx 8
169 (III) 73 86 102
76-78, 79 16, 17, 18 19, 20a-d 10, 13, 14, 15 t-cx 6, 7, 8
80, 81 (I) 21a, b 16, 17
16 (I) 22 p-s 1
117(11) 37 45 56
160 (III) 66 73 92
103 (I-II) 31
61 (II-III)
44 45 p-s 2
102 (I-ll) p-s 4
13 (Chadw.) 13 (Zootermopsis) 24 (Tenodera) p-s 5 6
24 (Chadw.) 24 (Zoot.)
31 (Chadw.) 31 (Zoot.)
4 (I) (Chadw.) 4 (Zoot.) p-s 6 6
17 (II) (Chadw.) 83
28 (Ill) ( .. )? 28 (Zoot.)
35 (I-II) 44 p-s 9
189 (II-2A) p-s 13
39 (II) 57 60 p-ti(cx) 1
68 (III) 83 93
122, 123 (II) 54 61 p-ti(cx) 2 7
164, 165 (Ill) 80 94
124 (II) 55 63 p-ti(cx) 3 7
166 (III) 81 95
75 26 19 p-cx4 7
126 50 56 64b
167 79 82 96b
22 (I) 26 19 p-cx5 7
127 (II) 47-49 58, 59 64a, c
168 (III) 76-78 84, 85 96a, c
97 (I) p-cx 7(x)
106 (I-II) 33 42 sll
152 (II-III)
104 (!-II) 30 34 43 s 12
151 (II-III) 60
191? (III-2A)
100 (I-II) 28 s 13
148 (!I-III) 59 67 84
192 (III-IIA)?
99 (I) 6 s 14
101 (I) 29 s 15
fu3-fu3 (Chadw.) s 16
193 (III -2A) s 20
TABLE VII (Continued)
Mantis Stagmomantis
Periplaneta O.lontotermes (LaGreca & (Levereault Muscle
(Carbonell 1947) (Maki 1938) Raucci 1949) 1938) designations Remarks
105 (II) 40 s-ex 1
173 (III) 69
20 (I)? s-ex 2
133 (II) 46 52 69
170 (III)
74, 75 79 100
82 (I)
20 25 12 s-ex 3
171 (III)
99
98 (I) 19 s-ex 4
134 (II)
45 53 67
15 23 11 s-ex 5
41 50 66
70 77 98
83
24 18 s-ex 6
132 51 68
172
78 101
85a, b 23 27a, b, c 23c, d, e t-tr 1
135a
51 61 71c
177a 80 87 103c
85c, d (I) 24 28 23b p-tr 1 or 3 8
135c (II) 52 62 71d p-tr 2
177c (Ill) 81 88 103d
135b (II) 53 63 71b s-tr 1
177b (Ill) 82 39 103b
107 (II) 54 40 spiracle
147 (Ill) 83 58 occulsor 9
41 spiracle 9
57 dilator
39 sternal sp. m.
REMARKS (Table VII)
( 1) 69 in Periplaneta arises from the anterior part of the pronotum and goes
downward to its insertion on a fold of the ventral integument of the neck. This muscle
may be t-cv 1.
(2) 36 and 48 in Mantodea arise dorsally from the scutum and were described as
being inserted on the basisternum. Apparently, however, they are inserted on the pre-
episternal part of the definitive basisternum. Therefore, they probably represent t-p 6 or
possibly t-p 5. For 121 in Periplaneta, see remark 3.
(3) 121 in Periplaneta extends between the first phragma and the lateral margin-of
the basisternum, and no serial homologue is present in the metathorax. In Blattella, as
Matsuda (19S:6a) showed, the serial homologue of the muscle in the metathorax is
inserted on the proximal (ventral) anterior margin of the trochantin. If this point of
attachment is primary as Matsuda contended, the muscle is a t-ti 1 and agrees well in
position with 71 in Lepisma. If, however, the insertion on the basisternum is primary,
this muscle is a t-s muscle. The position of muscle 121 in the mesothorax, however,
does not exactly agree with any one of the t-s muscles. Another possibility is that this
muscle is t-p 5. This muscle appears late during postembryonic development (Tiegs
1955; Storch and Chadwick 1967); this fact supports the interpretation that the muscle
is t-p 5, since in other insects t-p 5 appears late during the postembryonic development.
( 4) 9 in Stagmomantis is a three-part muscle, 17 in Mantis a four-part muscle,
62a in Stagmomantis a two-part muscle, and 95a in Stagmomantis a three-part muscle.
All of them are inserted on the same point, the tip of the trochantin.
(5) 65 in Stagmomantis is a two-part muscle and it may correspond to 48 and 49
in Mantis.
(6): Muscles in Zootermopsis and Tenodera were described by Chadwick (1957);
those in the column of Periplaneta are muscles in other cockroaches described by
Chadwick (1957).
(7) p-ti 2, 3 arise from the dorsal area of the anepisternum in Periplaneta, and the
same muscles in Mantodea arise from the anterior basalare, which corresponds to the
dorsal part of the anepisternum in Periplaneta. p-cx 4, 5 arise from the secondary
anterior basalare in Mantodea, and this point of origin is secondary. No muscle is
attached to the small posterior basalare in any of the three orders.
(8) 28 in Mantis was described as a epimerotrochanteral muscle; 85c in Peri-
planeta arises partly from the epimeron.
(9) These spiracle muscles are not necessarily homologous.
In Nauphoeta cinerea Storch ( 1968) showed the following muscles that are absent
in Periplaneta americana: T 1-Cx 1 No. 10 (a posterior tergocoxal muscle), 1 Ph-POR
(op-t 3), PlR 1-Cx 1 Nos. 1 and 2 (p-cx 6), DCer-Tent (t-s(cv) 1), 1 Ph-1 LMemb
(t-p 17), SA 1-PlR (p-s 1), SA 1-1 LMemb (p-s 5), T 2-2 Ph No.2 (t 13?), SA 2-2
LMemb (t-p 17), T 3-3Ph No.2 (t 13?), SA 2-Cx 2 No.3 (s-ex 5), SA 3-SA 3 (s 16),
SA 3-Cx 3 No. 3 (s-ex 5).
THE GRYLLOBJ.ATTODEA
The neck region: In Grylloblatta the lateral cervical sclerite (Figs. 66, 67) consists
of a larger anterior lateral cervical sclerite that articulates with the occipital condyle, and
a smaller posterior lateral cervical sclerite which, according to Walker (1938), is con-
tiguous to what he called the coxopleurite, close to the junction of the latter with the
trochantin. As will be discussed below, the posterior point of articulation is the pre-
episternal area. Hence, the lateral cervical is presternal in origin.
The prothorax: The pronotum is a simple plate and laterally curved inward, thereby
covering the dorsal part of the propleuron.
The propleuron (Fig. 66) is fairly generalized. At its ventral end the pleural ridge
is provided with a well-developed condyle, which articulates with the coxa. The para-
coxal suture is considered lost. An imaginary line representing this last suture is shown
by a broken line that meets the anapleural suture, and the distal end of this imaginary line
also reaches the point of the pleural ridge where the pleural arm extends internally.
The anapleural suture is well marked. The area dorsal to this suture is the anepi-
sternum, and the preepisternum (basal pleural sclerite of Crampton, 1915; basal sclerite
of Walker, 1938) lying posterior to the anapleural suture is indistinguishably fused with
the katepisternum. As seen in Fig. 66, the posterior lateral cervical sclerite articulates
with the preepisternal area. The dorsal part of the anepisternum is invaginated into the
thoracic cavity beneath the pronotum. The large trochantin is separated from the
katepisternum proper; the epimeron is also well preserved.
The prosternum: The prosternum (Fig. 67) is extensively desclerotized. The basi-
sternum has a median longitudinal ridge, and the paired furcal pits lie posterior to the
basisternum. The first spina demarcates the posterior limit of the prosternum.
The meso- and meta-notum: Both the mesonotum and the metanotum are shorter
than the pronotum, and .without subdivision by sutures. The anterior margin is ridged
but lacks a phragma, and its lateral area is slightly deflected to cover the dorsal part of
the pleuron.
The meso- and meta-pleuron (Fig. 67): The pleural ridge runs obliquely; a T-
shaped expansion at its upper end is comparable with the pleural wing process. The
pleural arm is situated near the posterior (ventral) fourth of the pleural ridge in the
mesopleuron, and more anteriorly in the metapleuron. The pleural arm is barely
168
ANAPLEURAL SUTUR
PRONO
PLEURAL
RIDGE

PLEUROCOXAL
ARTICULATION
AN EPISTERNUM
PREEPISTERNUM
LATERAL
CERVICAL SCLERITE
PARACOXAL
SUTURE
KATEPISTERNUM
TROCHANTIN
FIG. 66. Lateral view of prothorax in Grylloblatta campodeiformis (from Walker).
connected in front with the pleural ridge, because it is chiefly a process of the epimeron.
This epimeral origin of the pleural arm is more evident in the metapleuron, judging from
the figure given by Walker ( 1938). The paracoxal suture cannot be recognized.
The anapleuron on the episternum consists of the preepisternum (precoxale of,
Walker) and the anepisternum, which are separated by the anapleural cleft. The
anepisternum is indistinguishably fused with the katepisternum along the pleural ridge,
owing to the loss of the paracoxal suture. The preepisternum is separated from the
basisternum by the longitudinal pleurosternal cleft.
The trochantin is large; a deep infolding of the cuticle exists along its anteromesal
margin, and the proximal part of this fold is thickened into a ridge that terminates at the
acute posteromesal angle in the articulation with the coxa.
The epimeron is almost membranous except over a rather narrow area along the
pleural ridge. Neither the subalare nor the basalare is differentiated.
The meso- and meta-sternum (Fig. 67): The border between the mesosternum and
the metasternum is marked by the second spina. Only the basisternal area is distinctly
sclerotized. The furcal pits lie along the posterior margin of the definitive, sclerotized
sternum. A remarkable fact is the presence of .the third spina in the metasternum.
The adult musculature: Walker's ( 1938) study on Grylloblatta campodei/ormis is
the only work on the thoracic musculature of this order. As seen in Table VI, the
characteristic features of thoracic muscles in Grylloblatta are the presence of t 14 and
p 3, which are flight muscles and are usually absent or only weakly developed in other
wingless pterygotes.'" A few t6rgopleural muscles, which connect the dorsal part of the
pleuron and the lateral part of the tergum in other insects, are also present, although
these muscles are absent in wingless Dixippus shown in the same table. The loss of the
posterior tergocoxal muscles is also noteworthy.
'"It should be recalled that flight muscles in wingless forms of Coleoptera, Heteroptera, etc., tend to be
reduced or lost.
169
LATERAL CERVICAL
BASISTERNUM
S C L E R I TE
TROCHANTIN
FURCA
SPIRACLE
PLEUROSTERNAL
CLEFT
PREEPISTERNUM
SPINA 2
FURCA
TROCHANTIN
FIG. 67. Ventral view of prothorax and mesothorax in Grylloblatta campodeiformis (from
Walker).
THE ORTHOPTERA
The neck region: In Orthoptera, according to Alicata ( 1962b), the anterior articula-
tion of the lateral cervical sclerite is either with a small sclerite that Alicata called the first
cervical scierite ( Cv 1 in Fig. 68A), or directly with the postocciput. The lateral cervical
sclerite (second cervical sclerite of Alicata) may or may not be divided into anterior and
posterior parts (Fig. 68B). The presternal origin of the lateral cervical sclerite is evident
in that it articulates posteriorly with the precoxale, which is anapleural (p. 170).
The small first cervical sclerite (Fig. 68A) appears to be a detached postoccipital
condyle, and it is present in Gryllacridioidea, Tettigonioidea, Tridactyloidea, and
Acridoidea (except Tetrigidae), as Alicata thought. op-cv 1 (1 of Alicata) is inserted
on this small plate. In Tetrix (Fig. 68B) no such detached plate occurs, and this repre-
sents a more primitive 'condition in Orthoptera.
Alicata also believed that the undivided lateral cervical sclerite represented a primi-
tive condition. Such an undivided lateral cervical sclerite is present in Dolichopoda,
Oecanthus, Tettigonioidea, etc.; in Acheta ( = Gryllus) it is incompletely divided
(Fig. 68A). Alicata's belief appears justified, since in primitive genera, such as Capnia
(Plecoptera) and Siphlonurus (Ephemeroptera), the lateral cervical sclerite is a simple
170 MEMOIRS OF TilE ENTOMOLOGICAL SOCIETY OF CANADA
undivided plate... Furthermore, as Alicata's study indicates, the undivided lateral
cervical sclerite occurs commonly in Grylloidea, Tettigonioidea, and Gryllacridioidea;
and its division into two parts in these superfamilies occurs only in a highly specialized
genus, Gryllotalpa. On the other hand, in Acridoidea, in which the thoracic structures
are generally highly specialized (as will be discussed later), the division of the lateral
cervical sclerite into two pieces occurs rather consistently. It appears, therefore, that in
Orthoptera the undivided lateral cervical sclerite represents a more primitive condition.
The division of the lateral cervical sclerite into two plates is present in the first-stage
nymph of Dociostaurus (Iannone 1939). This is comparable with Periplaneta (Storch
and Chadwick 1967), in which the division of the lateral cervical sclerite also occurs in
the early stage nymph.
The dorsal cervical sclerite in Dolichopoda serves as an anterior point of attachment
for a dorsal longitudinal muscle, cv(d)-t 1. Because of this attachment, Alicata thought
the dorsal cervical sclerite in Dolichopoda and Troglophilus was homologous with the
acrotergite. This homology is questionable, as already discussed (p. 9). A dorso-
lateral cervical sclerite occurs in Tetrix (Fig. 68B). The ventral cervical sclerite does
not occur commonly in Orthoptera; it is present in Acheta.
The pronotum: In Orthoptera the pronotum overgrows the pleuron (Fig. 69A-C).
The lateral margin of the pronotum is either the free marginal ridge, or it is simply
continuous with the pleuron through the intervening membrane between the pleuron and
the pronotum. Ander ( 1939) thought legitimately that, at least in lower Orthoptera, the
pronotum with the free lateral margin or edge is more primitive.
In many groups of Orthoptera the pronotum is laterally bent and is distinctly higher
than it is wide (e.g., Tettigonioidea, many Rhaphidophoridae, and Stenopelmatidae,
Ander 1939). Accompanied by such an elevation, formation of transverse ridges on the
pronotum commonly occur in Orthoptera. Thus, DuPorte (1919) showed, in Romalea,
four transverse internal sulci on the pronotum. He thought the first (anterior) sulcus
marked the farthest point to which the head might be retracted within the prothorax, and
hence this sulcus represented the anterior extremity of the primitive pronotum. DuPorte
also considered that the dorsal part of the second sulcus and the whole third sulcus had
been produced by the pull of muscles, and that the fourth represented the posterior
extremity of the primitive pronotum.
An excessive prolongation of the posterolateral part of the pronotum, especially in
Tettigonioidea and Acridoidea, frequently covers the base of the forewing; La Greca
(1947a) discussed how this enlarged area is raised when the wings are spread (see
p. 185). Uvarov and Thomas (1942) showed how phasic differences in the shape of the
pronotum in Locusta migratoria are produced by mechanical causes associated with
muscular pull.
The prop/euron: A large part of the propleuron is overgrown by the pronotum.
This hidden propleuron in Orthoptera was first discovered by Crampton (1918c) and
DuPorte (1919), and various authors since Carpentier (1921a, b; 1923) have called it
the cryptopleuron. As DuPorte showed, the almost completely hidden propleuron is
present in Tettix, Conocephalus, Microcentrum, etc. The cryptopleuron is also evident
in Gryl/oblatta; and according to Ander ( 1939), it also occurred in Protorthoptera."
The wall of the cryptopleuron has undergone varying degrees of desclerotization in
various groups of Orthoptera, and this is especially pronounced in the epimeron. How-
ever, the pleural ridge is always present. Both DuPorte (1919) and Ander (1939) have
traced the reduction of the pleuron in a series of groups of Orthoptera.
In a relatively well-exposed propleuron, such as that in Platycleis (Fig. 69A),
neither the anapleural suture (cleft) nor the paracoxal suture is present on the episternum,
but the trochantin is clearly present and the anapleural plate is indistinguishably con-
tiguous to the basisternum. The anterior margin of the anapleuron (precoxale) is
46A divided lateral cervical sclerite. however, is also commonly present in some other lower insects such as
Blattaria, lsoptera, Mantodea, etc. '
4
7'fhe cryptopteuron also occurs in the pterothorax of wingless Tachycines (Carpentier 1921b) and Gymno-
plectron (Richards 1955).
cv l
POSTOCCIPUT
ANTERIOR
LATERAL CERVICAL
SCLERITE
LATERAL
CERVICAL SCLERITE
A
FIG. 68. A, neck region in Gryllus campestris (from Alicata); B, neck region in Tetrix
subulata (from Alicata).
172
A
EPISTERNAL
APODEME-----L
PLEURAL
TROCHANTI
FURCA
EXTRA PLEURAL
SPACE
\
'
-
---i-f+------PRONOTUM
I
I
I
,, /
i ' ....... "'
--- \
EPIMERON
PRONOTAL CREST (PLATE)
SPINA
MESONOTUM METANOTUM
/EPISTERNUM
FURCA
FIG. 69. A, lateral view of prothorax in Platycleis intermedius (from Onesto); B, lateral
internal view of Gry/lotalpa vulgaris (from Carpentier); C, lateral internal view of Cylindro-
ryctes spegazzini (from Carpentier).
MATSUDA: THE. INSECT THORAX 173
provided with a well-developed ridge that is also continuous with the similarly ridged
anterior margin of the basisternum. Onesto ( 1963) called this marginal area the pre-
costal suture (sutura precostale). In a cave cricket genus, Gymnoplectron (=Macro-
pathus), Richards (1955) called this marginal ridge the verasternite. From observation
of this genus the author is convinced that this area in Gymnoplectron is a well-developed,
marginal reinforcing ridge, or the prepectus.
In Gryl/otalpa (Fig. 69B), Carpentier (1921a, 1936), Camerlengo (1936), and
Rakshpal ( 1954) found that the pleural ridge runs obliquely dorsocaudad from its
ventral condyle, and the epimeron is well retained. These authors thought that the
presumed epimeron was the pleural process (arm); Carpentier ( 1936) reinterpreted it
correctly as the epimeral process, and contended that if it is the pleural process,
Gryllotalpa would be the only genus of Orthoptera in which it occurs. Carpentier
(1936) showed that in G. gryllotalpa (Fig. 69B) a triangular notal crest is formed on the
dorsal part of the pleural ridge; this crest is continuous anteriorly as a narrow plate, and
becomes directly attached to the pronotum. In Cylindroryctes (Fig. 69C) the pleural
ridge as a whole is greatly expanded dorsally to be continuous with the endonotal ridge.
The prosternum: The anapleuron is proximally fused with the basisternum in the
prothorax. The furcasternum, which bears the furcal pits, is usually greatly reduced;
the furcal pits are often united by the internal transverse ridge or the sternacosta, as seen
in Dissosteira (Snodgrass 1929a) and Platycleis (Fig. 76B). In Orthoptera the pro-
thoracic spina lies generally close to the furcal base, as in the mesothorax. In Platycleis
(Fig. 76A, B) and Dissosteira (Snodgrass 1929a) the spina arises from a narrow trans-
verse plate, which Onesto ( 1963) and Snodgrass ( 1929a) called the spinasternum. In
Gryllotalpa (Fig. 69B) the greatly developed spina arises directly from the furcal base
and its distal eild is directed posteriorly; in Cylindroryctes (Fig. 69C) it extends even as
far as the metathorax (Carpentier 1936). In Catantopinae, a raised tubercle or spine on
the presternum between the legs helps coordinate the movements of the front legs
(Rowell 1961).
The mesonotum: In the anterior area of the mesonotum of Tettigoniidae, as Onesto
(1963) showed, the prescutoscutal suture traverses the entire notal surface as in Platycleis
(Fig. 70A), and the suture is medially lost in Tylopsis (Onesto 1963). In both genera
the prescutoscutal suture is not confluent either with the antecostal suture, or with the
anterolateral scuta! suture. In Acridoidea there is a tendency for the prescutoscutal
suture to lie more caudad at its lateral ends, and the prescutoscutal suture and the antero-
lateral scuta! suture tend to merge into a single continuous line. In Podisma (Fig. 70C)
and Anacridium (Fig. 71A) the two sutures are still distinct. In Dissosteira (Snodgrass
1929a), Schistocerca (Misra 1947) and Locusta (Albrecht 1953) the two sutures have
completely merged and the original point of union is lost.
The acrotergite is well defined by the antecosta, and the prealar arm represents the
lateral extension of the acrotergite in Orthoptera (Fig. 70A), as LaGreca (1947b) first
pointed out. In Lamarckiana, according to Thomas (1952), this acrotergal prealar arm
articulates with the first basalare (an episternum).
In Tylopsis (Onesto 1963) and Platycleis (Fig. 70A) the remnant of the scuto-
scutellar suture is apparently retained, although in Acridoidea it has become completely
or almost completely lost. Therefore, the division of the mesonotal surface into the
scutum and scutellum is absent in many species of Orthoptera.
In the posterior half of the mesonotum the recurrent scutoscutellar suture frequently
arises from the middle of the scutellar posterior margin (Fig. 70A). In Acridoidea
there is a characteristic subhorizontal suture, which approaches the scuta! margin between
the postmedian and posterior notal wing processes (Fig. 70C); Snodgrass (1929a) called
this suture in Dissosteira the secondary suture, Misra ( 1947) erroneously called the area
behind this suture in Schistocerca the lateral scutellum; and Thomas (1952) followed
Misra in the study of Lamarckiana.
The scutellum is laterally continuous with the axillary cord. A well formed post-
notum is absent. However, paired sclerites beneath the scutellar area in Orchelinum
174
PRESCUTOSCUTAL SUTURE
SCUTOSCUTELLARSUTURE
RECURRENT
SCUTOSCUTELLARSUTURE
POSTNOTUM
A
B
PRESCUTUM
ANTEROLATERAL
SCUTAL SUTURE
PRESCUTUM 3

PRESCUTOSCUTAL
SUTURE
c
SECONDARY SUTURE
FIG. 70. A, mesonotum of Platycleis intermedia (from Onesto); B, dorsal segmental border
between mesonotum and metanotum in Orchelinum sp. (original); C, right half of mesonotum
in Podisma pedestris (from LaGreca).
(Fig. 70B) and probably in some other genera may represent a remnant of the
postnotum.
The mesonotallateral margin: Compared with other lower hemimetabolous orders,
the mesonotal lateral margin in Acrididae and Tettigoniidae is more specialized. In
Anacridium (Acrididae), as LaGreca (1947b) showed (Fig. 71A), the large, lateral
prescutal area is differentiated into an articular lobe, and the head of the Sc vein fits into
the cavity formed by this lobe; a small apodeme lies near the articular lobe, and a tergo-
pleural muscle (t-p 4) is attached to it. In Schistocerca, according to Misra (1947),
ANTEROLATERAL
SCUTAL SUTURE
MEDIAN NOTAL
PROCESS
POSTMEDIAN NOTAL
PROCESS
AXILLARY CORD
TERGAL
FISSURE
POSTMEDIAN
. NOTAL PROCESS
SCUTUM
MATSUDA; TilE INSECT TIIORAX 175
A
B
-----PREALAR PROCESS (ARM)

,..--------TEGULA
ANTERIOR NOTAL
PROCESS
l'ST AXILLARY
SCLERITE
4TH AXILLARY
SCLERITE
ANTERIOR NOTAL
PROCESS
'ST AXILLARY
SCLERITE
Fm. 71. A, mesonotallateral margin of Anacridium aegyptium (from LaGreca); B, lateral
margin of Tettigonia viridissima (from La Greca),
the prescutum is divided into two areas by an intraprescutal suture, which gives rise to
two apodemes for the attachment of muscles, t-p 4 and t-p 5. In Schistocerca the distal
part of the prescutum has a vertical notch that accommodates the condyle of the costo-
subcostal sc!erite, and a horizontal notch that provides a socket for the curved anterior
process of the first axillary sclerite.
In Anacridium (Acrididae) the scutallateral margin (Fig. 71A) has roughly three
areas: the anterior and posterior lobes and the median concavity. The anterior,
antemedian, median, and postmedian notal wing processes articulate with the first axillary
sclerites. A small emargination, the tergal fissure, is present posterior to the antemedian
ANTEMEDIAN
NOTAL
PROCESS-
l'ST AXILLARY
SCLERITE
3' RD AXIL SCL
B
A
I
Anr
~
PROXIMAL ANTERIOR
PROCESS of 2'ND
AXIL SCL
LIGAMENT
FIG. 72. A, forewing base in Anacridium aegyptium (from La Greca); B, connections of
second axillary sclerite in Anacridium aegyptium (from La Greca).
notal wing process. Normally, the posterior notal wing process is represented by the
fourth axillary sclerite.
In Tettigoniidae the differentiation of the notal wing processes is much less
pronounced than in Acrididae. The lateral prescutal area is membranized. In
Tettigonia (Fig. 71B), according to LaGreca (1947b), a semilunar plate in the mem-
branized area has the same relative position as the articular lobe has in Anacridium; viz.
it lies near the tegula and is associated with the articular head of the subcostale.
In Gryllidae also (personal observation), a small prescutal sclerite occurs in a corres-
ponding position.
The small sclerite near the articular lobe in Anacridium, the semilunar sclerite in
Tettigonia, and the small prescutal sclerite in Gryllidae are homologous with the prealar
sclerite in some other orders.
The forewing base structures: In Anacridium the arrangement of the axillary
sclerites is shown in Fig. 72A. Noteworthy are the ligamentous connections between the
first axillary sclerite and the antemedian notal wing process, the articulation of the head
of the first axillary sclerite with the anterior margin of the second axillary sclerite, and
FIG. 73. A, proximal end of C + Sc vein in forewing of Anacridium aegyptium (from
LaGreca); B, connection of C and Sc veins in forewing with dorsal part of mesopleuron in
Zonocerus elegans (from La Greca); C, ventral connection of C + Sc vein in forewing with
dorsal part of mesopleuron in Anacridium aegyptium (from LaGreca); D, hind wing base in
Metrioptera sepium (from La Greca).
HUMERAL
PLATE
LIGAMENT
BASI COSTAL
SCLERITE
DISTAL MEDIAN
PLATE
3'RD AXILLARY
SCLERITE
BASI COSTAL
SCLERITE
B
177
ANTERIOR
PROCESS
ANTEBIOR
BASALARE
D
COM CAVITY of Sc
PRESCUTAL
ARTICULAR LOBE
l'ST AXILLARY
SCLERITE
2'ND AXILLARY
SCLERITE
POSTERIOR NOTAL
PROCESS (4TH
AXIL. SCL.)
the distal connection of the second axillary sclerite with the distal plate." The
distal part of the large, horizontal, third axillary sclerite may correspond to the basanale
in other insects, such as Forficula (Fig. 82D) and Blatta (Fig. 55A). The fourth
axillary sclerite is the detached posterior notal wing process. The ventral second axillary
sclerite (Fig. 72B) is articulated with the dorsal surface of the pleural wing process by a
tendinous ligament, and its apical process is connected with the subalare by a mem-
branous tendon.
In Anacridium (Fig. 73A-C), as La Greca (1947b) showed, the C vein of the
forewing has a long and robust elbow-shaped base, which starts from the point of union
with the Sc vein. It has a broad, membranous, anterior concavity in which the humeral
plate lies; it bears also the preapical anterior and posterior tubercles or processes.
Further, in Anacridium the proximal articular head of the subcostal base consists of the
dorsal and ventral lips (Fig. 73A, C); the dorsal lip has two apical obtuse projections.
When the wings are closed, the anterior projection of the dorsal lip fits into the apical
cavity of the prescutal articular lobe, and the posterior projection fits, by means of a
robust membrane, into the apical deflection of the neck of the first axillary sclerite.
When the wings are open, the anterior projection lies against the apex of the prescutal
articular lobe. With these connections, the dorsal lip plays an important role in the
opening and closing of the wing. The ventral lip is connected with the pleural wing
process by a tendinous ligament (Fig. 73C).
The anterior membranous concavity of the Sc vein base (Fig. 73A) is connected
with the distal margin of the prescutal articular lobe. Similarly, the smaller, mem-
branous posteri-or concavity is connected with the anterior margin of the second axillary
sclerite by a tendinous membrane. Such connections resist strong contraction when the
wings are closed.
Ventrally (Fig. 73C) the base of the C vein is isolated as a row of sclerites,
which are joined, by means of a robust tendinous ligament, with the posterior dorsal apex
of the anepisternum (anterior basalare). Another similar tendinous ligament connects
the posterior dorsal end of the basalare (posterior basalare) and the anterior margin of
the Sc vein base.
In Zonocerus the subcostal dorsal lip is less developed than in Acrididae, and does
not form an elbow. The connection of the bases of the C and Sc veins with the dorsal
part of the pleuron is shown in Fig. 73B.
The proximal and distal median plates are generally well developed in Acrididae
(Fig. 72A). The distal median plate is sometimes reduced to a short, thin sclerite, and
the proximal median plate is always more or less fused with the anterior margin of the
third axillary sclerite. In Tettigoniidae the proximal and distal plates are often mem-
branous or semimembranous, and are not clearly distinguished; when the wing is closed
the two plates fold up on each other along the line that separates them.
The metanotum: The metanotum is usually larger than the mesonotum, because it
bears the principal flight organs, the hind wing. As seen in Fig. 74A-C, the metanotal
surface is similar to the mesonotal surface in most Orthoptera; the postnotum is, however,
always present and IS often well developed. The second phragma, arising only from the
metanotal antecosta, is single walled. It is continuous anteriorly with the scutellar
inflection, and the third phragma arises posteriorly from the area where the postnotum
meets the first abdominal tergite. The lateral extension of the postnotum reaches the
epimeral area and forms the postalar bridge.
The function of the metanotal gland beneath the tegumina in Oecanthinae is
associated with copulation; Walker and Gurney (1967) discussed this structure from the
taxonomic viewpoint.
The hind wing base: Although not described by LaGreca (l947b), the hind wing
base in Metrioptera sepium (Fig. 73D) exhibits a similar arrangement of the axillary
sclerites to that in the forewing base. The first axillary sclerite is united with the base of
'
8
1 t is not immediately clear whether the distal median plate contains proximally the base of the R vein
(basiradiale), as in Blattaria, Mantodea, and other insects.
PRESCUTOSCUTAL
SUTURE
PRESCUTOSCUT AL
SUTURE
POSTERIOR NOTAL
PROCESS
PHRAGMA
SCUTOSCUTELLAR
SUTURE
POSTNOTUM
A
8
c
MUSCLE DISC.
ANTE COSTA
ACROTERGITE
POSTEROLATERAL
SCUTAL SUTURE
ANTE COSTA
PRESCUTOSCUTAL
SUTURE
. TERGAL FISSURE
4TH AXILLARY SCL.
SUTURE
179
FIG. 74. A, metanotum in Tylopsis /iliifolia (from Onesto); B, metanotum in Platycleis
intermedia (from Onesto); C, metanotum in Dissosteira carolina (from Snodgrass).
. 180
the Sc vein and is connected with the second axillary sclerite; it is also connected with
the third axillary sclerite by the tendinous ligament. The second axillary sclerite is con-
nected with the base of the R vein, as in other insects, and it is also associated distally
with the proximal median plate and posteriorly with the third axillary sclerite. The third
axillary sclerite articulates proximally with the fourth axillary sclerite (posterior notal
wing process), and is indistinguishably fused with the proximal median plate. The ill-
defined distal median plate is definitely associated with M and Cu veins.
The meso- and meta-pleuron: The pleural suture is well marked in the mesopleuron
and the metapleuron; the pleural arm arises from near the ventral end of the pleural
ridge. A complete paracoxal suture has not been found in Orthoptera.
In the mesothorax of Gryllidae (Fig. 75A) and Tettigoniidae (Fig. 76A, B), the
anapleural cleft clearly separates the preepisternum from the anepisternum; the pre-
episternum is separated from the basisternum by the true pleurosternal suture or cleft.
In the metathorax of these families the anapleural cleft is absent; in Tettigoniidae (Figs.
75B, 76A) the preepisternum is absent, due to membranization.
In Acridoidea the definitive pleuron is either a simple, elongate plate (e.g., Mare Ilia,
Carbonell 1959), or it may be an elongate plate provided with secondary sutures. What
Snodgrass (1929a), Misra (1947), Albrecht (1953), Carbonell ( 1959), and Yu and
Luh (1964) called the pleurosternal suture is the external manifestation of an internal
ridge, which serves for the attachment of muscles, t-p 5, 6. It is therefore quite
uncertain, as will be discussed further below, whether it represents a modified pleura-
sternal suture.
In the mesothorax of Schistocerca (Fig. 77A) the true pleurosternal suture is
probably what Misra (1947) called the basisternal suture, since its position agrees
approximately with the lateral margin of the basisternum in Platycleis (Fig. 76B). In
Schistocerca the pleurosternal suture of Misra (1947) or the secondary internal
strengthening ridge lies clearly distal (or dorsal) to this true pleurosternal suture in the .
mesothorax and metathorax. In the mesothorax of Dissosteira (Fig. 77B) the pleura-
sternal suture, comparable with that in Schistocerca, is evidently absent, and what
Snodgrass ( 1929a) called the pleurosternal suture corresponds to the secondary suture
(ridge) in Schistocerca.
In the metathorax of Acridoidea thus far studied, the pleurosternal suture is absent.
The sutures in the metathorax that have been called the pleurosternal sutures lie generally
even more distad than the counterparts in the mesothorax, and they are also clearly
secondary sutures that have developed for the attachment of the dorsoventral muscles.
It is possible that these ventral secondary sutures sometimes represent the modified
anapleural sutures, since they agree in position roughly with the anapleural cleft in
Tettigoniidae (Fig. 76A, B).
The trochantin is present in both the mesothorax and the metathorax of many
Orthoptera. Typically, an isolated trochantin lies in the membrane above the coxal
margin.
The epimeron is simple and undivided superficially. However, in the metathorax of
Tylopsis Onesto (1963) showed that an oblique internal ridge on the epimeron is con-
tinuous with the postnatal lateral extension, and the position of this internal ridge agrees
with the transepimeral suture (ridge) in some holometabolous insects. The meso-
thoracic spiracle lies at the ventrolateral angle of the epimeron in Tettigoniidae, Acrididae,
and many other families of Orthoptera. The unusual position and the structural details
of this spiracle in some Orthoptera were discussed by Carpentier (1924, 1925) . The
subalare is usually a small plate above the dorsal margin of the epimeron.
The meso- and meta-sternum: A more generalized condition of the pterothoracic
sterna, in which the mesosternum and the metasternum are clearly separated, is seen in
Tettigoniidae studied by Onesto ( 1963). In the mesosternum of Platycleis the sterna-
costa connecting the furcal bases is obvious externally, and internally it is ridged (Fig.
7 6B) . The anteriorly converging sutures from the furcal bases (the sterna costal sutures
of Onesto) form a triangular space, which is somewhat similar to the sternellum in
PLEURAL RIDGE
AN EPISTERNUM
ANAPLEUR AL C ~ L ~ E F;;T:-----_j
EPIMERON
PREEPISTERNUM
BASE OF SUBCOSTA
EPISTERNAL
LONGITUDINAL SUTURE-+'"+-----\
PLEURAL ARM
ANA PLEURAL CLEFT - - - r ~ ~
B
I
I
I
I
PLEURAL ARM
PLEUROSTERNALSUTURE
{CLEFT)
I
I
I
I
I
I
I
I
.. ..
I . . .
: L-+.:.,:..,:..:f----PLEURAL
I SUTURE
I
I
I
EPIMERON
181
Fro. 75. A, pleuron and sternum in mesothorax of Acheta (= Gryllus) pennsylvanica
(original); B, metapleuron in P/atycleis intermedia (from Onesto).
. 182
PLEURAL
STERNACOSTA
FU
FURCASTERNUM
l'ST ABDOMINAL
STERNUM
MARGINAL APODEME
FURCA

SPINA STERNUM ?
PLEURAL PROCESS-
TROCHANTIN-
A
EEPISTERNUM
+ BASISTERNUM
EPIMERON
COXA
PRESTERNUM
BASISTERNUM
STERNACOSTA
ANAPLEURAL
CLEFT
,---- ANTEPISTERNUM
FURCA
FIG. 76. Thoracic sternum in Platycleis intermedia (from Onesto). A, ventral view; B,
internal view.
Blattaria, Isoptera, etc. This triangular space is the Apophysengrube of Ander ( 1939).
The second spina is peculiarly continuous from the sternacosta, which connects the furcal
bases {Fig. 76B), and this condition is true also of Gryllidae (personal observation).
Evidently, therefore, the furcasternum and the spinasternum are so reduced that they are
represented only by the furcae, the sternacosta, and the anteriorly displaced spina. The
reduction of the posterior part of the mesosternum is thus correlated with the loss of the
postnotum dorsally. This reduction in the posterior part of the mesothorax and the
concomitant shift forward of the anterior part of the metathorax led to ever increased
consolidation of the two pterothoracic segments in many Orthoptera.
The large posterior triangular plate immediately behind the spina in Platycleis
(spinasternum of Onesto) is thus probably the metathoracic presternum. In the meta-
sternum of Tylopsis, this plate is completely isolated from the mesosternum and from the
rest of the metasternum. Onesto, therefore, called the plate the intersternite, but it is
clearly homologous with the metathoracic presternum in Platycleis. In Amblycorypha,
studied by Kramer (1944), the presternal area is continuous with the basisternum.
In the metasternum the formation of the posterior triangular space ( sternello of
Onesto) arising from the furcal bases is evident both in Platycleis and in Tylopsis. The
furcal pits represent the posterior end of the metasternum, which is followed by the first
abdominal sternite.
Turning now to the more specialized pterothoracic sterna in Acrididae (Fig. 77 A,
B), it is immediately evident that the membranous area surrounding the coxal base and
the sternum in Tettigoniidae has become completely sclerotized, the presternum is not
recognizable, and a secondary ridge from the furcal base extends posterolaterally to meet
the inner posterior angle of the coxal base.
In the mesosternum of Dissosteira (Fig. 77B) the posterolateral area Snodgrass
(1929a) called the sternellum corresponds to the membranous area between the meta-
thoracic presternum and the mesothoracic coxa in Platycleis (Fig. 76A, B). Clearly,
therefore, the ridge demarcating this area is a secondary formation for the firmer attach-
ment of a sternocoxal muscle (s-ex 5). Further, the median area behind the line
connecting the furcal pits corresponds to the metathoracic presternum in Tettigoniidae,
although its posterior margin has been lost. In the first-stage nymph of Dociostaurus, as
Jannone (1939) showed, the posterior margin of the metathoracic presternum is con-
tinuous laterally with the secondary ridge that demarcates the sternellum of Snodgrass.
In all adult acridids that have been studied, however, this suture is absent, although it may
be present in Chrotogonus and Lathicerus, judging from the figures given by Uvarov
(1966).
The term sternellum used by Snodgrass, and Albrecht ( 1953), is confusing, since
the term is sometimes applied to the median longitudinal area of the basisternum in
Blattaria and related orders. Misra (1947) called this area in Schistocerca the urea-
sternal lobe. However, the area is clearly not a part of the furcasternum, as is obvious
from the foregoing discussion. The area corresponds in position to the ventropleurite;
however, it is a secondarily produced area, and hence is not homologous with the primi-
tive ventropleurite, which is absent in other more primitive Orthoptera.
Compared with Platycleis (Fig. 76A, B), the metafurcal pits obviously lie more
cephalad in Schistocerca (Fig. 77A) and other acridids. Evidently, the shift forward of
the metasternum has proceeded farther in Acrididae than in Tettigoniidae. The plate
immediately behind the sternacosta of the metasternum in Locusta, Dissosteira, and
Schistocerca (Fig. 77A) is the first abdominal sternite, homologous with the first
abdominal sternum occurring in the corresponding position of Platycleis (Fig. 76A, B).
Snodgrass thought that this area was an extension of the first abdominal sternite and
called it the precosta; Arora and Asket Singh (1958) called it the acrosternite of
the first abdominal segment. Misra (1947) called it the anterior first abdominal basi-
sternum, and Albrecht ( 1953) called it the spinasternite. In Marellia, which Carbonell
( 1959) studied, such a forward extension of the first abdominal sternum is extremely
'n'fi:tere is also a good possibility that this plate is a secondarily detached anterior part of the metathoracic
bas1stemum.
PLEUROSTERNAL
SECONDARY
EPISTERNAL SUTURE------,.-4
SECONDARY
VENTROPLE URITE -----,-;;F--"'T.;
SECONDARY
EPISTERNAL SUTURE
SUTURE
SECONDARY RIDGE
I
ATKORACIC BASISTERNUM
l'ST ABDOMINAL STERNUM
ANEPISTERNUM
INFRACOXAL
LOBE of SNODGRASS
AN EPISTERNUM---II:,__ __
PLEURAL
ARM---JJ....:
SECONDARY
VENTRO-
PLEURITE
STERNACOSTA
A
B
SECONDARY RIDGE
SECONDARY
SUTURE
FURCA
FIG. 77. A, ventral view of pterothorax in Schistocerca gregaria (from Misra); B, internal
view of mesosternum in Dissosteira carolina (from Snodgrass).
narrow, being represented by a very narrow median longitudinal space. The lateral
triangular plate along the inner margin of the coxa (infracoxal lobe of Snodgrass) is
most probably also a secondarily formed area that cannot be homologized with the pleural
or sternal part in more generalized insects. In Marellia no such secondary formation
is present.
Uvarov (1922) used the width of the metathoracic presternum to distinguish the
two forms of Hierog/yphus banian. Uvarov ( 1923) also used the meso- and meta-sternal
characters to segregate two closely related species of Schistocerca. Roonwal (1946)
used the width of the anterior first abdominal sternum to segregate solitary and
gregarious phases of Schistocerca gregaria. While these authors studied sternal struc-
tures at the interspecific and intraspecific levels, considerable variations on these structures
appear to provide useful taxonomic characters, also at the levels of higher taxo-
nomic units.
Secondary sternal sutures of different natures are present in some other Orthoptera.
In Cylindrochaeta (Snodgrass 1929a) and Cylindroryctes (Carpentier 1936) the meso-
sternum is separated into anterior and posterior parts by a secondary transverse suture;
the mesosternum in Gryllotalpa is very much narrowed by the approximation of the
middle coxal bases. Ander (1939) traced the evolutionary development of what he
called the Mediansklerit throughout Ensifera. His Mediansklerit is the spina in certain
cases, and it is sometimes the metathoracic presternum (e.g., Lezina). As he showed,
the enlargement of the spina, associated with compression of the groove between furcal
bases, appears to be true at least in Rhaphidophorini and Ceuthophilini (Hubbell 1936,
cf. Ander 1939).
The anatomical basis of flight: Besides the anatomical details involved with
extension and flexion of wings already explained (p. 176), La Greca (194 7 b) further
explained the mechanism of extension and flexion of wings, as follows.
In extending the forewing, the costa-subcostal region is the only area that moves
actively. The pronator-extensor muscles (p 3, p-ti 2) act on the C-Sc through a
ligamentous connection and on basicostales (Fig. 73C). In this movement, the Sc vein
serves as a pivot (Fig. 78) on the point of articulation between the distal median plate
and the second axillary sclerite. The point of application of force lies on the anterior
ventral margin of the Sc vein base, where the tendinous connection lies with the pleural
wing process (Fig. 73C). This point lies very near the pleural fulcrum (process), and
the Sc vein functions as a lever on the horizontal plane. The pleural flexor ( t-p 14)
inserted on the third axillary sclerite is directly responsible for the flexion of the forewing.
For the folding mechanism see La Greca (194 7 b).
In some Orthoptera the posterolateral angle of the pronotum covers the base of the
forewing. For extension and flexion of the wing, this part of the pronotum must be
lifted. According to La Greca ( !947a), this is performed by a pleurosternal muscle
(probably p 3) and a pair of intersegmental ventral muscles. La Greca (1948) also
described the sensory and motor nerves of the flight organs in Acrididae.
Wilson (1962), in his experimental work on the flight muscles in Melanoplus and
Romalea, showed that some of the muscles function both as flight and leg muscles:
t 14, depressor of wing; t-p 5, elevator of wing; t-p 6, elevator of wing; t-ti 2, elevator of
wing and promotor of coxa; t-cx 7, elevator of wing and remotor of coxa; t-cx 6, elevator
of wing and remotor of coxa; p 3, pronator depressor of wing; p-ti 3, pronator depressor
of wing and promotor of coxa; t-cx 8, supinator depressor of wing and remotor of coxa;
t-tr 1, elevator of wing and depressor of trochanter.
The adult musculature: Studies of the adult thoracic musculature in Orthoptera
that have been made in this century include Voss (1905) in Gry/lus domesticus, DuPorte
(1920) in Gryllus assimilis, Carpentier (1923) on the dorsoventral muscles in Curti/la
gryllotalpa and Tachycines asynamorus, Snodgrass (1929a) in Dissosteira carolina,
Gunther and Deckert (1933) on leg muscles in Orthoptera, LaGreca (1938) in Gryllo-
talpa gryllotalpa, Maki ( 1938) in Locust a migratoria manilensis, Atractomorpha
ambigua, Xiphidion maculatum, and Brachytrupes portentosus, Jannone (1939) in
Dociostaurus maroccanus, Misra (1946, 1947) in Schistocerca gregaria, La Greca
(1947b) on some of the dorsoventral muscles in Acrididae and Tettigoniidae, Thomas
(1952) in Lamarckiana sp., Albrecht (1953) in Locusta migratoria migratorioides, Ewer
(1953, 1954c) on the thoracic muscular and nerv:ous systems in Acanthacris ruficornis;
Ewer (1954a) on some muscles in Zonocerus elegans; Ewer ( 1954b) on the nymphal
muscles in certain acridids, Ewer (1955) on prothoracic muscles in certain acridids.
Rakshpal (1955) in Gryllota/pa africana, Wiesend (1957) in Locusta migratoria migra-
torioides, Ewer ( 1958} in Lentula callani, Campbell (1961) on the muscular and
nervous systems in Locusta migratoria migratorioides, Alicata ( 1962a) on the muscular
and nervous systems in Eyprepocnemis plorans, Alicata (1962b) on the cervical muscles
in Orthoptera, Wilson ( 1962) on flight muscles in M elanoplus differentia/is and Romalea
microptera, Ewer (1964) in Bullacris longicornis and Pneumora inanis, Yu and Luh
(1964) in Locusta migratoria manilensis, and Blackith and Blackith ( 1967) on some
muscles in Morabinae.
. 186
HUMERAL PLATE
DISTAL
MEDIAN PLATE
PIVOT
NECK of l'ST
AXILLARY
SCLERITE
Z'ND AXILLARY
SCLERITE
ANTEMEDIAN
NOTAL PROCESS
PROXIMAL
MEDIAN
PLATE
3'RD AXILLARY
~ C L E R I T E
FIG. 78. Forewing base in Anacridium aegyptium (from La Greca).
Of these, the works on Gryllus by Voss ( 1905), on Dissosteira by Snodgrass
( 1929a), and on Xiphidion by Maki ( 1938) are shown in Table VIII, since they repre-
sent the major groups of Orthoptera and since the muscles are clearly described in
these studies. The sternocoxal muscles in Xiphidion are not included here.
The thoracic musculature in Orthoptera has remained relatively primitive, and some
important flight muscles (such as t-p 6, p 3) are rather poorly developed. In higher
Orthoptera, such as Dissosteira, tergopleural muscles have undergone considerable reduc-
tion in number. Noteworthy also is the obvious shift in points of origin of certain leg
muscles (s-ex 4, 5), which still maintain the primitive points of insertion on the
coxal margin.
Gross aspects of the postembryonic muscle development: In the studies by Ewer
(1954b) and Wiesend (1957) on the development of thoracic muscles in Orthoptera, all
or nearly all nymphal muscles develop continuously until the adult stage is reached. In
Gryllus, Voss (1912) showed a remarkable similarity between the nymphal and the adult
metathoracic muscles, only a small sternal muscle in the nymph being lost in the adult.
No new muscle is formed during the postembryonic development in Locusta migratoria
migratorioides (Wiesend 1957) and this is probably true also of other species, although
in Locusta the flight muscles increase in size very considerably after the third stage.
As Ewer (1954b), Thomas (1954b), and Wiesend (1957) have shown, some of the
muscles in the last nymphal stage or the newly emerged adult degenerate as the adult
matures. Such degenerating muscles in Acanthacris, Phymateus, and Locusta included:
mesothoracic and metathoracic phragma-pleural muscles ( t-p 3), mesothoracic and meta-
thoracic accessory tergopleural muscles ( t-p 8 or 9) , mesothoracic and metathoracic
anterior tergopleural muscles (t-p 4), mesothoracic and metathoracic accessory posterior
tergopleural muscles ( tp 15), meso thoracic pleuro-subalar muscle ( t-p 16), meta-
thoracic oblique dorsal muscles (t 12, 13), mesothoracic lateral oblique muscle (t-s 1),
and mesothoracic spina-pleural muscle (p-s 6).
MATSUDA: TIIE INSECT TIIORAX
187
Of these degenerating muscles in Acrididae, t-p 3, t-p 4, t-p 16, t-s 1, and p-s 6 are
present in the adult of Tettigoniidae or Gryllidae, or both (Table VIII). t-p 8 or 9 and
t-p 15 are present in other orders.
The nymphal muscles that are lost in the mature adult in Locusta migratoria migra-
torioides (Thomas 1954b) include: 1 t.p. (t-p 17); 2 t.p. (t-p 15?); 3 t.p. (t-p 15);
4 t.p. (t-p 4 or 11). In the same subspecies of the same species, Wiesend (1957) found
the following postimaginal degenerating muscles: II dlm 3 (t 12); II, III pm 4 (t-p 3);
II, III pm 9 (t-p 8); II, III pm 10 (t-p 4); II, III ism (p-s 3); II ism (p-s 6); II,
III bm 7 (s-ex 1); III dim 2, 3 (t 12, 13); III vlm (s 16).
Of these degenerating muscles in Locusta migratoria migratorioides, s-ex 1 is present
in other Acrididae (Ewer 1954b); t-p 3, t-p 4, and p-s 6 are present in the adult of
Tettigoniidae and Gryllidae; p-s 3 and s 16 are present in other species of Acrididae;
t-p 8 or 9, t-p 11, t-p 15, and t-p 17 are present in other orders.
The above facts show that generally the nymphal musculature is more primitive
than the mature adult musculature, since the degenerating muscles are often retained in
the adult of more primitive groups."' Thus, specialization of the thoracic musculature
in Orthoptera has involved mainly the loss of muscles.
Ewer (1954a) found that in the brachypterous form of Zonocerus elegans t 14,
t-p 5, 6, and p 3 are reduced in size. These muscles in wingless females of Lamarckiana
sp. are never more than strands (Thomas 1954a), and the indirect flight muscles are
completely absent in apterous Lentula callani (Ewer 1958) and apterous Morabines
(Blackith and Blackith 1967).
Thomas (1953) found that in the brachypterous female of Chorthippus parallelus
the flight muscles were poorly developed in the third and fourth stages. She suggested
that factors determining the brachypterous or macropterous condition of the adult
probably operate before or during the second nymphal stage. Atzinger (1952) also
studied the correlation between development of flight muscles and degrees of wing
development in 14 species of Orthoptera.
The postembryonic development and histology of flight muscles: As Tiegs (1955)
showed, the development of flight muscles and of some .other muscles in Gryllidae,
Tettigoniidae, and Acrididae is by enlargement of fibers in the earlier stages of nymphal
development. Later, the proliferation of fibers proceeds by cleavage. In earlier stages
the muscle fibers are of the tubular type, but in the adult they are of the close-packed
type. Chudakova and Messner-Bocharova ( 1965) also showed, in Acheta domestica,
the histological change from the tubular to the close-packed type during postembryonic
development.
According to Tiegs ( 1955), powerful flight muscles arise from rudimentary muscle
fibers. In Chortoicetes (Acrididae), for example, t 14 develops from six rudimentary
muscle fibers that are present in the first stage nymph, and in the adult t 14 comprises up
to 1300 fibers.
THE DERMAPTERA
The neck region: In Allostethus (Fig. 79A) the two-part lateral cervical sclerite
extends forward to articulate with the occipital ridge; on its way forward, the sclerite
passes the area immediately proximal to the reduced preepisternum. Hence, there is no
way of confusing the lateral cervical with the preepisternum, which becomes the lateral
cervical sclerite in many holometabolous insects; the lateral cervical sclerite in
Allostethus represents the lateral remnant of the presternum, as will be discussed
shortly (p. 190). In Echinosoma also, as Crampton ( 1926a) and Giles ( 1963) showed,
the association or union of the lateral cervical sclerite, along a suture, with the pre-
episternal plate is evident. In Dermaptera, therefore, the lateral cervical sclerite is
511
The possible postimaginal degeneration of thoracic muscles in more primitive groups of Orthoptera has not
been studied. Presumably, the postimaginal degeneration of muscles is less frequent in primitive groups with
soft body surface, since the degenerating muscles in Acrididae (with hardened skeleton) can still be functional
in the adult of the primitive groups.
. 188
TABLE VIII
Thoracic musculature in Orthoptera
Gryllus Dissosteira Xiphidion Muscle
(Voss 1905) (Snodgrass 1929a) (Maki 1938) designations Remarks
0 dvm d 7 op-cv 1
0 dvm c 50 9 op-cv 2
0 dvm b 51 10 op-cv 3
I dim la 56? 2 cv(d)-t 1
47 4 Op-t 1
0 dim 1 48 5 op-t 2
0 dim 2 49 1 op-t 3
0 ivdm 1 57 op-p 1 ? 1
0 ivdm 2 op-p 1
0 vim Sa op-p 2
0 ism t-cv 1
0 ism 2 53 11, 12 t-cv 2
52b t-cv 3
0 dvm a, a' 52a t-s(cv) 1
0 vim 3 cv-s 1
54 cv-s 2(x)
0 vlm 2 16 cv-cx l(x)
0 vim 4, 5 55 6 s 1, 2
I dim 3a, b t 12
II dim 4
III dim 3, 4
3 t 13
II dim 3 82 30
III dim la
I dim la 49, 56 1, 2 t 14
II dlm 1, 2 81 29
III dlm 1 112 58
58 (I) t-p 2
III pm 8 t-p 3? 2
II pm 10 t-p 3
III pm 10
II pm 11 37 t-p 4
III pm 11 62
II dvm 7 83 36 t-p 5
84 (I I)
113 (III)
t-p 6
II pm 9 t-p 10
III pm 9
38 (II) t-p 12
I pm 12?
t-p 14 3
II pm 12, 13 85 39
III pm 12 114 63
II pm 7
t-p 16
III pm 7 64
I ism 13 t-s 1
II ism
III ism
II dvm 6 t-ti(cx) 1
III dvm 6
I dvm 1 62 15 t-ti(cx) 2
II dvm 1 89 41
III dvm 1 118 66
TABLE VIII (Continued)
Gryllus Dissosteira Xiphidion Muscle
III dvm 4 t-cx 5
I dvm 7 64 18 t-cx 6
91 45
III dvm 3 120 70
I dvm 2 65 19 t-cx 7
II dvm 2 90 46
III dvm 2 119 71
I pm 6 63 20 (?) t-cx 8
II pm 6 99 47
III pm 6 129 72
II pm 5 de p 1
II pm 14 97
127 (III)
p 3
I Zm 14 p-s 1
II Zm 86 40
III Zm 115 65
34 (II) p-s 2 4
61 (III)
59 (I-II) p-s 3
II ifm p-s 6
II pm 5b 68a (I) 94, 95 p-ti 1
III pm Sb 125
II pm 1 42 p-ti 2
III pm 1 67
II pm 2 98 p-ti 3
III pm 2 128
I pm 4 68b 24? p-cx 4 5
II pm 4 96? 53
III pm 4 126? 78
I pm 5 23 p-cx5 5
II pm Sa 96? 51, 52
III pm Sa 126? 76, 77
I vlm 1 88 32 sll
I vlm 3 87 33 s 12
II vlm 3 117 60
I v\m 4, 5 60 (I-II) 31 s 13
II vlm 4, 5 116 (II-III) 59
I vlm 7 61 8 s 14
35 (II)
I vlm 6 s 15
Ill vlm 2-5 s 20
I vlm 2 s-ex 1
II vlm 2
II bm 5 101 s-ex 2
III bm 5 122
Ibm 2 66 s-ex 3
II bm 2
III bm 2 123
Ibm 7 67 s-ex 4 6
II bm 7 93
124 (III)?
Ibm 1 s-ex 5 7
II bm 1 92
III bm 1 121
TABLE VIII (Continued)
Gryllus
Dissosteira Xiphidion Muscle
Ibm 3
69
s-ex 6
II bm 3, 6 100
III bm 3 130
I dvm 5, 8 71d 25 t tr 1
II dvm 5
103 (pars) 54
III dvm 5
133 (pars) 79
1 pm 6a ? 71c 27 p tr 1
I pm 4a 71b 26 p tr 2
II pm 3
55
III pm 3
80
Ibm 4
28 s tr 1
II bm 4
103 (pars) 56
III bm 4 133 (pars) 81
REMARKS (Table VIII)
( 1) These muscles are inserted anteriorly on the neck membrane in Gryllus and
Dissosteira.
(2) III pm 8 in Gryllus is inserted on the episternum, ventral to the insertion of
III pm 10 (t-p 3).
(3) Points of origin on the pleuron of 39 and 63 in Xiphidion are not clear in
Maki's description.
( 4) Points of origin of these muscles in Xiphidion are the anterior ventral corner
of the episternum. They may possibly represent modified p-s 6 inserted on the meso-
sternal body, which is evaginated from the second spina (Thomas 1962).
( 5) 96 and 126 in Dissosteira represent either p-cx 4 or p-cx 5.
( 6) 124 in Dissosteira arises from the furcal base, probably owing to the loss of
the metathoracic spina.
(7) 92 and 121 in Dissosteira arise from the secondary ridge formed in the area
posterolateral to the furcal base. This is obviously due to the shift in point of origin
that accompanied the shift forward of the mesosternum and the metasternum.
Besides the muscles listed above, some of the muscles in Gryllotalpa (La Greca
1938) are apparently so specialized that they cannot be homologized with the muscles in
lower insects. Such muscles include IU-SF, which is a pleurocervical muscle;
TR-EM ' "
111
, which may be either p-tr 2 or p-tr 3; etc. In Lamarckiana, according to
Thomas ( 1952), a transverse tergal muscle (t 16) occurs; in Acanthacris (Ewer 1954b)
s 16 is present.
presternal in origin. The posterior end of the lateral cervical sclerite, which may be
fragmented, is often connected with the anterolateral angle of the basisternum.
On the ventral side of the neck between the lateral cervical sclerites, there are two
or three transverse ventral plates in Dermaptera. The hindmost plate immediately
anterior to the basisternum is, as Crampton (1926a) maintained, the presternum,
although this plate has been called the "posterior ventral cervical sclerite" by recent
workers (Henson 1953; Popham 1959; Giles 1963; Denis 1965b; Davies 1966). As
seen in Fig. 79 A, this plate lies clearly in line with the lateral cervical sclerites, and the
three sclerites together correspond to the presternum, as in Capnia (p. 8). Thus, in Allo-
stethus (Fig. 79A) two anterior ventral sclerites are ventral cervical sclerites. In
Echinosoma (Giles 1963) and Forficula (Henson 1953) the definitive median presternal
plate is considerably enlarged, due apparently to fusion of the presternum with the
posterior one of the two ventral cervical sclerites. In fact, in Echinosoma and Forficula
VENTRAL CERVICAL
SC!-ERITE
SPINA
INFOLDING of
BASISTERNUM
BASISTERNUM
PREEPISTERNUM
-PRESTERNUM
FURCA
TROCHANTIN
ANAPLEURAL CLEFT
------T-'r--+--- BASISTERNUM
+ PREEPISTERNUM
A
PLEURAL ARM
PREEPISTERNUM .J
PARACOXAL
PLEURAL SUTURE ---/--i;-/1[1.
ANEPIMERON
PLEURAL
SUTURE

TROCHANTIN. KATEPIMERON
/

FURCA 0 ANEPIMERON
FIG. 79. A, ventral view of thorax in Allostethus aegyptius (original); B, lateral view of
neck region in Forficula auricularia, showing sensory areas (from Popham); C, lateral view of
prothorax in Allostethus indicus (from Pantel); D, lateral view of prothorax in Labidura
reparia (from Pantel).
there are two ventral sclerites, instead of three as in Allostethus (Fig. 79A); in Forficula
the line of fusion is present, as seen in fig. 1 B of Henson ( 1953).
In Forficula Popham ( 1959) showed that the ridged membrane between the anterior
ventral cervical sclerite and the posterior ventral sclerite (ventral cervical sclerite plus
presternum) allows considerable expansion and contractoin of the neck region. Popham
( 1960a) further showed, in F orficula, the distribution of setae on the neck region
(Fig. 79B); the setae are proprioceptive organs that enable the insect to perceive the
position of the head relative to the thorax.
The pronotum: The pronotum in Dermaptera is a simple plate and is often provided
with a median longitudinal ridge. The anterior margin has a well-defined depressed
area, which Popham (1959) called the pretergum; Giles (1963) regarded the area as
serially homologous with the acrotergite in the pterothorax. In Echinosoma (Giles
1963) the paired dorsal cervical sclerites lie immediately anterior to the supposed acro-
tergite. The lobate posterior part of the pronotum covers a large part of the mesonotum.
The prop/euron: In A/lostethus (Fig. 79C) and Labidura (Fig. 79D) an oblique
suture runs nearly parallel to the basal (distal) suture of the trochantin; it extends
posteriorly across the pleural suture as far as the posterior margin of the epimeron. The
fact that this suture (the fissure divisante of Pantel, 1917) is confluent with the anapleural
suture near the anterior end of the trochantin leads us to interpret that it is the paracoxal
suture that is complete, both on the episternum and on the epimeron. The propleuron in
A 1/ostethus is thus divided into the katapleuron which consists of the katepisternum, the
trochantin and the katepimeron, and the anapleuron which consists of the reduced pre-
episternum, the anepisternum and the anepimeron. In Echinosoma also a complete
paracoxal suture is present, as Pantel (1917) suspected and as the figures of Giles (1963)
clearly showed. The occurrence of this suture only in more primitive genera of
Dermaptera leads us to suspect that the paracoxal suture is indeed archaic. In Labidura
(Fig. 79D) the paracoxal suture separates the katepimeron from the anepimeron,
although it is incomplete on the episternum. Among the Dermaptera Pantel (1917)
studied, the paracoxal suture in Aniso/abis and Calocrania is even less complete than in
Labidura. In Crampton's (1926a) figures this suture is completely absent on the
epimeron of and of Hemimerus.
The pleural arm of Allostethus (Fig. 79C) lies near the middle of the pleural ridge,
approximately where the paracoxal suture passes the pleural suture externally. The
episternal part of the pleural arm is fused with the sternal furca. In Dermaptera the
external pit for invagination of the pleural arm is often evident.
The prosternum: In Dermaptera the anteriormost presternal plate is, as already
discussed, the presternum that is usually isolated from the basisternum by a narrow
membranous area. In A/lostethus (Fig. 79A) the narrow sternacosta connecting furcal
bases separates the furcasternum from the basisternum. The spina is clearly present in
A/lostethus (Fig. 79A), Forficula and Echinosoma (Giles 1963).
The mesonotum: In A/lostethus (Fig. 80) the anterior transverse plate is thinly
sclerotized and subvertically sunken anteriorly, and it serves as the anterior point of
attachment of the dorsal longitudinal muscle ( t 14). Since the plate lies anterior to the
prescutoscutal suture, it can be regarded as a composite plate in which the original border
(antecosta) between the prescutum and the acrotergite has become lost. A small sclerite
immediately lateral to this plate must be the detached prealar arm. In Forficula
(Fig. 81A) the corresponding plate is a narrow sclerite that adjoins the tegula. Onesto
( 1961) regarded this sclerite as a part of the tegula. However, this sclerite is connected
with the dorsal margin of the episternum, as the prealar arm in other orders. Despite
its association with the tegula, therefore, this sclerite is not homologous with the prealar
sclerite in other orders. In the mesothorax of Dermaptera muscle t-p 4, which is inserted
on the prealar sclerite or the subtegula in other orders, is inserted on the anterolateral
part of the mesonotum. The origin of this sclerite, whether prescutal or acrotergal,
cannot be determined with certainty.
ACROTERGITE+ -----------DETACHED
PRESCUTUM_ - PREALAR ARM
-TEGULA
SUTURE ( ---
\
' ANTEROLATERAL
PARAPSIDAL L--:-SCUTAL SUTURE
SUTURE
POSTERIOR
NOTAL PROCESS
SCUTELLUM
DETACHED
PREALAR ARM----
Fro. 80. Dorsal view of pterothorax in Allostethus aegyptius (original).
In Allostethus (Fig. 80) the prescutoscutal suture is almost continuous laterally
with the anterolateral scutal suture, and the parapsidal suture arises from the prescuto-
scutal suture. The parapsidal suture is apparently absent in most other genera of
Dermaptera.
In Allostethus (Fig. 80) the scutoscutellar suture is absent, and a simple median
projection occurs on the posterior margin. In Forficula, however, the median projection
is continuous with anteriorly converging sutures, which Henson ( 1953) homologized with
the scutoscutellar suture in Blatta. In Echinosoma also the corresponding projection and
the suture occur. In Forficula an oblique lateral suture, which extends laterally from the
scutoscutellar suture, appears to represent the tergal fissure. In fact, in another species
of Forficula (F. mikado) the suture clearly reaches the scuta! lateral margin behind the
anterior notal process, as is the case with the tergal fissure (suture). In Allostethus the
median desclerotized plate beneath the median projection extends laterally on each side
to be continuous with the axillary cord (Fig. 80). The mesothoracic postnotum is
absent in Dermaptera.
The mesonotal lateral margin and axillary sclerites: In F orficula auricularia, studied
by Onesto (1961), the anteriormost structure involved with the articulation of the fore-
wing is the tegula, which is adjacent to the detached prealar arm already discussed
(Fig. 81A). The anterolateral angle of the mesonotum is the anterior notal wing process;
it articulates with the detached prealar arm and with the neck of the first axillary sclerite.
An inconspicuous median or postmedian process articulates with the posteromedial angle
of the definitive first axillary sclerite. The posterior notal wing process is present behind
a deep excavation on the postmedian scuta! margin.
The definitive first axillary sclerite (Fig. 81A) is actually a fusion product of the first
and second axillary sclerites; the posterolateral angle of this composite sclerite, according
to Ones to ( 1961), is the second axillary sclerite that has become indistinguishably
incorporated; this area has direct articulation with the third axillary sclerite. As seen
from the illustrations of the axillary sclerites in Spongiphora (Snodgrass 1909), Forficula,
194
AN EPISTERNUM
DETACHED
PREALAR ARM
ANTERIOR
NOTAL PROCESS
POSTERIOR
NOTAL PROCESS
EL
BASICOSTALE +
BASI SUBCOST ALE
SUBALARE
PLEURAL SUTURE
BASICOSTALE
+ BASISUBCOSTALE
3'RO AXILLARY
SCLERITE

3'RD AXILLARY
SCLERITE
Fro. 81. Elytral basal structures in Forficula auricularia (from Onesto). A, dorsal view;
B, lateral view.
Psalis, and Labidura (Pantel 1917), the degree of fusion of the two axillary sclerites
appears to differ in various earwigs. The small ventral second axillary sclerite (Fig. 81B)
articulates with the area that corresponds to the pleural wing process by a tendinous
ligament; it is laterally fused with the ventral sclerotization of the basicostale and basi-
subcostale, and posteriorly associated with the subalare.
FIG. 82. A-C, hind wing holding mechanism in Forficula auricularia (from Eastop). A,
elytron with a ridge bearing modified hairs; B, modified hairs on metanotum; C, posterior end
of ridge bearing modified hairs on elytron; D, dorsal view of hind wing base in Forficula
auricularia (from Onesto); E, lateral view of hind wing base in Forficula auricularia (from
Onesto); F, dorsal view of elytral base in Forficula auricularia (from Onesto); G and H,
diagrams showing elastic hind wing base in Forficula auricularia (from Kleinow).
I
I
I
I
I
I
I
I
I
I
ELYTRON--
JUGUM
BASICOSTALE
195
Z'ND AXIL. SCL. +
PROX. MED. PLATE
BASANALE
l'ST AXIL. SCL.
---------BASALARE
....... .... i. PLEURAL WING
. . . PROCESS
G
3'RD AXIL. SCL.
GROOVE
F
H
1 AX. SCL.
'------ BASALARE
BASICOSTALE
BASI STERNUM
FIG. 83. Internal view of mesosternum in Forficula auricularia (from Hamon and Ovazza).
The third axillary sclerite in Forficula (Fig. SlA) articulates proximally with the
posterior notal wing process, and anterolaterally with the posterior process of the second
axillary sclerite; its posterior margin is fused with the anal field along a line; its proximal
process is connected with the base of the first axillary sclerite by a tendinous ligament,
and when the wing is closed both sclerites fit together. The ventral sclerotization of the
third axillary sclerite (Fig. SIB) consists of two pieces; the larger piece articulates with
the subalare (Fig. 81B).
The mesop/euron (Fig. SIB): The mesopleuron lacks a well-defined pleural wing
process; its division into the episternum and the epimeron is obscured dorsally, the pleural
ridge being increasingly weakly marked dorsally. The pleural arm arises from the
middle of the pleural ridge. The basal are is not differentiated; in F orficula Ones to
( 1961) named the dorsal tip of the episternum processo basa/are, which implied that the
basalare has become incorporated into this area. The episternal longitudinal suture is
distinct, reaching caudally the base of the trochantinal plate. This suture is characteristic
of the mesopleuron of Dermaptera, although it occurs also in Mantodea and Orthoptera.
In Allostethus the paracoxal suture is represented by the anteromesal margin of the
definitive trochantin or the eutrochantin (Fig. 79A); it meets the anapleural cleft at the
anteromesal angle of the trochantin. The caudal part of the preepisternum extends
proximally.
The mesosternum: The presternum in Echinosoma is demarcated posteriorly by a
transverse suture or ridge, and Giles ( 1963) denied any morphological significance of
this suture. However, since the presternum is present in the prothorax, there is no
positive reason for the anterior marginal area not to be serially homologous with the
prothoracic presternum. An actual observation of this area in Echinosoma has shown
that this marginal area is a little too broad to be a secondary strengthening apodeme
(prepectus), which often occurs in other insects. In other genera that have been studied,
this presumed presternum has become lost or fused with the basisternum.
The basisternum is lobately expanded and infolded laterally (Fig. 79A); Matsuda
( 1960a) mistook this infolded part of the basisternum for the ventropleurite. The
furca arises from the posterior part of the lateral folded part of the sternum in Al/ostethus
(Fig. 79A), Echinosoma, and probably most other Dermaptera. The division of the
sternal lobe into the basisternum and the furcasternum cannot be recognized externally.
Internally, however, the furcal bases are connected by a straight horizontal ridge, which
can be regarded as the sternacosta that separates the basisternum from the furcasternum.
The mesothoracic spina, as Hamon and Ovazza (194S) clearly showed in Forficula,
arises from the internally folded posterior part of the sternum (Fig. S3) and is therefore
invisible externally. In Echinosoma examined, the spina arises in the same way as in
Forficula.
The metanotum: In Allostethus (Fig. 80) the anterior marginal strip is, as in the
mesothorax, the prescutum plus the acrotergite. The small, paired phragmata are con-
tinuous medially from this prescutoacrotergal area, which lies beneath the mesothoracic
scutellum.
In Allostethus (Fig. 80) a narrow sclerite associated with the tegula is apparently
serially homologous with the detached prealar arm in the mesothorax. In Forficula
Kleinow (1966) recognized a small sclerite that lies lateral to the phragma, which he
called the Phragmasklerit. Judging from Kleinow's description and figure of this sclerite,
it apparently differs from the detached prealar sclerite in Allostethus."' According to
Klein ow (1966), muscle t-p 4 (his muscle 143) is attached to it; and the sclerite is
probably homologous with the prealar sclerite in other orders.
The metanotum behindthe prescutoscutal suture is largely the scutum. The scuto-
scutellar suture (Fig. 80) is not posteriorly continuous with the axillary cord, but is
delimited posteriorly by a membranous fold that reaches the third axillary sclerite
(Fig. 82D).
A peculiar wing-holding structure on the mesonotum lies along the scutoscutellar
suture (Fig. 80). In Labia, Forficu/a, and Labidura Eastop (1953) found that the hairs
supported on the ridges on either side of the median longitudinal axis of the metanotum
are double-rooted, thick and curved (Fig. 82B). On the undersurface of the anterior
end of each elytron, near the inner edge, is a thickened longitudinal ridge (Fig. 82A)
bearing about 50 downwardly pointing, thickened, curved hairs with sharp apices
(Fig. 82C). These two groups of hairs come into contact when the elytra are closed.
Hardas (1961) also described the corresponding structures in Labidura riparia.
The posteriorly separated scutellum (Fig. 82D) carries the sclerotized, posterior
metanotal arm, which extends posterolaterally, supporting the bases of the A veins. This
arm must be homologous with the axillary cord in other insects. A similar modification
of the scutellum has often occurred to the median
1
bilobed posterior extensions in
Dermaptera. The metathoracic postnotum is represented by a transverse plate beneath
the median scutellar extension (Fig. 80); it has no invagination to form the third
phragma. In Forficula the metathoracic dorsal longitudinal muscle (t 14) is attached
posteriorly to the posterior part of the postnotum, in front of the anterior edge of the
first abdominal tergite.
The metascutallateral margin and hind wing base structures: The metascutallateral
margin in Forficula (Fig. 82D) is, according to Onesto (1961), provided, from the front
backward, with the anterior, antemedian, median, postmedian, and posterior notal wing
processes; in Allostethus (Fig. 80) the processes are less marked, and hence the scuta!
lateral margin is less specialized.
The first axillary sclerite in Forficula (Fig. 82A) is connected distally by its neck
with the basicostale; its proximal margin articulates with the anterior, antemedian,
median, and postmedian notal processes; and its posterior margin is connected with the
third axillary sclerite by a tendinous ligament.
The definitive dorsal second axillary sclerite (Fig. 82D) is connected proximally
with the first axillary sclerite, and its anterior end continues to the basiradiale. The
dorsal posterolateral angle of the definitive second axillary sclerite represents, as Onesto
(1961) maintained, the proximal median plate that has become indistinguishably fused.
This fusion. is evident from the fact that this sclerite is nearly quadrangular in shape, and
its posterolateral angle articulates with the distal process of the third axillary sclerite, as
the proximal median plate usually does. The small ventral second axillary sclerite
(Fig. 82) is connected to the pleural wing process by a tendinous ligament; it is
posteriorly associated with the subalare, and is laterally, partly fused with the ventral
third axillary sclerite.
"'The difference needs to be confirmed by dissecting fresh material of AllostethUJI.
The dorsal third axillary sclerite is characteristic in having its .proximal process
separated from the third axillary sclerite proper by a membrane; according to Onesto,
this condition is primitive. The proximal process is connected anteriorly with the second
axillary sclerite and with the base of the first axillary sclerite by a broad tendinous liga.
ment; the caudal end of the third axillary sclerite articulates with the posterior notal
process and its distal margin with the basanale.
The distal median plate is, according to Onesto, fragmented into three plates: the
anteriormost plate is indistinguishably fused with the base of the R vein, the hindmost
large plate is associated proximally with the detached basanale, and distally with the
M -Cu vein complex.
The metathoracic tegula is associated with the basicostale (humeral plate) laterally,
and articulates posteriorly with the neck of the first axillary sclerite, with the basi-
subcostale, and with the basalare.
The basisubcostale is ventrally in contact with the basalare and the pleural wing
process by means of tendinous ligaments. The basanale lies alongside the bases of the
A veins.
The metapleuron: The metapleuron (Figs. 79A, 82E) is separated by the pleural
suture into the episternum and the relatively broad, undivided epimeron. In the
episternal area the paracoxal suture is lost in the area distal to the base of the katepisterno-
trochantin or the eutrochantin; the anapleural cleft is oblique or often nearly vertical,
reaching the anterior end of the katepisternotrochantinal plate. The episternal longi-
tudinal suture is absent in the metapleuron. In most Dermaptera the preepisternum has
probably become indistinguishably fused with the basisternum to form a large, definitive
basisternal plate (Fig. 79A). According to Pantel (1917), the preepisternum (latero-
sternite of Pantel) is present in Labidura riparia. According to the same author (1917),
the clearly separated antecoxale is present in Dip/atys and Forcipula, and it is in-
completely separated in Allostethus indicus and Labidura riparia.
In Forficula the dorsal part of the anepisternum (Fig. 82E) has an extensive basalar
cleft which surrounds the basalare. In many Dermaptera the basalar cleft also almost
separates off the anterior dorsal part of the anepisternum. Onesto ( 1961) called this
area in F orficula the second basal are.
In most Dermaptera the definitive trochantin appears to be a fusion product ( eutro-
chantin) of the trochantin and the katepisternal antecoxale. However, the Katopleure
of Verhoef! (1902a) in Echinosoma occidentale, which lies anterior to the presumed
eutrochantin, may represent the antecoxale demarcated by the paracoxal suture. In
most Dermaptera the paracoxal suture is absent.
The metasternum: As already discussed, the broad definitive sternal plate is due to
incorporation of the preepisternum into its anterolateral area; it is probably not the
marginal evagination of the sternum, as in the mesosterna! plate. The furcal pits are
recognizable externally in Forficu/a (Henson 1953) and Echinosoma (Giles 1963),
although they are absent in Allostethus (Fig. 79A). The well-developed furcae are
always present. The metathoracic spina is absent.
The anatomical basis of flight: According to Klein ow (1966), the hind wing is
tilted upwards by the action of an elastic skeletal mechanism, as follows: Both ends of
the basalare, the processes of the humeral plate (basicostale) and of the subcostale, form
a push-button apparatus (Fig. 82G, H). When the processes of the humeral plate and
of the basisubcostale clasp the heads of the basalare from below and above, the hind wing
rests on the body; when the wing tilts upwards, both heads of the basalare are disengaged
from the processes. In the resting position the head of the basisubcostale lies outside the
concavity on the head of the first axillary sclerite; when the wing is tilted the head of the
basisubcostale slides posteriorly to fit the concavity in the first axillary sclerite.
The muscles responsible for tilting the hind wings in F orficula are: the meta-
phragmo-mesophragmal muscle ( 131 of Kleinow, t 14), the metaphragmo-metanotal
muscle (132 of Kleinow, t 12); the metanoto-trochanteral muscle (139 of Kleinow,
t-tr 1), and the merocoxo-metanotal muscle (140 of Kleinow, t-cx 6 or 7) that moves
both tbe legs and the wings. By pulling t-tr 1 and t-cx 6 or 7, the wings will tilt; i.e. the
anterior corner of the wing is held in front of the basalare, and at the same time the
wings are elevated. As is usually the case, t 14 depresses the wings and t 12 is the
antagonist of t 14. In Labia a fifth muscle, the basisterno-metanotal muscle (t-p 5 or 6,
58 of Maki, 1938) functions exclusively as a depressor of the metanotum, and hence it is
important in elevating the wings.
The anepisternum (basalare 2 of Kleinow) and the muscles attached to it are im-
portant in flight. The muscles involved are the superior metaepisterno-basicoxal muscle
(151 of Kleinow, p-cx 4) and the metapleuro-trochanteral muscle ( 152 of Kleinow,
p-tr 2). The pressure produced by a contraction of these bifunctional muscles causes
pronation of the hind wings.
The first and second axillary sclerites together form a single cooperative shield.
The first axillary sclerite articulates with the scuta! margin and, above the second axillary
sclerite, forms a rigid lever with the anterior margin of the wing. The second axillary
sclerite lies on the pleural wing process, which is a fulcrum for the wing movement.
Therefore, when the first axillary sclerite is lowered by muscle action, the rest of the wing
rotates upward around the pleural wing process.
The main part of the third axillary sclerite lies on the axis of rotation, which
extends from the posterior notal wing process to the posterior end of the second axillary
sclerite. When the proximal process of the third axillary sclerite is pulled downward
and medially by a muscle (t-p 14), the force is transmitted to the third axillary sclerite
proper, then to the above-mentioned axis of rotation behind the posterior margin of the
first axillary sclerite. The hind wings are then placed on the back of the body.
The elytra are raised if the mesoscutum is pulled downward. The dorsal end of the
pleural ridge functions as a fulcrum for this movement, and a groove on the mesoscutum
(Fig. 82F), into which the elytron fits when at rest, serves as a hinge socket for erection
of the elytron; the muscles responsible for this movement are the mesonoto-trochanteral
muscle (1 04 of Kleinow, t-tr 1), the merocoxo-mesonotal muscle ( 105 of Kleinow,
t-cx 6 or 7), and the praecondylo-mesopleural muscJe (108 of Kleinow, t-p 4).
For the anterolateral swinging of the elytron, the jugal field and the third axillary
sclerite (fused with the jugal field, Fig. 82F) perform a special function as a hinge. In
the resting position these areas represent the anterior angle of the elytron, and the short
side of the triangular jugum connects the jugal field with the mesoscutum. When the
jugal field flaps sideward, the elytron also swings sideward. Two muscles responsible for
this movement are attached to the shorter side of this jugal field-scutum hinge; they are
the mesopleuro-mesopteral muscle ( 112 of Kleinow, t-p 14) that rotates the elytron
backward, and the merocoxo-mesepimeral muscle (122 of Kleinow, t-cx 8) that moves
the jugal field outward. Further information about the movement of wings is found in
Verhoef! (1917), Henson (1951), and Kleinow (1966).
The adult musculature: Maki (1938) studied the thoracic muscles in Anisolabis
annulipes and Labia curvicauda, and Kleinow (1966) studied those in Forficula auri-
cularia; they are shown in Table IX.
The characteristic of the dermapterous thoracic musculature is the poorly developed
flight muscles; op-cv(v) 4 and p 4 are peculiar to this order. According to Mercier and
Poisson (1923), Forficula auricularia with fully developed hind wings cannot fly, because
the flight muscles (t 14 and t-p 5 or 6) are atrophied. Popham (1959) discussed the
function of cervical muscles in the head movement in F orficula.
THE COLEOPTERA
The neck region: The lateral cervical sclerites are often present in Polyphaga,
although they are never present in Adephaga (Larsen 1966). In Chauliognathus
(Fig. 84) each lateral cervical sclerite consists of an anterior elongate plate and a
posterior shorter plate. The anterior plate articulates with the occipital margin of the
200
TABLE IX
Thoracic musculature in Dermaptera
Forficula Anisolabis Labia Muscle
(Kleinow 1966) (Maki 1938) (Maki 1938) designations Remarks
57 3 3 op-t 2
55 1 1 op-t 3
61 8, 9 8 op-cv 2, 3
60 10, 11 9, 10 t--cv I
64 14 14 t--cv 3? 1
op-p?
58a 6 6 op-cv(v) 4
66 5 5 cv-s1
58 4 4 s I or 2
71 2 2 t 12
96 26 26
132 54 54
53 (I II) t 13
55 1 1 t 14
95 25 25
131 52 53
106 (II) 33 34 t-p 3
142 (III) 59 60
108 (II) 34 t-p 4
143 (III) 60 61
58 (Ill) t-p 5 or 6
144 (I II) 62 (III) t-p 11
35 (I I) 35 t-p 12?
61 (III)
112(1!) 37 t-p 14
146 (Ill)
109 (II) 36 t-p 15
85 (I) 13 12, 13 t-p? 2
87
12 11 t-s 1
125
32 33
156 58 59
78 (I) 15 15 t-ti(cx) 1
102 (II) 38 39
64 (III) 64
79 (!) 20 20 t-ti(cx) 2
81 17, 18 17, 18 t-cx 6, 7
105
41 42
140
68 68
122 (II) 42 43 t--cx 8 3
153 (III) 69 69
150a (III) 63 63 p4
124 (II) 36
62 (III)
38 p-s 1
75a (I) 30 30, 31 p-s 6
57 (II)
75c (I-III) 29 29 p-s 7 4
57 (Illi p-s 9 5
65 (III) 65 p-cx 2 or 3
82 . 21 21 p-cx 4
117 46 47
151 73 73
Forjicula
(Kleinow 1966)
84 (I)
150 (III)
77 (I-II)
99 (II-III)
74 (I-II)
98 (II-III)
7 5 (I)
76 (I-II)
75b (I-II)
99a (II-II I)
80
104
139
86 (I)
121 (II)
152 (III)
92 (I)
160 (III)
REMARKs (Table IX)
TABLE IX (Continued)
Anisolabis Labia Muscle
(Maki 1938) (Maki 1938) designations Remarks
19 19 p-ex 5
45 (II) 46
72 72
s 12
56 56
27 27 s 13
55 55
7 7 s 14
31 (II) 32
28 28 s 15
39 40 s-ex 1
66 66
43 (II) 44 s-ex 3
70 (III) 70
16 16 s-ex 5
40 41
67 67
44 (II) 45 s-ex 6
71 (III) 71
22 22 t-tr 1
47 48
74 74
23 23 p-tr 1
48 49 p-tr 2
75 75
24 24 s-tr 1
49 (II) so
76 76
( 1) All muscles arise from the dorsal part of the proepimeron. They may deserve
a new designation under op-p, or they may be modified op-p 1.
(2) These muscles are prothoracic tergopleural muscles, which cannot be homolo-
gized with t-p muscles in the pterothorax.
(3) Muscles 42 and 69 in Anisolabis, in the absence of the subalare, originate from
the posterior lateral margin of the tergum.
( 4) The posterior points of attachment of these muscles are on the anterolateral
angle of the metasternum (presternum in A nisolabis).
(5) The anterior point of attachment of 57 in Anisolabis is on the lateral angle of
the presternum.
head; the posterior plate lies folded beneath the former, and its end directed towards the
median longitudinal axis. In the shape and the manner of folding of the lateral cervical
sclerite Coleoptera is therefore similar to lower hemimetabolous insects such as Blattaria,
Isoptera, Dermaptera, etc.
Crampton ( 1926a) gave figures of the lateral cervical sclerite in Rhipiphorus, Silpha,
Staphylinus, Hydrous, Meloe, Lampyris, and Atractocerus. In all of these genera, the
lateral cervical sclerite lies anterior to the precoxal bridge, as in Chauliognathus (Fig. 84)
and in lower hemimetabolous orders. In Coleoptera (Fig. 85A, B) the precoxal bridge
is clearly anterior to the trochantin, and the bridge is the preepisternum (see p. 203).
Therefore, the lateral cervical sclerite in Coleoptera must be presternal in origin, as in the
202
LATERAL
CERVICAL
SCLERITE ------......_
PREEPISTERNUM
EUTROCHANTIN
COXA
ANAPLEURAL
SUT
AN EPISTERNUM
EPI
ANAPLEURAL
CLEFT
KATEPIMERON
KATEPISTERNUM
BASISTERNUM
RCA
PREEPISTERNUM
BASISTERNAL
AREA
TROCHANTIN
2
3
PREEPISTERNUM
+ BASISTERNUM
IMAGINARY
PARACOXAL
SUTURE
FIG. 84. Ventral view of thorax in Chauliognathus pennsylvanicus (from Campau).
hemimetabolous orders. According to Larsen ( 1966), the lateral cervical sclerite is
greatly reduced in Ips, and it is represented by a single plate in Oedemera and
Necrophorus.
Although no ventral cervical sclerite has been found in Coleoptera, the development
of the gula in certain Coleoptera is quite pronounced (DuPorte 1960; Evans 1965). If
DuPorte's theory (1962) on the origin of the gula (p. 9) is valid, the ventral cervical
sclerite in the ancestral Coleoptera must have been incorporated into the gula. In
Sphaeridium and Cercyon, according to Larsen (1966), the dorsal cervical sclerites
consist of median unpaired and lateral paired plates.
The pronotum: The most conspicuous feature of the pronotum in Coleoptera is its
lateral expansion, which covers the dorsolateral part of the propleuron. The prothoracic
pleuron in Coleoptera is thus comparable with the cryptopleuron in Orthoptera, as
Carpentier ( 1929) contended. The inflected lateral part of the pronotum or the
epipieuron in Coleoptera is often separated from the dorsal part of the pronotum by a
more or less sharp lateral edge. In larval Coleoptera the lateral pronotum is sometimes
separated from the dorsal pronotum by a membrane, as Quennedey ( 1965) showed in
Sphaeridium. Quennedey, however, called the area the dorsopleural sclerite. In larval
Riolus the area Marvillet ( 1960) called the epimeron may also represent the lateral tergite.
This isolated. lateral plate in the larva probably becomes the lateral pronotum or the
epipleuron, which covers the dorsolateral part of the propleuron in the adult. In fact, as
Crampton (1926a) showed, the ventrally placed notal carinae (c and d of Crampton) in
adult beetles appears to correspond to the proximal and distal margins of this isolated
lateral tergite in the larva.
The anterior margin of the pronotum is inflected and surrounds the occiput of the
head. The posterior evagination is usually pressed against the base of the elytra; and
the posterolateral expansion of the pronotum is sometimes conspicuous (e.g., Elateridae).
The propleuron: In a primitive coleopteron, Chauliognathus pennsylvanicus
(Fig. 85A), the pleural suture is traceable ventrally as far as its caudal end, which
articulates with the lateral coxal margin. The episternal area proximal (ventral) to the
pleural suture is represented by a broad subtriangular plate, which is the katepisternum
that has become indistinguishably fused with the trochantin ( eutrochantin). Distal
(lateral) to the pleural suture and beneath the inflected pronotum lie the narrow
katepimeron and a larger, more anterior epimeral plate. The precoxal bridge, which has
a direct connection laterally with the inflected ventral edge of the pronotum, must be
a fusion product of the anepisternum and the preepisternum that reaches the anterolateral
angle of the sclerotized sternum (the basisternum), since it lies clearly anterior to the
eutrochantin; it agrees in position with the precoxal bridge in Blatt ella (Fig. 54 A). The
paracoxal suture may be represented by a short, oblique suture arising from the distal
anterior margin of the eutrochantin. The anapleural suture is absent.
Crampton (1926a), comparing Lampyris, Meloe, and others with mantids, pointed
out a tendency for the pronotum to extend ventrally and meet the base of the precoxal
bridge along a line x (Fig. 85A, B) and that the definitive trochantin is the eutrochantin
(discussed above). He emphasized that the Cantharoidea (Lampyrioidea of Crampton)
have retained the most primitive condition in the prothorax among the major groups of
Coleoptera. Similarly, Ferris (1935) pointed out the primitiveness of the prothorax in
Photuris (Lampyridae). The preceding discussion on the propleuron in Chauliognathus
supports the contention of Crampton and Ferris. The ventrolateral area of the pro-
thorax in other Coleoptera, then, must have been derived from a condition similar to that
seen in Chauliognathus. In the following discussion Gyrinus (Fig. 85B) is chosen as a
representative of the more derived groups.
In Gyrinus (Fig. 85B), as Larsen (1966) showed, the landmark x of Crampton,
which represents the inflected ventral edge of the pronotum, can be readily identified by
comparing it with x in Chauliognathus (Fig. 85A). In Gyrinus, x takes a sharp turn at
the middle, although in Chauliognathus the corresponding area extends straight back-
wards. In Gyrinus, at the same time, the eutrochantinal area has evidently shifted
caudolaterally, accompanying the shift in direction of the ventral edge of the pronotum
( x) . This shift in position of the propleuron has resulted in: the loss of the pleurocoxal
articulation, which is already relatively weak in Chauliognathus; a shift laterocephalad of
the eutrochantin; the displacement forward of the a11terior piece of the epimeron (pleural
apophysis of Ferris, cryptopleuron and the cryptopleural ridge of Larsen); and the shift
of the proximal end of the precoxal bridge from the basisternal lateral margin to the
lateral margin of the sternum behind the furcal base. This rotation of the pleuron
appears to be the only interpretation by which the differences between Chauliognathus
and Gyrinus can be accounted for. Whether this rotation can be traced during develop-
ment remains to be investigated.
As a corollary of -this interpretation, the sternopleural suture (or the anapleural
suture of Larsen, 1966) in Gyrinus is actually the anterior margin of the eutrochantin,
the epimeron of Larsen is largely katepisternal, and the invaginated part of the propleuron
corresponds to the epimeron. The transverse line beneath the posterolateral margin of
the inflected pronotum (Fig. 85B) may represent the displaced pleural suture.
Crampton (1926a) applied his idea of the ventrally expanded tergum to other
Coleoptera also. He believed that the definitive propleuron in Silpha, Hydrous, Dytiscus,
PARA COXAL
SUTURE?
PRECOXAL BRIDGE
CPREEPISTERNUM t'
KATEPISTERNUM
+ TROCHANTIN
( EUTROCHANTIN)
FURCA---7
B
EPIMERON
KATEPIMERON
.14.-:.,..----+---- PLEURAL
A
SUTURE
PLEUROCOXAL
ARTICULATION
EPIMERON
PLEURAL
SUTURE?
FIG. 85. A, lateral view of prothorax in Chauliognathus pennsylvanicus (original); B, lateral
view of prothorax in Gyrinus marinus (from Larsen).
Harpa/us, and others is at least largely the pronotum that has overgrown the propleuron,
and that the secondary sutures (his c, d, e) on the ventrolateral surface of the prothorax
were therefore tergal sutures. Crampton's line e lies very near the leg base. Ferris
( 1935) rejected Crampton's idea of the so far ventrally expanded pronotum; he thought
that in Dineutus the ventrally inflected pronotal part was narrow and the definitive
propleuron was the true propleuron. This idea of Ferris was accepted by Hinton
(1939), Campau (1940), Evans (1961), Larsen (1966) and Doyen (1966).
The pleurocoxal articulation in higher Coleoptera, as is clear from the discussion on
Gyrinus, is often the trochantinocoxal articulation. In higher Coleoptera the acetabulum
is formed through the development of pleural and sternal supracoxal lobes. The lobes
are separated by the coxal cleft. According to Larsen (1966), in Hydrous the cleft is
very long, and in Gyrinus the two lobes are fused. The formation of the acetabulum
allows only rotatory movements and a very slight abduction of the coxa.
The prosternum: In Chauliognathus (Fig. 84) the presternum consists of the short
basisternum, which is continuous with the furcasternum bearing simple, paired furcae.
In Gyri nus and Carabus (Larsen 1966) the basisternal area is almost completely absent
and the precoxal bridge reaches the base of each furca. In the prothorax the furcal
bases are separated and the furca has no association with the pleural arm, which is absent.
The prothoracic spina has been found in several Adephaga and Meloidae (Larsen 1966),
in the larvae of Tenebrio (Josting 1942), Riolus (Marvillet 1960), Peltodytes (J aboulet
1960) and Sphaeridium (Quennedey 1965).
The mesonotum: In Coleoptera the mesonotum is usually much smaller than the
metanotum, both in width and length. Its anterior region is subvertically inflected. In
Creophilus (Fig. 86A), as Larsen ( 1966) showed, the prescutoscutal suture is retained
medially. The intersegmental membrane is attached to the area anterior to the prescuto-
scutal suture. The antecosta must be represented by the marginal line from which the
median longitudinal process and the lateral phragmata arise. The prealar arm arises
from each lateral end of the prescutal area. In Chauliognathus also, the positions of the
lateral phragma and the prealar arm relative to the intersegmental membrane is the same
as in Creophilus.
The shape of the first phragma varies considerably in different groups of Coleoptera.
In Chauliognathus, for instance, the median process is absent; in Gyrinus (Larsen 1966)
the first phragma bears two posteriorly directed processes, and so on. Muscle t-p 3 is
attached on the laterophragma, not on the prescutal prealar arm .
. The prescutoscutal suture may be partly or completely lost. According to Larsen
( 1966), the prescutal prealar arm is formed in Cantharidae (Fig. 86B), Scarabaeidae,
and Elateridae, but it is absent in Adephaga. The arm extends ventrally to the anterior
margin of the episternum, and in Aphodius it is indistinguishably fused with the basalare
(Larsen 1966). The tegula is absent in Coleoptera.
The remainder of the mesonotum has undergone profound modifications, primarily
to hold the resting elytra on its surface. In the relatively unmodified mesonotum of
Chauliognathus (Fig. 86B) the scutoscutellar suture takes a typical V-shape and is
traceable posteriorly as far as its lateral end where the axillary cord starts. In Chaulio-
gnathus, therefore, the division of the mesonotum proper into the scutum and scutellum
is quite clear. Moreover, the scutellum is already lobately prolonged posteriorly, and
this part is exposed between resting elytra. The basal margin of each elytron lies
roughly on the secondary oblique posterolateral line of the mesonotum. This line is not
ridged internally and it !llay correspond to the posterolateral scuta! suture in other orders,
In many beetles including Dytiscus (Fig. 86D), the inner anterior angle of each
elytron fits in the ridges a and b that are formed on the scutum. The median unpaired
plate, the scutellum in taxonomy, is exposed between resting elytra. A comparison of
this median plate with the corresponding area in Chauliognathus (Fig. 86B) shows that
this area was secondarily formed through the partial loss of the scutoscutellar suture and
the development of sutures a and b. In Dytiscus the lateral part of the scutoscutellar
INTERSEGMENTAL. MEMBRANE
PRESCUTUM
BASICOSTALE
ANTECOSTA
A
. PRESCUTOSCUTAL
SUTURE
ATERAL PHRAGMA
SCUTUM PREALAR ARM
SCUTELLUM
B
SCUTOSCUTELLAR
SUTURE
BASISUBCOSTALE (
BASALARE +
AXILLARY SCL.?
ALAR //\ LATERAL
APOPHYSIS POSTNOTUM
GROOVE n-PLEURAL SUTURE
EPIMERON C BASISTERNUM +
PREEPISTERN UM
PRESTERNUM 3
D
+ SPINATERNUM 2
BASI COST ALE
EPISTERNUM 3
ANAPLEURAL
SUTURE
\ BASISTERNUM
BASISUBCOSTALE + PREEPISTERNUM
J ::_

AXIL. SCL.
--e. . 3'RD AXIL. SCL.

I
ELYTRON LATERAL
POSTNOTUM
F
EPIMERAL
LOBE
PARACOXAL
SUTURE?
4TH ABD.
SEGMENT
EEPISTERNUM +
BASISTERNUM
PLEURAL
SUTURE
RON
ridge is still clearly retained and is continuous with the axillary cord. However, in many
other beetles the scutoscutellar suture has been lost (e.g., Creophilus).
The mesothoracic postnotum is greatly reduced and retained only laterally, although
in Creophilus, Coccinella, and Necrophorus the lateral postnotum almost reaches the
middle of the mesonotum (Larsen 1966). As Snodgrass (1909) first showed, the lateral
remnant of the postnotum (yoke plate) in Calosoma connects the lateral scutellar
extension with the second phragma that arises from the metanotum; Snodgrass suspected
it to be the postnotum (pseudonotum of Snodgrass). El Kifl (1953) also showed in
Tribolium a small sclerotized plate, which he homologized with the yoke plate of
Snodgrass ( 1909). The examination of beetles in this work has also revealed that a
yoke plate, which connects the lateral scutellar extension with the metanotal anterolateral
angle, is clearly present in Priacma, and that in Chauliognathus it has a ligamentous
connection with a small subalare.
Since the presence of the mesothoracic postnotum is a well-established fact, the
claims of Snodgrass (1935) and Doyen (1966) that the mesothoracic postnotum is
absent in the mesothorax is invalid. In any case, the presence of a reduced postnotum
in the mesothorax of Coleoptera is not surprising, considering the presence of such a
reduced postnotum in Hymenoptera, Lepidoptera, and some hemimetabolous orders.
The mesoscutallateral margin: In the mesothorax the lateral margin of the scutum
is strongly depressed, and hence the notal processes have become considerably obliterated.
The obtuse process at the middle of the scuta! margin has been called the posterior notal
wing process by many workers including Doyen ( 1966). However, this process appears
to correspond to the postmedian notal process, judging from its association with the
proximal caudal end of the first axillary sclerite (Figs. 86E, 88) and from the fact that
the apparent posterolateral scuta! suture starts at its caudal end. A true posterior notal
process is represented by the fourth axillary sclerite, which lies anterior to the axillary
cord in Gyri nus (Larsen 1966). The undetached, posterior notal process is probably
absent in many Coleoptera, although the scutellar process of Doyen ( 1966) in Tenebrio
may represent the posterior notal process.
Axillary sclerites in the elytral base: In ChauliognathJ,ts (Fig. 86E) the caudal end
of the small first axillary sclerite has a ligamentous connection with a median obtuse
process (postmedian notal process) on the scutallateral margin; the sclerite also connects
with the proximal end of the subcostal process of the elytron. The second axillary
sclerite lies distally along the basal margin of the elytron, posterior to the subcostal
process. The definitive ventral sclerotization of the second axillary sclerite lies above
the dorsal margin of the mesopleuron. However, the anterior part of this sclerite or the
whole sclerite may be the basalare. In the absence of the posterior notal process, the
third axillary sclerite has no obvious connection with the scuta! margin.
According to Larsen (1966), in Gyrinidae the first and second axillary sclerites are
closely attached to each other, the fourth axillary sclerite is apparently present, and the
posterior part of the first axillary sclerite is a free sclerite. As in Dermaptera, close
association of the first and second axillary sclerites is a general tendency in Coleoptera,
and often their original line of fusion may be difficult to recognize. In some beetles
(e.g., Tenebrio, Doyen 1966) the second axillary sclerite has a connection at its caudal
end with the subalare. Herbst (1952) described the elytral base in Rutelinae and
Cetoniinae in considerable detail ...
59His highly -Peculiar interpretation of structures of the elytral base are not discussed here.
FIG. 86. A, cephalic view of mesonotum in Creophilus maxillosus (from Larsen); B, dorsal
view of mesonotum in Chauliognathus pennsylvanicus (original); C, dorsal view of elytral base
in Chauliognathus pennsylvanicus (original); D, dorsal view of mesonotum in a Dytiscus sp.
(original); E, dorsal view of elytral axillary sclerites in Chauliognathus pennsylvanicus
(original); F, ventral view of pterothorax in Harpalus rufipes (from Larsen); G, ventral view
of mesothorax in Pterotus obscuripennis (from Campau).
The mesopleuron: The mesopleuron is divided by the pleural suture into the
episternum and the epimeron. At least in some Coleoptera including Chaulio'gnathus
(Fig. 86C), the pleural wing process continuous from the pleural suture is absent. In
Chauliognathus a small sclerite above the dorsal margin of the mesopleuron is isolated by
a narrow membrane. Campau (1940) called this small sclerite the pleural wing
process; however, as already noted, it may represent the basalare. Since the pleural
wing process is absent in Chauliognathus (which exhibits many primitive features), the
apparent pleural wing process in some Adephaga (Larsen 1966) has most probably
resulted from a secondary fusion of the basalare (or the ventral second axillary sclerite)
with the dorsal margin of the pleuron. Larsen's statement ( 1966, p. 110) that the
basalare is absent in Adephaga except Haliplidae strongly suggests such a possibility.
Furthermore, the tendency for the basalare to fuse with the dorsal margin of the pleuron
is highly pronounced in the metathorax of Coleoptera. In the mesothorax of Dermaptera,
which have a similar wing base, the basalare is also incorporated into the dorsal area of
the episternum. These facts strengthen the idea of the fusion of the basalare (or the
ventral second axillary sclerite) with the dorsal area of the episternum in the mesothorax
of higher Coleoptera. In higher Coleoptera the definitive pleural wing process (basalare?)
forms an elaborate socket, which receives the proximal end of the basal process of
the elytron.
The pleural ridge is continuous dorsally with the dorsal internal process that Doyen
(1966) called the alar apophysis (Transversalapophyse of Stellwaag, 1914a; pleural arm
of Larsen, 1966). This apophysis is most probably not homologous with the pleural arm
in most other orders, since it arises dorsally and externally lacks a pit. In Chaulio-
gnathus the apophysis also is continuous with the membrane surrounding the dorsal part
of the epimeron. .
The paracoxal suture is absent in the mesopleuron of all Coleoptera that have been
studied. The anapleural suture is clearly present in its typical position in Chauliognathus
(Fig. 84), and it divides the anapleuron into the anepisternum and the preepisternum.
The anterior (proximal) margin of the preepisternum thus represents the boundary
between the pleuron and the membranous sternum in Chauliognathus. As Campau
(1940) showed, the preepisternal part of the precoxal bridge is indistinguishably fused
with the sclerotized basisternal area in cantharoid genera, Calopteron, Lucidota, Pterotus
(Fig. 86G) and Podabrus.
In higher Coleoptera the anapleural suture (the pleural costa of Campau, 1940) has
been considered to be the pleurosternal suture by various workers including Larsen
(1966) and Doyen (1966). Thus, in Harpalus (Fig. 86F) or in Tenebrio (Doyen
1966), for instance, the claimed pleurosternal suture on both sides converge anteriorly
from the dorsal (posterior) pleurocoxal articulation, and thus the claimed sternum
surrounds a large part of the ventral coxal base. Actually, however, the claimed pleura-
sternal suture and the basisternum in Harpalus (Fig. 86F) are the anapleural suture, and
the preepisternum fused with the basisternum, respectively. This is immediately clear
in a comparison of these areas in Harpalus with the corresponding areas in
gnathus (Fig. 84) and Pterotus (Fig. 86G). In higher Coleoptera such as Harpalus
(Fig. 86F) the membranous supracoxal part of the pleuron and the membranous sternal
area seen in Chauliognathus and Pterotus have become extensively sclerotized and, at the
same time, the anapleural suture has extended posteriorly so far as to reach the pleural
suture near the pleurocoxal articulation. The definitive basisternum in higher Coleoptera
including Harpalus thus contains the preepisternum.
In higher Coleoptera, the anterior margin of the episternosternal area is often flanged
by a marginal plate. El Kifl. (1953) called this marginal area the preepisternum, and
Doyen ( 1966) doubted that this marginal strip was homologous with the preepisternum
of Ferris and of Matsuda. However, this area clearly corresponds to the secondary
marginal strip (or apodeme) in the corresponding area in many orders, which is called
the prepectus.
The free trochantin is clearly exposed in Chauliognathus (Fig. 84) and Pterotus
(Fig. 86G); in higher families it is often hidden beneath the pleurosternal evagination
(acetabi.U.um), although in Gyrinidae the trochantin is absent (Larsen 1966). The
ventral pleurocoxal articulation is absent, the coxal bases being widely separated.
The dorsal margin of the epimeron is modified into a ridged shelf that is continuous
from the pleural wing process (Fig. 86C); and the resting elytron fits into this shelf. As
already noted, the small subalare is often connected with the caudal end of the second
axillary sclerite.
The mesosternum: In Chauliognathus (Fig. 84) the only sclerotized sternal plate is
the posterior subtriangular plate, the basisternum being membranous. The furcae arise
from near the midle of its anterior margin. In a majority of Coleoptera, according to
Larsen (1966), the furcal arms do not arise from a common base, and there is usually no
muscular connection (p-s 1) between the furcal arm and the alar apophysis (pleural arm
of Larsen).
In higher Coleoptera the anterior membranous part of the sternum in Chaulio-
gnathus is sclerotiztd. Conspicuous modifications occur on the definitive mesosternum
in some beetles. Thus, in Elateridae, as Jeuniaux ( 1950) and Laurent ( 1961) el(plained,
the anterior part of the definitive mesosternum is provided with a median longitudinal
groove, into which the posterior median process of the presternum fits. The two struc-
tures together constitute a part of the jumping device peculiar to the family. In higher
Coleoptera the posterolateral part of the definitive sternum (acetabulum) above the coxal
base is largely the preepisternum that has become indistinguishable from the true sternum.
According to Larsen (1966), the mesothoracic spina occurs in most Carabidae and
is absent in other Adephaga, although the mesosterna! spina is present in larval Peltodytes
(Jaboulet 1960), Riolus (Marvillet 1960), and Sphaeridium (Quennedey 1965).
Apparently, the mesothoracic spina is common in larval Coleoptera; it most likely
degenerates during later periods of development. The loss of the spina in adult beetles
has often obliterated the border between the mesosternum and the metasternum, as
discussed later (p. 214).
For detailed information on the mesothor:tcic supracoxal lobe in various Coleoptera,
see Larsen ( 1966). Bell (1967) discussed the taxonomic significance of the coxal
cavity formed by the supracoxal extension in Adephaga.
The metanotum: In the metanotum of Chauliognathus (Fig. 87A), the inter-
segmental membrane is attached in the area just anterior to the prescutoscutal suture.
Note the presence of the parapsidal suture (Fig. 87B), which reaches anteriorly the pre-
scutoscutal suture as in other insects. The process lying lateral to the prescutoscutal
suture is therefore a prescutal structure. A small sclerite lying lateral to this process has
been called the muscle disc; this sclerite corresponds to the prealar sclerite in other
orders. Muscle t-p 4 is attached to this sclerite. It is not immediately clear whether
this sclerite is homologous with the Phragmasklerit in Dermaptera. In Necrobia
(Tremblay 1958) and Atomaria (Evans 1961) this prealar sclerite is supported by a
tendon.
In Chauliognathus (Fig. 87A) the antecostal suture is horizontal and the bilobed
phragma arises from it internally. This interpretation of the suture agrees with that of
Campau (1940), who also worked on the same species of Chauliognathus. Bostick
( 1945) in his study of Calosoma and Saxena ( 1955) in his study of Mylabris also
followed Campau's interpretations. Doyen's (1966) interpretation of the anterior part
of the metanotum in Tenebrio is also essentially in agreement with the one presented here.
Snodgrass (1909, 1935) overlooked the antecostal suture and the true prescutoscutaf
suture. Therefore, he designated the whole central region of the scutum, which is
bounded by the parapsidal sutures, and a narrow anterolateral area bearing the anterior
notal process as the prescutum. Larsen (1966) followed Snodgrass, with reservation
that the median part of Snodgrass' prescutum may not in its entirety be identical with
the prescutum in other insects.
PRESCUTOSCUTAL
SUTURE
PREALAR
SCL.

.
MEMBRANE
ATTACHMENT

\ GITE
ANTE COSTA
A
PHRAGMA
PRESCUTUM
3'RD AXIL. /)
SCLERITE

NOTAL
PROCESS C
LATERAL
POSTNOTUM SUBALARE
PRE EPISTERNUM
+ BASISTERNUM
PROXIMAL
MEDIAN
PLATE
ISTAL
MEDIAN
PLATE
MEMBRANE
ATTACHMENT
PRESCUTDSCUTAL
SUTURE
MUSCLE DISK
' /.RALARE
NOTAL
_........""""' PROCESS}
:'lll7'+"'+-'i""'----- ALAR
APOPHYSIS
SECOND AXILLARY
SCLERITE
\ SUBALARE
\--LATERAL
POSTNOTUM
ANEPISTERNUM
KATEPIMERON
PARACOXAL SUTURE
ANAPLEURAL
CLEFT
ANEPISTERNUM- /
KATEPISTERNUM
VENTRAL
CONDYLE
-'r--AN EPISTERNUM
ALAR APOPHYSIS
PARACOXAL
SUTURE
F
E
FIG. 87. A, cephalic view of metanotum in Chauliognathus pennsylvanicus (original); B,
dorsal view of metanotum in Chauliognathus pennsy/vanicus (original); C, dorsal view of hind
wing base in Chauliognathus pennsylvanicus (original); D, lateral view of hind wing base in
Chauliognathus pennsylvanicus (original); E, metapleurosternum in Necrobia rufipes (from
Campau); F, dorsal part of metapleuron in Gyrinus marinus (from Larsen).
According to Larsen ( 1966), the bilobed condition of the second phragma is often
obliterated, and in Gyrinidae it is highly reduced. In most Coleoptera the second
phragrria is exclusively metanotal. When the lateral remnant of the mesothoracic post-
notum (phragma) is connected with the metathoracic acrotergite (Fig. 87B), the second
phragma may be said to be laterally double-walled.
In Chauliognathus (Fig. 87B) and other beetles, the anterior.scutal area has, on each
side, an oblique ridge which coverges posteromesally; this ridge is a mechanical modifica-
tion that has developed in Coleoptera; Crampton ( 1918a) called it the alacrista. The
anterior lateral end of the alacrista reaches the anterior end of the anterolateral scuta!
suture; this part, therefore, may represent the lateral parapsidal suture. The median
longitudinal groove formed between the ridge on each side is somewhat widened anteriorly
for the reception of the posterior (anal) margins of the elytra at rest.
The scutoscutellar suture on each side coverges anteriorly and forms a well-
developed ridge internally; this ridge is well marked in all Coleoptera except Gyrinidae
(Larsen 1966). The anterolateral scuta! suture demarcates the anterior notal wing
process and extends to the caudal end of the detached, smaller prealar sclerite (muscle
disc) in Chauliognathus and in many other beetles. The anterior association of the
anterolateral scuta! suture with the prescutoscutal suture is more or less obscured, because
of the modification of the lateral prescutal area already discussed. The posterolateral
scutal suture is apparently present in Macro pus (Riischkamp 1927), although it is absent
in most Coleoptera. Another prevalent suture is a transverse, intrascutal suture that
arises from near the anterior end of the alacrista (Fig. 87B).
In Coleoptera the metathoracic postnotum is well developed, as in Orthoptera and
Dermaptera. In Chauliognathus (Fig. 87B) the postnotum is divided, by an oblique line
on each side, into the median plate and the lateral parts. The lateral postnotum extends
anteriorly as far as the caudal end of the subalar muscle disc and meets, on its way, the
dorsal margin of the epimeron. The third phragma invaginates from the posterior
margin of the postnotum. In other Coleoptera also, the postnotum is often divided into
the median unpaired part and the lateral parts. In Gyrinidae, according to Larsen
( 1966), the reduced median postnotum is infieBted below the scutellum. The associa-
tion of the anterior end of the lateral postnotum with the subalare is also common in
Coleoptera (e.g., Tenebrio, Doyen 1966; Tribolium, El Kifi 1953; Necrobia, Tremblay
1958).
The metanotallateral margin and axillary sclerites: The small, subtriangular anterior
notal wing process lies behind the detached prealar sclerite (or the muscle disc) at the
anterolateral angle of the scutum. Riischkamp (1927) found that the detached prealar
sclerite ( Fliigelgesimse) is often fused with the anterior notal wing process, and the line
of fusion is present in Dytiscus, Lucanus, and Macropus. Larsen (1966) called the
anterior notal process the lateral prescutallobe, which was in error.
The tergal fissure is generally inconspicuous. Hence, the differentiation of the
antemedian and the median notal wing processes is usually not well marked, or is absent
in the metathorax of Coleoptera. However, the posterior notal wing process is often
well developed.
In Chauliognathus (Fig. 87C) the first axillary sclerite articulates proximally with
the anterior notal process and distally with the base of the Sc vein at its head. According
to Larsen ( 1966), in Polyphaga the posterior margin of the first axillary sclerite is almost
straight or concave, and in Adephaga the sclerite is long. In Calosoma (Fig. 88), as
Tietze ( 1963) showed, the long first axillary sclerite is medially constricted and has its
typical association with the postmedian notal process at its posterior end; such a prolonga-
tion of the first axillary sclerite has also occurred in higher Lepidoptera.
In Chauliognathus (Fig. 87C) the dorsal second axillary sclerite articulates on its
proximal margin with the first axillary sclerite, and its proximal anterior margin is
connected with the R vein by the radial bridge as in Plecoptera, Lepidoptera, etc.,
although in some beetles (e.g., Tenebrio, Doyen 1966; Calosoma, Fig. 88) the radial
bridge is absent. In Chauliognathus the second axillary sclerite is continuous distally
MEDIAN
t-p 13.14
3'RO AXIL. SCL.
: ..
. . .
Fro. 88. Hind wing base in Calosoma sycophanta (from Tietze).
with the median plate, but in many beetles the two areas are often separated by a narrow
membrane. The caudal end of the second axillary sclerite articulates with the third
axillary sclerite; the ventral second axillary sclerite lies behind the emargination on the
posterior margin of the pleural wing process in C hauliognathus (Fig. 87D). A tendon
connects the second axillary sclerite and the subalar muscle disc in Tenebrio (Doyen
1966); the connection of the two sclerites probably occurs in many others.
In Chauliognathus (Fig. 87C) the proximal process of the third axillary sclerite is
connected with a small sclerite that lies between the first and third axillary sclerites.
According to Larsen (1966), this small plate serves as a point of attachment for the
flexor muscle (t-p 13, 14), which arises from the pleuron, and the presence of this sclerite
is characteristic of Coleoptera. However, in Atomaria, according to Evans ( 1961), no
such sclerite is present and the flexor muscle is attached to the proximal margin of the
third axillary sclerite.
It appears that a characteristic of the third axillary sclerite in Coleoptera is its
relatively more distal position, and this is due to distal expansion of the posterior notal
process, as typically seen in Calosoma (Fig. 88). The third axillary sclerite is proximally
attached or connected with the apex of the posterior notal process as in other orders.
In Chauliognathus (Fig. 87C) and Atomaria the posterior distal area of the third axillary
sclerite lies above the subalar muscle disc. The anterior distal angle of the third sclerite
is associated with the bases of the A veins. In Chauliognathus (Fig. 87D) the subcostal
process (basisubcostale) is connected with the tip of the pleural wing process and fits into
the small space formed in front of the latter, when the wing is folded.
The metapleuron: The pleural suture is always traceable dorsally from the ventral
point of articulation with the coxa to the pleural wing process. The direction of the
pleural suture is nearly horizontal because of the extension cephalad of the enlarged
metathorax. In Chauliognathus the alar apophysis, which is serially homologous with
the one in the mesothorax, is continuous from the pleural ridge (Fig. 87D).
The paracoxal suture in the metathorax appears to be represented by a transverse
suture that lies above the coxal base in some beetles. As Campau (1940) showed in
Enoclerus, Nacerda, Necrobia (Fig. 87E) and Cupes, the apparent paracoxal suture
arises from the ventral process (for coxal articulation) and extends laterally to the
anapleural cleft near its posterior end. In Priacma examined in this study, a transverse
trochantin, which is much larger than the one in Necrobia (Fig. 87E), lies immediately
MATSUDA: THE INSECT THORAX 2B
above the coxal base and below the paracoxal suture. Further, in Chauliognathus, as
Matsuda ( 1960a) pointed out, the paracoxal suture is retained also on the epimeron
(Fig. 87D). At first glance, the location of the paracoxal suture may appear to be too
posterior to be the true one, but this is due to the great enlargement of the metathorax,
especially of its anterior part, during the pupal stage. This is evident from the fact that
the metathorax is about as large as the metathorax in larval Peltodytes (Jaboulet 1960;
Fig. 89), Rio/us (Marvillet 1960), and Sphaeridium (Quennedey 1965). Such a
posteriorly located paracoxal suture is present also in some other orders, such as
Mecoptera, Diptera, etc.
The anapleural cleft usually remains membranous in the metathorax. It is usually
long and reaches the pleural suture near its posterior end; this elongation of the cleft (or
the suture) is also correlated with the enlargement of the metathorax. The course of
the anapleural cleft or suture in dytiscids is considerably more oblique than it is in other
Coleoptera, as Balfour-Browne ( 1964) pointed out. This oblique direction of the
relatively short anapleural suture is obviously correlated with the relatively short meta-
thorax in this family.
The anepisternum in the metathorax of most Coleoptera is thus a relatively long
plate between the pleural suture and the anapleural cleft (Fig. 87E). A distinctly
isolated basalare is present in Gyri nus (Fig. 87F). In many beetles, however, the
basalare has become indistinguishably fused with the dorsalpart of the pleuron (as in
the mesothorax), or with the pleural wing process. The posterior end of the basalare
or of the basalar area is connected internally with a cup-shaped alar apophysis, which is
also called the basalar disc because of its obvious connection with the basalare.
The preepisternum in the metathorax of Chauliognathus (Fig. 84) is probably
represented by a broad plate, anterior to the presumed original katepisternal area, on
each side of the median longitudinal suture. The more proximal part of the definitive
preepisternum (or the definitive sternum) along the median longitudinal suture is
probably the basisternal area, which, however, is no longer definable because of its fusion
with the preepisternum.'" In Mylabris Saxena ( 1955) interpreted the area in question as
the preepisternum.
In many other beetles, such as Harpalus (Fig'. 86F), Leptinotarsa (Rivnay 1928),
Hydrophilus (Richmond 1931), Staphylinidae (Blackwelder 1936), Ararderus (El Sayed
1940), Sandalus (Rings 1942), Ca/osoma (Bostick 1945), Tribolium (El Kifi 1953),
Necrobia (Tremblay 1958), Rhyncholus (Amzy 1958), Tenebrio (Doyen 1966), the
anterior margin of the definitive preepisternum is lost, and the preepisternal area and the
anteriorly placed sternum (spinasternum 2 + presternum 3 in Chauliognathus, Fig. 84)
are continuous.
In the metathorax, modifications of the area (mainly pleural) surrounding the coxal
base are generally less pronounced than in the mesothorax. When the metacoxae are
movable, the coxa has a ventral articulation on its mesal margin with a process that arises
from the furcal base.
The metathoracic epimeron is a simple, undivided plate in most beetles, although
rarely (e.g., Chauliognathus) the paracoxal suture separates off a small katepimeron from
the rest of the epimeron or the anepimeron. A padlike semimembranous area occupies
a dorsal area of the epimeron in beetles; the inner surface of the costal margin of the
resting elytron lies on this area. For more information about the wing-locking
mechanism, refer to Breed and Ball (1909), Stellwaag (1914a, b), Hatch (1926), Corset
(1931), Heberdey (1938), Lindroth (1948), Herbst (1952), and Larsen (1966).
The metathoracic subalare forms a well-developed, cup-shaped apodeme, to which a
flight muscle (t-cx 8) is attached. In some gyrinids the subalare is absent (Larsen 1966).
The metasternum: In Chauliognathus (Fig. 84) the subtriangular sternal plate
anterior to the definitive preepisternum is contiguous on each side to the posterior
extension of the mesothoracic epimeron, and its anteromesal end reaches the meso-
thoracic furcal bases. Nearly all authors have interpreted the corresponding area in
03
See p. 214 for further discussion of the border between the preepistemum and the basisternum.
PR
SPINA 1 ESTERNUM.
ANEPISTERNUM
BASISTERNUM
PARACOXAL
SUTURE?
EPIMERON
POSTCOXAL
BRIDGE
SPINA 2
PRE EPISTERNUM
ESTERNUM
+ AN EPISTERNUM
BASISTERNUM
EPIMERON
COXAL
CAVITY
FIG. 89. Ventral view of larval pterothorax in Peltodytes sp. (from Jaboulet).
other beetles as the mesothoracic sternum, although Campau ( 1940) regarded the area in
Chauliognathus as the metathoracic sternum.
The discrepancy, however, can be accounted for by referring to the ventral aspect
of the pterothorax in the larva of Peltodytes. In this larva (Fig. 89) the mesothoracic
spina clearly demarcates the boundary between the mesosternum and the metasternum.
Taking the spina as a landmark, the paired, transverse subtriangular plates anterior to
the spina can be interpreted as the spinasternum, as Jaboulet ( 1960) called them. In the
metasternal area paired anterior plates are the presternum in more generalized insects.
Although the basisternum is a narrow median longitudinal space, it maintains exactly the
same position relative to the preepisternum as in hemimetabolous orders, i.e., the ventral
(proximal) margin of the preepisternum directly faces the basisternum.
During later periods of the postembryonic development, the larval mesothoracic
spina most probably becomes lost in most beetles... At the same time, the metathoracic
presternum will either become lost or fused with the mesothoracic spinasternum. The
area under discussion in Chauliognathus (Fig. 84) and the corresponding area in other
beetles are, therefore, provisionally regarded as a composite area consisting of the
mesothoracic spinasternum and the metathoracic presternum.
If the narrow median longitudinal space between the preepisterna in the larva of
Peltodytes (Fig. 89) represents the basisternum, the median longitudinal suture in the
adult, which forms a ridge internally, is purely basisternal. Presumably the basisternal
area will be pressed into the thoracic cavity and the coxal bases will be brought closer
together towards the median longitudinal axis during later stages (pupal stage?) of the
postembryonic development. The degree of invagination of the basisternum and possibly
of the preepisternum must vary in different beetles; in some beetles the original basi-
sternal area is presumably present indistinguishably along the median longitudinal suture.
The metasternal furca exhibits a wide variety in shape, size, and position in various
groups of Coleoptera. Crowson (1938) thought that lymexylids exhibit the most
primitive condition of the furca in having free arms, extensive lamina, and anterior
tendons that arise close together and supported merely by a median thickening of the
lamina. Balfour-Browne (1961) classified five types of the furca; the simplest, hence
MAccording to Larsen (1966), the mesothoracic spina is not present in any adult Coleoptera except Carabidae,
whereas in many larval beetles this spina occurs (see p. 209).
215
perhaps the most primitive furca has a wide transverse base with an arm at each end.
Crowson ( 1944, 19 55) and Riha ( 19 55) also showed various types of furca.
Modifications of the pterothorax in apterous and brachypterous forms: Correlated
with reduction of the hind wings, various other parts of the pterothorax, especially the
metathorax, have undergone considerable modifications in some beetles. These modifica-
tions are mainly obliterations of various thoracic sutures and parts as a result of increased
sclerotization. Some muscles, especially the flight muscles, have become reduced. For
detailed information, refer to Riischkamp (1927), Jackson (1928, 1933, 1952, 1956a, b),
Leech (1942), Larsen (1954), Tietze (1963), and Smith (1964).
Phylogenetic relationships: Crampton ( 1918a) emphasized the close affinity between
the Dermaptera and Coleoptera. He pointed out the following structural similarities
between the two orders: the presence of the myodisc in the meta thorax, the similar shape
in the first axillary sclerite (np of Crampton), a pronounced tendency for the metanotal
region to become broader than long, a marked tendency for the scutellar region of the
mesonotum to become pointed posteriorly and to overlap the anterior portion of the
metanotum behind it, a marked tendency for the mesothoracic postnotum (postscutellum
of Crampton) to become reduced, the formation of two alar ridges on the metanotum,
the presence of a transscutal suture on the metanotum.
Crampton was correct in pointing out all these structural similarities except for the
similar shape in the first axillary sclerite, which is a matter of choice of specimens,
especially in Coleoptera; the other similarities are either synapomorphic or convergences.
Other probable synapomorphic characters in Dermaptera and Coleoptera which this
study has revealed are the tendency to form the gula, and the absence of the mesopleural
wing process. The presence of the presternal cervical sclerite in Coleoptera, Dermaptera,
and other hemimetabolous insects can be regarded as symplesiomorphic. All of these
characters support Crampton's idea that Dermaptera and Coleoptera are closely related,
and appears to preclude a possible close relationship of Coleoptera to other holo-
metabolous orders.
Tillyard (1931) noticed the similarity in the hind wing between Staphylinidae and
the permian Protelytroptera, which he regarqed as ancestral to Dermaptera; and he
regarded this as due to convergence. Similarly, Kleinow (1966) regarded the similarity
in the hind wing folding mechanism between Staphylinidae and F orficula as due to
convergence. The phylogenetic position of the Protelytroptera is, however, rather
uncertain. For instance, one of the major characters Tillyard used in erecting this order
was the presence of short cerci, which led him to think that the Protelytroptera are
probably not holometabolous. However, the cercus occurs in some Coleoptera,
Hymenoptera and some other Holometabo1a, especially in their larvae. The relation-
ships of Protelytroptera to Dermaptera and to Coleoptera require reconsideration.
For the discussion of Protocoleoptera, originally proposed as ancestral to Coleoptera,
refer to Till yard (1924), Forbes ( 1928) and Zeuner ( 1933). Based on certain larval
characters and the complete metamorphosis, Coleoptera have been regarded as relatives
of Neuroptera by many workers including Crowson (1960).
The anatomical basis of flight: In the mesothorax the e1ytral margin is locked into
the metanotallongitudinal groove by the action of muscle t 14 (Stellwaag 1914a; Herbst
1952); the apex of the mesoscutellar lobe is raised and the elytra are simultaneously
extended by the action oft 12 and t-p 3 (Stellwaag 1914a; Herbst 1952); by contraction
of t-p 16 elytra are brought rearward (Larsen 1966), and t-p 13, 14 are flexors of elytra;
t-tr 1 may also act as an elevator of elytra (Larsen 1966); p-cx 2 and t-cx 8 are also
probably flight muscles;because they are absent in the forms that have greatly reduced
flight organs (Larsen 1966) .
In the metathorax Larsen (1966) found that eight fibrillar muscles (p. 218) are
direct or indirect flight muscles. Of these fibrillar muscles, t-14 is a depressor of the
wing and acts antagonistically to t 12, t-p 5, 6, t-cx 1 or 2, and t-cx 3 (Larsen 1966).
As Menon and Joshi ( 1957, 1958) and Larsen (1966) convincingly showed, t-cx 1 or 2
is functionally an elevator of wings. Increased incorporation of the leg base into the
thoracic wall in some beetles has resulted in the change of point of origin of t-cx. 1 or 2
to a cup-shaped apodeme near the apex of the trochantin, and hence' its dorsal point of
attachment is functionally a point of insertion; contraction of this muscle tl;lerefore
elevates the hind wing, Larsen (1966) thought t-cx 3 was also a flight muscle, since it
becomes reduced in flightless forms, p-cx 2, as p 3, is a pronator and depressor of
wings, but it can also be an extensor of wings. t-cx 8 acts as a supinator, which depresses
the posterior margin of the wing during an upstroke; this is done by virtue of the
connection of the subalare (to which t-cx 8 is attached) with the second and third
axillary sclerites, Thus, in Coleoptera the action of t 14 is assisted by p 3, p-cx 2,
and t-cx 8.
Chadwick ( 1953) stated that in certain more modern insects modifications of the
flight mechanism involved the approximation of the leg bases beneath the center of
gravity, and the tendency for the coxal segment of the leg to become incorporated into
the body wall. Chadwick thought that this secondary development reversed the former
origins and insertions of several of the dorsoventral muscles, which made them highly
effective as depressors of the tergal margin and hence as elevators of the wings. In
Coleoptera t-cx 1 or 2, t-cx 3, and p-cx 2 represent this stage of modification; these
muscles have no influence on the leg movement.
Among the dorsal tergopleural muscles in the metathorax, t-p 4 bends the pleural
wing process medially; the action of t-p 8 must be correlated with the action of p 3 and
p-cx 2, opposing the pull of the latter (Larsen 1966); t-p 16 has a controlling effect on
the position of the second axillary sclerite (Larsen 1966); t-p 13, 14 are usual flexors of
the wing. p-s 1 probably has no effect upon the click mechanism in Coleoptera (Larsen
1966), although in a coccinellid Pringle (1957a) related the click mechanism top-s 1.
Besides the skeletal modifications involved with the movement of wings already
discussed, more detailed information on the anatomy and the function of the skeletal
parts concerned with flight are available in the works of Stellwaag (1914a, b),
Riischkamp (1927), and Herbst (1952).
The adult musculature: The gross aspects of thoracic muscles in the adult Coleop-
tera have been studied by Berlese ( 1909b) in Hydrophllus piceus, by Bauer ( 1910) in
Dytiscus marginalis, by Maki ( 1938) in Cicindela kaleea, Chlaenius naeviger, Borolinus
minutus, Epilachna vigintioctopunctata, Ceropria induta, Rhaphidopalpa femora/is,
Mime/a testaceiviridis, by Saxena ( 1953) in Mylabris pustulata, by Larsen ( 1954) in
Gyrinidae (flight muscles), by Donges ( 1954) in Cionus scraphulariae (prothoracic
muscles), by Jackson (1952) in Agabus (flight muscles), by Menon and Joshi (1957) in
Stenocera chrysis (flight muscles), by Menon and Joshi (1958) on a tergocoxal muscle
in beetles, by Tietze ( 1963) in Carabidae (flight muscles) , by Smith ( 1964) in some
Coleoptera (flight muscles), by Davydova (1966) in Lampyridae (flight muscles), by
Larsen (1966) in Coleoptera, and by Balfour-Browne (1967) on the metathoracic
muscles in Dytiscidae. Since Larsen's ( 1966) study is most comprehensive, the thoracic
muscles recognized by him are homologized in Table X.
The thoracic musculature in Coleoptera has undergone considerable reduction in
number of muscles, presumably as a result of increased sclerotization of the exoskeleton.
A strongly sclerotized exoskeleton simply cannot serve as a functional surface for small
muscles. Characteristically, the dorsoventral muscle (t-p 5, 6) occurs only in the meta-
thorax. As Larsen ( 1966) showed, t-cx 1 or 2, t-cx 3, and p-cx 5 are fibrillar and they
are flight muscles in Coleoptera.
Gross aspects of the postembryonic muscle development: A comparison of the
adult thoracic muscles in Dytiscus marginalis studied by Bauer (1910) with the larval
thoracic muscles in the same species studied by Speyer ( 1922) shows the following
development of the thoracic muscles. The tabulation relates to the mesothoracic muscles,
the metathoracic muscles being similar.
(1) Larval muscles that are found in the adult:
I 1 (op-t 3), I 2 (op-t 2), I 3a, b (t-12), I 4a (s 1 or 2), I 6 (op-cv 3), I 7a, b
(t-s (cv) 1), I 8 (t-s (cv)9), I 10 c (t-p muscle?), I 12 c (t-s 1), one of I 13a, b (t-cx 1,
2), one of I 13d, e (t-cx 6 or 7, 8), I 14d (p-cx 9?), II 1 (t 14), II 3 (t-12), II 4a (s 13),
II llb (t-s 1), II 13a, b (t-cx 1, 2), II 13d (t-cx 6, 7), II 14a, b (p-cx 4, 5?),
II 14d' (s-ex 3).
(2) Larval muscles that become lost:
(a) Those present in other adult Coleoptera include II 4e (p-s 3), II 10c (t-p 3),
II lOg (t-p 16?).
(b) Those present in the adult of other orders include I 5 (s 14), I lOa (op-p 2,
op-s 2 in adults?), I 12a, b (t-p 1, 2), I 13c (p-cx 9?), I 14a, b (p-cx 4, 5), I 14c
(p-cx 6?), I 15 (p-cx 7), I 16 (s-ex 4), II 4e (p 1), II 4f (p-s 5), II 4g (p-s 1), II lOb
(t-p 4?), II lla (t-s 3), II 12b (p-s 6?), II 12c (t-s 2? or t-p 17?), II 14c (s-ex 6?),
II 15 (p-cx 7), II 16 (s-ex 4), II 17 (s-ex 1).
(c) Those that are absent in adult insects include I 4b (cervicoprothoracic p-s 7?),
I 9 (t-s muscle for respiration), I lOc (t-s muscle), I lOb-f (t-p musch:s), I 11 (t-s 1?),
I 12d (p muscle), one or two of I 14 a-d (p-cx muscles), II 4b (p muscle), II 4c (p-s 8),
II 4d (p-s 7), II lOe (t-p muscle), II 10f (t-p muscle), II 12d (p muscle).
A striking fact is that the large number of p-s and p-cx muscles become lost during
later periods of development. These muscles are often present in the adult of lower
orders, such as Neuroptera, Blattaria, etc. Most of the muscles listed under 2c have
been produced caenogenetically. Muscle II 11a, which is listed under 2b, is supposed to
become attached to the first or second axillary sclerites in the adult Ephemeroptera.
Breed's ( 1903) study on the postembryonic development of metathoracic muscles in
Thymalus marginicollis provides some information which more recent studies have failed
to produce; important facts that his study revealed follow.
A dorsal longitudinal muscle, l.mt'nt' is t 12, which is absent in the old pupa and
the adult. t-cx 6 or 7 (fix. cox. mt'thx 2) consists of three bundles in the larva, and
they become closely approximated during the pupal stage. Similarly, t 14 (mt'nt) in
the larva exists as three distinct muscles, and later in the young adult it becomes a single
muscle. p 3 (ext. a!. mag. mt'thx) consists of three or four short fibers that lie above
the base of the larval leg. These fibers elongate very rapidly in the pupa, and fuse
completely at their ends; in the adult the muscle is even larger and splits ventrally into
two, spreading backward and downward to the lateral expanse of the definitive meta-
sternum (preepisternum). Thus, in Thy malus p 3 appears to arise as a p-cx muscle.
t-cx 8 (ext. al. pa. mt'thx) is a two-bundled tergocoxal muscle in the larva. During the
pupal stage the two bundles fuse and shift their dorsal attachment to the subalare
(posterior cupula of Breed). t-p 13 (fix. a!. mt'thx 1, 2) is a single mass of fibers in
the larva. p 1 ( e'stn.) forms during the pupal stage. s 20 (ext. cox. mt'thx 2) is a
muscle of the first abdominal segment; it becomes attached to the metafurca during the
pupal stage and becomes s 20. p-x 9 (v of Breed), which extends obliquely between the
dorsal part of the anterior interpleural area and the ventral end of the posterior inter-
pleural ar.ea, is one of the first larval muscles that disappears.
The postembryonic development and histology of flight muscles: In Calandra,
according to Murray and Tiegs (1935), the pleural and dorsoventral muscles (p 3 and
t-p 5, 6) develop from the myoblasts of the larval metathoracic transverse muscles; t 14
develops from the median dorsal muscle in the larva, and t 12 probably from the larval
dorsolateral muscle.
Murray and Tiegs (1935) described the earlier development of p 3 as follows: "In
the developing prepupa the myoblasts (freed from larval muscles) proliferate exceedingly
rapidly, and within about a day have formed a narrow band of cells adhering at this stage
to the thoracic wall in the position of the larval muscle-fibers, which they have replaced.
Within this band of myoblasts there are now differentiated three vertical columns of
structureless cytoplasm with scattered nuclei, each column being invested by a dense
layer of myoblasts. From each of these columns with its adhering myoblasts one of
the pleural muscles will form".
In the later prepupa, thickened, developing muscles begin to separate at their lower
ends, and in the 3-day pupa they attain their definitive position. At this stage the
myoblasts begin to invade the central core of cytoplasm within whose substance they
adopt a longitudinal polarity. Mitosis and chromatic globule extension occur as late as
the eighth-day pupa. A filament grows from opposite ends of each myoblast. These
filaments are the rudiments of the muscle fibrils.
According to Larsen (1966), eight pairs of muscles (t 14, t 12, p 3, t-p 5, 6, t 12,
t 14, t-cx 1 or 2, t-cx 8) are fibrillar muscles in Coleoptera. Darwin and Pringle (1959)
found that the average diameter of fibers of p 3 is 80 p. in Lucanus, 110 p. in Oryctes, and
140 p. in Melo/ontha. In Carabidae and Dytiscidae Smith ( 1964) found that the fiber
diameter is only 25-35 p., and in abnormal flight muscles as little as 2.5 p.. Jordan
( 1955) also studied striated muscles in Popilia japonica.
Degeneration of flight muscles: Degeneration of flight muscles is known to occur
in some adult Coleoptera. In Trypodendron /ineatum Chapman ( 1956, 1958) found
that flight muscles degenerate during gallery construction but regenerate before the beetle
leaves the gallery. Reid ( 1958) conducted a similar study on the changes of flight
muscle in the female of Dendroctonus monticolae. Atkins and Farris ( 1962) also
established the fact that in Dendroctonus pseudotsugae indirect flight muscles degenerate
during the establishment of brood galleries.
The above-noted degeneration of flight muscles in adult beetles must be distinguished
from the developmental retardation or failure of flight muscles in flightless macropterous
and brachypterous forms in some Coleoptera. Thus, Jackson (1933) found the flightless
macropterous form in Sitona. Jackson ( 1952, 1956a, b) further showed in various
aquatic Coleoptera that flight muscles are more or less atrophied in the macropterous
form, and in such cases some modifications of the metathorax and other parts of the
body occur. Jackson ( 1952) suggested that beetles with abnormal flight muscles are
individuals in which the normal development of the flight muscles and their supports has
been arrested at an early stage. Tietze's ( 1963) study on Carabidae and Smith's ( 1964)
study on Ptinidae and others, which involved detailed histological studies of flight muscles,
have convincingly supported Jackson's hypothesis.
An interesting fact that Tietze's ( 1963) study has revealed is that direct flight
muscles and other dorsal tergopleural muscles (t-p 3, 4, 8, 13, 16) persist when indirect
flight muscles are lost or greatly atrophied; and Davydova (1966) confirmed this tendency
in Lampyridae. Tietze ( 1963) also thought that degeneration of flight muscles repre-
sents the first evolutionary step towards rudimentation of wings. Larsen ( 1949b)
compared the atrophied flight muscles in the macropterous forms of some Carabidae with
the muscle resorption in some Heteroptera.
THE ZORAPTERA
No detailed study has been made on the morphology of the zorapterous thorax.
The following description and discussion are based on the observations of the external
structures of Zorotypus spp. by Crampton ( 1920a) and Delamare-Deboutteville ( 1947).
The neck region: The cervical sclerites in Zorotypus (Crampton 1920a) consist of
the well-developed dorsal cervical sclerite and the two-part lateral cervical sclerite
(Fig. 90A) which articulates with the episternal plate (lateropleurite of Crampton) .
. Since this episternal plate is, as discussed below, the anapleuron, the lateral cervical
sclerite must be presternal in origin.
The prothorax: The pronotum in Zorotypus (Fig. 90A), as illustrated by Crampton,
is a simple plate. Characteristically, the elongate anterior episternal plate is separated
by a membrane from the trochantin (Figs. 90A, B; 91). This episternal plate therefore
must be the anapleuron; its proximal area represents the preepisternal area. The
pleurosternal suture (or cleft) separates the preepisternal area from the lateral margin of
the basisternum (Fig. 91). The anapleural cleft (suture) is absent on this anterior
anapleural plate.
TABLE X
Thoracic musculature in Coleoptera
Coleoptera (Larsen 1966)
Muscle designations Remarks
1
op-t 2
2 op-t3
10
op--s 1 and op-cv 3
8
t-s(cv) 9
9 t-cv 1
7
t-cv 2 and t-s 1
6 cv-s 4
5
s 1 or 2
3, 4 t 12
29
61
2 t 14
28
60
13 (I) t-p 1
33 (II) t-p 3 1
67 (III) t-p 4 2
64 (Ill) t-p 5 or 6
69 (III) t-p 8
36 (II) t-p 13 and 14
71 (III)
68 (III) t-p 15 or 12
35 (II) t-p 16
70 (III)
34 (I I) t-p 16? 3
11 t-s 1
32
65, 66
14 t-ti(cx) 1 and 2
39
74
75 (III) t-ti(cx) 3
15 t-cx 6 or 7
40
76
17 t-cx 8
43
79
72 (III) p 1
73 (Ill) p3
37 (II) p-s 1 4
38 (HI) p-s 3
80 (Ill) p-ti(cx) 1
16 (I)
42 (II)
p-ti(cx) 2
78 (III)
41 (II) p-cx5
77 (III)
31 (I-II) s 12
63 (II-III)
30 (HI) s 13
62 (IHII)
Coleoptera (Larsen 1966)
45 (II)
82 (III)
19
46
83
18
44
81
4 7 (II)
84 (III)
48, 49 (II)
50 (II)
20 (I)
51, 52 (II)
85 (III)
REMARKS (Table X)
TABLE X (Continued)
Muscle designations
s-ex 2
s-ex 3
s-ex 5
t-tr 1
p-tr 1
p-tr 2
p-tr 3
s-tr 1
Remarks
5
( 1) 33 is inserted anteriorly on the anterolateral corner of the mesonotum or the
second phragma and posteriorly on the alar apophysis (pleural arm of Larsen).
(2) 67 extends between the prealar sclerite and the pleural ridge just below the
pleural wing process. Larsen ( 1966) homologized his 67 with 33.
(3) 34 was found only in Aphodius and Trox. It is a short muscle with a wide
attachment area on the epimeron and is inserted on the lateral margin of the mesonotum
just anterior to the lateral process of the second phragma. This muscle may be a
modified t-p 16.
( 4) 37 stretches between the furcal arm and the pleural ridge below the alar
apophysis.
(5) 20 arises from the cryptopleural ridge, which is the displaced epimeron.
Besides the muscles recorded by Larsen ( 1966), the following muscles are known
to occur in Coleoptera: 25 in Epilachna and Rhaphidopalpa (Maki 1938), which arise
from the lower portion of the mesothoracic episternum and are attached to the subalare
(t-p ?); 96 in Hydrophilus (Berlese 1909b), 32 in Rhaphidopalpa and 30 in Mimela
(Maki 1938), which are the epimerotrochanteral muscles (p-tr 3) in the pterothorax;
2 in the prothorax of Mimela (Maki 1938), which is t 13; Musculus levator capitis
verticalis in Dytiscus (Bauer 1910), which is op-cv 1, or possibly op-cv 2; Musculus
rotator capitis inferior in Dytiscus (Bauer 1910), which is op-s 2.
The paracoxal suture is absent on the episternum, but a distinct, subhorizontal ridge
near the posterior end of the epimeron (Fig. 90B) may represent the paracoxal suture
(ridge), or it may represent the caudally directed pleural arm as in Grylloblatta.
Delamare-Deboutteville ( 1947) thought that this ridge cuts the katepimeron ventrally.
The large, bipartite trochantin articulates anteriorly with the coxal margin.
The prosternum in Zorotypus (Fig. 91) consists of the anterior basisternum, which
meets the preepisternum along the pleurosternal suture on each side, and the posterior
membranous area, in which the sternellum (see p. 20) (justasternellum of Delamare-
Deboutteville 1947) lies. The furcae arise from a common base at the posterior end
of. the sternellum. The prothoracic spina is absent.
The meso- and meta-notum: In winged Zorotypus (Fig. 90A, D) the anterior
margin of the mesonotum is produced medially. The prescutoscutal suture is absent in
the figure given by Crampton. The well-developed prealar arm (Fig. 90A) reaches the
MATSUDA.; THE INSECT THOKA.X
PRONOTUM
LATERAL PARAPSIDAL
SUTURE
SUBALARE
EPISTERNUM
(ANAPLEURON)
DORSAL
CERVICAL
SCLERITE
LATERAL
CERVICAL
SCLERITE
VENTRAL
CERVICAL
SCLERITE
ANAPLEURAL SUTURE
RIDGE
AN EPISTERNUM
+ PREEPISTERNUM
PARACOXAL
PLEURAL ARM RIDGE
COXA/
'PARACOXAL RIDGE
(SUTURE)?
KATEPIMERON?
B
TROCHANTIN
POSTNOTUM PREALAR ARM
POSTNOTUM
TROCHANTIN
BASISTERNUM
PLEUROSTERNAL
SUTURE?
DORSAL
CERVICAL
SCLERITE
SCUTOSCUTE-
LLAR SUTURE
221
FIG. 90. A, lateral view of thorax in winged Zorotypus hubbardi (from Crampton); B, lateral
view of prothorax in wingless Zorotypus guineensis (from Delamare-Deboutteville); C, lateral
view of meso- and meta-thorax in wingless Zoroty pus guineensis (from Delamare-
Deboutteville); D, dorsal view of thorax in Zorotypus hubbardi (from Crampton).
episternum both in the mesothorax and metathorax; the arm appears to be a prescutal
extension, as in Plecoptera and Psocoptera. The suture arising from the base of the
prealar arm must be the lateral parapsidal suture. The scutoscutellar suture is present
both in the mesonotum and in the metanotum. The postnotum is, as in Plecoptera, also
clearly present beneath the posterior end of the scutellum in both pterothoracic segments
(Fig. 90A).
The pterothoracic tergal surface in Zorotypus (Fig. 90D) is similar to that in
Psocus (Fig. 93B). In Psocus also the lateral parapsidal suture and .the scutoscutellar
suture are the only sutures that are clearly present, and the anterior margin of the meso-
notum is produced medially.
The meso- and meta-pleuron: In the winged form of Zorotypus (Fig. 90A) the
anapleural suture is distinct, and the membranous cleft anterior to the trochantin roughly
corresponds in position to the paracoxal suture. The dotted line on the episternum,
which is roughly in line with the paracoxal cleft and reaches the pleural suture, probably
does not represent the paracoxal suture, since a more probable paracoxal suture on the
epimeron of the wingless form qf Zorotypus (Fig. 90C) lies far more ventrally. A
dotted, subhorizontal line below the anapleural suture (Fig. 90A) may represent the
pleurosternal suture.
In both segments of the wingless form (Fig. 90C) the epimeron has an oblique
suture or ridge, which may be the paracoxal suture, since its position agrees well with the
paracoxal suture on the epimeron in Dermaptera (Fig. 79C), Micropteryx (Fig. 147A),
etc. The pleural arm lies near the middle of the pleural ridge, and the ventral end of
the pleural ridge forms a condyle, which articulates with the coxa.
The meso- and meta-sternum: In winged Zorotypus (Fig. 90A) the basisternum is
separated by the indistinct pleurosternal suture from the preepisternum; in wingless
Zorotypus (Fig. 91) the pleurosternal suture has apparently become completely lost; the
definitive mesosternum and metasternum in the wingless form are, as in some Plecoptera,
anterolaterally expanded, due to inclusion of the preepisternal area. The furca arises
from the posterior end of the sclerotized sternal plate in both segments. The meso-
thoracic spina is absent.
Phylogenetic relationships: Crampton (1921, 1938) thought that Zoraptera are the
most primitive hemipteroid insects, and exhibit so many characters suggestive of a close
relationship with Isoptera, that this might be taken to indicate that they and hemipteroid
insects in general were derived from the same protorthopteroid ancestor as Isoptera.
Gurney (1938) and Ross (1955) also came to essentially the same conclusion.
The similarities in many thoracic structures among Zoraptera, Plecoptera, and
Psocoptera in the foregoing discussion, however, also support Gurney's (1938) statement,
"Regarding the thoracic sclerites, these are suggestive of relationship with Plecoptera and
other lower orders, as well as Corrodentia". Hennig ( 1953) was inclined to believe in
their close relationships with Psocoptera, Plecoptera, and Embioptera.
THE PSOCOPTERA
The neck region: In the relatively long, membranous neck region of Psocoptera
only the lateral cervical sclerite is present. So far, neither the dorsal cervical sclerite nor
the ventral cervical sclerite has been observed. The postoccipital condyle, with which
the lateral cervical sclerite articulates, is well developed in Psocoptera. In Stenopsocus,
as Badonnel (1934) showed, the postoccipital condyle is a slender process; in Psocus
venosus examined in this study and in another species of Psocus, which Crampton
( 1926a) studied, it is a short and robust process.
In Embidopsocus (Fig. 92), as Crampton ( 1926a) showed, the posterior point of
union of the lateral cervical sclerite is evidently with the preepisternal area. Hence, the
lateral cervical sclerite is of presternal origin. In Psocus (Fig. 94A) and Stenopsocus
(Fig. 96B) it is united with a small isolated episternal plate. Crampton (1926a), how-
ever, called the plate in Psocus the postcervicale; Cope ( 1940a) also called the same
plate in Psocus confraternus the cervical sclerite. What is called the posterior lateral
cervical sclerite in other insects lies posteroventral to the anterior lateral cervical sclerite,
whereas the corresponding plate in Psocus spp. lies posterolateral to it. Thus, the small
plate in question is not the cervical sclerite, as Badonnel ( 1934) pointed out in Stenop-
socus; it is the reduced preepisternum, as will be discussed later.
FORAMEN
MAGNUM
FURCA
LATERAL CERVICAL
SCLERITE
FURCA
223
FIG. 91. Ventral view of thorax in Zorotypus guineensis (from Delamare-Deboutteville).
The pronotum and the propleuron: The pronotum is greatly reduced in length, and
is fused laterally with the similarly reduced propleuron. In Embidopsocus (Fig. 92) and
Psocus (Fig. 94A) the line of union of the propleuron with the pronotum is clear,
although in Stenopsocus it has been lost.
The pleural suture is always present and its ventral end articulates with the coxal
margin. The preepisternal area in Embidopsocus reaches the anterolateral angle of the
basisternum. In other genera it is reduced to a small, isolated plate, with which the
lateral cervical sclerite is united or articulated. Neither the paracoxal suture nor the
anapleural suture is clear. The trochantin has no direct articulation with the coxal
margin. The epimeron is well retained. In Stenopsocus and Psocus a conspicuous
process invaginates peculiarly from the ventral margin of the episternum above the
trochantin (Fig. 94A); this pleural process is distally united with the tip of the furcal arm.
The prosternum: In Embidopsocus (Fig. 92) the prosternum is broad and well
sclerotized. Its division into the basisternum and the furcasternum by the sternacosta
224
PREEPISTERNUM
TERGO-
PLEURAL
SUTURE
BASI STERNUM
STERNACOSTA
FIG. 92. Neck and prothorax in Embidopsocus sp. (from Crampton).
is evident. Crampton ( 1926a) called the posterior triangular sternal plate the spina-
sternum, but this plate is probably a fusion product of the furcasternum and the
spinasternum. The definitive furcospinasternum is thus similar, both in shape and origin,
to the corresponding plate in Embioptera (Fig. 51 ) .
In Stenopsocus (Fig. 96B) and Psocus (Fig. 94B) the presternum is largely
membranous, leaving a small sclerotized area surrounding the furcal bases. In Psocus
confratemus (Fig. 94B) and in Psocus venosus examined in this study, the lateral ends of
the definitive sternum form condyles, which articulate with the inner coxal margin on
each side. In Stenopsocus Badonnel ( 1934) recognized a rudimentary prothoracic spina.
The mesonotum: The anterior area of the mesonotum is greatly expanded and
bulged in Psocoptera. In Psocus venosus (Fig. 93A) the anterolateral scuta! suture is
represented by a ridge along the anterolateral scuta! margin. Two sutures meet at the
presumed anterior end of the anterolateral scuta! suture. Of these, the anterior suture
runs subvertically to reach the antecostal suture. This suture must be the prescutoscutal
suture, because in Plecoptera and Embioptera the suture tends to run obliquely proximo-
cephalad, often as far as the antecosta (Figs. 4 7 A, 50), and because the prescutoscutal
suture often meets the anterolateral scuta! suture laterally. The exaggerated length and
direction of this suture in Psocus (Fig. 93A) is due entirely to the greatly enlarged
condition of the anterior part of the mesonotum. The large lateral plate demarcated by
this suture is the prealar arm; it. extends ventrally and unites with the anterior margin of
the episternum as in some Plecoptera, and is the only prescutal area in the mesonotum.
Apparently in Psocus confraternus (Fig. 94A) and Stenopsocus (Fig. 96A) the prescuto-
scutal suture is represented by a narrow membranous cleft; hence the prescutal area
(prealar arm) is isolated. Badonnel ( 1934) called this plate the tergopleural sclerite,
and Cope (1940a) regarded it as part of the anepisternum. Thus, both authors failed
to recognize this plate as the prescutal prealar arm.
Another suture (Fig. 93A) arising from the anterior end of the anterolateral scuta!
suture extends posteriorly on the mesonotum; it is clearly the lateral parapsidal suture,
because it meets the anterolateral scuta! suture and the prescutoscutal suture (membrane)
as in other insects.
ANTEROLATERAL
SCUTAL SUTURE
PRESCUTUM
(PREALAR ARM)
AN EPISTERNUM----+.'---
A
.... ..
115
LATERAL PARAPSIDAL
SUTURE
ANTECOSTA
:..___ ___ INTERPLEURITE
LATERAL
PARAPSIDAL SUTURE
SCUTOSCUTELLAR
SUTURE
POSTEROLATERAL
SCUTAL SUTURE?
POSTNOTUM
' r - - - ~ - L - - SCUTOSCUTELLAR
SUTURE
SUBALARE
POSTEROLATERAL
SCUTAL SUTURE
FIG. 93. A, cephalic view of mesonotum in Psocus venosus (original); B, dorsal view of
thorax in Psocus confraternus (from Cope).
LATERAL
PARAPSIDAL SUTURE
PREALAR ARM
AN
PLEURAL ARM PIT
LATERAL CERVICAL
SCLERITE TROCHANTIN
PREEPISTERNUM
LATERAL PARAPSIDAL SUTURE
SUBALARE
POSTNOTUM

___,F/4'+-ANEPISTERNUM
:::J_@+-j--KATEPISTERNUM
TERAL CERVICAL
. SCLERITE
.t------PREEPISTERNUM
r------PLEURAL ARM
RCA
ANAPLEURAL f SPIRACLE
SUTURE-----f::.,---,/
PLEURAL
EPIMERON
ANEPISTERNUM
PLEURAL
B
FIG. 94. Thorax of Psocus confraternus (from Cope).
ANEPISTERNUM
FURCA
FURCAL PIT
FURCA
STERNUM 3
POSTCOXAL
BRIDGE
A, lateral view; B, ventral view.
No phragma is formed beneath the antecostal suture. In Psocus the greatly
swollen anterior wall of the mesonotum serves as the anterior point of attachment of the
dorsal longitudinal muscle (t 14), and the phragma is unnecessary.
In the posterior half of the mesonotum in Psocus (Fig. 93B) the incomplete scuto-
scutellar suture is present. The posterolateral scutal suture is presumably present in the
mesonotum. The posterior notal wing process lies immediately anterior to the axillary
cord. The postnotum is greatly developed ventrally and laterally and serves as the
MATSUDA: TiiE INSECT THORAX 227
. . . . -
. . . .
l'ST AXIL. SCL.
FIG. 95. Dorsal view of forewing base in Stenopsocus stigmaticus (from Badonnel).
second phragma. Laterally, the postnotum is united with the epimeron along the
posterior margin of the latter.
A small sclerite between the anterodorsal margin of the anepisternum and the
prealar arm is the prealar sclerite, which serves as a point of attachment for muscle
t-p 4. Badonnel ( 1934) called this sclerite the sclerite prealaire, and Cope ( 1940a)
misidentified it as the basalare. This sclerite is often associated with the tegula in other
insects (e.g., subtegula in Leipidoptera). In Psocoptera, however, the tegula is absent.
Axillary sclerites of the forewing: In Stenopsocus Badonnel ( 1934) described the
axillary sclerites as follows (Fig. 9 5) : The anterior tip of the first axillary sclerite lies
near the proximal extremity of the Sc vein; its proximal (internal) margin is connected
with the scuta! lateral margin, including the anterior notal wing process; and its distal
margin rests on the proximal margin of the second axillary sclerite.
The second axillary sclerite has its dorsal surface continuous with the first axillary
sclerite; its anterior end rests on the base of the R vein; its median process a on the
posterior margin is in contact with the anterior head (process) of the third axillary
sclerite; laterally, the distal posterior end is articulated with an arc d' that is in turn
connected with the third axillary sclerite; internally the second axillary sclerite carries a
process b that distally appears to lie free in the membrane.
The third axillary sclerite, besides its anterior connection with a of the second
axillary sclerite, has its posterior base connected to the edge of the wing through an arc, e.
The third axillary sclerite is also connected with the proximal end of the A vein by
another process d, which is continuous with its branch d', which in turn articulates with
the second axillary sclerite. The posteroventral tip of the third axillary sclerite is
connected with the posterior subalar process by a ligament, and the dorsal crest of the,
third axillary sclerite lies on the posterior notal process when at rest.
The mesopleuron: The mesopleuron is divided into the anterior episternum and the
posterior epimeron by the nearly straight pleural suture, which carries no distinct pleural
arm internally. Both in Stenopsocus and Psocus the paracoxal suture is absent, although
a suture above the trochantin in Stenopsocus (Fig. 96B) may possibly represent the
paracoxal suture. The anapleural suture is ridged internally and reaches the pleural
228
MEMOIRS OF TilE ENTOMOLOGICAL SOCIETY OF CANADA
suture posteriorly. The anepisternum is relatively large. The externally isolated
basalare is absent. However, an internal apophysis arises from beneath the dorsal
margin of the anepisternum, where the basalare is normally separated (Fig. 94A).
The episternal area ventral (proximal) to the anapleural ridge (Figs. 94A, 96A, B)
is the fused preepisterno-katepisternum. In Psocus (Fig. 94B) this area is separated
from the basisternum by a membranous cleft, which probably represents the border
between the pleuron and the sternum; in Stenopsocus (Fig. 96B) the corresponding area
appears to be represented by the pleurosternal suture, although it may simply be the
external indication of the sternal invagination. In both genera the clearly differentiated
trochantin is continuous with the preepisterno-katepisternum, and apically articulates with
the coxal margin.
The epimeron is well retained and meets the lateral extension of the postnotum along
a suture. The subalare lies near the dorsal end of the epimeron, and c o n m ~ c t s with the
third axillary sclerite in Stenopsocus. In Stenopsocus a well-sclerotized postcoxal bridge
is formed along the posterior coxal margin.
The mesosternum: In Psocus (Fig. 94B) the basisternal area has a median longi-
tudinal suture, which Cope (1940a) called the discrimen; invagination occurs along this
suture, as in holometabolous insects. The small interpleurite below the mesothoracic
spiracle may represent the remnant of the presternum. In Stenopsocus the anterior
margin of the sternal area is secondarily ridged for reinforcement (prepectus) and isnot
the presternum. The furcal bases lie very close to the median longitudinal axis. The
area posterior to the furcal bases in Psocus is continuous with the metasternum. The
mesothoracic spina is absent.
The mesothoracic pleurosternal region in Psocoptera clearly resembles that in
typical holometabolous insects in having a median longitudinal invagination (the
discrimen), in the approximation of leg bases toward the median longitudinal axis, and
in the presence of a well-developed anapleural suture. However, the main difference is
in the absence in this order of the ventral condyle arising from the furcal base, which in
holometabolous orders articulates with the inner coxal margin. The function of the
anterior pleurocoxal articulation, which lies farther proximally (ventrally) than that in
other orders, is probably functionally similar to the ventral coxal articulation in holo-
metabolous orders.
The metanotum: In Psocus (Fig. 93B) the median anterior margin of the
metanotum is lobately produced and becomes continuous with the dorsal part of the
mesothoracic postnotum, without forming an infolding between them. The second
phragma is therefore merely a ventral extension of the mesothoracic postnotum. The
anterior area of the metanotum on each side of the median production is strongly, sub-
vertical! y bent ventrolaterally; the prealar arm and the lateral parapsidal suture are
absent. As in the mesonotum, the scutoscutellar suture is present. The postnotum
forms the third phragma posteriorly, and laterally the postalar bridge by union with
the epimeron.
The metapleuron: In the episternal region of Psocus (Fig. 94A) and Stenopsocus
(Fig. 96A) the distinct anapleural suture is absent, and the dorsal and ventral sclerotized
areas are separated by an intervening membranous area. The basalare is represented by
the intern! apophysis beneath the dorsal margin of the anepisternum. In Psocus venosus
(examined) the dorsolateral angle of the ventral sci erotized plate is united with the
posterolateral angle of the mesothoracic lateral postnotum, where the latter meets the
epimeron. This ventral episternal plate in Psocus and Stenopsocus is probably ex-
clusively katepisternal; the preepistemal area has evidently become lost, accompanied by
approximation of the middle and hind leg bases. Both in Stenopsocus and in Psocus the
subalare is present above the epimeron; the narrow postcoxal bridge encircles the
posterior coxal margin arrd reaches the furcasternum ventrally; and the slender trochantin
articulates with the anterior coxal margin.
LATERAL PARAPSIDAL
SUTU
PLEURAL
SUTURE
EP
ANAPLEURAL
RIDGE
LATERAL
CERVICAL
SCLERITE
COXA
CORPOTENTORIUM

ANEPISTE
229
PREALAR SCLERITE
ALARE
POSTNOTUM
EPIMERON
LEURAL RIDGE
AN EPISTERNUM
TROCHANTIN
PREEPIST ERNUM
+ KATEPISTERNUM
RIDGE

,.,__.,. ..
TROCHANTIN -----t---'nii;;:-
PLEURAL

POST COXAL
BRIDGE
'YI---i---PLEURAL RIDGE
r----c:DIMERON
POSTNOTUM
+PHRAGMA
r---RASISTERNUM

POSTNOTUM
FURCASTERNUM
Fro. 96. Thorax of Stenopsocus stigmatic us (from Badonnel). A, lateral view; B, ventral
view.
230
The metasternum: In Psocus (Fig. 94B) and Stenopsocus (Fig. 96B) the meta-
sternum is represented by a narrow longitudinal strip that is continuous from the
mesosternum. It branches behind the median furcal pit and encircles the posterior
coxal margin. Internally, the low median longitudinal ridge in the metasternum is
continuous with that of the mesosternum. Evidently, the anterior part of the meta-
sternum has, along with the mesospina, become lost, thereby completely obliterating the
original ventral segmental border between the mesothorax and the metathorax.
The anatomical basis of flight: Badonnel ( 1934) explained the mechanism of the
movement of the forewing as follows:
Flexion of the wing: When the extended forewing flexes to resume a folded position,
the posterior membranous area of the wing, which connects the mesonotum with the anal
field, sinks into a groove in which chitinous e and d (Fig. 95) come to lie; at the same
time the third axillary sclerite becomes inclined forward and inward, revolving around its
ventral tip, which is connected with the subalare; the R vein, turning around its proximal
articulation, comes to lie along the body; and finally the wings come to be applied against
the lateral walls of the body.
The forewing is flexed when the third axillary sclerite is depressed by the contraction
of muscle W 1 ( t-p 14), which is inserted on the third axillary sclerite. Acceleration of
the rotation of the R vein base is probably provided by the action of d' on the second
axillary sclerite, and perhaps also by the contact of the third axillary sclerite with the
apophyses, a and b, of the second axillary sclerite.
Extension of the wing: In many insects wings are extended by the contraction of the
subalar and the basalar muscles. In these insects the second axillary sclerite is connected
with the subalare. Therefore, when the subalare is pulled by the subalar-coxal muscle
(t-cx 8), the second axillary sclerite is depressed; the depression of the second axillary
sclerite in turn causes the extension of wings. In psocids, however, the subalare is not
connected with the second axillary sclerite, but with the third axillary sclerite. The
depression of the third axillary sclerite would lower the wing instead. The extension
mechanisms in psocids, therefore, requires a different explanation.
The psocids jump before they fly, by a vigorous action of the posterior legs. The
wings then expand so rapidly that the wing extension follows the wing elevation instantly.
The extension is therefore only a special phase of elevation, and is produced by depressing
the tergal base of the wing. This depression, which is transmitted to the first and second
axillary sclerites, turns the R vein in such a way as to bring its dorsal surface upward,
while the heads of the Sc and R veins move circularly from the front towards the tergal
base of the wing; the C vein lies perpendicular to the sagittal plane, and the wing remains
extended. For this action of extension and elevation of the wing, muscle W 3 ( t-p 11),
which is inserted on the first axillary sclerite, is mainly responsible.
As is clear from the foregoing discussion, depression of the tergal lateral margin of
the wing causes the lowering of the dorsal base of the wing, and hence elevation of the
wing. Conversely, depression of the pleural margin or elevation of the tergal margin
causes the lowering of the wing. The elevator muscles are: V (t-p 8), W 3 (t-p 11),
W 2 (t-p 10), I (t-cx 7), K (t-p 6), K' (t-ti 2), and DO (t 12). The depressor muscles
are E (p-cx 4), F (t-cx 8), F' (t-cx 8?), and DL (t 14).
The adult musculature: Badonnel ( 1934) and Maki ( 1938) have studied the
thoracic musculature in Stenopsocus stigmaticus and in Psocus tokyoensis, respectively.
The results of these studies are shown in Table XI. The sternocoxal muscles described
by Maki are omitted, because their exact positions are difficult to discern from Maki's
drawings. Also, Ribaga's (1901) sketchy description of thoracic muscles in Trichopsocus
dalii is not referred to here.
The anterior tergocoxal muscles are conspicuously scarce in this order, whereas the
muscles occurring in the pleuron are relatively abundant. The well-preserved sterno-
coxal muscles represent a primitive feature of the thoracic musculature in this order.
op-p 3 is peculiar to Psocoptera.
TABLE XI
Thoracic musculature in Psocoptera
Psocus Stenopsocus
(Maki 1938) (Badonnel 1934) Muscle designations Remarks
2 M. protergo-occ. obi. op-t 1
1
DPr cv(d)-t 1
'DPo op-t 3
DVA op-cv 1
5 DVL 1 op-cv 3
9 LL op-p 2
DVL 2 op-p 3
6 T,S t-s(cv) 1
7 t--;;v 1
13 X1 cv-cx 1(x)
3 Lv 1 s 2
19 (II) DO t 12
44 (III) DO
1 DPr, DPo t 14
18 DL
43 DL
23 (I I) t-p 3
4 7 (III)
p
25 (II) z t-p 4
49 (III) z
K (II)
K (III)
t-p 6
10 (I) G t-p? 1
24 (II)
48 (III)
v t-p 8
27 (II) W2 t-p 10
26 (II) W3 t-p 11
so (Ill)
28 (II) W1 t-p 14
51 (III) W1
8 u t-s 1
22 u
46 u
12 (I) L t-ti(cx) 2
29 (II) K'
14 c t-cx 6
31 c
54 c
14 c t-cx 7
32 I
55 I
33 (I I) F t--;;x 8
56 (III) F
34 (I I) F' t--;;x 8?
2
11 (I)
p-s 1
h 2 (I-II) p-s 2 3
h 3 (II-IH)
p-s 3 3
U (I-II) p-s 5? 4
U (II-III)
U (lil-IA)
37 (II)
e p-ti(cx) 1
59 (III)
e
TABLE XI (Continued)
Psocus StenoPsocus
(Maki 1938) (Badonnel 1934) Muscle designations Remarks
E p-cx 4
38 E
60 E
16 e p-cx 5
36 e'
58 e'
52 ? H (II-III) p-cx 8 5
LVM 2(!-ll) sll
21 (I-II) LVI 3 (II-III) s 12
20 (I-II) LVL 2 s 13
45 (II-III) LVL 3
4(1) d 1, d 2 s 14
LVI 2 (I-II) s 15
lv (III-2A) s 20
X 2 (I-II) s-ex 1
X 3 (II-III)
y (I) s-ex 2
y (II)
J(l)
s-ex 4
J (I I) s-ex 6? 6
J (I !I)
17 A t-tr 1
39 A
61 A
A' p-tr 2
40 A'
62 A'
41 (II) a s-tr 1
63 (III) a
REMARKS (Table XI)
( 1) These muscles are prothoracic tergopleural muscles, which are not homolo-
gized.
(2) Arising from the subalare, these muscles are attached on the remoter apodeme
on the posterolateral coxal margin.
(3) Badonnel ( 1934) thought that h 3 was serially homologous with h 2, but h 3
is obviously inserted on the furca, and h 2 is on the spina.
(4) U in the pterothorax of Stenopsocus arises from the ventral margin of the
phragma.
(5) In fig. 17 of Maki (1938) 52 appears to represent p-cx 8, although his
description does not agree with his figure.
(6) J in Stenopsocus appears to be s-ex 6 or 3, not s-ex 4.
THE MALLOPHAGA
Of the genera referred to in the following discussion, Tetrophthalmus (Cope 1941),
Trimenopon, and Myrsidea (Mayer 1954) belong to the suborder Amblycera; Esthiop-
terum (Cope 1940b), Bovicola, and Columbicola (Mayer 1954) belong to the suborder
Ischnocera.
The neck region: The more generalized lateral cervical sclerite in Columbicola
(Fig. 97C) articulates with a small anterior plate that is presumably the detached post-
occipital process; posteriorly the sclerite is connected with the episternum. Thus, in
Columbicola the lateral cervical sclerite is not very different from that in Psocoptera.
IIi. Esthiopterum (Fig. 97 A) the lateral cervical sclerite is represented by a small
sclerite that unites the head and the precoxal bridge of the propleuron. Peculiarly, in
this genus, an additional dorsolateral cervical sclerite is present. In Bovicola the lateral
cervical sclerite is absent.
In Tetrophthalmus (Fig. 98A), among Amblycera, the lateroventrally continuous
cervical sclerite appears to represent a fusion of the lateral cervical sclerite and the
dorsolateral cervical sclerite, which are present in Esthiopterum (Fig. 97 A). In
Trimenopon (Fig. 98E) and Myrsidea the lateral cervical sclerite is a small single plate,
which connects the postoccipital ridge with the anterior end of the dorsally displaced
pleural ridge. In these genera the sclerotized episternum and the epimeron have been
lost, and the lateral cervical sclerite has acquired a direct connection with the pleural ridge.
The pronotum: In Columbicola (Fig. 97C) and Esthiopterum (Fig. 97 A) the
pronotum is a simple plate without subdivision, and it is clearly separated laterally from
the pleuron. In Bovicola (Fig. 97B) a subtriangular plate is separated from the pronotal
anterior margin, and the pronotum is laterally indistinguishably fused with the
p_ropleuron.
In amblycerous Mallophaga the process of modification of the pronotum appears to
have been quite different. In Trimenopon (Fig. 98E), Myrsidea, and Tetrophthalmus
(Fig. 98A), the pronotum has a median transverse ridge; and the ridge in Trimenopon
is crossed by a longitudinal ridge.
In Tetrophthalmus (Fig. 98A) the pronotum is simply laterally fused with the pro-
pleuron, and the pleural ridge is bent caudad from a point near the presumed dorsal
anterior angle of the propleuron. In Myrsidea the pleural ridge lies obliquely caudad
and meets the transverse ridge of the pronotum. In Trimenopon (Fig. 98E) the dorsally
displaced pleural ridge demarcates the lateral edge of the definitive pronotum, and its
connection with the coxal margin has been lost. It is by its association with the furcal
end and with the lateral cervical sclerite (as in Trimenopon) that this apparent pronotal
lateral ridge can still be homologized with the pleural ridge. This pleural ridge appears
at first glance to be serially homologous with a lateral ridge on the metanotum of the
same species.
Mayer ( 1954) called the narrow area cut by the pleural ridge in Trimenopon and
Myrsidea the paranotum. This paranotum naturally cannot be homologized with the
paranotum in lower insects. The area may represent the proepimeron, since it is
demarcated by the pleural ridge. In Amblycera an unpaired sclerite (prothoracic post-
notum of Mayer) is present between the pronotum and the mesonotum.
The propleuron and prosternum: In Columbicola (Fig. 97C) the well-retained
episternum is a simple plate, being undivided by sutures. Its anterior articulation with
the coxa is either pleurocoxal or trochantinocoxal. The epimeron extends ventrally and
forms a condyle that articulates with the coxa. The furcal arm is continuous with the
posterior dorsal angle of the propleuron. In Bovicola (Fig. 97B) the strongly developed
pleural ridge represents the anterior margin of the propleuron, and its ventral end is
articulated with the coxal margin. A process from the posterior ventral margin of the
coxa fits into a socket on the ventral anterior margin of the strongly ridged posterior
margin of the epimeron, which merges with the sternum behind the coxa. A similar
postcoxal bridge is formed in Esthiopterum and Columbicola (Fig. 97C), and a process
from this area articulates with the ventral posterior margin of the coxa.
Among Amblycera; Tetrophthalmus (Fig. 98A) has a more generalized propleuron
in which the sclerotized episterum and epirneron are well retained. The furcal arm is
joined with the ventral part of the pleural ridge; the ventral articulation with the coxa
is absent. Cope ( 1941) showed that in this genus the first thoracic spiracle lies in tbe
proepimeron. In Myrsidea and Trimenopon the propleuron is nearly membranous. In
Myrsidea the pleural ridge articulates with the coxa, but the ventral articulation is absent;
234 MEMOffiS OF THE ENTOMOLOGICAL SOCIETY OF CANADA
I
PROEPISTERNUM
POSTCOXALE SPIRACLE
PROEPIMERON ----
PLEURAL------,;
RIDGE
PROFURCA -------;;'
PRONOTUM
POSTOCCIPITAL
RIDGE -----.::

POSTOCC.
PROC
A
B
LATERAL
CERVICAL SCLERITE
EPIMERON C
PLEURAL
RIDGE
FURCA
MESONOTUM
+METANOTUM
MESOSTERNUM
+METASTERNUM
PLEURAL
RIDGE
MESONOTUM
+METANOTUM
--'ffi--- --EPISTERNUM

FIG. 97. A, lateral view of thorax in Esthiopterum diomedeae (from Cope); B, lateral view
of thorax in Bovicola caprae (from Mayer); C, lateral view of thorax in Columbicola columbae
(from Mayer).
in Trimenopon (Fig. 98E) the reverse is true, there being only a ventral articulation
with the coxa.
In Trimenopon (Fig. 98D, E) the ventral side of the prothorax is provided with a
quadrangular sclerotization surrounding the median membranous area; a small median
process on its posterior margin fits into a socket formed on the anterior margin of the
anterior transverse bridge of the mesosternum, and a process on the lateral margin
PRONOTUM -----....'"-

EPIMERON
Cor!----METANOTUM

;J.- RIDGE
EPISTERNUM
:l>;----!ti"IMERON
PROFURCA
METAFURCA

MESOSl FURCA
MESONOTUM-
METANOTUM
NOTAL RIDGE
PROSTERNUM
LATERAL
CERVICAL
SCLERITE
METASTER-
NUM ---F""""'"-
A
MESOCOXA
B
c
MESOSTERNUM
235
FIG. 98. A, lateral view of thorax in Tetrophthalmus sp. (from Mayer); B, dorsal view of
meso- and meta-thorax in Trimenopon hispidum (from Mayer); C, ventral view in Tetroph-
thalmus sp. (from Cope); D, ventral view in Trimenopon hispidum (from Mayer); E, lateral
view of thorax in Trimenopon hispidum (from Mayer).
articulates with the inner coxal margin. The prothoracic furca arises from the lateral
corner of the anterior bridge of this definitive mesosterna! sclerotization (Fig. 98E). In
Myrsidea (Fig. 99B) the sternal sc!erotization is without a median membranization, and
it has a process on its median posterior end. In Tetrophthalmus (Fig. 98C) the furcal
bases lie more proximad.
In ischnocerous genera the ventral surface between the coxae is membranous. In
Bovicola and Esthiopterum (Fig. 99A) the furcal pit lies on the postcoxo-sternal
sclerotization.
The meso- and meta-notum: The division of the tergum into the prescutal, scuta!,
and scutellar areas is absent in the meso- and meta-notum of Mallophaga. In Tetroph-
thalmus (Fig. 98A) the meso- and meta-notum are distinctly separated by a membrane.
In Trimenopon (Fig. 98B) the small mesonotum is strongly sclerotized and has a hollow,
median apodeme which invaginates into the metanotum. The border between the
metanotum and the metapleuron is strongly ridged. In Trimenopon (Fig. 98E) Mayer
(1954) recognized the metathoracic prescutal arm (Tergalarm of Mayer), which is,
however, probably a secondary formation. In Myrsidea a sclerotized mesonotal bar
extends ventrally along the anterior margin of the mesopleuron to become continuous
with the base of the sclerotized sternal bar. In all of these three genera of Amblycera,
the mesonotum is smaller than the metanotum. In Ischnocera the mesonotum and
metanotum are completely fused. In Columbicola (Fig. 97C) and Esthiopterum a
median longitudinal ridge is present on this fused mesometanotum.
The mesopleuron: In Tetrophthalmus (Fig. 98A) the mesopleuron is fused with the
mesonotum. The pleural ridge is a strongly sclerotized bar which forms a condyle for
articulation with the coxa. The episternum is retained but without division. The
anterior articulation with the coxa probably represents the trochantinocoxal articulation;
in Myrsidea also, the anterior articulation probably represents the trochantiriocoxal
articulation. In Trimerwpon (Fig. 98D) the ventral articulation is with the longitudinal
sclerotized area along the inner margin of the coxa, and it does not seem homologous
with the anterior articulation in Tetrophthalmus and Myrsidea. Both in Trimenopon
(Fig. 98E) and in Myrsidea the pleural ridge becomes confluent with the transverse ridge
of the mesonotum. In all of these three genera the mesopleuron is distinctly retained.
In Columbicola and Esthiopterum a conspicuous process arises from the anterolateral
angle of the mesonotum (not shown in Fig. 97A). Mayer (1954) thought this process
was the pleural apophysis (arm). In position, however, the process appears to corres-
pond to the prescutal arm that occurs in the corresponding position in the metanotum
of Trimenopon (Fig. 98E). In Bovicola the pleural arm and the pleural ridge are absent;
they are apparently replaced functionally by the corresponding pleural arm and ridge of
the metathorax (Fig. 97B). The pleural articulation with the coxa is present in all three
genera of Ischnocera. In Bovicola (Fig. 97B) and Columbicola (Fig. 97C) a process
on the posterior margin of the definitive propleuron articulates with a socket on the
ventral anterior margin of the mesocoxa. In Esthiopterum, however, the corresponding
process is formed on an isolated plate behind the propleuron, and this condition is
probably more primitive than that in Bovicola and in Columbicola. If the isolated plate
in Esthiopterum becomes fused with the propleuron, the condition present in Bovicola
and Columbicola would result.
The mesosternum: In Tetrophthalmus, as Cope ( 1941) showed (Fig. 98C), the
episternal sclerotization extends ventrally along the inner margin of the coxa, and the
furcal pit lies at its caudal end. In Myrsidea (Fig. 99B) the sclerotized bar from the
pleuron is continuous along the inner margin of the coxa, and merges with the median
ventral sclerotization which unquestionably represents the sternum. In Trimenopon
(Fig. 98D) the sclerotization along the inner margin of the coxa is continuous with the
anterior transverse sclerotization.
In Esthiopterum (Fig. 99A) a small lateral plate along the inner coxal margin bears
the furcal pit; in Columbicola this lateral plate is continuous with the episternum; in
Bovicola (Fig. 98B) the mesosternum is continuous with the presternum. In both
Amblycera and Ischnocera the furcal arm is not fused with the pleural arm or ridge.
The metapleuron: In Tetrophthalmus (Fig. 98A) and Trimenopon (Fig. 98E) the
articulation of the pleural ridge with the coxa is dorsal or posterodorsal, and in Myrsidea
posteroventral. In Myrsidea the anterior articulation lies on the anterodorsal coxal
margin; the condyle for this articulation is formed on a narrow sclerite with a strong
anterior ridge that Mayer called the trochantin (Fig. 99B). The corresponding anterior
articulation with the coxa in Trimenopon and Tetrophthalmus lies more ventrad than
in Myrsidea.
ROSTERNUM
FURCAL PIT
MESOFURCA
MESOSTERNUM METASTERNUM
+ METASTERNUM
B
METAFURCA
FIG. 99. A, ventral view of Esthiopterum diomedeae (from Cope); B, ventral iew of
thorax in Myrsidea cornicis (from Mayer).
In Esthiopterum (Fig. 97A) the pleural ridge runs horizontally forward from the
dorsal point of articulation with the coxa. The large metapleuron below the ridge is
therefore the episternum. The anterior articulation with the coxa, which Cope ( 1940b)
did not show but Mayer ( 1954) showed, may be the trochantinocoxal articulation, as in
Amblycera. As Cope ( 1940b) showed, the second thoracic spiracle lies on the anterior
third of the dorsal margin of the pleuron (i.e. pleural ridge). In Columbicola (Fig. 97C)
the metapleuron is somewhat similar to that of Esthiopterum. According to Mayer
( 1954), however, a distinct pleural ridge is absent. The second thoracic spiracle lies on
the closed membrane between the metanotum and the metapleuron. In Bovicola
(Fig. 97B) the greatly developed pleural ridge extends into the tergal area, and the
anterior articulation with the coxa is absent.
The metasternum: In Tetrophthalmus (Fig. 98C) the episternal sclerotization
extends ventrally along the inner coxal margin and the furcal pit lies on this extension.
A triangular, anteromedian sternal plate lies between the lateral sclerotizations. In
Myrsidea (Fig. 5'9B) the median subtriangular sclerotization is much more extensive than
in Tetrophthalmus, and in Trimenopon (Fig. 98D) the lateral and median sclerotizations
have become indistinguishably fused.
In Columbicola and Esthiopterum (Fig. 99A) the fused, median mesometasternal
plate is separated from the lateral sclerotization continuous from the episternum. The
furca arises from the ventral margin of the latter. In Bovicola the sternal area is not
sclerotized.
The adult musculature: Mayer ( 1954) described the thoracic muscles of
Trimenopon hispidum, Myrsidea cornicis, Bovicola caprae, and Columbicola columbae.
They are shown in Table XII.
It is difficult to homologize these muscles with those in other orders, because of
shifts in points of origin and insertion. The most pronounced shifts have evidently
resulted from the great reduction of the rnesothorax, and often the mesothoracic muscles
have acquired new poin!5 of origin on the metathorax. The transsegmental distributions
of some other muscles in this order are also secondary.
Generally, the amblycerous genera have retained more muscles than the ischnocerous
genera. It is interesting to note that muscle p-s 5 is well retained, a feature that is also
shared by Psocoptera. The relative abundance of ventral muscles also indicates a close
relationship of this order to Psocoptera, although some of the muscles have shifted their
points of attachment.
238 MEMOIRS OF
THE ENTOMOLOGICAL SOCIETY OF CANADA
TABLE XII
Thoracic musculature in Mallophaga
Trimenopon Myrsidea Bovicola Columbicola Muscle
(Mayer 1954) (Mayer 1954) (Mayer 1954) (Mayer 1954) designations Remarks
I dim 2 I dim 2 op-t 1
I dim 1 I dim 1 I dim 1 I dim 1 op-t 3 or ev(d)-t 1
0, I lm 1
0, I lm 1 0, I lm 1 0, I lm ? op-p 1, op-s 2 1
0, I lm 2
0, I lm 2 0, I lm 2 t-cv 1 or 2 2
0, I trm ev-ex(x) 1
0, I vim 1 0, I vim s 1 or 2
I dim 4
I dim 4 I dim 2 I dim 3 t 12
III dlm 2 III dim 2 III dim
I dlm 2, 3 I dim 3 t 13
III dim 1
III dim 1 III dim
I tpm 1 I tpm I tpm I tpm t-p? 3
I tpm 2, 3, 4
t-p? 3
II tpm
t-p? 3
III tpm
III tpm t-p? 3
I, II ism 1 t-s 1 4
II, III ism 1
III tfm III tfm
I dvm 1 I dvm 1 I dvm 1 I dvm 1 t-ti(ex) 2 5
II dvm 1 II dvm 1 II dvm 1 II dvm 2(pars)
III dvm 1? III dvm 2 or dvm 1
II dvm 2 II dvm 1? t-ti(ex) 3 5
I dvm 2 I dvm 2 I dvm 2 I dvm 2 t-cx 6 or 7 5
II dvm 3 II dvm 2 II dvm 2 II dvm 2
Ill dvm 2 III dvm 1 III dvm 1 III dvm
II pm 2 II pro 2 t-cx 8?
III pro 1 III pm 2?
III pfm III pfm p-s 1
I, II ism I, II ism 2 I, II ism p-s 5
II, III ism II, Ill ism 2 II, III ism II, III ism
I pm 1 I pm 1 I pro 1 I pm 1? p-ti(ex) 2 or 3
II pm 1 II pm 1
Ill pm 1 III pm
I, II vim 1 I, II vim I, II vim I, II vim s 13
II, III vim 1 Il,IIIvlm1 II, III vim
III, Ia vim 1 III,Iavlm
II, Ia vim II, Ia vim s 15 ? or p-s 10 6
I trm I trm I trm s 16
II trm 1 II trm 1 II trm 1 II trm 1
III trm 1 III trm 1 III trm
Ill trm 2 III trm 2 s 20? 7
Ibm 2 Ibm 1 Ibm s-ex 3
III bm 2
II trm 2 II trm 2 II trm 2 II trm 2 s-ex 4 ? 8
Ibm 1 s-ex 5
II bm
III bm 1
II, III vim (ex) II, III vim II, III vim II, III vim s-ex 7
(ex) 1 (ex) 1 (ex) 1
II, III vim II, Ill vim s-ex 7 ? 9
(ex) 2 (ex) 2
I dvm 3 I dvm 3 t-tr 1
II dvm 5 II dvm 3 II dvm 3 II dvm 3
III dvm 3 III dvm 3 III dvm 2
TABLE XII (Continued)
Myrsidea Bovicola Columbicola Muscle
(Mayer 1954) (Mayer 1954) (Mayer 1954) (Mayer 1954) designations Remarks
I pm 2
I pm 2 I pm 2 I pm 2 p-tr 1
III pm 3
Ibm 3
Ibm 2
Ibm s-tr 1
II, III vlm
II, III vlm s tr 1 ?
(ex) 2
(ex) 3
10
REMARKS (Table XII)
( 1) 0, I lm in Columbicola is inserted ventrally on the coxal margin; 0, I lm 1 in
Bovicola is inserted on the pleural ridge; 0, I lm 1 in Trimenopon and Myrsidea is inserted
on the dorsally extended furca.
(2) 0, I lm 2 in Bovicola is inserted anteriorly on the postoccipital ridge. This
is probably due to a secondary shift that has accompanied the loss of the lateral cervical
sclerite.
( 3) I tpm muscles cannot be homologized.
( 4) II, III ism 1 in Myrsidea arises from the middle of the metanotum. Presum-
ably this is due to shift backward of the origin of the muscles. III tfm in Myrsidea
arises from near the lateral margin of the metanotum.
(5) Dorsal and ventral points of attachment of these muscles vary considerably
in different genera. Homologizations are based on the positions of muscles on the coxal
margin relative to the points of pleurocoxal articulation. The homologies proposed
here differ considerably from those proposed by Mayer ( 1954).
( 6) These muscles extend between the mesofurca and the first abdominal
sternum, and may be p-s 10. Usually s 15 extends between the profurca and the
second spina.
(7) III trm 2 in Trimenopon extends from the metafurca to the membranous
fold between the metasternum and the first abdominal segment; the same in Myrsidea
stretches the area between the metafurca and the median line of the metasternum.
( 8) II trm 2 connects transversely the mesothoracic coxae. This muscle is
probably the derivative of s-ex 4. The loss of the second spina has apparently resulted
in the direct connection of mesocoxae by this muscle.
(9) These muscles connect the basal part of the mesofurca and the posterior
margin of the metacoxa.
( 10) These muscles arise from the mesofurca and are inserted on the trochanteral
apodeme of the hind leg. The derivation by these muscles is quite unclear.
THE ANOPLURA
Ferris (1951) studied the thorax of Solenopotes capillatus, Pediculus humanus, and
Haematopinus suis. In all three species the tergum, the pleuron, and the sternum of
three thoracic segments are continuously fused and largely membranous. Ferris used
the internal phragmata, which clearly correspond to ridges in Mallophaga, as
the landmarks for determination of segmental boundaries, as well as for determination
of original divisions of a segment into the tergum, the pleuron, and the sternum. As
Ferris showed, the phragma (ridge) distally (laterally) articulates with the coxal margin
of all three segments (Fig. 100A, B). He, therefore, homologized it with the pleural
ridge in other insects.
In Solenopotes (Fig. 100A) the phragma (ridge) extends dorsally to meet its
counterpart from the other side on the median longitudinal axis of the body. Since,
according to Ferris, these ridges are homologous with the pleural ridge, Ferris had to
240
NOTAL RIDGE
TERGUM----
SPIRACLE-----
-------COXA 1
PLEURAL RIDGE
(PHRAGMA)
----PROFURCA
STERNUM
FIG. 100. A, dorsal view of thorax in Solenopotes capillatus (from Ferris); B, dorsal view
of thorax in Haematopinus suis (from Ferris); C, ventral view of thorax in Haematopinus
suis (from Ferris).
conclude that the terga have been almost entirely suppressed by this union of subcoxal
elements. In Solenopotes, therefore, the pronotum is represented by a narrow median
area, and the metanotum by a small median pit. Ferris applied the same interpretation
to other Anoplura (Fig. lOOB), and thought that the areas surrounding the spiracle were
isolated notal islands.
Comparing the thorax in Anoplura with the thorax in Mallophaga, it is immediately
clear that in Anoplura the pleural ridge can become continuous with the notal ridge, as in
Myrsidea, Trimenopon (Fig. 98E) and Bovicola (Fig. 97B). The dorsolaterally con-
tinuous ridges in Anoplura, therefore, must be homologous with the dorsolaterally
continuous ridges in Mallophaga. Thus, the phragma (ridge) in question cannot be
used as a landmark in determining the division of a segment.
A more reliable landmark in determining the border between the pleuron and the
tergum in Anoplura is the position of the spiracle. In Mallophaga, as already seen, the
spiracle lies on the dorsal part of the pleuron. It is safe to assume that in Anoplura also
the position of the spiracle roughly separates the tergal region from the pleural region.
Judging from the positions of the spiracles, then, the dorsal surface of the thorax in
Anoplura is largely the tergum.
In Mallophaga the dorsolateral ridges do not necessarily correspond to segmental
borders, and quite frequently the mesonotum and metanotum are fused. The apparent
division of the thorax into three segments by ridges in Anoplura is, therefore, secondary;
it is safe to suppose that in Anoplura the segmental borders have been lost.
The ventral side of the thorax is normally membranous, except for the single median
sternal sclerotization. In Haematopinus (Fig. lOOC) the sternal sclerotization has a
sternal furca on each anterolateral angle, and it is fused with the prothoracic pleural arm.
The adult musculature: Ferris (1951) illustrated the thoracic muscles in Haemato-
pinus, but his descriptions lack sufficient detail; homologies of the thoracic muscles in this
order, therefore, cannot yet be determined.
THE THYSANOPTERA
In the following discussion, reference is made almost exclusively to an anatomical
work on the thorax of Thysanoptera by Mickoleit (1961), since other works (Melis
1935a; Doeksen 1941; Jones 1954) are not comparable with it in the presentation of
anatomical details.
The neck region: In Thysanoptera only the single-lobed lateral cervical sclerite
occurs in the membranous neck region (Fig. lOlA, B). Anteriorly, the sclerite
articulates with a process that arises from the postoccipital ridge (Kehlsporn of
Mickoleit). Posteriorly, it articulates with the ventral part of the episternum (trochantin
of Mickoleit), which in Terebrantia is the reduced preepisternokatepisternal plate, as will
be discussed below. The lateral cervical sclerite is, therefore, presternal in origin. In
Phloeothrips it appears to connect also with the sternal sclerotizafion (my examination).
The pronotum and propleuron: In Tubulifera, including Phloeothrips and Haplo-
thrips (Fig. lOlA) the posterolater,'S). angle of the pronotum is excavated, and this area is
occupied by the propleuron. In Phloeothrips and Haplothrips the propleuron and the
pronotum are distinct from each other. In Phloeothrips (Fig. lOlA) the pieural ridge
dorsally reaches the posterolateral margin of the pronotum, thereby demarcating
posteriorly the narrow epimeron. The pleural arm lies at the ventral end of the pleural
ridge. The sclerotized episternum is divided into the anterior and posterior areas by an
intervening desclerotizeq area, which apparently allows smooth movement of the coxa.
The horizontal line that separates the anterior sclerotized area into the dorsal and ventral
plates is probably the anapleural suture, judging from a similar relative position of the
anapleural suture in the mesothorax. Thus, in Phloeothrips the anepisternum consists
of two plates which lie along the pronotal lateral margin, as labeled in Fig. lOlA; in
Aeolothrips (Fig. lOlB) and Thrips (Fig. 101E), which belong to Terebrantia, the
anepisternum is a single plate.
242
AN EPISTERNUM
PRONOTUM
AN EPISTERNUM
LATERAL
CERVICAL
SCLERITE
LATERAL
POSTNOTUM
PLEURAL SUTURE
L--ANIEPISTERN UM
COXA 1
PRESCUTUM
FURCASTER-
NUM l'ST PHRAGMA
--4.o+--- PROCOXA
B
SPINASTERNUM
TEGULA?
SPIRACLE
ANEPISTERNUM
EPIMERON
FURCASTERNUM
PRO COXA
D
ANAPLEURAL
SUTURE
PREEPISTERNUM
+ KATEPISTERNUM
OTUM
ACROTERGITE
c
EPIMERON
243
In Phloeothrips (Fig. lOlA) the area ventral to the anapleural suture is the fused
preepisternurri and katepisternurn (Fig. lOlA). In Aeolothrips (Fig. 101B) and Thrips
(Fig. !OlE) the preepisternokatepisternal area is represented by the definitive trochantin,
which lacks the typical articulation with the coxal margin, as in Psocoptera.
The prosternum: As in Psocoptera, the degree of sclerotization of the sternal area
varies considerably in different genera. The proboscis is placed on the sternum; hence,
the sternum is more or less membranized mesally. The basisternum is a thinly sclero-
tized median plate in Thrips (Fig. 102B), but it is represented by paired sclerites in
Haplothrips (Fig. 104A). The furcasternum (eusternum of Mickoleit, 1961) is a
medially constricted transverse plate in Thrips; it is represented by paired sclerites in
Haplothrips. The furca arises from the lateral margin of the furcal plate. The spina-
sternum in Thrips is a narrow transverse plate bearing the spina at the middle of its
posterior margin; the same in Haplothrips is a small, subtriangular plate which bears the
spina at its end.
The meso- and meta-notum: Characteristically, in Thysanoptera, the anterior part
of the mesonotum is separated from the rest of the mesonotum by an intervening
membranous area. This isolated anterior part, which Mickoleit ( 1961) called the inter-
tergite, is vertically deflected internally and is hidden beneath the posterior margin of
the pronotum. As Mickoleit showed (Fig. 101C), this part consists of the prescutum
and the acrotergite, and their division is marked by the presence of the :first phragma
below. The detached prescutal arm (Fig. lOlA) is discussed on p. 245.
The anterolateral scuta! suture in Phloeothrips and Thrips (Fig. 102A) is incomplete,
being represented by the thickened ridge on the anterolateral scuta! margin, as in
Psocoptera. Mickoleit (1961) called the oblong subtriangular plate along the antero-
lateral scuta! margin the tegula (Figs. lOlA, 102A). Despite its large size, this
presumed tegula occupies the same relative position as the much smaller prealar sclerite
in Psocoptera (Figs. 93A, 96A). However, the plate in Thysanoptera, unlike the prealar
sclerite in Psocoptera, does not receive muscle t-p 4.
The posterolateral scuta! area is d e p r e s ~ e d along the suture that appears to be the
posterolateral scuta! suture (Fig. 102A). According to Mickoleit ( 1961), the groove
anterior to the posterolateral angle of the mesonotum is ridged internally for the attach-
ment of muscles. The scutoscutellar suture is absent.
The description of the scuta! lateral margin behind the anterior notal process is
omitted here, because the connection of this margin with the axillary sclerites is not
clearly known."'
The mesothoracic postnotum deflects vertically from the posterior margin of the
mesonotum (scutellum) (Fig. 103). Therefore, it cannot be recognized externally as in
Psocoptera. In sagittal sections of this area (Fig. 103), the posterior extension of the
mesonotum (postnotum) is first folded beneath the mesonotum, then it extends vertically
to form the second phragma. The anterior marginal area of the metanotum is first
briefly folded vertically, then extends anteriorly to join the phragma continuous from the
mesopostnotum ( acrotergite of Mickoleit). Here the second phragma is double-walled,
if the infolding of the anterior metanotum is regarded as its posterior wall.
In Phloeothrips the anterolateral angle of the metanotum is the anterior notal wing
process. In Thrips (Fig. 102A) paired protuberances occur near the middle of the
55
Mickoleit did not study the axillary sclerites. Jones ( 1954) presented a sketchy drawing of the axillary
sclerites in Chirothrips. Bekker's (1958b) study on the wing base in Haplothrips is hard to understand.
Bekker recognized five axillary sclerites, but he did not homologize them with those in other insects.
FIG. 101. A, lateral view of thorax in Phloeothrips coriaceus (from Mickoleit); B, lateral
view of prothorax and neck region in Aeolothrips fasciatus (from Mickoleit); C, sagittal
section showing dorsal intersegmental relation between prothorax and mesothorax in Phloeo-
thrips coriaceus (from Mickoleit); D, lateral view of mesothorax in Thrips physapus (from
Mickoleit); E, lateral view of prothorax and neck region in Thrips physapus (from Mickoleit).
REMNANT of
PRE EPISTERNUM
+KATEPISTERNUM
FURCASTERNUM
SPINASTERNUM
+ PRESTERNUM
ANAPLEURAL
CLEFT----+-HY
AN EPISTERNUM
STERNUM +
EPISTERNUM
A
B
PRESCUTUM
(INTERTERGITE)
METANOTUM
l'ST ABO. TERG.
LATERAL CERVICAL
SCLERITE
BASISTERNUM
TRANSVERSE
RIDGE
STERNELLUM
(Mickoleit)
BASISTERNUM +
PREEPISTERNO-
KATEPISTERNUM
l'ST ABO.
STERNUM
FIG. 102. A, dorsal view of thorax in Thrips physapus (from Mickoleit); B, ventral view of
thorax in Thrips physapus (from Mickoleit).
metanotallateral margin; in Phloeothrips the protuberances are less marked. In Thrips
(Fig. 102A) a transverse cleft divides the metanotum into the anterior and posterior
parts; in Phloeothrips the metanotum is a single large plate without a subdivision. The
laterocraniales Tergit and perhaps also the laterocaudales Tergit of Mickoleit (Fig. lOlA)
in Phloeothrips probably represent the metathoracic postnotum.
The mesopleuron: The mesopleuron is divided by the pleural suture into the large,
anterior episternum and the posterior epimeron. The pleural ridge bears a well-
MESONOTUM
POSTERIOR
WALL of
PHRAGMA
L
------------VERTICAL
PROJECTION
FIG. 103. Sagittal section showing dorsal intersegmental relation between mesothorax and
metathorax in Phloeothrips coriaceus (from Mickoleit).
developed arm near its ventral end. In Phloeothrips (Fig. lOlA) two anterior episternal
plates (Episternit of Mickoleit) are probably homologous with the prealar arm and the
interpleurite in Psocus (Fig. 94A), although the dorsal one of these plates lies anterior'
to the spiracle in Phloeothrips and dorsal to the spiracle in Psocus. In Thrips
(Fig. !OlD) the two plates have become completely fused.
The rest of the episternum is comparable with that in lower insects. In Phloeothrips
the horizontal line, which nearly reaches the coxal base at a level below the ventral end
of the pleural suture, is considered to be the anapleural suture. Although this suture
does not have a connection with the pleural suture, it is probably the anapleural suture,
since in Psocus (Fig. 94A) the suture lies more ventrad than in many other insects, and
the slightly more ventral position of the suture in Phloeothrips does not preclude the
possibility that it is homologous with the anapleural suture. Apparently, in the phylogeny
of the Psocoptera-Thysanoptera complex, tho! suture has had a tendency to lie increasingly
more ventrad. In Thrips (Fig. !OlD) the suture lies even more ventrad than in
Phloeothrips, and it has completely lost its typical connection with the pleural suture.
Indeed, on the basis of morphological connections, this suture cannot be identified as the
anapleural suture; but on the basis of its apparent continuity in phylogeny, the above
homologizations are still justified.
The large episternal area dorsal to the anapleural suture is thus the anepisternum.
In Phloeothrips the basalare lies above the dorsal margin of the anepisternum.
In Phloeothrips (Fig. lOlA) the area ventral to the anapleural suture is, as in
Psocoptera, the preepisternum and the katepisternum that have become fused, as a
result of the loss of the paracoxal suture. The trochantin is basally indistinguishable
from the katepisternal area, but in Haplothrips its proximal end is clearly recognized as
the trochantinal end, because of its shape and position near the coxal margin (Fig. 104A).
In Thrips (Fig. 102B) the preepisternokatepisternal area has become indistinguishably
fused with the basisternUhl, as discussed more fully later.
In Phloeothrips (Fig. lOlA) the epimeron has a transverse ridge that separates the
dorsal epimeral plate. Both in Phloeothrips and in Thrips (Fig. lOlD) the anteriorly
displaced metathoracic basalare lies on the dorsal epimeral area, as will be discussed later.
Ventrally, in Phloeothrips (Fig. 104B) and Haplothrips (Fig. 104A), the pleura-
sternal cleft separates the broad sternum from the pleuron. In Thrips (Fig. 102B) the
pleurosternal suture is absent, and the ventrally located anapleural suture gives a spurious
LATERAL
CERVICAL SCLERITE ------,
PLEUROST
TROCHANTIN
INTERPLEURITE
TROCHANTIN
ANAPLEURAL
SUTURE - - ~ e : : J
BASI STERNUM
+VENTRAL
EPISTERNUM
}
I '
' \
I \
A
B
ASISTERNUM
CASTERNUM
LEU RITE
--------PRESTERNUM
FURCA
FURCA
FIG. 104. A, ventral view of meso- and meta-thorax in Haplothrips statices (from Mickoleit);
B, ventral view of meso- and meta-thorax in Ph/oeothrips coriaceus (original).
appearance of being the pleurosternal suture. In Thrips, therefore, the katepisterno-
preepisternal area has become a lateral part of the broad definitive mesosternum.
The metapleuron: The metapleural suture is oblique; it is nearly horizontal in the
anterior half in Phloeothrips (Fig. lOlA). The epimeron is greatly reduced. In the
episternal area of Phloeothrips the anapleural suture lies in the area that corresponds to
the position of the same suture in the mesopleuron; the suture is absent in Thrips
(Fig. 1 02B). In Phloeothrips the narrow, anterior marginal strip must be a strengthening
ridge, or the prepectus.
An unique feature of the metepisternum in Thysanoptera is the anteriorly displaced
basalare (Fig. lOlA, D). Its position relative to the dorsal (anterior) margin of the
anepisternum and the associated muscles (p 3, p-cx 2 or 3), however, clearly show that
this plate represents the metathoracic basalare (functional basalare of Mickoleit). In
Phloeothrips (Fig. lOlA) the basalare bears the second thoracic spiracle, although in
Thrips (Fig. lOID) this spiracle lies above the net-shaped basalare. In both, an
apodeme invaginates from this basalare into the thoracic cavity for the attachment
of muscles.
The meso- and meta-sternum: Unlike the pterothoracic sternum in Psocoptera, the
pterothoracic sternum in Thysanoptera is broad. This difference has probably arisen
directly from their different habitats. Crawling within highly restricted spaces (under
bark, husk, etc.) has probably necessitated the production of widely separated leg bases,
and hence a wide sternum in Thysanoptera.
In Thrips (Fig. 102B), according to Mickoleit (1961), the ventral process for
articulation with the middle coxa is formed by the lateral end of the transverse ridge;
Mickoleit called this ridge the sternocosta. However, in Thrips the mesothoracic
preepisternokatepisternal area is indistinguishably fused with the basisternum. The
articulation is therefore probably homologous with the proximally differentiated
trochantin of the preepisternokatepisternal plate in Haplothrips (Fig. 10'4A) and
Phloeothrips (Fig. 104B), which is not fused with the basisternum. The ,apparent
sternacosta in Thrips is therefore purely secondary... A narrow transverse strip
(sternellum of Mickoleit) lies posterior to the secondary transverse ridge in Thrips.
In Phloeothrips (Fig. 104B) and Haplothrips (Fig. 104A) this secondary transverse
ridge appears to have been lost. The definitive intersegmental suture between the meso-
sternum and the metasternum in these genera probably corresponds to the posterior
margin of the posterior marginal strip of the mesosternum (sternellum of Mickoleit) in
Thrips (Fig. 102B). The mesothoracic spina is lost in Thysanoptera, and this straight
intersegmental suture probably does not correspond to those in lower Pterygota.
In Haplothrips (Fig. 104A) the paired, small sclerites in front of the anterior
margin of the mesothoracic basisternum and between the lateral interpleurites (pre-
sterna?) are probably remnants of the pr,esternum. In Thrips (Fig. 102B) the
corresponding area is occupied by a single, transverse plate bearing the spina on its
posterior margin; this definitive spinasternum may contain the mesothoracic presternal
element. In Phloeothrips (Fig. 104B) three presternal sclerites occur in the corres-
ponding area.
Priesner's ( 1957) study on the pterothoracic sterna shows that the furcae lie
invariably on, or along, the median longitudinal axis of the sternum, as in Psocoptera,
and that the median longitudinal elevation is often present. Evidently, each furcal base
has not migrated laterally with the widening of the sternum, as in some parasitic Diptera
(p. 319). According to Priesner, this median longitudinal ridge is better developed in
good fliers and especial! y in good springers.
The adult musculature: Melis (1935a) described the thoracic muscles in Liothrips
oleae, Maki (1938) described those of Machatothrips artocarpi, and Mickoleit (1961)
those of Thrips physapus (Terebrantia) and Phloeothrips coriaceus (Tubulifera).
Homologies of muscles in the last two species are shown in Table XIII. Some muscles
have evidently undergone considerable shift in points of attachment, so that sometimes
the homologies are uncertain.
Peculiarities of the thoracic musculature in Thysanoptera appear to be in the
retention of the anterior tergocoxal muscles only in the prothorax, and in the presence
of t-p 20 and of a few other muscles that have apparently shifted their origins. The
thoracic musculature of this order is similar to that of Psocoptera in the relative
abundance of ventral muscles (despite the overall tendency towards reduction). The
presence of p-s 10, which extends between the mesofurca and the first and second
abdominal sterna, is a characteristic feature that is shared also by Mallophaga. The
presence of fewer trochanteral muscles in Phloeothrips is probably due to the dorso-
ventral flattening of the body.
56
By definition, the sternacosta connects the furcal b a s e s ~
Gross aspects of the postembryonic muscle development: In an incomplete work by
Melis (1935b), which compared the nymphal and adult thoracic muscles, seven nymphal
prothoracic muscles are present in the adult, and t-p 6 is already present as a distinct
muscle in the nymph.
THE HEMIPTERA (HOMOPTERA)
The neck region: Among auchenorrhynchous Homoptera, a small, undivided lateral
cervical sclerite has been found in Cicada (Taylor 1918; Crampton 1926a), Cyclochila
and Huechys (Larsen 1945c), and Magicicada ( = Tibicina) (Kramer 1950). The
sclerite articulates anteriorly with the occipital process, which is well developed as in
Psocoptera and Thysanoptera. In Cicada also Taylor (1918) showed paired dorsal and
dorsolateral sclerites on the neck membrane. Apparently, in auchenorrynchous Homop-
tera other than Cicadidae, the lateral cervical sclerite is absent. Lew ( 1960) found no
cervical sclerite in the old nymph of Magicicada; in Cyclochila Larsen ( 1945c) observed
that the imaginal lateral cervical sclerite is formed inside the nymphal skin during the
last molt into an adult.
In Cicada at least, as Crampton ( 1926a) clearly showed, the lateral cervical sclerite
has the posterior articulation with the precoxal bridge, which represents the preepisternum.
Hence, the lateral cervical sclerite in Cicada is probably presternal in origin as in
Psocoptera.
In sternorrhynchous Homoptera the sclerites in the neck region are somewhat
difficult to identify, because of the close approximation, or even union, of the ventral
prothorax with the head. In Aleurodes (Fig. 107C), according to Weber (1935), the
lateral cervical sclerite is a slender semilunar plate which, however, appears to be either
the reduced proepisternum (precoxal bridge) or the trochantin. Weber's muscle 0 ism 1,
which is inserted on this sclerite, is probably homologous with t-ti 2 in Thysanoptera.
This muscle is not inserted on the lateral cervical sclerite in other insects. Further, this
sclerite has no normal articulation with the head. All these facts do not support the idea
that the sclerite is the lateral cervical sclerite. In Aphis a doubtful sclerite that Weber
( 1928b) called the Kehlstiick is also not the lateral cervical sclerite. As Weber ( 1928b)
and Larsen ( 1945c) thought, its position is too anterior to be the lateral cervical sclerite.
In the male of Apiomorpha, which Theron ( 1968) described, however, the lateral cervical
sclerite is apparently present.
The pronotum: In Hemiodoecus (Peloridiidae, Coleorrhyncha), as Evans (1939)
showed (Fig. 105A), the pronotum has lateral (paranotal) extensions separated by a
longitudinal suture on each side. Evans regarded them as serially homologous with
wings. In Hemiodoecus the pronotum is fused with the head in the first instar, but later
they become distinct.
In auchenorrhynchous Homoptera modifications of the pronotum are often con
spicuous. A toothlike process occurs on the ventral lateral margin of the pronotum in
Stictocephala and Oxyrhachis (Membracidae), and it may appear to be a pleural structure
( Hasenoehrl and Cook 1965). Wilfried ( 1965) classified the development of the
pronotal lobe in Membracidae in relation to its selective value. In some groups of the
cicada, such as Tettigarcta (Myers 1928; Evans 1940) and Peregrinus (Mathur and
Joesph 1961), the posterior extension of the pronotum covers part of the mesonotum.
Among sternorrhynchous Homoptera the pronotum in Aphis (Fig. 106B) has paired,
elongated sclerites, which form bridges with the mesonotum. In Aleurodes (Fig. 107C)
the pronotum is completely fused with the propleuron, forming a dorsolateral pro
thoracic ring.
The propleuron: In Hemiodoecus (Fig. 105A) the propleuron is divided into the
episternum and the epimeron by the pleural suture. The episternum is further divided
into the preepisternum, the anepisternum and the trochantin by the anapleural cleft and
by the membranous area surrounding the trochantin.
MATSUDA! nm INSECT THORAX
TABLE XIII
Thoracic musculature in Thysanoptera
P hloeothrips
(Mickoleit 1961)
M. lev. cap. terg.
M. lev. cap. prphr.
M. compr. occ.
M. cerv.-postocc.
M. depr. cap. terg.
M. depr. cap. furc. lat.
M. dors. obi. (I)
M. dors. obi. (III)
M. metnot. abdtg.
M. lev. cap. prphr. (!)
M. dors. rect. (II, III)
M. pl.-terg. epist. caud. (II)
M. st.-not. (II, III)
M. pl.-pt. (II, III)
M. pl.-terg. epist. cran. (II)
M. pl.-terg. crist. pl. (II, III)
M. pl.-terg. candy!. (III)
M. prphr.-prfurc.
M. mesphr.-mesfurc.
M. metfurc.-metphr.
M. prom. cox. terg. troch. (I)
M. prom. cox. terg. bascox. (I)
M. rem. cox. terg. (I, II, III)
M. cox.-subalar. (Ill)
M. pl.-basalar. (III)
M. praecox.-basalar. (II, II I)
M. pl.-pt. basal. (III)
M. furc.-pl. (II, III)
M. mesfurc.-abdst.
M. abd. cox. pl. (I, II, III)
M. mesfurc.-profurc.
M.
M. metfurc.-abdst.
M. rem. cox. furc. (I)
M. rem. cox. furc. dors. (II, III)
M. rem. cox. furc. ventr. (II, III)
M. prom. cox. furc. (I I, II I)
M. add. cox. furc. (!)
M. depr. troch. pl. (I)
M. depr. troch. furc. (II, III)
Thrips
(Mickoleit 1961)
M. lev. cap. terg.
M. lev. cap. prphr.
M. cerv.-postocc.
M. depr. cap. terg.
M. depr. cap. furc. lat.
M. dors. obi.
M. dors. obi.
M. metnot. abdtg.
M. lev. cap. prphr.
M. dors. rect.
M. pl.-terg. epist. caud.
M. st.-not.
M. pl.-pt.
M. pl.-terg. epist. cran.
M. pl.-terg. crist. pl.
M. pl.-terg. candy!.
M. pl.-terg. (I)
M. prphr.-prfurc.
M. mesphr.-mesfurc.
M. metfurc.-metphr.
M. prom. cox. terg. troch.
M. prom. cox. terg. bascox.
M. rem. cox. terg.
M. cox.-subalar.
M. st.-pl. (II)
M. pl.-basalar.
M. praecox.-basalar.
M. pl.-pt. basal. (III)
M. furc.-pl. (I, II, III)
M. mesfurc.-abdst.
M. abd. cox. pl.
M. mesfurc.-prfurc.
M. prfurc.-prsp.
M. metfurc.-abdst.
M. rem. cox. furc.
M. rem. cox. furc. dors (II, III)
M. rem. cox. furc. ventr.
M. prom. cox. furc.
M. add. cox. furc.
M. depr. troch. terg. (I, III)
M. depr. troch. pl.
M. depr. troch. furc. (I, II, III)
Muscle
designations
op-t 2
op-t 3
op-cv 1
op-p 1
t-cv 2, 3
s 2
t 12
t 13
t 14
t-p 3
t-p 5, 6
t-p 14
t-p 17 ?
t-p 19 ?
t-p 20
t-p?
t-s 1
t-ti 2
t-ti(cx) 3
t-cx 6 or 7
f-cx 8
p 1
p 2 ?
p3
p 5 ?
p-s 1
p-s 10 or s 15
p-cx5
s 13
s 14
s 20
s-ex 2
s-ex 3
s-ex 5
s-ex 6
t-tr 1
p-tr 1
s-tr 1
249
Remarks
1
2
3
4
5
6
7
8
9
10
250
REMARKs (Table XIII)
( 1) These muscles connect the dorsal part of the interpleural sclerite (Episternit
of Mickoleit) and the membrane lateral to the mesonotum. .
(2) In the metathorax these are two-part muscles, which. connect the pleural
ridge with the posterior lateral margin of the metanotum. They may be modified t-p 19.
( 3) A prothoracic t-p muscle that is not homologized.
( 4) These are two-part muscles.
(5) These muscles arise from the notopleural membrane (Thrips), or from the
apodeme on the pleural ridge (Phloeothrips), and are inserted on the posterior coxal
margin.
(6) The muscle extends between the middle of the episternum and the near
lateral margin of the definitive sternum, where the anapleural suture is present.
(7) These muscles connect the anterior margin of the metepisternum and the
anteriorly displaced basalare. They may represent displaced p 2.
( 8) These muscles connect the anteriorly displaced meta thoracic basal are with
the wing articulation. The posterior point of their attachment is not clear in the original
description.
(9) These muscles connect the mesofurca with the ventrolateral anterior margins
(antecosta of Mickoleit) of the first and second abdominal segments. They may also
be s 15.
( 10) M. abd. cox. pl. in the prothorax of Phloeothrips may be t-cx 4.
In the old nymph of Magicicada (Fig. 105B), as Lew (1960) showed, the pre-
episternal area is thinly sclerotized, and the epimeron is divided into the dorsal and
ventral parts by the transepimeral suture as in the mesopleuron. This suture is also
clearly marked in the adult of a Magicicada examined. In adult Cicadidae and other
auchenorrhynchous Homoptera the episternal part of the anapleural ring is not divided
into the anepisternum and the preepisternum, the anapleural suture being absent. The
preepisternal area is often narrowed and reaches the sternum ventrally. In Scolops the
broad preepisternal area is indistinguishably fused with the sternum. The trochantin is
present in the prothorax of auchenorrhynchous Homoptera.
In Psyllidae (Weber 1929; Tremblay 1965) the pleural suture is nearly vertical and
straight. A small, but clearly differentiated trochantin is basally continuous with the
episternum (Fig. 106A). In the membranous area behind the epimeron of Psy/la
(Fig. 1 06A) a dorsal oblong sclerite and a ventral sclerotization surrounding the first
thoracic spiracle occur. In Trizona (Tremblay 1965) an additional sclerite is present
dorsally above the oblong sclerite. In Aleurodes (Fig. 107C) the ventral half of the
anterior margin of the definitive tergopleuron is the pleural suture, and the anterior
semilunar plate, as already discussed, is the remnant of the episternum. In winged
Aphis (Fig. 106B) the propleuron is more generalized than in other Sternorrhyncha,
being incompletely fused with the pronotum; the clearly defined episternum, epimeron
and trochantin are present. Characteristically, in all the above-mentioned genera of
Sternorrhyncha, the preepisternal area does not extend ventrally, because of the intimate
association of the ventral part of the prothorax with the head.
The prosternum: In the fourth-stage nymph of Hemiodoecus, according to Evans
( 1939), the small paired furcae are separated basally. In Cicadidae the prosternum is
represented by a single medially sclerotized plate bearing the furcae, which is often
medially modified into a trough for the reception of the rostrum at rest. In other
Auchenorrhyncha, as in Cicadidae, the sternum is often reduced and forms a trough.
In winged Aphis (Fig. 109C), despite close approximation of the front leg bases to
the head, three distinct sternal plates are recognized. Weber ( 1928b) called them the
presternum, the basisternum, and the furcasternum, respectively. In Psyllidae, as Weber
( 1929) showed, the labium and the prosternum are in such close union that the pro-
TROCHANTI
PLEURAL
SUTURE
PRESCUTO-
SCUTAL
SUTURE
PARAPSIDAL
SUTURE
A
B
c
ANAPLEURAL CLEFT
AN EPISTERNUM
EPIMERON
TRANSEPI MERAL
SUTURE
PARA COXAL
SUTURE
251
MEMBRANE ATTACHMENT
PHRAGMA.
PRESCUTUM
FIG. 105. A, ventral view of prothorax in Hemiodoecus fide/is (from Evans); B, lateral view
of propleuron in old nymph of Magicicada septendecim (from Lew); C, dorsal anterior part
of mesothorax in Magicicada septendecim (original).
252 MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CAN.\DA
LATERAL
PARAPSIDAL
SUTURE
TRANSANEP IST_.;..._..--
EitNAL SUTURE
ANEPISTERNUM
PROBOSCIS
ANAPLEURAL
SUTURE
PREEPISTERNOKATEPI
STERNUM+ BASISTERNUM
PRESCUTOSCUTAL
SUTURE-----f.
POSTTERGITE ---..,..,
PREALAR ARM
(PRESCUTAL) ----
TRANSANEPISTERNAL
A
B
l'ST ABO.
TERG.
c----UI'TANOTUM

PLEUROCOXAL
ARTICULATION
CHANTIN
POSTNOTUM 2
PHRAGMA 3
N------ POSTNOTUM 3

SUTURE
EPIMERON
FURCA
BASI STERNUM
FURCASTERNUM
Fro. 106. A, lateral view of thorax in female Psylla mali (from Weber); B, lateral view of
thorax in winged female Aphis fabae (from Weber).
l'ST ABO.
TERG.
LATERAL
PARAPSIDAL
SUTURE
PRONOTUM
+ PROPLEURON
PREALAR
PLEURAL
PROSTERNUM
TRANSANEPISTERNAL
SUTURE
LATERAL
PREALAR ARM PARAPSIDAL SUTUR
A
METASCUTELLUM
B
c
253
MESDSCUTELLUM
MET APOSTNOTUM
POSTNOTUM 2
PLEURAL WING .
PROCESS
METANOTUM
LATERAL
POSTNOTUM 2
EPIMERON
POSTNOTUM 3
+ METEPIMERON
l'ST ABO. SEG.
PLEURAL
SUTURE
METEPISTERNUM
FIG. 107. A, dorsal view of thorax in Psylla mali (from Weber); B, dorsal view of the thorax
in Aleurodes brassicae (from Weber); C, lateral view of thorax in Aleurodes brassicae (from
Weber).
sternum is modified into paired processes for grasping the base of the labial rostrum, and
the muscles inserted on the sternum control the movement of the rostrum. The furca
arises from the lateral margin of the furcasternum. In Aleurodes (Fig. 107C) the
prosternum is a narrow, transverse sclerite that nearly reaches the lateral end of the
pronoto-pleuron. The furca arise from each lateral end of the prosternal plate, as in
Thysanoptera and Aphis. The prothoracic spina has not been found in Homoptera.
The mesonotum: In the anterior part of the mesonotum of Magicicada (Fig. 105C)
the antecostal suture represents the anterior margin of the mesonotum, and the first
phragma arises from beneath this suture. The horizontal prescutoscutal suture is
retained only laterally and demarcates the narrow prescutum anteriorly. The prealar
arm, which is basally demarcated by a line, is clearly a prescutal part. The parapsidal
254
suture arises anteriorly from the prescutoscutal suture, as in other insects. The antero-
lateral scuta! suture and the lateral parapsidal suture are absent.
Among auchenorrhynchous Homoptera studied by Kramer (1950), Aulacizes
(Fig. 108B) has retained the complete prescutoscutal suture which runs transversely
behind the antecostal suture, the well-developed acrotergite anterior to the antecostal
suture, the parapsidal suture, and the prescutal prealar arm. What Kramer (1950)
called the "notaulices" in Ceresa (Fig. 108C) is probably the lateral parapsidal sutures.
The medial area bounded by the parapsidal sutures in auchenorrhynchous
Homoptera has often been called erroneously the prescutum, but a complete prescutum
demarcated by the prescutoscutal suture is present only in Aulacizes. In auchenor-
rhynchous Homoptera the prealar arm normally reaches the dorsal part of the episternum,
thereby forming a prealar bridge.
In winged Aphis (Fig. 106B) the oblique prescutoscutal suture is, as in Cicadidae,
continuous anteriorly with the parapsidal suture. The tergal fissure is well marked,
and the anterolateral scuta! suture is absent. In Aleurodes (Fig. 107B, C) the lateral
parapsidal suture, instead of the parapsidal suture, arises laterally from the base of the
prealar arm. The anterior of the two protuberances along the scuta! lateral margin is
probably homologous with the prealar sclerite in Psocoptera, since it lies posterior to the
prealar arm as in Psocoptera (Fig. 96A). The larger posterior protuberance is the
tegula, because it is associated with the wing base. In Psylla (Fig. 107A) a sub-
horizontal, transscutal suture that connects the membranous fissure formed behind the
base of the prescutal arm on each side may represent the lateral parapsidal sutures that
have become medially continuous. In Psylla, as in Aleurodes, the two protuberances
along the scuta! lateral margin are the prealar sclerite and the tegula, respectively. In
all three genera of sternorrhynchous Homoptera the prealar arm, which is of prescutal
origin, reaches the anepisternum.
Weber (1928b, 1929, 1935) called the area bounded by the parapsidal sutures or
by the lateral parapsidal sutures in these sternorrhynchous Homoptera the prescutum.
This usage of the term prescutum is not in accord with the definition of the prescutum
clarified in this work (p. 12). In Coccoidea also, as the studies of Ezzat ( 1956),
Giliomee (1967) and Theron (1968) show, the parapsidal suture commonly occurs.
On the posterior half of the mesonotum in Magicicada the posterolateral scuta!
suture is present. In Aulacizes (Fig. 108B) the same suture is grooved and extends as
far back as the posterior end of the mesonotum and forms a wing locking device, which
Kramer (1950) called the tergal wing groove. In Lepyronia (Kramer 1950) also, a
tergal groove is formed in a corresponding position.
The anteromesally produced scutoscutellar suture is clearly present in winged Aphis,
although the suture is more or less medially obliterated or absent in other Homoptera
that have been studied. In auchenorrhynchous Homoptera a median or postmedian
transverse suture occurs. In Lepyronia this suture reaches the tergal wing groove,
thereby forming a posteromedian subtriangular plate; in Aulacizes (Fig. 108B) the same
suture nearly reaches the tergal wing groove. Kramer (1950) called the postmedian,
subtriangular plate posteriorly demarcated by this transverse suture the scutellum.
Since the transverse suture does not represent the anterior limit of the scutoscutellar
suture, "the scutellum" of Kramer is not homologous with the scutellum in Aphis.
In Magicicada (Fig. 108A) the anterior margin of the scutellar area is nearly straight
and without a medial production; the postnotum is well exposed. In many other
Homoptera the median caudal production of the scutellum is more or less conspicuous,
and the postnotum is often hidden beneath the mesal production of the scutellum. In
most Homoptera the mesothoracic postnotum extends laterally to unite with the epimeron,
thereby forming a postalar bridge.
In Psy/la, as Weber (1929) clearly showed (his fig. 21), the anterior wall of the
second phragma, to which the mesothoracic dorsal longitudinal muscles (t 14) are
attached, is a simple invagination of the mesothoracic postnotum. A much smaller
invagination from the anterior margin of the metanotum forms a narrow slit behind the
POSTERIOR
NOTAL
PROCESS
POSTERO-
LATERAL
SCUTAL
SUTURE
SCUTOSCUTELLAR
DORSAL
tERVICAL
SCL
PRONOTUM
PRONOTUM
BASALARE
PLEURAL
MESONOTUM
SUBALARE
WING PROCESS
ANEPISTERNUM
PLEURAL
SUTURE ------f-'-l
EPISTERNUM -----t-
TROCHANTIN -----%
ANAPLEURAL
SUTURE
255
PRESCUTOSCUTAL
. SUTURE
ARAPSIDAL
SUTURE

'-. ARM
ANTERIOR
NOTAL
PROCESS
POSTERIOR
NOTAL
PROCESS
AXILLARY
CORD
8 TERGAL WING
GROOVE
(POSTEROLATERAL
SCUTAL SUTURE?)
LATERAL
-- ARAPSIDAL
SUTURE
POSTERO-
LAtERAL
SCUTAL
SUTURE
METASCUTUM
--11:----PLEURAL
WING PROCESS
:Il!:ifl'Lf-,!----PLE URAL
SUTURE
RON
TROCHANTIN
Fro. 108. A, dorsal view of thorax in Magicicada septendecim (from Kramer); B, meso-
notum of Aulacizes irrorata (from Kramer); C, mesonotum in Ceresa bubalus (from Kramer);
D, lateral view of thorax in Aulacizes irrorata (from Kramer).
obliquely placed mesothoracic postnotum, and represents the posterior wall of the second
phragma. In Magicicada and Aulacizes, judging from the description given by Kramer
( 1950), the second phragma is primarily the invagination from the mesothoracic post-
notum, while in Lepyronia the second phragma is the invagination of the anterior margin
of the metanotum.
Axillary sclerites: Weber (1929) schemetically showed the relationships of axillary
sclerites of the forewing with associated structures in Psylla and Aphis as follows:
Prescutum
Scutum
Scutellum
HORIZONTAL RELATIONSHIPS IN Psylla
-----Prealar arm ----Prealar sclerite
(polster 1 of Weber)
I
Tegula (polster 2 of
Weber)- Costa
I
/
Ax. sci. 1 ----(Subcostal
Anterior I
not. proc."'-
Ax. sci. 2 -----Radius+ media+ cubitus
I
-----Ax. scl. 3 ----Analis
-----Ligament -"'----Posterior margin of forewing
VERTICAL RELATIONSIDPS IN Psylla
Prescutum Scutum Scutellum----Postnotum
/ I
Anterior margin--Wing--Posterior margin
of wing \ of wing
/Ax. sci. 2 I
/ Subalare
// I
Basalare Epimeron
I
Episternum Pleural ridge
Coxa
HORIZONTAL RELATIONSIDPS IN Aphis
Prescutum --------Prealar arm
r T'"""""' ~ P >""'" ~ C " " "
// I
Ant. notal }
proc. /Ax. sci. 1 Subcosta
I \d. d' b.
Med. notal Ax. sci. 2 Ra ms + me 1a + cu ttus
proc. I
post. not.
proc. ---Ax. sci. 3 ------ Analis
Scutum
Scutellum --------.Ligament------- Posterior margin
VERTICAL RELATIONSHIPS IN Aphis
Prescutum Scutum ----Scutellum---- Postnotum
Anterior PostLior margin

Basalare
Epimeron
--Episternum
<------ Pleural ridge
Coxa
The mesop/euron: In Cicadidae the mesopleuron is generalized and resembles that
of Neuroptera. In Tibicen (Fig. 109A) the paracoxal suture extends the entire
episternal area and reaches the pit for the pleural arm. On the epimeron the
transepimeral suture extends from the pit on the pleural ridge to the ventral end of the
lateral postnotum. Matsuda (1960a) mistook this suture for the paracoxal suture (then
precoxal suture). The oblique anapleural suture divides the anepistemum into the dorsal
anepisternum and the ventral preepisternum. The basalare is separated from the
anepisternum by the membrane surrounding it. The preepisternum appears to meet its
counterpart from the other side along the median longitudinal suture and the lateral
margin of the anteriorly placed basisternum. Matsuda ( 1960a) also recognized a
narrow strip along the furcastemum, which he interpreted as the ventropleurite."' In
contrast to holometabolous insects, however, this presumed ventropleurite does not form a
process for articulation with the inner margin of the coxa. In the absence of the ventral
process Tibicen and other Homoptera are similar to fsocoptera. The katepisternum is
represented largely by the trochantin, which articulates with the coxa. The antecoxale
is a narrow strip between the paracoxal suture and the trochantin. The mesepimeron
is divided by the transepimeral suture into dorsal and ventral parts. The latter has a
lobate extension posterolaterally; ventrally, it forms a postcoxal bridge that becomes
continuous with the katepisternal element along the inner margin of the coxa."'
In Tettigarcta, according to Evans (1940), the mesepisternum also consists of the
anepisternum, preepisternum, and katepisternum; the mesepimeron is divided by the
transepimeral suture into dorsal and ventral parts, which Evans, however, called the
anepimeron and the katepimeron, respectively. In other Cicadidae the mesopleuron is
also apparently similar to that in Tibicen and Tettigarcta. In some cicadas, such as
Tibicen (Taylor 1918), the anepisternum is further subdivided by a transverse suture.
Taylor called the smaller ventral part of the anepistemum the median portion of the
episternum.
In the mesopleuron of many other auchenorrhynchous Homoptera the paracoxal
suture is absent. The anapleural suture is present in Scolops, Lepyronia, and Aulacizes.
The trochantin is present in the mesopleuron of all auchenorrhynchous genera studied by
Kramer (1950). A diagonal suture on the mesepimeron in Aulacizes (Fig. 108D) and
Ceresa (Kramer 1950) has the function of receiving the basal lip of the C vein of the
tegumen; Kramer called this suture the pleural wing groove. Judging from the figures
given by Taylor (1918), the pleural wing groove occurs commonly in auchenorrhynchous
Homoptera. Accordingto Taylor (1918), in Fulgoridae, Jassidae, and Membracidae
the free anterior basalare is absent, although the posterior basalare comparable with that
in Cicadidae is present.
67
As Snodgrass (1927) and Lew (1960; Fig. 109B) have shown, the subcoxal element in the last instar nymph
completely surrounds the leg base of all segments. In the adult the coxal bases come to lie closer to each
otber. Yet it is probable that the subcoxal (katepleural) element including the ventropleurite is present along
the inner coxal margins in the adult.
In the mesopleuron of winged Aphis (Fig. 106B) the paracoxal suture and the
trochantin are absent. Two oblique sutures meet near the ventral end of the pleural
suture. Weber ( 1928b) homologized correctly the ventral one' with the . ariapleural
suture in Perla (Fig. 48).
158
The preepisternum in Aphis has consequently' become
indistinguishably fused with the basisternum. The dorsal oblique suture is then homo-
logous with the transanepisternal suture in Tibicen, and the transverse triangular plate
(Lateropleurit of Weber) is homologous with the median portion of the episternum of
Taylor (1918). Further, the anapleural suture lies more ventrally than those in most
insects, but it lies in a position comparable with the anapleural suture in Thysanoptera.
The anapleural suture and the transanepisternal suture are present also in Chermes
(Gaumont 1967), and Psylla (Fig. 106A).
In Psylla (Fig. 106A) the pleural suture has a medial interruption. The trochantin
is absent, and the area immediately above the coxa is sclerotized. Weber (1929)
thought that an oblique suture reaching the pleurocoxal articulation ( anapleural suture in
Fig. 106A) separates the pleuron from the sternum," and Tremblay (1965) thought that
in Trioza the sternum and the pleuron are fused in the area dorsal to the suture in
question. These interpretations are not acceptable, because in no insect does the sternum
invade dorsally as far as the ventral end of the pleural suture. The suture in question is
probably the anapleural suture, considering the tendency for the anapleural suture to lie
more ventrad in Psocoptera, Thysanoptera, and Aphis.
In Aleurodes (Fig. 107C) the mesopleuron appears to be even more specialized
than in Aphis and Psyllidae. The pleural suture is only dorsally distinct. The
imaginary ventral part of the pleural suture is indicated by a vertical broken line that
reaches the pleumcoxal articulation. The transanepisternal suture is apparently present;
other sutures on the episternum cannot be homologized. The epimeron meets the lateral
postnotum along a suture. The transepimeral suture is apparently present.
In various male scale insects what Theron (1958), Giliomee ( 1967, 1968), and
Beardsley ( 1968) called the marginal ridge and the subepisternal ridge correspond to the
anapleural suture and the transanepisternal suture, respectively.
The mesosternum: In Tibicen (Fig. 109A) the mesosternum consists of the
anteriorly widened basisternum (presternum of Matsuda, 1960a), the median longi-
tudinal ridge, and the posterior small plate. The furcal base is continuous with the
median longitudinal invagination. The mesothoracic spina is absent. The border
between the mesosternum and the metasternum is, therefore, obscure. The definitive
mesofurcasternum may actually be the metathoracic basisternum that has migrated
forward, since the basisternal plate both .in the mesothorax and meta thorax apparently
shift their positions forward during the final molt into the adult. This is evident in a
comparison of Fig. 109B with Fig. 109A. Such a shifting forward of the metasternum
is also evident in Psocoptera (Fig. 94B). The ridged anterior margin of the basisternum
(prepectus) is continuous with the similarly ridged anterior margin of the episternum.
In Aulacizes and Lepyronia (Kramer 1950) the mesosternum also has an anteriorly
widened basisternum and a posterior narrow plate connected by a median longitudinal
suture. In Scolops (Kramer 1950) the widely separated furcal pits are connected by a
transverse line (sternacosta). The absence of the mesothoracic spina in these genera
makes the posterior limit of the mesosternum uncertain. As in Cicadidae, the oval
shaped definitive furcasternum present posterior to the furcal pit in these genera may be
the metathoracic basisternum.
In Psyllidae the mesosternum consists of an anteriorly dilated, oval-shaped basi-
sternum and its narrowed posterior end which bears furcae. The narrow area along the
inner margin of the coxalbase may well be pleural in origin. In wingless female Aphis
the coxal bases are widely separated; the mesofurcal pits lie wide apart from each other
58ln Weber's terminology. the anapleural suture is "Grenzleiste Lateropleurit-Laterostemit".
ssweber (1928b) homologized the corresponding suture in Aphis (Fig. 106B) with the anapleural suture in
Perla (Grenz;leiste Lateropleurit-Laterosternit). Weber (1928a, b) was aware that a subcoxal element (Latero-
sternit of Weber, or the preepisternum in this work) lies ventral to this suture. Yet he (1929) called the
corresponding suture in Psylla the "Grenznaht" between the sternum and the pleuron. '
PARACOXAL SUTURE
ANTECOXALE
FURCAL
B
FURCAL
PIT
FURCA EPIMERON
259
BASALARE
EPIMERON
PRESTERNUM
BASISTERNUM
FURCA
"'%---- BASISTERNUM
FURCASTERNUM
FIG. 109. A, ventral view of pterothorax in Tibicen bifida (from Matsuda); B, ventral view
of thorax in old nymph of Magicicada septendecim (from Lew); C, ventral view of thorax in
winged female of Aphis fabae (from Weber).
and they are connected by a transverse line (sternacosta?). The mesosternum is
therefore clearly divided into the anterior basisternum and the posterior furcasternum.
In the winged female of Aphis (Fig. 109C) coxal bases of the middle legs are even more
widely separated. A deep, internally ridged, median longitudinal suture extends nearly
the entire length of the mesosternum, and the furcae arise from a common base at the
caudal end of the ridge. A secondary transverse groove crosses the middle of the
longitudinal suture. Thus, there are great sexual differences in the mesosternum of
Aphis. In Aleurodes the median longitudinal invagination of the sternum is also con-
spicuous. In all of these sternorrhynchous Homoptera the mesothoracic spina is absent,
as in auchenorrhynchous Homoptera. The shift forward of the basisternal plate, how-
Fro. 110. A, mesonotum in Sea lops pungens (from Kramer); B, ventral view of metathorax in
Aleurodes brassicae (from Weber); C, same as B, showing episternal articulation with coxa
(from Weber).
ever, has not occurred in sternorrhynchous Homoptera with widely separated leg bases.
It appears probable, then, that the (presumed) shift forward of the basisternal plates in
auchenorrhynchous Homoptera occurs mainly as a mechanical consequence of the
approximation of leg bases during the final molt into the adult.
The metanotum: Among auchenorrhynchous Homoptera studied by Kramer
(1950), the relatively generalized metanotum in Scolops (Fig. llOA) is nearly as large
as the mesonotum. The antecostal suture is clearly present medially and, according to
Kramer, it gives rise internally to a poorly developed phragma. The scuta! area has a
membranized area on each side of the median quadrangular area. The broad postnatal
area is closely united with the scutellum, and laterally with the epimeron to form the
postalar bridge.
In many other auchenorrhynchous Homoptera the metanotum is considerably
reduced. In Magicicada the anterior margin of the metanotum is connected medially
with the mesothoracic postnotum, without forming a phragma; the metathoracic post-
notum is a narrow plate that is largely hidden beneath the scutellum, an\l it unites with
the epimeron laterally.
In Lepyronia the metathorax is large and plays an important role in flight; the
phragma that arises from the anterior margin of the metanotum is a functional part of
the phragma. The prescutum, which is demarcated laterally by the apparent prescuto-
scutal suture, sends out the prealar arm from its side. In Lepyronia and Ceresa the
scutoscutellar area has a median longitudinal sulcus.
In winged Aphis (Fig. 106B) the reduced metanotum is a simple, undivided plate.
Its lateral postnotum extends posterolaterally to unite with the epimeron. In Aleurodes
(Fig. 107C) the large postnotum forms the third phragma and is completely fused
laterally with the epimeron.
The metapleuron: In Tibicen (Fig. 109A) the paracoxal suture is absent both on
the episternum and on the epimeron, and the anapleural suture is indistinct. The
trochantin is present. In Tettigarcta (Evans 1940) the episternum is undivided, but a
line dividing the epimeron into anterior and posterior parts may represent the paracoxal
suture. In the male cicadas the epimeron is a broad lobe, sometimes called the
operculum; it forms a cover for the tympanum of the sound-producing organ. In
Cryptolympaoo (Taylor 1918) two basalaria are present.
In other auchenorrhynchous Homoptera sutures dividing the pleuron are, as in
Cicadidae, largely lost. In Ceresa the epimeron is divided by a suture into dorsal and
ventral parts. The development of the meron in some auchenorrhynchous Homoptera
is conspicuous and it has become solidly united with the pleuron. The trochantin is
usually present in auchenorrhynchous Homoptera. In many auchenorrhynchous
Homoptera, as will be discussed later, the proximal (ventral) end of the preepisternum
meets the counterpart from the other side along the median longitudinal suture.
In Apsylla, as Weber (1929) showed, the straight pleural suture divides the meta-
pleuron into the episternum and the epimeron; the trochantin is present. In Psy/la
(Fig. 106A), however, the pleural suture cannot be recognized, because of the enormous
dorsal development of the meron, and the trochantin is fused with the episternal area
along a thick ridge.
In Trioza (Tremblay 1965) also the meral area is greatly developed dorsad and the
pleural suture is lost. In both Psy/la and Trioza the imaginary original pleural suture is
nearly horizontal, as seen in Fig. 106A. In the winged female of Aphis (Fig. 106B)
and Aleurodes (Fig. 107C) the pleural suture is present and the trochantin is absent.
No conspicuous meron is formed in either of these genera. The episternum in Aleurodes
is strongly approximated towards the median longitudinal axis, as will be discussed shortly.
The metasternum: In Cicadidae (Fig. 109A) the metathoracic basisternum, as
already noted, has probably shifted its position anteriorly. The median longitudinal
suture on the ventral surface of the metathorax is therefore probably the line of union
of the proximal ends of the preepisterna. The furcae arise more anteriorly than in the
mesosternum. Hence, the area between coxal bases can be regarded as the furcasternum.
In many other auchenorrhynchous Homoptera also the median longitudinal suture is
probably the line of union of the preepisterna.
In winged Aphis (Fig. 109C) the furcal pits lie wide apart from each other. The
basisternum is fused with the episternum. The furcasternum is separated laterally by a
horizontal suture from a narrow, sclerotized pleural area along the inner margin of the
coxa. In Aleurodes (Fig. llOB) the episternum (metasternum of Weber 1935) on each
side meet along the median longitudinal axis, which in turn forms a high ridge internally,
and the furca lies at its anterior end. Apparently, therefore, the sternum has become
completely lost externally. The episternal plate forms a condyle at its distal end, and
articulates with the coxal margin (Fig. llOC). Weber (1935) called this articulation
the sternocoxal articulation. Hence, it was implicit in Weber's interpretation that the
episternal condyle was the ventral condyle present in many holometabolous insects.
However, the condyle probably represents the distal end of the trochantin, since the
condyle, unlike the true ventral condyle, lies far behind the furcal base. The more
ventral position of this presumed distal end of the trochantin is comparable with that in
Psocoptera (p. 228), and is due simply to the strong approximation of coxal bases
towards the median longitudinal axis of the body. In Psy/la the metasternum is a
narrow and well sclerotized plate; internally furcae are strongly developed.
The adult musculature: Berlese (1909b) studied the thoracic musculature in Cicada
(= Tibicen) plebia; Maki (1938) and Vasvary (1966) have studied the same in Huechys
sanguinea and Tibicen chloromera, respectively. For other auchenorrhynchous Homop-
tera Maki's ( 1938) study on the thoracic muscles in Cicadella ferruginea still stands
alone. In Table XIV the muscle numbers for Cicadidae are those used by Vasvary.
Those numbers with M and B at the end are the muscle numbers used by Maki
and Berlese.
Among the studies of thoracic muscles in sternorrhynchous Homoptera, only
Weber's works are tabulated in Table XV. They are those in Aphis fabae ( 1928b),
Psy/la mali ( 1929), and in Aleurodes sp. ( 1935). There are also less complete studies
262
on the thoracic musculature in Schizoneura americana (Miller 1933b), in the male of
Pseudococcus spp. (Make! 1942), in Dorsicha stebbingi (Abdul-Latif and Aziz Aslam
Khan 1959), in Dactylosphaera vitifolii (Rilling 1960), and in Chermes (Gatimont
1967). A considerable difficulty was experienced in homologizing the muscles that have
shifted their points of attachment, and some of the homologies suggested for the muscles
in the prothorax and the neck region are not certain.
Compared with the thoracic musculature in Psocoptera, the thoracic musculature in
both Auchenorrhyncha and Sternorrhyncha is much reduced in number. An unique
muscle in Sternorrhyncha is p-cx 9. The relative abundance of t-p muscles is charac-
teristic of Homoptera.
Gross aspects of the postembryonic muscle development: In the nymph of Psylla,
according to Weber ( 1929), the external structure and the musculature of the mesothorax
are similar to those of the metathorax. Weber described only the metathoracic muscles,
but he tabulated and figured the mesothoracic and prothoracic muscles in the last instar
nymph. Identifications of the muscles in these two segments can be based mainly on his
figures. Makel's ( 1942) study on Pseudococcus spp. was aimed at tracing the develop-
ment of individual muscles, but again, the very limited descriptions often makes it
difficult to identify them. Nevertheless, some gross aspects of the developmental pattern
of the thoracic muscles that are discernible from their studies are summarized below,
with some trepidation.
In Psylla mali
Nymphal muscles that become lost in the adult: Ibm 1 (s-ex 5), I pm 3, 4 (p and
p-cx muscles), I dvm 3-5 (t-cx muscles), I ism 1, 2 (t-s 1), II vim 1, 2 (s 13?), II dvm 4
(t-cx 6 or 7), III dim 2, 3 (t 12, 13?), III zm 1, 2 (p-s 1), III dvm 4, 5 (t-cx 6, 7).
Adult muscles that are formed either by fusion or by reduction of nymphal muscles:
II zm (p-s 1) from nymphal II zm 1, 2, 3; II ism ( t-s 1) from nymphal II ism 1, 2
(t-s 1?); III dvm (t-p 5 or 6) from nymphal III dvm 1, 2; III dim (t 14) from nymphal
dim 1, 2 (t 14).
Nymphal muscles that probably enlarge and split secondarily to produce a greater
number of muscles in the adult: Nymphal 0 ism that become 0 ism 1, 2 (t-s(cv) 9);
nymphal 0 vim that become 0 vim 1, 2, and 3 (s 1, 2); nymphal II pm 1-3 that become
II pm 1-7 (t-p 4, t-p 16?, t-p 19, p 3, p-cx 5); nymphal III pm 2 that become III pm 2, 3
(p 3); nymphal III vim that become III vim 2, 3 (s 20?), which Weber listed but did
not describe.
In Pseudococcus spp.
( 1) Muscles that undergo split, fusion, or shift in points of attachment:
20 (t-cv 1?), 23 (t-cx 6 or 7), 26 (s-ex 4), 41 (p-s 1?), 46 (t-cx 6 or 7), 47, 48, and
49 (t-p muscles), 50 and 51 (p-s muscles), 55 (t-p muscle), 56 (t-s 1), 57 a (t-p muscle),
57a (p-s muscle), 61 (spiracle muscle), 66 and 67 (t-p muscles), 73 (p-tr 1), 77 (pleural

(2) Nymphal muscles that are absent in the adult: 35 (t-p 5?), 42 (t-cx 6 or 7),
43 (t-cx 6 or 7), 53 and 54 (t-p 20?), 58 (p-s muscle), 59 (t-s 6?), 60 (p-s muscle?),
74 (p muscle).
(3) Muscles that transform by incorporating myoblasts into nymphal muscles
during development: 35' (t-p 5, 6), 36' (p 3).
( 4) Muscles that are new formations of the pronymphal or nymphal stage:
62 (t-p 14), 63 (t-p muscle), 64 (t-s muscle), 65 (t-p 4?).
In Psylla and Pseudococcus some muscles are lost in the adult. Whether these
muscles are present in the newly hatched adult and later become lost as in Orthoptera,
or whether they are lost during the final molt into the adult, is not known.
The postembryonic development and histology of flight muscles: The development
of a flight muscle (t 14) in Cyclochilla, according to Tiegs (1955), is through repeated
longitudinal fiber cleavages of a single immature nymphal fiber; this fiber, in its turn,
arose from a small group of about 12 myoblasts in the newly emerged nymph. The
TABLE XIV
Thoracic musculature in auchenorrhynchous Homoptera
Cicadidae Cicadella
(Vasvary 1966) (Maki 1938) Muscle designations Remarks
4 1 cv(d)-t 1
8 op-t 2
11, 12 op-cv 3
9 op-p 1
18, 19 op-p 2
4 op-s 1
13, 14 5, 6 t-s(cv) 1
7 t-s(cv) 9 1
10M t-cv 1
15 t-cv 2
10 cv-s 1 2
16 t-cv 3 ? 3
9 3 s 1 or 2
7 2 t 12
35 20
42
5, 6 t 13 ?
4 1 t 14
34 19
65 41
39 (I I) t-p 3
37 (II) 22 t-p 5, 6
67 (III) 42
40 (II) t-p 8
43 (II) t-p 10 or t-p 4 4
41 (II) 24 t-p 12
69 (Ill) 46
45, 46 (II) 25, 26 t-p 13
71, 72 (Ill) 47, 48 t-p 14
44 (II)
70 (III) ?
t-p 15
47 (II) t-p 16
20 (I) t-p? 5
42 (I-II) t-p ? t-s 2 ? 6
17 8 t-s 1
38 23
68 45
21, 22 10 t-ti(cx) 2
so 30
75 49
29 (I) 14 t-ti(cx) 3
24, 25, 26 11, 12 t-cx 6, 7
54, 55 32
78, 79 51, 52
27 t-cx 8
56 33
80 53
48 (I I) 28 p3
49 (II) 29 p-s 1
74 (III)
TABLE XIV (Continued)
Cicadidae Cicadella
(Vasvary 1966) (Maki 1938) Muscle designations Remarks
73 (II-III) 27 (HI) p-s 5
76 (III) p-ti 1 7
51, 52 (II) p-ti 2
60 (II) 36 p-ti 3
15 p-cx 4 8
46 M? 35
83 54 ?
30, 31 (I) 16 p--cx5
36 (HI) 21 s 13
66 (II-III) 43
104 B (I) s 14
28 13 s--ex 3
57, 58 34
81, 82
23 s--ex 5
53 31
77 50
59 (II) s--ex 6
32 17 t-tr 1 9
61 37
84, 85 55
33 (I) 18 p-tr 1
38 (II)
62 (II) p-tr 2
86 (III) 56
63 (II) 39 s-tr 1
87 (III) 57
REMARKS (Table XIV)
( 1) 7 in Cicadella is inserted anteriorly on the head outside the base of the
proboscis.
(2) Vasvary ( 1966) described the posterior point of attachment of muscle 10 in
Cicadidae as the pleural arm, although its homologue (according to Vasvary) in Huechys
(Maki 1938) is attached posteriorly on the furca.
(3) 16 in Cicadidae is inserted on the base of the tentorium. Therefore, this
muscle may be t-s(cv) 1.
( 4) 43 in Cicadidae connects the lateral margin of the mesonotum immediately
anterior to the anterior notal wing process with the pleural wing process.
(5) 20 in Cicadidae is a prothoracic tergopleural muscle that is not homologized.
( 6) 42 in Cicadidae connects the lateral margin of the mesonotum and the pro-
thoracic pleural arm. It is absent in both Huechys (Maki 1938) and Cicada (Berlese
1909b).
(7) 76 in Cicadidae may be p-ti 2. Its relatively anterior position may be due to
reduction of the pleuron.
(8) 54 in Cicadella and 46 M in Huechys arise from the anterior portion of the
episternum.
(9) 84 in Cicadidae arises from the second phragma.
TABLE XV
Thoracic musculature in sternorrhynchous Homoptera
Aphis Psylla A leur odes Muscle
(Weber 1928b) (Weber 1929) (Weber 1935) designations Remarks
I dim 1 I dim I dim 2 cv(d)-t 1
0 dim 0 dim 2 op-t 1 or 2
0 dim 1 op-t 2 or 3
I dim 1 op-t 3
0 dvm op-p 1 1
0 vim 2 0 dim op-p 2
0 vim 1 op-p 2 ? 2
0 vim 3 op-p 2 ? 3
0 dim 3 0 ism 2 t-s(cv) 1
0 ism 0 ism 1, 2 ? 0 ism 3 t-s(cv) 9 4
Ovim 4 0 vim 1-3 0 vim s 1, 2 5
I dim 2 t 12
II dlm 3 II dlm 2 I I dlm 2
III dlm
II dim 2 ? II dim 2 t 13 6
I dim 1 I dim, 0 dim 1 I dlm 2 t 14
II dlm 1 II dim 1 II dlm 1
III dim I I I dim
II pm 5 II pm 10 t ? 7
I dim 3, 4 t ? 8
III pm 3 t-p 3
II pm 12 II pm 7 II pm 9 t-p 4
Ill pm 7
II dvm 1, 2 II dvm 1, 2 II dvm 3 t-p 5, 6
III dvm 1 III dvm III dvm 1, 2
II pm 4 t-p 8
II pm 10, 11 II pm 7 ? t-p 10 9
II pm 5, 6 t-p 13 10
Ill pm 4, 5 t-p 14
II pm 4 t-p 16 ? 11
I ism t-p 17
II ism 12
II pm 3 t-p 19
1 pm 1 I pm t-p? 13
I dvm 2 t-s 1 14
II ism II ism
III ism III ism 1, 2
0 ism 1, I dvm 1 t--cx( ti) 2, 3 15
II dvm 3 II dvm 1
I dvm 1 I dvm 1, 2 t--cx 6 or 7
II dvm 3 II dvm 4 16
III dvm 2 Ill dvm 4
II pm 8 t--cx 8
III pm 6
II pm 6 p 1
II pm 3 p2 17
II pm 1-4 II pm 1, 2 II pm 1, 2 (pars) p3
III pm 1 III pm 1-3 III pm 1
Izm Izm Izm p-s 1
II zm II zm
III zm
TABLE XV (Continued)
Aphis Psylla Aleurodes Muscle
(Weber 1928b) (Weber 1929) (Weber 1935) designations Rem.arks
I pm 7
p-s? 18
II pm 7
p-ti(cx) 1
I pm 2 ? I pm 1 ?
II pm 2 (pars)
p-ti(cx) 2 19
I pm 3, 4
p-cxS
II pm 8 II pm 5, 6
III pm 2
I vim 2 p-cx8
I pm 5, 6 I pm 2 p-cx 9
II pm 9
III pm 3
II vim 1 s 12
I vim 1 ? I vim 1, 2 I vim s 13 20
II vim 2 II vim
III vim III vim ? III vim s 20 21
Ibm 3 Ibm 3 Ibm 3 s-ex 3 22
II bm 3 II bm 3 II bm 3
III bm 3 III bm 3 III bm 3
Ibm 1 Ibm 1 s-ex 5
II bm 1 II bm 1 II bm 1
III bm 1
I dvm 2 t-tr 1
II dvm 2
III dvm 3
III pm 2 p-tr 2
Ibm 2 Ibm 2 s-tr 1
II bm 2 II bm 2 II bm 2 22
III bm 2 III bm 1, 2 III bm 2
REMARKS (Table XV)
( 1) Weber ( 1928b) thought that the posterior point of attachment of 0 dvm in
Aphis was presumably on the gular sclerite (Kehlplatte) that had become indistinguish-
ably fused with the propleuron.
(2, 3) 0 vim 1 in Aphis connects the dorsal margin of the preepisternum with the
lateral margin of the head at the point where the dorsal tenorial arm ends; 0 vim 3 in
Aphis extends between the propleural ridge and the lateral part of the proximal labial
segment; this muscle may be a modified op-p 2.
( 4) 0 ism 1, 2 in Psylla are a single muscle in the nymph that arises from the
anterior margin of the pronotum, or from near the ventral end of the anteriorly placed
first phragma.
(5) 0 vlm 1-3 in Psylla are a single muscle in the nymph.
(6) II dlm 2 in Aphis extends between the posterior margin of the scutellum and
the. posterior corner of the area delimited posteriorly by the lateral parapsidal suture.
(7) These muscles extend from the prealar arm to the anterior lateral margin of
the mesoscutum.
(8) I dim 3 in Aphis connects the lateral posterior end and the lateral anterior
end of the sclerotized pronotum. I dim 4 is a short muscle that connects the posterior
margin of the pronotum proper with the posttergite of Weber.
(9) II pm 7 in Aleurodes connects the episternum and the median part of the
"Fliigelgelenk". Thus, Weber's description was not specific in the dorsal point of attach-
ment. t-p 10 usually arises from the pleural arm, which is lost in Aleurodes.
( 10) II pm 5, 6 in A leurodes extend from a secondary pleural ridge to the third
axillary sclerite (Analge/enkstiick of Weber, 1935).
( 11) II pm 4 in Psyl/a goes from the epimeral surface to the dorsal edge of the
epimeron.
(12)
(13)
I ism in Aleurodes may be t-s 1.
These muscles are prothoracic tergopleural muscles that are not homologized
here.
( 14) I dvm 2 in Aphis arises dorsally from the posttergite of the pronotum.
(15) Weber (1935) thought that 0 ism 1 in Aleurodes was a tergocervical muscle,
but it is t-ti 2 (p. 248).
( 16) I dvm 1 in Psy/la and A phis are inserted on the presternum and are retractors
of the labium.
( 17) II pm 3 in Aleurodes may be p 1.
(18) I pm 7 in Aphis extends between the posterior margin of the epimeron and
the lateral edge of the sternum near the furcal bases.
( 19) Prothoracic muscles in Aphis and Psylla may represent p-cx 4.
(20) I vim 1 in Aphis extends from the mesofurca to the presternum, to the
postcoxal bridge, and to the posterior margin of the presternum.
(21) Weber (1929) listed III vim in Psy/la, but did not describe it.
(22) In the metathorax of Psylla two sternal muscles (bm 1, 2) are inserted on
the trochanteral apodeme, and they are responsible for a strong jumping action in the
species of this genus. III bm 1 in Psylla was probably originally a s-ex 5, which has
become a sternotrochanteral muscle by shift in the point of insertion. In Pyrilla
(Fulgoridae, Sander 1956) a pleurotrochanteral muscle (p-tr 1 ?) is responsible for
jumping.
initial fiber cleavage in other auchenorrhynchous Homoptera studied by Tiegs is similar.
A remarkable feature of the histogenesis here is the incorpor.ation of free myoblasts into
the cleaving fiber-rudiments. Thus, in Ery throneura, for instance, the mesothoracic
dorsal longitudinal muscle ( t 14), which consists of five fibers, arises from free myoblasts
that lie below the tergal wall. By the time the nymph is 0.9 mm long the number of
myoblasts increases to about 30. Of these myoblasts, five become increasingly
filamentous and grow along a column of myoblasts until they extend from one end to the
other and their ends f.use with the epidermis. Free myoblasts other than the five pioneer
myoblasts are, according to Tiegs, gradually incorporated into the five muscle rudiments.
In Cyclochilla (Cicadidae), according to Tiegs (1955), the flight muscles are of a
specific type, with the fibers arranged in small bundles in a nucleated sheath and with the
fibrils enclosed in complex lamellae. The structural dimensions of the flight muscles are,
however, not significantly different from the tubular type, the fibrillar diameter measuring
less than 1 ~ " In Cercopidae and J assidae the flight muscles are of the pseudofibrillar
type, with fibrillar diameters ranging from 1 to 2 ~ "
Johnson (1957, 1959) showed the histological changes occurring during the break-
down of flight muscles in alate aphids, Acyrthosiphon pisum, etc.
THE HEMIPTERA (HETEROPTERA)
The neck region: According to Larsen (1945c), the cervical sclerites are always
absent in Heteroptera. .In Leptocorisa, however, Akbar ( 1957) found a pair of blunt
sclerites developed from the inner wall of the episternum. These apodemes, according
to him, articulate with the corresponding processes from the lateral angles of the rim of
the foramen magnum (the postoccipital condyles). In Ramphocorixa Griffith (1945)
described the condylar surfaces on the postocciput and on the episterna that serve as
fulcrum and buffer when the head moves. It is not immediately clear whether these
structures in Leptocorisa and Ramphocorixa represent lateral cervical sclerites; it appears
more probable that they are secondarily produced articulatory devices.
In Heteroptera the dorsal neck membrane is often covered by the anterior evagina-
tion of the pronotum. In some Heteroptera the sclerotized neck region is long and
exposed, and in Helotrephidae (Esaki and China 1928) the head and prothorax are
fused. Parsons ( 1963) reported in Saldula a prothoracic spiracle in the neck region,
but now rescinds this observation (pers. comm.).
The pronotum: In Heteroptera the pronotum often evaginates posterolaterally. A
transverse internal ridge indicates the point of evagination of the lobe thus formed
(Hinterlappen of Larsen, 1945c, d). The posterior lobe is laterally the epimeron that
has become fused with the pronotum; thus it covers the wing base laterally, and the
mesonotum posteriorly. The evagination tends to extend beyond the mesonotum in such
families as Gerridae, Tingidae, etc.; in such cases the mesoscutellum is not prolonged (or
evaginated) and the pronotal posterior lobe functionally substitutes for the meso-
scutellum. Khanna's study ( l963d), which traced the development of these ridges in
Dysdercus, shows that in the second nymphal stage the pronotum is marked off by
internal ridges into very narrow anterior and posterior sclerites and a large median
sclerite. When these sclerites are compared with the pronotum in the adult, it is clear
that the posterior prolongation of the pronotum occurred in the area behind the posterior
transverse ridge; therefore, this area is not homologous with the postnotum. In wingless
forms of Gerridae the pronotal evagination does not occur (Matsuda 1960c), and this is
apparently true of other related families (e.g., Veliidae, Esaki and Miyamoto 1955).
The pronotum is often provided with an anterior internal ridge, which is marked
externally by a transverse suture. This ridge demarcates anteriorly the collarlike plate
that covers the membrane between the head and the pronotum proper in such groups as
Saldula (Parsons 1963). In Lethocerus, according to Parsons (1968), the anterior
evagination of the pronotum (cervical lobe of Parsons) overlaps the postocciput, and
two pairs of tendons tie in the neck membrane, which attaches to the inner ridge of this
lobe; the shorter pair of the tendons provide attachments for dorsal muscles.
The propleuron: In Heteroptera the propleuron is often indistinguishably fused
with the pronotum dorsally, and with the sternum ventrally. Thus, the three parts
together often form a prothoracic ring.
In aquatic Hemiptera (Fig. 111) according to Parsons (1967), the supracoxallobes
conceal the ventral end of the pleural suture, and only that region of the pleural suture
lying dorsal to the lobes is visible externally. The visible part of the suture is usually
continuous with the dorsal end of the coxal cleft, C. Internally, the point at which the
supracoxal lobes start their evaginations, B, varies in different aquatic Hemiptera.
In Parsons' group 1 (Notonecta, Hesperocorixa, Gelastocoris, Ochterus) B starts
internally at the level of C. The dorsal end of the pleural suture A clearly lies dorsal
to B. In group 2 (Belostoma, Lethocerus, Ambrysus, Ranatra, Nepa), however, the
point of evagination of the supracoxal lobes B lies at or near the level of the dorsal end
of the pleural suture (ridge internally). According to Parsons, the point B has moved
dorsally in group 2, and the supracoxallobes are only partly separated in Belostomatidae
and Naucoridae; in Nepidae they have fused entirely, and the coxal cleft has been
entirely eliminated.
Thus, in Parsons' view, the pleuron extends dorsally beyond the dorsolateral edge of
the prothorax, which usually would be considered as the lateral limit of the pronotum.
This interpretation is drastically different from that of Larsen (1945c, d), who asserts
that, in N epidae, the definitive lateral wall of the prothorax is the tergum that has
descended as far as the leg base.
as Parsons showed in some aquatic Hemiptera including Nepa, the
pleural ridge extends dorsally, clearly beyond the level of the lateral edge of the
pronotum. If the position of the dorsal limit of the pleural ridge is taken as the sole
reliable criterion in determining the border between the tergum and the pleuron, Parsons'
view is certainly valid. However, Parsons did not give a positive reason why the lateral
MA.TSUDA.; THE INSECT THORAX
RIM of COXAL
:-CAVITY
......,l
NOTONECTA
B: c ~ ~ VENTRAL END M
) ~ ' PLEURAL SUTURE:: <> :: . :::
~ - - - - ,, ......
__ _., .....
'
HESPEROCORIXA
?
A&B
/--A
GELASTOCORIS
c
OCHTERUS
BELOSTOMA )
....
A&B
A&B
LETHOCERUS
}
)
: : <: .:: .'::. :::: ..
A&B
c
)
AMBRYSUS
RANATRA
NEPA
269
FIG. 111. Diagrammatic external views of anterolateral prothorax in various Hydrocorisae.
Heavy broken lines indicate lateral angle of body; heavy stippling indicates two-layered,
evaginated supracoxal lobes, bordered dorsally by rim of coxal cavity, which lies internally
(from Parsons).
edge of the pronotum should not be the true lateral limit of the pronotum. She merely
suggested that the lateral edge of the pronotum in Belostoma, Lethocerus, and Nepa
could be a secondarily acquired strengthening ridge. Furthermore, in Coleoptera
'(p. 202) and Orthoptera (p. 170) the cryptopleuron, or the pleuron overgrown by the
pronotum, is a well-established fact. It is therefore possible, or even probable, that in
some Heteroptera the cryptopleuron occurs. In fact, besides her main argument,
Parsons also alluded to such a possibility.
The episternal part of the supracoxallobe is continuous with the anterior part of the
sternum and forms a precoxal bridge; the pleurosternal suture has completely disappeared
in the prothorax of Heteroptera. Neither the paracoxal suture nor the anapleural suture
is present in the propleuron. The trochantin beneath the supracoxallobe may be small;
it is never firmly articulated with the coxal margin. The anterior margin of the precoxal
bridge may be evaginated in some Heteroptera (e.g., Dysdercus, Khanna 1963a; Saldula,
Parsons 1963). This evagination (prepectus) is purely secondary, and cannot be
homologized with a subdivision of the episternum in lower insects; although in Dysdercus
Khanna (1963a) attempted to do so. The epimeral part may or may not be fused with
the pronotal posterior lobe; it also extends ventrally and may or may not form a postcoxal
bridge. Internally the pleural ridge often gives rise to a conspicuous coxal process.
The pleural arm is often united with the sternal furca to form a pleurosternal bridge
(Larsen l945c).
The prosternum: In most Heteroptera the posterior part of the sternum is broad,
and the furcae lie wide apart near the inner margin of the coxal base. The furcal bases
are connected by a transverse ridge, the sternacosta, in Leptocorisa (Akbar 1957),
Dysdercus (Khanna 1963a), and Coridius (Gael 1967). In these forms, therefore, the
sternum is externally divided into the anterior basisternum and the posterior furcasternum.
Often the posterior part of the presternum is produced posteriorly at the middle, or it
may be provided with paired processes (e.g., Naucoris, Rawat 1939). Larsen (1945c)
thought that the two processes in Nepa, Belostoma, Corixa, and others might represent
the spina; Akbar ( 1957) thought that in Leptocorisa the spina was merged into the
posterior part of the presternum.
Larsen's interpretation was based on the presence of a mesofurcal muscle that is
inserted anteriorly on the area in question (32 of Larsen). This muscle may appear to
represent a spina-furcal muscle (s 12). However, in Homoptera s 12 is absent in the
pro- and meso-thorax. It is probably the furca-furcal muscle (s 13 ), which is present in
Homoptera, that has shifted its anterior point of attachment. This interpretation is
supported further by the fact that in Heteroptera Larsen did not recognize a definitive
furca-furcal muscle (s 13). Thus, Larsen's identification of the spina was based on
misidentification of the muscle. In fact, the prothoracic spina is absent in Homoptera,
and for this reason alone' it is difficult to accept the presence of the prothoracic spina in
Heteroptera. Further, considering the general tendency for thoracic parts to evaginate,
the structures in question appear to be nothing but the modified ( evaginated) posterior
parts of the prosternum.
The mesonotum: The anterior part of the mesonotum is nearly vertically bent
ventrad. In Nezara (Fig. 112B), as Maloeuf (1932) showed, the is
demarcated posteriorly by the antecosta, which gives rise to the bilobed first phragma.
The prescutoscutal suture is clearly retained laterally, and the prealar arm is the lateral
extension of the prescutal area. The parapsidal suture has lost the direct connection with
the prescutoscutal suture, which is lost medially.
The prescutoscutal suture is present also in Lethocerus (Lauck 1959) and
Ptilomera (Matsuda 1960c); in Dysdercus (Fig. 112A) Khanna (1963c) showed the
laterally retained prescutoscutal suture. In Gelastocoris (Fig. 112C) and Saldula
(Fig. 113B) what Parsons (1960a) called the prealar membrane agrees in position with
the prescutoscutal suture. In Gelastocoris the parapsidal suture is also represented by a
membranous cleft. Quite often, in other Heteroptera, the prescutoscutal suture is com-
pletely lost, although the parapsidal suture is usually present.
As Maloeuf ( 1932) first pointed out for Heteroptera, the true prescutum is not the
area bounded by the parapsidal suture as earlier workers before him (Tower 1913;
Taylor 1918) thought. Yet, Larsen (1945c) and Parsons (1960a)"' continued to refer
to the area in question as the prescutum. Akbar's view (1957) that in Leptocorisa the
prescutoscutal suture extends as far as the anterior notal wing process is incorrect, as
Gupta's (1965) actual examination of the same species indicated. Even on a priori
grounds, Gupta's observation is acceptable, because the posterior part of the presumed
prescutoscutal suture reaching the anterior notal wing process must correspond to the
anterolateral scuta! suture, which is absent or obliterated in other Hemipteroidea
including Psocoptera, Thysanoptera, Homoptera, and other Heteroptera.
Another suture that frequently occurs on the mesonotum is a transscutal suture
(Fig. 112A) that connects the tergal fissure on both sides. According to Larsen
( 1945a), this suture occurs in Lygaeidae, Pyrrhocoridae, Corixidae, Nepidae, Reduviidae,
aiid Aradidae; this suture is present also in Pyrrhocoris (La Greca 1949b), Tropidothorax
(LaGreca and Cesaro 1951), Dysdercus (Khanna 1963a). Akbar (1957), in error,
called this transscutal suture the scutoscutellar suture.
"'Parsons (1963) withdrew this interpretation.
SCUTUM
SCUTOSCUTELLAR
SUTURE?
ANTE COSTA
PHRAGMA
PHRAGMA
FOREWING
PRESCUTUM
EPIMERON
EPISTERNUM
271
PRESCUTOSCUTAL
SUTURE
TRANSSCUTAL
SUTURE
PRESCUTUM
AL
PARAPSIDAL CLEFT
ANTE COSTA
MEMBRANE
SUTURE
XYPHUS
FIG. 112. A, mesonotum of Dysdercus koenigii (from Khanna); B, cephalic view of meso-
notum in Nezara viridula (from Maloeuf); C, cephalic view of mesonotum in Gelastocoris
oculatus (from Parsons).
In Nezara (the suture limiting the scutum from the scutellum of Maloeuf, 1932),
Naucoris (the scuta! suture of Rawat, 1939), and Coridius (the scutoscutellar suture of
Goel, 1967) the transscutal suture does not reach the lateral margin of the scutum,
although in Coridius the suture is laterally directed toward the tergal fissure.
The presence of the true scutoscutellar suture in Heteroptera is rare. In Eurygaster
and Odontoscelis, Larsen (1945c) claimed to have seen the true scutoscutellar suture,
which arises laterally from the hinge formed by the scutellum and the postnotum. In
Saldula, as Parsons ( 1963) showed (Fig. 113A), the presumed scutoscutellar suture is
obliterated laterally, and lacks the direct association with the axillary cord that is typical
of the scutoscutellar suture in other insects.
In Dysdercus (Fig. 112A) there is another transverse suture behind the transscutal
suture. Khanna ( 1963a) called this suture the scutoscutellar suture. It is not im-
mediately clear whether this suture corresponds to the scutoscutellar suture in Saldula.
In many Heteroptera the mesonotum posterior to the transscutal suture strongly
evaginates and forms a typical triangular scutellum, which is exposed between forewings.
LATERAL
SCUTAL FISSURE
(TERGAL FISSURE) --------..
ACROTERGITE
PARAPSIDALI
SUTURE:_
POSTEROLATERAL
"SCUTAL SUTURE
POSTNOTUM 2
TERGAL WING
GROOVE-
SCUTOSCUTELLAR
SUTURE?
METANOTUM
SCUTELLUM
PARAPSIDAL
SUTUR
TERGAL
FISSURE
PLEURAL WING
PROCESS
ABDOMEN
PLEURAL
I
POSTNOTUM 3
ANTERIOR & ANTEMEDIAN
NOTAL PROCESSES
LATERAL SCUTAL
PROCESS
KNOB
PLEURAL WING
PROCESS 3
'----EPIMERON 3
AXILLARY
CORD
POSTNOTUM 2
METANOTUM
OTUM 3
PLEURAL
SUTURE
i----EPISTERNAL
LOBE 3
FIG. 113. A, dorsal view of pterothorax in Sa/du/a pa//ipes (from Parsons); B, lateral view
of pterothorax in Saldula pal/ipes (from Parsons).
MATSUDA: THE INSECT THORAX Z73
In Saldula (Fig. 113A) Parsons ( 1963) called a fissure arising from behind the
anterior notal .wing process the lateral scuta! fissure, and the area demarcated by this
fissure the lateral scuta! process. According to Parsons ( 1963), both the lateral scuta!
fissure and the lateral scuta! process are present in Hesperocorixa and Notonecta, although
they are absent in Gelastocoris (Parsons 1960a). In Saldula the suture lateral to this
fissure demarcates posteriorly the tergal wing groove. This suture is probably
homologous with the posterolateral scuta! suture in Homoptera that forms the tergal
wing groove. The tergal wing groove is, then, homologous with the posterolateral scuta!
area in Homoptera and other insects. When the insect is at rest, the groove receives the
posterior margin of the forewing. In Gelastocoris (Fig. 115B) the groove becomes
larger and membranous anterolaterally, and the membrane continues into the wing. In
other Heteroptera also the wing groove is formed in corresponding positions.
In Saldula (Fig. 113A) and Gelastocoris (Fig. 115B) a narrow process from the
anterior end of the tergal wing groove extends ventrally and is fused with the subalare.
These findings of Parsons support Larsen's (1945c) generalization that in Heteroptera
the scutellar process" fuses with the suba!are. In Pyrrhocoris (LaGreca 1949b) a small
process (processo spiralto, Fig. 114B) apparently connects with the subalare; in Coridius,
according to Goel (1967), the scutellar process gives support to the subalare and the
axillary cord.
Often in Heteroptera a median longitudinal suture of varying length is present on
the mesonotum. It may extend anteriorly even to the first phragma (Fig. 112B). The
dorsal median longitudinal muscle (t 14) lies along the ridge formed by this suture
internally.
The well-developed postnotum is medially hidden by the scutoscutellum; however,
it is often laterally exposed (Fig. 113B) and joined to the epimeron to form the postalar
bridge. In Gelastocoris it is also united with the metathoracic episternum and in Saldula
(Fig. 113B) it is fused, along the intersegmental suture, also with the metepimeron.
Posteriorly the postnotum continues into the anterior wall of the second phragma; this
phragma is usually greatly developed on either side of the midline.
The subalare, according to Larsen ( 1945 c), is present in most terrestrial Heteroptera,
although in most aquatic Heteroptera it is absent; Notonecta was the only aquatic genus
in which Larsen found the subalare, but it occurs also in Gelastocoris (Fig. 113A).
The subalare has a well-developed apodeme, to which t-cx 8 is attached.
The mesoscuta/lateral margin and the forewing base structures: In Tropidothorax
(Fig. 114A) all notal wing processes except the posterior notal process can be recognized.
The tergal fissure (Tergalspalt of Larsen, emarginazione of La Greca) behind the
antemedian notal process continues into the transscutal suture. In Pyrrhocoris
(Fig. 114B) the postmedian and posterior notal wing processes are not differentiated.
In aquatic Heteroptera the anterior three notal wing processes are recognized. A
feature common to all Heteroptera is the absence of the posterior notal wing process, as
Larsen (1945c) and Parsons (1960a, 1963) maintained. The idea of Rawat (1939),
Akbar (1957), and Khanna (1963a) that the lateral limit of the lateral scutellar process
represents the posterior notal process is invalid. The tegula is absent in the mesothorax
of Heteroptera.
In Tropidothorax (Fig. 114A) and Pyrrhocoris (Fig. 114B) the first axillary sclerite
articulates dorsally with the anterior and median notal wing processes on the scuta!
lateral margin; in Gelastocoris (Fig. 115B) and Saldula (Parsons 1960a, 1963) the
small first axillary sclerite also has the same association with the scuta! lateral margin.
In all four genera referred to, the dorsal second axillary sclerite is fused with both the
basisubcostale (humeral plate of Parsons) and the median plate. This fusion is a
characteristic of the forewing base in Heteroptera. The third axillary sclerite (fourth
axillary sclerite of Parsons) is fused or connected with the subalare in Heteroptera. In
61
The posterior margin of the tergal wing groove may be considered as scutellar, since it is continuous with
the posterior margin of the forewing base, and corresponds to the scutellar arm which continues to the
axillary cord in other insects.
BASIRADIALE
l'ST AXIL. SCL.
3'RD AXIL. SCL.
2'ND AXIL. SCL.
TERGAL
WING GR OrOVE
BASI RADIALE -------::::-----,
MEDIAN
PLATE
TERGAL ~ -
WING G R O O V E ~
BASICOSTALE
SUBALME
EPIMERON
A
ANTERIOR
NOTAL PROCESS
ANTEMEDIAN
NOTAL PROCESS
POSTMEDIAN
NOTAL PROCESS
BASISUBCOSTALE
-ANTERIOR
.;:--<'$'?t.=----- NOTAL PROCESS
B
c
2'ND AXIL. SCL.
TRANS SCUTAL
SUTURE
PROCESSO
SPIRAL TO
BASI RADIALE
VENTRAL
2'ND AXIL. SCL.
3'RD AXIL. SCL.
FIG. 114. A, forewing base in Tropidothorax leucopterus (from La Greca and Raucci); B,
dorsal view of forewing base in Pyrrhocoris apterus (from La Greca); C, lateral view of
forewing base in Pyrrhocoris leucopterus (from La Greca).
Pyrrhocoris (Fig. 114C) the usual association of the ventral second axillary sclerite with
the pleural wing process occurs.
The mesopleuron: The mesopleuron in most Heteroptera is indistinguishably fused
with the mesosternum. However, Taylor (1918) thought that in Notonectidae,
Reduviidae, Emesidae, Cimicidae, Phymatidae and Tingidae, a faint line demarcated the
sternum from the pleuron. Such a faint pleurosternal suture appears to be present in
Cimicidae (Larsen 1945a; Usinger 1966). Matsuda (1962) also thought that an
oblique suture on the ventral surface of the mesothorax in Notonecta (Fig. 116A) and
Hesperocorixa was a suture that separates the pleuron from the sternum. Whether these
claimed pleurosternal sutures really represent the pleurosternal sutures, however, remains
open to question. The apparent pleurosternal suture in Gerridae may also be a
secondarily produced suture. However, the suture is more distinct and complete in
more primitive genera in this family (Matsuda 1960c).
The pleural suture and its internal ridge are well formed, especially in some aquatic
Heteroptera. In some semiaquatic Heteroptera and many terrestrial Heteroptera, only
the pleural (coxal) process, the pleural arm, and the pleural wing process are present
(Larsen ( 1945c).
As in the propleuron, the ventral part of the mesopleuron has become evaginated to
form the supracoxal lobes covering the coxal base. This area is split by a longitudinal
fissure (coxal fissure) that gives, as Taylor (1918) first demonstrated, a spurious appear-
ance of being the ventral part of the pleural suture. The pleural arm arises further
dorsally on the pleural ridge than in most orders, often near the dorsal end of the ridge.
Neither the paracoxal nor the anapleural suture has been identified with certainty
in the mesoepisternum of Heteroptera. In Saldula (Fig. 115A) the t-ridge of Parsons
(1963) is a free internal ridge from the level of the pleural (coxal) process that merges
ventrally with the sternum. The suture formed by this ridge externally represents the
area where the supracoxal evagination starts. Another short ridge (Fig. 115A) lying
anterior to the t-ridge may possibly represent the remnant of the paracoxal ridge.
According to Larsen ( 1945c), the well-isolated basal are is absent in the mesothorax:
of Heteroptera. A partially isolated dorsal plate in Notonecta (Fig. 116A) was called
the basalare by Matsuda (1962); Larsen (1945c) also recognized the basalare in
Notonecta. The apparent basalare in Sa/dula (Fig. 115A), according to Parsons ( 1963),
fits under the pleural wing process and articulates with the humeral plate. In many
Heteroptera the dorsal part of the episternum is incompletely separated or marked off by
the internal ridge from the pleural wing process.
The trochantin lies beneath the supracoxal lobe; it is present even in a highly
specialized, parasitic family, Polyctenidae (Ferris and Usinger 1937).
In the mesothorax of Heteroptera, the epimeron is usually much smaller than the
episternum and does not form the postcoxal bridge. The sutureon the mesepimeron of
Xenocorixa (Fig. 116B), which Matsuda (1962) considered to be the paracoxal suture,
is probably homologous with the transepimeral suture on the epimeron in the cicada
(Fig. 109A); in Corixidae, according to Hungerford ( 1948), this transepimeral suture
occurs only in this genus. In Saldula (Fig. 115A) the ridge m may correspond to the
transepimeral suture.
In Saldula (Fig. 113B) the free edge of the evaginated epimerallobe curves sharply
upward, producing a high ridge. The ridge ends in a knoblike projection that lies in the
region of the postalar bridge between the epimeron and the postnotum; and the knob fits
into a depression in the costal margin of the resting forewing. Comparable wing-holding
mechanisms occur in o t ~ e r Heteroptera. For more information about the wing-holding
mechanism, see Poisson (1924), Weber (1930), Larsen (1949a), Gaumont and Moreau
(1961), and Ewald (1963).
In Corix:idae the epimeron is folded into two lobes, outer and inner. The outer
lobe (evagination) often completely covers the inner lobe, which bears the tympanum
62
fi2'Jbe tympanum in Corixidae was first found by Hagemann (1910). This specialized sense organ is a scolo-
pophorous organ. Larsen (1957) and Parsons (1962) have summarized the knowledge of this organ which is
also present in the metathorax and abdominal segments of Heteroptera. '
EPISTERNUM BASALARE
+ BASISTERNUM
FURCA
COXAL CAVITY
PREALAR MEMBRANE
ANTERIOR &
ANTEMEDIAN
PROCESS
TERGAL FISSURE
-----EPIMERON
-EPIMERAL LOBE
A
l'ST AXIL. ACL. HUMERAL PLATE
\
3'RD AXIL. SCL.
POSTERIOR NOTAL PROCESS
FIG. 115. A, ventral view of pterothorax in Saldula pallipes (from Parsons); B, dorsal view
of pterothorax in Gelastocoris oculatus (from Parsons).
and spiracle. However, in a primitive Australian genus of Corixidae, Diaprepocoris
(Fig. 117D), the outer lobe covers incompletely the inner lobe (Matsuda 1962).
The mesosternum: Apart from the possible pleurosternal suture mentioned earlier,
the approximate lateral limit of the mesosternum is recognized by the position of the
furcal pits, which lies along the widely separated inner coxal margins. Hence, in most
Heteroptera the presumed mesosterna! area is relatively large.
The anterior limit of the mesosternum is sometimes delimited by a secondary
evagination along the ridged anterior margin, and is continuous with the prepectus of the
pleuron: Khanna ( 1963a) described this condition in Dysdercus, erroneously calling it
the presternum. In most Heteroptera the anterior sternal surface is the basisternum,
which is indistinguishably fused laterally with the pleuron. This area is often depressed
to accommodate the forelegs, and Gupta ( 1963) called this depressed area in Saldula
and Aepophilus the presternum.
The basisternum is often provided with a median longitudinal ridge, as can be seen
in Belostomatidae, Naucoridae, Notonectidae, Corixidae (Larsen 1945a), Saldula
(Parsons 1963), etc. According to Larsen ( 1945a), the species without the ridge or
with a weakly developed longitudinal ridge tend to have widely separated furcae.
Conversely, in species with a strongly developed longitudinal ridge the distance between
the furcae tends to be shorter. The furcal bases are often connected by the transverse
sternacosta, as in Nezara (Maloeuf 1932), Saldula (Parsons 1963), etc. The furca has
one or two branches; in some forms it is fused with the second phragma.
In Heteroptera, as in Homoptera, the second spina has been lost. Hence, the furca
and the sternacosta (when present) represent the posterior limit of the mesosternum, as
Larsen (1945c) maintained. In most Heteroptera the posterior portion of the meso-
sternum is produced posteriorly at the middle, and is called the xyphus.
The metanotum: The metanotum is nearly always smaller than the mesonotum, and
in many groups it is hidden beneath the mesoscutellum. The anterior margin is medially
continuous with the posterior wall of the second phragma, thereby often forming a deep
cleft behind the latter (Larsen 1945c). Contrary to Larsen's (1945c) generalization
that the prescutum is absent in the metathorax in Heteroptera, a reduced prescutum is
retained laterally in Pyrrhocoris (Fig. 117B), as La Greca ( 1949b) has pointed out.
Larsen ( 1942) found that in Mesocerus the lateral part of the metanotum is isolated
as a triangular plate from the rest of the metanotum by a narrow membranous slit.
This plate is connected anteriorly with the posterior wall of the second phragma and
laterally with the wing base. Four muscles, one from the tergum and three from the leg,
are inserted on the plate. According to Larsen 1945c), the separation of the antero-
lateral part of the metanotum occurs in most terrestrial Heteroptera and in Ranatra. A
comparable separation of the metanotal part in Saldula (Parsons 1963) is the antero-
lateral edge, which can flex upon the rest of the metanotum. Parsons ( 1960a) also
noted in Ge/astocoris a similar separation of the anterolateral angle of the metanotum.
In many terrestrial Heteroptera, the greatly reduced medial part of the metanotum
is fused in varying degree with the second phragma (Larsen 1945a), or with the meso-
thoracic postnotum (Pyrrhocoris, La Greca 1949b). In some Heteroptera the medial
reduction of the metanotum is highly pronounced. For instance, in Mesocerus (Larsen
1942), the scutoscutellum is represented by a transverse ridge that connects the lateral
triangular plate on each side.
In some aquatic Hemiptera, in which the metanotum is well retained medially, a
median longitudinal sulcus often diverges posteriorly into the transverse suture and
delimits the postnotum. In Ramphocorixa, as Griffith (1945) showed (Fig. 117A), the
posteriorly diverged part of this suture is principally the line that demarcates the post-
notum posteriorly. In Gerridae (Matsuda 1960c) the median longitudinal part of the
corresponding suture may be obliterated posteriorly, but its posterior divergences are
merely the line between the metanotum and the first abdominal tergite. Larsen ( 1945c)
and Parsons ( 1960a) called this suture (ridge) in some aquatic Heteroptera the scuto-
scutellar suture. The ridge is laterally continuous with the axillary cord as the
scutoscutellar suture, but medially it does not assume the typical position and shape of
the scutoscutellar suture.
The metathoracic postnotum is often larger than the rest of the metanotum.
Laterally, it forms the postalar bridge in many Heteroptera. The third phragma is con-
tinuous from the postnotum, and is well developed in some aquatic Hemiptera.
278
The metascutal lateral margin and the axillary sclerites of the hind wing: Apart
from the anterior notal wing process, which may be detached along the anter-olateral
corner of the metanotum (already discussed), the lateral margin of the metanotum is
generally straighter than the mesonotal lateral margin, the median and postmedian notal
wing processes being absent or undifferentiated. Larsen (1945c) denied the presence
of the undetached posterior notal process, which was, however, found in Gelastocoris
(Fig. 115B; Parsons 1960a) and Saldula (Parsons 1963). In these genera the acute
posterolateral angle of the metanotum, which represents the posterior notal wing process,
articulates with the proximal end of the third axillary sclerite. According to Larsen
( 1945c), the fourth axillary sclerite, which represents the detached posterior notal wing
process, is present in Notonecta.
The axillary sclerites in Pyrrhocoris (Fig. 117B) are similar to those in the forewing
in having a large definitive humeral plate, which presumably consists of the basicostale,
basisubcostale, and the anterior part of the second axillary sclerite, and the undivided
median plate; the dorsal third axillary sclerite is large. In Gelastocoris (Fig. 115B) and
Saldula, as Parsons ( 1960a, 1963) showed, the third axillary sclerite is also large as in
Pyrrhocoris; the definitive humeral plate also is large in these genera, and in Gelastocoris
the median plate appears to consist of a small sclerite and the strongly thickened base of
wing veins. Schaefer (1962) described, in some Coreidae, Pentatomoidea and Larigidae,
a spine arising from the third axillary sclerite, and denied its stridulatory function, which
had been suspected. Puchkova (1961) called the definitive humeral plate in Holosalda
erroneously the first axillary sclerite.
The metapleuron: The pleural ridge is well formed in some aquatic and semiaquatic
Hemiptera; in some terrestrial Heteroptera the ridge is often weakly developed, being
represented externally by the dorsal part of the pleural suture and by the pleural wing
process. According to Larsen ( 1945c), the pleural arm is absent in the meta thorax of
Heteroptera.
The episternum is much larger than the epimeron, whcih is often greatly reduced.
In most Heteroptera the original ventral limit of the episternum has been lost by fusion
with the metasternum. However, in Dysdercus, Khanna (1963a) showed a longitudinal
suture, which, by its position, appears to be the well-marked pleurosternal suture.
In some Heteroptera the stink (scent) groove appears to lie approximately on the
area that corresponds to the paracoxal suture. Brindley ( 1934) thought that the location
of the scent groove in Cimicidae corresponded to the area (or suture) between the
precoxale (anepisternum plus preepisternum) and the antecoxale in lower insects. In
the terminology used in this work, this area corresponds to the paracoxal suture. On
the other hand, however, Carayon ( 1962) suggested that the stink groove was primitively
an unpaired, ventral invagination on the intersegmental membrane between the meta-
thorax and the abdomen.
The locations of the stink gland opening and of the groove vary greatly in different
Heteroptera. In a primitive corixid genus, Diaprepocoris from Australia (Fig. 117D)
and Tenagobia, the groove extends farther anteriorly than in Notonecta (Fig. 116A),
and this anterior extension does not correspond to any suture in lower insects. In
Saldula (Fig. 115A), as Parsons (1963) observed, the stink groove is horizontal as in
many other terrestrial Heteroptera, and the stink apparatus
03
has a single, medial opening
as in most Amphibiocorisae. In Gelastocoris (Fig. 117F) the stink groove is anteriorly
continuous with the smooth posterior edge of the mesothoracic posterior epimeral lobe.
This edge is so placed that the secretion emerging from the stink groove flows out upon
it; Parsons ( 1960b) thought that it would probably serve as an evaporating surface for
the secretion. In Anthocoridae, according to Brindley (19 34), a small pocket on the
anepisternal area serves to retain the scent secretion. Variations of the stink groove
OSThe stink apparatus consists of a pair of tubular g l a n d s ~ which communicate with a reservoir of variable
form that may or may not be furnished with an accessory gland that opens to the exterior, typically through
two orifices lying on the metapleuron ventral to the anterior margin of the metacoxal cavities. Detailed
anatomical studies of the stink gland were made by Henrici (1940) and Betten (1942). For variations, refer
to Brindley (1930), Wells (1954), Gupta (1961), Carayon (1962), Waterhouse and Gilby (1964).
PLEURa-
STERNAL
SUTURE?
ANAPLEURAL
CtEFT?
STINK GROOVE
BASISTERNUM
-+EPISTERNUM
COXA
PRESTERNUM
+BASISTERNUM
KATEPI
BASALARE
TROCHANTIN
KATEPISTER-
NAL LOBE?
TRANSEPIMERAL
SUTURE
STINK GROOVE
279
FIG. 116. A, ventral view of pterothorax in Notonecta lobata (from Matsuda); B, ventral
view of pterothorax in Xenocorixa vittipennis (from Matsuda).
within Aradidae were shown by Usinger and Matsuda (1959). In some genera of
Gerridae ( Gigantometra, Brachymetra, etc.), the stink groove is continuous from each
side to the median single opening, the omphalium; in other genera of Gerridae the groove
is absent and only the omphalium is present (Matsuda 1960c). According to China
(1955), some other water striders, such as Hebroveliidae, Oce/lovelia, and Heterocleptus,
have dual openings.
The clearly identifiable anapleural cleft, or suture, is absent in the metapleuron of
Heteroptera. However, in Notonecta (Fig. 116A) an oblique, short, membranous cleft
may, judging from its position, be the anapleural cleft. In most Heteroptera the
trochantin is present beneath the supracoxallobe. In Gelastocoris (Parsons 1960a) the
pleural process has an extension that articulates with the trochantin. The basalare is
absent, at least in most Heteroptera.
The metepimeron is often greatly reduced. Matsuda (1962) showed a sequence of
reduction of the epimeron and the concomitant formation of the air chamber" in Noto-
nectidae and Corixidae. In Notonecta (Fig. 117C) the pleural suture separating the
anepisternum from the epimeron is distinct, and the epimeron is relatively broad. In a
primitive corixid genus, Diaprepocoris (Fig. 117D), the anepisternum and the epimeron
are narrower than in N otonecta, but they are distinctly separated by the oblique pleural
suture. In Hesperocorixa (Fig. 117E) the greatly narrowed anterior half of the
epimeron and the similarly narrowed anepisternum together form a groove, which in turn
continues to the space formed below the outer epimeral lobe of the mesothorax to form
the air chamber. The pleural suture is recognizable in this groove.
In Hesperocorixa (Fig. 117E) the posterior part of the epimeron is prolonged and
distinct from the anterior (or often dorsal) part of the epimeron which is narrowed.
This secondary separation of the epimeron apparently occurs in some other Heteroptera
(e.g., Ranatra, Larsen 1945a). In Pyrrhocoris (La Greca 1949b), Gelastocoris and
Saldula (Parsons 1960a, 1963), and probably in many others the epimeron joins the
postnotum posterolaterally to form the postalar bridge.
The postcoxal bridge is not formed by the epimeron; according to Brindley (1938),
the first abdominal segment forms the definitive postcoxal bridge.
The metasternum: In some Heteroptera such as Leptocorisa (Akbar 1957), the
metasternum is externally divided into anterior and posterior parts by the sternacosta,
which connects the metafurcal bases internally. In Gelastocoris (Parsons 1960a) and
many other Heteroptera, the posterior portion of the metasternum is often medially
produced to form the xyphus. According to Larsen (1945a), the metat!:10racic basi-
sternum in Heteroptera lacks the median longitudinal ridge; in Dysdercus (Khanna
1963a), however, it is clearly present. According to Larsen (1945c), in most aquatic
Hemiptera the metathoracic furca is as large as the mesothoracic furca, whereas in
terrestrial Heteroptera the metafurca is usually smaller than the mesofurca. In Ranatra
(Larsen 1954d) a few additional sternal apophyses occur.
Modifications in short-winged and wingless forms: Larsen's ( 1950) comparison of
thoracic structures in the macropterous, hypomacropterous (forewing reaches the sixth
abdominal segment in the female), and micropterous forms in Aphelocheirus aestivalis
shows that the secondary sclerotization in the micropterous form includes mainly the
membranous area between the tergum and the pleuron and the membranous wing base;
increased sclerotization in the short-winged (brachypterous) forms also obliterates the
fissures and sutures on the tergum, and the reduction in sizes of structures includes the
phragma, prealar bridge, etc. LaGreca's (1949b) comparison of the winged form with
the brachypterous form of Pyrrhocoris apterus showed increased sclerotization in the
axillary region. In Gerridae, as Matsuda ( 1960c) showed, the definitive intersegmental
sutures between the mesothorax and the metathorax in wingless forms of the different
major groups represent different combinations of the sutures that are present in
winged forms.
The anatomical basis of flight: In Heteroptera the fore- and hind-wings function
simultaneously, since they are fastened together by special mechanical devices (Poisson
1924; Larsen 1949a; Gaumont and Moreau 1961; Ewald 1963). Therefore, the move-
ment of the hind wing is affected to a large extent by the action of the mesothoracic
muscle. In Ranatra, according to Larsen ( 1949a), muscle t 14 in the metathorax is not
a flight muscle, but is the levator of the abdomen.
Contraction of indirect flight muscles in the mesothorax (t 14, t-p 5 or 6), according
to Larsen ( 1949a), disengages the forewing from the wing-holding devices, and causes
the forewing to spread sideward. The hind wing also spreads passively, because of its
MFor more information about the structures and physiology of the air chamber, see Dogs (1908), Bracher
(1909). Hoppe (1912), Ege (1915), Larsen (1924), Rawat (1939), Griffith (1945), and Popham (1960b).
EPISTERNUM
AXILLARY
CORD
EPIMERON
SCUTUM+
SCUTELLUM
POSTNOTUM 3
ANTECOSTA
FURCA
A
c
OUTER LOBE
of EPIMERON 2

PLEURAL
SUTURE 3
E
EPISTERNUM 2
PIRACLE
PLEURAL SUTURE 2
STINK
GROOVE
COXAL CAVITY
ANTEROMEDIAL
FLAP of STINK
GROOVE
XYPHUS
F
281
PRESCUTUM
8
3'RD AXIL. SCL.
'
SUBALARE
OUTER EPIMERON
INNER EPIMERON
TYMPANUM
AN EPISTERNUM 3
PLEURAL
SUTURE 3
D
SPIRACLE
EVAPORATING
SURFACE
COXAL CAVITY 3
FIG. 117. A, metanoturn in Ramphocorixa acuminata (from Griffith); B, hind wing base in
Pyrrhocoris apterus (from LaGreca); C, lateral view of meso- and meta-pleuron in Notonecta
lobata (from Matsuda); D, lateral view of meso- and meta-pleuron in Diaprepocoris brachy-
cepha/a (from Matsuda); E, lateral view of meso- and meta-pleuron in Hesperocorixa laevigata
(from Matsuda); F, ventral view of meta thorax in Gelastocoris oculatus, showing stink groove
(from Parsons).
282
connection with the forewing. Barber and Pringle (1966) have found experimentally,
however, that in belostomatids t 12 is primarily responsible for unlocking the wing; t 14
and t-p 5 or 6 are responsible for wing unfolding. In some Heteroptera p 3 is an
extensor and promotor of the wing. Maloeuf ( 1932) thought that in Nezara p-tr 2
substituted for the function of p 3, which is absent. In Ranatra (Larsen 1949a), how-
ever, p-tr 2 cannot replace p 3 functionally.
As in Orthoptera, the wing base, hidden beneath the posterior lobe of the pronotum,
must be exposed before the wings can spread. This is made possible by the contraction
of a prothoracic muscle, op-t 3, which raises the pronotal lobe. When the wings are
brought back to the resting position, contraction of the prothoracic cv (d) -t 1 brings the
pro notal lobe down against the wing base. According to Larsen (1949a), t-p 13 in
Ranatra cannot by itself function as a flexor, and he suggested that it would control
direction in the movement of wings during flight.
The metathoracic muscle concerned with movement of the wing is 57 of Larsen
(t 14?), which in some Heteroptera is inserted on the anterolaterally or laterally isolated
metanotal plate. This isolated area is connected with the wing base in such a way that
the pull of this muscle causes sideward spread of the wing and elevation of its anterior
margin. In Ranatra, according to Larsen ( 1949a), this muscle is absent, although the
anterolateral metanotum is isolated. Larsen has stated that in Ranatra, t-tr 1 and t-cx 2,
which are attached to this area, replace the function of the dorsal muscle (t 14?).
Thus, some leg muscles in Heteroptera, as Larsen ( 1949a) pointed out, are bifunctional.
The adult musculature: The literature on the thoracic musculature in adult Heterop-
tera is as follows: Maloeuf (1932) in Nezara viridula, Maki (1938) in Sigara substriata
and Eurostus valid us, Rawat (1939) in Naucoris cimicoides, Larsen ( 1945d) in 29
species of Heteroptera, Larsen ( 1949a) in Ranatra /inearis, Short (1953) on the leg
muscles of Corixa punctata, Sprague ( 1956) in Hydrometra martini, Akbar ( 1957) in
Leptocorisa varicornis, Lauck ( 1959) in Lethocerus spp., Parsons ( 1960a, b )''in Ge/asto-
coris oculatus, Brinkhurst ( 1960) in Gerris najas, Khanna ( 1963b) in Dysdercus
koenigii, Parsons (1963) in Saldula pallipes, Goel ( 1967) in Coridius janus, Parsons
( 1968) on the prothoracic muscles in Lethocerus. In Table XVI homologies of the
muscles recognized by Larsen ( 1945d) are shown.
The thoracic musculature in Heteroptera is a reduced version of the musculature in
auchenorrhynchous Homoptera. Flight muscles are fewer in number. The muscle
that dilates the stink gland irt terrestrial Heteroptera is a unique muscle, which probably
represents a modified sternal or sternocoxal muscle.
Gross aspects of the postembryonic muscle development: Larsen (1945c)
generalized that the nymph always has more muscles than the adult. In Ranatra,
according to him, M 9 (t-s 1), M 32 (s 13), M 35 (t-s 1), 39 (p-s 1), 56 (t 14), and
60 (t-s 1) in the nymph become lost; in Notonecta, Nepa, and Naucoris also six
nymphal muscles become lost (Larsen 1945c); in Pentatomidae (Maloeuf 1932; Larsen
1945c) s 13 in the nymph becomes lost in the adult.
In Gerris (Larsen 1945c) M 32 (s 13), M 33 (p-s 5), and M 35 (t-s 1 or 2) in the
nymph are present in the young adult, but they become lost in the older adult. Similarly,
in Pentatomidae (Larsen 1945c) s 13 is present in the nymph and the young adult, but
it soon disappears.
In the nymph of Notonecta a metathoracic muscle connecting the pleural arm with
the metanotum disappears in the adult. Larsen (1945c) thought this muscle was
homologous with 66 of Make! ( 1942) in Pseudococcus, which is an elevator of the hind
wing.
Khanna ( 1963c) observed in Dysdercus koenigii that, during growth of the fifth
instar, the mesal dorsal muscle becomes t 14, and the external muscle becomes t 12; the
ventral oblique muscles (p-s 5, p-s 9) in the nymph are lost during the final molt; a
tergopleural muscle (lateral muscle of Khanna) in the mesothorax splits into two during
the final molt, and forms a tergoepisternal muscle (t-p ?) and the axillary muscle
(t-p 13 ?) ; a tergotrochantinal muscle (t-ti 2 or 3) shifts its ventral point of attachment
to the sternum after the third stage to become t-p 5 or 6; a tergofurcal muscle (t-s 1) is
lost in the adult; and a posterior tergocoxal muscle splits into two, and the anterior part
becomes attached to the subalare ( t-cx 8). These observations clearly indicate that there
is a tendency for muscles to become reduced in number in the adult stage, due to the
degeneration of muscles during the postembryonic development and even after the adult
stage is reached (as in Orthoptera).
No caenogenetic trend is evident, since nearly all nymphal muscles are homolo-
gizable with adult muscles of Heteroptera, or of other orders. In the nymph of
Dysdercus, p-s 9 is present only rarely in adult insects. It may be said that in Heterop-
tera the nymphal musculature is generally more primitive than the adult musculature,
because these degenerating muscles are present in the adults of other Heteroptera and
lower orders including Homoptera.
It is known that some fully winged Heteroptera are unable to fly. In Nepa and
Ranatra, Ferri ere ( 1914), Brecher ( 1916), and Poisson ( 1924) found that flight
muscles are more or less reduced in some individuals of the macropterous form. Larsen
(1950, 1955) also observed reduction of flight muscles in fully winged flightless
individuals of many heteropterous genera, including Naucoris, Nepa, Corixa, and some
terrestrial Heteroptera.
60
More recently, Young's (1961, 1965a) studies on Corixidae
and Notonectidae have shown that the muscles t 14, t 12, and t-p 5, 6, are reduced in
flightless forms. Young (1965c) has further shown that in Corixidae the reduced flight
muscles result from failure of development after eclosion rather than from degeneration
of normally developed muscles; he ( 1965b) thought that temperature was a contributing
factor in the reduction.
Histology and development of flight muscles: In Gerris lacustris, according to
Poisson ( 1924), the dorsoventral flight muscle ( t-p 5, 6) is formed from myoblasts in
the old third nymphal instar. During this stage two dorsoventral bridges of connective
tissue appear on both sides of the digestive tube in the mesothorax, which direct the
formation of the muscles. The myoblasts themselves assemble first near the intestine,
not along the bridges. They then multiply by mitosis before becoming fibers in the
direction of the bridges; during the fourth stage they form a syncytium. During the
fifth stage, myoblasts become very abundant and fuse with fibers that have already
been produced. Thus, the dorsoventral flight muscle is formed by fusion of myoblasts.
Young's (1965c) study of Sigara shows that the development of flight muscles in the
teneral adult is by enlargement of the fibrils already present at eclosion.
Poisson's (1924) figures show that the dorsal longitudinal muscles (t 14) in
Ranatra and Notonecta are of the fibrillar type. In various nymphal Heteroptera
Larsen (1945c) found that M 30 (t 14), M 31 (t 12), M 34 (t-p 5 or 6), M 37 (p 3),
M 57 (t 14?), M 43 (t-cx 8), M 67 (t-cx 8), M 38 (t-p 13), M 61 (t-p 13),
M 39 (p-s 1), and M 36 (t-p 8?) are atypical, nonstriated, and nonfunctional.
THE NEUROPTERA
The neck region: In Neuroptera the lateral cervical sclerite is of different origins in
different groups. In Sialis (Fig. 118A), in which neither the prothorax nor the neck
region is especially prolonged, the lateral cervical sclerite is obviously the preepisternum
that is continuous from the anepisternum, and it has become detached by a narrow
membranization from the katepisternum. It is connected anteriorly with the ventro-
lateral angle of the postocciput. The location of the sclerite relative to the propleural
parts and to the head is thus similar to that of Micropteryx (Fig. 144A), where it also
appears to be nothing but a modified preepisternum. In Micropteryx, however, the
preepisternal lateral cervical sclerite is separated by the anapleural cleft from the
anepisternum.
05
Larsen (1950) in Ranatra and Parsons (1960a) in Gelastocoris found that some macropterous individuals
with fully developed flight muscles can never fly.
'284
TABLE XVI
Thoracic musculature in Heteroptera
Heteroptera (Larsen 1945d) Muscle designations Remarks
4 cv(d)-t 1
op-t 1
2 op-t 2
3 op-t 3
10 op-p 1
7 op-s 1
6 s 1 or 2
5(I) t 12
31 (II)
3, 4 t 14 1
30
56, 57
34 (I I) t-p S or 6
36 (II) t-p 8
38 (II) t-p 13
61 (III)
5 (I) t-p 17 ?
ll(I) t-p ?
9 t-s 1 or 2
35
60
13 t-ti(cx) 2
40
63
14 (I) t-ti(cx) 3
16 t-cx 6 or 7
41
64
15 t-cx 8
43
67
37 (II) p3
12 p-s 1
39
62
33 (II-III) p-s 5
65 (III) p-ti(cx) 1
42 (II) p-cx4 2
66 (III)
17 (I) p-cx 5
32 (HI) s 13 3
58 (II-III)
8(I) s 16
80 (III-2A) s 20
59 (III) s-stg. 4
19 s-ex 3
45
69
18 s-ex 5
44
68
Heteroptera (Larsen 1945d)
20
46
70
21 (I)
47 (II)
71 (III)
22
48
72
REMARKS (Table XVI)
TABLE XVI (Continued)
Muscle designations
t-tr 1
p-tr 1
p-tr 2
s-tr 1
( 1) 57 is inserted anteriorly on the movable lateral metanotal plate.
285
Remarks
(2) These muscles arise from the anterior area of the episternum, but judging
from their insertion near the pleural process, they are probably p-cx 4.
(3) 32 is inserted anteriorly on the mesothoracic furcal arm or on the xyphus.
( 4) 59 arises from the metafurcal arm and is inserted on the stink gland sac, and
functions as a dilator of the sac; this muscle may be a modified s 16. The muscles of the
stink gland sac in Gelastocoris (Parsons 1960b) and in Saldula (Parsons 1963) are not
homologous with 59 of Larsen.
Besides the muscles recognized by Larsen (1945d), the following muscles have
been found in Heteroptera: 10 Bin Gelastocoris (Parsons 1960a), which is op-p 2; 21b
in Saldula (Parsons 1963), which is p-tr 3; 61 in Saldula (Parsons 1963), which is
t-p 14; a dorsoventral muscle from the metanotum that controls the metathoracic scent
gland in Saldula (Parsons 1963), and M. glandulae thoracicae in Gelastocoris (Parsons
1960b). In Lethocerus (Parsons 1968) 6A is s 1, but is unique in its anterior insertion
on the hypopharyngeal wing.
In Plega (Fig. 118C) and Nemopterella (Acker 1958), in which the neck region is
somewhat lengthened, the lengthened lateral cervical sclerite is directed strongly forward
to maintain connection with the head. Even so, the sclerite still can be interpreted as
the preepisternum, since it maintains the same position with respect to the head and the
pleuron as in Sialis. In Pal pares figured by Czihak ( 1956), and Myrmeleon figured by
Sundermeier ( 1940), the lateral cervical sclerite, though somewhat enlarged, is also
similar to those in Plega and Nemopterella. In all these genera, the sclerites on the two
sides are clearly separated ventrally by a membrane between them.
In Cha.uliodes (Maki 1936), Corydalus (Kelsey 1954), and Aguila (Ferris and
Pennebaker 1939; Matsuda 1956a), all belonging to the suborder Megaloptera, the head
is of the prognathous type, and the neck region and prothorax are greatly lengthened.
In these genera, as will be discussed below, the preepisternum has no connection with the
head, and an entirely different lateral cervical sclerite connects the preepisternum and
the head.
In Aguila (Fig. 118B) the narrow posterior lateral plate, labeled as the pre-
episternum, is the same as the lateral cervical sclerite in Planipennia in its position relative
to the rest of the episternum; the median plate between the preepisterna is the basi-
sternum. Here the preepisternum faces the entire lateral margin of the basisternum, as
in the pterothorax of lower insects. The tripartite plate anterior to the preepisterno-
basisternum is the lateral cervical sclerite that has the connection with the head. In
Chauliodes (Maki 1936) and Corydalus (Crampton 1926a; Kelsey 1954) also the
posterior plate is the preepisternum plus the basisternum as in Aguila. In Corydalus
286
KATEPISTERNUM
& TROCHANTIN
LATERAL
CERVICAL
SCLERITE
BASISTER-
NUM
c
A
PRONOTUM
AN EPISTERNUM
SPINASTERNUM
ANAPLEURAL
CLEFT
AN EPISTERNUM
PLEURAL
INVAGINATION
POSTPLEURAL
SCLERITE
EPIMERON -------t:r l f l ~ " - '
PLEURAL SUTURE
CHANTIN
FURCAL
PIT
8
V ENTR 0 LATERAL
CERVICAL SCLERITE
(PRESTERNUM)
PREEPISTERNUM
BASISTERNUM
FURCA
ANEPISTERNUM
FURCA
FIG. 118. A, pro thorax and neck region in Sialis flavilatera (from Czihak); B, ventral view
of thorax in Aguila adnixa (from Matsuda); C, ventral view of prothorax and neck region in
Plega signata (from Ferris).
the anterior plate is ventrolaterally continuous as in Aguila, but has no direct connection
with the head; in Chauliodes the paired anterior plates are not continuous medially.
The anterior plate in Corydalus, Chauliodes, and Aguila is, then, clearly not
homologous with the lateral cervical sclerite in Planipennia and Sialis (Megaloptera).
The only source of derivation of this plate in Corydalus, Chauliodes, and Aguila is,
therefore, the presternum, which forms the lateral cervical sclerites in hemimetabolous
insects. In other words, in these genera belonging to Megaloptera the lateral cervical
sclerite is formed from the presternum as in hemimetabolous insects, and at the same
time, the preepisternum is detached from the rest of the episternum, as in many holo-
metabolous insects. Since in Aguila (Fig. 118B) and Corydalus the presternal lateral
cervical sclerites are ventrally continuous, the whole continuous plate may be called the
ventrolateral cervical sclerite.
The anterior plate was called the jugulare by Crampton ( 1926a) and Kelsey ( 1954).
Ferris and Pennebaker ( 1939) called the anterior plate (jugulare) and the posterior
plate (preepisternum plus basisternum) together the cervical sclerite.
In Chauliodes and Corydalus a small median unpaired plate and a smaller sclerite
on each side of the median plate are present behind the ventral posterior margin of the
head. The dorsal cervical sclerite is present in Corydalus (Kelsey 1954) and Nernop-
terella (Acker 1958).
The pronoturn: The pronotum is greatly prolonged in Aguila, Corydalus, and Plega
(Ferris 1940a). In Aguila the anterior part of the pronotum covers the prolonged
ventrolateral cervical sclerite; in Plega the prolonged posterior part of the pronotum
arches ventrad behind the pleuron on each side to form a dorsoventrally fused, complete
ring in front of the mesothorax.
The prothoracic pleurosternurn: The pleural suture in the prothorax of Neuroptera
is largely obliterated. This is due mainly to invagination of the posterior part of the
pleuron, which becomes fused with the furcal arm (Fig. 118A); this invaginated posterior
part perhaps contains the pleural arm. The pleural process for coxal articulation
is retained.
As already discussed, in all Neuroptera the preepisternum is completely isolated
anteriorly. The rest of the proepisternum consists of anepisternal and katepisternal
areas. In Sialis (Fig. 118A) the definitive katepisternum is the eutrochantin that has the
apically differentiated trochantin; in some genera (Aguila, Nernopterella, etc.) the
trochantin is absent. The paracoxal suture is absent in the propleuron of Neuroptera.
In Corydalus what Kelsey (1954) called the precoxal arm traverses the ventral surface of
the prothorax, immediately in front of the coxae; this area must be katepisternal in
ongm. In Corydalus and probably in other Megaloptera, the katepisternum has, con-
comitant with the shift forward of the basisternum, come to occupy the area where the
basisternum lies normally.
The basisternum in Planipennia including Ascalaphus, Myrrneleon, Palpares, Plega
(Fig. 118C), and Nernopterella has a median longitudinal ridge, and sclerotization does
not extend much beyond the level of the coxa. The furcae lie wide apart on the lateral
edges of the posterior sclerotized sternal plate. The spina lies far behind the furcal pits,
and is present in Myrrneleon, Ascalaphus, Chauliodes, Corydalus, and Aguila. In Aguila
(Fig. 118B) the spina is on the middle of the posterior margin of a narrow transverse
plate that Ferris and Pennebaker (1939) and Matsuda (1956a) called the mesothoracic
presternum. Taking the position of the spina as a landmark, this plate represents the
ventral limit of the prothorax, and the plate should be called the spinasternum.
68
A few postpleural.sclerites are present in the intersegmental area (s. lat.) between
the propleuron and the mesopleuron. In Corydalus, for instance, a small prespiracular
sclerite lies anterior to the first spiracle; below this sclerite lies a large intersegmental
pleurite bearing an apodemal invagination. Posterior to the intersegmental pleurite there
is a subspiracular plate. These last two sclerites serve as points of attachment for some
muscles, and they may be the remnants of the mesothoracic presternum.
66
1t is possible that the spina has shifted its position posteriorly to lie on the presternum (see p. 20).
288
The meso- and meta-notum: In many Neuroptera the mesonotum and metanotum
are similar in shape and size. The following discussion is limited, therefore, to the
mesonotum, unless stated otherwise.
In Corydalus (Fig. 1190) the antecosta bearing the first phragma internally is
present in the larval, pupal, and adult stages. In Chauliodes (Maki 1936) the antecostal
suture also demarcates the narrow acrotergite (precosta of Maki). In other studies of
Neuroptera this suture has not been referred to. In Plega (Ferris 1940a) the pronotum
is supported by the modified mesothoracic prescutum and the acrotergite.
The prescutum is clearly marked off from the scutum by the prescutoscutal suture
only during the larval stage in Corydalus (Kelsey 1957), as in many other immature
holometabolous insects (Crampton 1918b). In adult Neuroptera the prescutoscutal
suture is absent, although a short suture that demarcates the base of the prealar arm in
Corydalus (Fig. 1190) may represent the prescutoscutal suture.
The prealar arm (median prealar sclerite of Maki 1936, Tergalarm of Sundermeier
1940, laterophragma of Kelsey 1957) is the prescutal extension, which projects into the
intersegmental membrane; usually it does not reach the episternum, although in
Myrmeleon it does reach the anterior dorsal margin of the anepisternum. In Myrmeleon
Sundermeier ( 1940) pointed out that the arm passes anterior to the mesothoracic spiracle,
and it connects the pronotum and the mesonotum by a stiff ligament. In Chauliodes
(Maki 1936) and Corydalus (Fig. 1190), a small prealar sclerite lies posterolateral to
the prealar arm. The anterolateral scuta! suture is always well marked and demarcates
the suralare (anterior tergallever of Kelsey 1957).
The lateral parapsidal suture that arises from the base of the prealar arm posteriorly
becomes confluent with the counterpart from the other side, and forms a median anterior
lobe of the scutum in Plega (Fig. 119A; Ferris 1940a), Myrmeleon (Sundermeier 1940),
Ascalaphus and Palpares (Czihak 1956), and Nemopterella (Acker 1958). The suture
is absent in Aguila (Ferris and Pennebaker 1939; Matsuda 1956a), Sialis (Czihak 1953),
and in Corydalus (Kelsey 1957). Sundermeier ( 1940) and Czihak (1956) have called
the median area demarcated laterally by the lateral parapsidal sutures (their parapsidal
sutures) the prescutum. Crampton (1919) in Nemoptera also called the same area the
prescutum. The term prescutum as clarified in the present work (p. 12) is, however, a
narrow anterior strip bearing the prealar arm.
The anteriorly produced scutoscutellar suture (Fig. 119A, D) is always well marked.
In the pterothorax of Neuroptera, therefore, the lateral end of the well-defined scutellum
is continuous with the axillary cord. The posterolateral scuta! suture is also frequently
present; it arises from the lateral scuta! margin just behind the median notal process
(Fig. 1190) and reaches the base of the posterior notal wing process (Corydalus,
Chauliodes), or the scutoscutellar suture (Myrmeleon).
The postnotum is separated from the mesothoracic scutellum by a conjunctival
membrane; laterally, it meets the dorsal extension of the epimeron to form a postalar
bridge; in Corydalus it also reaches the posterior end of the subalare.
The second phragma is usually not conspicuous in Neuroptera. In Myrmeleon,
according to Sundermeier ( 1940), the second phragma is a bilobed lamella beneath the
ridge of the mesothoracic postnotum, and its posterior lobe (wall?) passes directly over
the metanotum. In Corydalus the medially ridged posterior margin of the postnotum is
continuous directly with the metanotum, and the single-walled second phragma arises
from the line of union between the mesopostnotum and the anterior margin of the
metanotum. It is difficult to determine, in this case, whether the phragma is meso-
postnatal or metanotal in origin.
In some Neuroptera, such as Nemopterella (Acker 1958) or Myrmeleon
(Sundermeier 1940), the metanotum is considerably reduced in size. Concomitant with
this reduction, the sutures present in the mesonotum are more or less obliterated in the
metanotum. Riek (1967) described, in many Neuroptera, a possibly stridulatory surface
on each lateral area of the metascutum. In Corydalus the third phragma arises from the
line of union between the metathoracic postnotum and the first abdominal tergite.
LATERAL
PARAPSIDAL
SUTU .
l'ST ABO.
. TERG.
SURALARE
A
TEGULA POSTMEDIAN NOTAL PROCESS
SICOSTALE
(HUMERAL PLATE) POSTEROLATERAL.
SCUTAL SUTURE
... .<
. . BASIRADIALE
c
POSTERIOR
NOTAL
PROCESS
B
MEDIAN NOTAL
PROCESS
RADIAL BRIDGE
PLEURAL RIDGE
TERGAL
FISSURE
SCUTELLUM
SCUTOSCUTELLAR
SUTURE
289
FIG. 119. A, dorsal view of pterothorax in Plega signata (from Ferris); B, lateral view of
forewing base in Rhaphidia sp. (from Sharplin); C, dorsal view of forewing base in Corydalus
cornutus (original); D, dorsal view of mesonotum in Corydalus cornutus (original).
The mesoscutal lateral margin and the fore wing base structures: In Corydalus
(Fig. 119D) the anterior and the antemedian notal processes on the suralare are not
distinctly differentiated. The suralare is followed by the inconspicuous tergal fissure,
which is followed by the small median and postmedian notal processes. The posterior
notal process is large i!l Neuroptera; it may be detached to form the fourth axillary
sclerite, as in the metathorax of Corydalus.
In Corydalus (Fig. 119C) the dorsal first axillary sclerite has association with four
anterior notal wing processes; anteriorly it is connected with the basisubcostale. The
dorsal second axillary sclerite lies along the distal margin of the first axillary sclerite;
anteriorly it is fused with the base of the R vein. The third axillary sclerite is distally
fused, along a suture, with the proximal median plate; anteriorly it is connected with the
posterior end of the second axillary sclerite; proximally it has the usual connection with
the tip of the posterior notal wing process, and a ligamentous connection with the caudal
margin of the tirst axillary sclerite.
The ventral second axillary sclerite in Raphidia (Fig. 119B), as Sharplin (1963a)
showed, has ligamentous connections with the pleural wing process and with the subalare.
The meso- and meta-thoracic pleurosternum: The following descriptions refer to
the mesothoracic pleurosternum, unless otherwise noted.
The pleural suture is clearly present; the pleural arm is formed internally near the
middle of the pleural ridge and projects caudad; the caudal projection of the pleural arm
is especially pronounced in Ascalaphus and Myrmeleon (Sundermeier 1940). The
epimeron is well retained.
The paracoxal suture is always present on the episternum, reaching the middle of
the pleural suture. It also becomes confluent with the anapleural cleft distally. In
Sialis (Fig. 120A), as Weber (1928a) and Matsuda (1960b) showed, the paracoxal
suture extends into the epimeron, dividing the latter into the broad, dorsal anepimeron
and the ventral, small katepimeron. The one occurring in the mesepimeron of Nemop-
terella (Acker 1958) may also represent the paracoxal suture. The paracoxal suture is
strongly ridged internally, and Ferris called it the pleural costa. According to Kelsey
(1957), the suture in Corydalus is formed from the adult tissue at the final stage of
development.
In the metathorax of Aguila and Plega (Fig. 120B) the transepimeral suture reaches
caudally the first abdominal spiracle. The same suture is represented by a horizontal,
membrane in the mesothorax of Plega. Ferris ( 1940a) thought this membranous area
to be a continuation from the anapleural cleft. He designated the area ventral to the
transepimenil cleft as the katepimeron. Clearly, however, the transepimeral suture in
Plega (Fig. 120B) and the epimeral paracoxal suture in Sialis (Fig. 120A) are not
homologous. Hence, the areas demarcated by these two different sutures are not
homologous; the katepimeron of Ferris in Plega is dorsally the ventral part of the
anepimeron in Sialis. The transepimeral suture occurs also in Ascalaphus (Czihak 1956).
The anapleural cleft or suture is always present and meets the paracoxal suture
distally. The ventral margin of the an episternum demarcated by this suture is sometimes
ridged internally. In Ascalaphus it is a straight ridge that reaches the anterior end of
the basalare.
In some Neuroptera the basalare is not externally detached. The internal apophysis
for the attachment of muscles, which Ferris and Pennebaker ( 1939) and Matsuda
(1956a) called the pleural apophysis in Aguila, occurs beneath the dorsal area of the
an episternum (where the basal are is detached in other insects). In Ascalaphus the
anepisternum is largely membranous and the ingrown basalar apophysis lies beneath the
sclerotized dorsal margin of the anepisternum; in Corydalus the basalare is demarcated
externally by a narrow membrane.
The preepisternum extends ventrally to meet the ventral median longitudinal ridge.
The pleurosternal suture has not been found in adult Neuroptera, the basisternum being
completely invaginated. The ventral process (Fig. 121) for coxal articulation, which is
the modified ventropleurite, is formed along the median longitudinal ridge at a level
anterior to the furcal pit.
The antecoxale along the paracoxal suture is not differentiated in Neuroptera. The
undivided trochantin is distally indistinguishably fused with the katepisternum.
The dorsal margin of the epimeron is emarginated, and the well-formed subalare is
always present in the membrane between the tergum and the epimeron. The postcoxal
bridge (Fig. 121) formed by extension of the ventral end of the epimeron, if present, is
narrow along the posterior margin of the coxa.
In Corydalus, as Kelsey's (1957) observation on the development of the ventral
thoracic area has shown, the basisternum is nearly completely invaginated in the adult;
the same is probably true of other Neuroptera. The furcal bases are closely approxi-
AN EPISTERNUM
ANAPLEURAL
SUTURE
TROCHANTIN
BASALARE
AN EPISTERNUM
PLEURAL WING
PROCESS

-KATEPIMERON
A
PREE?ISTERNUM-----T
- KATEPISTERNUM
B
291
FIG. 120. A, lateral view of mesothorax in Sialis sp. (original); B, lateral view of metathorax
in Plega signata (from Ferris).
mated, and appear externally as a single median pit. The mesothoracic spina is present
in Chauliodes and Corydalus.
The adult musculature: The adult thoracic musculature in Neuroptera was studied
by Miller ( 1933a) in Chrysopa plorabunda, by Maki ( 1936) in Chauliodes formosanus,
by Korn ( 1943) in Myrmeleon europaeus, by Czihak ( 1953) in Sialis flavilatera, by
Kelsey (1954, 1957) in Corydalus cornutus, by Czihak (1956) in Ascalaphus
macaronius, Myrmeleon europaeus and Palpares libelluloides, and by Matsuda ( 1956a)
in Agulla adnixa. In Table XVII the muscles in Chauliodes, Corydalus, Sialis and
Myrmeleon are homologized. The first two genera are long-necked Megaloptera and
PREEPISTERNUM
AN EPISTERNUM
VENTRAL

PR
FURCA
KATEPISTERNUM
ANT IN
FIG. 121. Ventral view of pterothorax in Plega signata (from Ferris).
Sialis is a short-necked representative of the same suborder. Myrmeleon belongs to
another suborder, Planipennia.
The abundance of muscles attached to the pleuron (t-p, p-s, p-cx, p muscles) in
Neuroptera is definitely a primitive feature for the Pterygota, although most of these
muscles have not been inherited from the wingless ancestor.
Noteworthy is the presence of t-s 10? in Corydalus, which connects the anterolateral
part of the pronotum and the posterolateral part of the prothoracic basisternum; this
muscle agrees in position with t-s 10 in Odonata which is, however, inserted on an
apodeme of the basisternum. Thus far, p-s 7 in the cervico-prothoracic region has been
found only in Corydalus among adult insects. Some relatively rare muscles, such as
p-cx 7(x), p-cx 8, p-cx 9, and s-ex 8, also occur in Neuroptera.
Gross aspects of the postembryonic muscle development: Kelsey (1954, 1957)
traced the development of thoracic muscles throughout the postembryonic stages of
Corydalus cornutus, and Korn ( 1943) described the larval and adult thoracic muscles in
Myrmeleon europaeus. These two studies are summarized below.
In Corydalus cornutus
(I) Larval muscles that become lost or change their positions during development.
( 1) Muscles that become lost only in a certain segment or segments, and their
serial homologues are present in the other segments.
86a (t-ti 2), 123 (p-s 9?), 133 (t-s 1 or 2), 135 (t-p 17), 178 (s 12?),
179 (p-s 9)."' 2 sps- 3 ils (p-s 7) of Chadwick (1959b)?
(2) Muscles that are lost in the adult of Corydalus, but are present in the adult
of other Neuroptera.
86b (t-ti 3), 96 (s-ex 4), 117 (t 11), 131 and 155 (t-ti 3), 167 (s-ex 1),
171 (s-ex 8(x)), 176 (s-ex 4?).
( 3) Muscles that are fused or may become fused with a neighboring muscle.
112 and 113 (t 14), 130 (t-ti 2) and 131 (t-ti 3), 138 (t-p 3?).
( 4) Muscles that become, by reorganization, adult muscles with different muscle
designations. 175 (p-tr 1 or 2) that becomes 165 (t-p 5?); 136 (t-p 5?)
that may become 128, 129 (t-p 5, 6).
"''n the larva 179 and 184 are p-s 9; only 179 becomes lost.
MATSUDA: THE INSECT THORAX 7.93
(5) Muscles that are lost in the adult and absent in other adult Neuroptera.
Muscles marked with an asterisk are palaeogenetic.
120 (p-s 8),* 134 (t-s 3), 139 (t-p 27),* 140 (t-p 28),* 147 and
149 (t-p 29), 152 (p-s 14), 155 (t-cx 2 or 3), 162 (p-cx 9), 177 (p-s 5),
2 sps - 3 ils (p-s 7) * of Chadwick ( 1959b).
( 6) Questionable muscles.
Larval muscles 146 and 148 are said to become lost, but they appear to
become 84 (t-p 1) in the adult.
Of the muscles under category 5, t-p 29 and p-s 14 are probably purely caenogenetic,
since such muscles have not been found in adult insects. p-cx 9 may be present in the
adult of Myrmeleon.
(II) Adult muscles that develop during the pupal stage.
86c (p-ti 1), 128 (t-p 6), 129 (t-p 5), 154 (p 3), 163 (t-p 13), 164 (t-p 14),
165 (t-tr 1) from 175 (p-tr 1), 166 (p-tr 2), 168 (s-ex 5), 169 (s-ex 6).
In Myrmeleon europaeus
Korn ( 1943) did not trace the development of individual muscles and this makes it
often difficult to discern the larval states of adult muscles. It is clear, however, that in
Myrmeleon the caenogenetic trend of muscles is more pronounced than in Corydalus.
For instance, in the neck region at least four muscles (0 ism 4, 6-8) out of eight 0 ism
Korn recognized cannot be homologized with neck muscles in adult insects.
None of the ten t-p muscles in the adult mesothorax can be traced back to muscles
in the larva. Presumably, most or all of the adult t-p muscles are formed during the
pupal stage.
Some larval muscles, which become lost during development, are present in the
adult of other Neuroptera. In the prothorax, for instance, I vim 2 (s 14), I vlm 2 (s 12),
I ism (p-s 9), and I ism 3 (p-s 6) are present in other adult Neuroptera.
Regarding the shift in points of attachment during postembryonic development,
Korn found that larval III pm 1 (p-cx 5) becomes adult III pm 2 ( p-cx 4), that larval
III pm 2 (p-cx 9?) shifts its insertion from the basicoxa to the trochanteral apodeme in
the adult to be p-tr 3, and that larval III pm 9 (t-cx 8) shifts its origin from the dorso-
lateral pleural (?) plate to the subalare.
THE MECOPTERA
The neck region: In Mecoptera including Merope (Figs. 122, 124A), Panorpa
(Fig. 123A), Bittacus and Boreus (Fig. 123B), the lateral cervical sclerite is a sub-
horizontally placed narrow plate that connects the anepisternum and the postoccipital
process. It is therefore probably of preepisternal origin, as in some Neuroptera (Sialis,
Myrmeleon, Plega, etc.). Miyake (1913, cf. Hasken 1939) and Mutuura (1961)
interpreted the lateral cervical sclerite as a detached part of the proepisternum. A small
dorsal cervical sclerite or sclerites are present in Panorpa, as shown by Hasken (1939)
and Ferris and Rees (1939).
The pronotum: An unique feature of the pronotum in Mecoptera is its connection
with the anterior margin of the mesopleuron by an extension of its posterolateral angle
(Figs. 122, 123A, B); in Boreus (Fig. 123B) the first thoracic spiracle lies on this
connection. The bridge so formed appears to replace the function of the mesothoracic
prealar arm, which is absent in most Mecoptera. In Panorpa (Fig. 123A), Merope
(Fig. 122), Bittacus, and probably in many others, the pTOnotum has a transverse internal
ridge along the posterior margin, and an additional internally ridged transverse suture
divides the rest of the pronotum into anterior and posterior parts. In Merope (Fig. 122)
the pronotum covers the dorsal neck membrane and the posterior part of the head,
although in Panorpa (Fig. 123A) the dorsal neck membrane is well exposed. In Merope
the pronotum also covers the propleuron laterally, and a median, unpaired longitudinal
suture and a lateral suture on each side meet the posterior transverse suture.
294
MEMOffiS OF THE ENTOMOLOGICAL SOCIETY OF CANADA
TABLE XVII
Thoracic musculature in Neuroptera
Corydalus Chauliodes
(Kelsey (Maki Myrmeleon Muscle
1954, '57) 1936) Sialis (Czihak 1953) (Korn 1943) designations Remarks
56, 57 53 0 dim 1 (pars)? cv(d)-t 1
83 59 M. pron.-occ. lat. op-t 1
57 60 M. pron.-occ. lat. 0 dlm 2 op-t 2
58, 59 M. meson.-occ. obl. 0 dlm 1 (pars) op-t 3
n 69 M. cerv. torq. 0 dvm 2 op-cv 1
70, 71 70, 71 M. cerv. torq. 0 ism 1 op-<:v 2 1
60 M. pleur.-occ. op-p 2 2
67 op-s 2
M. pron.-occ. ten. t-s(cv) 1
67 M. meson.-tent. 0 ism 2 t-s(cv) 9
66 72? 73? M. pron.-pl. long. I ism 1 t-cv 1
68? 72? 73? M. pron.-cerv. 0 dvm 1 t-cv 2
64, 65 64,65,66 t-cv 3
62 M. furco-cerv. 0 ism 3 cv-s 1
73 cv-s 3
87
84 M. cerv.-cox. cv-cx 3
62 p-s(cv) 7
61, 63 61 M. pl. furco-tent. 0 vim 1, 2 s 1, 2
M. meson. occ. lat. (I) t11
115(II) 104
115 (III) 154
76, 77 58 Mm. meson.-pron. t 12
116 106 Mm. dors. obl. II dlm 3
116 156 Mm. dors. obl. III dlm 3
69 54-57 Mm. meson.-pron .. t 13
114 105 Mm. scutello-postn. II dim 2
114 155 M. scutello-postn.
56,57,58,59 53 M. meson.-occ. obl. ? 0 dim 1 t 14
112, 113 102, 103 Mm. dors. recti. II dlm 1
112, 113 152, 153 Mm. dors. recti. III dim 1
84 (I) 78 M. pron.-postpl. t-p 1
85 (!) 79 t-p 2
137 (II) 114 M. noto-epist. brev. II pm 12 t-p 3
137 (III) 164 M. noto-epist. brev. III pm 12
142 (II) 115 M. pleuroc.-prenot. II pm 11 t-p 4 3
142 (III) 165 M. pleuroc.-prenot. III pm 11
129 (II) 113 M. nota-stern. II dvm 1 t-p 5
129 (III) 163 M. nota-stern. III dvm 1
79? t-p 6
128 112 M. nato-stern. II dvm 1
128 162 M. nota-stern. Ill dvm 1
141 (II) 116 M. noto-epist. long. II pm 9, 10 t-p 8 4
141 (III) 166 M. noto-epist. long. III pm 9, 10
153 (II) 117 M. noto-pleur. long. t-p 12
153 (III) 167 M. noto-pleur. long.
163 (II) 119 b M. epist.-alar. II pm 8? t-p13 5
163 (III) 169 b M. epist.-alar. Ill pm 8?
164 (II) 119a M. pleuroc.-alar. II pm 13, 14 t-p 14
164 (III) 169 a M. pleuroc.-alar. III pm 13, 14
TABLE XVII (Continued)
80-82 (I) 76, 77 Mm. pron. pl. I pm 2-5 t-p? 6
161 (II) 120 M. epim.-subal. II pm 6 t-p 16
161 (II I) 170 M. epim.-subal. III pm 6
135 (HI)
M. meson.-postpl. t-p 17
143 (II)
118 M. pleurae. brev. II pm 7 t-p 15
143 (III) 168 M. pleurae. brev. III pm 7
74?
68 t-s 1 7
150
111 M. furco-phragm. II pm 16
150
161 M. furco-phragm. III pm 16
75 (HI)
t-s 2 7
78 (I) 74, 75 t-s 7
79 (I I)
t-s 10 ? 8
80 M. pron. cox. t-ti(cx) 2
130
127 M. noto-troch. II dvm 2
130
167 M. noto-troch. III dvm 2
81 (I) Mm. pron. cox. t-ti(cx) 3
94
t--cx 5 9
158?
158?
92
82 M. pron.-cox. ext. I pm 1 ? t--cx 6 9
159a 128 Mm. not. mer. II pm 4?
159a
178 Mm. not.-mer. III pm 4?
159b (II) 129,130 Mm. not. mer. II dvm 4 t--cx 7 9
159b (III) 179,180 Mm. not. mer. III dvm 4
M. pron.-cox. int. (I)? t-cx 8
160 (II) 138 M. epim.-subal. II pm 5
160 (III) 186 M. epim.-subal. III pm 5
145 (II) 123 M. intraep. p 1
145 (III) 173 M. intraep.
144 (II) 122 p 2
144 (III) 172
154 (II) 124 M. epist.-st. II pm 1 p3
154 (III) 174 M. epist.-st. III pm 1
151 (II) 125 Mm. furc.-pleuroc. II zm p-s 1
151 (III) 175 Mm. furc.-pleuroc. III zm
M. epist.-sp. (I-II) p-s 2
lip m 17 p-s 3
119 (I II) 109 spin. seh. (I I I) p--s 6
121 (II-III) 159
184 (II-III) 108 (HI)
158 (II-III)
p-s 9
122 (HI) p-s 10
132 (I I) 135 M. epist.-troch. p-ti(cx) 3
132 (III) 185 M. epist.-troch.
89-91 87 M. pl.-cox. (1) I dvm 1 p--cx4
156 134 M. epist.-cox. long. II pm 2
156 184
M. epist.-ocx. long. III pm2
88
I dvm 2, 3 p--cx5
157 133 M. epist.-cox. brev. II pm 15
157 183 M. epist.-cox. brev. III pm 15
M. postpl.-cox. tr. (I) p-cx 7(x)
Ibm 3(HI) p-cx 8
93, 95(1) 83 (I)
I dvm 4 p--cx 9? 10
TABLE XVII (Continued)
M. epist.-troch. (II, III) p-cx?
11
122 (I-I 1)
s-11 or p-s 10
127 (I-II) M. furco-spin. s 12
180 (II-III) M. furco-spin.
124,!25,126 107 I vim s 13
(I-I I)
!8!,182,183 157 Mm. furco-furcal. II vim
(II-III)
118 (!) M. pl. furc.-sp.
M. pl. furc.-sp.
s 14
185 (III) M. pl. furc.-sp.
63 s 18 "12
110
160
spin.-cox. s-ex 1
170 (II) 126 II bm 4 s-ex 2
170 (III) 176 III bm 4
95? 86? Mm. furcopl.-cox. I ism 2 s-ex 3 10
173 132 M. stern.-mer. II bm 1
173 182 M. stern.-mer. III bm 1
spin.-cox. (I) s-ex 4
172 (II)
86c 85 M. stern.-cox. troch. s-ex 5
168 131 M. stern.-cox. II bm 2
168 181 M. stern.-cox. Ill bm 2
93 (I) ? Ibm 2 s-ex 6
169 (II)
169 (III) 10
Ibm 1 s-ex 8
165 (II) 137 M. abd. fern. not. II dvm 3 t-tr 1
165 (Ill) 187 M. abd. fern. not. III dvm 3
97 (I) p-tr 1
166 (II) 138 M. abd. fern. epist. II pm 3 p-tr 2
166 (III) 188 M. abd. fern. epist. III pm 3
88 Mm. abd. fem. furcopl. I dvm 5 s-tr 1
174 (II) 139 M. abd. fern. furc. II pm 3
174 (Ill) 189 M. abd. fem. furc. III bm 3
REMARKS (Table XVII)
(1) 71 in Corydalus may be op-cv 3.
(2) 60 in Corydalus is inserted dorsally on an apodeme in the dorsal cervical
membrane.
(3) The posterior point of attachment of 142 in Corydalus is on the pleural arm;
in Aguila (Matsuda 1956a) the corresponding muscle is a tegula-pleural wing process
muscle. The anterior attachment is on the prealar sclerite in Corydalus and Chauliodes.
( 4) pm 9 in Myrmeleon may represent t-p 7.
(5) pm 8 in Myrmeleon is inserted dorsally on the second axillary sclerite of
Korn but this muscle may actually be inserted on the third axillary sclerite, since Korn
( 1943) described his Pterale 2 as Analgelenkstuck.
(6) These muscles are prothoracic tergopleural muscles, which are not
homologized.
(7) The anterior point of attachment of 74 in Corydalus was described as the
pleural arm; 75 may be t-s 1.
MII.TSUDII.: THE INSECT THORII.X 297
(8) 79 in Corydalus connects the anterolateral portion of the pronotum and
posterolateral. portion of the basisternum.
(9) In Corydalus 158 may represent t-cx 6, and 159a and 159b together may
represent t-cx 7.
(10) 93 and 95 in Corydalus and 83 in Chauliodes arise dorsally from the
posterior surface of the pleural arm, and they may actually be sternocoxal muscles;
I dvm 4 in Myrmeleon arises from the posterior wall of the pleural ridge, and is attached
to the pleural process that articulates with the coxa.
( 11) Czihak (1953) did not figure his M. epist.-troch.
( 12) 110 and 160 in Chauliodes arise from the furcal base and each divides into
two branches, one extending forward and the other backward; these muscles form a
common ventral muscular net. 63 is also similar to 110 and 160.
The propleuron: In Merope and Panorpa the posterior part of the propleuron is
invaginated; this invaginated part, according to Hasken ( 1939), Ferris and Rees (1939)
and Mickoleit (1967), is fused with the furca (Fig. 124B), as in Neuroptera. However,
in Boreus (Fig. 123B) the propleuron is clearly divided by the pleural suture into the
anterior episternum and the posterior epimeron, which is exposed.
In Merope the episternum has a short extension ventrad (the precoxal bridge of
Mickoleit, 1967), with which the lateral cervical sclerite articulates. This ventral
extension may be the remnant of the katepisternum. The trochantin is clearly present
in Merope (Fig. 124) and Panorpa lugubris (Crampton 1926a). In Panorpa communis
(Fig. 123A) the episternum is a simple, undivided plate bearing the pleural process,
which articulates with the coxa at its ventral end. In Boreus the ventral margin of the
episternum is ridged.
The prosternum: The prosternum has no direct connection either with the
propleuron or with the coxa. In Merope the longitudinal invagination leads posteriorly
to a rhomboidal furcal pit, from which furcae arise (Fig. 124). In Panorpa and Bore us,
however, the furcal bases are rather widely separated. No clear division of the pro-
sternum by the sternacosta into the basisternum and the furcasternum occurs in
Mecoptera. The well-formed prothoracic spina has not been found. However, Fuller
BASALARE
TEGULA
AN EPISTERNUM 2
SPIRACLE
LATERAL
CERVICAL
SCLERITE
EPISTERNUM 1
PLEURAL SUTURE
PARACOXAL SUTURE
PRE EPISTERNUM 2 KATEPISTERNUM 2
SUBALARE 3
LATERAL
POSTNOTUM 3
ANEPIMERON 3
P ARACOXAL
KATEPIMERON 2
SUTURE
MERON 3
MERON 2
COXA 3
COXA 2
FIG. 122. Lateral view of thorax in Merope tuber (from Mickoleit).
298
BASALARE 2
LATERAL
CERVICAL
SCLERITE
COXA 1
MUSCLE DISC
LATERAL
CERVICAL
SCLERITE
EPISTERNUM 2
SUBALARE 2
BASALARE 3
l
ANAPLEURAL SUTURE
EPIMERON 3
KATEPISTERNUM
- PARACOXAL
SUTURE
A
PARACOXAL
SUTURE
POSTNOTUM 3
EPIMERON 3
EPISTERNUM 3
B
FIG. 123. A, lateral view of thorax in Panorpa communis (from Hasken); B, lateral view of
thorax in male Bore us westwoodi (from Fuller).
(1954) thought that a small process elevated posteriorly from the furcal base in Bore us
might represent the remnant of the spina, and Crampton ( 1926a) showed a vestigial
spina in Chorista and Panorpa lugubris.
The mesonotum: In Bittacus (Fig. 125) the anterior part of the mesonotum is
considerably prolonged. The anterior margin of the mesonotum invaginates beneath the
posterior margin of the pronotum, and the antecostal suture is hidden beneath the latter.
The first phragma from the antecosta extends forward. The anterolateral marginal area
is demarcated by a suture, which is the posteriorly extended prescutoscutal suture; such
extension is easily conceivable, because the anterior part of the mesonotum is greatly
ANEPISTESTERNUM
PLEUROSTERNAL
SUTURE?
ANAPLEURAL
SUTURE
MERON 3
COXA 3
PLEURAL
RIDGE
PLEUROCOXAL
ARTICULATION
A
LATERAL CERVICAL
SCLERITE
TROCHANTIN
STERNUM 1
FURCAL PIT
EPIMERON 2
_2..---- BASISTERNUM 2
B
PLEURAL SUTURE
BASISTERNUM 3
+PREEPISTERNUM 3
VENTRAL PROCESS
299
FIG. 124. A, ventral view of thorax in Merope tuber (from Mickoleit); B, prothoracic sterno-
pleural region and coxa in Me rope tuber (from Mickoleit).
PRONOTUM ~ SPIRACLE
PARAPSIDAL ~ - - ~
SUTURE - ~ - ----PRESCUTOSCUTAL
< ~ > . ~ v ~ > SUTURE
SCUTOSCUTELLAR
SUTURE
POSTERIOR
NOTAL
PROCESS
AXILLARY
CORD
TERGAL
FISSURE _____ _.,
SCUTELLUM 3 -----.Q
3'RD PHRAGMA
TRANSSCUTAL SUTURE
~ " : : : : : : : : : :
\---J,. _ _::.,J---:=;t:_:_------ POSTNOTUM 2
FIG. 125. Dorsal view of thorax: in Bittacus pilicornis (original).
prolonged. Crampton's (1919) figure of Bittacus sp. and Mickoleit's (1968) figure of
Bittacus pilicornis ( 1968) also show the presence of the prescutoscutal suture. An
oblique suture arising from the middle of the prescutoscutal suture (Fig. 125) is the
parapsidal suture. The prescutum demarcated by the prescutoscutal suture in Bittacus
is a simple marginal strip lacking the prea!ar arm. In Bittacus a posterior transscutal
suture arises from the scuta! lateral margin anterior to the posterior notal wing process.
In other genera the mesonotum is not prolonged and the sutures present in Bittacus
are either absent or obliterated. In Panorpa (Fig. 126) the anterior margin of the
mesonotum is clearly separated from the pronotum by a membrane, and the antecostal
suture is medially visible. The paired, small, first phragma is formed beneath the
antecostal suture. The prescutoscutal suture demarcates a narrow marginal strip on the
anterolateral angle, and the parapsidal suture is absent. A marginal ridge behind the
prescutoscutal suture represents the anterolateral scuta! suture, and the posterior lateral
ridge represents the posterolateral scuta! suture.
In Merope the anterior part of the mesonotum is similar to that of Panorpa, and a
small process on the anterolateral angle of the mesonotum, according to Mickoleit
PRESCUTO-
SCUTAL SUTURE
TERGAL FISSURE
SCUTUM 2
ANTEROLATERAL
SCUTAL SUTURE
SCUTOSCUTELLAR
SUTURE
\ I
PRESCUTOSCUTAL :t- 2
SUTURE _'I
------------- ___::-=;rp----__ POSTNOTUM
f
SCUTAL
4'TH AXILLARY \
SCLERITE I-
-- ;::o--
.-----
SCUTELLUM 3 -
POSTNOTUM 3
SCUTUM 3
PARAPSIDAL
SUTURE
SCUTOSCUTELLAR
SUTURE
FIG. 126. Dorsal view of thorax in Panorpa nuptialis (original).
(pers. comm.), corresponds to the prealar arm in Chorista, which Crampton ( 1925b)
discovered. In Bore us (Fig. 127), as Fiiller (1954) showed, the anterior part of the
mesonotum is hidden beneath the well-developed pronotum, and the first phragma is not
formed. The male mesonotum is T-shaped because of membranization; this narrowing
of the mesonotum is associated with the peculiar position of the fore wing, which has
been modified into an accessory copulatory organ (Steiner 1937). The posteriorly
divergent scutoscutellar suture demarcates the pentagonal scutellum.
In all genera the postnotum is well developed and is divided by a suture on each
side into the median postnotum and the paired lateral postnota. The lateral postnotum
meets the epimeron to form a postalar bridge. In Boreus the postnotum consists of
two plates: the anterior is attached to the scutellum, and the posterior is attached to the
anterior margin of the metanotum. In Mecoptera (other than Boreus) the second
phragma arises from the line of union of the mesothoracic postnotum and the anterior
margin of the metanotum, and is single walled as in Corydalus.
The mesoscutal lateral margin and the forewing base structures: The tergal fissure
in Merope, Bittacus (Fig. 128A), and Panorpa is rather inconspicuous. Hence, the
anterior, postmedian, and posterior notal processes are differentiated. The postmedian
notal process is identified by its association with the proximal posterior angle of the first
axillary sclerite. The tegula is present in a typical position, i.e. anterior to the forewing
base. The subtegula of Mickoleit ( 1967, 1968), which supports the tegula, is
homologous with the prealar sclerite in Neuroptera. In the dorsal surface of the
forewing base in Bittacus (Fig. 128A; Sharplin 1963a), the association of axillary
sclerites with wing vein bases are clear-cut and generalized, since they approximate a
.302
SCUTUM 2
SCUTUM 3
TERGAL
FISSURE
TERGAL FISSURE
l'ST ABO. TERG.
FIG. 127. Dorsal view of thorax in male Boreus westwoodi (from Fiiller).
primitive condition of the wing base in the Pterygota (Fig. 4). The proximal median
plate, which is theoretically the axillary sclerites of the M and Cu veins, has no confusing
association with other axillary sclerites; the first and second axillary clearly
detached basal parts of the Sc and the R, respectively.
On the ventral surface of the forewing base in Panorpa (Fig. 128B), the ventral
wall of the second axillary sclerite has ligamentous connections with the pleural wing
process and with thesubalare; the subalare has, in turn, connection with the pleural wing
process. Sharplin ( 1963a) pointed out that in Panorpa the ventral sclerotization of the
second axillary sclerite is, as in Neuroptera, connected by a simple cuticular bar to the
dorsal sclerotization of the second axillary sclerite, which is in turn fused to the R vein.
In Boreus the articulation of the forewing is vertical to the body, instead of being
parallel to it, and the axillary sclerites have undergone consequent changes in their
positions. Fiiller ( 1954) described the axillary sclerites without figures, and it is
therefore too difficult to summarize his study here.
The mesopleurosternum: In Mecoptera the internally ridged pleural suture divides
the mesopleuron into the anterior episternum and the posterior epimeron. Internally,
the pleural ridge bears the pleural arm near its middle; the pleural process at the ventral
end of the ridge is somewhat conspicuous. The paracoxal suture occupies a relatively
"more ventral (posterior) position than in most other orders. In Merope (Fig. 122) the
paracoxal suture also appears to be present on the epimeron (x of Mickoleit 1967),
although it is not directly continuous with the paracoxal suture in the episternum; in
Bittacus, as Matsuda (1960a) and Mickoleit (1968) have shown, the suture reaches a
point on the pleural suture more dorsal than in Merope, and the same is true of Chorista
(Crampton 1925b); in Panorpa the suture is incomplete, and does not reach the pleural
suture; in Boreus (Fig. 123B) the suture is absent.
-J The ana pleural suture in Chorista, as Crampton's ( 1925b) figure indicates, nearly
reaches the dorsal end of the paracoxal suture as in other orders; in other genera studied
the anapleural suture is more or less obliterated and its posterior end does not lie near the
lateral end of the paracoxal suture.
The trochantin is represented by a muscle disc that lies near the ventral end of the
paracoxal suture in Panorpa (Fig. 123A); in Bittacus the apical part of the trochantin
I'ST AXIL. SCL.
RADIAL PLATE
(BRIDGE)
-----3'RD AXIL. SCL.
POSTERIOR NOTAL PROCESS
2'ND AXIL. SCL.
303
FIG. 128. A, dorsal view of forewing base in Biltacus sp. (from Sharplin); B, inner lateral
view of right second axillary sclerite and associated structures in Panorpa communis (from
Sharplin).
is well retained (Mickoleit 1968), The basalare is incompletely separated from the
episternum in front of the pleural wing process. As Hasken (1939) and Fiiller (1954)
have shown in Panorpa and Boreus, the anterior margin of the episternum is reinforced
by the marginal ridge (Vorderapodem of Hasken).
The preepisternum in Merope is apparently incompletely separated from the sternum
by the presumed (or possible) pleurosternal suture (Fig. 124A}, which is comparable
with an oblique suture occurring in a corresponding position of some Trichoptera and
Diptera (Fig. 136A).
08
In other Mecoptera this suture is absent, and hence the pleuron
is indistinguishably fused with the basisternum.
The dorsal part of the epimeron in Boreus (Fig. 123B) is fused with the meta-
thoracic episternum, and the pleural intersegmental line is obliterated dorsally. The
subalare is present above the epimeron in Panorpa, Bittacus, and Merope.
The division of the sternum into the basisternum and the furcasternum is absent.
The basisternal area has a median longitudinal invagination that leads posteriorly to the
common furcal base. As Hasken (1939) showed in Panorpa, the reinforcing ridge on
the anterior margin of the episternum is continuous with the median sternal invagination.
The ventral process tha( articulates with the coxa is formed by the ventral extension from
beneath the median longitudinal invagination. The mesothoracic spina is absent in
Mecoptera.
The metanotum: In Bittacus (Fig. 125) the anterior median quadrangular space on
the metanotum is formed by convergence of the parapsidal sutures, which arise from the
ussee p. 37 for the discussion of this suture.
prescutoscutal suture. In Panorpa (Fig. 126) the prescutoscutal suture is obliterated
anteriorly, but is continuous posteriorly to the anterolateral scuta! suture, and the
parapsidal suture is absent; in M erope, judging from Mickoleit's (1967) figure, neither
the prescutoscutal nor the parapsidal suture is present; in Boreus (Fig. 127) also these
two sutures are absent.
In the posterior half of the metanotum the posterolateral scuta! suture is clearly
present in Bittacus (Fig. 125); it may be represented by a marginal ridge in Panorpa, and
it is absent in Merope. The scutoscutellar suture is always present, and in Boreus
(Fig. 127) it is strongly produced anteriorly. The posterior notal wing process is
represented by the detached, fourth axillary sclerite in all genera that have been studied.
The well-developed postnotum extends laterally to form a postalar bridge in all four
genera. As in the mesothorax, the postnotum is often divided by ridges into the median
plate and the lateral plates. The third phragma arises from the ridge formed on the line
of union of the postnotum with the anterior margin of the first abdominal tergite. In
Bittacus (Fig. 125) the first abdominal tergite is largely membranous, and the phragma
appears to be a continuation from the postnotum.
The metapleurosternum: The noteworthy differences between the metapleuron and
the mesopleuron are that the anapleural suture is clearly present in Merope (Fig. 122),
Panorpa (Fig. 123A), and Bittacus; and that the presumed pleurosternal suture is absent
in all genera studied. In Boreus (Fig. 123B) the first abdominal segment is fused with
the metathoracic epimeron.
The adult musculature: The thoracic musculature in Mecoptera was studied by
Maki ( 1938) in Neopanorpa ophthalmica, by Hasken (1939) in Panorpa communis, by
Fiiller ( 1955) in Boreus westwoodi, by Mickoleit ( 1968) in Bittacus pilicornis and
Harpobittacus nigriceps, and by Storch and Chadwick (1968) in Bittac_us strigosus.
The muscles described by the first three authors are homologized in Table XVIII.
Setty (1940), Chadwick (1959b), and Mickoleit (1967) have also described certain
muscles in Mecoptera.
Compared with Neuroptera, the number of muscles in Mecoptera is considerably
smaller. The presence of p-tr 4 is a characteristic feature, shared also by Diptera. Two
muscles from the pleuron (t-p 13, 14) are inserted on the third axillary sclerite, as
in Neuroptera.
THE DIPTERA
The neck region: The membranous neck region is always provided with a lateral
cervical sclerite on each side. In Tipula (Rees and Ferris 1939; Bitsch 1955; Fig. 130B)
the lateral cervical sclerite consists of three sclerites, which apparently have resulted from
membranization of the originally single plate. In another tipulid, Limnophila (Fig.
129A), the lateral cervical sclerite is a single plate that is narrowed anteriorly. In these
tipulids the posterior part of the sclerite is comparable, in its relative position, with the
detached preepisternum that forms the lateral cervical sclerite in Micropteryx (Fig. 144A)
and Trichoptera (Fig. 141A). In eriopterine tipulids (Crampton 1925a), Protoplasa
(Williams 1933) and others, the lateral cervical sclerite is an undivided, single plate,
which is also apparently of preepisternal origin.
In some forms, as will be shown later, the posterior point of articulation of the lateral
cervical sclerite may be with the ventrally shifted anterior margin of the pronotum.
The lateral cervical sclerite is still recognizable, even in such a highly specialized genus
as Basilia (Fig. 129B). In Chionea Bitsch (1955) showed paired openings of a
glandular canal along the inner margin of the lateral cervical sclerites (Fig. 137 A).
In Tabanus (Bonhag 1949), Musca (Crampton 1942), and Asilus (Sara and
Smerdel 1953) paired or unpaired sclerites occur anterior to the basisternum; Crampton
and Bonhag called them the presternum. However, these sclerites are probably the
TABLE XVIII
Thoracic musculature in Mecoptera
Bittacus and
H arpobittacus Panorpa Neopanorpa Boreus Muscle
(Mickoleit 1968) (Hasken 1939) (Maki 1938) (Fuller 1955) designations Remarks
6 0 dlm 1 0 d!m 1 cv(d) -t 1
8 op-t 1
7 0 dlm 2 op-t 2
5, 9 ? op-t 3 1
0 ism 1 6 0 ism 1 op-s 2
11 op-p 1
0 dvm 1 5 0 dvm op-cv 2
3 0 dvm 2 5 0 dvm op--cv 3
2 0 ism 2 0 ism 2 t-s (cv) 1 2
1 0 ism 3 7, 8 t-cv 2, 3 or 3
t-s (cv) 9
22 0 vlm 1, 2 2 Ovlm s 1, 2
4 0 ism 4 3 cv-s 1
0 ism 5 13 0 ism 3 cv--cx 1(x) 4
10 I dlm I dim t 12 or 13
28 II dlm 2 24 t 12
28 III dim 2 54
II dlm 3 t 13
III dlm
27 II d\m 1 23 t 14
27 Ill d\m 1 53
13 I pm 3 12 I pm 3 t-p? 5
12 I pm 4 10, 11 I pm 4 t-p? 5
44 II pm 11 30 t-p 3
44 III pm 11 60
46 II pm 8 31 t-p 4
46 III pm 8 61
29 II dvm 1 28 t-p 5 or 6
29 Ill dvm 1 58
45 II pm 15 29 t-p 8
45 III pm 15 59
40 II pm 5 t-p 10
40 III pm 5
II pm 6 32 t-p 12
III pm 6 62
41 II pm 9 33 t-p 13
41 III pm 9 63
42 34 II pm 3 ? t-p 14 6
42 64 Ill pm 3 ?
43 II pm 7 t-p 15
43
39 II pm 12 36 II pm 12 t-p 16
39 III pm 12 66
II pm 16 35 II pm 5 t-p 19
Ill pm 16 65
I ism 9 I ism t--s 1
30 II dvm 2 42 t-ti(cx) 2
30 III dvm 2 71
TABLE XVIII (Continued)
Bittacus and
Harpobittacus Panorpa Neopanorpa Boreus Muscle
(Mickoieit 1968) (Hasken 1939) (Maki 1938) (Fuller 1955) designations Remarks
15
I dvm 15, 16 I dvm t-cx 7
34
II dvm 4 44
34
Ill dvm 4 73
38
II pm 4 45 II pm 2 t-cx 8
38
Ill pm 4 74 Ill pm 2
3Sb
I I pm 13 37 p 1 ?
35b
Ill pm 13 67
35a
II pm 14 38 p2
35a
III pm 14 68
36
II pm 1 39 p3
36
III pm 1 68
so
II zm 40 p-s 1
50
III zm 69
53 (I-I I)
p-s 6
53 (II-III)
56 (II-III)
II pm 4
III pm 4
p-ti ? 7
14
I pm 1 19 I pm 1 p-ti 2 or 3
37
II pm 2 48 II pm 1
37
III pm 2 77 Ill pm 1
II pm 10 47 p-cx5
III pm 10 76
26 (HI) s 12
52 I vim 25 I vim s 13
II vim 55
23 I ifum 4 I ifum s 16
27
57
III vim Ill vim s 20
51 II bm 1 41 s-ex 2 8
51 III bm 1 70 III bm 4
21 Ibm 3 18 Ibm 3 s-ex 3
49 II bm 4
49 III bm 4
19 Ibm 1 ? 14 Ibm 1 s-ex 5 9
47 II bm 3 43 II bm 2
47 II I brn 3 72 III bm 2
20 17 s-ex 6
48 II bm 5 46? II brn 3
48 II brn 5 75 ? II bm 3
31 II dvrn 3 49 II dvm t-tr 1
31 Ill dvm 3 78 III dvm
17 (I) 21 p-tr 1
16 I pm 2 20 I pm 2 p-tr 2
32 II prn 3 50
32 III prn 3 79
56 II cxrn 1 II cxm 4 p-tr 4
56 III cxm 1 III cxm 4
18 Ibm 2 22 Ibm 2 s-tr 1
33 II bin 2 51 II bm 1
33 II bm 2 80 Ill bm 1
REMARKS (Table XVIII)
( 1) 9 ip. Bittacus is attached anteriorly on the neck membrane.
(2') These muscles may be regarded also as t-cv 2.
307
( 3) The anterior attachment of these muscles is on the middle of the lateral
cervical sclerite. 8 in Neopanorpa arises from the anterolateral corner of the meso-
notum, and 0 ism 3 in Panorpa arises from the lateral phragma, as t-s(cv) 9.
( 4) 13 in N eopanorpa is crucial, whereas 0 ism 5 in Panorpa and 0 ism 3 in
Boreus are not.
(5) These are prothoracic tergopleural muscles, which cannot be homologized.
(6) In Boreus pm 3 is inserted dorsally on the Hauptgelenkstuck, which, according
to Fiiller (1954), is a fusion product of the subalare and the second axillary sclerite.
(7) pro 4 in Boreus may be p-cx 5, which is present in other genera.
(8) These muscles are attached to the ventral pleural process.
(9) Ibm 1 in Panorpa is, according to Hasken (1939), inserted on the posterior
margin of the coxa. Mickoleit (1968) homologized it with 19 in Bittacidae, which
is s-ex 5.
In Bittacus strigosus Storch and Chadwick (1968) found an anterior tergopleural
muscle of the prothorax (14 of Storch and Chadwick), the mesothoracic s 13 (62 of
S. and C.) and t-p 19 (45 and 76 of S. and C.), which were not found in Bittacidae
by Mickoleit (1968).
secondary ventral cervical sclerite, since in Mecoptera and lower Nematocera the pre-
sternum is absent.
The pronotum: In Limnophila (Fig. 129A), as Crampton (1925b) showed, two
transnotal sutures on the pronotum exactly compare with the two transnotal sutures in
Panorpa (Fig. 123A). The posterior transnotal suture is continuous with the pleural
suture in Limnophila; in Tipula (Fig. 130B), erioptt!rine tipuloids (Crampton 1925a),
and many nontipuloid Nematocera (Crampton 1925b) also, the posterior transnotal
suture is continuous with the pleural suture, as in Limnophila.
In Limnophila (Fig. 129A) the pronotum lies clearly anterior to the mesonotum; in
many other Nematocera (Fig. 130B) the pronotum is pressed downward by the greatly
enlarged anterior part of the mesonotum. Evidently accompanied by the shift in position
of the pronotum, the posterolateral angle of the pronotum has lost the connection with
the mesothoracic episternum, which is present in Mecoptera.
In Tabanus (Fig. 130A) the anterolateral and posterior parts of the pronotum are
conspicuously swollen to form calli, yet the pronotal origin of these calli can readily be
recognized by comparing them with the pronotal subdivision in Tipulidae (Fig. 130B).
As in Tipula, the pleural suture is continuous with the posterior transnotal suture. In
Tabanus, intrusion cephalad of the mesonotum is more advanced than in Tipula. How-
ever, Bonhag ( 1949) interpreted the anterior (ventral) pronotum (callus) as pro-
episternal in origin, and he disagreed with Bromley ( 1926), who thought this callus was
the lower lobe of the anterior pronotum. Bonhag's argument was based on the fact that
internally the pleural apophysis extends dorsally and partially circumscribes the upper
limits of the callosity (anterior pronotum) .
Bonhag's pleural apophysis (arm) is, however, questionable. In Tipula reesi
(Fig. 130B) the pleural ridge (suture externally) is continuous with the ridge formed
internally beneath the posterior transnotal suture, and the pleural arm or apophysis is
absent. Therefore, what appears to be the pleural ridge in Tabanus is most probably
the fusion of the pleural ridge and the posterior transnotal ridge. The pleural apophysis
of Bonhag is, in any case, not serially homologous with the mesothoracic pleural
apophysis, which lies farther ventrad and is differently shaped. As will be discussed
(p. 328), the associated muscles also support these reinterpretations of the sutures
in Tabanus.
PRONOTUM
TRANS NOTAL

COXA 1
PRONOTUM +
PROPLEURON
METANOTUM +
METAPLEURON
COXA 3
PROSTERNUM
----------MESONOTUM
SPIRACLE 1
B
FIG. 129. A, lateral view of prothorax in Limnophila macrocela (from Crampton); B, dorsal
view of thorax in Basilia nana (from Nussbaum).
As a corollary of the above interpretation, the lateral cervical sclerite in Tabanus
articulates posteriorly with the pronotum, not with the propleuron! In Tipula (Fig.
130B) :the basal part of the lateral cervical sclerite lies on the proepisternum near the
displaced anterior margin of the pronotum. It is naturally expected that in some higher
Diptera with an even more ventrally displaced pronotum the lateral cervical sclerite
would have the connection with the pronotum.
The mode and degree of fusion of the pronotum, the propleuron, and the mesonotum
vary in different groups of Diptera. In Psorophora (Snodgrass 1959), Macrochile
(Crampton 1926b), and Asilus (Sara and Smerdel 1953) the anterolateral pronotum is
delimited from the proepisternum by a suture, and the callous posterior pronotum is
fused ventrally with the proepimeron; in Asilus, however, Sara and Smerdel thought the
anterolateral part of the pronotum was the anepisternum. In the typical thorax of
Culicidae Komp (1937) showed a suture that separates the posterior pronotum from the
proepimeron. In Drosophila (Fig. 131), as Zalokar (1947) and Ferris (1950) showed,
the anterolateral pronotum is not callous and is indistinguishably fused with the pro-
episternum; the close association of the swollen posterior pronotum (humerus) with the
mesonotum led Ferris to call the former erroneously the prescutum.
In a pupiparous genus, Basilia (Fig. 129B), the dorsally displaced prothorax is a
horseshoe-shaped sclerite. Its division into the tergum and the pleuron is completely
obliterated; its articulation with the coxal base and its position relative to the first spiracle
and to the lateral cervical sclerites, however, readily identify it as the prothorax. In
Trichobius, another pupiparous genus, the leg bases are not displaced dorsally, and the
prothoracic parts are fused together; according to Zeve and Howell ( 1963), the episternal
part has become fused with the mesonotum, and the pronotal area lies anteromedially
between the coxal bases in the ventral view. This interpretation is difficult to accept,
because, according to Zeve and Howell, the tergum lies ventral to the pleuron, This
discrepancy apparently arose from the fact that they accepted Bonhag's ( 1949) interpreta-
tion that the anterolateral part of the pronotum was the episternum. The pronotum of
Zeve and Howell is most probably the anterior end of the mesonotum.
In a nematocerous genus, Chionea (Fig. 132B), in which the forewings are vestigial,
the pronotum and the propleuron lie anterior to the mesothorax as in Mecoptera. This
reversal to a primitive condition has obviously resulted from reduction of the mesonotum,
which was accompanied by a great reduction of the forewings. However, in Basilia,
which represents one of the most specialized groups of Diptera, the loss of wings has
not reversed the unique evolutionary trend for the dipterous prothorax to retract caudad.
The propleuron: In lower Diptera the propleuron is separated by the pleural suture
into the anterior episternum and the posterior epimeron. There is no subdivision of the
episternum by a suture or sutures. The precoxal bridge in Johannsenomyia (Crampton
1925b), Sciara (Crampton 1942), Drosophila (Zalokar 1947; Ferris 1950; Fig. 131),
and Asilus (Sara and Smerdel 1953) is ventrally continuous with the anterolateral angle
of the basisternum. This precoxal bridge in Diptera is, as in some Lepidoptera (Fig.
149A, B), obviously a result of the secondary sclerotization of the membranous
katepisternal area in Mecoptera (Figs. 123A, 124A).
A well-formed trochantin has not been found in the prothorax of Diptera. The
epimeron, unlike that of most Mecoptera, is often externally exposed, and it forms a
postcoxal bridge in such forms as Tipula, Chionea, Tabanus, etc.
The prosternum: In Diptera (Fig. 136A, B) the prothoracic sternum is often
divided into the anterior basisternum and the definitive posterior furcasternum by a
suture, which connects the furcal bases. This apparent sternacostal suture merges
mesally into the median longitudinal suture of the basisternum. The definitive urea-
sternum or the posterior presternum is often triangular in shape with a pointed posterior
tip, and the furcal pits lie on its anterolateral angles.
In Asilus, as Sara and Smerdel ( 1953) showed, the vestigial spina lies between the
furcal bases. It is therefore probable that in most Diptera including Tipula, which lack
PRESCUTOSCUTAL
SUTURE
AN EPISTERNUM
POSTCOXAL
BRIDGE
ANAPLEURAL
SUTURE
BASALARE LATERAL
PARAPSIDAL
SUTURE
PLEURAL SUTURE 2 PREEPISTERNUM EPIMERON
TERGAL FISSURE
PREALAR LOBE (PREALAR TERAL
PRESCUTOSCUTAL
SUTURE
BASALARE
ARM) SCUTALSUTURE
SCUTUM ----,L----
PARAPSIDAL
SUTURE
POSTERIOR
PRONOTUM
PLEURAL
SUTURE
B
SUTURE
EPISTERNUM 3
EPIMERON
TRANSEPIMERAL
SUTURE
SCUTELLUM
POSTNOTUM
SPIRACLE
PLEURAL
SUTURE
AN EPISTERNUM
PREEPISTERNUM
FIG. 130. A, lateral view of thorax in Tabanus su/cijrons (from Bonhag); B, lateral view of
thorax in Tipu/a reesi (from Rees and Ferris).
MATSUDA: TilE INSECT TI!ORAX
PRONOTUM +
EPISTERNUM--.="';--
BASISTERNUM
FIG. 131. Ventral view of thorax in Drosophila sp. (from Ferris).
311
the spina, the definitive posterior presternum is a fusion product of the posterior pro-
sternum (presumably largely furcasternum) and the anterior part of the mesosternum.
In Tipula (Fig. 136A) the furca unites with the pleural arm and forms a continuous
brace. This is true also of eriopterine Tipulidae (Crampton 1925a), Drosophila
(Zalokar 1947), Tabanus (Bonhag 1949), and probably many others. In Chionea
(Fig. 137A) the coxal bases are widely separated and the sternal area is corres-
pondingly large.
The mesonotum: In Tipula (Fig. 130B) the narrow, anterior marginal strip is the
acrotergite; it is separated from the pronotum by a narrow membrane. The first
phragma is not formed from beneath the antecosta, and the strongly bulged anterior wall
of the mesoscutum serves as the anterior point of attachment for the powerful dorsal
longitudinal muscle (t 14). In Tabanus (Fig. 130A) the acrotergite of the mesonotum
(anterior pronotum of Bonhag) lies anterior to the pronotum and the first phragma, as
Bon hag ( 1949, his figs. 6, 7) showed, is continuous from the base of the acrotergite and
from the anterior margin of the posterior pronotum (humeral callus of Bonhag); in
Asilus also the first phragma arises from a corresponding position.
In Tipula reesi (Fig. 132A) four sutures arise from the lateral margin of the meso-
notum. The anteriorrnost suture is unmistakably homologous with the parapsidal suture
in Bittacus (Fig. 125), .. since it arises from the middle of the posteriorly extended
prescutoscutal suture. Judging from the studies of Crampton ( 1925a, b) and Williams
( 1933) the parapsidal suture associated with the prescutoscutal suture occurs rather
commonly in Nematocera. In Tipula (Fig. 133A) the anterolateral scuta! suture may
be present.
In some higher forms of Diptera the posterior pronotum (humerus) is incorporated
into the anterior lateral wall of the mesonotum, in conjection with the disappearance of
the parapsidal suture. In such cases the posterior limit of the humerus may still be
retained and simulate the parapsidal suture (e.g., Tabanus, Fig. 130A).
In Tipula the prescutum delimited by the prescutoscutal suture is long and its
posterior end is lobate. Rees and Ferris ( 1939) have called this lobate end the pre-
scutellar lobe. In another species of Tipula studied by Bitsch (1955) and in Tipula
manhatta examined in this study, this prescutellar lobe reaches the anterior margin of the
89
Crampton (1919) pointed out similarities in the tergal sutures of Bittacus and Tipula.
312
basalare. In its relative position, therefore, the lobe agrees exactly with the prealar arm
in other insects, although the prealar arm occurs only rarely in Mecoptera (Chorista).
The prescutellar lobe occurs commonly in Diptera. In Drosophila it has been called the
epine suturale (Zalokar 1947), and in Asilus Sara and Smerdel ( 1953) called it the
spina suturale.
The second intrascutal suture in Tipula (Fig. 132A) is the lateral parapsidal suture,
since it arises from the posterior end of the prescutoscutal suture (or the base of the
prescutellar lobe). The lateral parapsidal suture is highly persistent in Diptera, although
it has not been found in Mecoptera. In some higher forms it is internally strongly ridged.
The tergal fissure in Tipula reesi (Figs. 132A, 133A) lies behind the antemedian
notal wing process. In Asilus (Sara and Smerdel 1953) and Drosophila (Zalokar 1947;
Ferris 1950) this cleft extends more proximally and is a line of flexion. This fissure is
apparently absent or obscure in Tabanus (Bonhag 1949), Sphaeroceridae (Kim and
Cook 1966), and some lower Diptera. In Mecoptera the tergal fissure is very
weakly developed.
The posteriormost transscutal suture in Tipula reesi (Fig. 132A) is homologous
with the posterolateral scuta! suture present in the metathorax of Bittacus (Fig. 125); it
arises from the lateral scuta! margin behind the postmedian notal process. This suture
is present in many Diptera. '
The scutoscutellar suture in Tipula (Fig. 132A) is medially produced, as in
Mecoptera. In Tabanus (Fig. 130A) the scutoscutellar suture is lost. The posterior-
most transverse suture is probably homologous with the posterior margin of the scutellum
in Tipula (Fig. 130B). In Tabanus the original posterior margin of the scutellum has
become a transscutellar suture, because of evagination of the scutellum. This definitive
transscutellar suture occurs commonly in Diptera that have the evaginated scutellum.
As Crampton ( 1942) showed, the modifications ( evaginations) of the scutellum are
conspicuous in some Diptera such as Dicranophora, Celyphus, etc.; such scutellar
modifications in Diptera have occurred quite independently of essentially the same
modifications in Coleoptera and Hemiptera.
The large postnotum in Tipula (Figs. 130B, 132A) slants caudad a little and lies
posterior to the scutellum. As in Mecoptera, the postnotum is divided by a suture on
each side into the unpaired median postnotum and the lateral postnotum on each side.
The second phragma is continuous from the well-marked posterior margin of the post-
notum. The metanotal anterior margin is attached on the posterior margin of the
median postnotum.
In Tipula (Fig. 130B) the lateral postnotum and the lateral phragma are united,
along a suture, with the posterior margin of the epimeron, thereby forming a robust
postalar bridge. In many other nematocerous flies the suture that separates the phragma
from the postnotum has become lost. This lateral postnotum (or plus laterophragma)
is obviously homologous with the lateral postnotum in Mecoptera and in other insects .
that have the postalar bridge. Therefore, the appelation pleurotergite used by Crampton
( 1925a, b, 1942), Williams {1933), Bitsch ( 1955), and Byers ( 1961) for this area is
misleading. Rees and Ferris (1939) called this area the laterotergite, which is more
acceptable.
In many higher Diptera the postnotum is completely overgrown by the scutellum,
yet its division into the median and lateral parts is apparently maintained in most cases.
In Chionea the subdivision of the mesonotum is absent.
The mesoscutal lateral margin and axillary sclerites of the forewing"': In Tipula
(Fig. 133A) three notal wing processes that are clearly recognized externally include the
robust anterior notal wing process, the antemedian notal wing process lying immediately
anterior to the tergal fissure, and the posterior notal wing process separated by a narrow
membranous cleft from the scutallateral margin. Peculiarly the postmedian notal wing
70
Rees and Ferris (1939), Bonhag (1949), and Mickoleit (1962) have given clear illustrations of the mesonotal
lateral margin and the axillary sclerites. The following description is based partly on the observation of
Tipula spp, and Tabanus gracilis by the author.
MATSUDA: THB INSECT THORAX
PRESCUTO-
SCUTAL SUTURE
'
'
LATERAL
PARAPSIDAL
SUTURE
POSTEROLATERAL
SCUTAL SUTURE
POSTERIOR
NOTAL PROCESS
A
LATERAL
CERVICAL
SCLERITE
PROEPISTERNUM
SPIRACLE
8
EPISTERNUM Z
313
PRESCUTUM
PARAPSIDAL
SUTURE
SCUTUM
ANTERIOR
NOTAL
PROCESS
TERGAL FISSURE
POSTMEDIAN
NOTAL
PROCESS
SCUTELLUM
HALTER
POSTNOTUM
(PHRAGMA)
MESONOTUM
HALTER
METANOTUM
FOREWING
EPIMERON Z
SPIRACLE
PLEURAL
SUTURE
FIG. 132. A, dorsal view of pterothorax in Tipula reesi (from Rees and Ferris); B, lateral
view of thorax in Chionea lutescens (from Bitsch).
ANTERIOR
NOTAL PROCESS
l'ST. AXIL. SCL.-
MEDIAN
2'ND AXIL. SCL.
BASI COST ALE
(HUMERAL
PLATE)
TEGULA
PLEURAL
WING
PROCESS ----.r
BASALARE -----'
A
B
ANTEROLATERAL
SCUTAL SUTURE?
ANTEMEDIAN
NOTAL PROCESS
3'RD AXIL. SCL.
SUBALARE
EPIMERON
FIG. 133. A, dorsal view of forewing base in Tipula reesi (from Rees and Ferris); B, lateral
view of forewing base in Tipula vernalis (from Mickoleit).
process lies folded beneath the inner surface of the scuta! lateral margin. In Tabanus
(Fig. 130A) the tergal fissure is obliterated; the fourth axillary sclerite is formed by the
distal detachment of the posterior notal wing process. What Bonhag ( 1949) called the
posterior notal wing process (in his fig. 20) is actually the postmedian notal process. In
the same way, in Musca and Sarcophaga what Mihalyi (1936) and Boettiger and
Furshpan ( 1952) called the posterior notal process is actually the postmedian notal
process. In these genera the proximocaudal end of the first axillary sclerite articulates
with this process, as in Tipula; and this fact identifies the process as the postmedian notal
process. The same interpretation applies also to Asilus and probably to many other
Diptera.
The first axillary sclerite in Tipula (Fig. 133A) articulates with the anterior notal
wing process and, as already noted, with the postmedian notal wing process lying beneath
the scuta! margin. The head of the first axillary sclerite articulates distally with the base
of the Sc vein (regarded by Ferris and Rees as the humeral plate).
In Tipula (Fig. 133A) and Tabanus (Fig. 134) the second axillary sclerite is a
subtriangular plate facing the distal margin of the first axillary sclerite. The base of the
R vein lies above the dorsal margin of the second axillary sclerite, and the median plate
lies separated distally from the second axillary sclerite.
In Tipula and Tabanus the third axillary sclerite overlies the second axillary sclerite
and the median plate; its proximal process is connected with the posterior notal wing
process, and its distal end is associated with the thick base of the A vein.
Ventrally, in Tipula (Fig. 133B), the second axillary sclerite is connected with the
posterior part of the pleural wing process. The base of the R vein fits into an articulatory
sclerite, which apparently constitutes the anterior part of the pleural wing process. The
ventral third axillary sclerite lies closely above the subalare.
The mesopleuron: In Tipula (Fig. 130B) the pleural suture is oblique and nearly
straight; the pleural arm lies approximately at the middle of the pleural ridge, as in
Panorpa. In higher Diptera there is a strong tendency for the pleural suture to make a
sharp horizontal turn cephalad at the middle, as seen in Tabanus (Fig. 130A), Droso-
phila, Musca, etc. At least in Tabanus and Drosophila the pleural arm arises from the
middle of the pleural suture. The ventral end of the pleural ridge forms a conspicuous
ventral pleural arm for the attachment of muscles. In Basilia (Fig. 137C) the meso-
notopleuron is displaced dorsally, and the ventral end of the pleural process articulates
with the coxa, which is fragmented into a few sclerites.
The anapleural suture is persistent in Diptera. The suture arises from near the
middle of the anterior margin of the episternum and runs subhorizontally towards the
pleural suture in Tipula (Fig. 130B) and other Nematocera (Crampton 1925a, b); in
some Nematocera this suture is apparently absent or partially lost, especially anteriorly.
In Culicidae (Komp 1937; Crampton 1942; Snodgrass 1959) the posterior half of the
anapleural suture has probably been lost, and it has become continuous with the basalar
cleft; in Tabanus (Fig. 130A) the anapleural suture is internally ridged; in Drosophila
(Ferris 1950) and Calliphora (Lowne 1892) it is anteriofly lost, and in Musca (Hewitt
1914) it is absent.
In Tipula (Fig. 130B) the two-part basalare is separated from the anepisternum by
a transverse suture, and from the pleural wing process by a membranous cleft. In many
nontipulid Nematocera the dorsal area of the anepisternum has a large, membranous cleft
surrounding the basalare. In Tabanus (Fig. 130A) the basalaria are surrounded by
relatively narrow membranous clefts.
The preepisternum is the area ventral to the anapleural suture; in most flies the
preepisternum is, as in other Holometabela, indistinguishably fused with the basisternum.'
In Gnophomyia (Fig. 136B), as Crampton (1925b), Hendel (1928), Hennig (1941),
and Matsuda ( 1960a) showed, the pleurosternal suture is evidently present, and it
separates the basisternum from the preepisternum as in lower insects (e.g., Perla,
Fig. 48). Hennig ( 1941) maintained that the suture in Gnophomyia was merely a
weak internal ridge, and hence was not homologous with the pleurosternal suture. As is
often the case with other sutures, such as the paracoxal sutures, the anapleural suture,
etc., however, a suture can be ridged internally in varying degree. Therefore, there is no
positive reason why the suture cannot be the pleurosternal suture as well as a
strengthening ridge.
In Tipula reesi (Fi'g. 136A) Rees and Ferris (1939) showed an incomplete, oblique
suture on each side of the median longitudinal suture; although they did not refer to it.
The presence of an exactly comparable suture in Tipula manhatta was confirmed in the
present study. This suture in Tipula spp. is apparently homologous with the presumed
pleurosternal suture in Merope (Fig. 122). Whether this suture truly represents the
pleurosternal suture awaits further studies (see p. 37).
2'ND AXILLARY
SCLERITE
3'RD AXILLARY
SCLERITE
4'TH AXILLARY SCLERITE
(POSTERIOR NOTAL PROCESS)
R HUMERAL PLATE
TEGULA
1'ST AXILLARY
SCLERITE
~ ~ , _ _ __ t-p 13
.___ ___ t-p 14
:::::;;:::== t -p 15
POSTMEDIAN
NOTAL PROCESS
FIG. 134. Dorsal view of forewing base in Tabanus sulcifrons (from Bonhag).
In most Diptera the leg bases lie closer together along the median longitudinal axis
than they do in Tipula, and the presumed pleurosternal suture is absent; in these forms
it can be assumed that the narrow basisternal element is present indistinguishably along
the median longitudinal suture.
The true paracoxal suture on the episternum was shown by Crampton (1925b)
71
to
be present in Simulium, Eucorethra, Dixa, Plecia, Ceratopogon, Psorophora, and
Johannsenornyia. This suture demarcates the pleurotrochantin of Crampton (katepi-
sternum) on the episternum. In Eucorethra underwoodi (Fig. 135), which was
examined in this study, the paracoxal suture extends onto the epimeron. The paracoxal
suture in Tipulidae, claimed by Rees and Ferris (1939) and Matsuda (1960a), lies too
ventral to be the true paracoxal suture.
Because ot'the absence of the paracoxal suture, the katepisternum is not well defined
in most Diptera. Only in some nontipulid Nematocera mentioned above the dorsal
margin of the katepisternum is clearly recognized, and the trochantin is apically dif-
ferentiated from the katepisternum proper; the two areas are completely fused, and
together they form the eutrochantin. In Tipula (Fig. 136A) and Gnophomyia (Fig.
136B) the area surrounding the coxa is membranous and without a trochantin. The
membranous area is sclerotized in higher groups of Diptera, due partly to approximation
of leg bases. The ventral process, which articulates with the coxa, is visible externally
in Gnophomyia, in which the sternum is well exposed (Fig. 136B).
In Tipula (Fig. 130B) the epimeron lies above the moderately developed meron,
and is divided by an oblique ventral suture into large dorsal and small ventral areas.
This suture is probably the transepimeral suture, since it reaches posteriorly near the
metathoracic spiracle; its anterior end near the ventral end of the pleural suture is, how-
ever, rather atypical for the transepimeral suture. A conspicuous evolutionary tendency
of the .posterior part of the mesothoracic pleuron in Diptera, which is clear from studies
by Young (1921) and Crampton (1925a, b), is the strong development of the meron in
higher Diptera. The meron, which is displaced dorsally in varying degree, becomes
indistinguishably fused with the ventral part of the epimeron to form a larger, definitive
meron or the meropleurite of Crampton ( 1925b).
71
Crampton did not name this suture.
KATEPISTERNUM
'
"
I '
I '
\
(J
//$1(;;;
r__,p.,..:"J.
PARACOXAL
SUTURE
TROCHANTIN
FIG. 135. Mesopleuron and leg base in Eucorethra underwoodi (original).
317
Among the Nematocera Crampton ( 1925a) studied, the line of demarcation
(anepimeral suture of Crampton) between the definitive meron (meropleurite of
Crampton) and the epimeron in Protoplasa, Bruchomyia, and Chrysopi/us appears to
agree with the transepimeral suture in Tipula (Fig. 130B), since the line in these forms
reaches near the metathoracic spiracle. In these forms, therefore, it is probable that
the lower portion of the epimeron and the meron have become fused to form a
definitive meron.
In some Nematocera, such as Dolichopeza (Byers 1961) and Mycetobia (Crampton
1925b), a transepimeral suture is continuous from the anapleural suture on the
episternum and reaches the posterior margin of the epimeron at the level dorsal to the
postalar bridge. Therefore, this suture may not be homologous with the transepimeral
suture, which lies more ventrad in most Diptera."
Among nonnematocerous Diptera the dorsal margin of the definitive meron appears
to be represented by the transepimeral suture in Tabanus (Fig 130A); in Asilus (Sara
and Smerdel 1953) the meron does not reach the transepimeral suture, as in Tipula; in
Drosophila (Zalokar 1947; Ferris 1950) and Musca (Crampton 1942), the dorsal limit
of the meron lies more dorsad than the transepimeral suture in most flies. In most
Diptera the well-developed subalare lies above the dorsal margin of the epimeron.
The mesosternum: In Tipula (Fig. 136A), as already mentioned, the presumed
pleurosternal suture demarcates the basisternum, which has a median longitudinal suture.
Internally the furcal base lies on the posterior end of the median longitudinal ridge
(suture externally), and clearly anterior to the coxal base; this anterior position of the
furcal base has evidently resulted from the shift forward of the anterior mesosterna!
element, a characteristic feature of the dipterous thorax. The furcal arm is firmly united
with the pleural arm. The narrow median sclerite posterior to the furcal base is the
furcasternum; the ventral process extends laterally from the posterior part of the urea-
sternum and articulates with the inner coxal margin.
In Gnophomyia, in which the sternum is well exposed (Fig. 136B), the area
posterior to the furcal bases is divided into two areas. Crampton ( 1925b) and Hennig
( 1941) called them the anterior furcasternum and the posterior furcasternum, res-
pectively. However, the supposed posterior furcasternum probably includes the
metasternal element, judging from the tendency for the anterior thoracic sterna to
shift forward.
In Tabanus, Drosophila, and many other higher Diptera, the pleurosternal suture
has been lost, the coxal bases lie closer to the median longitudinal axis of the body, and
7
21'his suture can be homologous with the transepimeral suture, if its dorsal position is proven in phylogenetic
studies to be due to a continuous shift in position.
PROSTERNUM LATERAL CERVICAL
(BASISTERNUM) SCLERITE
PREEPISTERNUM
BASISTERNUM 2
PLEUROSTERNAL
SUTURE ?
POSTCOXALE 2
PRECOXALE 3
VENTRAL
PROCESS
BASISTERNUM 1
BASISTERNUM 2
FURCASTERNUM 2--
MESOSTERNUM+
META'STERNUM
PLEUROSTERNAL
SUTURE
PRO EPISTERNUM
CAL PIT
FURCASTERNUM 1
+ MESOSTERNUM
FURCA 2
FURCASTERNUM 2
PROCESS
METASTERNUM
FURCA 3
A
PROSTERNUM +
MESOSTERNUM
8
FIG. 136. A, ventral view of thorax in Tipula reesi (from Rees and Ferris); B, the same in
Gnophomyia sp. (from Hennig).
MATSUDA: TilE INSECT 'ITIORAX 319
the area corresponding to the furcasternum in Gnophomyia has invaginated into the
thoracic cavity.
In' Chionea (Fig. 137A), which has widely separated coxal bases, the mesosternum
is again exposed and considerably shortened; and the widely exposed pleurosternum
meets its counterpart along the median longitudinal axis of the body. It is noteworthy
that the common furcal base and the median longitudinal ridge have remained in their
original positions on the median longitudinal axis. In other words, the separation of the
leg bases has not influenced the position of these structures in Chionea, although the
position of the furcal base with respect to the leg base has shifted caudad.
According to Hennig (1941 ), the leg bases were primitively close to the median
longitudinal axis, as seen in Musca. In Hippobosca (Fig. 137B) the metasternal furcae
on the median longitudinal axis have shifted their positions forward and have united with
the mesosterna! furcae, while the median and posterior leg bases became widely separated.
In a parasitic genus, Basilia (Fig. 137C), as Nussbaum (1960) showed, the leg
bases have shifted onto the dorsal lateral margin of the thorax. The completely fused
thoracic sterna consists of the anterolateral part on each side separated by an oblique
membranous cleft and the posterior median sternum. Nussbaum (1960), on the basis of
the musculature, concluded that the paired processes arising from near the membranous
cleft are the profurcae. Based on the pattern of innervation, further, she attempted to
divide the fused sternal plate into its original components.
In both Basilia and Hippobosca a median longitudinal ridge extends through the
entire length of the thoracic sternum. Peculiarly, in Basilia, however, the only common
base of the meso- and meta-furcae lies on the posterior end of the median longitudinal
ridge. It is thus possible that, unlike in Hippobosca, the mesothoracic furcae have
migrated cauded and fused with the metafurcae, or else the mesothoracic furcae simply
have been lost. What Nussbaum ( 1960) called the vordere Leiste in Basilia, which
articulates with the coxa, may be the remnant of the pleural process.
In all of these forms without wings or with greatly reduced wings, the sternum is
widely exposed and they are forced to walk or to remain stationary in restricted spaces.
Under such circumstances, widely separated leg bases must- certainly be more advan-
tageous than medially approximated leg bases, which their winged ancestors must have
had. The same adaptation has occurred at least in Thysanoptera (p. 247) and in some
parasitic Hymenoptera (p. 368). In the cave-dwelling genus Trichobius, the leg bases
are also widely separated, although they have well-formed forewings (Zeve and
Howell 1963).
The metanotum and the halter base: In Tipula vernalis (Mickoleit 1962; Fig. 138B)
the scuta!, scutellar, and lateral postnotal areas reaching the epimeron are recognized.
As Young ( 1921) showed, the metanotum is also well retained in Psychoda. In higher
Diptera the metanotum, lying beneath the mesoscutellum, is greatly reduced; its lateral
part, which represents the lateral postnotum or the laterophragma, often reaches the
metepimeron (e.g., Drosophila, Tabanus), as in Tipula.
In Tipula (Fig. 138A), as Mickoleit (1962) showed, the first, second, and third
axillary sclerites in the halter base assume typical positions relative to one another; the
first axillary sclerite articulates with the anterior notal wing process, and its posterior end
reaches what may represent the postmedian notal wing process. In the lateral view
(Fig. 138B), the pleural wing process fits into a small cavity formed at the base of the
humeral plate.
The metapleuron: In the metathorax of most Diptera the distinct pleural suture
separates the anterior episternum from the epimeron. In Tipula vernalis (Fig. 138B)
the dorsal part of the episternum is isolated by a transverse, membranous cleft, and its
ventral anterior border is distinct from the mesoepimeron and the mesothoracic meron.
In the species of Tipula studied by Bitsch (1955), the episternal area is narrow and the
trochantin is present. Young (1921) recorded the presence of the katepisternum
320
LATERAL CERVICAL
SCLERITE
BASISTERNUM 1
FURCAL PIT
BASISTERNUM 3
STERNAL RIDGE
MESOSTERN
VENTRAL PROCESS ?
METASTERNUM
COXA 1
PLEURAL
PROCESS?
STERNAL RIDGE
FURCAE 2,3
.. . ..
:---------FURCA 1
FURCA 2
FURCA 3
B
PROFURCA
COXA 2
MESOSTERNUM
+ METASTERNUM
COXA 3
FIG. 137. A, ventral view of thorax in Chionea lutescens (from Bitsch); B, dorsal inner
view of thorax in Hippobosca sp. (from Hennig); C, dorsal inner view of thorax in Basilia nana
(from Nussbaum).
ANTERIOR
NOTAL PROCESS
POSTMEDIAN
NOTAL PROCESS
HALTER
SCUTUM
SPIRACLE-
EPISTERNUM 3
MERON 2
321
-----TEGULA
1 'ST AXIL. SCL.
3'RO AXIL.SCL.
POSTNOTUM
SCUTELLUM
POSTNOTUM3
'"'""--" I --------l'ST ABO. TERG.
B
FIG. 138. Halter base in Tipula vernalis (from Mickoleit). A, dorsal view; B, lateral view.
(pleurotrochantin of Young) in Rivellia, Leria, etc., which, however, needs to be
confirmed.
The metasternum: In Tipula (Fig. 136A) the metasternum is greatly reduced.
The internally ridged median longitudinal suture passes through the entire ventral surface
of the metathorax. Rees and Ferris (1939), Bitsch (1955), and Mickoleit (1962) have
shown that the sternal furcae arise at the level of the ventral process and articulate with
the coxa. Taking the location of the furcae as a landmark, the basisternum roughly
occupies the anterior half of the metasternum in Tipula. In Tabanus and Drosophila
the metasternum is even more reduced, but the furcae are still present. In Chionea
(Fig. 137 A) the metasternum is well retained, and the ventral process lies definitely
posterior to the level of the furcal base.
The anatomical basis of flight: Boettiger and Furshpan's ( 1952) and Pringle's
(1957a) explanations of the mechanics of flight in Sarcophaga bullata are given first,
using terminologies consistent with those used in the preceding discussion.
Most of the sutures present in other Diptera are also found in Sarcophaga (Fig.
139A). Noteworthy anatomical facts concerned with flight are: the anterior end of the
postmedian notal process ( scutellar lever of B. and F.) that articulates with the first
axillary sclerite; formation of a parascutal shelf by extension caudad of the sural are; the
hinge point i where the postnotum articulates with the base of the postmedian notal
process. Note also that the prescutoscutal suture and the lateral parapsidal suture meet
at point j, as in other Diptera.
Pringle ( 1957a) summarized the click mechanism of the wing in the same species as
follows. If the wing is in the up position (Fig. 139B), the lateral force produced by
the dorsal longitudinal muscle, t 14 (through the parascutal shelf and the first and second
axillary sclerite) forces apart f on each side, but here it is resisted if the pleurosternal
muscle (p-s 1) is also contracted. At the same time, however, if the longitudinal force
through the postmedian notal process is sufficiently strong to raise the first axillary
sclerite at the point z and carry y (anterior notal process) nearly to the line joining x
and f, the elastic energy stored in the tergum is released by the rapid rotation of the
second axillary sclerite about the pleural wing process, and the wing clicks into its
down position.
On the upstroke (Fig. 139C), the first effect of contraction of the dorsoventral
muscles ( t-p 5, 6) is to force apart f and, again, little movement takes place, since y is
above the line xf. When the downward force transmitted through the postmedian notal
process again reaches the critical point, the wing clicks back into the up position; this
mechanism only works if the lateral movement of point f is resisted by contraction
of p-s 1.
Boettiger and Furshpan (1952) and Pringle (1957a) did not explain the actions
of some other muscles, including the direct flight muscles inserted on the axillary
sclerites, which are especially abundant in Diptera (see Table XIX). Mainly on the
basis of muscle attachments, Bonhag ( 1949) explained the actions of the direct flight
muscles in Tabanus, as follows. Direct muscles of the first and third axillary sclerites
and those of the postmedian notal process (Fig. 134) are the effectors of the flexion
movement, since, by their concerted contraction, the wing is drawn onto the back of the
fly when at rest; the basalar-pleural muscle (p 2), by pulling the basalare ventrad and
inward, accounts for the ventral deflection and forward thrust of the anterior margin of
the wing during the down-stroke; deflexion of the posterior margin of the wing during the
upstroke is accomplished by tergopleural muscles ( t-p 15) inserted on the postmedian
notal wing process; an anterior tergopleural muscle ( t-p 3) causes extension of the wing
from the resting position over the back, and the function of a posterior tergopleural
muscle ( t-p 9) appears to be antagonistic to t-p 3; a tergopleural muscle ( t-p 4)
apparently stabilizes the pleural wing process during flight. Williams and Williams
(1943), Boettiger and Furshpan (1952), and Smart (1958) suggested that the primary
function of the tergotrochanteral muscle ( t-tr 1) is jumping, and hence it is a starter
in flight.
Schneider (1953) and Danzer ( 1956) found in Calliphora that the mesothoracic
muscles propel the halter during flight. Mickoleit (1962) thought that in Tipula,
Tabanus, and Musca the function of the four muscles of the halter (t-p 5 or 6, p 2, t-cx 8,
t-p 8) are concerned with the movement of the halter. Of these, the tergosternal
(pleural) muscle (t-p 5 or 6) is the elevator, and the pleurosubalar muscle (t-cx 8) is the
depressor of the halter in Tipula. The elevator and the depressor of the halter in
Calliphora (Pringle 1948; Schneider 1953) and in Tipula are homologous.
The adult musculature: Besides earlier works by Kiinckel (1875, 1881) on Eristalis
tenax and by Lowne (1890-1895) on Calliphora erythrocephala, more recent works on
the thoracic musculature in Diptera include those in Calliphora vomitaria (Ritter 1911),
Ctenacroscelis mikado, Ptecticus latifascia, Lathyrophthalmus obscuritarsis, Calobata
sinensis and Orthellia claripennis (Maki 1938), Drosophila melanogaster (Zalokar 1947)
Tabanus sulci frons and Tabanus americanus (Bonhag 1949), Drosophila melanogaster
(Miller 1950), Anisopus fenestra/is (Smart 1959), Basilia nana (Nussbaum 1960), and
Tipula vernalis (Mickoleit 1962).
As seen from Table XIX, the thoracic musculature in Diptera has an overall
similarity to that of Mecoptera. However, in Diptera there is a significant decrease in
LATERAL PARAPSIDAL
SUTURE
PRESCUTOSCUTAL
SUTURE
ANTERIOR
NOTAL PROCESS
PLEURAL
ARM
WING ------,'--
A
MATSUDA: TilE INSECT TiiORAX 323
PARASCUTAL
SHELF
(SURALARE)
POSTMEDIAN
NOTAL
PROCESS
---------SCUTUM
z (1 AX.)
FIG. 139. Diagram showing click mechanism of forewing in Sarcophaga bullata (from
Boettiger and Furshpan). See text for A. B. C.
the number of muscles, especially in cervical muscles, and in those attached to the pleuron
and the sternum. Strangely, the reduction of cervical muscles is more pronounced in
Tipula than in other Diptera.
Some tergopleural muscles (t-p 10, 13, 14, 15), however, have increased in the
number of muscle-fiber bundles, due most probably to enlargement and subsequent
splitting of muscles that were originally smaller in size. These muscles and the pleuro-
basalar muscle (p 2), which has also become enlarged in Diptera, are direct flight
muscles, and the presence of such powerful direct flight muscles in this order represents
a culmination of their development in the evolution of winged insects.
p-cx 8 and s-ex 4 in Diptera have not been found in Mecoptera. It remains to be
seen whether they represent new formation in Diptera, or if their absence in Mecoptera
is due to the lack of sufficient knowledge in the musculature of that order.
The larval musculature: A summary of Dirkes' (1928) study of the larval
musculature in Psychoda alternata follows. This work is the only one that adequately
describes and illustrates the thoracic musculature of any dipterous larva.
In the mesothorax (Fig. 140A) nine dorsal longitudinal muscles extend the entire or
nearly the entire length of the segment, and three shorter dorsal muscles are present;
four of these 12 muscles, which are shown by broken lines, become the dorsal longi-
tudinal muscles in the adult. Six vertical dorsoventral muscles are confined to the
lateral (pleural) wall of the mesothorax; the seventh lateral muscle is an oblique inter-
segmental muscle that connects the mesonotum and the metasternum; one of the two
other oblique lateral muscle is t-p 30 that is present in other larval holometabolous insects.
Six ventral muscles extend throughout the entire length of the mesosternum; two oblique
ventral muscles, one from the anterior border and another from the posterior border of
the mesothorax, meet on the middle of the sternum. The metathoracic musculature are
very similar to the mesothoracic musculature. In the prothorax there are more muscles
as seen in Fig. 140A.
Most of these larval mesothoracic muscles cannot be homologized with muscles in
adult insects including Thysanura, i.e. they are caenogenetic. The most conspicuous
feature of this larval musculature is the abundance of pleural muscles, which are only
poorly represented in Thysanura. The caenogenetic muscles degenerate later and have
nothing to do with production of adult muscles.
The postembryonic development of the flight muscles: In Simulium (Hinton
1959a), among Nematocera, the imaginal discs of the indirect flight muscles and that of
tergal depressor of the trochanter ( t-tr 1) are present in the second stage larva, and these
muscles develop quite independently of the larval muscles in all postembryonic stages.
In Psychoda Dirkes (1928) and Schmidt (1929) have found that four out of 12 dorsal
muscles present in the larva become the dorsal longitudinal muscles in the adult; the
dorsoventral flight muscles (t-p 5, 6) and the tergotrochanteral muscle (t-tr 1) develop
later from imaginal rudiments. What Weismann (1866) found about the development
of flight muscles in Corethra was similar to that in Psychoda in that indirect flight muscles
develop, through the histological differentiations, from the rudiments that were present
in the embryo. In Culex Hulst (1906) found that the flight muscles originated from
imaginal discs by proliferation of the embryonic cells, which has persisted undifferentiated
during the larval stage.
An important feature common to the development of flight muscles in all of these
Nematocera, as the preceding discussion indicates, is that they develop independently
throughout the postembryonic development.
Among Brachycera, imaginal muscles in the larva of Pachygaster are laid down as
three dorsal rudiments (Strange of Hansel) connected by a peculiar network (Fig. 140B).
According to Hansel ( 1913), the imaginal muscles arise from this system by in-
corporating some elements of larval muscles. In the fully grown larva of Strationyidae,
according to Jusbaschjanz ( 1910) the imaginal flight muscles are represented by longi-
tudinal and transverse rudiments (Strange). Jusbaschjanz ( 1910) showed that these
MESOTHORAX
.A
B
FIG. 140. A, prothoracic and mesothoracic muscles in larva of Psychoda sp. (from Dirkes);
B, diagram showing distribution of imaginal thoracic muscles in old larva of Pachygaster
meromelas (from Hansel).
two sets of rudiments of the imaginal muscles arise from the rudiment already present
in the embryo.
Among higher Diptera, the indirect flight muscles and leg muscles in Musca
vomit aria, according to Weismann (1864), originate from the decay of larval tissues,
and represent an entirely new formation during the pupal stage. Viallanes ( 1882) found
in Musca also that the indirect flight muscles arise from a special rudiment which was
formed from the fat cells. Kiinckel (1875, in Volucella), Ganin (1876, in Anthomyia),
and Lowne (1893-1895, in Calliphora) were the earlier workers who maintained the
origin of indirect flight muscles from the imaginal disc.
Perez's (1910) more careful study on Muscidae, however, showed that the indirect
flight muscles arise from two different elements, larval muscles and myoblasts. Accord-
ing to Perez, the myoblasts, which have persisted since the embryonic stage, penetrate
three degenerating larval dorsal longitudinal muscles to form the dorsal longitudinal
muscles in the adult.
In Drosophila Tiegs (1955) found that free myoblasts adhering to the outer surface
of larval muscles invade the larval muscles fibers to produce flight muscles. Shatoury's
( 1956) experimental work on Drosophila has further revealed that cells migrating from
the developing trochanteral depressor ( t-tr 1) enter the three paired larval dorsal longi-
tudinal muscles ( t-14), and that these cells are responsible for division of the three larval
muscles on each side into six.
Clearly in higher Diptera, participation of other cells outside the degenerating dorsal
longitudinal muscles is indispensible for the development of the imaginal dorsal longi:
tudinal muscles. This contrasts with the development in Nematocera, in which the
homologous muscles develop independently, without participation of other cells.
Degeneration of adult flight muscles: As in some other orders, the postimaginal
degeneration of muscles (t 14, t-p 5, 6) occurs also in some Diptera. In Lipoptena
cervi, according to Mercier (1924), the indirect flight muscles (t 14, t-p 5, 6) completely
326
degenerate after the wings are broken off, and the insect becomes an t<Ctoparasite, while
at the same time the development of leg muscles ensues. Bursell ( 1961) found in adult
Glossina that indirect flight muscles ( t 14) continue to grow after the first blood meal;
according to Glasgow and Glasgow (1962), the increase of the same muscles is by the
enlargement of muscle tissue already existing on emergence.
Histology: As Tiegs' (1955) study showed, the indirect flight muscles in Diptera
are of the fibrillar type, and the fibers composing them are commonly of large diameters,
reaching a maximum of 1.8 mm in Rutilia (Tachinidae). In nematocerous families
fiber numbers may be as great as 450, but in higher Diptera the number of fibers is often
very small. In higher Diptera, when there are only a few fibers, they tend to be giants;
and when the body is large, they tend to have a large diameter. Earlier, Partmann
(1948) came to essentially the same conclusion in saying "Insegesament wird also
deutlich, dass die dorsalen Langsmuskeln der gut fliegenden Dipteren bei Erreichen einer
bestimmten absoluten Grosse ihren kompakten, geschlossenen Aufbau aufgeben, zu
einer Umkonstruktion der Muskeln schreiten (i.e., giant fibers). Bei den Nematoceen
wird diese durch Zerlegung in Langsstrange erreicht." In fact, many of Partmann's
figures show giant fibers. The fibril diameter of flight muscles in most flies ranges from
1.6 to 3.5 !1. (Tiegs 1955).
The direct flight muscles are of the tubular type (Williams and Williams 1943).
Smart (1957) found in Anisopus and Ptychoptera that the subalar-coxal muscle (t-cx 8)
is of the fibrillar type. Grant ( 1960) found that in Psychoda phalaenoides the tergal
depressor of the trochanter (t-tr 1) is of the tubular type, although this muscle is usually
of the fibrillar type in another psychodid genus, Pericorna, and in Simulium (Hinton
I959a).
THE TRICHOPTERA
The neck region: In Trichoptera the lateral cervical sclerite (Fig. 141A) articulates
anteriorly with a process on the postoccipital ridge, and posteriorly with the ventral
anterior margin of the anepisternum. The shape of the lateral cervical sclerite and its
position relative to the proepisternum are very similar to those of Micropteryx (Fig.
144A). As in Micropteryx, therefore, this lateral cervical sclerite in Trichoptera most
probably represents a modified preepisternum.
73
Tindall (1965) unquestioningly in-
terpreted this sclerite in Limnephilus as the preepisternum.
In Neuronia (Fig. 141A) a setigerous plate is detached dorsally from the narrowed,
anterior partof the lateral cervical sclerite. In N euronia and in a species of Trichoptera
studied by Crampton (1926a), the posteroventral angle of the lateral cervical sclerite
meets the anterolateral angle of the sternal plate.
The pronoturn: As Schultz (1914) showed, the pronotum in Trichoptera has paired,
oval, setigerous protuberances on its anterior dorsal surface. They are homologous
with the patagia, which occur in the corresponding area of the pronotum in Lepidoptera
(Figs. 144C, 150A).
The propleurosternurn: In Neuronia (Fig. 141A) the pleural suture is cut through
near its ventral end by a narrow, oblique, membranous cleft that is serially homologous
with the paracoxal suture in the pterothorax. This cleft divides the episternum into the
anepisternum and the katepisternum. The latter differentiates apically as a narrow
trochantin, which, in Lirnnephilus (Tindall 1965), does not reach the coxal margin.
The epimeron is more or less greatly reduced or desclerotized. In Neuronia (Fig. 141A)
and Lirnnephilus the sclerotized epimeron is represented by a narrow, oblique bar.
The prosternum in Trichoptera is largely membranous. The median narrow sclero-
tized plate may not be continuous with the narrow posterolateral angle of the lateral
cervical sclerite on each side. Internally, the furca is united laterally with the posterior
end of the sclerotized epimeral area. The prothoracic spina is present in Trichoptera; in
Lirnnephilus it is long and projects into the mesothorax.
r.JSee p. 334 for discussion on the lateral cervical sclerite in Micropteryx.
327
TABLE XIX
Thoracic musculature in Diptera
T>pula Tabanus Orthellia Anisopus Muscle
(Mickoleit 1962) (Bonhag 1949) (Maki 1938) (Smart 1959) designations Remarks
1 1 cv(d)-t 1
2c, d op-t 2
2a, b 2, 3 op-t 3
3 9, 10 op-cv 2
2
4 op-cv 3
5 12 t-cv 1
3
8,9 11 t-cv 2 or 1
t-s 1
1
t-cv 3 2
17 13 cv-cx 3
5
6, 7 7, 8 (?) cv-s 1 3
4
10, 11 4, 5, 6 s 1, 2
30 (II) 28 22 2 t12or13
29 (II) 27 20, 21 1 t 14
42 (II) 42 7b t-p 4 4
64 (III) 59 49
27 (II) 29 23 4 t-p 5 or 6
60 (III) 56 48
44 (I I) 31 25 8a t-p 7 or 3 5
46 (II) t-p 8
45 (II) 32 24 t-p 9
34 (I I) 34b 28
62 (I II) 50
9a t-p 10 6
33 (I I) 34a 27 9a t-p 10? 7
35 (II) 35a 29 9b, i t-p 13
36, 37 (II) 35b 30 9b, ii t-p 14
63 (III) 60a, b 51
38, 39, 40 (II) 36, 37, 38 31,32,33,34 9c, i, ii t-p 15 8
43 (I I) t-p 15 ? 8
65 (III)
26 (II) 7a? t-p 18 9
16 (I) t-p? 10
15 (!) t-cx 7 1l
28 (II) 30 40 5
32 (II) 10 t-cx 8 12
62 (III) 61 52
31 (II) 33 35 8b p2
61 (I II) 57 53
58 (I II) p3
47 (II) 40 36 11a p-s 1
66 (III) 63 54
9 (I) 16 17 p-cx 4 or 13
t-cx 3
52 (II I) 66- 58 p-cx 4
65 (II I) p-cx 5
15 (I) p-cx8
41 (II-III) 43 47 s 13
17 (I)
s ? 14
TABLE XIX (Continued)
Tipula Tabanus Orthellia Anisopus Muscle
(Miekoleit 1962) (Bonhag 1949) (Maki 1938) (Smart 1959) designations Remarks
48 (II) 41 37 llb s-ex 2
67 (Ill) 64 55
7 13 16 s-ex 3
19 44 41
50 68 57
14b s-ex 4 15
6 12 14 s-ex S
18 45 38, 39 14a
49 67 56
8 14 s-ex 6
20
51
11 (I) 19a, b t-tr 1 16
46a
54 (III) 69a
22 (II) p-tr 1
55 (III)
10 (I) p-tr 2
24 (I I) 47a 44 13a p-tr 4
57 (III) 72a 61
12 19 s-tr 1
21 46b 12
53 69b 60
REMARKS (Table XIX)
( 1) Mickoleit (1962) described 3 in Tipula as arising from the pleuron, but it is
probably from the tergal area. Similarly, Bonhag ( 1949) described 8 and 9 as arising
from the propleuron. As already pointed out, the area that Bonhag called the anepi-
sternum is actually the anterolateral part of the pronotum.
(2) 1 in Tipula arises from the first phragma, due probably to the shift in origin.
( 3) 8 in Orthel/ia is said to be inserted on the posterior end of the ventral cervical
sclerite.
( 4) These muscles arise from or beneath the prescutal margin, not from a
prealar sclerite.
(5) In Mecoptera t-p 3 is present, but t-p 7 is absent. t-p 7 in Diptera may be
t-p 3 that has shifted its anterior point of attachment. In Anisopus Sa was described as
arising from the anterodorsal margin of the anepisternum, but in Smart's (1959) figure
it comes from the tergum.
(6) Metathoracic 62 in Tabanus and 50 in Orthel/ia extend between the anterior
notal wing process and the pleural wing process.
(7) These muscles are large, fan-shaped muscles that arise from the episternum
and are inserted on the first axillary sclerite. They may be derivatives of t-p 10.
(8) In Tipula 38, 39, and 40 are inserted on the fourth axillary sclerite, which
represents the detached distal end of the posterior notal wing process; in Orthellia also
homologous muscles are inserted on the fourth axillary sclerite; in Tabanus 36 and 38
are inserted on the definitive posterior notal process, which is, however, the postmedian
notal process; and 37 is attached to the fourth axillary sclerite. 43 in Tipula is dorsally
inserted on the posterolateral scuta! margin between the third and the fourth axillary
sclerites; this muscle may therefore be homologous with one of t-p 15 in Tabanus, which
is inserted on the postmedian notal process.
(9) 7a in Anisopus may be t-p 9.
(10) 16 in Tipu/a is a prothoracic t-p muscle, which is not homologized.
(11) 15 in Tabanus was described as a pleural remotor of the coxa, but the actual
point of origin is on the tergum.
(12) The muscles in the metathorax are attached to the ventral pleural arm.
( 13) 9 in Tipula has a broad attachment on the cranial side of the tergopleural
ridge (pleural ridge of Mickoleit), and 17 in Orthellia has a wide origin on the tergo-
pleural area. 16 in Tabanus originates on the swollen anterolateral pronotum (pro-
episternal callus of Bonhag) and appears to be a tergocoxal muscle. These muscles
may, therefore, represent t-cx(ti) 3.
( 14) 17 in Tipula arises from the apophysis of the presternum of Mickoleit and is
inserted on the mediocranial process of the profurca. This muscle may be a cv-s muscle.
(15) 14b in Anisopus arises posteriorly from the spinasternum and is inserted on
the posterior rim of the coxa. This muscle is, therefore, s-ex 4, which is absent in
Mecoptera.
( 16) Prothoracic muscles in Tipula and Tabanus were described as pleural muscles.
The mesonotum: In Neuronia (Fig. 141C) the anterior part of the mesonotum is
vertically bent and is not visible in the dorsal view. The prescutoscutal suture delimiting
the prescutum is not recognized, but the prealar arm, which unites with the basalare, is
probably prescutal in origin. The intersegmental membrane is attached along the
horizontal line shown in Fig. 141C; the area ventral to the line may be considered as the
first phragma. In any case, the vertically deflected part of the prescutal area functions
as the first phragma in Trichoptera. In the dorsal view (Fig. 141B), the median longi-
tudinal suture extends from the anterior end of the mesonotum to the scutoscutellar
suture. The anterolateral scuta! suture in Neuronia (Fig. 141B) is well marked, and it
extends posteriorly almost to the base of the median notal wing process. The postero-
lateral scuta! suture, which is distinctly ridged internally, almost reaches the lateral end
of the scutoscutellar suture.
The mesothoracic postnotum in Neuronia (Fig. 141D) is clearly separated from the
scutellum by a membrane. A narrow, lateral postnotum is separated on each side from
the rest of the postnotum by a longitudinal ridge. The lateral postnotum extends
anteriorly beneath the wing base, and the ridge that separates the lateral postnotum
articulates with the dorsal tip of the epimeron. The posterior margin of the postnotum
is deflected to form the anterior wall of the second phragma.
The mesonotal lateral margin and axillary sclerites: In Neuronia (Fig. 141C) the
large setigerous tegula lies dorsal to the prealar arm, and is connected with the basalare
by a subtegular arm. All the processes on the scuta! lateral margin that have been
recognized (p. 14) are present, as shown in Fig. 141B. The anterior and the antemedian
processes form a rather thick suralare, and the tergal fissure is narrow. The posterior
notal wing process is narrow and long; this process is detached to form the fourth axillary
sclerite in the pupa of Dicosmoecus (Goodrich 1935).
The first axillary sclerite, as Sharplin ( 1963a) has shown in Limnephilus, has
ligamentous connections with the scutallateral margin (Fig. 142A). The ventral second
axillary sclerite in Limnephilus (Fig. 142B), according to Sharplin ( 1963a), is connected
with the base of the R vein through a sclerotized flange; it also has a ligamentous connec-
tion with the pleural wing process and with the subalare. The third axillary sclerite has
the usual connection with the distal end of the posterior notal wing process, and also has
a ligamentous connection with the first axillary sclerite. The complex connections of
the ventral third axillary sclerite require a thorough study in the future.
The mesopleurosternum: An oblique pleural suture separates the anterior episternum
and the posterior epimeron. No pleural arm is differentiated internally on the pleural
ridge. In the episternal area the anepleural cleft becomes confluent with the paracoxal
suture laterally (Fig. 143A). Matsuda ( 1960a) regarded the latter suture in Arctopsyche
330
STERNUM
TROCHANTIN
PLEURAL _..,
PROCESS (for COXA)-"'
PREALAR ARM
~
----- , . ~ - - ~
SCUTOSCUT ELLAR
B
POSTEROLATERAL
SCUTAL SUTURE
LATERAL
POSTNOTUM
A
MEDIAN POSTNOTUM
& PHRAGMA
D
VENTRAL
PROCESS ----1-1--U
---- EPIMERON
PARACOXAL
CLEFT
TEGULA
POSTERIOR
NOTAL
PROCESS
'::f-.Z::::>-1-f--LATERAL
POSTNOTUM
EPIMERON
FURCA 2
FIG. 141. Thoracic structures in Neuronia regina (original). A, propleuron and lateral
cervical sclerite; B, dorsal view of anterior part of mesonotum; C, cephalic view of mesonotum
and mesopleuron; D, dorsal view of posterior part of mesonotum and its connection with
epimeron and sternum.
ANTERIOR &
ANTEMEDIAN
NOTAL PROCESSES---
MEDIAN NOTAL
PROCESS
RADIAL BRIDGE
PLEURAL WING
PROCESS----'
POSTMEDIAN
L - - - - - ~ NOTAL PROCESS
A
B
MEDIAN PLATE
VENTRAL
2'ND AXIL.
SCL.
Fro. 142. Forewing base structures in Limnephilus bimaculatus (from Sharplin). A, connec-
tion of first axillary sclerite of forewing with mesoscutal lateral margin; B, inner lateral view
of right second axillary sclerite.
as the basal (distal) line of the katepisternotrochantin. An oblique paracoxal suture in
Micropteryx (Fig. 147A), which extends onto the epimeron, has apparently become
lost on the epimeron in Trichoptera.
In Neuronia (Fig. 143A) the anepisternum is connected with the prealar arm, and
the basalare with the tegula. The preepisternum on either side apparently meets the
median longitudinal suture, which is the line from which the sternum. invaginates. In
Arctopsyche (Matsuda 1960a) the katepisternum is almost totally sclerotized and the
apically differentiated trochantin articulates with the coxal margin, although in N euronia
(Fig. 143A) the area above the coxal margin is membranous. The narrow ventropleurite
(ventral process) articulates with the inner coxal margin (Fig. 141D). The epimeron
(Fig. 141D) is enlarged dorsally by sclerotization. This definitive epimeron articulates,
at its anterodorsal angle, with the postnotum to form the postalar bridge. In Trichoptera
the bridge is formed mainly by extension of the epimeron, the postnotum being reduced
laterally. Furthermore, the sclerotized dorsal part of the definitive epimeron is in-
distinguishably fused with the dorsally elevated sternal furca, which forms the ridged
posterior border of the mesothoracic segment (Fig. 141D).
The basisternum is represented mainly by an external shallow, median longitudinal
line (Fig. 143A). Presumably the originally basisternal area has become indistinguish-
ably fused with the proximal area of the preepisternum. The pleurosternal suture is
absent, at least in Neuronia. The posterior half of the mesosternum is deeply invaginated
to form a lateral skeletal framework (the lateral furca), which is indistinguishably fused
with the epimeron (Fig. 141D); another furcal branch arises more proximally. In
Limnephilus Tindall (1965) showed a medially divided, heart-shaped sternal plate
exposed between the coxae and called it the anterior sternal sclerite; this area corresponds
to the anterior part of the deeply invaginated posterior sternum, or the furcasternum in
Neuronia (Fig. 143A). The mesothoracic spina is not known to occur in Trichoptera.
The metanotum: In Neuronia the anterior part of the metanotum is semidetached
from the rest of the metanotum by the membranization that surrounds it (Fig. 143C).
This part of the metanotum bears the poorly developed prescutal prealar arms and is
folded over the mesothoracic postnotum to form the posterior wall of the second phragma.
The anterolateral scuta! suture and the scutoscutellar suture are similar to those in the
mesonotum. The posterolateral scuta! suture cuts off a small posterolateral scuta! area
bearing the posterior notal wing process. The lateral margin of the scutum in N euronia
is provided with all notal wing processes except the postmedian process (Fig. 143C).
The metathoracic postnotum is detached from the rest of the metanotum and is
united with the anterior margin of the first abdominal tergite. This posteriorly displaced
postnotum assumes a vertical position and is functionally the third phragma. Further, as
Goodrich (1935) pointed out, the postnotum has a lateral connection (lateral post-
notum) with the metepimeron (Fig. 143B) as in the mesothorax. Therefore, Tindall's
( 1965) claim that the structure is the invagination of the first abdominal terg(te is invalid.
The metapleurosternum: In the metapleuron of Neuronia the dorsal margin of the
basal are has a direct association with the humeral plate ( basicostale) ; in the resting
position the latter fits into a concavity of the former. In Limnephilus, as Tindall (1965)
showed, the proximal end of the narrow preepisternum meets the medially divided
triangular plate, and the oblique, lateral line of this median triangular plate appears to be
a pleurosternal suture comparable with the presumed pleurosternal suture in Tipula
(Fig. 136A). Tindall thought this plate was serially homologous with the similarly
shaped area in the mesothorax already noted. The two areas, however, differ in
position, and the sclerite on the metathorax is possibly the exposed basisternum (see
p. 37). In Neuronia and Limnephilus a slender trochantin is well differentiated apically
from the katepisternum ( eutrochantin), and articulates with a slender bar that arises
from the coxal margin.
The adult thoracic musculature: Maki (1938) studied thoracic muscles in
Stenopsyche griseipennis and Tindall ( 1965) studied those in Limnephilus marmoratus.
They are homologized in Table XX.
Compared with the thoracic muscles in Mecoptera, the number of muscles in
Trichoptera is somewhat reduced. However, p-cx 9 is absent in Mecoptera and present
in N europtera, Trichoptera, and Lepidoptera.
Gross aspects of the postembryonic muscle development: Tindall (1963, 1964)
studied muscles in the larvae of Limnephilus sp. and Triaenodes bicolor. Unfortunately,
the fate of these muscles during later developmental stages was not traced. Although
many of these muscles can be homologized with adult muscles, some of them are
obviously purely larval (caenogenetic) muscles that disappear later, and some others are
Fro. _143. Thoracic structures in Neuronia regina (original). A, mesopleuron; B, connection
of third phragma and metepimeron; C, metanotum and first abdominal tergite.
PREALARARM
AN EPISTERNUM
ANAPLEURAL CLEFT
FURCASTERNUM-----
TROCHANTIN-
VENTRAL
PROCESS
ANTEROLATERAL
SCUTAL SUTURE
ANTERIOR
NOTAL PROCESS
ANTEMEDIAN
NOTAL PROCESS
MEDIAN
NOTAL PROCESS
POSTEROLATERAL
SCUTAL SUTURE
LATERAL
POSTNOTUM
l'ST .ABO.
TERGITE
333
~ - - - - - - - - - TEGULA
BASALARE
EPIMERON
PLEURAL SUTURE
PREALAR
ARM
TERGAL
FISSURE
SCUTOSCUTELLAR
SUTURE
c
EPIMERON 3
. 334
apparently palaeogenetic muscles inherited from the wingless ancestor that are retained
only during the larval stage. Some of the muscles that belong to the last category follow.
3.40 (op-t 4) in Limnephilus extends from the middle of the mesonotum to the
posterior dorsal region of the head capsule; it is probably homologous with 9 in Lepisma.
5.31 in Limnephilus is probably t-p 1, which occurs in the adult of Corydalus and
Lepisma.
5.34 in Limnephilus and Triaenodes connects the anterior (or medial) lateral part
of the tergum and the posterior ventrolateral part of the intersegmental fold; this muscle
is t-p 30, which may be present in some Apterygota. This muscle appears to be present
also in Limnephilus fiavicornis, although Winkler (1959) did not number it.
7.13-7.16 in the pterothorax of Lirnnephilus and Triaenodes are tergopleural muscles
that are inserted on the anterodorsal edge of the pleuron; they are t-p 27 and t-p 28,
which are present in the Apterygota and larval Corydalus.
THE LEPIDOPTERA (ZEUGLOPTERA AND DACNONYPHA)
The neck region: In Micropteryx (Fig. 144A) the only cervical sclerite is the lateral
cervical sclerite, which is obviously a modified preepisternum, as is clear by its association
with neighboring sclerites. It is separated laterally by the anapleural cleft from the
anepisternum, and its caudal margin lies anterior to the katepisternum, as is typically the
case with the preepisternum; only its narrow anterior part represents a conspicuous
modification for articulation with the head. The same relationships between the lateral
cervical sclerite and the adjacent episternal plates can also be seen in the figure of
Micropteryx given by Hannemann ( 1956) and in the figure of Saba tinea by Crampton
(1926a).
The prothorax: In Micropteryx (Fig. 144C) the short pronotum has paired trans-
verse membranous spots that are homologous with the patagia in other Lepidoptera. In
Eriocrania, as Schultz ( 1914) showed, the patagia are rounder and larger than those in
Micropteryx.
The propleuron in Micropteryx articulates dorsally with the process formed on the
anterolateral angle of the pronotum (Fig. 144C). The pleural suture is distinct. In
the episternum the anapleural cleft, as already noted, separates the preepisternal lateral
cervical sclerite from the anepisternum.
The katepisternum (Fig. 144A) is separated from the preepisternal area by a
membranous cleft, which corresponds in position to the paracoxal suture in the meso-
thorax (Fig. 147A). The lateral (distal) base of the katepisternum is marked by a
straight and oblique suture, which corresponds to the part of the paracoxal suture that
extends into the epimeron in the mesothorax. The trochantinal plate is detached from
the proximal end of the katepisternum, and it articulates with the coxal margin. The
epimeron is a narrow plate along the pleural suture.
In Micropteryx the sternum (Fig. 144A) is represented by a small sclerotized plate
between the coxae; this plate is provided with a median longitudinal sulcus. In the dorsal
view (Fig. 144B) the internal ridge of the median longitudinal sulcus is continuous
laterally with the furcal frame work, which is in turn laterally continuous with the
posterior end of the narrow epimeron near the pleurocoxal articulation. The spina
projects dorsally from the median posterior end of the sternum.
The mesonoturn: The anterior part of the mesonotum in Micropteryx (Fig. 144D)
is vertically deflected as in Trichoptera, and it serves as the area for attachment of the
dorsal longitudinal muscles. No differentiation of the first phragma occurs anterior to
this area. The narrow membranous cleft that separates the prealar arm probably corres-
ponds to the prescutoscutal suture, since such a membranized prescutoscutal suture
occurs in some other insects (e.g. Heteroptera). The prealar arm does not reach the
episternum. The large tegula is supported by a thin, semilunar subtegula (Fig. 145),
which, as Sharplin (1963a) maintained, is a detached prescutal sclerite and appears
homologous with the prealar sclerite in other insects.
TABLE XX
Thoracic musculature in Trichoptera
Stenopsyche Limnephilus
(Maki 1938) (Tindall 1965) Muscle designations Remarks
2?
1.54 cv(d)-t 1 1
4 1.55, 1.56 op-t 1, 2 2
3
1.01-1.08, op-t 3
1. 51-1.53
8 3.03, 3.04 op-cv 1, 2
15
op-p 1
9 3.05 op-s 2
10 3.01 t-s(cv) 1 3
1.3 4.10 t-s(cv) 9 4
11 t-cv 1 5
12 t-cv 2
3.02 t-cv 3
6 cv-s 1
2.51 cv-s 4 ? 6
5 2.01, 2.02 s 1 or 2
29 (II) 1. 57 t 12
57 (III) 1.57
1 1.09-1.11 t 13
28 1.54
56 1.54
2? 1.54 t 14
17 1.01-1.05
55 1.01
16, 17 4.03-4.08 t-p (prothoracic)
35 (I I)
61 (III)
t-p 3 7
37 (II) 4.01 t-p 4
63 (III) 4.01
33 (II) 3.03 t-p 5 or 6
59 (Til) 3.03
36 (II) 5.01 t-p 7
62 (III) 5.01
4.02 (III) t-p 10
38 (II)
64 (III)
p-t 12
39 (I I) 6.01 t-p 13
65 (III) 6.01
40 (II) t-p 14
66 (III)
4.02 (II) t-p 15
5.52 (II) t-p 16
5. 52 (III)
14 3.51 t-s 1
34
60 3.51
45 (II) 3.04 t-ti 2
70 (!II) 3.04
20 7.52 t-cx 7
47 7.52
72 7.52
TABLE XX (Continued)
Stenopsyche Limnephilus
(Maki 1938) (Tindall1965) Muscle designations
Remarks
48 (I I) 5.51 t-ex 8
73 (III) 5.51
44 (II)
69 (III)
p 1
67 (III) p 2
41 (II) p3
42 (II) 4.51 p-s 1
32 (HI) p-s 6
50 (I I) 5.02 p--<:x 2 or 3
76 (III) 5.02
24, 25 (!) 7.03, 7.04 p--<:x4 8
75 (III) p-ex 5
21 (I) 7.51 P--<:X9
31 (I-II) 2.02 s 12
30 (HI) 2.51 s 13 9
58 (II-III) 2.01, 2.02
7(!) 2.01 s 14
43 (I I) 4.52 S--<:X 2
68 (Ill) 4.51
23 7. 53, 7. 54 s-ex 3
49
74
22 (I) 7. 55?, 7. 56 S--<:X 4 10
18, 19 7.01, 7.02 s-ex 5
46 7.11, 7.12?
11
71
7.13 (II)
7.13 (III)
s-ex 6
51 (II) 8.01, 8.02 t-tr 1
77 (Ill) 8.01, 8.02
26 (I) p-tr 1
52 (I I) 8.03 p-tr 2
78 (II I) 8.03
53 8.11 s-tr 1
79 8.11
8.11
REMARKS (Table XX)
( 1) 2 in Stenopsyche arises anteriorly from the anterolateral part of the
pronotum.
(2) 1.56 in Limnephilus is op-t 1; others are op-t 2.
(3) 3.01 in Limnephilus is attached to the posterior tentorial bridge.
( 4) 13 in Stenopsyche connects the mesothoracic prealar arm? and the lateral
cervical sclerite.
(5) 11 in Stenopsyche stretches between the anterolateral corner of the pronotum
and the middle of the inner portion of the lateral cervical sclerite.
(6) 2.51 in Stenopsyche connects the median anterior sternal sclerite between
the lateral cervical sclerites with the profurca.
(7) There is an homologue in the mesothorax of Limnephilus that Tindall
( 1965) did not number.
(8) These muscles may be p-cx 2 or 3.
(9) 2.01 and 2.02 in Limnephilus may be p-s 9.
(10) 7.55 in Limnephilus is a crucial muscle.
337
(1 I) 7.11 in Limnephilus is attached to the anterior median edge of the sclero-
tized part of the coxa; 7.12 is attached on an anterior inflexion of the coxal rim.
On the scutum (Fig. 144E) the anterolateral scuta! suture separates the suralare, and
the posterolateral scuta! suture extends to the scutoscutellar suture. The scutoscutellar
suture is broadly produced anteriorly and is continuous laterally with the axillary cord.
The postnotum lies subvertically behind the mesonotum proper and forms the anterior
wall of the second phragma, as in Trichoptera; its anterolateral angle (lateral arm of
Sharplin, 1963b) articulates with a process on the lateral posterior margin of the
scutellum. The lateral postnotum (laterophragma, or laterpostnotum) in Micropteryx
is detached as a narrow plate and extends anteriorly into the area beneath the wing base.
The plate articulates with the sci erotized epimeral plate (Fig. 147C), as in Trichoptera.
The mesonotal lateral margin and axillary sclerites of the forewing: On the scuta!
lateral margin of Micropteryx (Fig. 144E) the anterior and antemedian notal processes
are not well differentiated, and together form a robust suralare; the median, postmedian,
and posterior processes are clearly differentiated, as in Trichoptera.
In Micropteryx, as Sharplin (1963a) showed (Fig. 146A), the first axillary sclerite
is connected by ligaments with the scuta! margin on the anterior, antemedian, median,
and postmedian notal processes; its head has a ligamentous connection with the basi-
subcostale (Fig. 146B). The second axillary sclerite (Fig. 146B) has connections
anteriorly with the base of the R vein, and distally with the proximal median plate.
Proximally it articulates with the first axillary sclerite. Ventrally (Fig. 145), the second
axillary sclerite is connected with the subalare and the tip of the pleural wing process,
and with the proximal median plate by means of a bending cuticle (p. 16) on the ventral
wing membrane. The third axillary sclerite (.Fig. 146C) is connected anteriorly with
the proximal median plate, distally with the A vein, and proximally by a short ligament
with the semidetached, posterior notal wing process.
The mesopleurosternum: The pleural ridge in Micropteryx is about the same
thickness throughout, and the pleural arm is absent. The dorsal end of the ridge forms
an unswollen conical pleural wing process, and the ventral end, which articulates with
the coxa, forms a shallow socket.
In the ventral view (Fig. 147A) the paracoxal suture is distinct. Its lateral (distal)
half is sharply bent posterolaterad across the pleural suture, and almost reaches the
posterolateral angle of the epimeron, as in Sialis (Fig. 120A). In Mnemonica, as
Crampton (1920b) showed, the paracoxal suture is absent on the epimeron; in Microp-
teryx arunce/la, also shown by Crampton ( 1920b), the same suture is present on the
epimeron.
The anapleural cleft divides the anapleuron into the preepisternum and the anepi-
sternum, and it meets the paracoxal suture distally. The basalare (Fig. 147A) is
separated from the anepisternum by a narrow cleft. An apodeme arising from the
pleural ridge (tergopleural apodeme of Sharplin 1963a) extends toward the subtegula
(Fig. 145). The preepisternum appears to reach the median longitudinal axis of the
mesothoracic ventral surface, where the sternum invaginates, the pleurosternal suture
being absent. The sclerotized katepisternum is proximally continuous with the
trochantin, which articulates with the coxal margin. The ventral process (condyle) is
more clearly seen in the dorsal view (Fig. 147C); its continuation with the median
posterior end of the internal sternal skeleton is evident. The katepimeron is a narrow,
subtriangular area along the posterior margin of the epimeron. The dorsal margin of
the anepimeron is deeply concave, and the subalare lies on the membranous area above it.
ANAPLEURAL
CLEFT
SCUTOSCUTELLAR
SUTURE'-
POSTMEDIAN
NOTAL PROCESS
POSTEROLATERAL
SCUTAL SUTURE
EPISTERNUM
B
/
' /
c
. .. .. .: ::.:: . .
LATERAL CERVICAL
SCLERITE
(PREEPISTERNUM)
KATEPISTERNUM
TROCHANTIN
PLEURQN
ANTEROLATERAL
SCUTAL SUTURE
& ANTEMEDIAN

, / TERGAL FISSURE
FIG. 144. Thoracic structures in Micropteryx calthel/a (original). A, ventral view of pro-
thorax and neck region; B, dorsal internal view of prothorax, propleuron is somewhat rotated;
C, dorsal view of pronotum; D, cephalic view of anterior part of mesonotum; E, dorsal view
of mesonotum.
. 2'ND AXIL. SCL.
TERGOPLEURAL
APODEME
PLEURAL
WING PROCESS
BASAL ARE
SUBALARE
EPIMERON
FIG. 145. Lateral view of forewing base in Micropteryx calthella (from Sharplin).
The mesosternum is represented externally by a short, median longitudinal suture
that invaginates. In the dorsal view (Fig. 147C) the sternal furca is highly complex; its
anterior arm is united with the posterior (dorsal) epimeral lobe.
The metanotum: In Micropteryx the median anterior part of the metanotum is
superposed upon the mesothoracic postnotum, as in Trichoptera. The anterior margin
of the metanotum has a prescutal apodeme and the prescutal prealar arm (Fig. 148).
The prescutal apodeme of Sharplin ( 1963b) is probably serially homologous with the
subtegula in the mesothorax; muscle t-p 4 is attached to this apodeme. The prealar arm
is connected by a ligament with the dorsal margin of the anepisternum (Fig. 148).
On the metanotum all the sutures of the mesonotum can be recognized (Fig. 1470),
and similarly the notal processes of the scutum. As in Trichoptera, the metathoracic
postnotum is displaced posteriorly and is united with the first abdominal tergite to form
the third phragma; the phragma in turn articulates laterally with the swollen dorsal end
of the metepimeron as in Trichoptera. For convenience, the axillary sclerites of the
hind wing base in Micropteryx (Fig. 154A) are referred to in the discussion on the hind
wing base in Monotrysia and Ditrysia.
The metapleurosternum: In Micropteryx the metepisternum (Fig. 147B) differs
from the mesepisternum in having a secondary marginal evagination (hypopteron,
prepectus) on the anepisternum, a greatly reduced preepistemum, and an incomplete
paracoxal suture. In addition, the katepisternum is somewhat thickened along its
posterior margin. A narrow trochantin articulates basally with a projection on this
thickened katepisternal area, and apically with a slender process arising from the coxal
margin. The lateral furcal arm extends cephalad and unites with the dorsally extended
epimeron.
THE LEPIDOPTERA (MONOTRYSIA AND DITRYSIA)
The neck region: In, some Monotrysia the preepisternal origin of the lateral cervical
sclerite can easily be surmised, although the membranous area separating the
cervical sclerite from the episternum is generally more extensive than in Micropteryx
and Trichoptera. In Hepialus (Fig. 149A), for instance, the caudal end of the lateral
cervical sclerite almost touches the anterolateral angle of the sternal plate, and its distal
margin clearly lies anterior to the katepisternum, as in Micropteryx.
In Papilio (Fig. 149B), as Weber (1924a) and Crampton (1926a) showed, the
lateral cervical sclerites on the two sides are connected by a transverse sclerotization,
340 MEMOIRS OF THE ENTOMOLOGICAL . SOCIETY OF CANADA
ANTERIOR
NOTAL PROCESS l'ST AXIL. SCL.
TERGAL FISSURE
POSTMEDIAN
NOTAL PROCESS
A
RADIAL PLATE
BASISUBCOSTALE
M + Cu 1
2'ND AXIL. SCL. PROX. MED. PL.
FIG. 146. Forewing base structures in Micropteryx calthel/a (from Sharplin). A, connections
of first axillary sclerite with scutal margin; B and C, dorsal view of forewing base.
. .
\1';---PLEURAL
WING PROCESS
PLEURAL
SUTURE
----l,---ANEPIMERON
341
----r
ANEPISTERNUM . :7
PARACOXAL
SUTURE
TROCHANTIN
VENTRAL
PROCESS
EPIMERON
FURCA
/KATEPIMERON
.../------- HYPOPTERON
-1/.--/-;,L-------- EPIMERON
KAT EPISTERNUM
TROCHANTIN
B
SCUTUM
ANTEROLATERAL
SUTURE SCUTAL SUTURE
/ -----ANTERIOR &
-' ANTEMEDIAN
) NOTAL
PROCESSES
/ MEDIAN NOTAL
!. PROCESS
/
POSTMEDIAN
EPIMERON
D
3'RD PHRAGMA
FIG. 147. Pterothoracic structures in Micropteryx calthella (original). A, ventral view of
mesopleuron; B, ventral view of metapleuron; C, dorsal inner view of mesopleurosternum; D,
dorsal view of metanotum and first abdominal segment.
342
BASALARE
FIG. 148. Lateral view of hind wing base in Micropteryx calthella (from Sharplin).
thereby comptetely losing the connection or association with the sternum; here the
preepisternal origin of the lateral cervical sclerite is not directly evident. Weber (1924a)
called this transverse bar the "Kehlspange" and Crampton (1926a) called it the pre-
sternum. Since, however, the presternum as a primary division of the sternum is absent
in Trichoptera, Zeugloptera, and in a primitive genus H epialus, the transverse sclerite in
Papilio, must be a secondarily produced ventral cervical sclerite within the Lepidoptera;
Srivastava (1961) also thought it was the ventral cervical sclerite. In Papilio demoleus
an additional ventral cervical sclerite occurs; according to Ehrlich ( 1958b), the ventral
cervical sclerite occurs in some other genera of Papilionidae, Historis, and Apatura.
The pronotum: The pronotum is typically composed of a median dorsal plate,
paired dorsolateral membranous areas called the patagia, and paired lateral plates con-
tinuous with the dorsal margin of the propleuron (Fig. 150A). As Wellman ( 1894)
and Schultz ( 1914) showed, differentiation of these pronotal parts in higher Lepidoptera
is much more pronounced than in Zeugloptera. Schultz ( 1914) distinguished four types
of patagia; viz., vesicle-shaped, pocket-shaped, balloon-shaped and wing-shaped. In
some groups an additional protuberance lies posterior to the patagium, and is called the
parapatagium. The dorsal plate is also a highly elaborate structure in higher Lepidoptera.
The origin of the lateral plate (Fig. 150A), whether it is tergal or pleural in origin,
has been disputed. Weber (1924a) contended that it is episternal in origin, since in
Cossus the borderline between the lateral plate and the pronotum is recognizable only
dorsally. Michener (1952) was right, however, in maintaining that it is clearly a
pronotal structure in Hepialus, Phassus, etc.; in Micropteryx (Fig. 144C) the corres-
ponding area is also clearly pronotal.
The propleuron: The episternum in the prothorax of higher Lepidoptera is more
strongly sclerotized than in Micropteryx, and its division into the dorsal anepisternum
and the ventral katepisternum is more or less obliterated. In Hepialus (Fig. 149A) the
projection on the ventral margin is probably the reduced katepisternum (Lateropleurit
of Weber, 1924b), since the position of this area relative to the preepisternal lateral
LATERAL
CERVICAL
SCLERITE
AN EPISTERNUM
EPIMERON
LATERAL CERVICAL
SCLERITE------Jl
VENTRAL CERVICAL
SCLERITE
KA TEPISTERNUM
EPISTERNUM
PLEUROSTERNAL SUTURE?
A
B
KAT EPISTERNUM
!.-----ANAPLEURAL
CLEFT
PARACOXAL
SUTURE
------AN EPISTERNUM
FIG. 149. Ventral view of neck, prothorax and mesothorax in Lepidoptera (from Weber).
A, Hepialus humuli; B, Papilio machaon.
cervical sclerite and the sternum agrees with that of the katepisternum in Micropteryx
(Fig. 144A). In many Lepidoptera the katepisternal projection has become in-
distinguishably fused with the anterior part of the sternum to form a transverse bridge,
as seen in Papilio (Fig. 149B), Protoparce (Madden 1944), Eacles (Michener 1952),
and some higher Diptera (Fig. 131). This transverse bridge may be called the pre-
coxale, but this definitive precoxale is not homologous with the precoxale in hemi-
metabolous insects, which is formed by participation of the preepisternum.
The trochantin is aosent in most Lepidoptera. Madden ( 1944) showed in Proto-
parce a small, triangular plate which may represent the trochantin. The ventral process
(ventropleurite) is absent, and hence there is no ventral articulation with the procoxa in
Monotrysia and Ditrysia, as in Micropteryx and Trichoptera. In the prothorax the
pleurocoxal articulation occurs only between the ventral end of the pleural ridge and the
coxal margin. This relatively weak pleurocoxal articulation is related to the atrophy of
the front leg in Rhopalocera. The epimeron is usually narrow.
DORSAL PLATE
;__---LATERAL PLATE
(PRONOTUM)
B
FIG. 150. A, posterior internal view of prothorax in Danaus plexippus (from Ehrlich); B,
tegula of Cat ocala elocata (from Hering).
The prosternum: The sclerotized sternal plate either becomes prolonged anteriorly
to meet the inner posterior angle of the lateral cervical sclerite as in H epialus (Fig.
149A), or its anterior end, as already noted, becomes indistinguishably fused with the
katepisternal extension (Fig. 149B).
Internally, the furcal arm is united with the ventral end of the epimeron as in
Micropteryx, although its exact point of fusion may be obscure in higher forms. Weber
( 1924b) thought that it was united with the pleural ridge. As his figures of Hepialus
and Cossus show, however, the furca is united with the ventral end of the epimeron,
which Weber regarded as the ventral end of the pleural ridge. Dierl (1964) appears to
be alone in describing the epimeron in Fumea as the area of union with the furca, and
his is most probably correct. The prothoracic spina is perhaps always
present in Lepidoptera.
The meso no tum: The anterior part of the mesonotum in Monotrysia and Ditrysia
is more or less vertically bent downward, as in Trichoptera and Zeugloptera. Hence,
the powerful dorsal longitudinal muscles are attached to this area. The development of
the first phragma is correspondingly small, and it may even be difficult to recognize.
Srivastava (1962), however, recognized in Papilio the antecostal suture and a narrow
first phragma anterior to it.
The prescutoscutal suture traversing the anterior part of the mesonotum commonly
occurs in Lepidoptera. It connects the base of the prealar arm (prescutal arm of
Sharplin) on both sides. The position of this suture agrees with that of the transverse
prescutoscutal cleft in Micropteryx (Fig. 144D).
The anterolateral scuta! suture demarcates the sura!are of Michener ( 1952) and
Ehrlich ( 1958b), which bears the anterior notal wing process. The suture is strongly
ridged internally, and reaches the base of the prescutal arm anteriorly, as is clearly seen
in Telea (Niiesch 1953), Anthocaris (Fig. 151A), Papilio (Srivastava 1962), and
probably many others. The scutum may have a median longitudinal suture, as seen in
Hepialus (Weber 1924a). The posterolateral scuta! suture is not well marked. The
scutoscutellar suture in higher Lepidoptera is medially strongly produced and the
narrow lateral scutellar area is continuous with the axillary cord.
In many Lepidoptera the mesothoracic postnotum is placed vertically or subvertically
beneath the posterior margin of the scutellum. Sharplin ( 1963b) called this area the
phragma, implying that actually this postnotum (acrotergite of Sharplin) functions as a
phragma for the attachment of dorsal longitudinal muscles. As in Micropteryx, the
detached laterophragma (lateral postnotum) extends anteriorly beneath the wing base; in
Danaus, according to Ehrlich (1958a), the anterior end of the laterophragma articulates
with the socket of a protuberance from the ventral edge of the scutum, immediately
behind the caudal end of the postmedian notal wing process ( adnotale of Ehrlich) .
Apparently, the relationship between the laterophragma and the posterior scuta! lateral
margin in Eacles (Michener 1952) is similar to that in Danaus. According to Sharplin
(1963b), the lateral part of the laterophragma extends forward in higher Ditrysia, and
may be fused with the underside of the postmedian notal wing process (median notal
wing process of Sharp lin) . Sharp lin further thought that this would give the dorsal
longitudinal muscles direct leverage on the first axillary sclerite of the forewing.
The mesonotal lateral margin: The greatly developed tegula is a characteristic
feature of Lepidoptera. Its vertex lies on the anterior costal margin of the forewing; its
smooth inner surface is, as Hering ( 1956) showed, provided with a cavity, through
which the tegula maintains connection with the wing base (Fig. 150B). The dorsal and
ventral edges of the tegula are often prolonged to form processes. The dorsal process is
attached to the mesonotum, and the ventral process lies on the underside of the forewing
base; such processes are, however, absent in some families. The tegula is supported by
a detached prescutal plate or subtegula (sclerite accessorio of Onesto 1959b), and by the
tergopleural apodeme (Fig. 151A), which are discussed later. Hering (1956) suggested
that the tegula, besides its presumed function as a sensory organ can substitute for the
axillary sclerite and its main function in Frenatae is to reinforce the wing articulation.
Jordan (1928) and Hering (1956) discussed the taxonomic value of the tegula in
Lepidoptera.
On the scuta! lateral margin of the mesonotum of many Lepidoptera the anterior
and antemedian notal processes together form a robust process, with which the narrowed
anterior part of the first axillary sclerite has a ligamentous connection (Figs. 151B,
152A). This robust process is followed by a rather deeply excavated membranous area,
or the tergal fissure. In Stigmella (Fig. 151B) and Hepialus (Sharplin 1963a) the tergal
346
PRESCUTUM
SURALARE
ANTERIOR
NOTAL PROCESS
POSTMEDIAN
NOTAL PROCESS
MEDIAN PROCESS ?
4'TH AXIL. SCL.-
A
SUBTEGULA
-------TEGULA
TERGOPLEURAL
APODEME
PROX. MED. PL.
2'ND AXIL. SCL.
3'RD AXIL. SCL.
l'ST AXIL. SCL.
:
TERGAL FISSURE \\ (.:.-.::_ ?S.
------r/': .. l'ST AXIL. SCL.
POSTMEDIAN
NOTAL
PROCESS
B
FIG. 151. A, forewing base in Anthocaris cardamines (from Onesto); B, connection of first
axillary sclerite with scuta! lateral margin in Stigme/la basalella (from Sharplin).
fissure is followed posteriorly by the median and postmedian notal processes, as in
Micropteryx (Fig. 146A). In some higher Lepidoptera (Fig. 152A), however, dis-
tinction between the median and postmedian processes is somewhat obliterated, due
obviously to enlargement of the postmedian process. In these forms the presumed area
of the postmedian notal process still maintains the ligamentous connection with the
proximal caudal end of the first axillary sclerite, just as in Micropteryx and Stigmella.
This enlarged area produced by the fusion of the two notal processes is the adnotale
of some workers; Sharplin ( 1963a) called this area the median notal process.
Irr Anthocaris (Fig. 151A), as Onesto (1959b) showed, the postmedian notal
process (processo vertice scutale of Onesto) extends forward, thereby almost closing the
tergal fissure. Furthermore, the base of this process lies where the posterior notal
process becomes detached to form the fourth axillary sclerite. In Papilio also the
presumed postmedian notal process (median notal process of Srivastava, 1962) shares
its base with the posterior notal process; and this fact led Srivastava to believe that the
postmedian notal process was a part of the posterior notal process. This peculiar
development of the postmedian notal processes in Anthocaris and Papilio represents a
highly specialized condition.
The forewing base structures: As Sharplin (1963a) showed, the first axillary
sclerite in the forewing of Lepidoptera is distally connected with the second axillary
sclerite, and typically has four ligamentous connections with the scuta! margin as seen
in Stigmel/a (Fig. 151B); its anterior arm (head) is always associated with the Sc vein.
Sharplin further showed that when the area produced by fusion of the median and post-
median notal processes (median notal process of Sharplin) is large in advanced forms,
the first axillary sclerite is also large. A thinly sclerotized part of the head of the first
axillary sclerite marks the, point of attachment of a tergopleural muscle (t-p 10).
The dorsal second axillary sclerite (Fig. 152C) in Lepidoptera is connected with
the R vein, and articulates with the first axillary sclerite and with the proximal median
plate (first median plate of Sharp lin) . The ventral second axillary sclerite is tied to
the subalare and to the pleural wing process by ligaments, and is also connected with the
proximal median plate either by a bending cuticle or by a ligament.
In Trichoptera and Lepidoptera except Hepialoidea, according to Sharplin ( 1963a),
connection of the dorsal and ventral second axillary sclerites occurs through invagination
of a flange from the base of the R vein. This flange or peg fits into a shallow socket
formed on the ventral second axillary sclerite (Fig. 142B).
The third axillary sclerite articulates proximally with the posterior notal wing
process and is associated distally with the vannal and jugal, areas of the wing. In
Lepidoptera, according to Sharplin (1963a), the third axillary sclerite is either articulated
or fused with the proximal end of the proximal median plate and it has no direct
articulation with the second axillary sclerite. In Anthocaris- (Fig. 152B), however,
Onesto (1959b) showed that the definitive, third axillary sclerite is directly connected
along a line with the second axillary sclerite. Most probably, what Onesto called the
third axillary sclerite in Anthocaris is anteriorly the proximal median plate that has
become indistinguishably fused with the third axillary sclerite.
The posterior distal margin of the third axillary sclerite in lower Lepidoptera lies
along the base of the jugum and is concerned with jugal folding. In higher Ditrysia,
according to Sharplin (1963a), however, the corresponding area is not concerned with
jugal folding; instead, it is hinged to the remaining part of the third axillary sclerite by
means of a bending cuticle, and it also articulates with the posterior notal wing process.
This distal part of the definitive third axillary sclerite may correspond to the area which
Onesto ( l959a) called the basanale in Blattaria (Fig. 55A). Sharplin (1963a), how-
ever, called this area the fourth axillary sclerite in error. Onesto ( l959b) found in
Anthocaris a tendinous connection between the first and third axillary sclerites (Fig.
151A), which is present also in other insects.
According to Sharplin (1963a), the proximal median plate (Fig. 152B, C)
penetrates ventrally from near its junction with the third axillary sclerite to the lower
surface of the wing, where it is joined to the second axillary sclerite.
The distal median plate (Fig. 152C) articulates with the distal end of the proximal
median plate and is fused with the M vein, when this vein is clearly recognized; this
condition is present in Micropteryx and H epialus and is primitive. The base of the
second Cu vein overlies the distal end of the distal median plate in Incurvarioidea and
348
R- PLATE
l'ST AXIL.
SCL.
2'ND AXIL
SCL.
PROXIMAL MEDIAN
PLATE
A
PROX. MED.PL.
8
--Cul
DISTAL MEDIAN
PLATE+ Cu 2
Flo. 152. A, connection of first axillary sclerite of forewing with scutal lateral margin in
Plusia gamma (from Sharplin); B, lateral view of forewing base in Anthocaris cardamines
(from Onesto); C, dorsal view of forewing base in Hepialus lupulinus (from Sharplin).
Stigmelloidea; the same vein is fused to the radial plate in all Ditrysia except the
Papilionoidea (Sharplin 1963a).
The folding mechanism of the forewing, including the wing base structures, was
succinctly expiained by Sharplin (1964a), and a further explanation is not attempted
here. Sharplin ( 1964b) also discussed the taxonomic value of the wing base structures.
The mesopleuron: The pleural ridge separating the episternum and the epimeron
is always externally distinct. The dorsal end of the pleural ridge, or the pleural wing
process, is strongly bent caudad in higher forms, evidently in association with the more
posterior position of the wing base. The pleural arm lies relatively more ventrad and
has a broad base. At least in higher forms, the ventral end of the pleural ridge does not
form a typical condyle for articulation with the coxa. In Papi/io (Weber 1928a), for
instance, the pleural ridge is ventrally directly continuous with the coxal ridge (Fig.
153A) ,n and in Zygaena (Weber 1928a) the ventral end is continuous with ridges
formed on the ventral margin of the episternum and epimeron. In Danaus (Ehrlich
1958a) also, the pleurocoxal connection is complex, a ridge from the pleural ridge being
continuous to the median longitudinal ridge or discrimen.
In Hepialus (Fig. 149A) and Phassus (Matsuda 1960a) the paracoxal suture is
complete on the episternum; in no species of Monotrysia and Ditrysia is the paracoxal
suture known to cross the pleural suture; the suture is often incomplete (Fig. 153B),
even on the episternum, or completely absent, as seen In many figures given by
Shepard (1930).
The anapleural cleft is nearly always present and it may be represented by a suture.
Typically, it extends to the pleural suture near the middle and meets the paracoxal suture,
when the latter is present. In the dorsal area of the episternum immediately anterior to
the pleural wing process, the basal are (second basal are of Sharplin, 1963a) is always
present. Sharplin also recognized another basalare (first basalare of Sharplin), which
is an incompletely detached anterior dorsal area of the anepisternum.
A conspicuous structure in the anepisternal area is what Sharplin ( 1963a) called
the "tergopleural apodeme," which runs underneath the basalare to reach the pleural
ridge (Fig. 151A). The origin of the tergopleural apodeme in Lepidoptera, including
Zeugloptera, can be traced back to a tergopleural muscle which Sharplin ( 1963a)
called the third tergopleural muscle ( t-p 4). In the meta thorax of Trichoptera and
metathorax of Lepidoptera the muscle occurs in the corresponding position (Fig. 148),
but no tergopleural apodeme is formed. In the mesothorax, according to Sharplin
(1963a), this apodeme is distinct from the subtegula in Micropterygoidea, Eriocranioidea,
and Hepialoidea, the two structures being connected by a muscle ( t-p 4) ; they touch one
another in Incurvarioidea and are fused in Ditrysia. Thus, the gradual replacement of
t-p 4 by the tergopleural apodeme is evident.
Apparently in most Lepidoptera the preepisternum extends ventrally, as far as the
median longitudinal invagination of the sternum, as seen in Hepialus (Fig. 149A). The
pleurosternal suture has been recognized in H erse (Rothschild and Jordan 1902) , in
Choristoneura (Freeman 1947) and in Phassus (Matsuda 1960a), in which the basi-
sternum is exposed.
The preepisternum has a tendency to form the marginal prepectus (hypopteron), as
in some other orders. In lower Lepidoptera, such as Hepialus (Fig. 149A) and
Phassus, this secondary sclerotization is absent; it is present in Papilio (Fig. 153A) and
in many other Lepidoptera. Madden (1944), Michener (1952), Ehrlich (1958a), and
Srivastava (1962) have called this secondary area the preepisternum. In the species
studied by these authors the true paracoxal suture is absent, and they have mistaken
the anterior margin of the preepisternum that demarcates the prepectus for the paracoxal
suture. This mistake led them to interpret the prepectus as the preepisternum. In
7In Trichoptera such a fusion with the coxa apparently does not occur. Instead, the basal part of the coxa,
according to Crampton (1920b), is separated by a narrow membrane to form the basicoxite of Crampton.
Crampton thought that this feature was of great diagnostic value for determining whether an insect is
lepidopterous or trichopterous.
350
PLEURAL RIDGE
HYPOPTERON
(PREPECTUS)
MARGINAL
RIDGE OF COXA
EPISTERNUM
PARACOXAL SUTURE
MEMOmS OF THE ENTOMOLOGICAL SOCIETY OF CANADA
PLEURAL RIDGE
A

EPIMERON

PLEURAL RIDGE
B
FIG. 153. A, inner view of pterothorax in Papilio machaon, sterna removed (from Weber);
B, lateral view of pterothorax in Catocala concumbens (from Shepard).
these species what they called the katepisternum, or the sternopleurite (Ehrlich 19 5 Sa),
is the fused katepisternum and preepisternum.
The katepisternum, when it is well defined anteriorly by the paracoxal suture (Fig.
149A), is a relatively narrow sclerite above the coxal margin, and it has a well formed
trochantin in lower forms (e.g., Hepialus, Phassus); the trochantin is absent in most
Lepidoptera. The ventral condyle for articulation with the coxa is, as Weber ( 1928a)
and Matsuda ( 1960a) have shown, continuous from the furcal base.
The dorsal margin of the epimeron is more or less excavated by the membranous
area above, and the large subalare is always present in this membranous area. A con-
spicuous modification of the mesepimeron in Lepidoptera is the formation of the pre-
epimeron in some forms (Fig. 153B). The anterior part of the epimeron is often
included in the bend of the pleural ridge and becomes separated by a furrow or suture
from the main part of the epimeron.
The mesosternum: In Herse (Rothschild and Jordan 1902), Choristoneura
(Freeman 1947), and Phassus (Matsuda 1960a), as already noted, the mesosternum is
not completely invaginated and the pleurosternal suture is recognizable externally. In
351
Papilio, as Weber ( 1928a) showed, the invaginated sternal ridge becomes increasingly
higher posteriorly and is continuous with the furcasternum, which bears paired furcae.
As :in Mitropteryx and Trichoptera, the furcasternum bears two pairs of furcal
arms, of which the lateral pair is fused with the posterior margin of the epimeron.
Weber ( 1928a) thought that in Lepidoptera this union of the furcal arm (lateral furcal
arm) with the epimeron replaced a pleurosternal muscle p-s 1, which connects the furcal
arm and the pleural arm. Chadwick (1959a) suggested that such a union took place
as a result of sclerotization that has advanced along the course of furcoepimeral or furco-
postcoxal muscles or ligaments. If Chadwick's suggestion is valid, the muscle in question
is probably p-s 5, which connects the interpleurite and the furca of the preceding segment.
It should be pointed out, however, that in Trichoptera the line of union is extensive and
internally strongly ridged. Such an extensive line of union is somewhat unlikely to
represent the former course of a muscle, since p-s 5 is a relatively weak muscle.
The meso thoracic spina in Phassus is, as Matsuda ( 1960a) showed, attached to the
posterior wall of the invaginated furcasternum. It is apparently absent in a great
majority of Lepidoptera.
The metanotum: As Sharplin's ( 1963b) figures show, the anterior part of the
metanotum has undergone varying degrees of membranization in various groups of
Lepidoptera. As in Trichoptera and Zuegloptera, a sclerotized posterior wall of the
second phragma is absent.
According to Sharplin (1963b), the prealar arm (prescutal arm) is retained only in
more primitive Monotrysia; in Hepialoidea muscle t-p 4 is inserted on the proximal end
of the prealar arm, although in Zeugloptera and Eriocranioidea t-p 4 is inserted on a
separate prescutal apodeme, which is probably serially homologous with the subtegula in
the mesothorax. The prescutal apodeme remains attached to the prescutum in Stig-
melloidea and Incurvarioidea. In primitive Ditrysia the apodeme is detached but has a
ligamentous connection with the prescutum. This ligament is lost in higher Ditrysia; in
these forms the prescutal apodeme is attached to the laterophragma (lateropostnotum)
of the mesonotum by a short and thick ligament.
The reduction in size of the metanotum is mainly due to reduction of the scutum,
the scutellum having been relatively well pr.eserved. The acutoscutellar suture is
medially produced in lower forms, but it often becomes flattened or nearly horizontal in
higher forms. The posterolateral scuta! suture, when present, demarcates the posterior
notal wing process, as seen in Hepialus (Weber 1924a). In Anthocaris Onesto ( 1959b)
showed the anterolateral scuta! suture that separates the suralare.
As in Trichoptera and Zeugloptera, the metapostnotum is displaced posteriorly and
is united with the first abdominal tergite; since Weber (1924a) nearly all authors have
followed this interpretation. The lateral postnotum (phragma) has the usual connection
with the epimeron; in some moths this lateral connection is involved with sound reception.
Heitmann (1935) showed various modes of connection of the lateral postnotum with the
epimeron in various moths.
The metascutal lateral margin and the hind wing base structures: Generally the
median and postmedian notal processes are not conspicuous. The posterior notal process
is well developed, and it may be detached or partially detached from the meta-
notum proper.
The first axillary sclerite has three points of ligamentous connection with the scuta!
margin; the hindmost one is connected with the postmedian notal process; the connection
with the median notal process is consistently absent in the figures given by Sharplin
(1963b).
The second axillary sclerite is, according to Sharplin ( 1963b), a single, solid sclerite
joining the upper and lower wing membranes. Dorsally, its proximal edge has a broad
articulation with the first axillary sclerite; anterodistally it is connected with the radial
bridge. The dorsal articulation between the proximal median plate and the second
axillary sclerite has been lost in the hind wing of most Lepidoptera, although it is present
in Micropteryx (Fig. 154A) and Eriocranioidea. Ventrally the second axillary sclerite
is tied to the subalare and to the pleural wing process by ligaments.
In Micropteryx (Fig. 154A) the base of theM vein appears to be fused with the base
of the Cu 1 vein, which runs into the wing base just posterior to the R vein. The
proximal end of the M + Cu 1 vein forms a loop of bending cuticle, which rejoins R at
its junction with the radial bridge (plate); the base of Cu 2 and the distal end of the
distal median plate are fused to this loop. Note here the primary association of the M
and Cu veins with the distal median plate, which is clearly present.
In other Lepidoptera, according to Sharplin ( 1963b), the base of the M vein is
detached to form the median arm (Fig. 154B); the arm projects from the base of the
radial bridge (plate) toward the Cu 2 vein and the median plates. Here the primary
connection of the M vein and the Cu 1 vein with the distal median plate is obliterated.
In higher Lepidoptera the median plates are reduced to a single sclerite and are closely
associated with the third axillary sclerite. In Incurvarioidea and Ditrysia the median
arm is expanded to reach the base of the enlarged Cu 2. These two plates together,
spanning the gap between the third axillary sclerite and the R vein, take over the function
of the median plates which are highly reduced (Fig. 154B).
The third axillary sclerite in Monotrysia is present only dorsally and articulates
proximally with the posterior notal wing process, and anterodistally with the proximal
median plate. In Ditrysia the third axillary sclerite appears on the ventral wing
membrane near its articulation with the posterior notal wing process. An anal plate is
present in Zeugloptera (Fig. 154A), Monotrysia, and primitive Ditrysia.
The metapleurosternum: As in the mesothorax, the ventral end of the pleural ridge
does not form the ordinary condyle for articulation with the coxa. In Papilio (Fig.
153A), for instance, Weber (1928a) showed that the internal ridge extends along the
ventral edge of the episternum, and is continuous with the similarly ridged anterior margin
of the episternum.
Judging from the figure given by Shepard ( 1930), the ana pleural cleft is present in
Phassus; it is probably absent or represented by an incomplete suture in the metathorax
of a great majority of Lepidoptera. An incomplete paracoxal suture is apparently
present in some Lepidoptera that Shepard (1930) figured, although it has been lost in
many species. The apically poorly differentiated trochantin may be present in some
lower Lepidoptera. In many forms the prepectus (hypopteron) is formed along the
anterior margin of the episternum.
As is clear from Shepard's (1930) figures of many species, the anterior dorsal part
of the episternum is demarcated by a transverse suture that may or may not reach the
dorsal part of the pleural suture. Shepard called it the basalar pad, and in many cases it
corresponds to the first basalar sclerite of Sharplin ( 1963b). In Eriocranioidea the
posterior basalare immediately anterior to the pleural wing process (the second basalare
of Sharplin) is, according to Sharplin ( 1963b), joined to the prealar arm (prescutal arm
of Sharp lin) by a ligament; in Hepialoidea the prealar arm is bent and the distal half is
fused to the first and second basalaria of Sharplin. A remarkable fact Sharplin
(1963b) found was that in Acossus the apparent second basalare is a fusion product of
the basalare and the detached prealar arm. Sharplin inferred further that the definitive
second basalare in Ditrysia, in which the prealar arm is absent, might also be formed by
fusion of the detached prealar arm with the basalare. The tergopleural arm is not
formed in the metapleuron, the apodeme being represented by a muscle, t-p 4.
The dorsal margin of the epimeron is engulfed by a broad membranous area, in
which lies the subalare. This membranous area is the tympanal membrane in some
moths." The posterior dorsal end of the epimeron is joined to the postnotum.
In the metasternum the lateral furcal arm extends posteriorly and is joined to the
postcoxal bridge or to the ridged posterior margin of the epimeron. The ventral condyle
for coxal articulation is, as in the mesothorax, continuous from the furcal base.
'"The thoracic organ for sound reception has been formed through modifications of the epimeron, the post-
notum, and the first abdominal segment. For more information regarding this organ see Forbes (1916)
Eggers (1919), Richards (1932), Heitmann (1935), Kiriakoff (1952, 1963), and Treat '(1959). '
HUMERAL PLATE
DISTAL
MED. PL.
POSTERIOR
NOTAL PROCESS
ANAL PLATE
RADIAL PLATE
2'ND AXIL.
SCL
MATSUDA: TifE INSECT TIJ:ORAX
BASISUBCOSTALE
A
BASISUBCOSTALE
353
3'RD AXlL.
SCL.
MEDIAN
PLATE
FIG. 154. A, dorsal view of forewing base in Micropteryx calthella (from Sharplin); B, dorsal
view of hind wing base in Tortrix viridana (from Sharplin).
The adult musculature: The lepidopterous thoracic musculature was studied by
Berlese ( 1909b) in Sphinx canvoluvuli, by Weber ( 1928a) on some' dorsoventral and
ventral muscles in Papilio machaon, by Maki (1938) in Plutella maculipennis,
Adoxophyes privatana, Papilio thaiwanus, Milionia zonea and Amata lucerna, by Nliesch
(1953) in Telea polyphemus, by Hannemann (1956) on the cervical muscles in
Micropteryx calthel/a, by Treat (1959) on metathoracic muscles in Crymodes devastator,
by Srivastava (1961, 1962) in Papilio demoleus, by Ehrlich and Davidson (1961) in
Danaus plexippus, by Ehrlich and Ehrlich ( 1963) in various butterflies, by Sharp lin
(1963a, b) and Mickoleit (1966) on some tergopleural muscles in various Lepidoptera,
by Dierl (1964) in Fumea casta, and by Kristensen (1968) on the cervical muscles in
Eriocrania haworthi. Denis ( 1965a) made comments on the study of Danaus by
Ehrlich and Davidson ( 1961). Of these studies, the muscles in Papilio (Srivastava
1961, 1962), the winged male of Fumea (Dierl 1964), Danaus (Ehrlich and Davidson
1961), and Telea (Niiesch 1953) are homologized in Table XXI. Sternocoxal and
pleurocoxal muscles are studied only for Fumea and Telea because of the clarity in the
given figures and descriptions.
The thoracic musculature in Lepidoptera is generally similar to that of Trichoptera,
and no obvious tendency for the reduction in number of muscles is evident. s-cx(cv) 1
and cv-cv 1 are unique muscles that have been found only in this order.
Besides the muscles listed in Table XXI, other muscles found in other Lepidoptera
include 13 of Hannemann (1956) in Micropteryx and 2 of Kristensen (1968) in
Eriocrania, which are op-p 1; prothoracic p-cx 9 and p-tr 1 in five species Maki (1938)
studied; 8 of Kristensen ( 1968) in Eriocrania, which connects the anterodorsal corner
of the propleuron and the anterior part of the lateral cervical sclerite.
The larval musculature: The larval mucles of Lepidoptera were first described and
illustrated by Lyonet ( 1760), who worked on Cossus cossus. More re.cent studies
include the work on Malacosoma americanum and others by Forbes (1914), the work on
Lasiocampa quercus and Catocala spp. by Barth (1937, 1948), and the work on Mala-
cosoma americanum by Snodgrass (1961). In none of these works, however, was the
points of attachment of muscles adequately described. An exact comparison of larval
muscles in Lepidoptera with those in Trichoptera (Tindall 1963, 1964) is, therefore,
not attempted here. It can be said briefly, however, that the larval Lepidoptera are also
provided with numerous muscles as in Trichoptera, and that their musculature is
generally similar to that in Trichoptera. It is also obvious that many of these larval
muscles are caenogenetic, their occurrence being confined purely to the larval stage. In
Ephestia kuhniella Blaustein ( 1935) traced the histolysis of a mesothoracic muscle which
helps evagination of the wing.
The postembryonic development of indirect flight muscles: Blaustein (1935) des-
cribed histogenesis of an intersegmental dorsoventral muscle" in Ephestia as follows:
Imaginal cells or myoblasts in the embryo lie where the imaginal muscle is to be. By
mitotic division they form a syncytium on the first day after pupation, then the syncytium
grows. About 90 hours after pupation, the nuclei scattered in the plasma start moving
to the periphery and arrange themselves on the margin of each syncytium strand; this
marks the beginning of the histological differentiation to produce a muscle bundle. In
the next 2 days the syncytium attains its final size, and then the sarcolemma is formed.
The development of the muscle is completed by the end of the twelfth day after pupation.
In A ntheraea, according to Niiesch ( 1965), the dorsal longitudinal muscle ( t 14)
is formed from the myoblasts, which arise from decomposed larval muscles.
THE HYMENOPTERA
The neck region and the propleuron: The neck region in Hymenoptera is charac-
terized by the absence of a well-isolated lateral cervical sclerite. Instead, the anterior end
of the definitive proepisternum directly articulates with the posterior end of the head.
76Blaustein_ thought it to be an indirect flight muscle. but it cannot be identified. It may perhaps be t 12.
355
TABLE XXI
Thoracic musculature in Lepidoptera
Papilio Dana us
(Srivastava Fumea (Ehrlich and Telea Muscle
1961, 1962) (Dierl 1964) Davidson 1961) (Niiesch 1953) designations Remarks
35 ? d! 1 ? cv(d)-t 1 1
28 28 dS op-t 1
29 29 d 2 op-t 2
30, 31 Lb dim 1 30, 32 d 1?, d 3 op-t 3
Lb ism 3 d4 op-cv 1 2
18 op-cv 2 2
26 19, 20 Op---{;V 3
32 Lb ism 2 27 d6 op-s 2
21 ?, 22? t ? t-s(cv) 1 3
cd t-s(cv) 9
34 Lb ism 1? t---{;V 2 4
35 Lb vim 2 25 cv cv-s 4
36 I vim 3 26 cv-s 1
so Ibm 2 I st a cv-cx 3 5
27 cv-cv 1 6
49 Lb ism 4 23 s---{;x(cv) 1
33 Lb vim 1 24 v s 1 or 2
41 33, 34 Id! 2 t 12
70 II dim 2 61 ? II dl 2
99 III dim 2 92 III d! 2
38?' 39, 40 I dim 1 31 I dl 1 t 13
64 49 II d! 3
III dl 3
65-67 II dim 1 S0-54 II d! la-e t 14
100 III dim 1 93 III dl Ia, b
71 II pm 3 68 II pd 1 t-p 3 7
III pm 3
77 II pm 4 67 II pd 4 t-p 4 8
107 112 II p 2
69 II dvm 1-3 56-58, II dv Ia, b t-p 5 or 6
62, 63
98 III dvm 1, 2 88,94 III dv 1
II pm 5 II pd 5 t-p 7 9
II pd 3 t-p 10 10
103 102 lli pd 3
75 II pm 6 78 II pd 2a, b t-p 13
104 III pm 4 110 III pd 2b
74 II pm 7 82 II pd 2c t-p 14
105 III pm 5 111 Ill pd 2a, c
76 (II) 81
III p 1
t-p 16
42, 43 (I) I dvm 1 36, 40 I pd a-c t-p (proth.)
44 (I-II) I ism 39 I dv t-s 1 11
79 89, 95 II dv 2 t-ti(cx) 2
109 III dv 2 or/and 3
45? 37 t-cx 7 12
83 II dvm 5 60, 65 II dv 4, 5
113 III dvm 4 91 III dv 4, 5
TABLE XXI (Continued)
Dana us
Fumea (Ehrlich and Telea Muscle
Papilio
(Srivastava
1961, 1962) (Dierl 1964) Davidson 1961) (Niiesch 1953) designations Remarks
82
112
101 (III)
72
73
102
106 (II-III)
80
110
68
97
37
87b, c
116b, c
87a?
116a?
55a
87d
116d
II pm 8 79, 80
III pm 6 97
II pm 1
III pm 1
69 (II)
99
II zm 84
III zm
III vim 2 (III -Ia)
I pm 1
II vim 3
I vim 1
II vim 1, 2
I vim 2
III vim 1
Ibm 3
Ibm 1 ?
II bm 1
Ill bm 1
II dvm 4
III dvm 3
II pm 2
III pm 2
Ibm 4
II bm 2
IVbm 2
98
38
70
100
83
109
47
85
46
114
103?
59, 64
90, 96
71
101
44
77
87, 107
REMARKS (Table XXI)
(1) These muscles may represent t 13.
II pv 4, 5
III pv 4, 5
III pv 1
II pv 1
II pv 7
III pv 7
II pv 8
II pv 2
III pv 2
I pv 6
II pv 6
III pv 6
I pv a
II vi 2
I v\1
II vi 1
I vi 2
II I vi 1
I st b
II st 3
III st 3
I st c
I st 1
II st 1
II I st 1
II st 4 ?
II dv 3
III dv 3
II pv 3
III pv 3
. II st 2
I II st 2
(2) All muscles listed may be homologous.
t-cx 8
p2
p3
p-s 1
p-s 3
p-ti(cx) 2
or 3
p-ti(cx) 4
p-ti(cx) 5
p-cx9
s 12
s 13
s 14
s 20
s 16?
p-s 5?
s-ex 3
s-ex 4
s-ex 5
s-ex 6
t-tr 1
p-tr 2
s-tr 1
13
14
(3) tin Telea arises from the anterior edge of the episternum and is inserted on
the tentorial bridge; 21 and 22 in Danaus arise from the dorsal part of the dorsal arm of
the lateral cervical sclerite. These muscles may be homologous with 8 of Kristensen
( 1968) in Eriocrania (p. 354), and they may deserve a distinct muscle designation.
( 4) Lb ism 1 arises posteriorly from the mesoprephragma.
(5) These muscles arise anteriorly from the definitive presternal area (ventral
cervical sclerite) .
357
( 6) 27 in Papilio connects the posterior ventral cervical sclerite and the posterior
ventral surface of the lateral cervical sclerite.
(7) These muscles are 1 t-p Mu. of Sharplin (1963a).
(8) These muscles are 3 t-p Mu. of Sharplin (1963a); varying degrees of the
apodemal formation from this muscle occur in various Lepidoptera.
(9) These muscles are 2 t-p Mu. of Sharplin (1963a).
(1 0) According to Mickoleit (1966), t-p 10 occurs in about one-half of the
lepidopterous species he studied.
( 11) I ism in Fumea is attached to the prothoracic epimeron.
( 12) The exact position of 45 in Papilio is not clear. In Papilio thaiwanus (Maki
1938) the corresponding muscle appears to represent t-cx 7. 60 and 65, dv 4 and dv 5
are regarded as representing single muscles that have become secondarily split.
(13) 103 in Danaus connects the metafurca (lateral) and the primary (?)
metafurcal arm. This muscle may be p-s 5.
(14) 87 a in Papilio arises from the subalare.
Clearly the lateral cervical sclerite in other holometabolous orders is contained in the
definitive proepisternum in Hymenoptera. The question is, then, whether the inclusion
of the lateral cervical sclerite in the proepisternum is due to fusion of the former with the
proepisternum, or if the lateral cervical sclerite is simply not differentiated from the rest
of the proepisternum.
In the prothorax of Schizocerus (Fig. 155B), an oblique ridge reaching the ventral
end of the pleural ridge is most probably serially homologous with the anapleural suture"'
in the pterothorax (Fig. 157E), which occupies the same relative position. The area
anterior to the anapleural suture in the prothorax of Schizocerus, therefore, consists of
the preepisternum and the katepisternum, and it is the anterior end of the preepisternal
area that articulates with the head, just as the lateral cervical sclerite does in other holo-
metabolous orders.
In the larvae of sawflies, as Crampton ( 1926a) showed (especially in his fig. 102),
the katapleuron (including the katepisternum) and the anapleuron (including the pre-
episternum) are separated by the paracoxal cleft. In the adult the paracoxal cleft
becomes lost, and, at the same time, the katepisternum becomes indistinguishably fused
with the preepisternum; and the anapleural suture appears. Obviously, then, the super-
ficially undifferentiated lateral cervical sclerite in the adult Hymenoptera is due to fusion,
as is clear in the postembryonic development of sawflies. In the absence of the isolated
lateral cervical sclerite in the adult, Hymenoptera resemble Lepismachilis.
Crampton (1909, 1926a) and Martin (1916) called the ventrolateral area of the
propleuron the cervicopleuron, implying that the lateral cervical sclerite has become
fused with the proepisternum. This idea of the origin of the cervicopleuron was accepted
by Weber (1927a), Duncan (1939), and Arora (1953). The facts in the sawflies dis-
cussed above support such an origin.
The interpretation of Snodgrass (1910), Weber (1925), and Alam (1951) that the
preepisternal area (the lateral cervical sclerite) is the proepisternum is also correct, but
these authors did not refer to the development (fusion) of the area in question.
Snodgrass (1942) thought that the anterior, constricted part of the definitive pro-
episternum in Apis was truly cervical in origin. Similarly, Richards (1956) thought
that the anterior part of the cervicopleuron in many sawflies studied by Ross (1937) was
the true cervical sclerite: However, this anterior area appears to be purely a secondary
constriction adapted for articulation with the occipital condyle from the posterior margin
of the head. Such a constriction of the anterior end of the cervicopleuron occurs in
many Hymenoptera, including Vespula (Duncan 1939), Apis (Snodgrass 1942), and
"The anapleural suture occurs also in the propleuron of some other sawflies, such as Arge (Snodgrass 1910)
and Dolerus (Crampton 1926a), although these authors interpreted the anapleural suture as the pleural suture
(Snodgrass), or as the line of union of the lateral cervical sclerite with the proepistemum (Crampton).
PRONOTUM
AN EPISTERNUM 2 ----+\-
A
OCCIPITAL
PROCESS
SPIRACULAR LOBE---.
c
M&"SO
-----METANOTUM
FIG. 155. A, connection of pronotum and mesothoracic episternum in Pamphi!ius luteicornis
(original); B, lateral view of cervicopleuron in Schizocerus sp. (from Weber); C, anterior
view of pronotum in Apis mellifera (from Snodgrass).
Stenobracon (Alam 1951). In Lestodryinus, according to Subba Rao (1953), it is the
internal process from the anterior constriction (cervical apodeme of Snodgrass, or the
posterior ramus of the occipital process of Duncan) that articulates with the occipital
condyle.
In higher Hymenoptera the anapleural suture has become lost, and the proximal
development of the ventral (proximal) margin of the cervicopleuron is extensive.
359
The pleural ridge lies lateral to the cervicopleuron (preepisternum). As Weber
(1927a) <lemonstrated, the thick pleural ridge in Schizocerus (Fig. 155B) and in other
tenthredinids -gives rise to the pleural arm; the arm in turn lies close to the profurcal arm
in Schizocerus. In Dolerus (Crampton 1926a) also, the anapleural suture and the
pleural suture occur at the same time.
Snodgrass' misidentification of the anapleural suture as the pleural suture in A rge,
which was already discussed (foot note, p. 357), led him to misinterpret the area posterior
to the anapleural suture as the epimeron. According to Snodgrass, the process borne on
this area is an epimeral process, not the ordinary pleural process or arm. In Apis
Snodgrass ( 1942) made the same interpretation that the anterior, weaker suture is the
pleural suture. As is clear in Fig. 156E, it is the posterior, thick ridge that is the pleural
suture; and the epimeron is lost in Apis.
Duncan (1939) noticed the absence of the proepimeron in Vespula, and he thought
that when the epimeron is so reduced, the epimeral arm of Snodgrass would appear to
arise from the episternum and would become virtually equivalent to the pleural arm.
As is clear from the preceding discussion, this reservation of Duncan is unnecessary.
At least in most Hymenoptera the proepimeron is absent.
The trochantin has been described in Tremex (Snodgrass 1910), in Schizocerus
(Weber 1927a), and in Urocerus (Richards 1956). In more specialized sawflies and in
higher Hymenoptera, the trochantin is either reduced to a minute vestige, or more often
it has disappeared.
The pronotum: In Hymenoptera the pronotum is dissociated, in varying degree,
from the rest of the prothorax. Jn Pamphilius (Fig. 155A) the ventral extension of the
pronotum is not at all pronounced and its posterior margin only loosely covers the
anterior margin of the mesonotum; its posterolateral angle is connected by a narrow
bridge with the anterior margin of the mesothoracic anepisternum, just as in Mecoptera.
In Ichneumonidae and Pompilidae (Weber 1925) the pronotum is undivided, but its
downward shift is pronounced. In Vespa (Weber 1925) the pronotum is divided into
the median unpaired part, or the collar, and the paired triangular lateral lobes. The
same division occurs in Vespula (Duncan 1939) and Stenobracon (Alam 1951). In
Apis (Figs. 155C, 156A) the pronotallateral lobes meet ventrally to form a ring, and
the cervicopleuron lies anterior to this pronotal ring. In this process of dissociation of
the pronotum from the rest of the prothorax, the pronotum has become displaced
posteriorly. However, at least in most Hymenoptera, fusion of the pronotum with the
metathorax has not occurred.
The prosternum: In many sawflies the anterior end of the presternum is free from
the posterior margin of the cervicopleuron. In higher forms, however, the cervico-
pleuron on the two sides may become continuous, and the presternum may even be partly
overgrown by the cervicopleuron (e.g., Vespula, Duncan 1939). An external division
of the presternum into the basisternum and the furcasternum by the sternacosta is absent.
In higher Hymenoptera the ingrowth of the furcal arms assumes highly elaborate shapes
(Fig. 156C).
In sawflies (Weber 1927a; Ross 1937), the prothoracic spina lies widely separated
from the furcal bases; it is present also in Astymachus (Compere 1962). Richards
( 1956) thought that the anterior discriminal pit of Michener (1944) in higher Hymenop-
tera was an indication of a vestigial spina. The fact that the prothoracic spina is present
in the larva of A pis (Daly 1964) suggests that in many Hymenoptera the spina
degenerates in later stages of development.
The mesonotum: In Hymenoptera the anterior part of the enlarged mesonotum lies
beneath the pronotum. In Pamphilius (Fig. 157A) the unpaired first phragma extends
from beneath the middle of the prescutoscutal suture, and the acrotergite is not dif-
ferentiated. In other Hymenoptera studied (Weber 1925; Snodgrass 1942; Alam 1951)
the first phragma is likewise a simple inflection from the anterior margin of the
mesonotum.
360 MEMOIRS OF TIIE ENTOMOLOGICAL. SOCIETY OF CANADA
OCCIPITAL PROCESS
B
TROCHANTIN----'
CERVICAL
APODEME
A
c
NEURAL
FORAMEN
PLEURAL ARM?
PLEURAL RIDGE
.... LATERAL
CERVICAL
SCLERITE
(PREEPISTERNUM)
FIG. 156. A, venral view of thorax in Apis mel/ifera (from Snodgrass); B, anterior end of
lateral cervical sclerite in A pis mellifera (from Snodgrass); C, dorsal view of inner sternal
skeleton in Apis mellifera (from Snodgrass); D, lateral view of cervicopleuron in Arge sp.
(from Snodgrass); E, lateral view of cervicopleuron in Apis melli/era (from Snodgrass).
As in Diptera, the prescutoscutal suture is a persistent suture in Hymenoptera, and
d e l i m i t ~ the narrow prescutal area along the median and the anterolateral parts of the
mesonotum. Because of the close association of the pronotum with the mesothorax, the
prealar arm is not formed in the mesothorax of Hymenoptera. In Pamphilius (Fig.
157A) the parapsidal suture arises from the middle of the posterolaterally extended
prescutoscutal suture, and the sutures on the two sides become confluent medially. As
Snodgrass (1910), Weber (1925), Reeks (1937), Ross (1937), Arora (1953), Rivard
(1955), Bracken (1961), Tait (1962), and Wong (1963) have shown, the parapsidal
sutures meet the median longitudinal suture in many sawflies; they are clearly present also
in a protohymenopterous genus Sycopteron, which Tillyard (1927) studied. What
Tulloch (1929) called notauli in Xiphydria, and perhaps in Ampu/ex, Myopone, and
Paraponera, are also probably the parapsidal sutures. The median area bounded by the
parapsidal sutures has often been called erroneously the pre scutum (see p. 13).
The anterolateral scuta! suture in Pamphilius (Fig. 157A) is externally indistinct,
although it is ridged internally. In some Hymenoptera the suralare demarcated by the
anterolateral scuta! suture extends posteriorly as far as the transscutal suture; anteriorly,
the suture meets the prescutoscutal suture. In Stenobracon, as Alam (1951) showed,
the suralare and the prescutum are continuous, although Alam did not recognize the
fusion of the two areas. He called the whole anterolateral marginal strip anterior to the
transscutal suture the prescutum. In Vespula this combined prescutoscutal area is
marginally ridged and is the apodemal fold of Duncan (1939) in Vespula; such a
marginal thickening occurs also in ants. In ants Tulloch ( 1935) called the dorsally
extended part of the ridge above the tegula (which probably represents the anterolateral
scuta! suture) the parascutal ridge, and named the somewhat expanded suralar area
delimited by this ridge the parascutum.
The scutoscutellar suture occurs commonly in sawflies (Fig. 157A), and the
scutellum is laterally continuous with the axillary cord. In higher Hymenoptera the
scutoscutellar suture meets the transscutal suture, as discussed below.
In Chalastogastra the transscutal suture is, when present, only weakly developed.
In many higher families the transscutal suture connects the tergal fissures (Fig. 158D),
and it perfectly agrees in position with the transscutal suture inDiptera and Heteroptera,
although it is not homologous with those in Diptera and Heteroptera.
In many Hymenoptera the transscutal suture lies clearly anterior to the scutoscutellar
suture. In A pis (Fig. 158D) and some other higher Hymenoptera, however, the postero-
lateral triangular part of the scutum is separated by the transscutal suture that meets the
scutoscutellar suture mesally; this area is called the axilla in taxonomy.
In the scuta! area another common suture is the median longitudinal suture. In
A pis Daly ( 1964) found that the median line is actually composed of two lines forming
a furrow. In the pharate pupa of Apis Daly found that the anterior edge of the meso-
scutum was deeply indented and had a shallow median furrow leading caudad toward
the transscutal suture. This indentation is the first phragma, and the furrow marks the
progressive caudal migration of the anterior attachment of muscle 1ph-2ph (t 14).
Daly called the furrow the anteroadmedian lines, as distinct from the median line
that is present at the very anterior edge of the mesonotum and poorly defined in A pis.
According to Daly ( 1964), the anteroadmedian lines are equivalent to the median
mesoscutal suture of Michener ( 1944), which was shown to be a single line. It is
probable that many sutures described as a median longitudinal line or suture may be of
the same nature as the anteroadmedian lines. Daly further says that the admedian lines
in many other Hymenoptera are separated and lie lateral to the true median line.
In Pamphilius (Fig. 157A) the posterolateral scuta! suture extends anteriorly to the
base of the postmedian notal process, as in other insects. The same suture is apparently
present also in other sawflies. In many higher Hymenoptera, in which the transscutal
suture is present, this posterolateral scuta! suture is apparently absent.
In Clistogastra another frequent suture is what is called the parapsidal furrow, which
extends anteriorly from the lateral part of the transscutal suture; it lies lateral to the
parapsidal suture (Tulloch 1929), and according to Reid (1941) the furrow nearly
always commences at, or close to, the point where the transscutal suture crosses the
parascutaJ ridge (anterolateral scuta] suture). The parapsidal suture (notauli) and the
parapsidal furrow may be mistaken for one another when either one of them is present
and the anterior (parapsidal suture) or posterior (parapsidal furrow) end is lost.
In Cimbex (Snodgrass 1942) and Pamphilius (Fig. 157A) the postnotum is well
retained, and the lateral postnotum meets the epimeron as in other insects; the second
phragma is continuous from the postnotum. In Clistogastra the postnotum is often so
greatly reduced medially that its connection with the scutellum is often restricted to its
lateral sides (Fig. 157D), and the second phragma is a well-developed lamella continuous
from the lateral postnotum.
Among sawflies, the lateral postnotum in Pamphilius (Fig. 157A) is modified into
the axillary lever, upon which the intersegmental tergofurcal muscle (t-s 1) is attached;
its lateral union with the epimeron clearly suggests that the lever is indeed a modified
lateral postnotum. In Sirex (Fig. 159A) the homologous structure is a mere process or
lobe of the lateral part of the second phragma; it is also laterally united with the epimeron.
In A pis (Snodgrass 1942), among higher Hymenoptera, a modified postnatal plate
or the axillary lever (Fig. 159C) is an elongate triangular sclerite lying against the mesal
surface of the stalk of the second phragma, upon which muscle t-s 1 is attached, and the
structure assists in wing flexion. Snodgrass (1942) showed a similar lever in Vespula
and in Sphecius (Fig. 159B); its union with the supporting sclerite of the subalare, which
must be the epimeral plate, still suggests that the lever is a modified lateral postnotum.
In Stenobracon (Alam 1951) the corresponding structure is a small projection, similar
to that in M egarhyssa shown by Snodgrass ( 1942).
The mesoscutal lateral margin: In Pamphilius (Fig. 157A) the mesoscutal lateral
margin behind the large tegula is the suralare, which is represented largely by the anterior
notal wing process. The antemedian notal process projects caudad, extending beyond
the level of the median notal process; the median notal process is an inconspicuous
process separated anteriorly by the narrow tergal fissure; the postmedian notal process
projects cephalad and has connection with the proximal caudal area of the first axillary
sclerite; the posterior notal process is represented by the detached fourth axillary sclerite.
The preceding description applies also to a species of Arge examined.
In Clistogastra, in which the transscutal suture nearly always occurs, the scuta!
lateral margin is usually strongly ridged and more or less bent downwards. Differentia-
tion of the notal processes is less marked, and the shorter distances among themselves are
obviously correlated with the shortened wing-base. The posterior notal process is
represented by the fourth axillary sclerite, as in sawflies. In A pis what Pringle ( 1957a)
called the scutellar lever is the postmedian notal process.
The forewing base structures: In the forewing base of Pamphilius (Fig. 157B) the
humeral plate (basicostale) consists of two plates. The connection of the anterior end
of the first axillary sclerite with the base of the Sc vein is clearly traceable. The base of
the R vein comes to lie on the dorsal margin of the second axillary sclerite. The median
plate lies along the distal margin of the second axillary sclerite, and its division into the
proximal and distal median plates is somewhat indistinct. The distal association of the
median plate with the A veins is characteristic of Hymenoptera, as Weber ( 1925) pointed
out; its primary association with the M and Cu veins is nowhere evident. The anterior
part of the third axillary sclerite apparently has a ligamentous connection with the distal
FIG. 157. A, dorsal view of Pamphilius luteicornis (original); B, dorsal view of forewing
base in Pamphilius luteicornis (original); C, dorsal view of forewing base in A pis me/lifera
(from Snodgrass); D, lateral view of left lateral arm of mesoscute!lum and corresponding stalk
of second phragma in A pis mel/ifera (from Snodgrass); E, cephalic view of pronotum and
mesothorax in Pamphi/ius luteicornis (original).
4'TH AXIL. SCL.
(POST. NOT. PROC.
SCUTELLU
MEDIAN
PRE EPISTERNUM
+ KATEPISTERNUM
+MESOSTERNUM
MATSUDA! THE INSECT THORAX
363
B
ANAL PLATE
- ~
3'RD AXIL. SCL.
D
364
BASALARE
BASALAR
TENDON
PLEURAL
RIDGE
EPISTERNUM
BASALAR
TENDON
EPIMERON
A
SUB ALAR
TENDON
B
TRANSPLEURAL
SUTURE
SUBALAR PIT
EPISTERNUM
TRANSSCUT AL SUTURE
SCUTUM 2
PRESCUTOSCUTAL
SUTURE+ ANTERO-
LATERAL SCUTAL SUTURE

--.,.,- METANOTUM
--+--1-
1
----l'ST ABO. TERG.
TEGULA ------+-----...,t,{L
\-----++---SECONDARY SUTURE
3
;:;: EPISTERNUM 2
C ERVICOPLE URON
PLEURAL SUTURE 3
PLEURAL SUTURE 2
SPIRACULAR LOBE
Fro. 158. A, inner view of mesopleuron and mesosternum in Sirex sp. (from Weber); B,
inner view of mesop!euron and mesosternum in Rhyssa persuasoria (from Weber); C, lateral
view of mesop!euron in Apis mellifera (from Snodgrass); D, lateral view of thorax in Apis
mellijera (from Snodgrass).
EPIMERON 2 ~ . -------
t-s 1
Z'ND PHRAGMA
AXILLARY LEVER
B
4'TH AXIL. S C L . - - - - - ~
365
Z'ND
PHRAGMA
AXILLARY LEVER
FIG. 159. Second phragma and lateral muscle lobes in Hymenoptera (from Snodgrass). A,
Sirex abottini; B, Sphecius speciosus; C, Apis mellifera.
end of the median plate and with the base of the A vein. The third axillary sclerite
articulates proximally with the fourth axillary sclerite (the posterior notal wing process).
In Apis (Fig. 157C) the basicostale (humeral plate) has apparently become enlarged
or fused with the basisubcostale, and the original relationships of the first and the second
axillary sclerites with the bases of anterior veins and with the median plate are difficult to
discern. Peculiarly, the first axillary sclerite is posteriorly connected with the fourth
axillary sclerite. The third axillary sclerite and the median plate have direct connections
with the base of the A vein.
In Pamphilius the ventral second axillary sclerite is connected with the basalare and
also with the tip of the pleural wing process. The ventral third axillary sclerite is
connected with the distal end of the elongate subalare. The latter is in turn connected
with the ventral second axillary sclerite and with the pleural wing process. In Vespula
(Duncan 1939) the third axillary sclerite is connected with the subalare; behind the
pleural wing process is a fossa, in which the third axillary sclerite turns during move
ments of the wing. In Apis (Fig. 158C) the ventral second axillary sclerite articulates
with the pleural wing process and with the subalare.
The mesopleuron: In Sirex (Fig. 158A) the oblique pleural ridge is clearly traceable
from the pleural wing process down to the pleural process that articulates with the coxa.
The pleural arm is absent internally, and the relatively large epimeron is present. In
Apis (Fig. 158C) the pleural ridge has not been correctly identified in the past, and
hence the dorsal part of the episternum has been variously interpreted. Snodgrass
( 1925) thought that the suture he later ( 1942) called the recurrent groove was the
pleural suture. In 1942 Snodgrass thought that the pleural suture consisted of a ventral
suture passing the pleural arm and the anterior extension of the recurrent suture.
According to Snodgrass ( 1942), two epimeral areas lie on the dorsal area of the meso-
pleuron above his pleural suture, and further, the pleural wing process is an epimeral
structure, and the basalare lies above the epimeron.
If the location of the pleural arm is the only criterion in identifying the pleural
suture, the pleural suture of Snodgrass is certainly the true pleural suture. Wheri, how-
ever, the epimeron is already so much reduced in Vespa (Weber 1925), in Rhyssa (Fig.
158B), and in Stenobracon (A! am 1951), the presence of such a large epimeral area and
in such an atypical position in Apis is difficult to conceive. Furthermore, the pleural
arm is absent in Sirex and Pamphilius; the nature of the pleural arm in Apis must there-
fore be seriously questioned.
Daly's (1964) study on the developmental anatomy of the honey bee provides a
decisive clue, with which to clarify this confused dorsal area of the mesopleuron in Apis.
Daly found in the pharate pupa of Apis that an invagination (episternal scrobe of Daly),
which later became the pleural arm of Snodgrass, appears in front of the pleurointer-
segmental suture (the true pleural suture plus the intersegmental suture). The sutures
that are associated with this scrobe (internally an arm) are formed later. Therefore, he
thought that these sutures should not be included in the course of the pleural sulcus,
or suture.
If these sutures are not part of the pleural suture, the only logical dorsal part of the
pleural suture is the dorsal posterior margin of the pleuron that reaches the pleural wing
process as in Rhyssa (Fig. 158B), and the epimeron has become nearly completely lost
in higher Hymenoptera including Apis. As in Psocoptera, the pleural arm in Hymenop-
tera is not necessarily a part of the pleural ridge (suture) . AI am (19 51) showed in
Stenobracon a pleural arm that occurs on the pleural suture where a seconcfary suture
(transpleural suture of Duncan) meets; in Vespa Weber ( 1925) showed a comparable
arm that is unconnected with the pleural ridge and is united with the furcal arm.
The transpleural suture, the preepisternal suture, and the scrobal sulcus of Daly on
the dorsal part of the pleuron are secondary formations in higher Hymenoptera.
According to Duncan ( 1939), the pleural suture arises from the subalar pit and reaches
the middle of the mesopleural posterior margin in Vespula; the area cut off dorsally by
this suture is the mesepimeron. It is obvious that such an interpretation is erroneous.
In many Hymenoptera a common suture on the episternum is an oblique suture,
which Matsuda (1960a, b) called the ana pleural suture. This suture (Fig. 157E) often
corresponds to the suture q of Snodgrass ( 1910). Its posterior end, unlike the typical
anapleural suture in holometabolous insects, reaches or nears the ventral end of the
pleural suture. Therefore, there is some uncertainty about its identity as the anapleural
suture. Another interpretation of this suture is that it is a combination of the anapleural
suture and the distal margin of the eutrochantin or of the paracoxal suture as seen on the
episternum in Trichoptera (Fig. 143A) and Lepidoptera (Fig. 147A).
It should also be pointed out that the line q ( sternopleural suture) of Snodgrass
(1910) is not always the same as the anapleural suture here identified. Thus, in his
fig. 23 (Erymotylus) and 27 (Tragus), q reaches ventrally near the anterior margin of
the coxal base, and in fig. 43 (Microterys) q extends anterolaterally from the lateral
coxal base, instead of running proximally. Compere (1962) applied Snodgrass' in-
terpretation of q ( sternopleural suture) in M icroterys to the corresponding sutures in
Scelioencyrtus (Fig. 160A), Astymachus, and Encyrtus, which belong to Encyrtidae.
It is immediately evident from the figure that the suture in question is the pleural suture;
it is neither the ana pleural suture nor the sternopleural (or pleurosternal) ridge as
Compere claimed, because the suture (ridge) leads dorsolaterally to the pleural wing
process. In Encyrtidae and probably in related families the anapleural suture is repre-
sented by a horizontal suture, which reaches the middle of the pleural suture (Fig. 160A).
PLEURAL
ANAPLEURAL
SUTURE?
WING PROCESS
VENTRAL
PROCESS
FURCA 2
COXA2
COXA 3
B
367
BASI STERNUM
PLEUROSTERNUM 2
STERNUM 3
ABO. STERN.
FIG. 160. A, mesothoracic pleuron and sternum in Scelioencyrtus myrmaricoides (from
Compere); B, internal sternal skeletons in A pis mellijera (from Snodgrass).
The opinions regarding the identity of the paracoxal suture in Hymenoptera have
diverged greatly. In Cephus, according to Michener ( 1944), the paracoxal suture (his
precoxal suture) starts from the discrimen (median longitudinal line), immediately in
front of the mesofurcal pit, bends sharply forward to the dorsal corner of the coxal
cavity, runs, partly obliterated, to the dorsal anterior corner of the episternum. Richards
( 1956) suggested a paracoxal suture in Hymenoptera, which runs from the ventral end
of the pleural suture to the dorsal anterior area of the episternum. In other insects the
paracoxal suture never reaches the anterior dorsal part of the episternum. It is therefore
obvious that the sutures proposed by these authors are not homologous with the para-
coxal suture in other insects. As Daly ( 1964) pointed out, the paracoxal suture of
Michener is an assemblage of the subpleural sulcus (already noted), the epicnemial
sulcus (along which the prepectus is formed anteriorly), and the suture anterior to the
coxal margin.
Matsuda ( 1960a, b) proposed an imaginary paracoxal suture in Arge, which extends
anterolaterally beyond the anapleural suture and meets the supposed paracoxal suture
which has been left on the epimeron. This interpretation is also invalid, since a para-
coxal suture never crosses beyond the anapleural suture. Furthermore, the presumed
paracoxal suture on the epimeron is connected with the suture along which the postnotum
meets the epimeron; the suture on the epimeron is therefore not the paracoxal suture, but
is the transepimeral suture present in Neuroptera and Diptera. In some other sawflies,
such as Schistocerus (Weber 1927a), the transepimeral suture is also present. The
above discussion leads us to conclude that in Hymenoptera the paracoxal suture is either
absent or, as already discussed, the distal part of the paracoxal suture on the episternum
is continuous with the anapleural suture.
In a typical ventrolateral part of the hymenopterous mesopleuron (Fig. 157E), the
area proximal to the anapleural suture and above the coxa must contain the preepisternum
and the katepisternum. The ventropleurite is represented by the ventral condyle that
articulates with the coxa. The trochantin has not been found in the mesothorax of
Hymenoptera.
The basalare is represented by a single small sclerite, which lies immediately
anterior to the pleural wing process in the mesothorax; it has an internal tendon for
muscle attachment (Fig. 158A, B). In Vespula the basalare rocks in and out in the
emargination on the dorsal margin of the episternum. In Stenobracon an emargination
similar to that in Vespula occurs. In Apis (Fig. 158C) the basalare is held by a wide
lobe lying dorsal to the pleural wing process.
The subalare is small; it maintains connection with the second axillary sclerite in
Apis, and in Vespula it has a ligamentous connection with the third axillary sclerite.
In higher Hymenoptera, especially in Ichneumonoidea and Chalcidoidea, the
anterior margins of the episternum and the sternum have a continuous secondary
evagination, the prepectus. Another secondary suture in higher Hymenoptera is a
short, ventral suture, which has been called the subpleural signum. According to
Michener ( 1944) and Leclercq ( 1957), it represents the remnant of the paracoxal
suture. Daly's (1964) study, however, has clearly shown that the short suture, or the
signum, merely marks the point of attachment of a dorsoventral muscle.
The mesosternum: In some Chalcidoidea the sternal area is widely exposed between
the episterna. In Encyrtidae (Fig. 160A), as Compere (1962) showed, the furcal pits
are widely separated from each other, and the ventral processes articulate with the widely
separated ventral leg bases. Thus, the plates above the furcal pits are obviously the
sternum. Although Compere called this plate the furcasternum, it is actually the basi-
sternum, since it faces laterally the preepisternal part of the composite preepisterno-
katepisternum (laterosternite of Compere). In Hymenoptera the mesothoracic furca-
sternum posterior to the furcal bases has become lost."
One of the species Compere studied, Scelioencyrtus mymaricoides, crawls between
the sheaths and culms of graminaceous plants. For insects that crawl in a space with
such closely suppressed surfaces, the closely approximated, typical hymenopterous legs
simply would not function properly as an organ of locomotion. Hence, the presence of
the widely separated leg bases and of the widely exposed sternum in some parasitic
Hymenoptera (including Scelioencyrtus) is conceivable.
In Eupelmus, also, as Richards ( 1956) showed, the sternum is also clearly exposed.
This exposure of the sternum is correlated with the abnormal course of the pleural suture,
which runs posterolaterally from its point of articulation with the coxa, as in
Scelioencyrtus (Fig. 160A). In Eupelmus, however, the furcal pit is single, despite
the widely exposed sternum; in this respect Eupelmus resembles some parasitic Diptera
(p. 319).
Typically, the border between the pleuron and the sternum has become completely
lost in the mesothorax of Hymenoptera, as in most holometabolous insects. Daly
( 1964) has shown in A pis that the internal median longitudinal ridge in the basisternal
area is produced by a migration or a proliferation of cells from a low midventral ridge.
This precludes the possibility that the basisternum is completely invaginated to form the
median lamella; the presence externally of an indefinite area of the basisternum on either
78Reduction of the mesothoracic furcasternum has occurred also in Orthoptera, Hemiptera, and some higher
orders.
ANAPLEURAL
SUTURE
TRANSEPIMERAL
SUTURE
369
PRE EPISTERNUM
+KATEPISTERNUM COXA
FIG. 161. Ventral view of metathorax in Arge sp. (original).
side of the median longitudinal line ( discrimen) is therefore probable, as Richards
-(1956), Compere (1962), and Daly (1964) contended.
79
Posteriorly the ventropleurites
(ventral processes) lie so closely along the median longitudinal axis that the posterior
sternal element must be present externally in a very narrow area between the ventral
processes and the furcal pit. The mesothoracic spina has become lost, concomitant with
the reduction or loss of the furcasternum.
The metanotum: In Chalastogastra the metanotum is well exposed. Laterally the
postnotum and the epimeron are either indistinguishably fused or fused along a suture.
Posteriorly, the postnotum is distinct from the first abdominal tergite; incorporation of
the first abdominal segment into the functional thorax is therefore not pronounced.
The third phragma is not formed, at least in Pamphilius examined.
In Wasps (Vespa, Vespula, and Stenobracon) the metanotal anterior margin is
loosely connected with the posterior margin or marginal flange of the mesothoracic
scutellum. Laterally, in Vespula (Duncan 1939) and Stenobracon (Alam 1951), the
metanotum is provided with a lateral carina, which loosely separates a small, lateral area;
Duncan and Alam called this area the lateral ramus. Both in Stenobracon and in
Vespula there is another detached anterior sclerite, which Duncan called the humeral
plate; it bears the anterior notal wing process, and in Stenobracon it connects with the
second phragma. The metapostnotum is represented by lateral remnants in Vespa
(metaphragma of Weber 1925); in Vespula they form crests that surmount portions of
the metapleura and the propodeum.
In Apis the metanotum is a narrow, mesally constricted plate interpolated between
the mesonotum and the propodeum (Fig. 15 8D) . It has a lateral, semidetached area
that corresponds to the metanotal ramus in the wasps; a smaller independent sclerite in
front of the metanotal ramus appears to be the humeral plate present in Vespa, because
the head of the first axillary sclerite rests on this plate. The third phragma, according to
Snodgrass ( 1942), is absent in bees.
The metapleurosternum: In Chalastogastra the pleural suture is distinct throughout
its entire length, from the dorsal pleural wing process to the pleural process that
articulates with the coxa. In Arge (Fig. 161) the anapleural suture is present, as in the
mesothorax. Matsuda (1960a, b) thought that the anterior margin of the membranized,
area above the coxa and an oblique suture on the epimeron was the paracoxal suture.
The former is not a suture and hence it cannot be the paracoxal suture; the suture on the
epimeron cannot be the paracoxal suture, for the same reason given in discussing the
corresponding suture on the mesothoracic epimeron.
"'As already discussed (p. 21), the presence externally of the sternum is more probable in a greatly enlarged
thoracic segment, such as the mesothorax in Apis.
In Clistogastra the pleural suture (sulcus) has become more or less obliterated. In
A pis (Fig. 158D), for instance, only the lower half of the true pleural suture is retained;
a secondary suture is continuous dorsally from this suture and a pit lies on this suture.
Another secondary, oblique suture divides the metapleuron into dorsal and ventral parts
in Vespula and Apis (Fig. 158D); a secondary suture in Vespa (Weber 1925), which
bears internally a well-developed apodeme, probably corresponds to this suture. The
basalare and subalare, though small, are recognized in the metathorax of Hymenoptera.
Fusion of the metapleuron with the first abdominal tergite is usually evident by the
line of their fusion. When they are completely fused without a line, the presence of the
first abdominal spiracle easily demonstrates the fusion of the two areas. The line of
fusion may be internally grooved, and may be split up into dorsal and posterior parts.
The episternum is indistinguishably fused with the sternum in the metathorax. A
claimed pleurosternal suture in Stenobracon (Alam 1951) may be the anapleural suture.
The ventral side of the metathorax has the median longitudinal groove, as in the
meso thorax.
In higher Hymenoptera the internal median longitudinal ridges in the meso- and
meta-sternum are continuous, as seen typically in Apis (Fig. 160B). The furcal arms
of both the mesosternum and metasternum arise from a common sternal inflection in
Apis (Fig. 160B) and Vespa. The furcal arm is either loosely united with the meta-
pleural arm as in Stenobracon, or fused with the pleural arm as in Vespula, or solidly
united with a thick ridge following the external suture between the metapleuron and the
propodeum, as in Apis.
Phylogenetic relationships of Hymenoptera: Thus far no close phylogenetic relation-
ship between Hymenoptera and any particular order has been established, although
Crampton (1919) suggested their affinities with Neuroptera and Mecoptera, and Ross
( 1936), based on the similarities in wing venations, maintained their derivation from the
Megaloptera-Mecoptera stem.
In certain thoracic structures the affinity of Hymenoptera with Mecoptera is
evident. They are: ( 1) a loose connection of the posterolateral angle of the pronotum
with the mesepisternum in Mecoptera and in a primitive hymenopterous genus
Pamphilius; (2) the absence of the prealar arm in most Mecoptera and in all Hymenop-
tera; and (3) the presence of t-p 19 in Mecoptera and Hymenoptera (and not in other
orders). All of these characters can be regarded as synapomorphic. (1) suggests that
Hymenoptera lost this character early in their evolution. The absence of muscle t-p 10
and the presence of t-p 12 are symplesiomorphic characters that are shared by Neurop-
tera and Hymenoptera.
Based on the above evidence, Hymenoptera may be regarded as a sister group of
Mecoptera. Hymenoptera appear to have departed early from the common stock, and
their structural differentiation has been rapid enough to obscure its relationship to
Mecoptera and Neuroptera. As Snodgrass (1910) pointed out, several lines of pro-
gressive modifications of thoracic structures within the Hymenoptera (autapomorphic
characters)"' are responsible for the origin of the order Hymenoptera.
The anatomical basis of flight: The flight mechanism in A pis mellifera was explained
by Stellwaag (1910), Snodgrass (1956), and Pringle (1957a, 1961). In Apis (Fig.
BOThe autapomorphic characters are: the s_eparation of the pronotum from the propleuron and its attachment
to the front of the mesothorax; the separation of the mesonotum into an anterior and a posterior plate by a
transverse membranous suture ( transscutal suture); concealment of the mesopostnotum and its phragma by
invagination within the cavity of the thorax; the reduction of the metanotum to a simple transverse plate .
carrying the hind wing; the fusion of the metapostnotum with the propodeum or the first abdominal segment;
the fusion of the first abdominal segment with the metathorax, and its complete incorporation into the
thoracic division of the body in Clistogastra; the formation of a single large mesopleural plate on each side
by the narrowing of the mesepimeron and the suppression of the mesopleural suture, and its secondary
division into an upper and. a lower plate; the formation of the prepectal plate in the mesothorax cut off from
the anterior parts of both the mesosternum and the mesopleuron; the obliteration of the metapleural suture
resulting in the formation of a- single metapleural sclerite, which becomes divided again into an upper and
lower parts.
Besides these autapomorphic trends, the reduction of tergocoxal muscles, the loss of the association of
median plate with the M-Cu vein, and the fusion of the lateral cervical sclerite with the proepisternum
be adc!ed as characters of Hymenoptera. The secondary separation of the mesonotum
mto the anter10r and postenor parts by a transscutal suture has occurred also in Diptera.
MATSUDA! THE INSECT THORAX
371
157C) the first axillary sclerite is well developed anteroposteriorly. It has articulations
anteriorly with the anterior notal wing process, and with the humeral complex; its caudal
end has a ligamentous connection with the fourth axillary sclerite. This unique develop-
ment of the first axillary sclerite is important in the wing movements of Apis melli/era.
According to Pringle ( 1957a), mesothoracic indirect flight muscles (t 13, 14,
t-p 5, 6) are responsible for the basic movements of pronation and supination, as well as
for depression and elevation of wings. In the mesothorax of Apis, according to Pringle
(1957a), the force of contraction of the dorsal longitudinal muscle (t 14) is transmitted
to the base of the posterolateral scuta! area ( scutellar arm of Pringle), which carries the
postmedian notal process (posterior notal process of Pringle). The presence of the
transscuta! suture (scuta! fissure of Snodgrass, mesonota! suture of Pringle) allows the
postmedian notal process to slide forward and upward, under the lateral wall of the
scutum bearing the anterior notal wing process.
This motion is transmitted to the wing base by the first axillary sclerite. When the
transscutal fissure closes, it turns the caudal end of the first axillary sclerite upwards, and
the upper end of the first axillary turns forwards and downwards. Since the first axillary
sclerite is attached to the humeral complex, the wing is thereby pronated as well as
depressed. The movement of the posterolateral scuta! area rotates the first axillary
-sclerite round the second axillary sclerite, which pivots forward about the pleural wing
process, lifting its posterior arm. When further contraction of the dorsal longitudinal
muscles moves the anterior notal process outwards and upwards, the second axillary
sclerite rotates further, so that the wing is depressed in a pronated attitude. At the
beginning of the upstroke, contraction of the vertical indirect muscles (t-p 5, 6) moves
the scutum downwards and backwards, which has the effect of reversing the movements
of the first and second axillary sclerites.
Further, according to Pringle ( 1961), in A pis the basalar, subalar, and scutel!ar
(scuta!?) muscles of the mesothorax alter the setting of the sclerites so that the amount
of pronation varies at different instants during the downstroke, giving control of balance
in the planes of pitch and yaw; the upstroke is ~ ! m o s t invariant, with full supination.
The pleurosterna! muscle (p 3) and the axillary lever muscle (t-s 1) control the power
generated by fibrillar indirect muscles and thus the amplitude of wing beat.
As in Hemiptera, the coupling mechanism of the fore and hind wings is important in
the flight mechanism of Hymenoptera. In Vespa Weber ( 1925) described this
mechanism in detail. Unlike in Hemiptera, however, the dorsal indirect flight muscles
are present in the metathorax of Hymenoptera. According to Pringle ( 1961), power for
moving the hind wing is drawn from the mesothoracic muscles to an extent controlled by
the mesophragmal retractor muscle ( t 14) of the meta thorax.
In contrast with some specialized orders (Orthoptera, Coleoptera, Lepidoptera,
etc.), the tergocoxa! and tergotrochanteral muscles have undergone a great reduction in
Hymenoptera. The functional change of these muscles from primarily leg muscles to
flight muscles has, therefore, never taken place during the evolution of Hymenoptera.
It is perhaps partly because of this peculiar evolutionary trend of the leg muscles that the
indirect flight muscles, as Pringle (1957a, 1961) has shown, have come to assume such
a dominant role in the flight mechanism of higher Hymenoptera, such as Apis.
The adult musculature: The thoracic musculature in Hymenoptera has been studied
by Lubbock (1879) in Lasius flavus (worker), Berlese (1909b) and Weber (1925) in
Vespa crabro, Morison (1927) in Apis mellifera, Weber (1927a) in Tenthredo sp. and
Schizocerus sp., Maki (1938) in Euthomostethus formosanus, Philopsyche sauteri and
Vespa ducalis, Duncan (1939) in Vespula pensylvanica, Snodgrass (1942) in Apis
mellifera, Alam (1951) in Stenobracon deesae, Wille (1956) in bees, Saksena (1959) in
Polistes hebraeus, Daly (1964) in Apis mellifera, and by Miirkl (1966) in Apis mellifera,
Formica polyctena and Vespa vulgaris. Out of these studies, the muscles in Euthomo-
stethus formosanus (Maki 1938), Stenobracon deesae (Alam 1951), Vespula pen-
sylvanica (Duncan 1939), Apis me/litera (Snodgrass 1942), and Formica polyctena
(Mark! 1966) are homologized in Table XXII. In the table cv-cx(x) 1 in Apis is a
muscle found by Daly ( 1964).
The most conspicuous feature of the hymenopterous thoracic musculature. is the
great scarcity of the tergocoxal muscles; only t-cx 8 is present in most Hymenoptera.
Such reduction of the tergocoxal and tergotrochanteral muscles has not occurred in
other major orders. Another unique feature is the wide occurrence of t-p 19, which
elsewhere occurs only in Mecoptera. The occurrence of t-s 4 in Euthomostethus
requires confirmation, since this muscle occurs only in Ephemeroptera.
In Table XXII homologies of cervical muscles are based on the conclusion that the
lateral cervical sclerite is not differentiated as a separate plate in Hymenoptera. The
cervical musculature thus identified shows no significant deviation from the corresponding
musculature in other insects,
Gross aspects of the postembryonic muscle development: Daly's (1964) study on
the postembryonic development of thoracic muscles in Apis mel/ifera has revealed the
following major facts: In the larva the thoracic muscles consist of ventral external longi-
tudinal muscles ( ve.), ventral internal longitudinal muscles (vi.), lateral external oblique
muscles (/e.), lateral internal oblique muscles (li.), transverse ventral muscles (tv.),
cruciate bands (cr.), dorsoventral external muscles (dve.), dorsoventral internal muscles
( dvi.), dorsal external oblique muscles (de.), and dorsal internal longitudinal muscles
(di.).
In the early pharate pupa the major groups of larval muscles are still recognizable.
Many imaginal muscles are formed at this stage in association with larval fibers; most of
them persist till the adult stage is reached. They are: ve in the prothorax becomes
fu 1-poc b ( s 1 or 2) and fu 1-pl 1 ( cv-s 4), and in the pterothorax it becomes fu 2-fu 1
and fu 3-fu 2 (s 13); prothoracic le becomes fu 1-poc a (s 1 or 2); li becomes fu-ph
(t-s 1) in all three thoracic segments; tv becomes fu 2-1 sps (s 12); cr becomes
pl l-ex lc(x) ( cv-cx(x) 1); dve p is associated with pi-poe a, b, c ( op-p 1, op-cv 3); dvi 1
becomes pi 2-t la, b (t-cv 3) and dvi 2 becomes t 2-pl2a (t-p 5, 6); de 1 becomes a part
of 1 ph-t (t 13) and de 3b becomes t 3-3ph (t 14); prothoracic di forms poc-1ph a, b, c, d
(op-t 3); di 2b forms 1 ph-2ph (t 14); and di 2a forms t 2-t 3 (t 13?).
In the same stage new imaginal muscles are formed, without obvious relationship to
larval muscles. They are largely lateral muscles that are concerned with flight. They
include: pl2-ba 2a, b (p 3), ex 2-sa 2 (t-cx 8), pl 3-ba 3a, b (p 3), ex 3-sa 3 (t-cx 8),
pi 2-3 ax 2a, b, c (t-p 13), pl3-3ax 3a, b, c (t-p 13), p12-t 2b (t-p 12), pi 2-t 2c (t-p ?),
pi 3-t 3a, b, c (t-p 5, 6?), p! 3-t 3d (t-p 5, 6?), pl 2-sp 2 (spiracle muscle), pl 1-t lc, d
(prothoracic t-p muscles), t l-ex 1 (t-cx 8). The vi muscles disappear progressively
during this stage.
During the pharate pupal stage the musculature is repositioned along with the
changes in body form to resemble the adult arrangement of muscles. le 2 and le 3
(t-p 30) disappear during this stage. According to Daly (1965), 12 degenerating pupal
muscles in Apis are homologous with adult muscles in Sirex and Xyela.
In the pupa, despite drastic changes in forms of the integument, the system of
muscles in the pupa is nearly the same as that in the pharate pupa. Alterations, how-
ever, include fu 1-1 ph (t-s 1), which degenerates; fu 2-1 sps (s 12), which loses the
median connection with the spina, becoming fu 2-fu 2 (s 16) one day after ecdysis, and
eventually degenerates; fu 3-pl 3 (p-s 1), which is reduced in width; and t I-t (t ?),
which disappears. Both t 2-pl 2a (t-p 5, 6) and 1 ph-2 ph (t 14) undergo two periods
of enlargement. In the early pharate adult fu 1-1 ph (t-s 1), t 2-pl 2c (t-p 4?), fu 2-pl 3
(p-s 3 or 5), and pl 3-t 3d (t-p ?) degenerate.
In the pharate adult all the muscles have approximated their adult size and form.
As a result of the increases in the areas of the origins, the former relationships of muscle
rudiments to the developing integument are observed, i.e. muscles come to occupy some-
what different points of attachment. At the same time, however, some muscles
degenerate during the early pharate adult. They include: fu 1-1 ph (t-s 1), t 2-pl 2c
(t-p 4), fu 2-pl 3 (p-s 3?), and pl 3-t 3d (a t-p muscle).
From the above account of the muscle development in Apis, it is dear that larval
vi and larval meso- and meta-thoracic le (t-p 30) are caenogenetic muscles, which
disappear and have nothing to do with the adult musculature of insects, and that, as in
Orthoptera and Coleoptera, the postimaginal degeneration of muscles occurs in
Hymenoptera.
The fact that lateral muscles concerned with flight appear in the pharate pupa
without any relationship with larval muscles suggests that their emergence is considerably
delayed, as compared with homologous muscles in hemimetabolous insects.
The postembryonic development of flight muscles: Tiegs (1922) described the
development of flight muscles (t 14 and t-p 5, 6) in Nasonia as follows: Although most
of the larval muscles degenerate, three or four pairs of dorsally situated thoracic muscles
become enveloped by the myoblasts, which were already present in the earliest larva,
lying near the muscles. The proliferated myoblasts extend forward from the meta-
thoracic segment and the rear of the mesothoracic segment. The extension proceeds in
a pair of great columns partly upon, partly independently of the larval muscles, drawing
the neighboring longitudinal muscles together as they advance. The myoblasts, several
hours later, begin to penetrate the tough sarcolemma, and work their way onto the
degenerating muscle substance; 8 hours after defecation the whole muscles are riddled
-with myoblasts; 122 hours later, no trace of larval muscles remain; in their place there
occurs now a pair of bands of myoblasts lying some distance below the integument.
Earlier, Perez (1912) described, in Polistes, the development of the dorsal longi-
tudinal muscle ( t 14), which was very similar to that in N asonia (Tiegs 1922). Perez
(1912) pointed out that the only significant difference in the mode of development of the
flight muscles between Polistes and Calliphora (Perez 1910) was the fact that the site
of emergence of myoblasts is external to the larval muscle, while in Polistes they lie close
to the larval muscles during development.
According to Janet ( 1907), the dorsal longitudinal muscle ( t 14) in the queen of
ants degenerates after the nuptial flight.
Histology: Daly's (1963) investigation of 18 indirect and direct flight muscles in 58
species of Hymenoptera has shown that almost one fourth of the species has fibrillar
muscles (fibrillar diameter over 1.5 I") in both segments of the pterothorax and that
one sixth have fibrillar direct flight muscles attached to the basalare (p 2, p 3, p-tr 2,
p-cx 2 or 3). For more detailed information on the histology of thoracic muscles in
Hymenoptera, refer to Morison (1928), Jordan (1933), and Darwin and Pringle (1959).
THE STREPSIPTERA
The following discussion applies exclusively to the winged male.
The neck region: The ventral neck region in Halictophagus is totally membranous.
According to Kinzel bach ( 1967), the lateral cervical sclerite is absent in Strepsiptera; in
Halictophagus the head is directly pressed on the ventral surface of the prothorax in the
resting position. Dorsally, in Halictophagus, the membrane between the head and the
pronotum is provided with sclerites, as shown in Fig. 162A; Kinzel bach ( 1967) called
the anterior dorsal sclerites the Vertexplatte.
The prothorax: The prothorax in Strepsiptera is greatly reduced. The pronotum
and the propleuron are fused in varying degree to form a ring. In Halictophagus the
pronotopleuron extends anterolaterally, and may appear at first glance to be the lateral
cervical sclerite. The pleural suture is recognizable in Halictophagus (Fig. 162B);
according to Pierce (1918), it is distinct only in Pyrilloxenos, another genus of
Halictophagidae.
On the ventral side of the prothorax in Halictophagus (Fig. 162C), a membranous
cleft extends posteriorly as far as the pleural process, roughly where the anapleural cleft
might occur. The median T-shaped sclerite is the pleurosternum, and its original com-
ponents are indistinguishably fused; its anterior margin is strongly ridged, and a small
sclerite anterior to the middle of this ridge is the prepectus.
TABLE XXII
Thoracic musculature in Hymenoptera
Vespula A pis Formica
Euthomostethus Stenobracon (Duncan (Snodgrass (Mark! Muscle
(!Vlaki 1938) (Alam 1951) 1939) 1942) 1966) designations Remarks
43, 44 op-t 1, 2
40, 41 40+41 op-t 3 1
4 41 42b, e 42 op-cv 3
7 42 32 42a? op-p 1
5
40 33 43 43 op-s 2
50 37 48 48 t-ev 2 2
6 38 47 46+47 t-cv 3
51 46 t-s(ev) 9 3
2 54 36 51 51 ev-s 4
10 49 pl 1- cv-cx(x) 1
ex leX
16 (I) 55 44 53 53 ev-ex 3
I
45 34, 35 44 44 s I, 2
43 (I !I) tll
47 (!) 51 t 12
21 (II)
42 (Ill)
46 (I) 50 45 45 t 13
20 (II) 70 72 70
18, 19 (II) 62 56 71 t 14
41 (Ill) 75 96
69 (I!) t ? 4
28 (I I) 74
47 (III)
t-p 4? 5
24 (II) 111 57
45 (III)
72 72 t-p 5 or 6 6
97?, 98?, 99?
29 (II) 68? 62 75 t-p 12 7
48 (Ill) 84?, 85?, 79
86?
66, 67 (II) 60, 61 76 t-p 13
49 (III) 88 100
30, 31 (II)
50 (III) 89 77
t-p 14
68 (I I)
t-p 15 ? 7
27 (II)
46 (II I)
t-p 18 8
51 (Ill) 82
64 (II)
78 102
t-p 19
8, 9 49,52,53 39-42, 51 49, 50 49, so t-p(proth.) 9
25, 26 (HI) 63 58 78 t-s 1
45a (II-III)
52 (Ill) t-s 4 10
12? 57? 48 55? 55? t-cx 8 11
36 72 82 82
57 81 80 105
65 (I I)
p2
32 (II) 64 59 77 p3
53 (III) 87 76 101
33, 34 (II) 69 70 79 p-sI
54, 55 (III)
TABLE XXII (Continued)
Euthomostethus Stenobracon
Vespula
(Duncan
A pis
(Snodgrass
(Maki 1938) (Alam 1951) 1939) 1942)
59 (III)
77a (Ill)
59 47 57
38 (II) 73 80
77b 84 103?
22, 23 (HI) 48 53, 54 52
44 (II-III) 71 73
3 (I)
79 (Ill) 71
14, 15? 58 46 56
37 75 66
58 79 82
13 (I) 60
80 (III) ?
52 58
11 56 43 54
35 74 65 81
56 78 81 104
75 (II) 68 83
106 (III)
17 (I) 61 45 61
83 (pars)
39 (I I) 76 67 86
60 (III) 83 (pars) 83 109
REMARKS (Table XXII)
( 1) These muscles may be cv(d)-t 1.
(2) 48 in Formica may be t-cv 1.
(3) These muscles may be t-cv 3.
Formica
(Mark!
1966)
80
52?
56
54
81
83
106
61
86
109
375
Muscle
designations Remarks
p-t(cx) 2 or 3
p-ex4
p-ex 5
s 13 12
s 14
s--ex 2
s-ex 3
s--ex 4
s-ex 5
s--ex 6 or part
of s--ex 3
p-tr 1
s-tr 1
( 4) 69 in V espula is a tiny depressor of the tegula that arises from the meso-
thoracic scutum and is inserted on the inner surface of the tegula.
(5) 74 in Apis arises from the mesepisternum and is inserted on the ventral side
of the humeral complex of the forewing.
(6) Metathoracic muscles in Apis connect the metafurcal arm and the metanotal
lateral plate.
(7) 68 in Stenobracon is dorsally attached to the posterior notal wing process.
It may therefore be t-p 15.
(8) 27 in Euthomostethus may be t-p 8 or t-p 10.
(9) 39 and 40 in Vespula and 49 in Stenobracon may be t-cv 1.
(10) 52 in Euthomostethus was described as connecting the mesofurca and the
subalar apodeme.
( 11 ) Homologies of prothoracic muscles suggested are not certain.
(12) 52 in Formica may be p-s 3.
Besides the muscles shown in Table XXII, the following muscles have been recorded
in Hymenoptera: 63 of Vespula pensylvanica (Duncan 1939), which is an intrinsic
muscle of the third axillary sclerite; I zm in Schizocerus sp., Tenthredo sp., and Vespa
crabro (Weber 1925, 1927a), which is p-s 1 in the prothorax; 17 of Vespa ducalis
(Maki 1938), which is a prothoracic pleural remotor of the coxa (p-cx 9); II pm 2 of
Tenthredo sp. (Weber 1927a), which may be p-cx 9; t 2-cx 2 of Daly (1963) in Sirex,
Cephus, and Xyela, which is t-cx 6 or 7; t 2-tr 2 of Daly ( 1963) in Sirex, Cephus,
Xyela, and Chalcidoidea, which is t-tr 1; t 3-tr 3 of Daly ( 1963) in Sirex, Xyela,
Chalcidoidea, and Cynipoidea, which is t-tr 1. Wille's ( 1956) and Daly's ( 1963)
studies provide information about the distribution of some thoracic muscles in various
taxa of Hymenoptera.
376
PLEURAL U
WING PROCESS
2'ND AXIL. SCL.
(SUBALARE?)
ALULA
D
F
A ANEPISTERNUM
I
PLEURAL
SUTURE
STERNUM
61
-r----_COXA
f{jj
B
SURALARE ?
l'ST AXIL. SCL.
+ 2'ND AXIL. SCL.
POSTMEDIAN
NOTAL PROCESS
3'RD AXIL.SCL.
""l..------FORE-WING
PREALAR ARM?
EPISTERNUM
+EPIMERON
FIG. 162. A, dorsal view of neck region, pronotum and mesonotum in Halictophagus sp.
(original); B, lateral view of prothorax in H alictophagus sp. (original); C, ventral view of
prothorax in Ha/ictophagus sp. (original); D, forewing base in Sty lops sp. (from Ulrich);
E, lateral view of forewing base in Sty lops sp. (from Ulrich); F, lateral view of mesothorax in
Sty lops sp. {from Ulrich).
The median longitudinal ridge and the narrow area along the ridge is at least largely
basisternum. The posterior end of this area bifurcates to form paired furcae, which
project inward. What Pierce claimed to be the trochantin is probably the furca; the true
trochantin is absent. Pierce ( 1918) showed, in Triozocera, four divisions of the pro-
sternum, including the posteriormost spinasternum. Whether the prothoracic spina
occurs in the spinasternum of Triozocera or of other Strepsiptera remains to be seen.
371
The mesonotum: In Halictophagus (Fig. 162A) the mesonotum appears to consist
of transverse plates, a small anterior plate and a larger posterior one that are
separated by a membrane. The anterior plate probably corresponds to the prescutum of
Ulrich (1930) in Stylops. The scutoscutellar suture is apparently absent in the meso-
thorax of Strepsiptera. In Halictophagus and Stylops (Fig. 162F) the posterolateral
extension of the mesonotum is fused with the epimeral part of the mesopleuron, thereby
forming a postalar bridge. - Therefore, the posterolateral part of the mesonotum can be
regarded as the postnotum.
The mesoscutal lateral margin and axillary sclerites: In Stylops (Fig. 162D) the
anterolateral angle of the mesoscutum, demarcated by an oblique line, may represent the
suralare," since the first axillary sclerite ( U nterer Teil des vorderen dorsa/en Axillarstiick
of Ulrich 1930) lies lateral to it. The posterior scuta! process (obere Fliigelfortsatz of
Ulrich) is probably the postmedian notal wing process, because it is connected with the
caudal end of the first axillary sclerite."' The second axillary sclerite may be completely
fused with the first axillary sclerite, since it lies closely along the distal margin of the first
axillary sclerite in the hind wing (Fig. 163D); a small anterior distal plate is, according to
Ulrich, the dorsal part of the elongate, proximal plate. The third axillary sclerite
(hinteres dorsales Axillarstiick) lies near the posterior margin of the reduced forewing.
Ventrally (Fig. 162E) the Praeparapterum of Ulrich is associated with the anterior
basal swelling of the forewing (vorderer Basalwulst), and extends to the dorsal margin
of the episternum. It is not certain whether this plate represents the basalare, which
Ulrich's term Praeparapterum implies. Judging from its shape and position, this sclerite
may represent a detached prealar arm. Another sclerite ( vorderes ventra/es Axillar-
stuck) has two divergent tips; the homology of this plate is unclear. The supposed
subalare (Postparapterum of Ulrich) lies posterior to the pleural wing process, at the
base of the posterior basal lobe (hinterer Basallappen), which Ulrich suspected to be
homologous with the alula in Coleoptera. This supposed subalare appears, by its
position, to be the ventral second axillary sclerite.
The mesopleuron: In Pyrilloxenos Pierce (1918) showed (Fig. 163A) the pleural
suture that separates the episternum from the epimeron. In Halictophagus (Fig. 163B)
the posterior margin of the mesopleuron may ventrally represent the pleural suture, since
the ventral end of this ridged margin forms a part of the pleural process that articulates
with the coxa. In many Strepsiptera including Stylops (Fig. 162F) the mesopleuron
below the pleural wing process is a simple plate, owing to the loss of the pleural suture.
On the ventral side of the mesothorax of Pyrilloxenos (Fig. 163A) the pleurosternal
suture separates the median subtriangular sternal plate from the lateral mesopleuron. An
oblique suture from the pleurosternal suture reaches laterally the pleural process that
articulates with the coxa, and this suture may be the anapleural suture. In the absence
of the paracoxal suture, the episternum appears to consist of the anterior anepisternum
and the posterior preepisternum, although the latter probably contains the katepisternum,
which has become incorporated into this area. The area above the leg base is mem-
branous, and the trochantin is absent. In Halictophagus (Fig. 163B) the pleurosternal
suture assumes exactly the same relative position in the Pyrilloxenos, and the presumed
anapleural suture is absent.
The mesosternum: Both in Pyrilloxenos (Fig. 163A) and in Halictophagus (Fig.
163B), the sternum consists of the anterior undivided part and the posterior part, which
is divided into two narrow lobes by the median longitudinal suture. The anterior
triangular plate is to be the presternum and the basisternum that have become
fused, or it may be represented by the basisternum alone. The posterior half is the
furcasternum. The posterior end of this area bifurcates and protrudes into the thoracic
cavity; they are, therefore, obviously furcae. The furca may appear to be the ventral
"The figure shows structures under a strongly pressed position. Therefore, this identification of the sclerite
is not certain.
62
In the metathorax the first axillary sclerite is caudally associated with the median notal process.
378
PRE EPISTERNUM
+ KATEPISTERNU .
SCUTUM
FOLDED
MEMBRANE
ANTERIOR
NOTAL
PROCESS
ANAPLEURAL
SUTURE?
PLEURAL
SUTURE
BASI STERNUM
INTERSEGMENTAL
FOLD
EPISTERNUM -------t-
PREALAR ARM
PARAPSIDAL
SUTURE
ANTERIOR
N ~ J & h s ?
MEDIAN
NOTAL
PROCESS
3'RD AXIL. SCL.
E
D
--i!f-.--SCUTUM
LATERAL
POSTNOTUM
POSTNOTUM
Fm. 163. A, mesosternum in Pyrilloxenos compactus (from Pierce); B, mesosternum in
Halictophagus sp. (original); C, cephalic view of metanotum in Halictophagus sp. (original);
D, dorsal view of hind wing base in Halictophagus sp. (original); E, dorsal view of metanotum
in Halictophagus sp. (original).
process for the coxal articulation, but it does not articulate with the coxa. Pierce (1918)
mistook the furca for the trochantin.
The metanotum: In the front view of the metanotum in Halictophagus (Fig. 163C),
the second phragma arises from the antecosta. The lateral arm is directed caudally and
its end represents the anterior notal wing process; it is not the lateral phragma that is
present in Coleoptera. The apparent prescutoscutal suture (Fig. 163C) is the line of the
membranous intersegmental fold, and the prescutoscutal suture itself is absent. How-
ever, the line on which the membrane is attached is probably where the prescutoscutal
suture could normally occur, since the parapsidal suture arises from this line (suture).
The area posterior to the antecostal suture and underneath the membranous fold is,
therefore, probably the prescutal area. The semidetached prealar arm, which lies lateral
to the prescutal area, extends ventrally to the dorsal portion of the episternum; its
position relative to these neighboring structures is typical of the prescutal prealar arm.
In many Strepsiptera, as Pierce's (1918) figures show, the lateral parapsidal suture
is present and the parapsidal suture is absent. In a species of Stylops examined, the
lateral parapsidal suture extends laterally to the base of the narrow suralare. The tergal
fissure in Halictophagus is membranous and extends posteriorly toward the scutoscutellar
suture. The lateral scuta! area demarcated posteriorly by the tergal fissure (suture) was
called the parascutellum by Pierce ( 1918). In Triozocera and some other genera, the
parascutellum of Pierce ( 1918) is incomplete, not reaching the scutoscutellar suture.
The scutoscutellar suture is always well marked and forms a subtriangular scutellum.
The definitive postnotum is enormously developed in Strepsiptera. Jeanne! ( 195 L)
thought that the first abdominal tergite is incorporated into it. A postalar bridge is
formed by the union, along a suture, of the lateral extension of the postnotum with the
epimeron (hypoepimeron) and with the hind coxal base (Fig. 164B).
The metascutallateral margin and axillary sclerites: In Halictophagus (Fig. 1630)
the scuta! process immediately behind the tergal fissure is the median notal process;
besides this process, only the anterior notal wing process is differentiated.
The axillary sclerites of the hind wing in Halictophagus (Fig. 1630) dorsally
consists of the first, second, and third sclerites. The first axillary sclerite, besides its
connection with the scuta! lateral margin (anterior and median notal wing processes), is
connected by its anterior end with the base of the Sc vein. The second axillary sclerite
is a slender, linear sclerite that lies along the distal margin of the first axillary sclerite; its
usual connection with the base of the R vein is absent. The third axillary sclerite lacks
the connections with the scuta! lateral margin. The distal ends of the first and second
axillary sclerites are intimately connected with the third axillary sclerite.
In Halictophagus (Fig. 164A) the base of the C vein fits in the notch on the anterior
end of the basalare in the resting position. The ventral second axillary sclerite is
connected with the pleural wing process. The long and slender process from the subalare
extends far into the axillary membrane, and is associated with the axillary sclerites (Fig.
164A, B, C). Kinzelbach (in press) showed the tegula and a ligamentous connection of
the subalare with the first axillary sclerite in Stylops.
The metapleuron: In Halictophagus (Fig. 164B) the pleural suture is continuous
from the oblique coxal margin, bent laterally near its dorsal end, and it carries a robust
pleural wing process at its end. The episternum lacks the usual division into three
parts. The proximal margin of the sclerotized episternum faces a broad membranous
area, which roughly agrees in position with the anapleural cleft in Coleoptera, in which
also the metathorax is prolonged and the course of the cleft is nearly cephalocaudal.
The preepisternal area in Halictophagus is membranized. In Halictophagus (Fig. 164B)
the apically notched basalare lies anterior to the pleural wing process and its base is
connected with the dorsal margin of the anepisternum, although the basalare is isolated
in Stylops (Fig. 164C). The epimeron is well retained along the pleural suture. In
Halictophagus (Fig. 164B) a detached subquadrangular area (hypoepimeron) lies along
the posterior part of the pleural suture, and this area is united with the coxal base and
380
A

:r
PLEURAL
SUTURE
..
...
. .
. . . :.
\ ...

STERNUM ).'.'
SUBALARE
POST NO TUM
B
FiG. 164. A, lateral view of hind wing base in Halictophagus sp. (original); B, ventral view of
metapleurosternum in Halictophagus sp. (original); C, lateral view of metapleurosternum in
Stylops pacificus (original).
381
with the lateral postnotum along a suture. The trochantin is absent. The pleural
suture is absent near the coxal margin in Triozocera (Pierce 1918). As Kiozelbach
(in press) has' shown, the coxa has become fused with the pleurosternum in varying
degree in various genera.
In Sty lops Ulrich (1930) showed the subalare (postparapterum of Ulrich), to
which a coxal muscle (t-cx 8) is attached. In the species of Stylops examined (Fig.
164C) the subalare has a straight, anteriorly projecting process that connects with the
axillary sclerites. However, in Halictophagus (Fig. 164B) the corresponding area (the
subalare) is continuous from the postalar bridge; probably this condition has resulted
from a secondary fusion of the subalare with the lateral postnotum. Thus, in H alicto-
phagus the lateral process may appear to be homologous with the lateral postnotum iD
Coleoptera (Fig. 87D), but this is spurious. In Coleoptera the anterior part of the
lateral process is not the subalare, but is the postnotum.
The metasternum: The definitive metasternum in Halictophagus (Fig. 164B) is the
area proximal to the membranous cleft along the episternum, the pleurosternal suture
being absent. The metasternum is sclerotized only in the posterior half, and the sclero-
tized area has the internally ridged median longitudinal suture (Fig. 164B). The paired
simple furcae have a narrow common base at the posterior end of the longitudinal ridge;
hence, there is no external division of the sternum into the furcasternum and the basi-
sternum. Neither the ventral nor the trochantinal process for articulation with the coxa
occurs; the firm union of the metapleuron (definitive metasternum) with the coxal base
occurred concomitant with the loss of these structures. In some Strepsiptera a faint,
oblique suture occurs on either side of the middle of the sternum.
The adult musculature: Ulrich (1930) described thoracic muscles in the male of a
Stylops sp. However, his description does not allow reliable homologizations of many
of the muscles he recorded. This is partly due to the somewhat peculiar musculature
in this order.
Phylogenetic relationships: The systematic position of Strepsiptera has been
disputed. Among more recent workers, J eannel ( 1945) assigned them a position near
Hymenoptera, and Ulrich ( 1943) assigned them near Coleoptera. Arnett's ( 1960)
inclusion of Strepsiptera in Coleoptera aroused opposition from Pierce ( 1964), who
maintained that Strepsiptera deserve the ordinal rank independent of Coleoptera.
Kinzelbach (in press) has pointed out, among other characters, the presence of
muscle t-p 10 and the absence of t-cx 6 in Strepsiptera which constitute synapomorphic
characters with the Panorpoidea (pp. 63, 68) and he was led to believe that Strepsiptera
are early offshoots of the Panoq,oidea, and therefore they are not, closely related to
Coleoptera. In Coleoptera t-p 10 is absent and t-cx 6 may be present or absent. On
the basis of these characters, therefore, Kinzelbach's conclusion is fairly well founded.
In any case, it does not seem appropriate to include Strepsiptera in the order Coleop-
tera, because at least in the thoracic structures Strepsiptera have a set of autapomorphic
characters, which include gradual incorporation of the hind coxa into the pleurosternum, ..
the caudally directed peculiar furca which simulates the ventral process, the reduction of
the forewing, .. the absence of the posterior notal wing process, the fusion of the meta-
thoracic postnotum with the first abdominal tergite (as in Hymenoptera), and the
presence of the tergal fissure on the metanotum. Furthermore, as already discussed,
some apparent similarities in the thoracic structures between Coleoptera and Strepsiptera
are spurious. A clearer establishment of a sister group relationship between the two
orders, if such exists, requires further studies.
aaxn Coleoptera the hind coxa may be immovably fused with the metepisternum, but the former always
remains recognizable .
.. Reduction of tbe forewing has occurred in some beetles.
THE SIPHONAPTERA
In the study of the thorax morphology of fleas, Snodgrass' ( 1946) study still stands
alone as a comprehensive work. Unless otherwise noted, therefore, the following
descriptions and discussions on the external structures are based on his study.
The neck region: The pronotum lies immediately behind the head, and its anterior
margin is overlapped by the head. The neck membrane is therefore extremely restricted
in extent. Yet, in Ctenocephalus (Wenk 1953) and Hystrichopsylla (Fig. 165A) the
lateral cervical sclerite connects the ventrolateral posterior margin of the head and the
anterior margin of the episternum near the lower anterior angle of the pronotum.
The prothorax: The prothorax in Siphonaptera consists of the dorsal pronotum
and the ventrolateral pleurosternum. In many fleas the posterior margin of the pronotum
carries the ctenidial processes (Fig. 165A).
The border between the pleuron and the sternum has become obliterated. In some
fleas an apparent pleural suture separates the propleural area into the episternum and the
epimeron. The pleurocoxal articulation in the prothorax consists, unlike in other insects,
of a pivot on the upper edge of the small basal foramen of the coxa and a receptive
acetabulum on the pleuron (Fig. 165E). On the broad posterior end of the pleuron lies
the spiracle-bearing plate, which links the prothorax and the lateral extremity of the first
phragma (Fig. 165A). The homology of this link plate is quite unclear, because it is
peculiar to Siphonaptera. The sternum bears, on its posterior margin, an apodeme with
paired projections (Fig. 165B).
The mesothorax: The mesonotum is a simple undivided plate and bears posteriorly
a flange and anteriorly a large, thin vertical phragma, upon which muscle fibers from the
postoccipital ridge are attached. The mesonotum has no trace of a sulcus, or ridge,
which divides the surface into the scutum, scutellum, etc.
In Pulex (Fig. 165C) the medially free, internal bar of the mesopleuron connects
the upper margin of the pleuron and coxal base. This bar therefore substitutes for the
function of the pleural ridge, which forms a process at its ventral end in other insects;
Snodgrass ( 1946) thought this bar was the detached pleural ridge; according to Wagner
( 1939), the detached pleural ridge is present in the mesopleuron of all fleas except
Sarcopsyllidae, Pulicinae, and the genus Corypsylla. In many fleas including Pulex
(Fig. 165C), therefore, the mesopleuron is not divided into the anterior episternum and
the posterior epimeron by the pleural suture. According to Fox ( 1941), however, the
ridge and the external suture for the ridge are present in nearly all fleas, although the
ridge and the suture are absent in Pulex, Corypsylla, and Corypsilloides. In Dactylo-
psylla, Opisodasys, and others an internal peglike process from the anterior thickening of
the lower tergal margin is secured in a pit of the pleuron on the supposed pleural ridge.
In Pulex (Fig. 165C), Ctenocephalides, and others the link plate between the meso-
thorax and the metathorax has its anterior attachment on the ventral posterior part of the
mesopleuron, at the base of the inner surface of the flange. The lower anterior part of
the mesopleuron in Ischnopsyllidae is, as Fox ( 1941) showed, bordered by an oblique
ridge, which reaches the ventral end of the pleural ridge. Fox ( 1941) thought that the
lower anterior area is mesosterna! (Fox labeled it the laterosternite), but the area is
probably the fused katepisterno-preepisternal area, and the ridge may represent the
ana pleural suture (Fig. 165C).
FIG. 165. Thoracic structures in Siphonaptera (from Snodgrass). A, lateral view of pro-
t4orax in Hystrichopsylla gigas; B, prothoracic pleurosternal plate in Hystrichopsyl/a gigas;
C, lateral view of mesothorax in Pulex irritans; D, anterior view of mesothorax in Pulex
irritans; E, base of left procoxa disarticulated from pleuron in Hystrichopsylla gigas; F,
mesosternum and sternal articulation of coxa in Pulex irritans; G, lateral view of metathorax
and first abdominal segment in Ctenocephalides felis; H, anterior view of metathorax in Pulex
irritans.
PRONOTUM
-----CTENIDIUM
PLEURAL SUTURE
MESOPLEURO
LINK PLATE J I
ANAPLEURALJ7 .-- C
RIDGE
PREEPISTERNUM
+ KATEPISTERNUM

COXA
MESOSTERN
ARTICULAR
SCLERITE ...,..._,'=....,
l____ E
FURCA
RIDGE
1 ABO. TERG.
LATERAL
ANTE COSTA METANOTUM
EPISTERNUM
COXA
PLEURAL PLEURAL
RIDGE SUTURE
383
D
2'ND PHRAGMA
-METANOTUM
STERNUM
H
In Pulex (Fig. 165F) the sternal area has a median longitudinal sclerotization, and
the membranous area along this sclerotized plate is probably the basisternum. An unique
feature appears to be the ventral process for coxal articulation, which arises from the
base of the basisternum (articular sclerite in Fig. 165F) and not from the base of the
furca. It is not certain, therefore, whether the ventral process is homologous with the
ventral process in other insects. The paired furcae, which arise from the posterior end
of the median longitudinal basisternal plate, have no association with the pleural ridge,
and unite at their ends in a vertical plate. The elongate foramen enclosed by the furcal
arms gives passage to the nerve connection between the mesothoracic and metathoracic
ganglia.
The metathorax: The metanotum in Ctenocephalides (Fig. 165G) is bordered
posteriorly by a flange that overlaps the first abdominal tergite; it forms a transparent
lobe on the posterior ventral margin, and overlaps the pleuron. The surface of the
metanotum has two transverse grooves, and on the lower part of each side lies a longi-
tudinal groove that connects the transverse grooves; they form ridges internally as in
Mallophaga. The anterior transverse suture is the antecostal sulcus that forms internally
the second phragma; the other two grooves have no relation to the tergal division in
winged insects. The mesonotum does not participate in the formation of the second
phragma. In some fleas the posterior notal ridge terminates laterally in an oval
expansion that overlaps and firmly clasps the upper end of the pleural ridge. The third
phragma is the anterior inflection of the first abdominal tergite.
The metapleuron of Ctenocephalides (Fig. 165G) is divided by the apparent pleural
ridge into the anterior episternum and the posterior epimeron. The lower part of the
episternum is usually strengthened by the submarginal thickening, and its ventral anterior
angle is united with the expanded anterior end of the sternum. A remarkable fact about
the epimeron is its larger size. In most species it has invaded the region of the first
abdominal segment, and overlaps the second segment. The expanded epimeron carries
on its dorsal margin the tracheal trunk of the first abdominal spiracle.
The pleural ridge (Fig. 165G) rests ventrally on the upper end of the lateral ridge
of the coxal rim; it is inflected from the pleural suture, and is never partially detached, as
in the mesothorax. If this ridge is homologous with the pleural ridge in other insects,
it is peculiar in that the ridge is well developed in the segment lacking wings.
The widened anterior part of the definitive metasternum is united laterally with the
episternum, usually without a line of demarcation between them (Fig. 165H). The
intercoxal arm of the sternum, which is probably serially homologous with the meso-
thoracic ventral process, articulates with the mesal basal margin of the hind coxa. The
sternum gives rise also to the paired apodemal processes, or furcae.
When fleas jump, the large metathorax sustains the principal upward and down-
ward thrust of the legs against the body, since the large hind legs unquestionably furnish
the principal leaping force. Jacobson (1940) showed that jumping in Aphaniptera is
attributable mainly to the articulation of the metacoxa with the trochanter-femur, and to
the action of the dorsoventral muscles attached to the trochanteral apodeme. Jacobson
pointed out that this mechanism resembles that of jumping Homoptera, and not that of
Saltatoria, in which the femur-tibia articulation is important in jumping.
The adult musculature: Jacobson (1940) studied the muscles concerned with
jumping in Ceratophyllus styx, Wenk ( 1953) showed cervical muscles in Ctenocephalus
canis, and Lewis ( 1961) described the thoracic musculature in Xenopsylla cheopis. In
Table XXIII muscles recognized by Lewis are homologized.
Despite the winglessness, t-14 is well retained in the meso- and meta-thorax. The
greatly reduced thoracic musculature in Siphonaptera is highly characteristic and difficult
to homologize.
TABLE XXIII
Thoracic musculature in Siphonaptera
Xenopsylla (Lewis 1961) Muscle designations
22 op-t 3
21b op-cv 2
21a op-cv 3
24 s 1 or 2
27 (I) t12or13
28 (II)
22 t 14
25
26
23 (I) t-s 1 ?
34 (III-2A) p-s 5?
38 (II-III) p-cx8?
43 (III) p-cx9
32 (I-II) s ?
33 (II-III) s 13
31 (I) s 16 ?
29 (II) s-ex 2
30 (III)
35-37 (I) s-ex?
39 (II) s-ex?
44 (III)
40 (II) s-ex?
45 (III)
41 (II) s-ex?
46 (III)
42d (II)
47d (Ill)
t-tr 1
42c (II)
47c (Ill)
p-tr 1 or p-tr 2
42b (II) s-tr 1
47b (III)
REMARKS (Table XXIII)
385
Remarks
1
2
3
4
5
6
6
6
6
( 1) 23 stretches between the first phragma and the lateral lobes of the posterior
inflection of the prothoracic pleurosternum.
(2) 34 extends between the metathoracic furca and the ventrolateral angle of the
second abdominal sternite.
( 3) 43 arises from the large area of the metepimeron and is inserted on the
posterior margin of the coxal base.
( 4) 32 stretches between the mesosterna! furca and the posterior face of the pro-
sternal inflection.
( 5) 31 is a transverse muscle, which stretches across the "trough" in the prosternal
inflection.
( 6) Homologies of these muscles are quite Unclear.
THE ODONATA
The dragonfly thorax has evolved along peculiarly specialized lines, while main-
taining some primitive features. The mesothorax and the metathorax are fused by a
peculiar development of thoracic parts to form a single functional unit (synthorax).
The neck region: The main cervical sclerites are three lateral cervical sclerites
(Fig. 166A, B), which Asahina (1954) called Cv 1, Cv 2, Cv 3, respectively. Cv 1 is
often J-shaped and articulates anteriorly with the occipital condyle. Cv 1 and Cv 2 are
often fused, and together they are called the eucervicale. Cv 3 is the postcervicale of
some authors which embraces the base of the eucervicale. When the postcervicale is
well developed, it articulates with the pronotal margin and the anterior margin of the
pleuron. When the postcervicale is not well developed, as in Davidius (Fig. 166A),
it articulates only dorsally with the anterolateral angle of the pronotum.
The postcervicale ( cv 3) appears to represent the detached anepisternum, since it
lies dorsal to the katepisternum and below the anterior pronotum. Essentially this was
the view held by Crampton (1926a), who thought the postcervicale in Prophlebia was
the precoxale.
Since the eucervicale (Cv 1 and Cv 2) is separated posteriorly from the detached
anepisternum ( Cv 3) by a suture that probably represents the ana pleural suture (Fig.
166A), and the eucervicale lies in front of the katepisternal area, the position of the
eucervicale relative to the neighboring areas agrees with the lateral cervical sclerite in
Micropteryx (Fig. 144A). The eucervicale in Odonata is, therefore, preepisternal
in origin, as in many holometabolous insects and in Lepismachilis.
Additional small cervical sclerites ( Cv 4, Cv 5) often occur on the dorsal and
dorsolateral cervical membrane (Fig. 166A) . The ventral cervical sclerite also occurs
in some forms.
The pronotum: The pronotum (Fig. 166D) is typically differentiated into the
anterior, median and posterior notal lobes, separated by two furrows where ridges are
formed internally. The first protergal apophysis (Fig. 166C, D) arises from the median
pit of the anterior furrow; the presence of the apophyses is peculiar to Odonata, as will be
discussed more fully later (p. 388). In nearly all Odonata the median lobe has a median
longitudinal sulcus, which connects the anterior and posterior furrows. In Epiophlebia
the median and posterior lobes are not well differentiated and this condition probably
represents a primitive condition in Odonata.
The propleuron: The propleuron is fused with the pronotum along a complete or
incomplete notopleural suture (Fig. 166B). The pleural suture is often distinct above
the pleurocoxal articulation. The sclerotized episternum in Epiophlebia and Davidius
(Fig. 166A) and Onychogomphus (Fig. 166E) is the katepisternum, which is enlarged
proximally. As in some holometabolous insects, it is the katepisternum that reaches the
anterolateral area of the basisternum in Odonata. In many Odonata the membrane
separating the katepisternum from the basisternum is lost, owing to sclerotization.
The paracoxal suture and the trochantin are always absent in the prothorax of
Odonata. The postcoxal bridge is not formed.
The prosternum: In most Odonata the basisternum is a large, subquadrangular plate.
In a Gomphus sp. observed, the anterior margin of the basisternum is secondarily
thickened (prepectus), as in some other orders. In Hemianax Hakim (1963) showed a
clear transverse suture that separates the basisternal plate into anterior and posterior
parts; he called the anterior part the presternum. In Ictinogomphus (Hakim 1963) the
anterior plate is narrow, and appears to be the prepectus. The border between the
basistemum and the furcasternum is often represented by the anteriorly produced sterna-
costa which connects the furcal pits (Fig. 166E). The paired furcae (Fig. 166C) are
conspicuous. The spinasternum (postfurcasternum of Crampton, 1926a; intersternite
of Asahina, 1954, and Hakim, 1963) is either paired or a single continuous plate. It is
connected mesally with the posterior margin of the furcasternum, and its lateral ends are
connected with the anterior margin of the mesosternum. According to Asahina ( 1954),
NOTOPLEURAL
SUTURE
EPIST
DORSAL CERVICAL
SCLERITE
TERGAL
APOPHYSEAL PIT
Cv 1
D
E
STERN A COST A
Cv 3 (ANEPISTERNUM)
- EPIMERON
ANAPLEURAL
SUTURE
KATEPISTERNUM
387
MEDIAN LOBE
of PRONOTUM
TERGAL
APOPHYSIS 1
EPIMERON
FURCA
BASI STERNUM
VENTRAL
CERVICAL SCLERITE
EUCERVICALE
BASISTERNUM
POSTCERVIC
ALE (ANEPI
STERNUM?)
PIT
FURCASTERNUM
FIG. 166. A, lateral view of prothorax and neck region in Davidius nanus (from Asahina);
B, lateral view of prothorax and neck region in Onychogomphus ardens (from Chao); C,
posterior view of prothorax in Onychogomphus ardens (from Chao); D, dorsal view of pro-
thorax in Onychogomphus ardens (from Chao); E, ventral view of prothorax in Onycho-
gomphus ardens (from Chao).
the prothoracic spina is absent in Odonata, although Crampton (1926a) claimed it
was present in Epiophlebia.
The meso- and meta-notum: In Odonata, as a result of the dorsal extension of the
mesepisternum (Fig. 167 A), the anterior part of the mesonotum is displaced anteriorly.
This isolated mesonotal plate bears the second tergal apophysis internally. Asahina
traced the gradual detachment of this sclerite from the rest of the mesonotum during
postembryonic development, and interpreted it as the acrotergite. As Tannert's (1958)
study clearly showed, the anterolateral angle of the mesonotum represents the, anterior
notal wing process. Therefore, it is probable that the area that corresponds to the
prescutum in other insects must be in the anteriorly detached mesonotal plate
(Fig. 167A).
In the literature of odonate morphology (Grandi 1947a; Chao 1953; Asahina 1954;
Hakim 1964), however, the prescutum is the anteriormost part of the mesonotum that is
demarcated by a transverse suture. The suture passes mesally the point of invagination
of the third tergal apophysis, and its lateral end lies on the mesonotal lateral margin
behind the anterior notal wing process. Clearly, therefore, the suture is not homologous
with the prescutoscutal suture, and the supposed prescutum is not homologous with the
prescutum in other insects. They are the pseudoprescutum and the pseudoprescutoscutal
suture, as discussed below.
It appears probable that the tergal apophyses in Odonata are homologous with the
pseudophragmata in Lepismachilis (Machilidae), because both in Odonata and Lepis-
machilis the invaginations are intrasegmental, and they are obviously not homologous
with the phragma in other pterygote orders and the endonotum in Lepisma. Further,
muscles t-s 10, t 18, and t 19 associated with the tergal apophysis in Odonata agree in
their positions with the muscles associated with the pseudophragma (pp. 60, 66).
These facts support the homologization of the tergal apophysis with the pseudophragma.
As already discussed (p. 104), the pseudophragma in I apygidae is traversed
externally by the pseudoprescutoscutal suture, and the pseudophragma and the tergal
apophysis are probably homologous. Therefore, the transverse suture crossing the
tergal apophysis in Odonata is probably homologous with the pseudoprescutoscutal suture
in Japygidae, Dilta, and Nicoletia."" The anterior tergal area demarcated by the
pseudoprescutoscutal suture in Odonata is therefore probably the pseudoprescutum, as in
Japygidae, Dilta, and Nicoletia.
The mesonotum behind the pseudoprescutoscutal suture is the scutum. In some
dragonflies, such as Ictinogomphus (Hakim 1964) and Anax examined in this study, an
internally ridged, transscutal suture occurs. This suture lies immediately posterior to
the tergal apophyseal pit; it is not homologous with the transscutal suture in other
insects, since the latter represents a specialized condition of the membranous tergal
fissure, which is absent in Odonata. ,
The mesoscutellum consists of three parts: a mesal bulge and two lateral depressed
areas. The presumed scutoscutellar suture is not strongly produced anteromesally as in
other insects. The axillary cord extends from its posterolateral angle.
The large mesopostnotum is connected anteriorly with the mesoscutellum by a
narrow membrane; and its posterior margin is folded over the anterior margin of the
metanotum via a membrane, and in Zygoptera it is fused with the anterior margin of the
metanotum (Hakim 1964). In the context of Snodgrass' theory (p. 23), Mathur ( 1956)
thought that this isolated postnatal plate in Odonata represented a stage of evolution of
the postnatal plate that had been unrecognized. The postnatal plate does not form the
anterior wall of the second phragma by invagination of its posterior area; in this
respect, the mesothoracic postnotum in Odonata is peculiar, and resembles those in the
Apterygota.
The metathoracic pseudoprescutum is often clearly demarcated by the pseudo-
prescutoscutal suture, which reaches laterally the base of the anterior notal wing process
Sl"fhere is a possibility that this suture in Nicoletta is homologous with the prescutoscutal suture in
Lepisma (p. 104).
389
TERGAL APOPHYSEAL
PIT 2
MIDDORSAL
CARINA
COLLAR CARINA
EPISTERNUM
/:.
__ .-;:.._.::.. ../
ANTEALAR
ANTERIOR NOTAL
- PROCESS 2
l'ST AXIL. SCL.
LATERAL PROCESS
of ANT. NOT. PROC.
MESOSCUTUM-----__:::
MESOSCUTELLUM -----:=
MESOPOSTNOTUM -----

PSEUDOPRESCUTIJM
METASCUTUM---
METASCUTELLUM-----
METAPOSTNOTUM -----
ANTEALAR SINUS
t ANEPISTERNUM)
A
ANTEALAR CARINA
DISTAL HUMERAL
PLATE
SCUTELLOALAR
CONNECTION
ARTICULAR PROCESS
of l'ST AXIL. SCL.
FIG. 167. A, dorsal view of anterior mesonotum and mesopleuron in Anotogaster sieboldii
(from Asahina); B, dorsal view of right half of pterothorax in Aeschna mixta (from Tannert).
(Fig. 168), as in the mesothorax. The fourth tergal apophysis, which arises from the
median pit of the pseudoprescutoscutal suture on the metanoturn, is vestigial in Davidius,
and has disappeared in Orthetrum (Asahina 1954).
In Pseudagrion, Ictinogomphus, and Crocothemis, as Hakim ( 1964) showed, a
complete or medially incomplete transscutal suture occurs near the posterior margin of
the metascutum. The metascutellum, like the mesoscutellum, consists of the median
plate and the lateral lobes that bear the axillary cords. The posterior part of the meta-
thoracic postnotum extends laterally to meet the epimeron, thereby forming a narrow
postalar bridge (Hakim 1964). The posterior margin of the postnotum is united with
the first abdominal tergite.
TERGAL APOPHYSEAL
PIT
__ DOPRESCUTUM 3
.!:-J7 -
SCUTELLUM 3
FIG. 168. Dorsal view of mesopostnotum and metanotum in Ictinogomphus rapax (from
Hakim).
The wing base structures: The wing base,. consists of the proximal and distal
humeral plates and the axillary plate (Fig. 167B). The proximal humeral plate
(proximale Costa/platte of Tannert, 1958) is a detached proximal part of the C vein, and
corresponds to the humeral plate in other insects. The distal humeral plate ( distale
Costa/platte of Tannert) is a small intermediate plate that connects the proximal
humeral plate and the proximal ep.d of the C vein proper. The R-M vein base is
recognizable on the dorsal surface of the axillary plate (Radius-Analisplatte of Tannert);
and on its ventral surface lies the Cu-A vein base. The proximal humeral plate and the
axillary plate are connected by an intervening triangular plate.
In Odonata only the first axillary sclerite (Figs. 167B; 169A, C) was recognized by
Snodgrass (1909), Chao (1953), and Tannert (1958). Since, however, this first axillary
sclerite has no direct association with the base of the Sc vein and since the phylogenetic
position of Odonata still remains somewhat enigmatic (p. 398), this definitive first
axillary sclerite can not be homologous with the fiTst axillary sclerite in other insects.
It is almost fused with the anterior notal wing process, and its lateral hook articulates
with the proximal humeral plate. A second axillary sclerite is not differentiated in
Odonata. However, the connection of the axillary plate with the pleural wing process
suggests, as Chao (19 53) thought, that the second axillary sclerite in other insects must
correspond to a part of the axillary plate in Odonata. Similarly, the connection of the
axillary plate with a posterior notal wing process"' (Ventralausliiufer des Scutum of
Tannert, 1958) suggests that the third axillary sclerite in other insects corresponds to the
posterior part of the axillary plate. The homologies of the wing base structures in
Odonata with those in a palaeodictyopterous insect, Ostrava nigra, were already
discussed (p. 43).
The tergal articulations with wings: In the forewing the anterior scutoalar articula-
tion involves the anterior notal wing process, the first axillary sclerite, and the proximal
humeral plate. According to Tanneit (1958, 1961), they articulate among themselves
by processes, and each component is fused with others in varying degree in various
SOApplies both to the fore and hind wings.
8
7This .Posterior notal wing process is not necessarily homologous with the posterior notal wing process in
other msects.
HUMERAL PLATE
ANTERIOR
NOTAL PRO .. ~ : . ~ -
VENTRAL
PROCESS
A
c
INNER WALL of
1'ST AXIL. SCL.
VENTRAL PROCESS
of 1'ST AXIL. SCL.
PROCESS
t-p .22
CONNECTING LAMELLA
of 1 AXIL. SCL.-AXILLARY PLATE
391
ANTERIOR NOTAL PROCESS
B
TERGAL
APOPHYSIS 3
FIG. 169. Anterior scutoalar articulations in forewing (from Tannert). A, forewing base in
Calopteryx virgo; B, connections of tergal apophysis, anterior notal process, and first axillary
sclerite in forewing of Aeshna mixta; C, forewing base in Aeshna mixta; D, muscles inserted
. on humeral plate and first axillary sclerite in Anisoptera.
dragonflies (Fig. 169A, C, D). In Zygoptera the degree of fusion is more advanced
than in Anisoptera, or the degree of differentiation of the two areas is less marked in
Zygoptera.
Note (Fig. 169A, C, D) the lateral and ventral processes (Auslaufer) of the first
axillary sclerite, which articulate with the proximal humeral plate and with the anterior
notal wing process. In.Anisoptera (Fig. 1690) they receive steering muscles (t-p 11,
t-cx 4); in Zygoptera the ventral process is absent and t-p 11', instead of t-p 11 (see
p. 399), is attached to the tergal apophysis; t-cx 4 is, however, present. The proximal
humeral plate also receives two muscles (t-p 22, 23), as shown in Fig. 1690.
The posterior scutoalar articulation with the wing is the connection between the
ventral extension of the scutum (the posterior notal wing process?) and the proximal
392
margin of the axillary plate. The structural details and the points of muscle insertion
on the axillary plate are shown in Fig. 170A, B.
The scutelloalar articulation (Fig. 170A) consists of the connection between the
caudal end of the proximal margin of the axillary plate and the internal ridge of the
scutellum. Caudal to the scutelloalar articulation lies the chitinized, elastic, axillary
cord, which extends into the wing. It has a small sclerite at its distal end for the attach-
ment of muscles t-p 21 and t-p 25.
Hatch (1966) showed, in Aeshna, six points of connection between the mesonotum
and the forewing base as follows: Connection 1 is of flexible cuticle between the scutum
(prescutum of Hatch) and the anterior lobe of the proximal humeral plate. Connection
2 is of cuticle, between the scutum and the humeral plate. Connection 3 is a tough
ligament, from the lateral ridge of the scutum to the posterior end of the humeral plate.
Connection 4 is of flexible cuticle, between the third semidetached scuta! plate and the
anterior mesal part of the axillary plate. Connection 5 is a strong, flexible cuticle
connection between the posterior ridge of the scutum and the axillary plate. Force is
transferred from the tergum to the wing through a resilin ligament and a flexible over-
lapping extension of the axillary plate. Connection 6 is a flexible cuticle connection
between the hardened lateral extensions of the scutellum and the posterior end of the
axillary plate. Unfortunately, these connections were not compared with the connections
which Tannert (1958) described in detail. The pleural articulation with the wings is
discussed on p. 394.
The meso- and meta-thoracic pleuron: The extension dorsad of the mesepisternum
and the extension ventrad of the metepimeron have obscured the original structural
constitution of the pleuron of the synthorax. Nevertheless, parts of the pleuron can be
homologized with those of lower pterygote orders.'"
The intersegmental suture between the meso- and meta-thorax on the pleural surface
(interpleural suture) is complete in Calopterygidae (Fig. 171A), although it is incomplete
even in an otherwise highly primitive genus, Hemiphlebia (Tillyard 1917; Asahina
1957). In two species of the Mesozoic dragonflies, Liassoph/ebia magnifica and a
supposed anisozygopteron, Asahina ( 1957) showed a complete or nearly complete inter-
segmental suture, which, in a living anisozygopterous genus, Epioph/ebia, is retained
only ventrally. As a result of the reduction of the intersegmental suture, the mesepimeron
and the metepisternum are fused in varying degree in most dragonflies.
The course of the mesothoracic pleural suture in Odonata is more or less crooked,
with its lower portion nearly perpendicular to the longitudinal axis of the body. Its
dorsal end, as Grandi (1947a) and Tannert (1958) showed, splits characteristically into
the anterior and posterior heads (processes). Needham and Anthony ( 1903) defined
the degree of skewness (obliquity) as the acute angle between the suture and the
imaginary line perpendicular to the longitudinal axis of the body, and defined the angle
of tilt of the wing bases as the acute angle between a line drawn through the wing bases
and the longitudinal axis of the body. Since then, various authors have measured the
degree of skewness and the angle of tilt of various dragonflies. It can be safely said that
the greater the skewness the greater the degree of specialization of the pleural suture,
because the pleural suture must have been nearly straight and perpendicular to the longi-
tudinal axis of the body primitively (Chao 1953), as seen in some lower pterygote insects.
The paracoxal suture, which reaches or nearly reaches the pleural suture, is present
in the metepisternum (Fig. 171A). In the metathorax the anapleural suture is a trans-
verse suture that passes through the spiracle, which lies more ventrad than in most other
orders. As in other insects, the paracoxal and anapleural sutures are nearly confluent on
the pleural suture. The areas that are defined by these sutures are, therefore, the
katepisternum, the preepisternum, and the anepisternum (Fig. 171A). Crampton
(1926a) and Asahina (1954) pointed out the presence of the presumed trochantin as
88
Many sutures on the pleuron, as already discussed, are homologous throughout the entognathous Apterygota,
the Thysanura, and the lower Pterygota. Therefore, despite the obscure phylogenetic position of Odonata,
it is safe to assume that correspondences in the positions of the sutures in Odonata with those in other
winged insects provide a basis for homologization.
AXILLARY
PLATE
POSTERIOR SCUTOALAR

PROXIMAL RIDGE of
AXILLARY PLATE
CONNECTING LAMELLA of
SCUTUM -AXILLARY PLATE
POSTERIOR HEAD
of PLEURAL
WING PROCESS
ARTICULAR PROCESS
t-p
c
R +M
D
SCUTAL VENTRAL
PROCESS
393
FIG. 170. A, posterior scutoalar articulation in Aeshna mixta; B, scutelloalar articulation in
forewing of Aeshna mixta; C, connection of posterior head of pleural wing process with
posterior cap tendon in Cordulia aenea; D, tip of anterior head of pleural wing process in
Cordulia aenea. All figures from Tannert.
minute sclerites (trochantinelles) in the membrane above the coxal base during the
larval stage.
In the mesepisternum the anapleural suture reaches the pleural suture (Fig.
171A, B). The paracoxal suture is absent. The area ventral to the anapleural suture
must be the preepisternum that has become fused with the katepisternum, although
Asahina (1954) and Matsuda (1960a) interpreted the area as wholly katepisternal.
The anteriorly isolated small plate bearing the spiracle (Fig. 171B) is a secondarily
differentiated part of the preepisternum, which is often called the stigmal plate. This
area is membranous in Epiophlebia (Asahina 1954) and sclerotized in others; it is com-
parable with the prepectus formed on the anterior margin of the mesopleuron in
other orders.
Especially during the final molt into the adult, according to Sargent ( 1937), the
metepimeron greatly expands ventrally, thereby pushing the metasternum backwards.
The metasternum moves caudad through successive molts, away from its normal position
near the coxa, while the metepimeron expands downwards and fills this space (Sargent
19 3 7) . It is probable that the paracoxal suture on the epimeron follows this expansion
of the ventral epimeron. A short suture from near the ventral end of the pleural suture,
which is displaced ventrocaudally, to the median longitudinal axis (Fig. 171D) may well
be the displaced paracoxal suture, and the narrow ventral area demarcated by the para-
coxal suture is the katepimeron, which forms a postcoxal bridge. The large ventral
surface immediately behind the metathoracic katepimeron is thus the displaced
anepimeron, which may meet the counterpart on the other side of the pleuron along the
ventral median longitudinal suture (e.g., Pseudagrion, Crocothemis, Hakim 1964). A
secondary suture or sutures may subdivide the posterolateral part of the expanded
anepimeron.
The dorsally expanded mesothoracic anepisternum, which is formed by marginal
growth of the episternum (Sargent 1937), meets the counterpart on the other side of the
pleuron along the middorsal longitudinal carina (Fig. 167A). Asahina (1954) sus-
pected that the imaginal middorsal line might be tergal in origin.
89
The antealar carina
(Fig. 167A, B) on each side, which is purely an imaginal structure (Sargent 1937),
reaches the caudal end of the middorsal longitudinal carina. The triangular anepisternal
space anterior to the antealar carina is the antelar sinus.
Although Snodgrass (1935) believed that there are no epipleurites (basalare and
subalare) in Odonata, Forbes (1943), Chao (1953), Asahina (1954), and Hakim
( 1964) homologized the anterior and posterior caplike tendons with the basalare and
subalare, respectively. The homologies proposed by these recent workers, however,
require careful reexamination. The posterior cap tendon, which was homologized
with the subalare, is an internal structure in Odonata, and is directly continuous with the
R + M vein and the pleural wing process, whereas the subalare is always exposed in other
insects. The basalare, with which the anterior tendon was homologized, can be hidden
beneath the anepistemal wall in some insects (e.g., Neuroptera) as it is in Odonata, but
the muscles (t-p 22, t-p 23) attached to the anterior tendon in Odonata are not
homologous or not necessarily homologous with those attached to the basalare in other
insects and the cap tendon itself is connected with the humeral plate (Fig. 169D).
These facts lead the writer to conclude that these tendons are probably not homologous
with the subalare and the basalare in other winged insects. A study on the development
of these tendons will throw more light on this confused problem.
The pleural articulation with wings: The anterior head of the pleural wing process
is directly connected with the proximal humeral plate that lies above, and this connection
forms the anterior pleuroalar articulation. The ventral side of the humeral plate fits in
t):le groove formed on the apex of the anterior head (Fig. 170D). The posterior
pleuroalar articulation is the articulation of the posterior head of the pleural wing process
with the articular process of the R-M vein (Figs. 170C, 172B); the posterior cap tendon
SDCarpentier a1;1d Carpentier's (1952) study on fossil meganeurids (Protodonata) revealed that the mesepisterna
on the two sxdes do not extend far enough dorsally to meet each other. This finding tends to support
Asahina's contention.
ANEPISTERNUM 2
INTERPLEURAL
SUTURE
SPIRACLE
/
ANAPLEURAL
SUTURE
BASISTERNUM 1
FURCAL PIT
ANEPIMERON
SPIRACLE
PARACOXAL
SUTURE 3
A
c
395
INTERPLEURAL
SUTURE
PLEURAL SUTURE 3
KATEPISTERNUM 3
FURCAL PIT
FURCAL PIT 2
KATEPIMERON 3
POSTSTERNUM
PIT of PRE EPISTERNAL
APODEME
FIG. 171. A, lateral view of thorax in Mnais strigata (from Asahina); B, lateral view of
pterothorax in Onychogomphus ardens (from Chao); C, ventral view of thorax in nymphal
Davidius nanus (from Asahina); D, ventral view of thorax in Mnais strigata (from Asahina).
extends from the articular process of the R-M vein (Fig. 170C). These pleural
articulations form a fulcrum for the movement of wings. According to Hatch (1966),
they are made elastic by the pad of resilin, a protein rubber discovered by Weis-Fogh
(1960).
The meso- and meta-sternum: In the fully grown nymph of Davidius (Fig. 171C)
the pleural element occupies a considerable area between the furcal base and the inner
coxal margin, and the ventral process that articulates with the coxa can be katepimeral in
origin. The sternacosta connects the furcal pits in the mesothorax.
In the adult (Fig. 171D) the much narrower area between the furcal base and the
inner coxal margin can still be safely regarded as pleural in origin, since the corresponding
area in the nymph is pleural. The ventral process becomes lost in the adult. In the
mesothorax the division of the narrow sternum into the basisternum and the furcasternum
is often absent or obliterated, although in Pseudagrion (Hakim 1964) the horizontal
sternacosta connects the furcal pits. The basisternum is widened anterolaterally, and
meets the ventrally extended katepisterno-preepisternal area.
The border between the mesosternum and the metasternum has become completely
obliterated, the mesothoracic spina being always absent. In Mnais (Fig. 171D) the
metathoracic basisternum has extended considerably forward between the mesocoxal
bases; in Epiophlebia the shift forward of the basisternum is less pronounced than
in Mnais.
The metafurcasternal area is generally even narrower than in the mesosternum, and
is represented by a median longitudinal suture, from which furcae arise internally.
Among the species Asahina (1954) studied, only Mnais (Fig. 171D) has clearly
separated furcal pits. As already noted, the displaced posterior part of the sternum
(poststernum of Cowley, 1941 and Asahina, 1954) lies behind the ventrally invaded
katepimeron. In Pseudagrion and Crocothemis (Hakim 1964) the poststernum
(anepimeron of Hakim) is clearly separated from the broad ventral anepimeral area,
which has a median longitudinal suture.
Concomitant with the shift forward of the metathoracic basisternum, the intrusion
of the katepimeron between metathoracic coxal bases, and the displacement of the post-
sternum during development, the hind leg bases come to lie immediately behind the
middle leg bases in adult Odonata. According to Carpentier and Carpentier ( 1952), a
fossil species, Meganeurula conjusa, resembles the nymphal Anisoptera more than it
does the adult dragonflies in that the hind leg bases are widely separated from each other
and from the middle leg bases.
The pleural and sternal internal skeletons: The highly complex internal skeletal
system in the dragonfly thorax is associated with the musculature, which is unique.
According to Sargent (1937), all internal skeletal structures are formed from the adult
epidermis.
In Anisoptera the pleural apodemes as wing braces and pivots are formed upon the
meso- and meta-thoracic pleural sutures as cuticular infoldings (Fig. 172A). In general,
the shape of apodemes with cross bars somewhat resemble a structural steel I-beam.
Sargent called the apodemes in Aeschnidae the scalariform type, because of their resem-
blance to a ladder. Correlated with the formation of this type of apodeme, the pleural
suture is straight in Aeschnidae. In Libellulidae the apodemes are all well developed,
but without cross bars, and the sigmoidal curving of the mesopleural suture is strongly
marked.
What Asahina ( 1954) called the preepisternal apodeme in the mesothorax (Fig.
172C) is an invaginated disc from the preepisternum; in Epiophlebia it is large and
disclike, giving a point of attachment for the anterior cap tendon muscle ( t-p 23) . The
posterior serial homologue of this apodeme in Epiophlebia arises from a pit that lies on
the intersegmental area between the mesothorax and the metathorax, at the ventral end
of the preepisternal plate (Asahina 1954).
The prefurca (Fig. 172C), in both the mesothorax and the metathorax, arises from
the lateral corner of the basisternum or along the anterior margin of the coxal cavity; a
BASISTERNUM 1
BASISTERNUM 2
FURCA
KATEPIMERON 3
t-s 1
s 19?
ARTICULAR
PROCESS of R + M
SCUTAL VENTRAL
PROCESS
POSTERIOR HEAD of
397
POSTERIOR
CAP-TENDON
FURCA 1
PREEPISTERNAL APODEME 2
PREFURCA 2
PRE EPISTERNAL
APODEME 3
PREFURCA 3
s 22
Fro. 172. A, pleural endoskeleton in Aeshna cyanea (from Russenberger and Russenberger);
B, pleural and posterior scutoalar articulations in Odonata (from Mickoleit); C, internal view
of thorax in Epiophlebia superstes (from Asahina).
tergosternal muscle, t-s 10, is inserted on this apodeme. The prefurcai apodeme is
probably homologous with the endosternal tendon h in Lepismachilis and other Thysanura
(Fig. 37), which also lies on the lateral margin of the basisternum. The furcae (Fig.
172C) arise from the furcal pits in the mesothorax. The metathoracic furca is not well
differentiated. According to Asahina (1954), the metafurca is supposed to occupy the
anterior portion of the long, reflexed ridge of the metathoracic katepimeron. All of the
sternal internal skeletons together tend to form a more or less continuous floor of the
pterothorax.
398 MEMOffiS OF THE ENTOMOLOGICAL SOCIETY OF CANADA
Phylogenetic relationships: As already discussed (p. 43 ), the wing base structures
in Odonata have nothing to suggest a close phylogenetic relationship with Ephemeroptera.
The same is true with regard to the . musculature (p. 399). Homologous structures"'
which occur in Machilidae, Diplura and Odonata, however, include: the tergal apophysis
in Odonata, which is homologous with the pseudophragma in Japygidae and Machilidae;
the pseudoprescutoscutal suture in Odonata, which occurs also in Nicoletia, Dilta, and
Japygidae; the prefurca in Odonata, which is homologous with the endosternal process,
h, in Thysanura and Diplura; the muscles (t 18, t 19, t-s 10) associated with the tergal
apophysis and the prefurca, which are homologous with muscles occurring in the corres-
ponding areas of Lepismachilis; and the lateral cervical sclerite, which is a modified
preepisternum in Lepismachilis and Odonata.
The presence of the above mentioned characters in these groups is probably
symplesiomorphic. Yet, it suggests that Odonata could have arisen from a Machilidae
or Japygidae-like ancestor. It is probable that the strong autapomorphic trend in
Odonata has obscured their phylogenetic relationship with the Machilidae and Diplura.
This idea is strengthened further by the fact that these plesiomorphic characters are
absent in the rest of the Pterygota and Lepismatidae (except for the endosternal process h
in Lepisma) and that no sister group relationship of Odonata with any order of the
Pterygota is evident."'
The anatomical basis of flight: Besides the peculiarities in the wing base and its
associated structures already mentioned, the unique flight muscles recognized by Tannert
(1958) include (Figs. 169B, D; 170A; 172B): M. depressor anterior (t-p 23), M.
depressor median us (t-p 24), M. depressor posterior (t-p 25), Regia anterior musculi
elevatoris (t-s 10), Regia posterior musculi elevatoris (t-s 11), M. gubernator costae
(t-p 22), M. gubernator processus ventralis pt. 1 (t-p 11), M. gubernator processus
lateralis pt. 1 (t-cx 4), M. gubernator r+m-c (t-cx 4'), M. gubernator ana/is la, b
(t-p 14?), M. gubernator analis 2 (tp 21), and M. tensor latera/is (t-p 11').
These muscles are well defined by their actions in two categories: those concerned
with elevation and depression, and those responsible for supination and pronation.
Among the depressors, the anterior depressor ( t-p 23) is attached dorsally to the anterior
cap tendon, which is in turn connected with the median area of the humeral plate (-Fig.
169D). The median depressor (t-p 24) is attached to the articular process of the R+M
veins through the posterior cap tendon (Fig. 172B), and the posterior depressor (t-p 25)
is inserted on the posterior margin of the axillary plate (Fig. 170A). The contraction
of these muscles directly causes depression of the wing, because of their more lateral
positions.
The anterior elevator ( t-s 10) is attached dorsally to the base of the tergal apophysis
(Fig. 169B), and the posterior elevator ( t-s 11) is attached to the ventral process of the
scutum (Fig. 170B). The contraction of these muscles indirectly causes elevation of
the wing, since they lie proximal to the depressors.
Neville ( 1960) showed experimentally that t-cx 4 (anterior coxoalar muscle of
Neville) is a supinator during the last part of the downstroke and during the upstroke;
t-p 21 (third subalar muscle of Neville) is a supinator; and downstroke pronation
supination balance is controlled by t-p 22 (second basalar muscle of Neville), t-p 23 (first
basalar muscle of Neville) and t-p 25 (second subalar muscle of Neville). Of these last
three muscles, t-p 22 and t-p 23 cause the large pronation during the last part of the
normal upstroke and during the downstroke, and t-p 25 causes supination control of the
pronation resulting from the action of t-p 22 and t-p 23 during the forewing downstroke.
In Zygoptera, according to Tannert (1958), the forewing sinks deeply into the
thorax in the resting position, and the inner margin of the humeral plate comes to lie
vertically on the anterior head of the pleural wing process (Fig. 170D); this is performed
by the action of the two elevators of the wing (ts 10, t-s 11), and t-p 11' functions as a
OOBased on moiphological correspondences.
91
This presumed phylogenetic position of the Odonata supports the idea of the diphyletic origin of tho
wing (p. 42).
399
lateral support of the wing base, because of its connection with the pleural wing process
and t h ~ tergal apophysis.
In the forewing of Zygoptera, muscle t-p 11, which is inserted on the ventral process
of the first axillary sclerite in Anisoptera, has become t-p 11', which is inserted on the
tergal apophysis. Tannert ( 1958) thought that this change in the point of muscle attach-
ment is presumably the reason why Zygotera are, as Magnan ( 1934) found, slower in
movement than in Anisoptera. In the hind wing of Anisoptera and Zygoptera t-p 11',
instead of t-p 11, is present. Tannert (1958), therefore, supported Magnan's (1934)
hypothesis that in Anisoptera the forewing and the hind wing function differently. In
anisopterous dragonflies Neville (1960) proved experimentally that a constant phase
difference exists between the contractions of the mesothoracic and metathoracic flight
muscles, and also a resulting phase difference between the movements of the wing base
sclerites and the wings in the two pterothoracic segments.
Despite some functional specializations, both wings are still highly efficient flight
organs. Odonata are indeed remarkable, as Tannert ( 1958) pointed out, in that they
have developed such an efficient flight mechanism without sacrificing the function of one
, of the two pairs of wings (usually the hind wing) by reduction. It should be remembered
that the highly efficient flight in higher Diptera, for instance, is accompanied by the
nearly total reduction of the hind wing.
Russenberger and Russenberger's ( 1959-60) work was concerned primarily with
the mechanics of flight in dragonflies, and Weis-Fogh (1964) discussed the effect of
respiration on flight in dragonflies.
The adult musculature: Earlier works on the adult thoracic musculature in Odonata
since Chabrier ( 1820-1822) include Poleta!ew ( 1881), Lendenfeld ( 1881), Luks
(1883}, Amans (1883, 1884), Calvert (1893}, and Berlese (1909b). More recently,
Tillyard (1917) showed thoracic muscles in Sympetrum, and Marcus (1920),
Heidermann ( 19 31), and Cremer (19 34) made anatomicophysiological studies on
(
thoracic muscles in Crocothemis servilla and Psolodesmus mandarinus. Clark (1940)
studied the anatomy in anisopterous dragonflies. Grandi (1947a) described muscles in
Calopteryx splendens. Asahina ( 1954) described and discussed the thoracic musculature
in Odonata. Tannert's ( 1958) study dealt with flight muscles in representatives of
Zygoptera and Anisoptera. Russenberger and Russenberger's (1959-60) study dealt
mainly with the physiological aspect of flight muscles in Aeschna cynea. Neville ( 1960)
studied the function of direct flight muscles in anisopterous dragonflies. Hakim ( 1963,
1964) described thoracic muscles in adult Odonata. Hatch's (1966) study dealth with
the function of flight muscles. Homologies of muscles recorded by Asahina ( 1954),
and some additional notes, are given in Table XXIV.
Of the thoracic muscles in Odonata, the occurrence of cv-cv?, t 18, t 19, t-p 21,
t-p 22, t-p 23, t-p 24, t-p 25, t-p 26, t-s 10, t-s 11, s 19?, s 21, s 22 is confined to Odonata.
The muscle identified as t-p 14 may also be peculiar to Odonata. Muscle t-cx 4 occurs
only in Odonata and Ephemeroptera. The occurrence of such a large number of unique
muscles is coincidental with the unique skeletal structures. Some common flight muscles
in other winged insects (including Ephemeroptera), such as t 14, t-p 5, 6 and p 3, are
absent in Odonata. The transsegmental distribution of the dorsal muscles is apparently
a reduced version of the dorsal thoracic musculature in the Apterygota.
Gross aspects of the postembryonic development and histology of muscles:
Maloeufs (1935a) study on the postembryonic development of thoracic muscles in
Plathemis lydia revealed that no new muscle is formed during transformation into the
adult. Many nymphal muscles become lost later; these degenerating muscles can be
classified, as follows:
Nymphal muscles that are present in the adult of other Odonata: 3 (op-t 3),
9 (cv-cx 1(x)?}, 31 and 53 (t-p 26), 37 and 60 (t-cx 6 or 7), 42 (s 19}, 'and 65 and
66 (s 21).
TABLE XXIV
Thoracic musculature in Odonata
Odonata (Asahina1954) Muscle designations Remarks
1 op-t 1 or 2
3 op-t 3
4, 5 op-p 1 1
7 t-cv 2
6 cv-cv ?- 2
9, 10 cv-cx 1 or 2(x)
11, 11' s 1, 2
2 (I) t 18
25 (II) t 19
45 (III)
12 (I) t-p (proth.)
28 (II) t-p 11 3
50 (III)
M. tensor lat. (II, Ill) t-p 11' 4
29, 30 (II) t-p 14 ? 5
51, 52 (III)
34 (II) t-p 21
56 (III)
21 (I I) t-p 22
43 (III)
22 (II) t-p 23
44 (Ill}
32 (I I) t-p 24
54 (III)
33 (II) t-p 25
55 (II I)
31 (II)
53 (IV)
t-p 26
67 (Ill) t-s 1 ?, p-s 5 ? 6
23 (I I) t-s 10
46 (II I)
23' (II) t-s 11
46' (III)
13 (I) t-cx 2 or 3
26 (I I) t-cx 4 7
48 (III)
27 (II) t-cx 4' 8
49 (I II)
14 t-cx 6 or 7
37
60
36 (II) p-cx5
58 (III)
41 (I-I I) s 13
42 (64) s 19
66 (Ili-2A) s 21
68 (II I) s 22
16 s-ex 3
38
61
17 (I) t-tr 1
39 (II)
p-tr 1
62 (II I)
Od6nata (Asahina 1954)
18 (!)
19
40
63
REMARKS (Table XXIV)
TABLE XXIV (Contnued)
Muscle designations
p-tr 3
s-tr 1
401
Remarks
(1) These muscles connect the postocciput and Cv 3, which is probably a detached
part of the proepisternum. Therefore, they are regarded as op-p 1.
(2) 6 connects Cv 3 and Cv 4. If Cv 3 is a detached episternal part, the muscle
is a cv-p muscle. No such muscle occurs in other orders.
(3) According to Asahina (1954), 28 and 50 arise from a flexible plate, lateral to
the scutum (first axillary sclerite?), and are inserted on the lower portion of the pleural
wing process. These muscles are M. gubernator processus ventralis pt. 1 of Tannert
(1958), which extends between the base of the pleural wing process and the ventral
process of the first axillary sclerite.
( 4) M. tensor latera/is of Tannert, in the forewing of Zygoptera and in the hind
wing of Zygoptera and Anisoptera, is attached proximally to the tergal apophysis. t-p 11'
is, according to Tannert, homologous with t-p 11.
(5) These muscles are M. gubernator ana/is 1a, b of Tannert (1958), which
stretch between the caudal side of the pleural wing process and the anal-vein area of the
axillary plate. This muscle may be t-p 14 in other insects.
( 6) 67 connects the dorsal portion of the posterior ridge of the metathoracic
epimeron (Asahina 1954), or the antecosta (anterior to?) of the first abdominal segment
(Maloeuf 1935a) with the posterior end of the furcal invagination.
(7) These muscles are M. gubernator proc. lat. pt. 1 of Tannert (1958), which are
inserted dorsally on the lateral process of the first axillary sclerite. Asahina ( 1954)
regarded these muscles as coxobasalar muscles.
(8) These muscles are M. gubernator r+m-c of Tannert (1958), which are
inserted dorsally on the axillary plate. Asahina ( 1954) regarded them as coxosubalar
muscles.
Besides the muscles Asahina ( 1954) recorded, the following muscles are known to
occur: Ibm 1 (s-ex 5) in the prothorax of Anisoptera (Clark 1940), a ventral trans-
verse muscle (p-s 6) in the promesothorax of Crocothemis and Psolodesmus (Maki
1938), and an additional prothoracic tergopleural muscle in Crocothemis (Maki 1938).
Nymphal muscles that are present in the adult of orders other than Odonata:
6 (cervical t-s 1), 15 (t-cx 9?), 20 (promesothoracic t-s 1), 24 and 47 (t-p 3 or 4), 35
and 57 (p-s 1 ?) , and 45 (t 14?).
Muscles that are peculiar to the nymphal stage of Odonata: muscle 10 that connects
the prosternum and the cervical sclerite on the opposite side; 59 that arises from the
intersegmental sternopleural process (between the meso- and meta-thorax) and is inserted
on the apodeme arising from the anterior portion of the mesothoracic (?) pericoxal
membrane, and 69 that connects transversely the intersegmental pleurosternal process on
each side. In the future, these muscles may be found in the Apterygota.
According to Maloeuf (1935a), in a 14 mm long nymph of Plathemis lydia the
diameter of a single fiber of nonfunctional muscle is 4.8 p., it is unstriated, and has
numerous nuclei in the superficial sarcoplasm; a functional muscle fiber in a 14 mm long
nymph is 20.8 p. in diameter, is striated, and has elongated nuclei in the superficial
sarcoplasm. For information about histogenesis of flight muscles, see Holmgren (1910)
and Cremer ( 1934).
In the adul! all flight muscles are tubular, and myofibrils are radially arranged in
fibers of small dtameter. In Libel/ula sp. the diameter of the fiber is 20-25 p., and the
diameter of the fibril is 0.7 p. (Holmgren 1910).
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INDEX TO TERMINOLOGIES
PAGB
Acetabulum .. . . . . .. . . .. .. . . . . . . . . . . . . . . . . 209
Acrotergite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11, 12
Adana! pterale . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 347
Alacrista . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 211
Alar apophysis . . . . . . . . . . . . . . . . . . . 208, 212, 213
Alula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377
Anapleura1 ring . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Anapleural suture . . . . . . . . . . . . . . . . . . 33, 36, 39
Anapleuron . . . . . . . . . . . . . . . . . . . . . . . . . 28, 33, 35
Anepistemum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 36
Antealar carina . . . . . . . . . . . . . . . . . . . . . . . . . . . 394
Antealar sinus . . . . . . . . . . . . . . . . . . . . . . . . . . . . 394
Antecosta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11, 12
Antecoxale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37, 38
Antemedian notal wing process ....... 12, 13, 14
Antenotum ......................... 11, 12, 103
Anterior basalare . . . . . . . . . . . .. . . . . . . . . . . . . . . 36
Anterior condyle (for coxal articulation) . . . . 46
Anterior notal wing process . . . . . . . . . . . 12, 13, 14
Anterior subalare ..................... , . . . . 39
Anteroadmedian line .. .. . . .. . .. . .. .. . .. . . . . 361
Anterolateral scuta! suture . . . . . . . . . . . . . . . . . 13
Apotome ................... \....... . . . . . . . 18
Autapomorphic ............. \. . . . . . . . . . . . . 143
Axilla .................... \. .. .. . . .. .. .. 361
Axillary lever ............... .'. . . . . . . . . . . . . 362
Axillary plate .. .. .. .. .. .. .. .... .. .. . .. 15, 390
Axillary sclerites . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Basal fold . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Basalar apophysis . . . . . . . . . . . . . . . . . . . . . . . . . 290
Basalar cleft . . . . . . . . . . . . . . . . . . . . . . . . . . 151, 157
Basalar suture . .. .. .. .. .. .. .. .. .. .. . .. .. . .. 36
Basalare . . .. . . . . . . .. .. . . .. . . .. . .. . . . .. . . . . 36
Basanale . .. .. .. .. . .. .. . . . .. .. .. . . .. .. 150, 198
Basicosta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Basicostale .................... ; . . . . . . 197, 198
Basicoxite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Basijugale . . . . . . . . . .. .. . .. .. .. . .. .. . .. . . . 150
Basipodite (of Borner) . . . . . . . . . . . . . . . . . . . . . 29
Basiradiale . . . . . . . . . . . . . . . . . . 130, 160, 161, 197
Basisternum . . . . . . . . . . . . . . . . . . . . . 18, 19, 20, 37
Basisubcostale . . . . . . . . . . 128, 150, 160, 161, 194
Biogenetic law . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
Bulla . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 120
Caenogenetic muscles . . . . . . . . . . . . . . . . . . . . 81, 83
Cap tendon (anterior) . . . . . . . . . . . . . . . . . . . . . . 36
Cap tendon (posterior) . . . . . . . . . . . . . . . . . . . . 394
Cervical apodeme . . . . . . . . . . . . . . . . . . . . . . . . . 358
Cervical muscles (Pterygota) . . . . . . . . . . . . . . . 56
Cervical sclerites (origin and evolution) . . . . . 7
Cervical sclerites (theories) . . . . . . . . . . . . . . . . 9
Cervicopleuron . . . . . . . . . . . . . . . . . . . . . . . . . . . . 357
Cervicoprothoracic muscles (Apterygota) . . 76, 77
Cervicoprothoracic muscles (Pterygota) . . . 76, 77
Close packed muscles . . . . . . . . . . . . . . . . . . . . . . 86
Coxa .. .. .. .. . .. . .. ....................... 46
Coxal apophysis . . . . . . . . . . . . . . . . . . . . . . . . .. . 106
Coxal suture . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Coxopleuron . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
Coxosternal suture (of DuPorte) . . . . . . . . . . . . 39
Coxostemum (of DuPorte) . . . . . . . . . . . . . . . . . 39
Cryptopleuron . . .. . . . . .. . . .. .. . .. . 170, 202, 265
Cryptostemum . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
Czyptothorax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 135
Discrimen .... , . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
Distal median plate . . . . . . . . . . . . . . . . . . . . . . . 16
Dorsal cervical sclerite (origin) . . . . . . . . . . . . 8
Endonotum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Endostemal processes .. .. . .. .. .. .. . .. .. .. .. 108
............... : . . . . . . . . . . . . . . 17
Ep>meron . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38, 39
Episternal longitudinal suture . . . . . . . . . . . . 36, 39
Episte"!'um . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Eucerv1cale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386
Eucoxa . . . . . . . . . . . . .. .... .. ............... 27
Eupleuron (of Crampton) . . . . . . . . . . . . . . . 27, 39
Eutrochantin .......................... : . . 37
Fibrillar. muscles . . . .. . . . . . .. . .. .. . .. .. .. .. 86
First axillary sclerite . . . . . . . . . . . . . . . . . . . 15, 16
First basalare .......................... : . . 36
PAGE
Fourth axillary sclerite . . . . . . . . . . . . . . . . . .. . 14
Furca . . . . . . . . .. .. . . . . . . . . . . . . . . . .. . . . . . 17, 18
Furcal pit . . . . .. . . . . . . . . . . . . . . . . . . .. . . . . . . . 126
Furcastemum . . . . . . . . . . . . . . . . . . . . . . . . :iS, 19, 20
Furcilla .. . .. .. .. . .. .. .. .. .. .. .. . .. .. .. .. . 24
Gula (origin) .. .. .. .. .. .. .. .. .. . . .. . . . . .. .. 9
Halter . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 319
Heterochrony . . . . . . . . . . . . . . . . . . . . 81, 82, 84, 86
Humeral plate ...................... 16, 36, 389
Hypoepimeron . . . . . . . . . . . . . . . . . . . . . . . . . . 39, 379
Hypopteron . . . . . . . . . . . . . . . . . . . . . .. . . . . . 19, 39
InciSura submediana . . . . . . . . . . . . . . . . . . . . . . . 160
Intemotum . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19, 94
InterpleUrite . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19, 24
Intersegmental relationship . . . . 21, 23 (theories)
Intersternum (lntersternite) . . . . . . . . . . . . . . 19, 94
Intratergal apophysis .. . .. .. .. . .. .. . .. . .. . . 45
Intertergite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Jugulare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 287
Jugum ................. , . . . . . . . . . . . . . . . . . . 45
Katapleural ring . . . . . . . . . . . . . . . . . . . . . . . . . . 3 8
Katapleurocoxal articulation . . . . . . . . . . . . . . . . 25
Katapleuron . . . . . . . . . . . . . . . . . 26, 28, 29, 35, 38
Katepimeron . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35, 38
Katepistemum .......................... 37, 38
Lateral cervical sclerite (origin) . . . . . . . . . . . . 7, 8
Lateral parapsidal suture . . . . . . . . . . . . . . . . . . 13
Lateral plate ............................. , 342
Lateral postnotum . .. . . . . . . . . . . . . . . . . . . .. . 36
Lateropleurite .. .. .. .. . . . .. .. .. .. .. . . . .. 36, 39
Laterosternite . . . . . . . . . . . . . . . . . . . 24, 29, 38, 39
Laterotergite .. . .. . .. . .. . .. .. .. .. .. .. .. . .. . 24
Linea ventralis .............. , . . . . . . . . . . . . . 92
Link plate .. .. .. . .. .. . .. .. .. .. . .. . . .. . .. .. 382
Maxilliped .. . .. .. .. . .. .. .. .. .. . .. . .. .. .. .. 2S
Median notal wing process . . . . . . . . . . . . . . . . 14
Median plate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Membrana dorsalis . . . . . . . . . . . . . . . . . . . . . . 26, 32
Membrana ventralis . . . . . . . . . . . . . . . . . . . . . . . 27
Merocosta .. .. .. .. . .. .. .. . .. .. .. .. .. . .. .. . 46
Meron . . . .. ..... ................. ... ..... 46
Meropleurite .. .. . . .. .. .. .. . .. .. . .. .. . . .. . . 316
Metazona . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157
Multiphased morphogenesis . . . . . . . . . . . . . . . . 28
Myofibrill . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86
Myogenic rhythm . . . . . . . . . . . . . . . . . . . . . . . . . . 87
Neala . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Neck region (origin) . . . . . . . . . . . . . . . . . . . . . . 6
Neck region (theories) . . . . . . . . . . . . . . . . . . . . 9
Neogenetic muscles ................... 81, 83, 84
Notauli . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 361
Notopterale .. .. .. . .. . .. . . .. . .. . . . . .. . . . .. . 39
gccipital condyle . . . . . . . . . . . . . . . . . . . . . . . . . . 358
mphalium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279
Operculum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 261
Other muscles (Lepisma) . . . . . . . . . . . . . . . . . . . 56
Palaeogenetic muscles . . . . . . . . . . . . 81, 82, 83, 84
Paracoxal suture . . . . . . . . . . . . . . 33, 35, 36, 37, 38
Pparanotal theory .. . .. .. .. .. .. .. .. .. .. . .. .. 40
aranotum . . . . . . . . . . . . . . . . . . . . . . . . 88
furrow .............. : .. : : :: : :: 361
araps>dal suture . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Parascutellum . . . . . . . . . . . . . . . . . . . . . . . . . 39, 379
. . . .. . . . .. .. .. .. . . .. . . . . .. .. . .. 361
ii ... .. :....... . .. . . . . .. . . .. . . . . .. 342
ragma (ongm) .. .. .. .. .. .. .. 21, 22, 23 24
(Odonata) ................ '395
p a! apophysis . . . . . . . . . . . . . . . . . . . . . . . . . 291
leural arm . . .. .. . .. .. .. .. .. . . .. . .. .. .. .. . 35
Pleural cleft .. . .. .. .. . .. . .. . .. .. .. .. .. . .. . 36
condyle .. .. .. .... . .. .. .. . .. .. . .. .. .. 37
ural costa . . . . . . . . . . . . . . . . . . . . . . . . . . 35, 290
Pleural muscles (evolution) . . . . . . . . . . . . . . . . 80
Pleural muscles (Pterygota) . . . . . . . . . . . . . . . . 70
::ural P!'rts (homologies) . . . . . . . . . . . . . . . . . 33
eural pit .. .. .. . .. .. .. .. . .. .. .. .. .. . .. .. . 35
PAGE
Pleural ridge . . . . . . . . . . . . . . . . . . . . . . . . . . . 34, 35
Pleural suture .. .. .. .. .. .. .. .. .. .. .. .. .. .. 34
Pleural wing groove . . . . . . . . . . . . . . . . . . . . . . . 257
Pleural wing process .. .. .. .. .. .. .. .. .. .. . 34, 36
Pleuron (hypothetically primitive) . . . . . . . . . . . 35
Pleuron (origin) ...................... 24, 27, 28
Pleuropodite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
Pleurostemal muscles (Lepisma) ....... 51, 78
Pleurosternal muscles (Pterygota) . . . . . . . . 70, 78
Pleurotergite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Pleurotrochantin .. . .. . . . . .. . .. . . . . . .. .. . . . 316
Pleurotrochantinal (coxal) muscles (Lepisma) 55, 78
Pleurotrochantinal (coxal) muscles (Pterygota)
.. . . .. .......................... 72, 78
Pleurosternal suture . . . . . . . . . . . . . . . . . . . . . . . . 37
Podopleuron . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28, 40
Postalar bridge . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38
Postcervicale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 386
Postcoxal bridge . . . . . . . . . . . . . . . . . . . . . . . . . 30
Postcoxale . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Posterior basalare . . . . . . . . . . . . . . . . . . . . . . . . . 36
Posterior condyle (for coxal articulation) . . . . 46
Posterior notal wing process . . . . . . . . . . . . . . . . 13
Posterior subalare . . . . . . . . . . . . . . . . . . . . . . . . 39
Postfurcasternum . . . . . . . . . . . . . . . . . . . . . . . . 20, 39
Postmedian notal wing process . . . . . . . . . . . . 14
Posterior trochantino-coxal articulation . . . . . . 37
Posterolateral scuta! suture . . . . . . . . . . . . . . 13, 14
Postnotum .. .. .. . .. .. .. .. . .. .. .. .. .. 11 (origin)
Postphragma .. . . . . . . .. . . .. .. .. . .. .. . .. . . . . 11
Poststernum . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18, 20
Poststemum (Odonata) . . .. . . . . . . . . . . . . . . . . 396
Posttergite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11, 39
Prealar arm . . . . . . . . . . . . . . . . . . . . . . . . . . . 14, 36
Prealar bridge . . . . . . . . . . . . . . . . . . . . . . . . . . 14, 36
Prealar sclerite . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Prealare (prealar arm) . . . . . . . . . . . . . . . . . . . 12, 14
Precosta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 94
Precoxal suture . . . . . . . . . . . . . . . . . . . . . . . . . . . 35
Precoxal bridge ...................... 30, 37, 39
Precoxale . . . . . . . . . . . . . . . . . . . . . . . . . . . . 143, 169
Precoxopodite . . . . . . . . . . . . . . . . . . . . . . . 25, 26, 27
Preepimeron . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 350
Preepistemal apodeme . . . . . . . . . . . . . . . . . . . . . 396
Preepisternum . . . . . . . . . . . . . . . . . . . 30, 33, 36, 37
Prefurca .. . . .. . . . . . .. . .. .. .. .. .. .. . .. .. . 18, 45
Prepectus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Prephragma ................... ,. . . . .. . .. . . . 11
Prescutal arm ................. ; . . . . . . . . . . . 14
Prescutellar lobe . .. .. . .. .. .. .. .. .. .. .. .. .. 311
Prescutoscutal suture . . . . . . . . . . . . . . . . . . . . . . 12
Prescutum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
Presternum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18, 19
Presternum (Presternite 1) . . . . . . . . . . . . . . 97, 110
Presternum (Presternite 2) . . . . . . . . . . . . . . . . . 99
Pretergite . . . . . . . . . . .. .. .. . .. .. .. .. . . . . . .. 39
Promesonotum . . . . . . . . . . . . . . . . . . . . . . . . . . . . 91
Proximal median plate . . . . . . . . . . . . . . . . . . . . 16
Prozona . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157
Pseudofetus (Dilta) ................ 13, 103, 104
Pseudonotum . .. . .. . .. . .. . .. .. .. .. . .. .. .. . 207
Pseudophragma . . . . . . . . . . . . . . . . . . . . . . . . 11, 104
Pseudoprescutoscutal suture . . . . . . . . . . . . 97, 104
Pseudoprescutum . . . . ........... 97, 103, 104
Pseudopteralia (of Grandi) . . . . . . . . . . . . . . 45, 116
Pteropleuron . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Recurrent scutoscutellar suture ............ .
Resilin
Sarcolemma ............................. .
Sarcostyle ................................ .
Scutellar lever ........................... .
Scutellum ................................ .
Scutoscutellar suture ...................... .
Scutum ............ , ............... .
Second axillary sclerite .................. ..
Second basalare .......................... .
Secondary segmentation ................... .
14
16
154
153
362
12
13
12
16
36
23
PAGE
Spina ......................... 17, 18, .19, 20, 22
Spinasternum . . . . . . . . . . . . . . . . . . . . . . . . . . . 18, 20
Spiracles (positions and evolution) . . . . . . . . . . 46
Spiracles (theories) ...................... . 48
Stenothorax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 132
Sternacosta . . . . . . . . . . . . . . . . . . . . . . 19, 20
Sternal muscles (evolution) . . . . . . . . . . . . . . . . 81
Sternal muscles (Lepisma) . . . . . . . . . . . . . . . 49, 79
Sternal muscles (Pterygota) . . . . . . . . . . . . . . 73, 79
Sternellum . . . . . . . . . . . . . 20, 151, 157, 183
Sternocoxal muscles (evolution) . . . . . . . . . . . . 81
Sternocoxal muscles (Lepisma) . . . . . . . . . . . 51, 79
Stemocoxal muscles (Pterygota) . . . . . . . . . . 74, 79
Sternopleurite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 350
Sternum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Sternum (divisions of, Apterygota) . . . . . . . . . 18
Sternum (divisions of, Pterygota) . . . . . . . . . . . 19
Stink apparatus . . . . . . . . . . . . . . . . . . . . . . . . . . . 278
Stink groove . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278
Subalare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28, 29
Subcoxa .' ...................... 24, 26, 27, 28, 29
Subcoxal theory .......... , . . . 24, 25, 27, 28, 29
Substitution . . . . . . . . . . . . . . . . . . . . . . . . . . . 29, 86
Subtegula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Supracoxal lobe . . . . . . . . . . . . . . . . . . . . . . 209, 268
Suralare . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13, 39
Synapomorphic . . . . . . . . . . . . . . . . . . . . . . . . 141, 143
Synthorax . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21, 23
Tegula . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
Telopodite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
Tergal . . . . . . . . . . . . . . . . . . . . . . 386, 388
Tergal fissure . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Tergal muscles (evolution) . . . . . . . . . . . . . . . . . 79
Tergal muscles (Lepisma) . . . . . . . . . . . . . . . . 49, 77
Tergal muscles (Pterygota) . . . . . . . . . . . . . . 60, 77
Tergal sutures (Apterygota) . . . . . . . . . . . . . . . 11
Tergal sutures (Pterygota) ............... , . 12
Tergal wing groove . . . . . . . . . . . . . . . . . . . 254, 273
Tergocoxal muscles (Lepisma) . . . . . . . . . . . . . 53
Tergopleural apodeme . . . . . . . . . . . . . . . . . 337, 349
Tergopleural muscles (evolution) . . . . . . . . . . . . 79
Tergopleural muscles (Lepisma) . . . . . . . . . 55, 77
Tergopleural muscles (Pterygota) . . . . . . . . . 60, 77
Tergosternal muscles (Leplsma) . . . . . . . . . . 51, 78
Tergostemal muscles (Pterygota) ...... 65, 78, 80
Tergotrochantinal (coxal) muscles (evolution). 80
Tergotrochantinal muscles (Lepisma) . . . . . . 53, 78
Tergotrochantinal (coxal) muscles (Pterygota)
............................ 66, 78
Tergum (divisions of) .. . .. .. . .. .. .. .. .. .. .. 11
Tergum (modifications) . . . . . . . . . . . . . . . . . . . . 14
Third axillary sclerite . . . . . . . . . . . . . . . . . . . . !S, 16
Transanepistemal suture . . . . . . . . . . . . . . . . . . . 258
Transepimeral suture . . . . . . . . . . . . . . . . . 36, 38, 39
Transscutal suture . . . . . . . . . . . . . . . . . . . . . . . 13
Transscutellar suture . . . .. . . . . . . .. .. .. . .. . . . 312
T-ridge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
Trochanteral muscles (Lepisma) . . . . . . . . . 53, 79
Trochanteral muscles (Pterygota) . . . . . . . . . 75, 79
Trochantin ........................... 28, 37, 38
Trochantinocoxal articulation . . . . . . . . . . . . 25, 30
Tubular muscles .. .. .. .. .. .. .. .. .. .. . .. .. .. 86
Ventral cervical sclerite (origin) . . . . . . . . . . . . 9
Ventral condyle (for coxal articulation) . . . 38, 46
Ventrolateral cervical sclerite . . . . . . . . . . . . . . 287
Ventropleurite ....................... 29, 30, 38
Veracoxa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Verasternite . . . . . . . .. . . . . . . . .. .. . . . .. . . . . . . 173
Wing (origin) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Wing (prothoracic) . . . . . . . .. . .. . . . .. . . . . .. 40
Wing (types) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Xyphus
277
Y-ridge . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19, 20, 46
MATSUDA.; 1'RB lNSBCT 1'RORA.X 429
INDEX TO AUTHORS
PAGE
Abdul-Latif and Azlz Aslam Khan . . . . . . . . . . 262
Acker, T. S. . . .. .. .. .. .. . . .. . 285, 287, 288, 290
Ahrens, W. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 157
Akbar, S. S. . . . . . . . . . . . 9, 267, 270, 273, 280, 282
Alam, S. M. . . 357, 359, 361, 3q2, 366, 369, 370,
371, 374, 375
Albrecht, F. 0. . . . . . . . . . . . . . 173, 180, 183, 185
Alexander, R. D. and W. L. Brown . . . . . . . . . 41
Alicata, P. .. .. .. .. .. .. .. .. .. . 8, 57, 58, 169, 185
............................ m
Amzy, N. M. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
Ander, K. . ....................... 170, 183, 185
Ando, H. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
A rgi!as, A. . . . . . .. . .. . . . .. .. .. . .. .. .. .. . . .. 111
Arnett, R. H. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 381
Arora, G. L. . . . . . . . . . . . . . . . . . . . . . . . . . . 357, 361
Arora, G. L. and Asket Singh . . . . . . . . . . . . . . 183
Asahina, S. . . . 386, 388, 389, 392, 394, 395, 397,
399, 400, 401
Atkins, M. D. and S. H. Farris . . . . . . . . . . . . 218
Atzirtger, W. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 187
Badonnel, A. . . . . . . . 222, 224, 227, 230, 231, 232
Balfour-Browne, F. A. . ............ 213, 214, 216
Barber, S. B. and J. W. S. Pringle . . . . . . . . . . 282
Barlet, J, ... 7, 11, 17, 18, 24, 28, 48, 49, SO, 51,
52, 53, 54, 57, 72, 99, 103, 104, 106, 110, 111,
112, 113
Barlet, J. et F. Carpentier .. 11, 18, 19, 25, 26, 48,
94, 97, 99
Barth, R. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 86, 354
Bauer, A. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216, 220
Beardsley, J. W. . . . . . . . . . . . . . . . . . . . . . . . . . . 258
Bekker (Becker), E. G. . . . . . . 24, 30, 40, 42, 116,
120, 243
Bell, R. T. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Berlese, A. . .. 66, 88, 90, 216, 220, 261, 264, 354,
371, 399
Betten, H. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278
Bitsch, J ................. 304, 311, 312, 319, 321
Blackith, R. E. and R. M. Blackith . . . . . . . . !85
Blackwelder, R. E. . . . . . . . . . . . . . . . . . . . . . . . . 213
Blaustein, W. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 354
Boas, J. . . . . .. . . .. . . .. . . . . .. . . .. . . . . . . . . . . 48
Bocharova-Messner, 0. M. . .. 42, 43, 80, 120, 122
Boettiger, E. G. . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
Boettiger, E. G. and E. Furshpan .. 314, 321, 322
Bonhag, P. F ... 304, 307, 309, 311, 312, 314, 321,
322, 327, 328
Bomer, c. .. ................ .... .. .. .. .. .. 29
Bostick, R. V. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
Bracken, D. F. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 361
Breed, R. S. .. .. .. . .. .. .. . .. . .. . 65, 68, 70, 74
Breed, R. s. and E. F. Ball . . . . . . . . . . . . 213, 216
Bretfeld, G .......................... 28, 90, 94
Brindley, M. D. H. . . . . . . . . . . . . . . . . . . . . 278, 280
Brirtkhurst, R. 0. . . . . . . . . . . . . . . . . . . . . . . . . . . 282
Bracher, F ............................ 280, 283
Bromley, S. W. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 307
Burmeister, H. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Bursell, E. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 326
Byers, G. W. . .. .. .. . .. .. .. .. .. . .. . .. .. .. .. 312
..............................
Camerlengo, A. . . . . . . . . . . . . . . . . . . . . . . . . . . . 173
Campau, E. J. . .......... 205, 208, 209, 212, 214
Campbell, J. I. . . . . . . . . . . . . . . . . . . . . . . . . . . . 185
Carayon, J. .. . . . .. . .. . .. . .. .. .. . . . .. .. . . . . 278
Carbonell, C. S. . . 6, 151, 164, 165, 166, 180, 183
Carpenter, F. M. . . . . . . . . . . . . . . . . . . . . . . . . . . 46
Carpenter, F. M. and E. S. Richardson . . . . . . 43
Carpentier, F. 17, 18, 19, 24, 25, 28, 30, 33, 34, 39,
46, 70, 90, 92, 94, 99, 106, 108, 170, 173, 180, 184,
. 185, 202
Carpentier, F. et J. Bar!et .... 17, 26, 32, 97, 104
Carpentier, F. et M. L, Carpentier . . . . . . 394, 396
Chabrier, J. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
Chadwick, L. E. . ... 24, 71, 72, 73, 85, 151, 167,
216, 304, 307, 351
Chao, H. . . . . . . . . . . . . . . . . 66, 388, 390, 392, 394
Chapman, J. A. .. .. .. .. .. . .. .. .. . .. .. .. .. . 218
Chen, T. Y. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
Cholodkowsky, N. . . . . . . . . . . . . . . . . . . . . . . . . 48
China, W. E. .. .. .. .. .. .. . .. . .. .. .. .. .. . . . 279
Chopard, L. . . . . . . . . . . . . . . . . . . . . . . . . . . 139, 157
P.\GB
Chudakova, I. V. I 0. M. Bocharova-Messner 187
Clark, H. W. . . . . . . . . . . . . . . . . . . . . . . . . . . 399
Cochran, D. G. and M. H. Ross . . . . . . . . . . . 41
Compere, J. , . . . . . . . . . . 359, 366, 368, 369
Cope, 0. B. . .. 222, 224, 227, 228, 231, 233, 236,
. 237, 238, 239
Corset, J. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
Cowley, E. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 396
Crampton, G. C . .. 9, 10, 13, 19, 20, 24, 36, 40, 82,
92, 122, 126, 128, 132, 135, 141, 143, 148, 154,
167, 170, 187, 190, 192, 201, 203, 211, 215, 218,
222, 224, 248, 285, 287, 288, 297, 298, 300, 301,
302, 304, 307, 309, 311, 312, 315, 316, 317, 326,
334, 337, 339, 342, 349, 357, 359, 370, 386, 387,
388, 392
Crampton, G. C. and W. H. Hasey .. 37, 45, 151
Cremer, E. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 399, 401
Crowson, R. A. . . . . . . . . . . . . . . . . . . . . . . 214, 215
Czihak, G. . .... 30, 285, 288, 290, 291, 294, 295,
296, 297
Daly, H. V. . . 21, 74, 85, 359, 361, 366, 367, 368,
369, 371, 372, 373, 375
Danzer, A. . . . . . . . . . . . . . . . . . . . . . . . . . 322
Darwin, F. W. and J. W. S. Pringle . . . . 218, 373
Davies, R. G. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 190
Davies, W. M. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Davydova, E. D. . . . . . . . . . . . . . . . . . . . . . . 216, 218
De Beer, G. . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
Delamare-Deboutteville, C. . . . . 20, !57, 218, 220
Denis, J. R. . . . . . . . . . . . . 18, 90, 94, 97, 99, 190
Dierl, W. . . . . . . . . . . . . . . . . . . . 344, 354, 355, 356
Dirkes, L. .. .. . .. .. . .. .. . .. .. .. .. .. .. .. .. . 324
Doeksen, J. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 241
Dogs, W .................................. 280
Donges, J. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 216
Doyen, J. T. . . . . . . . 205, 207, 208, 209, 211, 2!3
Duncan, C. D. 357, 359, 361, 365, 369, 371,
374, 375
DuPorte, E. M. . . . . . 9, 36, 38, 40, 130, 139, 170,
185, 202
DUrken, B. .. ............... 45, 47, 65, 120, 125
Eastham, L. E. S. . . . . . . . . . . . . . . . . . . . . . . . . . 47
Eastop, V. F. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 197
Edmunds, G. F. and J. R. Traver . . . . . . . . . . . 120
Ege, R. .. .. .. .. .. . .. .. .. .. .. .. .. .. .. .. . .. . 280
Eggers, F. . . . . . . . . . . . . . . . . . . . . . . . . . . 352
Ehrlich, P. R. .. .. .. . .. . .. . .. .. .. . 342, 345, 349
Ehrlich, P. R. and S. E. Davidson . 354, 355, 356
Ehrlich, P. R. and A. H. Ehrlich . . . . . . . . 354
El Klfl, A. H. .. .. .. .. .. .. .. . 207, 208, 211, 213
El Sayed, M. T. . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
Enderlein, G. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 99
Esaki, T. and W. E. China .. .. . .. .. .. .. .. . 268
Esaki, T. and S. Miyamoto . . . . . . . . . . . . . . . . 268
Evans, J. W. .. .. .. .. .. .. . .. . 248, 250, 257, 260
Evans, M. E. G. . . . . . . . . . . . . . 202, 205, 209, 211
Ewald, 1. . . . . . . . . . . . . . . . . . . . . . . . . . . 275, 280
Ewer, D. W. . . . . . . . . . . . . . . . . 185, 186, !87, 190
Ewer, D. W. and L. S. Nayler . . . . . . . . . . . . . !51
Ewing, H. E. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
Ezzat, Y. M. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 254
Ferriere, c. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 283
Ferris, G. F. . ... 30, 32, 35, 36, 37, 46, 203, 205,
239, 241, 287, 288, 290, 309, 315
Ferris, G. F. and P. Pennebaker . 285, 287, 288,
290
Ferris, G. F. and B. E. Rees . . . . . . . . . . 293, 297
Ferris, G. F. and R. L. Usinger . . . . . . . . . . . . 275
Feuerborn, J. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Forbes, W. T. M ......... 45, 215, 352, 354, 394
Fowler, J. W. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Fox, I. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382
J. . ......................... 48, 88, 90
Freeman, T. N. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 350
FUller, H . .... 24, 32, 33, 297, 301, 302, 303, 304,
305, 306
Fuller, C. . .. . .. .. .. .. .. .. .. 154, 155, 156, 157
Gangrade, G. A. . . . . . . . . . . . . . . . . . . . . . . . . . . 139
Ganin, M. .. .. . .. .. . . .. . .. .. .. . .. .. .. . . . .. 325
Gaumont, R. . . . . . . . . . . . . . . . . . . . . . . . . . . 258, 262
Gaumont, R. et R. Moreau . . . . . . . . . . . . 275, 280
PAGE
Geigy, R. . . .. . . . . . . . . . . . . . . . . .. . . . . . . . . . . . 29
Giles, E. T. . ............ 187, 190, 192, 196, 198
Giliomee, J. H. .. .. .. .. .. . . .. . . .. . .. .. 254, 258
Glasgow, J. P. and S. Glasgow . . . . . . . . . . . . 326
Gael, S. C. . . . . . . . . . . . . . . . . . . 270, 271, 273, 282
Goldschmidt, R. B. . . . . . . . . . . . . . . . . . . . . . . . . 40
Goodrich, A. L. . . . . . . . . . . . . . . . . . . . . . . . 329, 332
Gouin, F. I. . ............... 28, 30, 40, 85, 130
Grandi, M. . ... 28, 44, 45, 70, 75, 115, 116, 118,
120, 122, 125, 128, 130, 131, 388, 392, 399
Grant, V. J. I. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 326
Grasse, P. P. .. .. .. .. .. . .. . .. .. .. .. .. .. 154, 157
Griffith, M. .. .................... 267, 277, 280
Gruner, H. E. . . . . . . . . . . . . . . . . . . . . . . . 24, 25, 32
Gunther, K. and K. Deckert . . . . . . . . . . . . 185
Gupta, A. P. . .. .. .. .. .. .. .. . 270, 276, 277, 278
Gurney, A. B. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 222
Haeckel, E. H. P. A. . . . . . . . . . . . . . . . . . . . . . . 83
Hagemann, J. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 275
Hakim, Z. M. . . . . . . . . . . . . . . . 386, 388, 389, 396
Hamon, J. et M. Ovazza . . . . . . . . . . . . . . . . . . 196
Handlirsch, A. ...................... 46, 86, 104
Hannemann, H. J. . . . . . . . . . . . . ... . . . . . . 334, 354
Hansel, S. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 324
Hansen, H. I . ................. 24, 25, 29, 30, 92
Hanson, J. F. .. .. .. .. . .. .. .. .. .. .. .. .. .. .. 131
Hardas, K. S. G. . . . . . . . . . . . . . . . . . . . . . . . . . . 197
Hasenoehrl, G. and P. P. Cook Jr. . . . . . . . . . . 248
Hasken, W. . ... 293, 297, 303, 304, 305, 306, 307
Hassan, A. A. G. . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Hatch, G ..................... 17, 392, 396, 399
Hatch, M. H. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
Haupt, H. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Heberdey, R. F. . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
Heider, K. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Heidermann, C. . . . . . . . . . . . . . . . . . . . . . . . . . . . 399
Heitmann, H. . . . . . . . . . . . . . . . . . . . . . . . . . 351, 352
Hendel, F. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 315
Hennig, W. . . . . . . . . . . 45, 83, 222, 315, 317, 319
Henrici, H. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278
Henry, L. M. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Henson, H. . ............. 190, 192, 193, 198, 199
Herbst, H. G. . .. .. .. . .. .. .. .. 207, 213, 215, 216
Hering, E. M. von . . . . . . . . . . . . . . . . . . . . . . . . 345
Hewitt, C. G. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 315
Heymons, R. . . 24, 26, 27, 28, 29, 30, 31, 32, 47,
139, 141
Hinton, H. E. . ...... 41, 48, 83, 85, 96, 205, 326
Holdsworth, R. P. . . . . . . . . . . . . . . . . . . . . . . . . . 130
Holmgren, E. . . . . . . . . . . . . . . . . . . . . . . . . . 6; 9, 401
Holmgren, N. .. . .. .. . .. . . .. . .. .. . .. . .. . . . . 157
Holste, G. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Hoppe, J. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 280
Hubbell, T. H. . . . . . . . . . . . . . . . . . . . . . . . . . . . 185
Hulst, F. A. . . .. .. . . . .. .. .. .. .. .. .. . .. . . .. 324
Hungerford, H. .. .. .. .. .. . .. . .. . .. . .. .. . . . . 275
Huxley, H. E. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 87
Ibrahim, M. M. . . . . . . . . . . . . . . . . . . . . . . . . . . . 28
Ide, F. P. . . .. .. . . . . . . . . . .. .. .. .. . . . .. . . 42, 111
Imms, A. D. .. . . . . .. . . .. .. .. .. .. .. . . . . . . .. 90
IvanovaKasas, 0. M. . . . . . . . . . . . . . . . . . . . . . 47
Jaboulet, M. C. . . . . . . . . . . . . . . 205, 209, 213, 214
Jackson, D. I. .. ... ............... 215, 216, 218
Jacobson, H. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 384
Janet, C ......... ........ , .. .. .. .. .. .. .. .. . 373
Janin, M. . . .. . .. . . . . . . . . . ... .. .. . . . . .. . . .. 90
Iannone, G. .. .................... 170, 183, 185
Jeanne!, R. ................ .' 44, 45, 46, 379, 381
Jeuniaux, C. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Jeziorski, L. ..... 58, 139, 141, 143, 144, 145, 146
Jesting, E. A. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 205
Johnson, B. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267
Jones, T. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 241, 243
Jordan, H. E. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 373
Jordan, K. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 345
Jusbaschjanz, S. . . . . . . . . . . . . . . . . . . . . . . . . . . . 324
Kawai, T ................................. 131
Kellin, D. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47, 48
Keler, S. von . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 56
Kelsey, L. P. . . . . 21, 65, 70, 285, 287, 288, 290,
291, 292, 294, 295, 296
Kessel, E. L. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Khanna, S. . 68, 268, 269, 270, 271, 273, 276, 278,
280, 282
PAGE
Kim, c. w. . . .. . . . . . .. . . .. .... "" ..... " 29
Kim, K. C. and E. F. Cook .. .. .. .. .. . .. .. . 312
Kinzelbach, R. K. .. .. .. .. . .. . .. .. .. .. . 373, 381
Kiriakoff, S. G. . . . . . . . . . . . . . . . . . . . . . . . . . . . 352
Kleinow, W. . 69, 80, 197, 198, 199, 200, 201, 215
Knox, V. . .. 47, 62, 111, 114, 115, 116, 120, 122,
123, 124
Kolbe, H. J. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Komp, W. H. W. . . . . . . . . . . . . . . . . . . . . . . 309, 315
Korn, W. . ...... 71, 291, 292, 293, 294, 295, 296
Kramer, S. . ......... 151, 183, 248, 254, 258, 260
Kristensen, N. P. . . . . . . . . . . . . . . . . . . . . . . 354, 356
Kryzanowsky, S. G. . . . . . . . . . . . . . . . . . . . . . 83
Kukalova, J. . ........................ 42, 43, 44
Kiinckel, d'Herculais, J. . . . . . . . . . . . . . . . 322, 325
Kuske, G. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
La Greca, M. . . . . . 14, 17, 36, 40, 151, 157, 162,
163, 170, 173, 174, 175, 178, 185, 190, 270, 277, 280
I.a Greca, M. e R. Cesaro . . . . . . . . 270
I.a Greca, M. e A. Raucci . . 157, 162, 163, 164,
165, 166
Lameere, A. . . . . . . . . . . . . . . . . . . . . . . . 91
I.arsen, 0. . .. 46, 68, 70, 199, 202, 203, 205, 207,
208, 209, 211, 212, 213, 215, 216, 217, 218, 219,
220, 248, 267, 268, 270, 271, 273, 275, 277, 278,
280, 282, 283, 284, 285
I.aurent, L. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 209
Lauck, D. R. . . . . . . . . . . . . . . . . . . . . . . . . . 270, 282
I.eclercq, J. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 368
I.eech, H. B. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 215
Lefeuvre, J. C. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
I.ebmann, F. E. . . . . . . . . . . . . . . . . . . . . . . . . . 47, 48
Lemche, H. . . . . . . . . . . . . . . 40, 41, 42, 43, 44, 45
Lendenfeld, R. .. .. .. .. .. .. .. .. .. . .. .. . .. .. 399
Leston, D, . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Levereault, P. . ..... 157, 158, 159, 161, 163, 164,
165, 166
Lew, G. L. . . . . . . . . . . . . . . . ... . . . . . . . . . . 248, 250
I.ewis, R. E. ........................ , . 384, 385
I.indroth, C. H. . ................... ... . . .. . 213
Littig, K. S. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 141
Lowne, B. T. . .................... 315, 322, 325
I.ubbock, J. . ..... , . . . . . . . . . . . . . . . . . . . . . . . . 371
Luks, C. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 399
Lyonet, P. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 354
Mackerras, I. M. . . . . . . . . . . . . . . . . . . . . . . . . . . 45
Madden, A. H. . . . . . . . . . . . . . . . . . . . . . . . 343, 349
Magnan, A. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 399
Mlikel, M. .. .. .. . .. .. .. .. . .. .. . .. . 64, 262, 282
Maki, T. . ... 7, 48, 60, 64, 65, 66, 68, 69, 74, 78,
92, 94, 99, 110, Ill, 120, 123, 124, 131, 133, 134,
137, 139, 141, 143, 144, 145, 146, 151, 163, 164,
165, 166, !85, 186, 188, 189, 190, 199, 200, 201,
216, 220, 230, 231, 232, 247, 261, 263, 264, 282,
285, 288, 291, 294, 295, 296, 304, 305, 306, 322,
327, 328, 332, 335, 336, 354, 371, 374; 375
Maloeuf, N. S. . ...... 71, 74, 85, 270, 271, 277,
282, 401
Mark!, H. .. . .'. .. . . . .. . .. . . 6, 371, 372, 374, 375
Marcus, H. . .. . .. . .. .. . .. . .. . . .. . . .. . . .. . . 399
Marquardt, F. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Martin, J. F. . . . . . . . . . . . . . . . . . . . . . . . . . 126, 357
Martynov, A. V. . . . . . . . . . . . . . . . . . . . . . . . . . 41
Marvillet, C. .. .. . .. .. .. . .. . .. .. .. . 203, 209, 213
Mathur, A. C. . . . . . . . . . . . . . . . . . . . . . . . . . 387
Mathur, P. N. and A. N. T. Joseph . . . . . . . . 248
Matsuda, R. . . . . 6, 9, 18, 21, 28, 30, 32, 34, 35,
36, 38, 45, 46, 47, 80, 85, 86, 111, 115, 116, 120,
123, 124, 136, 143, 147, 148, 152, 157, 163, 166,
196, 205, 251, 258, 268, 275, 276, 277, 279, 280,
285, 287, 290, 291, 296, 302, 315, 316, 329, 331,
349, 450, 351, 366, 367, 369, 370, 394
Mayer, C. . . . . . . . . . . 232, 233, 236, 237, 238, 239
Melis, A. . . . . . . . . . . . . . . . . . . . . . . . . . 241, 247, 248
Menon, K. R. and N. K. Joshi ......... , 215, 216
Mercier, M. L. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 325
Mercier, L. et R. Poisson . . . . . . . . . . . . . . . . . . 199
Michener, C. D. . 35, 342, 343, 345, 349, 367, 368
Mickoleit, G. . . 63, 68, 74, 75, 241, 242, 243, 244,
245, 247, 249, 297, 300, 301, 302, 303, 304, 305,
306, 307, 312, 319, 321, 322, 327, 328, 354, 357
MthaJyi, F. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 314
Miller, A. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 262, 322
Miller, F. W. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 291
Miller, P. L . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
.Misra, S. D. . ........... 173, 174, 180, 183, 185
PAGE
Miyake, T. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 293
Morison, G. D. . . . . . . . . . . . . . . . . . . . . . . . 371, 373
Miiller-Biumenau, W. . . . . . . . . . . . . . . . . . . . . . . 48
Murray, F. V. and 0. W. Tiegs . . . . . . . . . . . . 217
Mutuura, A. . . . . . . . . . . . . . . . . . . . . . . . . . . . 8, 293
Myers, J. G. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248
Needham, J. G. and M. H. Anthony . . . . . . . . 391
Nelson, J. A. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Neville, A. C. . . . . . . . . . . . . . . . . . . . . . . . . 398, 399
Niiesch, H. . ............... 7, 345, 354, 355, 356
Nussbaum, R. . . . . . . . . . . . . . . . . . . . . . . . . 319, 322
Onesto, E ... 16, 128, 129, 148, 149, 150, 151, 159,
160, 173, 180, 192, 193, 196, 197, 198, 345, 347,
351
Paclt, J. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88, 90
Pantel, J. . . . . . . . . . . . . . . . . . . . . . . . . 192, 193, 198
Parsons, M. C. . . . . . 268, 269, 270, 271, 273, 275,
277, 278, 280, 282, 285
Partmann, W. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 326
Perez, c. . . . . . . . . . . . . . . . . . . . . . . . . . . 86, 325, 373
Pierce, D. W. . .......... 373, 376, 377, 379, 381
Pipa, R. I. and E. F. Cook . . . . . . . . . . . . . . . . 6, 7
Poisson, R. . . . . . . . . . . . . . . . . . . . . . . . 275, 280, 283
Poletaiew, N. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 398
Popham, E. J. . . . . . . . . . . . . . . . 190, 192, 199, 280
Prell, H. . . . . . . . . . . . . . . . . . . . . . . ... . . . 48, 88, 90
Priesner, H. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247
Pringle, J. W. S. . . . . 87, 216, 321, 322, 370, 371
Puchkova, L. V. . . . . . . . . . . . . . . . . . . . . . . . . . . 278
Quennedey, A. . . . . . . . . . . . . . . . 202, 205, 209, 213
Ragge, D. R. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 139
Rakshpal, R. . . . . . . . . . . . . . . . . . . . . . . . . . 173, 185
Raw, F. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Rawat, B. L. . . . . . . . . . . . . . . . . 270, 273, 280, 282
Reeks, W. A. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 361
Rees, B. E. and G. F. Ferris .. 304, 311, 312, 315,
316, 321
Reid, J. A. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 362
Reid, R. W. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 218
Remane, A. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83, 85
Rensch, B. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 83
Ribaga, G. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 230
Richards, A. G. . ..................... :. . . 352
Richards, A. M. . . . . . . . . . . . . . . . . . . . . . . . . . . 173
Richards, 0. W. . . . . 170, 357, 358, 367, 368, 369
Richards, 0. W. and R. G. Davies . . . . . . . . 48
E. A. . . . . . . . . . . . . . . . . . . . . . . . . . . 213
p: F: ::::::::::::::::::::::::::::::
Riter. w. A. . ........................ :. . 9, 10
Rilling, G. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 262
Rings, R. W. . ........................ . . . 213
Ritter, W. . ........................... . . . 322
Rivard, I. ............................. : . . . 361
Rivnay, E. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 213
Rodendorf, B. B. . . . . . . . . . . . . . . . . . . . . . . . . 45, 46
Roonwal, M. L. . ............ 27, 29, 32, 47, 184
Ross, H. H. . ............ 222, 357, 358, 361, 370
Ross, M. H. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40, 41
Rothschild, W. and K. Jordan . . . . . . . . . 349, 350
Rowell, H. F. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 173
Riischkamp, P. F. . ................ 211, 215, 216
Russenberger, H. and M. Russenberger . . . . . . 399
Saksena, R. D. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 71
Sander, K. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 267
Sara, M. and A. Smerdel . . . . 304, 309, 312, 317
Sargent, W. D. . . . . . . . . . . . . . . . . . . . . . . . . 394, 396
Saxena, 0. N. . ................... 209, 213, 216
Schaefer, C. . . . . . . . . . . . . . . . . . . . . . . . . 278
Schmidt, E. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 324
Schrrutt, I. B. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Schneider, G. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 322
Schultz, H. . ............ :. . . . . . . . . 326, 334, 342
B. N. . . . . . . . . . . . . . . . . . . . . 44, 45
Seher, R. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41
Setty, L. R. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 304
Sewertzov, A. N. . . . . . . . . . . . . . . . . . . . . . . . . . . 83
Shafiq, S. A. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Sharov, A. G. . ...................... 24, 45, 46
Sharplin, J. . . . . 16, 290, 301, 302, 334, 337, 339,
345, 347, 349, 351, 352, 354, 357
Shatoury, H. H. El . . . . . . . . . . . . . . . . . . . . . . . . 325
Shepard, H. H. . . . . . . . . . . . . . . . . . . . . . . . 349, 352
PAGE
Short, J. R. T. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 282
Smart, J. . . . . . . . . . . . . . . . . . . . 322, 326, 327 328
Smit, W. A. . ............................. ' 154
Smith, D. S. . . . . . . . . . . . . . . . . . 87, 215, 216, 218
Smreczinsk.i, S. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Snodgrass, R. E . .. 6, 9, 10, 11, 12, 13, 14, 15, 16,
18, 19, 20, 23, 24, 25, 30, 31, 32, 41, 42, 46, 47,
48, 69, 83, 94, 116, 125, 143, 148, 150, 159, 173,
180, 183, 184, 185, 186, 188, 189,,190, 193, 207,
209, 309, 315, 354, 357, 359, 361, 362, 365, 366,
370, 371, 374, 375, 382, 390
Speyer, W. . ................. 64, 65, 71, 73, 216
Sprague, I. B. . . . . . . . . . . . . . . . . . . . . . . . . . . 282
Srivastava, K. P. . . . . 342, 345, 349, 354, 355, 356
Steiner, P. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 301
Stellwaag, F. . ........... 208, 213, 215, 216, 370
Storch, R. H. . .................... 151, 153, 167
Storch, R. H. and L. E. Chadwick . . 68, 153, 166,
170, 304
Subba Rao, B. R. . . . . . . . . . . . . . . . . . . . . . . . . . 358
Sundermeier, W. . . . . . . . . . . . . . . 47, 285, 288, 290
Tait, N. N. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 361
Tanner!, W. . ..... 44, 64, 66, 388, 390, 392, 398,
399, 401
Taylor, L. H. . . . . . . 248, 257, 258, 261, 270, 275
Theron, J. G ...................... 248, 254, 258
Thomas, J. G. . . . . . . 41, 173, 185, 186, 187, 190
Tiegs, 0. W. . ... 27, 60, 65, 70, 72, 80, 86, 153,
154, 163, 166, 187, 262, 267, 325, 326, 373
Tietze, F. . . . . . . . . . . . . . . . . . . . 211, 215, 216, 218
Tillyard, R. J. . . . . . . . . 45, 66, 215, 361, 392 399
Tindall, A. R. . . . . 8, 62, 326, 332, 335, 336: 354
Tower, G. D. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 270
Treat, A. E. . . . . . . . . . . . . . . . . . . . . . . . . . . 352, 354
Tremblay, E. . . . . . . . 209, 211, 213, 250, 258, 261
Tulloch, G. S. . . . . . . . . . . . . . . . . . . . . . . . . 361, 362
Tuxen, S. L. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 88
Ulrich, W. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377, 381
Usinger, R. and R. Matsuda . . . . . . . . . . . 275, 279
Uvarov, B. P. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 184
Uvarov, B. P. and J. G. Thomas . . . . . . . . . . . 170
Vasvary, L. M. . ................. : 261, 263, 264
Verhoelf, K. W .. 9, 27, 29, 39, 132, 135, 198, 199
Viallanes, S. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 325
Voss, F. . . . . . 16, 47, 56, 185, 186, 188, 189, 190
Wada, S. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . 10
Wagner, J. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382
Walker, E. M. . . . . . . 144, 145, 146, 167, 168 169
Walker, I. J. and A. B. Gurney ............ ' 178
Waterhouse, D. F. and A. R. Gilby . . . . . . . . . 278
Weber, H. . ... 11, 13, 14, 19, 20, 21, 24, 29, 30,
32, 36, 38, 40, 46, 47, 48, 85, 94, 130, 131, 248,
250, 254, 256, 258, 261, 262, 265, 266, 267, 275,
290, 342, 344, 345, 349, 350, 351, 352, 354, 357.
. 359, 361, 362, 366, 368, 369, 371, 375
We1s-Fogh, T. . . . . . . . . . . . . . . . . . . . . . . 16, 396 399
Weismann, A. . ..................... 6, 324' 325
Wellman, M. H ............................ ' 342
Wells, M. J. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 278
Wenk, P. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 382, 383
Wheeler, W. M. . . . . . . . . . . . . . . . . . . . . . . . . . . . 48
Wiesend, P. . ....................... 70, 185 186
Wiesmann, R. . ........................... ' 7
Wigglesworth, V. B. . . . . . . . . . . . . . . . . . . . . . . . 41
Wilfried, W. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 248
W!lle, A. . ......................... 45, 371, 375
Wlllem, V. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 90
Williams, C. M. and M. V. Williams . . . . 322, 326
Williams, I. W. . .................. 304, 311, 312
W!Json, D. M. . . . . . . . . . . . . . . . . . . . . . . . . . 80, 185
Wmkler, D. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 334
Wittig, G ... 7, 40, 62, 63, 64, 70, 74, 80, 85, 122,
131, 132, 133, 134
Wong, H. R. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 361
Woodland, J. T. . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
Woodring, J. P. . . . . . . . . . . . . . . . . . . . . . . . . . . . 40
Young, B. P. . . . . . . . . . . . . . . . . . . . . . . . . . 316, 319
Young, E. C. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 283
Yu-Pei-yu, and Chin-jen Luh . . . . . . . . . . 180, 185
Zalessky, Iu. M. . . . . . . . . . . . . . . . . . . . . . . . . . . . 44
Zalokar, M. . ............ 309, 311, 312, 317 322
Zeuner, F. . ............................... ' 215
Zeve, V. H. and D. E. Howell ......... 309, 319
, ~ - - - ~ - - - - ~
ERRATA
On page 5, in the last paragraph, Dr. J. R. McAlpine should
read Dr. T. R. McAlpine.
On page 17, in FIG. 5, Mesocutide should read Mesocuticle.
On page 358, in FIG. 155A, Metanoturn should read Mesonotum.
- - - ~ - - - - - - - - - - - - - - - - - - - ~ - - - - - - - - - - ~ - - - - - - - ~ - - ------

Insect Morphology

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Insect Morphology

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MORPHOLOGY AND EVOLUTION

OF THE INSECT THORAX

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