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_ AUCHEN ORHYNCHOUS
HOMOPTERA

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By
SOL KRAMER

_ THE UNIVERSITY OF ILLINOIS PRESS
Bab? 4 Hg URBANA, 1950

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[Continued inside back y 0

THE MORPHOLOGY AND PHYLOGENY OF
AUCHENORHYNCHOUS HOMOPTERA

Illinois Biological Monographs, Vol. XX, No. 4
Published for the Graduate College, University of Illinois

BOARD OF EDITORS

Harley Jones Van Cleave
William R. Horsfall
Leland Shanor

The
Morphology and Phylogeny
of
AUCHENORHYNCHOUS
HOMOPTERA
(Insecta)

By
SOL KRAMER

tik ONEV ERS DY OF TLLINOIS PRESS
URBANA, 1950

COPYRIGHT 1950 BY THE UNIVERSITY OF ILLINOIS PRESS

MANUFACTURED IN THE UNITED STATES OF AMERICA

ACKNOWLEDGMENTS

The writer wishes to acknowledge his gratitude and appreciation to Dr.
William P. Hayes, under whose direction this study was undertaken. He has
shown an enthusiastic interest in the progress of this study from its inception
and contributed a critical and valuable reading of the original manuscript.
Further thanks are extended to Dr. Hayes for his help in obtaining financial
aid and other assistance which enabled the writer to devote a maximum of
time to studies and research during two years at the University of Illinois.

Appreciation is also expressed to Dr. H. H. Ross of the Illinois State Natural
History Survey, to Dr. Charles D. Michener of the American Museum of Nat-
ural History, to Dr. J. W. Evans of the British Museum and Louise M. Russell
of the U.S.D.A. Bureau of Entomology and Plant Quarantine for providing
specimens used in this study; to Dr. P. W. Oman of the U. S. National Museum
for specimens and identifications; and to Dr. Kathleen C. Doering of the Uni-
versity of Kansas for determinations.

Assistance in interpreting the homologies of the mouthparts of several
Homoptera was provided by R. E. Snodgrass during the writer’s studies in his
laboratory at the U. S. National Museum during the summer of 1947, and this
assistance is gratefully acknowledged. Sincere thanks are also expressed to
Miss Ruth Keller for typing the major portion of the original manuscript.

An additional note of thanks is due to Dr. Evans for a critical reading of the
manuscript, much helpful discussion and other kindnesses, during the writer's
stay in England in 1948-49, while preparing this study for publication. Con-
siderable additional information was thus made available.

CONTENTS

MIRE NNITNIRE UVONE OL Si nt Sot ae Nie oo Bee Coa a ek Ree 2 Ne 1

EP LDS. CORE ASSTn PTAG C0 <a a ce 3

— LAT SUES Ia Ce eS 6
Tue Screrites, APPENDAGES, AND Muscies oF THE Heap OF

RMR EAE EBRREEN NCEA eS machen eco ons Me A actrees cles oh eR den ee ee 8

mpolaps pungens Germar (Fulgoridae) .. 2.2.0.0 oc os. t<neesan = 8

iybicmna seprendecim (Linn.) (Cicadidae))......5..s0.025002 54 «00% > ii!

Lepyronia quadrangularis (Say) (Cercopidae).................. 13

gailacizes srrorata (Fabr.) (Cicadellidae)2 .. 3.0.22 os dan cca ee ne om. 14

Ceresa bubalus (Fabr.) (Membracidae)........... Sch tes ie ie 16

Tue Tuoracic Scvierites, WiNncs, AND Lecs oF AUCHENORHYNCHA......... 18

RECEYaN (DEINE PINIL OC TINE tre rer St ae Cees Ng eS eae ee Fel 18

ETE ND ACSC DE CIMA CCITIEN. ANG 3 St idea ie te oe ces, eS a ee 21

eRe QUAD TAN ODIAT IS 0 vo or slay SO raves Bee em CR baer 23

DAE RG EZ OSMIE ROTTEN cio oe hn, i se oe Rr TE 25

TOS PRUIEV GUISE es ts Symes ee eae tn at ae Te ee oF,

ne sPsorAcic Muscurature or Cerésa bubalus. os. o5 06d. sn ndot cdi cose 31

ABDOMINAL SEGMENTS OF AUCHENORHYNCHA........2.cceccececccceceeee 37

AOI I OONIS Hare eo nee «ete, He IT ae | tee ee 37

MUOMBACS ODI CWAC CIBC S a Iecy s orks Be cers ee Sl ee SOO 40

Pe PVT OMIT) GUDATUICUIATIS ah ockre Sohne Seo N ER Soe Cee ene Ca 43

PAAACE ESA ITT OVAL iss. She oe ee, ee ws ts a ne 45

SOOTS SULTS, COIR Cc, Seas hg pep he ay ee eae Se ia on the 47

Tue ABDOMINAL MuscuLatTureE oF Ceresa bubalus.. 2.0... cece cece ccc cees 51

eee usLOGENY (Or ACUCHENORHYNCHA: «sc 6.esse+sccsee ce coce cee btetaeke 54

Phylogenetic Considerations of Previous Authors................ 54

MED ESE SSURHIA Noyce ee ta ae ee I ete ie a one ns nS ee de ae 62

EUS BO Er aga Se ee ae ea a ne ee Ace ae ee Me eeen ont ae, e 71

PRAT HERE VEN EMOM Se!) Gly Se a gost ok Rnd OO ee See ae 76

LEDS OPEL ER en Sr O01 Coe ec as Pe Ne 79

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INTRODUCTION

Comparative studies of insect structure and resulting classifications
based on phylogeny have been few in comparison to the amount of mate-
rial available for study. Handlirsch (1906-08) was able to establish
the minimum geologic age of many groups of insects through paleonto-
logical studies, and he considerably modified the prevalent conceptions of
his day concerning the ordinal relationships of insects. Although Tillyard
(1930), Carpenter (1930), and others have attacked this problem, fossil
insects are meager in comparison to other animal fossils and many gaps
still exist in this line of evidence. Consequently, the insect taxonomist
must depend on morphological, physiological, ecological, and zoogeo-
graphical sources for his evidence of insect relationship. It should be
added that the vast numbers of insect species and groups, ordinarily a
handicap, is of decided advantage in phylogenetic studies. Insects have
retained in varying degree many primitive characters and their large
numbers increase the possibility that there may be derived from existing
forms, composite pictures of ancestral types which will clarify relation-
ships.

Just such an attack on the problem of insect phylogeny, through exten-
sive comparative studies of present-day forms, has been made by Cramp-
ton (1924). During the past forty years he has undertaken studies of rela-
tionship throughout the Insecta, based upon diverse characters, such as
the head capsule, thoracic sclerites, wing articulation, abdominal segments
and appendages, and genitalia, as well as on the characters of wing vena-
tion and mouthparts previously relied upon. His studies, which provided
important new concepts of insect phylogeny, have done much to lay the
groundwork for future investigation. It should be noted, however, that
Crampton’s studies were devoted primarily to the broader relationships
of insects, and his resulting classification concerns the relationships of the
higher categories such as subclasses, superorders, and orders. The prob-
lems of phylogeny within the orders of insects, concerning families and
lower groupings, remain virtually unsolved.

This comparative study of the Auchenorhyncha is an endeavour to
examine one such problem of relationship. One species from each of five
families of Auchenorhyncha has been selected for detailed study. Some
of the material presented here has been dealt with by others, but much of
it is new. Relationships of structure have been indicated by many original
drawings, and these studies of the structure of representative Aucheno-

1

2 MORPHOLOGY OF AUCHENORHYNCHA

rhyncha have for the first time been incorporated into and compared in
a single work. The writer has further included a detailed account of the
musculature of the membracid representative, Ceresa bubalus.

This investigation represents only a preliminary effort toward the un-
derstanding of relationships within the Auchenorhyncha. Through it, the
writer has been impressed with the little that is known concerning the
functional significance of particular structural modifications. Snodgrass
(1935), Weber (1933), and others have done a good deal to emphasize
the importance of ascertaining the functional value of anatomical facts,
and the value, wherever possible, of supplying reasons for existing struc-
tures. Such comparative functional studies should be integrated with
morphological studies within the lower categories of insects, if we are to
approach some understanding of what has been taking place in the evo-
lution of insects. In this connection it would not be amiss to point out
that comparative studies of behavior patterns in some groups of arthro-
pods, such as bees, wasps, spiders, etc., have provided valuable charac-
teristics for the systematist. Such work is based on the assumption, as
Lorenz (1939) has pointed out, that “the concept of homology, useful in
morphology, is just as applicable to inherited behavior-patterns as to
bodily characteristics.” Sound classifications should be based on the inte-
gration and evaluation of all ascertainable phylogenetic evidence.

The large numbers of insect species have in some measure precluded
these studies by systematists. It is nevertheless true that the systematist
who specializes in a particular group of insects is in the best position to
answer such questions, for he is most intimately acquainted with the
many species represented and their diverse modifications of structure and
behavior. Future attempts to provide reasons for structure, even though
our efforts be faulty, will lead the more quickly to an understanding of
what has been taking place in the diverse auchenorhynchous groups of
insects, and to a greater comprehension of the Homoptera in general. It is
hoped that this study will stimulate questions among taxonomists in these
insect groups, and will provide some morphological basis for their
answers.

NOTES ON CLASSIFICATION

In a work on the general classification of animals, Dumeril (1806)
separated the Hemiptera into six families. Two of these families, the
Collirostres or Auchénorinques,* comprising the cicadas, membracids,
cicadellids, fulgorids, and cercopids, and the Plantisuges or Phytadelges,
comprising the aleyrodids, chermids, aphids, and coccids, represent in
part the Homoptera as we know them.

Westwood (1840), adopting Latreille and Burmeister’s view as to the
division of Homoptera into three primary sections, named these sections
on the basis of differences in the number of tarsal joints as follows:

1. Trimera. Tarsi 3-jointed: antennae minute, setigerous, wings areo-

late (Cicada, Linnaeus).

2. Dimera. Tarsi 2-jointed: antennae moderate, filiform, 5-10 jointed:

wings subareolate (Aphis, Linn. and Psylla, Geoffr.).

3. Monomera. Tarsi 1-jointed: antennae 6-25 jointed: wings not areo-

late (Coccus, Linn.). |
The Dimera contain the families Psyllidae, Aphidae, and Aleyrodidae; the
Monomera contain the Coccidae; while the Trimera contain the Cicadi-
dae, Fulgoridae, and Cercopidae.

Amyot and Serville (1843) in their work on the classification of Hemip-
tera separated the Homoptera as a suborder, and divided the latter into
sections, Auchénorhynques and Sternorhynques. The former section con-
tains the Trimera of Westwood, while the latter comprises both the
Dimera and Monomera.

Buckton (1889) grouped the cicadas, membracids, fulgorids and jassids
into a single family, Tettigiidae, which he separated from the Aphidae,
Coccidae, etc. Following Latreille, who had divided the Cigalles or
Cicadae into two groups separating the singing from the nonsinging
forms, “les Chanteuses” and “les Muettes,” Buckton separated the Tet-
tigiidae into two subfamilies. The subfamily Stridulantia contained the
tribes Tibicinae and Cicadinae, while the subfamily Silentia contained
the tribes Membracinae, Fulgorinae, Cercopinae, and Jassinae. In this
connection it is interesting to note a recent comment by Evans (personal
communication, 1948) concerning a broadcast given by Ossiannilsson
from Stockholm of the amplified songs of jassids and cercopids, which
sounded like the songs of cicadas and frogs. He adds that Ossiannilsson

*This name refers to the apparent origin of the beak or proboscis, from the “neck”
or cervical region, as contrasted with the Sternorhyncha in which the proboscis seem-
ingly arises from the prothoracic sternal region.

3

4 MORPHOLOGY OF AUCHENORHYNCHA

had previously published a paper (Opusc. entom., 1946; 82) in which he
gives a preliminary note on sound-producing organs in the Homoptera.
Evans (1946b) had not seen this paper when he called attention to a
supposed tymbal in an aetalionid, Darthula hardwickii Gray.

Distant (1906) has pointed out that Westwood recognized only three
families of Auchenorhyncha: the Cicadidae, Fulgoridae, and Cercopidae.
Stal divided this group into four families: Stridulantia, Cercopida, Jassida
(including membracids), and Fulgorida, and in this view he was sup-
ported by Hansen. Although Distant treats the Membracidae as a sep-
arate, fifth family of the Auchenorhyncha, he adds that both Stal and
Hansen may possibly be right, on morphological grounds, in relegating
the membracids to a subfamily of the Jassidae.

According to China (1927), the family Peloridiidae was erected for a
primitive insect Peloridium hammoniorum taken on Navarin Island, Tierra
del Fuego, described by Breddin in 1897, and assigned to the Heterop-
tera. In 1924 a new genus and species of this family, Xenophyes cascus,
taken in New Zealand, was described by Bergroth. In the same year
another new genus and species, Hemiodoecus leai, taken at Hobart, Tas-
mania, was described by China.

Myers and China (1929) concluded that the Peloridiidae were more
closely allied with the Homoptera. This conclusion was primarily on the
basis that the only character clearly separating the Heteroptera from the
Homoptera was the presence of the gula, absent in the Homoptera and
the Peloridiidae. They suggested a new series, Coleorhyncha, for the
Peloridiidae, to be placed at the base of the Homoptera. The classifica-
tion of Hemiptera, based on key characters, was modified by them as
follows:

Galapsoresent feeaeert ts eighty Velaictetateten= be arc ete etek setts HETEROPTERA
Cuilagalnsentiae vis sate eae ce Dp ct BORIC Ace ea cae chee HOMOPTERA
Rostrum arising from base of head
Base of rostrum sheathed by propleural structures...... COLEORHYNCHA
Base of rostrum entirely free from prothorax........ AUCHENORHYNCHA
Rostrum arising between or caudad of fore coxae....... STERNORHYNCHA

More recently Evans (1946a), as a result of extensive taxonomic and
morphological studies of the leaf-hoppers and related Auchenorhyncha,
proposed a separation of this group into three divisions, as follows:

1. Tegula almost always present; middle coxae elongate, articu-
lated far apart, free and capable of lateral movement; hind

coxae immobile; wing without a marginal vein...... Fulgoromorpha
Tegula absent; middle coxae short, articulations near together;
hind coxae mobile; wing with a marginal vein.................. 2

2. Head with a swollen postclypeus, the crown with a small tri-
angular frons bearing a median ocellus............ Cicadomorpha

NOTES ON CLASSIFICATION 5

Head with a large but not necessarily swollen postclypeus,
which may or may not be distinct from the frons; frons
Matboubra median OGellUsi-0 i. hs. «asc s saa due elacaes Mh '@ oh Jassidomorpha
He then subdivides the Jassidomorpha into two superfamilies as fol-
lows:
In the head, the anterior arms of the tentorium fused with the pos-
terior arms; hind coxae short, conical, not transversely
dilated; hind tibiae with one or a few spurs but lacking
SIRS athe PTAs easton Schade, Sune ne ees cite Bas Teh Cercopoidea
In the head, the anterior arms of the tentorium not in contact with
the posterior arms; hind coxae transverse, laterally dilated;
hind tibiae armed with spines, some of which may be
Mmomnted om ‘enlarged’ DASES sy. a. 5005 ssc Blew Aes ae es Jassoidea
(includes the Membracidae)
In his most recent study Evans (1948) reiterated his concept that the

membracids belong to the jassoid complex, and divided the membracids
into three families, the Nicomiidae, Lamproteridae, and Membracidae.

MATERIALS AND METHODS

The procurement of properly preserved and adequate numbers of
specimens constitutes a major problem for such a study. Proper preser-
vation, essential for the study of insect musculature, is sometimes obtained
by chance when live specimens are dropped into 70 per cent alcohol. But
for the most part this method is inadequate. The musculature of some
insects used in this study, first dropped into steaming water and then into
70 per cent alcohol, was fairly well preserved. By far the best method of
preservation, on the basis of comparative tests with Ceresa bubalus, is to
drop live specimens into steaming water and then transfer to a 5 per cent
solution of chloral hydrate where the insects may be stored indefinitely.
Steaming water, followed by Bouin’s fluid for 24 hours, then storage in 70
per cent alcohol gives almost equally good results.

A variety of dissecting tools were made by grinding, on carborundum
and oilstone, stainless steel, surgical needles into extremely fine points,
probes, or scalpels and placing these into lightweight, dental broach hold-
ers. A pair of microdissecting scissors, the blunt ends of which were
ground down almost to needlepoint, proved very useful.

Dissections were performed underneath 50 per cent alcohol in metal
pillboxes, % inch high by 2 to 3 inches in diameter, bedded with paraffin
wax. As many as a score of dissections were made for each figured draw-
ing. The drawings were first made on cross-section paper, but later trac-
ing paper, underneath which a cross-section guide had been placed, was
used. An ocular micrometer proved to be an invaluable aid for this work.

For the dissection of the smaller insects in this study, an adaptation of
a method described by Guyer (1947, p. 92) proved very helpful. Longi-
tudinal, or cross-sections of small insects, or of parts of insects such as the
head, thorax, etc. were imbedded in paraffin wax. In imbedding, a small
area of paraffin is first melted with a hot needle or scalpel. The section
of the insect which has been removed from 70 per cent alcohol, and blot-
ted to remove excess liquid, but allowed to remain moist, is immersed in
the liquid paraffin up to the upper surface of the insect section. The
paraffin around the insect section is again melted so that it securely grips
the section when it sets. The dissecting pan is then flooded with 50 per
cent alcohol. Any paraffin which has run into the muscles or other internal
parts may be readily picked away with a needle. Dissections of minute
muscles and parts, which might have been extremely difficult, were
greatly facilitated by the use of this technique.

Another method which was of value in muscle study was an eosin
staining and oil of wintergreen clearing procedure which I have recently

6

MATERIALS AND METHODS £

described (Kramer, 1948). This technique enables one to view the posi-
tion and relationship of muscles in the thorax and abdomen directly
through the body wall of the insect. It was particularly useful when used
to ascertain the position of muscles in such smaller structures as the
labium, and segments of the leg of Ceresa bubalus.

THE SCLERITES, APPENDAGES, AND MUSCLES OF
THE HEAD OF AUCHENORHYNCHA

Scolops pungens Germar
(Fulgoridae )

Long considered as a single family Fulgoridae, the fulgorids have
more recently been given superfamily rank (Fulgoroidea) and have been
considered to represent as many as eighteen families. The modifications
of the head in this group are of particular interest since the head has been
interpreted by past writers to represent both specialized and generalized
modifications in the Homoptera.

The head of Scolops pungens is shown in frontal view in figure 7. The
median striated area is the postclypeus (pclp), beneath which is the
anteclypeus (aclp). The small labrum (Im) is suspended from the ante-
clypeus, and the lorae (lor) lie laterad of the clypeal area. Above, the
frontal region or frons (fr), prolonged dorsally into a cephalic process,
is separated from the postclypeus by the epistomal suture (est). The
frons contains three ridges or carinae medially and a pair laterally, the
latter extend down through the clypeal region. The prominent compound
eyes (e), lateral ocelli (oc, Fig. 6) and antennae (ant, Fig. 8) all lie
laterad of the frontal region.

The elongate region, posterior to the lorum and anteclypeus is termed
the maxillary plate (mxp, Fig. 8). The labium (lb) is suspended by a
membrane from the posterior edge of the maxillary plate and the hypo-
pharynx (not seen). Both the lorum and the maxillary plate are separated
from the genal region above (ge) by a definite internal ridge which ap-
parently represents the subgenal suture (sgs). The presence of the sub-
genal suture in this fulgorid, partly present or absent in other fulgorids,
is particularly noteworthy. The line separating the lorum and anteclypeus
from the maxillary plate has sometimes been referred to as the genal
suture, but this is a misnomer since the maxillary plate is merely closely
approximated to the margins of the lorum and anteclypeus and is in no
way joined to them.

The mandibular stylet (mds), sometimes called mandibular seta, or
mandibular bristle, is shown in figure 12. It consists of a comparatively
broad basal portion and a narrow bristle-like terminal portion. The re-
tractor muscle (rmds), originating on the vertex of the head capsule,
is inserted on the base of the mandibular stylet by means of an elongate
tendon. The base of the stylet gives rise to another arm, termed the lever

8

SCLERITES, APPENDAGES, AND MUSCLES OF THE HEAD 9

(lvr,), which articulates with the lorum at the point where the subgenal
suture meets the epistomal suture. On this lever is inserted the protractor
muscle (pmds) which originates on the ventral and anterior margins of
the lorum.

The maxillary stylet (mxs, Fig. 15) is likewise broader at the base and
narrows to a long terminal bristle. It possesses a long retractor muscle
(rmxs) originating on the vertex of the head and a protractor muscle
(pmxs) originating on the maxillary plate. A membrane ensheaths the
protractor muscle and forms an enclosed lobe with the maxillary plate.
The base of the stylet is enclosed in a pouch formed by this maxillary
lobe, the hypopharyngeal wing plate (hwp), and the posterior tentorial
bridge (tb), but the stylet is actually outside the head capsule. A sclero-
tized lever (Ivr,) runs from the base of the stylet to an apodeme of the
head capsule, apparently a part of the posterior tentorial bridge. The
muscles and levers of both the maxillary and mandibular stylets are thus
within the head capsule; whereas the stylets alone are in pouchlike cham-
bers outside the head capsule. It should be noted that the mandibular and
maxillary stylets of this fulgorid each possess single protractor and re-
tractor muscles.

The labium (Fig. 17) appears to consist of four parts. A basal segment
invaginates inwardly to form an apodemal process. Beyond this basal seg-
ment there are a pair of sclerites on the posterior wall of the labium and
these are followed by two long segments. Anteriorly, the labium possesses
a groove along its entire length in which the bristles of the mandibular
and maxillary stylets lie. Apart from a narrow, median, sclerotized rod,
this groove is membranous. Protractor muscles (plb) and _ retractor
muscles (rlb) of the labium, originating on the margins of the hypo-
pharyngeal wing plates, respectively, are inserted on the apex and the
base of the apodemal process of the first segment. A median section of the
labium (Fig. 14) reveals a pair of muscles (mlb,) going from the paired
sclerites of the posterior wall to the median sclerotized process of the
anterior groove, a second pair (mlb, ) from the median sclerotized process
of the groove to the posterior wall, another small pair (mlb,) from the
lateral edge of the anterior groove to the posterior wall, and a fourth
group of muscles (mlb,) running from the median sclerotized process to
the posterior wall.

The hypopharynx is best seen in a median section of the head (Fig. 13).
Laterally the hypopharynx sends a pair of chitinous plates, termed hypo-
pharyngeal wing plates, dorsally where they meet the tentorial bridge
(hwp, Fig. 9). Muscles (dsyr) originating on the hypopharyngeal wing
plates are inserted on an apodemal process of the salivary syringe (syr)
which is located in the hypopharynx. A pair of salivary ducts (sld) from

10 MORPHOLOGY OF AUCHENORHYNCHA

the salivary glands unite to form a common median duct which empties
into the salivary syringe. The saliva is discharged through the ventral
tip of the hypopharynx into the salivary channel formed by the maxillary
stylets which come together at this point.

The dorsal or anterior wall of the hypopharynx forms the posterior or
ventral wall of the sucking pump (pmp). As Snodgrass has shown this
is not the true mouth but a portion of the preoral cavity termed by him
the cibarium. The anterior wall of the sucking pump is formed by the
inner epipharyngeal wall of the clypeus. This anterior wall of the suck-
ing pump gives off a series of tendons medially on which are inserted
the large dilator muscles (dlclp) originating on the postclyeus.

Of considerable significance are the lateral arms of the hypopharynx
‘lh) which extend to the loral plates (Figs. 9 and 10). These connecting
sclerotized arms between the hypopharynx and lorae are very small in
comparison to the broad connection existing in the other homopteran
families to be considered.

The tentorial structure consists only of a posterior tentorial bridge
(tb, Figs. 9 and 16) extending from the dorsal tips of the hypopharyngeal
plates across the posterior opening of the head capsule. Unlike the
cicadid, cercopid, cicadellid, or membracid, there is an apodemal con-
nection between the head capsule and this tentorial bridge. In other
Homoptera, as far as has been ascertained, the posterior bridge extends
across the occipital foramen, but the original connections with the head
capsule have been lost. In Oliarus aridus (Fig. 11) an actual invagination
or tentorial pit (ptp) appears to exist on either side of the tentorial
bridge between the hypopharyngeal wing plate and the maxillary plate.

The antenna (Fig. 7) consists of a small basal segment or scape (scp),
followed by a large bulblike pedicel (pdc) and a threadlike, single seg-
mented flagellum (fl). Three antennal muscles (mant), inserted on the
scape, have a common origin on the tentorial bridge (Fig. 9).

It is interesting to note the various modifications of the postclypeus
and the loral areas found in the fulgorids. As indicated previously, Evans,
Spooner, and others have considered the loral areas to be lateral modi-
fications of the postclypeus. Figures 2, 3, and 5 show some of these modi-
fications. In these fulgorids the lorum, more or less, completely merges
with the postclypeus. Of significance in homologizing this loral area
throughout the Homoptera is the fact that the protractor muscles of the
mandible always originate on this region of the head however it may
be developed, as shown in the lateral views of an undetermined species
of Acalonia (Fig. 1) and Ormenis pruinosa (Fig. 4). This will be a more
fully discussed in a later section on phylogeny. Oliarus aridus is shown
(Fig. 6) to indicate the presence of a median ocellus. A median ocellus is

SCLERITES, APPENDAGES, AND MUSCLES OF THE HEAD 11

apparently present among the fulgorids only in the subfamily Cixiinae,
and only in the Cicadidae of other Auchenorhyncha.

Tibicina septendecim (Linn. )
(Cicadidae )

Because of their large size, the head and mouthparts of the Cicadidae
have been the subject of more studies than any of the other Homoptera,
so that this group is fairly well known. Marlatt (1895), Snodgrass (1921,
1927), Myers (1928), Evans (1938, 1940), Butt (1943), and others have
made studies of representatives of this family. A description of Tibicina
septendecim together with additional drawings, however, is included
here for comparative study and interpretation.

The most conspicuous feature of the head is the greatly enlarged post-
clypeus (pcelp, Fig. 19). A strong internal ridge is present medially and
it is demarked from the anteclypeus (aclp) by a very definite suture. It is
at once apparent that in this respect the head of this cicada has reached
a far greater development than shown among the fulgorids. The domi-
nance of the clypeal area in this and the remaining families of Auchen-
orhyncha is in marked contrast to that found in the fulgorids where the
clypeal area, although fairly well developed, occupies a considerably
smaller portion of the total head area. The loral areas (lor) extend dor-
sally almost as far as the postclypeus itself. Internally a strong apodemal
ridge (clr, Fig. 25) is present between the lorum and postclypeus. The
labrum (lm) fits over the basal portion of the median groove of the
labium (lb).

On the dorsal surface of the head (Fig. 26) three ocelli (oc) are pres-
ent. A median ridge runs from the back of the head to just behind the
median ocellus and sends two, somewhat faintly divergent ridges ante-
riorly. The triangular region (fr) containing the median ocellus evidently
represents the frontal area. The anterior dorsal dilator muscle of the
pharynx (phym,, Fig. 18) originates on the anterior margin of this trian-
gular frontal area.

The mandibular stylet (Fig. 28) possesses retractor muscles (rmds)
inserted on an internal tendon given off by the elongate arm of the base,
which go to the dorsal part of the head. Mandibular protractor muscles
(pmds) originating on the lorum are inserted on the considerably en-
larged lever and apodeme of the stylet. The lever (Ivr,, Fig. 25) articu-
lates with the dorsal-most portion of the lorum just below the point where
the anterior tentorial arm (at) invaginates inwardly.

The maxillary stylet (Fig. 23) varies considerably from that found in
the fulgorid. Instead of one retractor muscle there are now two (rmxs,
and rmx. ), with separate insertions on the base of the stylet and sep-

2h MORPHOLOGY OF AUCHENORHYNCHA

arate origins on the dorsal wall of the head. In addition to the protractor
muscle (pmxs) originating on the maxillary plate, the strongly sclerotized
lever of the maxilla (Ivr.) also bears a protractor muscle (pmxl) originat-
ing on the maxillary plate and a retractor muscle (rmxl) originating on
the posterior tentorial bridge.

The labium (Fig. 21) consists of three segments: a basal segment, an
expanded middle segment, and an elongate terminal segment. A pair of
median protractor muscles (plb) inserted on the apex of the apodemal
process and a pair of lateral retractor muscles (rlb) inserted on the basal
segment both run to the hypopharyngeal wing plate. A median section of
the labium (Fig. 22) indicates that there are three pairs of muscles, one
pair (mlb,) extends from the terminal margin of the first segment to the
sclerotized rod and apodeme of the anterior median groove, a second
pair (mlb,) from the sclerotized rod to the lateral wall of the middle
segment, and a third pair (mlb, ) from the median rod to the dorsal wall
of the middle segment. No muscles were seen in the terminal segment.

The hypopharynx (hph, Fig. 18) contains the salivary syringe (syr)
and gives rise posterolaterally to the greatly enlarged hypopharyngeal
wing plates (hwp, Fig. 20) upon which the dilator muscles of the syringe
lever (dsyr) are attached. The comparative size of the hypopharyngeal
wing plates is seen in figure 27. United with the main body of the wing
plate laterally is a thin sclerotized plate which appears to be an apodemal
process (mxap, Fig. 20) of the maxillary plate. The lever of the maxillary
stylet articulates with this apodeme, and not directly with the maxillary
plate as has heretofore been figured and described by some writers. These
two posterior plates, together with anterior membranes which meet where
the dorsal tip of the hypopharyngeal wing plate unites with the lateral
extremity of the posterior tentorial bridge, form the pouch which contains
the mandibular and maxillary stylets.

The anterior wall of the hypopharynx forms the ventral, cup-shaped
floor of the sucking pump. To the roof of the sucking pump, which nests
concavely in the ventral floor, are attached the large dilator muscles
(dlclp). Contraction of these muscles arches the roof convexly and
creates the pressure which sucks the food into the preoral cavity. As can
be seen by lifting or removing the anteclypeus, the hypopharynx is
broadly joined laterally to the lorae. The tentorium in this cicada (Fig. 25)
consists of a posterior tentorial bar or bridge (tb) from which two arms
(at) run anteriorly to the anterior tentorial pit (atp, Fig. 19). Each of
these anterior arms gives off a dorsal arm (dt), to the apex of which the
antennal muscles are attached. The antenna consists of a basal scape,
a pedicel, and a terminal threadlike flagellum.

SCLERITES, APPENDAGES, AND MUSCLES OF THE HEAD 13

Lepyronia quadrangularis (Say)
(Cercopidae )

The study of the head and mouthparts of this family of Homoptera,
aside from that included in the study of the external structure and biology
of Lepyronia quadrangularis made by Doering (1922), has received al-
most no attention.

The postclypeus (pclp, Fig. 29) of Lepyronia quadrangularis is again
characteristically striated. There is no suture separating the anteclypeus
(aclp) from the postclypeus as occurs in the cicada, but its distinctness
is indicated by lateral indentations. The anteclypeus is elongated ven-
trally beyond the head capsule over the basal segment of the labium (lb).
The labrum (lm) suspended from the anteclypeus lies over the median
groove of the middle segment of the labium. Dorsally the postclypeus
continues into a quadrangular area (fclp, Fig. 31) in which is incor-
porated the frontal region or frons. That this quadrangular area repre-
sents some part of the frontal region is substantiated by the anterior dorsal
dilator muscle of the pharynx (phym,, Fig. 30), which takes its origin
from this region. The vertex (vx, Fig. 31) contains a pair of ocelli (oc).
The head of Lepyronia quadrangularis is actutely angular, when viewed
from the side, with the frons and vertex making up the dorsal surface.
The ventral surface, comprising the clypeal area and labrum, slants pos-
teriorly (Fig. 30).

The mandibular stylet (Fig. 34) is similar to that found in the cicada.
The protractor muscles (pmds) originating on the lorum are inserted on
the lever of the mandible (lvr,) and the retractor muscle (rmds) is
inserted on the narrow arm of the base of the stylet. The lever articulates
with the upper corner of the lorum, but it is situated a considerable
distance beneath the anterior invagination of the tentorium.

The maxillary stylet (Fig. 33) is drawn with the protractor. muscles
(pmxs) of both the base of the stylet and lever (lvr.) enclosed within
the membranous sheath of the maxillary plate to show the relationship of
these parts. A pair of retractor muscles (rmxs) is inserted on the base of
the stylet. The lever also possesses a retractor muscle similar to that in
the cicada and in fact the entire musculature of the maxillary stylet is
simliar to that of the cicada. The lever of the stylet articulates with the
apodemal plate of the maxillary lobe (mxap, Fig. :32) and not to the
maxillary lobe itself.

The three-segmented labium (Fig. 37) is comparatively short and
thickened. The protractor (plb) and retractor muscles (rlb) are similar
to those already described in the fulgorid and cicada. Five pairs of trans-
verse muscles are seen in the basal and middle segments but there are
apparently no muscles in the terminal segment.

14 MORPHOLOGY OF AUCHENORHYNCHA

The mouthparts (Fig. 30) are very similar to those described for the
cicada. The salivary syringe (syr), however, is well developed and com-
paratively large. The dilator muscles of the salivary syringe (dsyr) are
of unusual size and are seen to occupy a very large proportion of the
posterior head opening (Fig. 32).

The tentorium, as in the cicada, consists of a posterior tentorial bridge
(tb, Fig. 32), and a pair of anterior arms (at, Fig. 34), each with a short
dorsal arm (dt). The antennal muscles (mant) originate on the apex of
the dorsal tentorial arm.

The left half of the head of Lepyronia quadrangularis, with the lower
part of the postclypeus, the anteclypeus, labium, and labrum dissected
away, is shown in figure 35 to reveal the relationship of the lorum (lor)
to the hypopharynx (hph). There is a broad ridge, normally covered by
the anteclypeus, which joins the lorum to the hypopharynx. This con-
nection between the lorum and hypopharynx is similar to that which is
present in the cicada. In addition there exists an internal ridge or apodeme
where the lorum and postclypeus meet, termed the clypeal ridge (clr,
Figs. 34 and 35).

A cross-section of the head through the sucking pump, stylet pouches,
and muscles of the mouthparts is shown in figure 36 to indicate their
relationships.

Aulacizes irrorata (Fabr.)
( Cicadellidae )

Evans has made a comprehensive taxonomic and morphological study
of this group over a period of ten years and in 1938 provided a valuable
evaluation of intrarelationships of this family based on an analysis of their
morphological characters. He considers that the external structures of the
head are second in value only to the venation of the wings for purposes
of comparative studies and the determination of phylogenetic relation-
ships. Although this group is here designated as a family for purposes of
uniformity, he considered it to be a superfamily, the Jassoidea, and di-
vided it into twenty-one separate families. Later (1948) he reduced the
familial divisions to eight, including three families of membracids.

The head of Aulacizes irrorata is in many respects similar to the cerco-
pid, Lepyronia quadrangularis. The clypeal region, however, is a little
larger and bulges out considerably more than in the cercopid. In ventral
view (Fig. 38) the large striated postclypeus (pclp), although merged
with the anteclypeus (aclp) beneath, is demarked from it by lateral
indentations. Laterad of the anteclypeus are the loral regions (lor).
The small labrum (lm) fits over the median groove of the short, compact
labium (Ib). Dorsally the postclypeus merges with the frons to form a
frontoclypeal region (fclp, Fig. 39). Although it is not possible to deter-

SCLERITES, APPENDAGES, AND MUSCLES OF THE HEAD 15

mine the exact extent of the frons, it should be noticed that the anterior
dilator muscle of the pharynx (phym,, Fig. 40) originates just ventrad of
the dorsal surface so that the postclypeus probably does not extend be-
yond this point. The posterior limits of the frons may possibly be marked
by the prominent lateral indentations. The vertex (vx) bears a pair of
ocelli (oc). The antenna consists of a basal scape and pedicel and a long
threadlike flagellum which shows marks of segmentation basally.

The musculature of the mandibular stylet (mds, Fig. 46) is uniquely
different from that of the preceding forms. In addition to the protractor
muscle (pmds) running from the lever (Ivr,) to the lorum and the re-
tractor muscle (rmds,) going from the base of the stylet to the vertex,
there is a second retractor muscle (rmds,) which goes to the anterior
tentorial arm (at).

The base of the maxillary stylet (mxs, Fig. 42) has one large retractor
muscle (rmxs) which originates on the vertex and a pair of protractor
muscles originating on the maxillary plate. One protractor muscle (pmxs)
is inserted near the base of the stylet, the other (pmxl) is inserted on the
lever (lvr.). The lever is broadly articulated to the apodemal plate of the
maxillary (mxap). |

The short, compact labium (Fig. 43) consists of three segments. Two
pairs of small triangular plates are present posteriorly on the terminal
segment, one pair located basally, one pair apically. Evans (1937) has
suggested that these paired plates may represent the glossae and that the
surrounding terminal segment is derived from the paraglossae. Likewise,
the basal and middle segments may represent the “mentum” and “pre-
mentum. These latter terms are of little value, however, for it has so
long been the custom to give names indiscriminately to various divisions
of the labium that the actual homology of the parts to which such terms
are assigned is doubtful. The usual pair of protractor (plb) and retractor
muscles (rlb) originating on the hypopharyngeal plates are inserted on
the apodemal process and base of the labium. A median section of the
labium (Fig. 44) reveals six pairs of transverse muscles.

The parts of the hypopharynx and the sucking pump mechanism are
very similar to those parts already described in previous forms. It should
be noted, however, that the sucking pump cavity when fully distended is
unusually large, and together with the well developed dilator muscles
(dlclp) indicates that this form possesses a powerful sucking apparatus.

The tentorium consists of a posterior tentorial bridge (tb, Fig. 41)
and a pair of anterior tentorial invaginations (at, Fig. 46) to which the
dorsal tentorial arms (dt) are attached. The pair of anterior tentorial
arms usually running to the posterior bridge is absent. Instead, where the
anterior tentorial arm might be expected there is found the tendon from

16 MORPHOLOGY OF AUCHENORHYNCHA

which arises the retractor muscle of the maxillary stylet. This tendon may
in part be a vestige of the anterior tentorial arm. The dorsal tentorial arm
can be identified by the antennal muscles (mant) which have their origin
on the apex of this part.

Ceresa bubalus (Fabr. )
(Membracidae )

The head and mouthparts of the Membracidae have been treated in
the earlier comprehensive works on the taxonomy, biology, and morphol-
ogy ot this group by Branch (1914), Funkhauser (1917), and Lawson
(1922). These have provided very desirable and basic information con-
cerning the Membracidae, but certain inaccuracies exist in these studies
because the homologies of the head and mouthparts were not as well un-
derstood then as they are now. These works have not provided any con-
sideration of the musculature.

The head of Ceresa bubalus is considerably flattened anteroposteriorly.
Seen in ventral aspect (Fig. 47) the conspicuous median area is the post-
clypeus (pclp), beneath which is the anteclypeus (aclp). The labrum
(Im), suspended from the anteclypeus, fits over the median groove of
the three-segmented labium (lb). The lorae (lor) are located on the sides
of the clypeal areas. The postclypeus is continuous on the conspicuous
median protuberance of the anterior surface of the head (Fig. 48). At
least a small dorsal portion of this protuberance representing the frontal
region of the head combines with the postclypeus to form a frontoclypeus
(fclp ), since the anterior dilator muscles of the pharynx (phym,, Fig. 49)
have their origin here. The vertex (vx), divided by a median ridge, pos-
sesses a pair of ocelli (oc). The vertex, which lies dorsally in the previous
forms discussed, is anterior in position in a line with the metopidium of
the pronotum, while most of the clypeal region, normally anterior, lies
ventrally.

The mandibular stylet (Fig. 54) has a large group of protractor muscles
(pmds) inserted on the lever (Ivr,) going to the lorum, and three re-
tractor muscles. The first of these muscles (rmds,) originates on the
dorsal region of the head capsule, the second (rmds.) on the dorsal arm
of the tentorium and the third (rmds,) originates on a lateral internal
edge of the hypopharyngeal wing plate where it meets the margin of the
head capsule. All three are inserted on the basal portion of the mandibular
stylet. |

The maxillary stylet (Fig. 53) is greatly elongated and its pouch has
been invaginated beyond the tips of the hypopharyngeal wing plates
(Fig. 50), so that it is separated from the vertex of the head by a pair of
short retractor muscles. There are two protractor muscles originating on

SCLERITES, APPENDAGES, AND MUSCLES OF THE HEAD ke

the maxillary plate (mxp), one (pmxs) inserted at the base of the stylet,
the other (pmxl) inserted on the lever (Ivr.). The lever itself appar-
ently articulates with the lateral edge of the hypopharyngeal wing plate
(hwp ). It should be noted, however, that the hypopharyngeal wing plate
has merged with the apodemal plate of the maxillary (mxap). This makes
it appear that in this membracid the wing plate is formed from both the
lateral elongation of the posterior of the hypopharynx and an invagina-
tion of the maxillary plate.

The three-segmented labrum (Fig. 51) resembles that of the leaf-
hopper Aulacizes, in that the posterior surface of the terminal segment
bears a pair of almost triangular plates basally. Paired protractor muscles
(plb) and retractors (rlb) originating on the hypopharyngeal wing plate
are inserted on the apex of the apodemal process and the basal segment,
respectively. A median section of the labium (Fig. 52) indicates that
four pairs of muscles are present within its three segments.

The hypopharynx (hph, Fig. 49) contains a small salivary syringe
(syr), the dilator muscles of which (dsyr, Fig. 50) are attached to the
hypopharyngeal wing plates. The sucking pump (pmp) is very similar to
those of the previous forms. The epipharyngeal wall upon which the
dilator muscles (dlclp) are inserted forms the anterior wall of the pump,
while the anterior wall of the hypopharynx forms its posterior wall. The
pump leads into the pharynx. It should be noted that the frontal con-
nective nerve from the brain runs anterior to the anterior dorsal dilator
muscles of the pharynx. As Snodgrass (1947) has shown, this nerve and
its ganglion separates the preoral muscles, which include the dilator
muscles of the sucking pump (dlclp), from the postoral muscles, which
include the hypopharyngeal muscles and pharyngeal dilator muscles.
Behind the brain there are two pairs of dorsal dilator muscles of the
pharynx (phym., phym,) going to the top of the head capsule and a
lateral group (phym,) going to the tentorium.

The tentorium consists of a posterior tentorial bar, or bridge (tb), and
the short dorsal arm of the tentorium (dt, Fig. 54) which arises from the
anterior invagination of the tentorium. The antennal muscles (mant), as
usual, have their origin on the dorsal arm of the tentorium. The anterior
arm of the tentorium, partly incorporated into the base of the dorsal arm,
has otherwise disappeared. The antenna consists of two basal segments,
the scape (scp) and pedicel (pdc), and a terminal flagellum (fl).

THE THORACIC SCLERITES, WINGS, AND LEGS
OF AUCHENORHYNCHA

The thoracic sclerites of the Auchenorhyncha, and the Homoptera gen-
erally, have received relatively little study. Aside from the general studies
of Crampton (1926) and Snodgrass (1927) which contain references to
individual Homoptera, the most extensive comparative studies of the
thoraces of the Homoptera is that of Taylor (1918). The wings, however,
have received better treatment and there exist good comparative studies
of the wings of the Fulgoroidea, Cicadellidae, and Cercopidae by Met-
calf (1913, 1917a, 1917b) and of the Membracidae by Funkhauser
(1913). Myers (1928) has discussed the thorax and wings of the Cicadi-
dae in his comparative study on the morphology of this family.

Scolops pungens

Prothorax. The head is closely joined to the prothorax by the cervical
membrane. There are no distinct cervical sclerites present. The prothorax
(Fig. 55) consists mainly of a saddle-shaped pronotum which covers the
dorsum and extends down over most of the pleural region. The pleural
region is considerably reduced. A portion of the episternum (es,) extends
beyond the ventral margin of the lateral lobe of the pronotum, but the
epimeron (epm,) is completely covered. Between these pleural sclerites
is the pleural suture (pls,) which extends into the coxal process (cxp,),
a pleural articulation for the coxa of the leg.

The prosternum (st,, Fig. 60) consists of a small sclerite connected to
the episternum on each side by a narrow, elongated precoxal bridge or
precoxale (pcx,). The trochantin (tn) is a small sclerite which forms a
point of articulation for the leg between the episternum and precoxale
and the coxa.

A pair of invaginations or furcal pits (fp,) give rise to a pair of sternal
apodemes or furcae (fu, Fig. 63). Each furcal apodeme unites with a
pleural apodeme (plap.) represented externally by the plural suture.
Together, these sternal and pleural invaginations represent the prothoracic
endoskeleton.

Mesothorax. The tergum of this wing-bearing segment (Fig. 56) is com-
posed of several distinct regions, the most conspicuous of which is the
large scutum (sc,). Anteriorly a narrow ridge represents the precosta or
pretergite (pc.). Behind the scutum is a small scutellum (scl,), the pos-
terior margin of which forms a fold or reduplication (rd, ). This posterior
fold of the scutellum continues into the axillary cord (axc.), forming a

18

THORACIC SCLERITES, WINGS, AND LEGS 19

connection with the posterior margin of the wing. The postnotum (pnt. )
although morphologically a part of the mesothorax is more closely asso-
ciated with the metathoracic tergum. Laterally it unites with the meso-
thoracic epimeron (epm.) by means of the postalar bridge (pw.). An-
teriorly the prealar bridge (aw.) unites with the episternum.

The dorsal region of the forewing is articulated to the metathoracic
tergum by means of a basal membranous area within which are a number
of articular sclerites, collectively termed the pteralia. The first of these
sclerites, the first axillary (lax., Fig. 56) articulates mesally with the
anterior notal wing process (anp.) and laterally with the second axillary
(2ax.). The second axillary in turn articulates with the common base of
the subcosta and radius of the wing veins and with the third axillary
(3ax.), while the latter axillary is united with the base of the vannal
region of the wing. The basal margin of the vannal region forms a sclero-
tized fold or lip which fits into a groove formed in the scutellar region of
the tergum, locking the wings in place when they are folded at rest.

The pleuron (Fig. 55) is composed of the episternum (es,) and the
epimeron, separated by the pleural suture (pls.). Dorsally these areas
continue along the pleural suture to form the pleural wing process or
articulation for the wing (pw-.), while the pleural coxal process (cxp,)
articulates ventrally with the coxa. The mesothoracic spiracle (sp.) lies
in the membranous region below the prealar bridge.

The membranous ventral area of the wing which unites the wing to
the pleuron contains several small sclerites, which serve for the insertion
of important wing muscles. These are termed the alaria by Crampton and
the epipleurites by Snodgrass. Anterior to the pleural wing process is
the basalar sclerite (ba.) united to the episternum. Behind the pleural
wing process is a narrow elongate subalar sclrite (sa,) which apparently
unites with the postalar bridge.

On the ventral side, the sternum (st, ) is broadly joined to the epister-
num by means of the precoxale (pcx.). The trochantin (tn,) lies be-
tween the precoxale and the coxal cavity. A pair of furcal pits (fp.) form
invaginations of the sternum which give rise to branched furcal apodemes
internally. The lateral branches of these apodemes unite with the pleural
apodemes as in the prothorax. The mesothoracic pleural apodeme is con-
siderably developed dorsally and extends into the body cavity just below
the tergum.

Metathorax. The mesothoracic scutellum overlies a considerable portion
of the metathoracic tergum, and the normal position of the two wing-
bearing segments has been displaced in figure 56 to reveal the divisions
of the metatergum. As already noted the postnotum (pnt, ) belonging to
the mesothoracic segment, is closely joined to the prescutum (psc,) of

20 MORPHOLOGY OF AUCHENORHYNCHA

the metathorax. The antecostal suture (acs,) which separates the post-
notum from the prescutum is the external demarcation of the poorly de-
veloped metathoracic phragma. The prescutum is separated fram the
scutum (sc;) by elongate membranous areas on each side. The scutellum
(scl, ) is poorly developed. Its posterior margin folds over to form a redu-
plication and continues into the posterior margin of the wing by means
of the axillary cord (axc;). A broad postnotal region (pnt;) is closely
united to the scutellum. The postnotum gives off a pair of narrow arms
laterally along the posterior margin of the pleuron to meet on the venter,
forming a postcoxal bridge (pocx;). The postcoxal bridge on each side
unites with a median triangular sclerite. The apex of this triangle sends
an elongate sclerotized process anteriorly between the large membranous
area of the coxae, which bifurcates into the body cavity.

The dorsal basal membranous area of the hind wing has a number of
articular sclerites. A small first axillary (lax;) lies between the anterior
notal process (anp;) and the larger second axillary (2ax;). Posteriorly
the third axillary (3ax,) articulates with a prominent posterior notal
process (pnp;).

The metathoracic pleuron (Fig. 55) presents a specialized structure,
apparently correlated with the development of jumping hind legs. The
episternum (es,), demarked from the epimeron (epm;) by the plural
suture (pls;), is cleft ventrally by membrane. Both the episternum and
the epimeron have been shifted in position so that these sclerites lie
almost horizontally, rather than vertically. Consequently the pleural wing
process (pwp;) has been pushed to the extreme anterior point of the
pleuron. The coxa has become rigidly united with the pleuron and al-
though the coxa (cx;) and meron (mr;) may be broadly distinguished
from the episternum and epimeron, considerable comparative study of
this region among the fulgorids would be required to determine the exact
boundaries of these sclerites. Taylor (1918) has indicated, however, that
in species of Ormenis and Fulgora, a comparable region bears muscles
extending to the subalar region, and since such muscles connect the
subalar sclerite and meron in other insects he concluded that the region
in question is a part of the coxa, and not of the pleuron.

The ventral basal membrane of the wing does not possess an externally
visible basalar sclerite anterior to the pleural wing process as does the
mesothoracic segment. Posteriorly, however, there is an elongate subalar
sclerite (sag).

On the venter the sternum is united both to the mesothoracic sternum
anteriorly and to the episternum laterally through a precoxal bridge
(pex;, Fig. 60). A wide membranous area separates the precoxal bridge
from the coxae. The origin of the small sclerotized areas within this mem-

THORACIC SCLERITES, WINGS, AND LEGS 21

branous region is uncertain, but may have been derived from the tro-
chantin.

Wings. The forewing is elongate-oval in outline and is somewhat
thickened, forming a tegmen-like covering for the membranous hind
wing. A vannal fold (vf) divides the hind wing into an anterior remigium
and a posterior vannus. The basic wing venation is noted in figures 79

and 80.

Legs. The mesothoracic leg (Fig. 80) consists of a basal coxal segment
(cx,), a short trochanter (tr), the femur (fe), tibia (ti), and a three-
segmented tarsus. The tarsus (Fig. 92) bears a pair of ungues or claws
(cl), between which is a membranous pad or pulvillus (pu). The basal
rim of the coxa is demarked at the basicoxa (bex) and a small lateral
portion forms the meron (mr). The prothoracic leg (Fig. 89) is almost
identical to the mesothoracic leg in structure, except that the meron (mr, )
is less developed. The larger metathoracic leg (Fig. 91) is modified for
jumping and consists of a coxal segment (cx), the meron (mr;) of which
is rigidly joined to the pleuron. There is a small trochanter, a femur, and
a tibia with a posterior row of toothlike spines, and a border of spines
apically. The apical ends of the first two tarsal segments are similarly
spined.

Tibicina septendecim

Prothorax. The cervical region of the cicada, Tibicina septendecim, con-
tains a pair of median dorsal cervicals (dc, Fig. 57) and a pair of lateral
cervical sclerites (lc).

L‘ke the fulgorid, the principal sclerite of the prothorax of the cicada
consists of a saddle-shaped pronotum which covers most of the pleural
region. In other respects, too, it is quite similar, except that there is a
postcoxal bridge (pocx,, Fig. 61) ventrally as well as a precoxal bridge
(pex). In addition, the median sternal region (st,) is modified into a
troughlike structure in which the labium of the head lies in repose.

Mesothorax. The main feature of this segment is that it bears the prin-
cipal burden of flight of the cicada. As a consequence this segment is
greatly enlarged and presents a number of modifications primarily corre-
lated with the flight mechanism. The tergum consists of an anterior pre-
scutum (psc., Fig. 57) with the scutum (sc.) lying posteriorly and lat-
erally. There is a narrow precostal region (pc,) present anteriorly, which
is bent back over the prescutum, and this relationship can best be seen in
longitudinal section shown in figure 59. The phragma (ph.) which is
fairly well developed is not suspended vertically, but is directed anteriorly
in a horizontal direction and can be seen dorsally beneath the prothoracic
intrasegmental membrane.

22 MORPHOLOGY OF AUCHENORHYNCHA

The postnotum (pnt.), although closely united with the metathoracic
tergum medially, is also joined to the reduplication (rd.), or posterior
fold of the scutellum (scl. ), laterally. The invaginated plate arising from
the posterior margin of the postnotum gives rise to a large, well developed
phragma (ph., Fig. 62) upon which arise the powerful oblique dorsal
muscles of the mesothoracic tergum. The phragma is thus anchored both
to the mesothorax laterally and to the metathoracic segment medially.
Four axillary segments, in the basal membranous region of the dorsal
surface of the forewing, provide a part of the tergal articulating mech-
anism.

Laterally (Fig. 58), the episternum and epimeron are respectively
divided into upper anepisternal (aes,) and anepimeral (aepm,) regions
and below into katepisternal (kes.) and katepimeral (kemp.) regions.
Connected to the episternum in the basal membranous area of the ventral
surface of the wing is the basalar sclerite (ba,) while the subalar sclerite
(sa,) is behind the pleural wing process (pwp.).

The mesosternum consists of a broad basisternum (bst., Fig. 61) sep-
arated from the prosternum by a membranous region, but joined laterally
to the katepisterna by precoxal bridges (pcx. ). The medioposterior fur-
casternum (fst.) contains a pair of furcal pits (fp) which invaginate to
form the furcal apodemes internally. Postcoxal bridges (pocx.) join the
furcasternum to the katepimera.

Metathorax. This segment is considerably reduced in comparison to
the mesothorax, and although the hindwing which it bears apparently
plays a lesser role in the flight of this insect, it does nevertheless actively
enter into the flight mechanism. The scutellum of the mesothorax overlies
half of the metathoracic tergum, but the latter has been withdrawn in
figure 57 to reveal its parts. To be noted is the fact that medially the
scutum (sc;) is separated from the scutellum (scl;) by a membranous
cleft. A small postnotum (pnt,) is present behind the scutellum. The
basal membranous region of the dorsal surface of the wing contains four
axillary sclerites.

Ventrally the basisternum (bst;) is rigidly united with the mesoster-
num anteriorly and joined to the pleuron by precoxal (pcx;) and post-
coxal bridges (pocx,;), while the furcasternum (fst, ) is posteriorly united
with the sternum of the first abdominal segment (I).

Wings. The forewing (Fig. 81) is almost twice as large as the hind
wing (Fig. 82) and is apparently the principal flight organ, although as
has been mentioned the hind wing is undoubtedly an equally active but
lesser organ of flight. This is emphasized since the development and
sclerotization of the wing-bearing segments is closely correlated with the
degree of activity of both the fore and hind wings. Care must be taken

THORACIC SCLERITES, WINGS, AND LEGS 23

not to mistake structural correlations with the flight mechanism for phylo-
genetic relationship.

Attention should be called to the nodal line of the fore wing (nl)
which forms a break in the sclerotization of some of the principal vein
branches, previously thought to be present only in the Cicadidae. Evans
(1948) has shown that a nodal line is also present in the jassoid family,
Hylicidae. The basic venation, noted in figures 81 and 82, is well discussed
by Myers (1928) for the Cicadidae as a whole.

Legs. The legs of the cicada bear the same segmentation as noted in the
fulgorid. The prothoracic leg (Fig. 93) is turned anteriorly and is so
drawn. The femur (fe) is considerably thickened and bears several spines
on its ventral margin. The mesothoracic leg (Fig. 94) shows no particular
modification. The meron (mr,) of the metathoracic leg (Fig. 95) is elon-
gate and pointed, forming a projecting spinelike structure. The tibia bears
several spinelike, movable setae and a row of spines apically. The distal
segment of the tarsus (Fig. 96) bears a pair of claws (cl) but no pulvillus.

Lepyronia quadrangularis

Prothorax. The pronotum (prnt, Fig. 67) of the cercopid, Lepyronia
quadrangularis, is typically saddle-shaped, covering most of the pleuron
laterally. Posterodorsally it overlies the anterior part of the mesothoracic
tergum, but the pronotum has been displaced anteriorly in figure 66.
The epimeron is completely reduced and concealed by the lateral lobe of
the pronotum. The sternum (st,, Fig. 68) forms a trough for the recep-
tion of the labium. In other respects it varies but little from the forms
already discussed.

Mesothorax. A narrow precosta (pc,) is demarked anteriorly on the
tergum, while the prescutum (psc,) behind is separated laterally from
the scutum (sc.) by a pair of narrow membranous clefts. The conspicuous
scutellum (scl.) is triangular in shape and its posterior half normally
overlies the metathoracic tergum (displaced in Fig. 66). As in the ful-
gorid the posterior lateral margins of the scutellum are grooved to form
a locking device (twg.) with the bases of the posterior margin of the
tegmina.

Laterally, the episternum is divided into an anepisternum (aes,) and a
katepisternum (kes.). The epimeron (epm,) and a part of the meta-
thoracic episternum (es;) is grooved to receive the base of the costal
margin of the tegmen. The episternum is broadly united to the sternum.
The sternum is differentiated into an anterior prebasisternite (pbs.), the
large basisternite (bst.), and a furcasternum (fst.) between the coxal
cavities. The furcasternite may possibly be a combination of the furcas-
ternite and the spinasternite found in the orthopteroid orders.

24 MORPHOLOGY OF AUCHENORHY NCHA

Three axillary sclerites are present dorsally in the basal membrane of
the wing, while ventrally there is a well developed subalar sclerite (sa. )
and a considerably reduced basalar sclerite. The mesothoracic spiracle
(sp, Fig. 67) is located in the membranous region of the pleuron between
the pro- and mesothorax.

Metathorax. This segment is well developed and apparently plays an
important role in the flight mechanism. In this connection it should be
noted that the well developed phragma is an invagination of the anterior
margin of the prescutum (psc;) and is primarily a functional part of the
metathorax, in contrast to the cicada in which the well developed
phragma is functionally a part of the mesothorax. Further, there is no
pronounced development of a postnotum between the mesothoracic and
metathoracic terga. The scutum (sc;) is large and contains a median
suture. This median suture (ms,) is the external demarcation of a well
developed median tergal apodeme which unites anteriorly with the
phragma. The scutellum (scl;) forms a narrow posterior margin of the
tergum.

Laterally the anterior margin of the tergum gives off two branches. The
anterior of these is the postalar bridge (pw.) of the mesothoracic seg-
ment and unites with the mesothoracic epimeron, while the posterior
branch forms the prealar bridge (aw; ) of the metathoracic segment and
unites with the metathoracic episternum (es;). The epimeron (epm;) is
united to the postnotum (pnt;) by a postalar bridge (pw,). The meta-
thoracic spiracular opening is concealed in the dorsal angle between the
mesothoracic postalare and the metathoracic prealare.

The sternum (st;) is rigidly joined to the mesothoracic furcasternite
anteriorly and united with the episternum by a narrow bridge. Although
a pair of membranous clefts demark the central region, there is no dis-
tinct demarcation of the lateral limits of the sternum.

The basal region of the wing contains three axillary sclerites dorsally,
the first axillary sclerite being considerably reduced. Ventrally there is
present a subalar sclerite (sa,;) but there is no external evidence of a
basalar sclerite. The metathoracic spiracle, although not readily visible,
is present in the grooved area formed between the mesothoracic epi-
meron and the metathoracic episternum in the upper region of the
pleuron.

Wings. (Figs. 83 and 84). The fore wing or tegmen is opaque and retic-
ulated, so that some of its venation is obscured. Doering (1922) has been
able to delineate the forewing venation by clearing and the venation
which she has obtained through this procedure is shown in broken line.
A projecting ridge or keel on the underside of the subcosta of the tegmen
latches on to a sharp crest of the metathoracic epimeron and helps to lock

THORACIC SCLERITES, WINGS, AND LEGS PA

the tegmen in place when at repose. The membranous hindwing bears a
row of spines or hooks on the costal margin. The basic venation of both
fore- and hindwings is indicated in figures 83 and 84.

Legs. The prothoracic (Fig. 97) and mesothoracic legs (Fig. 98) are
generally similar, except that the meron (mr.) of the middle leg is pro-
duced into a flat spinelike structure. The hind legs (Fig. 99) are consid-
erably modified. The meron (mr;) forms a ball-like structure which fits
into a socket formed by the epimeron, so that although not united to the
epimeron, it is closely associated with it. The tibia (ti,;) has two large
spines along its outer edge and a double row of spines at its apex. It is
this character of the tibia which is used to distinguish the Cercopidae
from the Cicadellidae. The first two tarsal segments also bear an apical
row of spines. The last tarsal segment of all the legs bears a pair of claws
(cl) and a notched, but single lobed pulvillus (pu), as shown in the
tarsus of the mesothoracic leg (Fig. 100). The hind leg of the cercopid
bears a considerable resemblance to that of the fulgorid in these respects.

Aulacizes irrorata

Prothorax. The prothoracic tergum takes the shape of a saddle-shaped
pronotum as in the previous forms described. It is shown in figure 65 in
dotted outline in its normal position covering the mesothoracic tergum.
The lateral and ventral aspects (Figs. 64 and 69) of this segment do not
vary significantly from that of Lepyronia quadrangularis above, except
that there is no development of a troughlike structure on the sternum as
in the cercopid.

Mesothorax. The mesothoracic precosta (pc, Fig. 65) consists of a
narrow sclerite, separated from the prescutum (psc.) by the antecostal
suture (acs). The antecostal suture is represented internally by a ridge,
not by the development of a phragma. The major portion of the tergum
is occupied by the scutum (sc.). The posterior tergum consists of the
scutellum (scl,), bearing a V-shaped groove (twg,) for the reception of
the basal lip of the posterior margin of the tegmen. The postnotum
(pnt. ), completely covered by the scutellar region of the tergum, gives
off a postalar arm (pw:.) laterally to the epimeron, whereas the anterior
portion of the tergum is united to the episternum by the prealar bridge
(aw, ). The membranous base of the wing contains three axillary sclerites
dorsally and both a basalar (ba.) and subalar sclerite (sa.) ventrally.

The episternum of the pleuron is divided into an anepisternum (aes»,
Fig. 64) and a katepisternum (kes.). The diagonal line (pwg), dividing
the epimeron (epm,) into two triangular areas, represents a groove into
which fits the basal lip of the costal margin of the tegmen, similar to the
arrangement found in the cercopid. The mesothoracic spiracle (sp.) is

26 MORPHOLOGY OF AUCHENORHYNCHA

located in the upper pleural membranous region between the pro- and
mesothorax.

The mesothoracic sternum is also similar to that of the cercopid, there
being an anterior prebasisternite (pbs., Fig. 69), a large basisternite
(bst.) and a posterior furcasternite (fst.) between the coxal cavities.
The basisternite is divided into two by a median suture, and its posterior
margin is demarked by another suture. The furcasternite bears the furcal
pits (fp.) which form an internal furcal apodeme. Laterally the sternum
is broadly joined with the pleuron.

Metathorax. The mesothoracic postnotum (pnt., Fig. 65) lies beneath
the posterior fold of the mesothoracic scutellum. Morphologically the
postnotum is a part of the mesothorax, but as in the cicada it is medially
united with the metathorax and laterally with the mesothorax. The large
phragma to which the postnotum gives rise through an invagination of its
posterior margin is more closely allied with the mesothorax functionally.
In this respect it differs from the intersegmental phragma between the
meso- and metathoracic segments of the cercopid, which phragma is inti-
mately associated with the metathorax.

The metathorax has no definite prescutal area. The tergum is composed
of a large scutum (sc;) and a smaller scutellum (scl, ). The metathoracic
postnotum (pnt,) behind the scutellum is only sclerotized laterally. A
postalar bridge connects the postnotum to the epimeron. Anteriorly, how-
ever, there is no distinct prealar bridge to connect the metathoracic ter-
gum to the episternum.

The pleuron consists of a distinct episternum (es,, Fig. 64) and epi-
meron (epm,) separated by the plural suture (pls;). The metathoracic
spiracle (sp;) is located in the membranous region of the pleuron, be-
tween the meso- and metathorax. The mesothoracic sternum (bst;, Fig.
69) has a precoxal connection laterally with the episternum. It sends a
median process posteriorly between the coxae, where it unites with a
postcoxal bridge given off by the epimeron. The trochantin (tn,) of the
coxa is quite large and apparently united with the episternum.

Wings. The forewing, or tegmen, is reticulated and thickened. Like
the tegmen of the cercopid it is divided by the vannal fold (vf, Fig. 85)
into an anterior remigial portion and a smaller posterior vannus. The
hindwing (Fig. 86) is membranous, and quite similar, in venation and
the presence of a series of hooks on the costal margin, to that of the
cercopid. In addition to the vannal fold (vf) it has a jugal fold (jf) which
demarks the jugum of the wing. The basic wing venation is indicated in
figures 85 and 86.

Legs. The pro- (Fig. 101) and mesothoracic legs (Fig. 102) are com-
paratively small and similar in appearance. The metathoracic leg (Fig.

THORACIC SCLERITZS, WINGS, AND LEGS 27

103) is several times larger. Its tibia (ti) is square in cross-section, pos-
sesses a row of spines along each of four edges, and is fringed with a
double row of spines apically. As with the cercopid and fulgorid, the first
two tarsal segments are fringed with spines apically. The claws (cl, Fig.
105) borne apically by the third tarsal segment are adherent laterally to
the bilobed pulvillus (pu, Fig. 104) in all three legs.

Ceresa bubalus

Prothorax. The membranous cervical region of Ceresa bubalus does not
contain any cervical sclerites. There are present, however, the paired,
triangular, occipital condyles (occ, Fig. 50) borne laterally by the occipi-
tal margin of the head.

The unusual development of the pronotum is characteristic of the
Membracidae. This structure, important taxonomically, takes on a re-
markable variety of shapes throughout the family, developing spines,
horns, crests, bulbous structures, and other developments, posteriorly,
anteriorly, dorsally and at the humeral angles. Unusual developments of
the pronotum are particularly evident in many of the tropical and sub-
tropical forms of this group. According to Funkhauser, the anterior,
dorsal, and humeral structures vary haphazardly among the species, and
although of almost no value in the differentiation of the higher taxonomic
categories, are of considerable value in the delineation of species. The
posterior structures, on the other hand, are of a more constant nature and
are frequently used for the differentiation of genera.

The cephalic portion of the pronotum, or metopidium (mtp, Fig. 72),
of the pronotum, of Ceresa bubalus is on a plane continuous with the
vertex of the head, and though marked with impunctations, does not pos-
sess any special structures. The slightly protruding areas on the lateral
margins of the prothorax just above the bases of the forewings are
termed the humeral angles (ha), and just above the humeral angles are
the lateral projections of the metopidium known as the suprahumeral
horns (sh). Obliquely posterior from the suprahumeral horns, a pair of
folds meet mesally to form a sharp median edge or carina. The posterior-
most portion of the median carina projects as a spinelike posterior process.

The pleural region (Fig. 75) consists of a definite episternal region
(es,) bounded laterally by the pleural suture (pls,). A well defined
epimeron does not exist, although the lateral lobe of the pronotum just
behind the pleural suture may include a portion of the epimeral region.
The episternum is turned inwardly immediately behind the head capsule
against which it lies. It is joined to the sternum by the precoxa (pcx, ).

A posterior view of the prothorax (Fig. 72) reveals a pair of furcal
pits (fp:) in the small rectangular sternal plate (st,). The furcal pits

28 MORPHOLOGY OF AUCHENORHYNCHA

mark the invaginations of the furcal apodemes (fu,) which possess two
branches. One branch runs dorsally, and the other curves laterally and
abuts against the pleural apodeme (plap,). The triangular trochantin
(tn,, Fig. 75) lies between the episternum and the coxa.

Mesothorax. This segment is large and well developed. The tergum
(Fig. 70) is primarily composed of a large anterior scutum (sc.) and a
posterior scutellum (scl.). The scutum possesses two pairs of lateral
sutures or internal ridges. One pair of such sutures or notaulices (no, )
arises anterolaterally and converges toward the median line, while the
second pair runs posteromedially from the anterior notal wing process
{anp,). A narrow precosts or pretergite (pc., Fig. 78) is set off from the
anterior margin of the scutum by the antecostal suture (acs, ). Posteriorly,
the scutellum covers the narrow postnotum (pnt,) which is united with
the metathoracic tergum. The bases of the wings are shown in figure 70,
slightly displaced from their normal position at rest. There are three axil-
lary sclerites (lax., 2ax,, 3ax,) in the basal membranous area of the fore-
wing.

The pleuron (Fig. 71) consists of an episternum (es,) and epimeron
(epm,) separated by the pleural suture (pls.). The extensions of these
sclerites dorsally along the pleural suture form the pleural wing process,
or articulation for the wing (pwp,), while the pleural coxal process, or
articulation for the coxa (cxp,) is formed by the extension of the epi-
sternum and epimeron at the ventral extremity of the pleural suture. As
in the cercopid and cicadellid previously described, the epimeron pos-
sesses a diagonal groove into which the basal costal lip of the fore wing
fits when the wings are at rest. A distinct basal region (ba,) is united
to the episternum dorsally, but a corresponding subalar region is not
demarked from the upper region of the epimeron. The pleuron is united
to the tergum by the prealar bridge (aw.) joining the episternum to the
scutum, and by the postalar bridge (pw.) joining the epimeron to the
postnotum. The mesothoracic spiracle (sp., Fig. 71) is in the membranous.
area just anterior to the prealar bridge.

The sternum (Fig. 74) consists of a central prebasisternite (pbs, )
flanked by the basisternal regions (bst.). (Compare with the sternum of
Aulacizes irrorata.) Even if these areas have a homologous value, it is
difficult to define their exact limits due to the broad fusion of the sternum
with the pleura and the consequent modifications of structure. Detailed
comparative studies of intrafamily modifications and relationships are
required to clarify the interpretation of these sternal structures. Posterior
to the precoxal bridge, a suture demarks a narrow marginal area which
Crampton (1915) has designated as the antecoxale (acx.), similar to
such a modification in Aulacizes irrorata (acx,, Fig. 69). Here again it

THORACIC SCLERITES, WINGS, AND LEGS 29

appears that the median part of this marginal antecoxale is a part of the
basisternal region. The furcasternite (fst. ) bearing the furcal pits (fp.),
lies between the coxal cavities. These pits invaginate to form the well
developed furcal apodemes (fu, Fig. 77), almost reaching to the pleural
apodemes (plap.). The remainder of the mesothoracic endoskeleton is
formed by a platelike phragma (ph., Fig. 76), bearing a deep median
notch. The mesothoracic phragma is formed by the invaginated folds
between the postnotum and the metathoracic scutum. A narrow post-
coxal bridge (pocx., Fig. 77) units the furcasternite with the epimeron,
behind the coxal cavity. The trochantin (tn,, Fig. 78) is a small sclerite
lying in the membranous area between the antecoxale and the coxa.

Metathorax. This segment is considerably smaller than the mesothoracic
segment. The major area of the tergum is occupied by the scutum (sc,,
Fig. 70), divided medially by a suture (ms;). The median suture is
represented internally by a median plate which unites anteriorly with
the mesothoracic phragma. The narrow posterior margin of the tergum
forms the scutellum (scl,), the lateral extensions of which articulate with
the posterior margin of the hind wing by means of the axillary cord
(axc,). Three axillary sclerites (lax,, 2ax;, 3ax,) are present in the basal
membranous area of the wing adjacent to the anterolateral margin of the
scutum.

The pleuron of the metathorax consists of an episternum (es;, Fig. 71)
and epimeron, the latter divided into a large anepimeron (aepm,) and a
smaller katepimeron (kepm,). The posterior portion of the mesothoracic
epimeron overlies the metathoracic episternum and the metathoracic
spiracle (sp;) is concealed in a small dorsal membranous area between
these segments. The pleural wing articulation (pwp;) and the pleural
coxal articulation (cxp.) are found at the respective dorsal and ventral
extremities of the pleural suture (pls,). A prealar bridge (aw,), joined
to the postalar bridge of the mesothorax, unites the epimeron to the
lateral remnant of the postnotum (pnt,). No distinct basalar or subalar
sclerites are present in the articulating membranous area of the wing and
pleuron.

The sternum (Fig. 74) is composed of an anterior basisternite (bs, ),
and a posterior furcasternite (fst,) which lie between the coxal cavities.
The basisternite is joined to the episternum by a precoxal bridge (pcx, ),
while the furcasternite is joined to the katepimeron by a postcoxal bridge
(pocx, ). The furcasternite bears a median pair of furcal pits which form
a small furcal apodeme (fu;, Fig. 73) internally. The trochantin (tns,
Fig. 76), closely approximated with the episternum, articulates with the
coxa. As seen in this posterior view of the metathorax the lateral rem-

30 MORPHOLOGY OF AUCHENORHYNCHA

nants of the postnotum, the postalar bridges, the postcoxal bridges, and
the sternum, are all united to form a ringlike sclerite around the
metathorax.

Wings. Both the fore- (Fig. 87) and hindwings (Fig. 88) are mem-
branous and apparently both pairs of wings are actively used in flight.
Funkhauser (1912) has made an excellent comparative study of the
wings of the Membracidae, and reference should be made to this study
for information concerning the venation. It should, however, be noted
that the forewing possesses a vannal fold (vf) along the course of what
is probably the second branch of cubitus, which separates the large
anterior remigial area from the narrow posterior vannal area, similar to
that which occurs in both the cercopid and cicadellid. Funkhauser refers
to this vannal fold as the claval suture. The venation* of the wings of
Ceresa bubalus is noted in figures 87 and 88.

Legs. The prothoracic (Fig. 111) and mesothoracic legs (Fig. 112) are
very similar. The coxae, trochanters, femora, tibiae, and three-segmented
tarsi of the first two pairs of legs possess no unusual features. The coxae
of the jumping metathoracic legs (Fig. 113) are considerably enlarged
and their mesal margins each possess an angular, toothed projection
which enables the coxae to lock in position against the sternum and each
other. The tibia of the hind leg is further modified by the possession of
four rows of spines along its length and an apical border of spines. The
first tarsal segment is about twice the length of the second tarsal segment,
whereas in the pro- and mesothoracic legs the second tarsal segment
exceeds the first segment in length. The claws (cl, Fig. 116) adhere to the
bilobed, membranous pulvilli (pu) or pads.

*The interpretation of the wing venation is largely based on the studies of
Metcalf (1913a, 1913b, 1917) and Funkhauser (1913). Evans’ interpretation
(1946a, 1948), in which Ri=Ria, Ro+3=Rw, and Rit;=R;, may be the more proba-
ble one if the position of the cross-vein r is accurately traced.

THE THORACIC MUSCULATURE OF
CERESA BUBALUS

The Cervical Muscles. The movement of the head is controlled by four
pairs of muscles (Fig. 50) as follows:

1. A pair of dorsal longitudinal muscles (21) running from the dorsal
occipital margin of the head to the lateral pretergal margin of the meso-
thoracic tergum.

2. A vertical muscle (22) from the dorsolateral occipital margin of
the head to the precoxa of the prothorax.

3. A muscle (23) from the occipital condyle to the dorsal, anterior
margin of the pronotum.

4. A short, thick muscle (24) from the lateral posterior margin of
the hypopharyngeal wing plate to the prothoracic furcal apodeme.

The Prothoracic Muscles. These muscles (Fig. 110) are roughly
divided into two groups, one with insertions on the furcal apodeme, the
other with insertions on the leg and origins on the pronotum. Besides
the muscle going from the furcal apodeme to the hypopharyngeal wing
plate, mentioned above, there are two other muscles connected to the
furcal apodeme.

1. An anterior muscle (31) from the apex of the furcal apodeme run-
ning dorsally to the lower region of the metopidium of the pronotum.

2. A posterior muscle (32) from the apex of the furcal apodeme run-
ning dorsally to the lateral margin of the mesothoracic pretergite.

Five prominent muscles, with origins on the lower region of the metv-
pidium of the pronotum are inserted on the trochantin, meron, coxa, and
trochanter. Posteriorly these are covered or enclosed by a sclerotized
partition (mp) shown in part in figure 72. The muscles (Fig. 110) are as
follows:

1. A large muscle (34) originating just beneath the ridge demarking
the lower region of the metopidium and inserted on the meron.

2. A muscle (35) originating just below the above one on the meto-
pidium and inserted on the mesal tip of the trochantin.

3. A muscle (36) from the upper lateral corner of the lower region
of the metopidium, inserted on the basal rim of the coxa just anterior
to the pleural articulation.

4. Similar to the above, but inserted on the basal rim of the coxa,
just posterior to the pleural articulation (37).

5. This muscle (38) is inserted mesally on the basal rim of the tro-
chanter. Its origin is near the above two muscles in the lateral corner of
the metopidium.

31

82 MORPHOLOGY OF AUCHENORHYNCHA

The mesothoracic leg will be used to demonstrate the leg musculature.
Since the musculature of the leg segments of the pro-, meso-, and meta-
thorax shows a general similarity, further description of the prothoracic
leg muscles, not originating on the pronotum, will be omitted at this
point.

The Mesothoracic Muscles. An examination of the musculature of this
segment at once indicates that of the two wing-bearing segments, the
function of flight is chiefly centered in the mesothoracic segment. Pre-
vious studies of the flight mechanism of insects have indicated that the
flight movements of the wings are performed by two groups of muscles,
the indirect wing muscles and the direct wing muscles. The indirect
wing muscles, by causing alternating changes in the shape of the thoracic
segment, are primarily responsible for the elevation and depression of
the wings. These are as follows:

1. Dorsal longitudinal muscles (41, Fig. 106). A broad pair of median
muscles stretched between the mesothoracic phragma and the arched,
anterior region of the scutum. These are the chief depressor muscles of
the wings.

2. Oblique dorsal muscle (42, Fig. 107). Laterad of the dorsal longi-
tudinal muscles, this muscle extends from the mesothoracic phragma to
the lateral region of the scutum. This muscle acts as a depressor in Ceresa,
but in other insects, where it becomes nearly vertical in position, it may
act as an elevator of the wing.

3. Tergosternal muscle (43, Fig. 107). This muscle, extending from
the anterolateral region of the scutum to the lateral region of the sternum
and the precoxal bridge, is the elevator muscle of the wing. (Figures
106, 107, and 108 have been drawn to reveal the muscles as they are
dissected from the median line to the lateral wall of the thoracic seg-
ments. )

4. This muscle (47a) extends from the dorsal surface of the projecting
pleural apodeme to the anterior region of the posterior notal process.

5. This short, thick muscle (47b) unites the lateral arm of the furca
with the pleural apodeme.

The direct wing muscles act as extensors and flexors of the wing. It is
these muscles which are primarily responsible for extending the wings
into flight position, and in combination with the natural flexibility of
portions of the wing, are responsible for bringing the wings back to rest.
Although the function of some of these muscles in this capacity is quite
apparent or easy to demonstrate, the exact function of others cannot
always be shown, for it is a complex integration of a number of these
muscles which is responsible for the various kinds of wing motion. Some

THORACIC MUSCULATURE OF Ceresa bubalus 33

of the indirect wing muscles are inserted upon the coxa and probably
play a role in the movement of the leg as well.

1. Basalar muscles. Two muscles are inserted upon the upper portion
of the episternum, or basalar sclerite, which invaginates into the body
cavity. The first of these (44, Fig. 109) arise from a lateral process of
the antecoxal sclerite, while the other (45a) arises from a tendon
attached to the basal rim of the coxa just anterior to the pleural articu-
lation. A branch of this latter muscle (45b) goes to the anterior margin
of the episternum.

2. Subalar muscle. As has been previously noted there is no distinct
subalar sclerite. The muscle which arises from the basal rim of the coxa
just posterior to the pleural articulation is the one which usually is in-
serted on the subalar sclerite in other insects where such a sclerite is
present. In this instance the muscle (46, Fig. 112) is inserted upon a
small invaginated process on the dorsal margin of the epimeron.

3. Muscle of the 2nd axillary. This muscle (49) arises broadly from
the anteroventral margin of the episternum and narrows to a tendon
which is inserted upon the 2nd axillary sclerite of the fore wing.

The Ventral Longitudinal Muscles. The ventral longitudinal muscles
(50, Fig. 106) of the mesothorax consist of a pair of small muscles ex-
tending from the anterior margin of the mesothoracic furca to the pos-
terior faces of the paired sternal apophyses of the prothorax. In addition
there is present a pair of short, thick muscles (51) which extend from the
raised posterior region of the furca to the lower posterior wall of the
phragma.

The Muscles of the Leg. Aside from the muscles mentioned above,
which arise on the coxa and are connected with the flight mechanism,
there are present a number of leg muscles originating on the thoracic
sclerites or their apodemes which are apparently connected with the
movements of the leg alone. These muscles are inserted on the coxal and
trochanteral segments of the leg and have their origins either on the
tergum or on the furcal arms of the sternum. Additional leg muscles are
concerned with the movement of the individual leg segments.

CoxaL MuscLEs (Fig. 112).

1. Tergal promoter of the leg (52, Figs. 107 and 112). This muscle
arises on the scutum and is inserted on the ventral end of the trochantin
by means of an elongated tendon.

2. Sternal remoter of the leg (53). This muscle arises on the furcal
arm of the sternum and is inserted on the posterior basal margin of the
coxa.

3. Pleurocoxal muscle (45b). This muscle has already been men-
tioned in connection with the basalar muscle (45a) of which it is a

34 MORPHOLOGY OF AUCHENORHYNCHA

branch. Since it arises on the anterior margin of the episternum and is
inserted on the basal margin of the coxa just anterior to the pleural
articulation, it is in all likelihood directly connected with the movement

of the leg.

4. Tergal remoter of the leg (48). This muscle, arising from the pos-
terior corner of the posterior notal process is inserted by means of a
tendon on the posterior basal margin of the coxa.

The remaining muscle with a coxal insertion is the subalar muscle
(46), inserted on the basal rim of the coxa just behind the pleural
articulation.

TTROCHANTERAL MUSCLES (Fig. 114).

Two groups of muscles are inserted on the basal margin of the tro-
chanter. The mesal articulation of the trochanter with the coxa bears a
long tendon upon which are inserted three muscles. The longest of these
muscles (54a) arises on the scutum; the second (54b) originates on the
furcal arm, while the third (54c) arises on the wall of the coxa. This
group of muscles acts as a depressor of the leg. The second group (55),
or extensors of the leg, consisting of several muscle bundles, all arise on
the outer wall of the coxa and are inserted on the lateral basal rim of the
trochanter.

FEMORAL MUSCLE (56, Fig. 114).

A single small muscle, inserted on the basal rim of the femoral seg-
ment, has its origin on the wall of the distal portion of the trochanter.

TIBIAL MUSCLES (Fig. 115).

1. Extensor muscle. This muscle is inserted on the outer basal margin
of the tibia by means of a tendon. The tendon bears two groups of
muscles, one originating on the wall of the distal portion of the trochanter
(57a), the other originating on the wall of the femur (57b).

2. Depressor muscles (58). A pair of short muscles inserted on oppo-
site sides of the inner basal margin of the tibia, arising on the distal
portion of the femur.

TARSAL MUSCLES (59, 60, Fig. 116).

These consist of two short bundles inserted on opposite sides of the
basal rim of the proximal segment of the tarsus, arising on the wall of
the distal end of the tibia.

PostTARSAL MUSCLE (6la, Fig. 116).

The unguitractor (unp) of the posttarsus continues into the tarsus and
tibia in the form of a long unguitractoral tendon (unt) upon which are
inserted two muscle branches, one arising on the wall of the middle por-
tion of the tibia (6la), the other arising on the wall of the proximal
portion of the tibia (not shown).

THORACIC MUSCULATURE OF Ceresa bubalus 35

The Metathoracic Muscles. The muscles of the metathorax bear very
little resemblance to those of the mesothoracic segment. The most striking
feature is the complete absence of those muscles termed the indirect wing
muscles, the dorsal longitudinals, the oblique dorsals and the tergosternal
muscles. The absence of these muscles precludes the possibility that the
flight mechanism of the hind wings is in any way similar to that of the
fore wings.

The most conspicuous muscles are those associated with the legs.
The leg, wing, and other muscles of the metathorax are listed as they are
seen in a median section of the thorax and dissected away toward the
lateral wall, as follows:

1. Trochanteral extensor of the leg (62a, Fig. 106). This is a large,
conspicuous muscle with a tendonal insertion upon the mesal, basal
margin of the trochanter, and originating partly on the posterior wall of
the mesothoracic phragma and partly on the scutum. A branch of this
muscle (62b) inserted upon the same tendon extends to the posterior
face of the mesothoracic furca. Another branch (not shown) originates
on the wall of the coxa.

2. Adductor of the coxa (63). A small muscle arising on the posterior
of the metasternal furca and extending, posteriorly and ventrally, to the
inner, posterior margin of the base of the coxa.

3. Ventral muscle (64). This is a short muscle extending from the
posterior surface of the metasternal furca to the postcoxal bridge.

4. Tergal remoter of the coxa (65, Fig. 107). A muscle arising on
the anterolateral corner of the scutum, inserted by means of a tendon
on the lateral rim of the base of the coxa just behind the pleural articu-
lation.

5. A broad, flat muscle (66) inserted on the anterior rim of the base
of the coxa; origin on the anterior margin of the episternum.

6. A muscle (67) extending from the trochantin, attached by a tendon
to the anterolateral corner of the scutum.

7. A muscle (68) extending from the precoxal bridge to the antero-
lateral corner of the scutum.

8. A short, narrow muscle (69, Fig. 108) extending from the pleural
apodeme to the anterior notal wing process.

The following muscles have their insertion upon the axillary sclerites
of the wing.

1. A muscle (70) extending from the lateral shelf of the base of the
coxa just behind the pleural articulation to a tendon inserted on the sec-
ond axillary sclerite.

36 MORPHOLOGY OF AUCHENORHYNCHA

2. A muscle (71, Fig. 117) extending from the rim of the base of the
coxa just behind the pleural articulation to the posterior lever of the third
axillary sclerite.

3. A muscle (72) arising on the ventral margin of the anepimeron and
inserted on the anter‘or lever of the third axillary.

ABDOMINAL SEGMENTS OF AUCHENORHYNCHA

The abdomen of auchenorhynchous Homoptera consists of eleven dis-
tinct segments, and the first eight abdominal segments bear spiracular
openings. In the past, the proper enumeration of the abdominal segments
has sometimes caused confusion, due in large part to the modifications
of structure of the two basal segments which form the articulation be-
tween the thorax and the remainder of the abdomen. A reduced sclero-
tization of the terga and sterna of these two segments, and the develop-
ment of extensive intrasegmental areas of membranization have been
largely responsible for the difficulty of delineating the true segmental
limits. The picture has been further complicated by the fact that the
metathoracic postnotum, usually separated from the thorax by a well
defined membranous area in other paurometabolous insects, frequently
forms a complete ring with the metathoracic postcoxal bridges which is
firmly united with the thorax.

The true segmental limits of the abdomen are marked internally by
the intersegmental ridges upon which the dorsal and ventral longitudinal
muscles are attached. Externally these intersegmental ridges may be
marked by lines or sutures, termed the antecostal sutures, which indi-
cate the primary segmentation. The flexibility of the primary segments
has been brought about by a comparatively narrow intrasegmental mem-
branization of the posterior portion of the primary segment, which has
resulted in a functional or secondary segmentation. Normally the pri-
mary segment between two antecostal sutures, or intersegmental ridges,
consists of a wide, anteriorly-sclerotized plate followed by a compara-
tively narrow, posterior membrane, which is in turn followed by a very
narrow rim of sclerite, sometimes absent, just before the succeeding
intersegmental ridge. It will be seen that in the basal segment of the
representatives of the Auchenorhyncha under discussion, the primary
segment sometimes consists of a narrow anterior rim of sclerite, followed
by a wide intrasegmental membrane, and a narrow or wide posterior
sclerite. Othér modifications of the first and second segments will be
discussed in the descriptions of structure which follow. In general, it may
be pointed out that where the enumeration of abdominal segments pre-
sents difficulty, it is possible to get one’s bearings by locating the eighth
abdominal segment which bears the last pair of abdominal spiracles.

Scolops pungens
Visceral Abdominal Segments. The partially distended abdomen of a
female is shown in lateral view in figure 119. The postnotum (pnt,) is a

37

88 MORPHOLOGY OF AUCHENORHYNCHA

broad sclerite stretching across the dorsum, broadly united to the epi-
meral regions (epm,) on each side. The first tergite (It) consists of a
narrow anterior margin of sclerite firmly united with the postnotum, and
a larger, bilobed posterior sclerite. These sclerotized regions are separated
by a narrow membranous aréa. The bilobed areas on either side are
united dorsomesally by a narrew’sclerite, while the anterior of the bilobed
sclerites unites with the anteriof rim of the first tergite, which is in turn
united to the postnotum. The first spiracle (sp,) is located in the mem-
branous tergal region just behind the posterolateral corner of the post-
notum. The second tergum (IIt) consists of a broad, anterior sclerite
separated from a smaller posterior sclerite by a membranous region. Lat-
erally, a small arm of sclerite unites the anterior and posterior tergites of
the second tergum. The second spiracle (sp.) is located in the antero-
lateral membrane of the second tergum. The narrow posterior sclerotized
portion of the second tergum is firmly joined to the broad sclerotized
region of the third tergum. The terga of segments three to eight are
uniform, each consisting of a broad, anterior sclerite, and a posterior
membrane. The spiracles of these segments are located in the mem-
branous area between the lateral margins of the median tergites and the
laterotergites (Itg).

The lateral sclerotized regions below the spiracles are regarded as
laterotergites in view of the fact that in other fulgorids the spiracles are
located in the upper halves of these sclerites. According to the studies
of Snodgrass, the dorsopleural line, which demarks the lower limits of the
dorsum, or tergal region, is always formed below the spiracles. Since the
spiracles do occur in these lateral sclerites in other fulgorids, it would
appear that these sclerites belong to the terga, and are consequently
here termed laterotergites.

Ventrally the first sternal segment (Is, Fig. 118) consists of a narrow
anterior semicircular sclerite which appears to arise from the region
where the posterolateral corner of the postnotum meets the epimeron,
and a narrow, median, posterior sclerite, separated by a membranous
region. The second sternal segment (IIs) likewise consists of narrow
anterior and posterior sclerites separated by a membranous region. Lat-
erally this posterior sclerite of the second sternal segment broadens and
fuses with the laterotergite. Sternal segments three to seven consist of
broad, anterior sternal plates, each followed by a membranous region.
It should be mentioned that these definitive sternal plates are evidently
composite plates, formed by the union of the primary sterna with the
lateral pleural regions. Morphologically then, these definitive sternal
plates are coxosterna or pleurosterna.

The Female Genitalia and Terminal Abdominal Segments. Examina-

ABDOMINAL SEGMENTS OF AUCHENORHYNCHA 39

tion of the structural parts of the ovipositor of Scolops pungens will
indicate that the fulgorids have retained an exceedingly primitive
form of this organ. The eighth tergum (VIIIt, Fig. 131), although re-
duced in size, possesses the normal structure of the preceding terga. The
definitive eighth sternal plate, however, has become modified. It consists
of two lateral halves (lvlf, Fig. 132) separated medially by a membranous
area. These give rise at their posteromesal angles to the arms or first
valvulae (lvl) of the ovipositor. It can be seen that the separate halves
of the definitive eighth sternal plate represent the highly modified first
valvifers of the more specialized ovipositors of other families of Auchen-
orhyncha.

The ninth tergal plate (IXt) gives rise at its lateral margin to a pair
of elongated sclerites which extend ventrally. The anterior of these lateral
branches of the ninth tergum bends posteriorly and unites with the dorsal
edge of the sclerotized portion of the first valvula. As will be seen later
this lateral branch of the ninth tergum represents the origin of the inner
ramus (lri) of the first valvula. The posterior, lateral branch of the ninth
tergum (2vlf) gives rise ventrally to the second valvula (2vl) and to
the broad bilobed flaps dorsally which form the third valvula (8vl). It
would appear that this posterior lateral arm of the ninth tergum repre-
sents the second valvifer of the specialized homopteran ovipositor.

The nature of the terminal abdominal segments in the fulgorids is
distinct from that found in the cicadas, cercopids, cicadellids, and mem-
bracids. Posteriorly, the ninth tergum forms a narrow ring which runs
ventrally beneath the tenth segment. The tenth segment (X) forms a
broad flaplike plate which extends over the dorsal margins of the third
valvulae. Its dorsal surface contains a hollowed area in which the cir-
cular eleventh segment (XI) rests. The eleventh segment bears a single
posterior projection which is termed the anal style (as). It has been
suggested that the anal style may represent the fused cerci of orthopteroid
insects. The anus opens caudally on the eleventh segment between the
base of the anal style and the overhanging posterior margin of the elev-
enth segment.

The Male Genitalia and Terminal Abdominal Segments. The ninth
segment (IX, Fig. 133) of the male consists of a single sclerotized plate
which forms a complete ring around the abdomen. Protruding caudoven-
trally from the membranous area behind the sclerotic ring of the ninth
segment is a pair of large, blunt parameres (pm), each with a hooked
projection on the dorsal margin. Above the parameres is the aedeagus
(aed). It consists of a basal ring of sclerite from which a pair of lateral,
apically bifurcating, arms are given off posteriorly. Above and below
these lateral arms, the aedeagus is membranous. The opening of the

40 MORPHOLOGY OF AUCHENORHYNCHA

aedeagus, or gonopore (gnp), lies in the membranous lobe above. Be-
tween the base of the aedeagus and the base of the parameres, there is
present a median, inverted Y-shaped sclerite, or basal plate (not shown).
The stem of the Y forms an elongate process which extends to the base
of the aedeagus, and each short arm of the Y-shaped basal plate articu-
lates with the base of a paramere. It should be noted that the bases of
the parameres do not possess apodemal processes which extend into the
body cavity as occur in the cercopid, cicadellid, and membracid. Above
the aedeagus is the flaplike tenth segment (X). Both the tenth and
eleventh (XI) segments are generally similar to the corresponding seg-
ments of the female.

An excellent, comprehensive study of the comparative structure of the
male genitalia of Rhynchota, including the Auchenorhyncha, has been
made by Singh-Pruthi (1924).

Tibicina septendecim

Visceral Abdominal Segments. The general structure of the abdomen
of the cicada has been adequately covered by other writers, but is in-
cluded here, for its comparative value, with the other representatives of
the Auchenorhyncha which are discussed. The female abdomen of Tibi-
cina septendecim is shown in lateral view in figure 121. The postnotum
(pnt, ) is a narrow sclerite extending to the epimera on each side. The first
abdominal tergum (It) consists of two sclerotized portions separated by
a membranous area. A narrow, anterior sclerotized margin of the first
tergum is united to the posterior margin of the postnotum, while the
wider posterior sclerite is joined to the anterior edge of the second ter-
gum. These sclerotized portions of the first tergum are joined laterally.
The first abdominal spiracle (sp,) lies in the anterolateral corner of the
first tergum immediately behind the postnotum.

Tergal segments two to eight consist of broad, arched plates of sclerite
anteriorly, followed by posterior segmental membranes. The eighth tergal
segment is wider than the terga of the preceding visceral segments.

The first abdominal sternum (Is, Fig. 120) consists of a median, pen-
tagonal plate fused with the posterior margins of the metathoracic post-
coxal bridges (pocx,;). It is divided into two parts, the posterior of
which folds into the tympanal cavity. The second sternum (IIs) also con-
sists of two parts. An anterior, median rectangular plate, united with
the posterior margin of the first sternal plate, likewise folds into the
tympanal cavity and sends out a pair of narrow arms laterally. The sec-
ond pair of abdominal spiracles (sp. ) is located in the lateral expansions
of the distals ends of these arms. Between these lateral arms of the anterior
half of the second sternum and the anterior margin of the posterior half

ABDOMINAL SEGMENTS OF AUCHENORHYNCHA 4l

lie the tympana (typ) or so-called “mirrors,” of the auditory organ.
The auditory capsule (au) which contains an innervated sense organ, or
chordotonal organ, is a tubercle-like, rounded swelling with a slit-shaped
membrane on the ventrolateral portion of the second abdominal tergite.

The first two abdominal segments of the male are generally similar in
structure to those of the female, except for those modifications of struc-
ture associated with the presence of the sound-producing organs peculiar
to the male. The ventrolateral portions of the first abdominal tergum
are modified to form the large, oval membranous areas, or tymbals
(tyb, Fig. 58), which bear conspicuous, parallel, riblike markings.

It has been previously mentioned that the metathoracic epimera of the
male are produced posteriorly to form the opercula (ope, Fig. 61) which
cover the tympanal chambers. The posterior half of the first abdominal
sternum is longer, and the membranous area between the lateral arms of
the first and second sterna is larger, than the corresponding parts in the
female, so that the tympanal cavity of the male is of greater dimensions
than that of the female. The V-shaped lateral arms of the anterior portion
of the second sternum are wider and stronger than the homologous arms
of the female. It should be noted that Myers (1928), in his comparative
studies of the Cicadidae, has followed the error of Vogel in considering
these arms as “differentiated anterior parts of the first sternite” and the
tympana as “differentiated posterior parts of the first sternite.” The cor-
rect homologies have been pointed out by Snodgrass (1933).

The song of the male is produced by the contractions of the powerful
V-shaped tymbal muscles which extend from either side of the median,
keeled, endoskeletal structure of the anterior portion of the second ster-
num to a pair of internal, sclerotized plates just beneath the tymbal
membranes. A short tendon extends from the center of each sclerotized
plate to the posteromesal sclerotized corner of each tymbal. According
to Myers the contraction and release of the tymbal muscles causes the
tymbal to produce a clicking sound. It is an extremely rapid series of
such clicks, said to be amplified by the large abdominal air sacs which
open directly to the exterior through the first abdominal spiracles, that
produces the “song” of the cicada. As in the female, there is an auditory
capsule present in the ventral portion of the second abdominal tergum.
In the female (Fig. 121), it may be seen that the sterna of segments
three to six consist of broad, anterior definitive sternal plates, each of
which is followed by a narrowed membranous area. Laterally the sternal
plates are united with the tergal plate, each forming a complete ring
around the abdomen. The spiracles of these segments are located in the
anterolateral corners of the sterna, just beneath the lateral margin of the

49, MORPHOLOGY OF AUCHENORHYNCHA

tergal plates. The sternal plate of the seventh segment is deeply and
broadly notched along its posterior margin.

Following a preliminary discussion of the nature and composition of
the abdominal segments of insects, Myers (1928) concludes that each of
the abdominal segments in a cicada consists of “a strongly over-arched
tergite meeting an entirely ventral sternite, which bears the spiracles,”
and apparently considers that the pleural areas are not included in the
definitive sternal plate. As has been noted, the first valvulae of the ful-
gorid ovipositor arise from the first valvifers, or limb base of the eighth
segment, which are homologous with the lateral portions of the definitive
sternal plates of the preceding segments. There is no question here that
the pleural regions are incorporated into the definitive sternal plates of
the fulgorid. Unless it can be shown that the cicadas represent a spe-
cial group among the Auchenorhyncha in which the definitive ele-
ments of the abdominal segments differ from those of the other families,
there would seem little reason to consider that the segmental elements
of the cicadas consist only of tergal and sternal portions.

The Female Genitalia and Terminal Abdominal Segments. The ter-
minal abdominal segments of the cicada are shown in lateral view in fig-
ure 134, with the sclerites somewhat displaced to show their relationship.
The eighth tergal plate (VIIIt) is generally similar to the preceding
tergal plates, except for the fact that it abruptly tapers at its lateroventral
margin. In the membranous region beneath the lateroventral margin of
the eighth tergal plate lies the eighth abdominal spiracle. Below this is
the first valvifer (lvlf), which gives rise at its anteroventral margin to the
first, or ventral valvula (lvl) of the ovipositor. Two rami, or sclerotic
rods, which run along the ventral and dorsal margins of the first valvula
strengthen it, and the dorsal or inner ramus forms a track along which
the second or inner valvula (2vl) of the ovipositor moves. The dorsal or
inner ramus of the first valvula is given off by the small plate (rp) be-
tween the first valvifer and the anteroventral angle of the ninth tergite,
which plate is united with the ninth tergal plate. The first valvula is thus
united with the ninth tergal plate by means of the inner ramus and ramal
plate.

The second valvifer (2vlf) is also joined with the ninth tergal plate.
The union, of the ramal plate and the second valvifer, with the ninth
tergal plate, is a condition apparently homologous with the union of the
corresponding parts of the fulgorid ovipositor with the anterior and pos-
terior lateroventral arms of the ninth tergal plate.

The second or middle valvulae (2vl) are borne by the anterior mar-
gins of the second valvifers, while the posterior margins give rise to the
third or dorsal pair of valvulae (3vl). The second valvulae, united along

ABDOMINAL SEGMENTS OF AUCHENORHYNCHA 43

their dorsal edges and grooved ventrally, fit within the first valvulae, and
together provide a passage for the eggs. The third valvulae are a pair
of flaplike structures into which the terminal portions of the ovipositor
are withdrawn when at rest. Snodgrass has pointed out that the genital
chamber of the female has two openings, an anterior large opening
(gnp, Fig. 134) in the membranous vestibulum behind the seventh ster-
nal plate, and a smaller posterior opening between the bases of the second
valvulae (not shown).

The terminal abdominal segments consist of a short annular tenth
segment (X, Fig. 137) which bears a ring of sclerite, widening ven-
trally, and the smaller eleventh segment (XI).

The Male Genitalia and Terminal Abdominal Segments. The eighth
tergal plate of the male (VIIIt, Fig. 136) has no special modification. The
eighth sternal plate (VIIIs), however, is a shovel-like structure extend-
ing considerably posterior of the eighth tergal plate. Between these plates
lies the eighth spiracle (sps). The ninth segment (IX) is represented by
a circular ring of sclerite sometimes termed the pygophor, or pygofer. The
ninth segment and aedeagus (aed) are shown in caudoventral view in
figure 138. The pygophor is seen to be broadly ringed dorsally and nar-
rowed ventrally, with toothed, caudoventral margins. The aedeagus is an
elongate, sclerotized tube arising in the caudal membranous region of the
pygophor. A basal plate (bp), bearing a pair of divergent arms, is united
with the base of the aedeagus. The genital opening or gonopore (gnp)
is located apically in the expanded membranous region of the aedeagus.
A pair of hooklike sclerotized structures are present in this membranous
area, one on either side of the gonopore.

The aedeagus is kept in place by the paired ventral claspers (vc) of
the tenth segment. The tenth and eleventh segments are shown laterally
in figure 130. The ventral claspers are large, hooked sclerotized appen-
dages, firmly united with the sclerotized ring of the tenth segment. The
eleventh segment (XI) is circled by a narrow band of sclerite and bears
the anal style (as). If the anal style represents the cercus of the eleventh
segment, the small oval sclerotized plate at its base may be the base of
the cercus, or basicercus. It should be noted that in the male genitalia
of the cicada, both the parameres and subgenital plates are absent,
although parameres are present in the primitive cicada Tettigarcta
tomentosa.

Lepyronia quadrangularis
Visceral Abdominal Segments. The sclerotized portion of the first ab-
dominal tergum (It, Fig. 123) of the female cercopid, Lepyronia quad-
rangularis, is mainly confined to the lateral area, in which a small anterior
plate united with the posterior margin of the postnotum is bridged with

44 MORPHOLOGY OF AUCHENORHYNCHA

a large posterior plate. The posterior plates of the first tergum on each
side are united by a narrow rim of sclerite across the mid-dorsal line. The
small, first abdominal spiracular opening (sp,) located in the membran-
ous area immediately behind the metathoracic postnotum is inconspicuous
and sometimes difficult to locate.

The anterolateral margin of the second abdominal tergal plate (IIt) is
united to the first tergal plate, and its posterior border is firmly joined
to the third tergal plate, so that no segmental movement is possible be-
tween the second and third terga. The second abdominal spiracle (sp, )
is located at the anterolateral corner of the second tergal plate. The third
tergal segment (IIIt) consists of a sclerotized, broad, anterior plate and
a posterior membrane. The third spiracle is located in the anterolateral
corner of the definitive sternal plate. In segments four to six there is a
partial membranization of the tergal plate laterally to form laterotergites
(Itg). The seventh and eighth tergal segments are of normal structure.
The fourth to eighth abdominal spiracles are located in membranous
folds between their tergal and sternal plates.

The first abdominal sternum (Is, Fig. 122), consisting of a sclerotized,
anterior, median plate and a posterior, narrow fringe of sclerite, is located
between the postcoxal bridges (pocx,) and posterior intersegmental line
(is). The lateral areas are membranous. The second abdominal sternum
(IIs) consists of a narrow anterior rim of sclerite which borders the
intersegmental line, joined by two lateral arms to a wider, posterior
sclerite. The definitive sternal plate of the third segment (IIIs) has a
median tubercle-like projection on its posterior margin. Sternal segments
four to seven are similar in structure, each consisting of a wide, anterior
plate of sclerite followed by a posterior, segmental membrane.

The Female Genitalia and Terminal Abdominal Segments. The struc-
tural elements of the female genitalia of the cercopid, Lepyronia quad-
rangularis, are similar to those of the cicada. In figure 139, the parts of
the ovipositor are shown in almost their normal resting position. In figure
128, these parts have been considerably displaced to show their relation-
ship. The first valvifers (lvlf) give rise ventrally to the paired, first val-
vulae (lvl). The second valvifers (2vlf), joined to the ninth segment
(IX), give rise anteriorly to the second valvulae (2vl), the basal halves
of which are united dorsally, and posteriorly to the flaplike third valvu-
lae (2vl). The second valvulae move along a sclerotized track on the
inner faces of the first valvulae formed by the inner ramal arms (Iri), of
the first valvulae. The inner ramal arm of each first valvula is joined to
the ramal plate (rp) which is in turn united with the anterior margin of
the sclerotized plate of the ninth tergite (IXt).

The terminal abdominal segments (Fig. 139) of the cercopid are

ABDOMINAL SEGMENTS OF AUCHENORHYNCHA 45

short. The tenth segment (X) is evidenced by a narrow ring of sclerite.
The eleventh segment (XI), in addition to a complete anterior ring of
sclerite bears a saddle-shaped sclerite, and a pair of small plates at the
base of the anal styles. The anus (an) lies between the bases of
the anal styles (as). Ventrally, between the paired anal styles, there is
present a median sclerite. All of these sclerotized portions of the terminal
segments of Lepyronia quadrangularis, unlike the corresponding parts of
some of the other Auchenorhyncha, are well sclerotized and distinct.

The Male Genitalia and Terminal Abdominal Segments. The ninth
segment (IX, Fig. 140) of the male consists of a complete ring of sclerite
to which the ventral subgenital plates (gp) are united. A small posterior
sclerite is separated dorsally from the main ring of the ninth segment.
Protruding caudally from the membrane above the subgenital plates are
the paired parameres, the basal plate and the aedeagus. The relationship
of these parts is shown in figure 129. The parameres (pm) are paired
structures, on either side of the basal plate (bp), each bearing several
hooklike spines or projections. The basal plate is triangular in shape and
bears a membranous area apically from which the base of the aedeagus
originates. The aedeagus bears a pair of bifurcating processes apically,
between the bases of which is the genital opening or gonopore. A slender
median membranous process lies between the two sclerotized bifurcating
processes.

The ringed, sclerotic plate of the tenth segment (X, Fig. 140) is larger
than that of the female, but in other respects the terminal abdominal
segments of the male are similar to those of the female.

Aulacizes irrorata

Visceral Abdominal Segments. The visceral segments of the abdomen
of the cicadellid Aulacizes irrorata, except for the two basal segments,
are subcylindrical and of an almost uniform width and breadth. The
first tergum (It, Fig. 125) consists of an anterior sclerotized region united
with the postnotum (pnt,) and a posterior sclerotized region united
laterally with the anterior plate by an oblique bridge of sclerite. The first
spiracle (sp,) lies in the membranous region of the anterolateral corner
of the first tergal segment just behind the postnotum. The second tergum
(IIt) consists of an anterior sclerotized region and a narrow posterior
rim of sclerite separated by a membranous area. The true limits of the
first and second tergal segments are marked by the intersegmental lines
(is). The second spiracle (sp.) is located in the membranous region im-
mediately below the anterolateral corner of the broad anterior sclerite
of the second tergum. The terga of segments three to eight are generally
uniform in structure, each consisting of a wide, anterior sclerotized plate

45 MORPHOLOGY OF AUCHENORHYNCHA

and a narrow, posterior segmental membrane. Each tergal plate of these
segments is differentiated into a laterotergite (ltg) bearing a spiracle in
its anterolateral corner. The laterotergite of the third segment is some-
what enlarged and articulates with the posterior margin of the meta-
thoracic epimeron.

The basal sternal segments are shown in figure 124. The first sternal
segment (Is) consists of a median sclerotized area, between the meta-
thoracic postcoxal bridges (poxc;) and the anterior bar of sclerite of the
second sternal segment, and lateral membranous areas. The anterior bar
of the second sternal segment (IIs) is followed by a membranous area
and a broad W-shaped sclerite. The anterior and posterior regions of
sclerite of the second sternum are united laterally. The third to sixth
segments consist of anterior, broad, rectangular plates of sclerite fol-
lowed by membranes. The seventh segmental plate of the female abdo-
men, which covers the valvifers and bases of the valvulae of the ovi-
positor, has a convex posterior margin.

The Female Genitalia and Terminal Abdominal Segments. The first
valvifers (1vlf, Fig. 141) bear the flat, first valvulae (lvl) of the oviposi-
tor. On the ventral edge of the first valvula is a thickened rod of sclerite,
the outer ramus (lro). The inner ramal arm (lri) which connects with
the ramal plate (rp) runs along the middle of the inner face of the first
valvula and gradually disappears toward the apex. The ramal plate is
broadly united with the anterolateral region of the tergal plate of the
ninth segment (IXt).

The second valvifers (2vlf) bear the second, or middle valvulae (2vl)
from their anterior regions and the third or dorsal valvulae (3vl) from
their posterior regions. The second valvulae are membranous basally,
flatly compressed, have strong saw-toothed dorsal margins apically, and
are separate. An outer ramus (2ro) runs along the ventral edge of each
second valvula. The dorsal half of the inner face of each first valvula is
closely appressed to the ventral half of the outer surface of each second
valvula, and it may be possible that the first and second pairs of valvulae
act in unison as one pair of appendages. In their resting position they
are sheathed by the third valvulae.

The tenth segment (X) is a short, ringed segment, only partially sclero-
tized ventrally. The eleventh segment (XI) bears an anterior ring of
sclerite, two small sclerites on each side and the apical, paired anal styles
(as) which are united along their median edges by a membrane.

The Male Genitalia and Terminal Abdominal Segments. The ninth
segment (IX, Fig. 142) of the male is completely ringed by sclerite.
Posteroventrally the paired subgenital plates are united with the ninth
annular sclerite. Exserted from the caudoventral membranous region of

ABDOMINAL SEGMENTS OF AUCHENORHYNCHA 47

the ninth segment are a pair of short, bluntly rounded parameres (pm).
Lying medially between the parameres and extending to the base of the
aedeagus (aed) is the single, elongate, basal plate (bp). Proximally this
basal plate bears a pair of diverging arms, and in this species, it can be
seen that the arms of the basal plates articulate with the inner, basal
parts of the parameres. The aedeagus bears a pair of diverging sclero-
tized, sharply pointed arms dorsally. Anterior to these arms and arising
at their base is a median projection, sclerotized anteriorly and laterally,
but membranous posteriorly, which bears the opening, or gonopore
(gnp), of the aedeagus apically. Between the aedeagus and dorsopos-
terior margin of the ninth annular sclerite on each side, and apparently
differentiated from it, is a pair of knoblike sclerotized plates. The tenth
abdominal segment (X) is almost entirely reduced to membrane, while
the parts of the eleventh segment are essentially identical to those of
the female.

Ceresa bubalus

Visceral Abdominal Segments. The abdomen of the female membracid,
Ceresa bubalus, shown in lateral view in figure 127, is highly arched,
somewhat compressed laterally, tapers posteriorly, and is suggestive of
the structure and shape of the abdomen of the cicada, Tibicina septen-
decim. The first tergum consists of an anterior rim of sclerite united with
the metathoracic postnotum (pnt,;), and a posterior sclerite united with
the anterior margin of the tergal plate of the second segment (IIt). These
sclerotized regions of the first abdominal tergum are joined by an oblique
bridge of sclerite laterally. The intervening areas are membranous. The
first abdominal spiracle (sp,) is located laterally in the anterior rim of
sclerite immediately behind the metathoracic postnotum.

The second tergal segment (IIt) consists of a single plate of sclerite,
the anterior half of which rises abruptly upward to meet the posterior
half of the plate. The latter half is bent at approximately right angles to
the anterior portion. The posterior margin of the second tergal plate is
rigidly united to the third tergal plate, there being no intervening seg-
mental membrane. The second abdominal spiracle (sp.) lies in the
membrane below the anterolateral region of the second tergal plate.

The third to seventh abdominal terga, except for a successive reduc-
tion in width, are uniform, each consisting of a highly arched anterior
plate of sclerite followed by the posterior segmental membrane. The
eighth tergum is wider than the preceding segment, and tapers antero-
ventrally. A ventrolateral region of each of tergal plates three to eight
bends underneath the abdomen, in a plane with the sternal plates, to
form the laterotergites (ltg). The spiracle of the third segment is located
in the anterodorsal corner of the laterotergite, while the spiracles of the

48 MORPHOLOGY OF AUCHENORHYNCHA

succeeding four segments are each located in the anteroventral corner of
their corresponding laterotergites. The eighth abdominal spiracle is
located dorsocentrally in the eighth laterotergite.

The first sternal segment (Is, Fig. 126) consists of lateral membranous
areas and a median, trapezoid-shaped plate of sclerite. The smaller an-
terior margin, and the wider posterior margin of this median plate re-
spectively are united with the metathorax and the second sternal plate.
The second sternal segment (IIs) consists of anterior and posterior plates
of sclerite separated by a middle region of membrane. The posterior plate
of the second sternum is united with the sternal plate of the third seg-
ment (IIIs). Each of segments three to six consists of an anterior rectan-
gular plate of sclerite separated by membranous regions from its own
laterotergites and the succeeding segment. The seventh definitive sternal
plate is longer than those of the preceding segments and its posterior
margin is notched medially.

The Female Genitalia and Terminal Abdominal Segments. A lateral
view of the genital segments and the parts of the ovipositor is shown in
figure 149. The seventh sternal plate partly covers the paired first valvi-
fers (lvlf). The first valvifer gives rise to an outer, ventral blade, or first
valvula (lvl) of the ovipositor. The anterior regions of the second valvi-
fers (2vlf) bear the inner middle blades, or second valvulae (2vl) of
the ovipositor. The second valvulae are united dorsally, forming a ven-
tral groove for the passage of the eggs, and have a dorsal, saw-toothed
edge apically. The flaplike dorsal, or third valvulae (3vl), which sheath
the blades of the ovipositor, arise from the posterior region of the second
valvifers. The relationship of the second and third valvulae to the second
valvifer is shown in figure 147.

Each ventral blade of the ovipositor possesses both an outer and inner
ramus, while the middle blade possesses only an outer ramus. The rela-
tionship of these rami, the other parts of the ovipositor and the ninth
tergite are best seen in internal view shown in figure 148. The outer
rami (lro) run along the ventral margin of the first valvulae and form a
track by means of which the first valvulae slide upon one another. The
inner ramus (lri) which runs along the dorsal edge of the first valvula
arises from the ramal plate (rp) which is united with the anterolateral
corner of the ninth tergite (IXt). The outer ramus (2ro) of the second
valvula runs along the ventral edge of the latter, and together with the
inner ramus of the first valvula forms a track, one on each side, by means
of which the inner valvulae slide on the outer valvulae. The second valvi-
fer is connected by a short thick bar of sclerite to the inner margin of
the anterolateral corner of the ninth tergite.

ABDOMINAL SEGMENTS OF AUCHENORHYNCHA 49

In the ventral view of the genitalia shown in figure 144 the seventh
sternal plate has been pulled back to expose the valvifers and the bases
of the valvulae. The genital opening, or gonopore (gnp) by means of
which the eggs pass from the median oviduct to the ovipositor, is shown
between, and at the base of, the second valvulae.

The terminal abdominal segments are shown in figure 145. These seg-
ments in the female present a rather unusual arrangement. The posterior
margins of the ninth tergal plate are bounded by an annular membranous
region which invaginates within the ninth segment and gives rise to a
sclerotized cylindrical tube which is apparently a part of the tenth ab-
dominal segment (X). Within this outer portion of the tenth segment is
another tube, somewhat less strongly chitinized, but its anterior margin
is rigidly united with the anterior margin of the outer, tubelike portion.
This inner, tubelike segment is apparently a posterior portion of the tenth
segment. The posterior portion of the tenth segment is followed by a
membranous section, followed in turn by another annular sclerite which
is a part of the eleventh segment (XI). The remainder of the eleventh
segment consists of the anal styles (as) and small paired, basal sclerites.

The Male Genitalia and Terminal Abdominal Segments. The male
genitalia are shown in lateral view in figure 151. The ninth tergum (IXt)
consists of a saddle-shaped tergal plate, broad dorsally and narrowing
laterally. On each side the caudoventral region of the ninth tergal plate
is differentiated into a flaplike plate bearing an immovable spine (ts).
Ventrally, the ninth segment forms a single subgenital plate (gp), which
is notched posteriorly. It is shown in ventral view in figure 154. The sub-
genital plates of other species of membracids, belonging to the genera
Aconophora, Platicotis, Campylenchia, Enchenopa, and others, consist
of either entirely separate paired genital valves and a separate sternal
plate, or show these parts in varying degrees of fusion. It therefore ap-
pears that the subgenital plate of Ceresa bubalus consists of the fused
genital valves and ninth sternal plate.

Above the subgenital plate are the elongate, paired parameres (pm).
Between the bases of the parameres and extending to the base of the
aedeagus is the median basal plate (bp). The aedeagus (aed) is a
median, hollow, rodlike structure with an elongate, vertical membranous
region caudally which bears the slit-shaped gonopore (gnp). The aedea-
gus is shown in figure 155, in caudal view. There is a median sclerotized
plate, apparently a part of the basal region of the aedeagus, extending
to the anteroventral margin of the tenth segment.

50 MORPHOLOGY OF AUCHENORHYNCHA

The terminal abdominal segments of the male differ from those of the
female. The tenth segment (X, Fig. 151) bears none of the modifications
of the female, but consists of an annular sclerite followed by a membran-
ous region. The eleventh segment (XI) also has an anterior annular
sclerite and paired anal styles (as). The relationship of the terminal ab
dominal segments and genitalia is shown in caudal view in figure 150.

THE ABDOMINAL MUSCULATURE OF
CERESA BUBALUS

Muscles of the Visceral Abdominal Segments. The muscles of the ab-
dominal segments of Ceresa bubalus are shown in figure 143 and may be
classed in three general groups. These are the dorsal longitudinal muscles,
the ventral longitudinal muscles, and the lateral muscles. The dorsal lon-
gitudinal muscles are in some segments subdivided as median dorsal
longitudinal muscles and lateral dorsal longitudinal muscles. These are as
follows:

DoRSAL LONGITUDINAL MUSCLES (d).

These muscles extend between the intersegmental ridges, or ante-
costae, of successive terga, the intersegmental ridges being present on the
anterior margins of the tergal plates. In the middle, visceral abdominal
segments these muscles are separated into two groups, the median dorsal
longitudinal muscles (dm) found on either side of the mid-dorsal line
of the terga, and the lateral dorsal longitudinal muscles (dl) extending
across the lateroventral portions of ventral longitudinal muscles (v).

VENTRAL LONGITUDINAL MUSCLES (Vv).

These muscles extend from the intersegmental ridges, or antecostae,
of the successive sterna, the intersegmental ridges being present on the
anterior margins of the definitive sternal plates. The muscles are located
in the lateral regions of the sternal plates.

LATERAL MUSCLES (1).

These muscles extend from the lateral margins of the definitive sternal
plates to the tergal plates. The lateral muscles extending from the lateral
margins of the definitive sternal plates of the typical abdominal seg-
ments consists of two groups, one a short, external, lateral muscle (le),
extending to the laterotergite, the other a longer internal muscle (li)
extending to the lower region of the tergal plate.

Within the first segment there are two bands of median dorsal longi-
tudinal muscles extending from the intersegmental ridge on the posterior
margin of the metathoracic postnotum to the succeeding intersegmental
line. A ventral longitudinal muscle (v) extends from the metathoracic
furcasternum to the anterior of the intersegmental line between the first
and second abdominal sterna. A lateral muscle extends from anterior of
the intersegmental ridge dorsally to the posterior margin of sclerite of the
first abdominal tergum. Behind this muscle is another smaller lateral

51

OF ILL. LB.

52 MORPHOLOGY OF AUCHENORHYNCHA

muscle (not shown) extending from the lateral intersegmental ridge of
the sternum to the intersegmental ridge of the second tergal plate.

The dorsal longitudinal bands of muscle of the second segment are
grouped together and not differentiated as median and lateral dorsal
longitudinal muscles. A lateral muscle extends from the anterolateral
margin of the second sternal segment to a ridge on the ventral region of
the third tergal plate. The dorsal, ventral, and lateral muscles of the
remaining visceral abdominal segments are typically as described in the
outline above, except that the dorsal longitudinal bands of muscle of
the seventh segment are grouped together, and in the eighth segment
they are hardly differentiated as separate bands of muscle. The ventral
longitudinal muscles of the seventh segment are absent. There are no
muscles associated with the spiracles.

Musculature of the Female Genitalia and Terminal Abdominal Seg-
ments. Two groups of muscles are associated with the first valvifer of the
eighth segment. The first (81), probably a modification of the lateral
muscles of the visceral segments, originates on the eighth tergal plate
and is inserted on a lateral or dorsal apodeme of the first valvifer. The
second group (82) consists of short, thick muscles extending from the
posterior wall of the first valvifer to the base of the outer ramus of the
second valvula and to the anterior wall of the second valvifer. These
muscles may be modifications of the ventral longitudinal muscles of the
visceral abdominal segments.

Two groups of muscle are associated with the second valvifer, both
taking their origin on the ninth tergal plate. The first, a single large
muscle (91), is inserted on an anterior apodeme of the lateral, dorsal
edge of the second valvifer. The second, consisting of several muscles
(92), is inserted on the posterior margin of the second valvifer and the
base of the third valvula. In addition there is a short lateral muscle (93)
extending from the wall of the apical region of the ninth tergal plate to
the caudoventral flap of the ninth tergal plate.

Three small muscles are associated with the tenth segment. These
muscles are inserted on the circular margin where the outer, anterior
tube and the inner, posterior tube of the tenth segment meet. The first of
these (101), inserted mediodorsally on the inner rim of the sclerotized
tube of the tenth segment, originates on the dorsal wall of the ninth
tergal plate. The second and third muscles (102) are inserted ventro-
laterally, and originate on the ventral sides of the ninth tergal plate.
These cylindrical sclerites of the tenth segment are apparently incapable
of being exserted, telescope fashion, but are limited in their protrusion
posteriorly by the membrane between the sclerotized portions of the

ABDOMINAL MUSCULATURE OF Ceresa bubalus 53

ninth and tenth segments. The eleventh segment is probably exserted by
the turgidity of the body cavity and the rectum.

Musculature of the Male Genitalia and Terminal Abdominal Seg-
ments. The muscles of the male genitalia are surprisingly complex and it
would therefore seem likely that the copulation of these insects is no mere
approximation of the genitalia of the male with those of the female. It
would seem, rather, that considerable maneuvering and adroit procedure
on the part of the male is required for the achievement of successful
mating.

The internal apodemal process of each paramere bears five distinct
muscles. These muscles, all of them paired except the last, are shown in
figures 152 and 153 and are as follows:

1. A large muscle (94) extending from the dorsal margin of the an-
terior region of the apodeme to the lateroventral region of the ninth
tergal plate.

2. A second muscle (95) from the dorsal margin of the anterior of
the apodeme, lying mesad of the one above, extending to the posterior, or
dorsal margin of the basal plate.

3. A muscle (96) extending from the ventromesal margin of the
anterior of the apodeme to the anterior margin of the subgenital plate.

4. A muscle (97) extending from the underside of the posterior region
of the apodeme to the inner face of the base of the subgenital plate.

5. A single transverse muscle (98, Fig. 153) uniting the anterior re-
gions of the apodemes.

The caudal membranous region of the ninth segment which lies be-
tween the base of the aedeagus and the base of the subgenital plate en-
circles the base of the parameres on each side. This circle of membrane
which is formed around the base of the paramere gives rise internally to
a tendon upon which is inserted a fan-shaped muscle (99, Figs. 152 and
153) which originates on the lateral wall of the ninth tergal plate. In
addition, a pair of muscles (100), on either side of the median sclerotized
process between the aedeagus and the base of the tenth segment, extend
to the caudoventral flaps of the ninth tergal plate.

Two pairs of muscles (104, 105, Fig. 153), inserted laterally on the
basal margin of the annular sclerite of the tenth segment, have their
origins in the dorsal wall of the ninth tergal plate.

THE PHYLOGENY OF AUCHENORHYNCHA

Phylogenetic Considerations of Previous Authors

Concepts of phylogeny, although almost entirely absent in the minds
of early nineteenth century insect taxonomists, were nevertheless re-
flected to some small extent in their systems of classification. Thus, the
general relationship of those groups of insects belonging to the Homop-
tera, and those belonging to the Heteroptera was embodied in West-
wood's classification of 1840. Similarly, Dumeril (1806) had indicated
the relationship within the Homoptera that existed among the Aucheno-
rhyncha and among those groups later known as the Sternorhyncha. It
must be remembered, however, that during the nineteenth century, insect
classifications were, like the classifications of other animals, based _pri-
marily on one or two key characters, such as the wings and mouthparts.
Some success was nevertheless achieved in delineating the ordinal and
subordinal relations of insects on this basis, but in attempting to extend
the use of one or two key charcters into familial and lesser categories the
consequent failure to represent phylogenetic relationship was more evi-
dent.

Osborn (1895) took issue with the prevailing system of his day of
placing the Coccidae as the lowest and presumably the simplest group
of the Homoptera. Separating the Homoptera into two subdivisions,
Sternorhynchi and Auchenorhynchi, he claimed that there is every reason
to consider the Sternorhynchi as the derivative form and the Aucheno-
rhynchi as the basal form of the phylogenetic tree. He wrote that “The
position of the rostrum upon the sternum, or, more properly, the coal-
escence of the rostrum with the sternum, must certainly be considered as
a more specialized condition than the free form and, in fact, the deriva-
tive form, the consolidation of the rostrum with sternum being the result
of the close approximation of beak and sternum resulting from their food
habits.”

Within the Auchenorhyncha, Osborn considered the Cicadidae to have
the most generalized condition of wing venation and body structure,
while the Membracidae, “except in the extremely specialized pronotum
are easily seen to be related to the Cicadidae, and naturally take their
position next to them. The Fulgoridae, while possessing specializations
of the head, are in thoracic structure and venation more generalized than
the remaining families, and while possessing many highly differentiated
subgroups, may very probably be interposed between the preceding fam-
ilies and the jassoid division. The Cercopidae in development of the

54

PHYLOGENY OF AUCHENORHYNCHA 55
scutellum and in texture of elytra, as well as in specialization of the tibiae,
show characters of rather high rank, and, if placed as subordinate to the
Jassoidea, they must at least be considered as a branch of nearly equal

or parallel rank.”

Since Osborn utilized such terms as “specializations of the head,” “gen-
eralized,” “characters of rather high rank,” “subordinate,” “nearly equal
or parallel rank,” etc. without specific meaning, it is not surprising that his
concept of relationships within the Auchenorhyncha, as indicated in

Chart 1, was entirely erroneous.

3yvait1Asd

COCCIDaAE we \
ee NO
~ \ I

Chart 1. Phylogenetic tree of Homoptera, after Osborn (1895).

Kirkaldy (1910b) pointed out that the interposition of the Fulgoridae
between the Cercopidae and the Membracidae is a misconception dating
from the earlier studies of Fieber, and was not in the least warranted by
the structure or habits of the groups in question. He correctly pointed
out that the Fulgoroidea were much more distinct from the rest of the
Auchenorhyncha than these latter groups were from each other. In an
earlier paper Kirkaldy (1906) concluded that the Fulgoroidea are the
“most specialized, highly organized and differentiated” of the Aucheno-
rhyncha, that the Cicadoidea as a whole are of “low estate,” and a “slight
degree of specialization” is shown by the Tetigonioidea (Jassidae,
Membracidae and Cercopidae ) in almost every point, and that the Mem-
bracidae are only Tetigonioidea with a highly specialized pronotum. His
concept of homopteran relationships is expressed in Chart 2.

As to the paleontological evidence, Handlirsch (1908) pointed out
that we can follow the Hemipteroids or Rhynchota up to the Paleozoic;
that in the Tertiary all essentially modern families were present and even
iu the Jurassic the principal groups already appear to be marked out, 30
that we can assign nearly all Mesozoic forms to the orders Homoptera
and Hemiptera (Heteroptera). He writes, “As regards the phylogeny of
the groups comprised under the Homoptera, it can in any case be main-
tained, that the present day forms of fulgorids have retained the most
primitive characters. They still have, for example, a simply convoluted

56 MORPHOLOGY OF AUCHENORHYNCHA

intestine without complicated loops (filter chamber), as occur in the
jassids and cercopids, etc. Typical fulgorids are already known to us
from the Liassic (Lower Jurassic), indeed in that period are also found
forms resembling the jassids* and a few species which are inclined
toward the cercopids and are consequently designated as Procercopidae.
Cicadidae first began to be found in the Cretaceous.

an
et)
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= uw <£ 9g a
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= = ~ —
fe || 2 bs}
= aoe
. <2 ~
- = SS
FULGOROIDEA
CICADOIDEA
APHOIDEA

PRO1O-HOMOPTERA
Chart 2. Phylogenetic tree of Homoptera, after Kirkaldy (1910).

“It will not be difficult, to derive the above named families, and the
jassids and cercopids as well, from the earlier fulgorids, by the interposi-
tion of forms which acquired the characteristic intestinal loops; per-
haps these forms were the Proceropidae. The Cicadidae may be de-
rived either from very early cercopids, or also from these Procercopidae,
since they too possess the intestinal loops and the characteristic antennae
of the families just mentioned. Consequently, I consider the suborder
Auchenorhyncha to be a monophyletic, natural one.”

The complete line of development including the Sternorhyncha, is,
according to Handlirsch, that “From the oldest fulgorids or from the
Protohemiptera there developed as early as the Triassic, more specialized

*Evans (1948) has indicated that “the available evidence would seem to suggest
that either true jassoids, or else their immediate ancestors, were in existence during
the Permian.”

PHYLOGENY OF AUCHENORHYNCHA 5g

types (perhaps the Procercopidae), from which presumably the jassids
and later the cercopids arose, as did the Psylloidea, and from these latter
presumably arose the Coccoidea during the Cretaceous. From cercopid-
like forms, may have arisen in the Malm (Middle Jurassic), or soon after,
the cicadas, while directly from the fulgorids in the course of the Jurassic
arose the Aphidoidea, and presumably the Aleurodoidea in the Cre-
taceous.”

In contrast to the Auchenorhyncha, Handlirsch maintained that the
Sternorhyncha do not represent a monophyletic group, but a polyphyletic
one, and proposed that each of the four families in this group be desig-
nated as suborders (though these have superfamily endings), Psylloidea,
Aleurodoidea, Aphidoidea, and Coccoidea, of equal rank with the Auch-
enorhyncha. His interpretation is expressed in the phylogenetic tree
shown in Chart 3.

Aucheno-
rhyncha
a

® 3
o vo 3 $3 8

ofszvegs
be] tao Tc
wUBUe Aa t Os
==zEzvtrorworyD
YJRs v4 oF
cae ng 5 2 5
vevvunwui cd

Present

Cenozoic

Cretaceous

U. Jurassic

M. Jurassic

L. Jurassic

Triassic

Chart 3. Phylogeny and geological distribution of the Homoptera, after
Handlirsch (1908).

Kirkaldy (1910a) took issue with Handlirsch’s views on the phylogeny
of the Homoptera, particularly as to the derivation of the Aleyrodidae
from the Fulgoridae, the placing of both in the same main branch as the

58 MORPHOLOGY OF AUCHENORHYNCHA

the Aphidae, the separation of all the aforementioned groups from the
Psyllidae and the Coccidae, and as to the derivation of the Cicadidae
from the Cercopidae, etc. Kirkaldy further disagreed as to the poly-
phyletic origin of the Sternorhyncha, maintaining that “if there is any
group which is apparently compact, it is the Sternorhynchous Homop-
tera.”

Taylor (1918) on the basis of a comparative study of the thoracic
sclerites of representative Heteroptera-Homoptera, indicated the follow-
ing relationships of Homoptera, shown by the thorax, without regard to
primitiveness:

Gicadidae.- a5. 2 fan | :

jessie ; Neuropteroid thorax... ./ Similar thoracic

NDDMIGHIGAC ot eas Beene ce Seats thn apane eet eee Gees Pee but few } similar
Syl dae fore. cay Sts cee Ra one onsets SOC UU CRNIOLNS . general
Membracidae........ Specialized) prothorax. 7.9.54. eet oe thoracic
Cercopidaeva-ae =. lan
Fulg ae Hed Peculiar fusion of metathoracic
Nise Bilas, 6505. sclerites’ and!coxaex-:s...Js 5 ss. ce ee ee

Tillyard (1919), in a paper describing fossil Homoptera, consisting of
tegmina from the Upper Triassic of Queensland, concluded that “It
would appear proved that the Homoptera became differentiated from a
single Palaeo-hemipterous stock, of which Prosbole is a representative,
in the Middle or Upper Permian. From the same stock the Heteroptera
became separated off at a somewhat later period, the oldest true Heterop-
tera known being the Dustaniidae from the Upper Trias of Ipswich.
Thus the Homoptera are older than the Heteroptera, as is evident on
morphological as well as palaeontological grounds. The separation of the
Sternorrhyncha from the Auchenorrhyncha must have taken place before
the Upper Trias. The oldest existing family of the Auchenorrhyncha ap-
pears to be the Jassidae; the oldest of the Sternorrhyncha are the Psylli-
dae. Venationally, but not in all other characters, the Jassidae are the
older of these two. It may also be pointed out that, after the jassids
became differentiated out, the old main stem of the Auchenorrhyncha
went on, and continued to be represented by many forms which, in cer-
tain directions, still preserved archaic characters which the jassids had
lost (e.g., the existence of a well developed subcostal vein). Thus there
is no difficulty in understanding how the Fulgoroidae and the Cercopidae
could have arisen later in point of time than the Jassidae, though pre-
serving certain archaic features which the jassids had lost. This is the
same problem as that which confronts us in studying the Panorpoid
orders, in which it is clear, palaeontologically, that the Mecoptera was
the first recent order to be differentiated out, though it is, in some

PHYLOGENY OF AUCHENORHYNCHA 59

respects, more highly specialized than other orders which arose from
the main panorpoid stem at later dates, such as the Megaloptera.”

Tillyard’s distinction, between one group of insects that may arise later
in point of time, but which preserve certain archaic features which an
earlier differentiated group has lost, is a most valuable one—even though
I consider his evaluation to be incorrect as applied to the fulgorids, cer-
copids, and jassids. His ideas on phylogeny are expressed in the diagram
in Chart 4.

er ee

JASSIDAE
MEMBRACIDAE

CICADIDAE
CERCOPIDAE

FULGOROIDEA

PSYLLIDAE
APHIDIDAE

COCCIDAE

ALEYRODIDAE

Chart 4. Phylogeny and geological distribution of the Homoptera, after Tillyard
(1919). Known fossil types are as follows: 1, Prosbole; 2, Permofulgor; 3, Scytinop-
tera and Permoscarta; 4, Dunstaniidae; 5, Eurymelidium, Mesojassus and Triasso-
jassus; 6, Mesoscytina, Mesodiphthera and Triassoscarta; 7, Ipsvicia; 8, Mesocixiinae;
9, Triassopsylla; 10, Archijassus; 11, Procercopis; and 12, Fulgoridium.

Singh-Pruthi (1925b), on the basis of an extensive comparative study
of the male genitalia of Rhynchota, considered that the “Cicadidae re-
semble Heteroptera in the form of the VIIIth segment, and in the non-
differentiation of the sub-genital plates; the Fulgoroidea also have no
sub-genital plates, and moreover, show different grades of aedeagus dif-
ferentiation parallel to those in Heteroptera. (Still more, most Fulgoroi-
dea, unlike the rest of Homoptera, resemble Heteroptera in not having
2 complete ovipositor.) But the structure and the position of the basal
plates, surrounding the basal foramen or lying on the segmental mem-
brane inside the body cavity, separate the two sub-orders quite dis-

60. MORPHOLOGY OF AUCHENORHYNCHA

tinctly without any exception. It seems that the heteropterous condition
is more primitive, but this cannot be asserted with any certainty unless
information about the condition of the basal plates in other orders of
insects is available.

“Thus the two sub-orders seem to have a common origin, and are not
distinct orders as suggested by some; but they are separated by a good
gap, and we must look to palaeontology for some Protohemiptera in
which the basal plates are in the intermediate condition.”

Jassidae

Membracidae

Cicadidae Sercopidae

Psyllidae

Fulgoroideae

Coccidae

Homoptera

Chart 5. Phylogeny of the Homoptera, after Singh-Pruthi (1925), based on a study
of the male genitalia.

Carpenter (1931) described a group of closely related, Lower Permian
insects of Kansas, belonging to the fossil family Archescytinidae. He
pointed out that the more primitive wing venation of the Sternorhyncha
las a reduced anal area and the second anal vein is absent. On the other
hand the antennae of Auchenorhyncha, consisting of two short segments
plus a bristle or style, are specialized, whereas the antennae of the Ster-
norhyncha consist of at least three segments and may have as many as
twenty-five. As far as the wing venation of the Archescytinidae is con-
cerned, they resemble the Auchenorhyncha, but Carpenter found one
species of fossil Archescytina which has long antennae consisting of at
least twenty-five segments. These insects also have a well developed
clavus and three-segmented tarsi, characteristic of Auchenorhyncha. Con-
sequently Carpenter considered that there was hardly justification for
placing these Permian insects with the Sternorhyncha, solely on the
basis of their antennae. He therefore proposed the erection of a third
division, the Paleorhyncha, of equal rank with the Sternorhyncha and

PHYLOGENY OF AUCHENORHYNCHA 61

Auchenorhyncha, “in order to avoid abolishing an outstanding char-
acteristic of the Auchenorhyncha by expanding the definition of that
division to include insects with multisegmented antennae.”

Evans (1943) accepted this new division and described a new genus
and species, Austroscytina imperfecta, belonging to the Archescyntinidae
within this division, from the Upper Permian of New South Wales. Later
(1948) he pointed out that though it is conceivable that the Archescyn-
tinidae lie close to the line of descent of the Jassoidea, they cannot pos-
sibly have been ancestral to the whole superfamily, since the media of
the tegmen of the Archescyntinidae is invariably three-branched, and
the common ancestor to all the jassoid families must have had a media
with four branches. According to him, the Archescyntinidae may possibly
have been a jassoid family which had already become specialized in cer-
tain directions by Lower Permian times.

o

o e ® §

Hy o 5 5 cr: eo 3S B
3s 3 eS CTE SE he Cy
= 3 < zs = £ £ £8 8g a
sae ee ee one a
=o 2S Pe yg 2g oe ST oe oy
OPO 4 es peek Ue eee Beigua Se
0 ey Fo; heehee ihe hie, ewe cke> Poste
oa »£ V <€ Od uU efF FE Ga ae

Cercopidae

Protoheteroptera Protohomoptera

Protohemiptera

Chart 6. Phylogeny of the Homoptera, after Spooner (1938), based on a study of
the head capsule.

Spooner (1938), on the basis of extensive comparative studies of the
head capsule of Heteroptera-Homoptera, also recognized the early, dis-
tinctly separate origin of the Fulgoridae from that of other families of
Auchenorhyncha, and derived all the remaining families of both Auchen-
orhyncha and Sternorhyncha from the Cercopidae. His conclusions re-
garding the relationships of the families, arrived at by a study of the
head capsule, are shown in Chart 6. _

62 MORPHOLOGY OF AUCHENORHYNCHA

Evans (1940) agreed with Spooner in the separate derivation of the
Fulgoridae and Peloridiidae from the Protohomopterous stem, a sugges-
tion which he himself had previously made (1938), but disputed the
derivation of the Sternorhyncha from the Cercopidae and the dual origin
claimed for the Jassoidea (Spooner’s Cicadellidae and Tettigonidae). He
indicated that there was no necessity of attempting to derive either the
divisions Auchenorhyncha or Sternorhyncha, one from the other. Further,
that while it was reasonable to suggest a dual origin of the Jassoidea on
the basis of head structure alone, other characters, such as the wing vena-
tion of the hind wings, would not support such a dual derivation. Evans’
views (1946) on the classification of the Auchenorhyncha are presented
in an earlier section of this paper (p. 4).

The above remarks, concerning the phylogeny of the Auchenorhyncha,
indicate that there still exists, among students of the Homoptera, con-
siderable contention regarding the relationships within this division. The
palaeontological evidence, while it gives some assistance in determining
the age of certain fossil Homoptera and the extent to which certain
characters were present in these forms, is comparatively meager and does
not, at the present time, substantially improve our knowledge of homop-
teran phylogeny. Although every source of evidence from this line is to
be valued, it is doubtful that our ultimate knowledge of relationship will
depend upon this source. Insects are fragile, and to date most of our
knowledge of fossil Auchenorhyncha and related forms depends to a
large extent, on the fossil wings of Homoptera, or parts of such wings. At
best, such evidence is one-sided and not conclusive. Tillyard (1919) in
describing fossil Homoptera, based on the tegmina, from the Mesozoic
of Queensland, noted “the initial difficulty, that the classification of the
Homoptera is based for the most part upon characters other than wing
venation” and that although “recent studies of the nymphal tracheation
in the various families have much improved our knowledge of the
venation,—they have not yielded a really satisfactory basis of classifica-
tion on venational characters alone.”*

Discussion

The table (p. 64) summarizes the principal similarities and differences
found in the representatives of the Auchenorhyncha studied herein. It
should be noted first that the species selected to represent the family
groups were chosen for availability and not for their primitive position
within the family. A conclusive picture of phylogeny cannot be drawn

*Evans stated his belief (in conversation, 1949), based on his own extensive
studies, that for the larger categories (families) the venational characters alone do
provide a satisfactory basis of classification.

PHYLOGENY OF AUCHENORHYNCHA 63

until one has compared the primitive representatives within the families,
but to know what is primitive some such investigation as I have
made here must be undertaken. Secondly, a series of representatives
within the families should ultimately be compared, but such an in-
vestigation is beyond the scope of this study. Many of the characters
have been compared, however, with other representatives of the families,
and with published descriptions. The subsequent remarks will therefore
have some worth, even though limited in extent.

The most striking feature of Scolops pungens is the primitive nature
of all aspects of the sucking pump mechanism in comparison to those of
the other Auchenorhyncha. While acknowledging that the sucking pump
and stylets of the Auchenorhyncha are in themselves highly specialized
mouthparts, those of Scolops, and all other fulgorids as far as I know,
have remained in a comparatively primitive state of development. To be
noted first in this connection are the small proportions of the preoral cly-
peal area to the remainder of the head capsule (excluding the cephalic
head process) in Scolops, compared to the much greater proportions of
this area in the cicada, the cercopid, the cicadellid, and the membracid.
Figure 8a, representing the lateral view of the head capsule of an un-
described species of Scudderia (Orthoptera), has been drawn for com-
parison with the essentially orthopteroid proportions of the clypeal area
to the remaining head capsule in Scolops pungens (Compare with Figs.
18, 30, 40, and 49).

In connection with the primitive nature of the sucking pump mech-
anism in Scolops pungens, it should be noted that the mandibular and
maxillary stylets both bear single protractor and retractor muscles. In
the other forms the musculature is always more complex, there being
two or three retractor muscles, and frequently additional protractor and
retractor muscles associated with the lever of the maxillary stylet as well
as with the stylet itself. There is no evidence, as far as I have been able
to ascertain, that the fulgorids have ever developed a sucking pump
mechanism comparable to that found in other Auchenorhyncha. It seems
sound to assume that the other Auchenorhyncha must have at one time
passed through a stage in which the sucking pump mechanism was in a
similar, relatively undeveloped state.

The ovipositor of Scolops pungens also represents an exceedingly prim-
itive stage of development. Particularly to be noticed is the vertical posi-
tion and lack of differention of the first and second valvifers and the
ramal plate (Fig. 131). In contrast, the valvifers and ramal plate of
Tibicina, Lepyronia, Aulacizes, and Ceresa are considerably modified,
their position having shifted in a strongly marked, diagonal, anteroventral
direction, and the first and second valvulae are highly specialized. The

MORPHOLOGY OF AUCHENORHYNCHA

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PHYLOGENY OF AUCHENORHYNCHA

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66 MORPHOLOGY OF AUCHENORHYNCHA

primitive nature of the ovipositor and the sucking pump, taken together
with the simple, unconvoluted condition of the intestinal canal reported
for the Fulgoridae by Handlirsch (developed into the specialized “Filter
chamber” in the remaining auchenorhynchous families), would indicate
to the writer that the Fulgoridae is a group which is distinctly separate
from all other Auchenorhyncha, and was the earliest to be differentiated
from the common base of the Auchenorhyncha.

Other characters which separate the Fulgoridae from the remaining
families are the fusion of the meron of the coxa of the metathoracic leg
with the epimeron, and the greatly modified second segment of the an-
tenna. These characters, however, represent specializations rather than
primitiveness. The Fulgoridae are also distinct in the possession of a
tegula, absent in other Auchenorhyncha, as far as I have been able to
ascertain.

Before proceeding further, it would be well to point out that the ovi-
positor of Scolops pungens is considerably more primitive than that
found in many representatives of the Orthoptera, which group has man-
dibulate mouthparts. Thus we can see that within two distinct orders we
have a combination of both primitive and specialized characters. The
more primitive mandibulate mouthparts are combined with highly spe-
cialized ovipositors in many groups of Orthoptera, whereas the fulgorid
combines specialized sucking mouthparts (compared to mandibulate
mouthparts) with a primitive ovipositor. This should make it clear that
insects by themselves are neither primitive nor specialized, and that
these terms can only be applied in a true, comparative sense to the indi-
vidual parts of insects. The fulgorids, for example, have been referred to
by some authors as being primitive insects, by others (Kirkaldy, 1906;
Funkhouser, 1917) as being specialized. Such terms offer no guide to
phylogeny, for it is an evaluation of separate characters which is im-
portant in the interpretation of relationships.

There has been considerable controversy concerning the origin of the
loral areas. Snodgrass (1938), in supporting his theory that the lorum
is derived from the hypopharynx, suggested that the loral areas in the
Fulgoridae represent a specialized, rather than a generalized condition.
In view of the fact that the clypeal, sucking pump region of the fulgorid
is a primitive one, in comparison with other Auchenorhyncha, it would
support the contrary concept that the outer part of the lorum is differ-
entiated from the lateral area of the clypeus, rather than from the hypo-
pharynx. Attention is called to the narrow lateral arm of the hypopharynx
(la, Figs. 9 and 10) which connects to the lorum. It seems likely that
this narrow arm between the hypopharynx and the lorum has been dif-
ferentiated from the hypopharynx, and has apparently expanded, in the

PHYLOGENY OF AUCHENORHYNCHA 67

other families of Auchenorhyncha, to form the broader connection. Thus
it might well be that although the loral areas are derived mainly from
the lateral regions of the clypeus, a part of the hypopharynx has formed
a bridge with the present structure.

The relationships and origins of the remaining groups of Auchen-
orhyncha is a much more difficult matter. Much of the difficulty arises
from the fact that when we consider the Cicadidae, Cercopidae, Cicadel-
lidae, and Membracidae, we tend to think in terms of the numerous,
recent representatives of these groups. These recent representatives have
a number of characteristics which, taken together, mark these families off
distinctly. Thus, the cercopid, cicadellid, and membracid possess an un-
mistakable similarity in the general plan of the thoracic sterna which
links these three forms together, and differentiates them from the cicada.
No doubt this similarity of the thoracic sterna is correlated with another
distinguishing feature, the presence of jumping hind legs, which sep-
arates the Cercopidae, Cicadellidae, and Membracidae from the Cicadi-
dae. In addition, the head of Cicadidae retains a small fronslike area
bearing a median ocellus, absent in Cercopidae, Cicadellidae, and Mem-
bracidae. It should be pointed out, too, that the loral areas lie alongside
the entire length of the postclypeus in the cicada, clearly the more an-
cestral condition as indicated in the fulgorids, whereas in the cercopid,
cicadellid, and membracid, the upper portions of the lorae are folded
into the head capsule or reduced, so that they are seen to lie adjacent to
the ventral portion of the postclypeus only. All these characters suggest
that the Cicadidae are distinct from the Cercopidae, Cicadellidae, and
Membracidae.

In this connection attention is called to the pleural wing grooves,
present in Lepyronia, Aulacizes, and Ceresa, which enable the bases of
the anterior margin of the tegmina to lock in place when at rest. This
device is absent in Tibicina. If such structures are consistent within Cer-
copidae, Cicadellidae, and Membracidae, they may represent another
reason for grouping these families apart from the Cicadidae.

Of these remaining three groups, the Cercopidae alone possess a com-
plete tentorium in which the anterior tentorial arms are connected with
the posterior arms (as is also the condition in the Cicadidae), whereas
in both the Cicadellidae and Membracidae the tentorial structure is re-
duced; the anterior arms no longer retain a connection with the posterior
arms. The hind legs of both Membracidae and Cicadellidae bear spines,
which are absent on the hind legs of Cercopidae. Evans (1946a) has rec-
ognized these affinities in grouping membracids and cicadellidids in one
superfamily, Jassoidea, as distinct from the Cercopoidea. In addition the
thoracic sterna of Aulacizes and Ceresa are exceedingly similar and both

68 MORPHOLOGY OF AUCHENORHYNCHA

possess a differentiated, mesothoracic antecoxal area (acx., Figs. 69 and
74), absent in Lepyronia.

It should be pointed out that a similar tendency toward the reduction
of the tentorial structure, already accomplished in the Cicadellidae and
Membracidae, is present also in the Fulgoridae. Although most members
of the Fulgoridae possess anterior tentorial arms, and in some the dorsal
tentorial arms are also present, the tentorial structure is reduced to the
posterior tentorial bridge in Scolops. The antennal muscles, normally
inserted on the dorsal tentorium in many insects, are here inserted on
the posterior tentorial bridge. This tendency toward the development or
reduction of particular structures within separate phylogenetic lines of
the Auchenorhyncha is a widespread one. We thus find that although
the modern representatives of these auchenorhynchous groups are suf-
ficiently distinctive and readily defined, certain more primitive repre-
sentatives show a remarkable number of features in common, and it is
for this reason that any attempt to determine when these familial groups
originated in point of time is such a difficult one.

The epistomal suture, lacking in some cercopids, the majority of
present-day jassids, and most membracids, is nevertheless present in
some representatives of all of these groups as well as in the fulgorids
and cicadas. The subgenal, or maxillary, suture, found in some of the
fulgorid representatives of this study, is completely present in some
jassids (Ulopinae and Stenocotini), and is vestigial in the prim-
itive cicada, Tettigarcta, some cercopids, and some jassids. It is absent
only in membracids. Further, Evans (1948) pointed out that the basic
membracid type of tegminal venation,* as represented by Xiphistes
tuberculatus, closely resembles the jassid type and differs only in that M
is not fused basally with R, and that probably the oldest representatives
of the membracids go back to very early times. This basic membracid
tegmen closely resembles Evans’ (1946a) reconstruction of the hypo-
thetical, ancestral tegmenal venation of the Jassoidea (Evans’ families
Aetaleonidae, Hylicidae, Eurymelidae, and Jassidae). Only when we
consider the more recent groups do we find specializations which are
of recent origin. Another character thought to be confined to the family
Cicadidae, namely the transverse nodal line of the tegmina, has been
found in the jassoid family Hylicidae. Apart from the Hylicidae, no trace
of such a line is retained in any other jassoids.

Additional evidence along these lines has recently been brought forth
in the interesting studies of Ossiannilsson (1949), who pointed out that

*According to Evans the arrangement of the veins of the tegmen offers a more
reliable character for the determination of relationships within the Membracidae, than
those of the highly variable pronotum (a much more recent specialization), hereto-
fore relied upon.

PHYLOGENY OF AUCHENORHYNCHA 69

the tymbal organs of Homoptera, generally considered to be present only
in the Cicadidae, are also present in cercopids, membracids (Centrotus),
some jassoid forms, and the fulgorid, Ommatidiotus. In cercopids and
certain jassoid genera the female possesses a functional sound-producing
organ of the same type as that of the male, though weaker. Moreover, the
female of Paropia has a distinctly striated tymbal which is, in the male,
only represented by traces. In Doratura both sexes have a sound-produc-
ing organ of the same type and equally well developed. Ossiannilsson
suggests that primitively both sexes possessed practically identical tymbal
organs, and that the present condition, wherein these organs are usually
confined to the males, has been brought about by reduction. He con-
cluded that the possession of a functional tymbal apparatus is general
among the Auchenorhyncha.

Evans (1940b) has called attention to the fact that not only the nymphs
of cicadas, but those of many cixiids (Fulgoridae) and certain cercopids
are subterranean, and that this may well be a primitive characteristic,
possibly associated with severe weather conditions prevailing at some
past period of geological history. The study of the habits and behavior
patterns of Auchenorhyncha is an almost unexplored field for the inves-
tigator and, if studied from the comparative point of view, could undoubt-
edly contribute much additional evidence to the solution of problems of
relationship.

All of this evidence supports the contention that only the numerous
modern representatives of the Cicadidae, Cercopidae, Jassidae, and
Membracidae are distinct and well defined; and that there are an im-
pressive number of instances in which primitive, less numerous, and little
known representatives of these groups show a variety of characters in
common.

It is interesting to note that the meso- and metathoracic segments of
Tibicina septendecim and Ceresa bubalus show certain superficial resem-
blances of shape and structure, as well as fore- and hindwings which are
both membranous. I have already pointed out that the affinity of the
membracid thorax lies with that of the cicadellid, and to a somewhat
lesser extent with that of the cercopid.

Lawson (1922) suggested that the main branch of the Cercopidae
arose a little earlier than that of the Cicadellidae and Membracidae,
based on the assumption that spittle-mass production must have required
a long time to perfect. Although there is morphological, and perhaps
palaeontological, evidence which might support an earlier origin of the
Cercopidae, such an earlier origin is not, in my opinion, supported by the
development of a specialization such as spittle-mass production. This is

70 MORPHOLOGY OF AUCHENORHYNCHA

but one example of the difficulties inherent in the attempts to reason out
phylogeny.

It is difficult to say whether there have been two “main” divisions of
development or three as Evans (1946) has recently suggested. If we are
to accept three, I would consider that the Cicadomorpha (Cicadidae )
and Jassidomorpha (Cercopidae, Cicadellidae, and Membracidae) are
more distinctly separated from the Fulgoromorpha (Fulgoridae) than
they are from each other, while acknowledging that the Cicadidae do show
distinct differences from the Cercopidae, Cicadellidae, and Membra-
cidae. The Fulgoridae were the earliest group to be differentiated from
the base of the auchenorhynchous stem, and are clearly distinct from
the remaining Auchenorhyncha. The cercopids, cicadellids, and membra-
cids are closely related, and the latter two groups show very marked
affinities. On the basis of this study, and our present knowledge of cicadas
on the one hand and the cercopids, cicadellids, and membracids on the
cther, the writer finds no conclusive evidence to support an earlier origin
for one or the other of these groups. Evans, however, has stated (in con-
versation ) that the modern cicadas were probably differentiated earlier
than modern jassids, but that the jassoid stem originated at an earlier
date than the cicadoid stem. As additional, primitive representatives of
these groups come to light, we may be able to make more valid conjec-
tures concerning their probable origin in time.

No attempt has been made to evaluate the taxonomic categories used
in this discussion. The family names have been used to indicate broad
groups and their general relationship. It is very likely that superfamily
and subfamily designations may properly replace certain family groups
as various writers have already suggested.

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1922. Ovipositors of Cicadellidae (Homoptera). Kans. Univ. Sci. Bull., 14:217-
298. 18 pls.

Russe 1, E. S.
1916. Form and function. A contribution to the history of animal morphology.
London, VI + 383 pp.
SincH-Prurut, H.
1924. The development of the male genitalia of Homoptera, with preliminary
remarks on the nature of these organs in other insects. Quar. Jour. Micr.
Sci., 69:56-96. 2 pls.
1925. The morphology of the male genitalia in Rhynchota. Trans. R. Ent. Soc.
London, 1925:127-267. 3 figs. 27 pls.

EIBLIOGRAPHY 13

Snoperass, R. E.

1921. The mouthparts of the cicada. Proc. Ent. Soc. Wash., 23:1-15. 15 figs.

1927a. The head and mouthparts of the cicada. Proc. Ent. Soc. Wash., 29:1-16.
11 figs.

1927b. Morphology and mechanism of the insect thorax. Smith. Misc. Coll.
(Publ. 2915), 80(1):108 pp. 44 figs.

1928. Morphology and evolution of the insect head and its appendages. Smith.
Misc. Coll. (Publ. 2971), 81(3):1-158. 57 figs.

1935. Principles of insect morphology. McGraw-Hill Book Company. 667 pp.
319 figs.

1938. The loral plates and the hypopharynx of Hemiptera. Proc. Ent. Soc.
Wash., 40:228-236. 11 figs.

1944. The feeding apparatus of biting and sucking insects affecting man and
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1947. The insect cranium and the “epicranial suture.” Smith. Misc. Coll. (Publ.
3896), 107(7):52 pp. 15 figs.

Spooner, C. S.

1938. The phylogeny of the Hemiptera based on a study of the head capsule.

Ill. Biol. Mon., 16(3):1-102. 398 figs.
TILLyArp, R. J.

1919. Mesozoic insects of Queensland. No. 7. Hemiptera-Homoptera; with a
note on the phylogeny of the order. Proc. Linn. Soc. N. S. Wales,
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1921. Mesozoic insects of Queensland, No. 8. Hemiptera-Homoptera (Cont'd. ).
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1926. Kansas Permian insects. Part 9. The order Hemiptera. Amer. Jour. Sci.,
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WALKER, FE. M.
1919. The terminal abdominal structures of orthopteroid insects: A phylo-
genetic study. Ann, Ent. Soc. Amer., 12:267-325. 75 figs.
1922. The terminal abdominal structures of orthopteroid insects. A phylogenetic
study. Part II. Male. Ann. Ent. Soc. Amer., 15:1-87. 107 figs.
WEBER, H.
1930. Biologie der Hemipteren. Julius Springer. Berlin. 543 pp. 329 figs.
1933. Lehrbuch der Entomologie, Jena. 726 + XII pp. 555 figs.
Witson, H. F. and Doner, M. H.
1937. The historical development of insect classification. John S. Swift & Co.,
133 pp.

LIST OF ABBREVIATIONS

1A —Ist anal

2A —2nd anal

3A -—8srd anal

aclp —anteclypeus

acs —antecostal suture
acx —antecoxal area

aed —aedeagus

aepm —anepimeron

aes —anepisternum

an —anus

anp —anterior notal wing process
ant —antenna

as _—anal style

at —anterior tentorial arm

atp —anterior tentorial pit, or
invagination

au —auditory capsule

aw -—prealar bridge

lax —first axillary sclerite

2ax —second axillary sclerite
38ax —third axillary sclerite

ba —basalare, or basalar sclerite
bex —basicoxite

bp —basal plate

br —brain

bst —basisternite

C —costa

cl —claw, or ungue

clr —clypeal ridge, or apodeme
cx —coxa

cxp -—pleural coxal process

Cu —cubitus

d —dorsal longitudinal muscle

de -—dorsal cervical sclerite

dl -—lateral dorsal longitudinal
muscle

dm —median dorsal longitudinal
muscle

76

diclp —dilator muscles of the clypeus
dsyr —dilator muscle of the salivary

syringe
dt —dorsal arm of the tentorium
e —compound eye

epm —epimeron
eph —epipharynx

es —episternum

est —epistomal suture

fe | —femur

fl —flagellum of the antenna
fp —furcal pit

fr —frons, or frontal region
fst | —furcasternite

fu. —furcal apodeme

ge —gena, or genal region

gnp —gonopore
gp —subgenital plate

ha —humeral angle of pronotum
hph —hypopharynx

hwp —hypopharygeal wing plate

is —intersegmental line

jf —jugal fold

kepm —katepimeron

kes —katepisternum

] —lateral abdominal muscle

Ib = —labium

Ie —lateral cervical sclerite

le —external lateral abdominal
muscle

Ih —lateral arm of hypopharynx

li —internal lateral abdominal
muscle

Im = —labrum

lor —lorum

Itg —laterotergite

Ivr, —lever of the mandibular stylet

LIST OF ABBREVIATIONS

—lever of the maxillary stylet
—media

—antennal muscle
—mandibular stylet

—muscle of the labium
—muscle maculum, or marking

—muscle partition formed by
apodemal wall of pronotum

—meron of coxa

—median suture of metathoracic
scutum

—metopidium of pronotum
—maxillary plate

—apodemal plate of the

maxillary
—nodal line of wing
—ocellus
—occipital condyle
—ocellar nerve
—operculum
—optic nerve
—precosta, or pretergite
—postclypeus
—pedicel of the antenna
—phragma
—pleural apodeme

—protractor muscle of the
labium

—pleural suture
—paramere

—protractor muscle of the
mandible

—sucking pump

—protractor muscle of the
maxillary stylet

—posterior notal wing process

—postnotum

—postcoxal bridge

—pronotum

—prescutum

—posterior tentorial
invagination

—pulvillus

pw

rmxs

rp

—postalar bridge

—radius

—reduplication of scutellum

—inner ramus, or sclerotized
thickening, of first valvula

—retractor muscle of the labium

—retractor muscle of the
mandibular stylet

—retractor muscle of the
maxillary stylet

—outer ramus, or sclerotized
thickening, of first valvula

—outer ramus, or sclerotized
thickening, of second valvula

—ramal plate

Is, Is. etc. —first abdominal

Se

syr

sternite, etc.

—subcosta
—scutum
—scutellum
—scape of the antenna
—subesophageal ganglion
—subgenal suture
—suprahumeral horn of

pronotum
—salivary duct
—salivary glands
—spiracle
—salivary syringe

It, IIt, etc. —first abdominal

tergite, etc.

ta —tarsus

tb _—posterior tentorial bridge
tg —tegula

thgng, —prothoracic ganglion
ti —tibia

tn —trochantin

tr _—trochanter

ts —tergal spine

twg —tergal wing groove
tyb —tymbal

typ —tympanum

—unguitractor plate

78

MORPHOLOGY OF AUCHENORHYNCHA

—ventral longitudinal muscle lvlf —first valvifer
—ventral clasper of 10th 2vlf —second valvifer
SOE MSI vst —vestibulum

—vannal fold
Vx —vertex

I, II, etc.—first abdominal
seginent, etc.

—first valvula
—second valvula
—third valvula

PLATES WITH EXPLANATIONS

PLATE I

Lateral view of head of Acalonia sp.

Frontal view of head of Acalonia sp.

Frontal view of head of Ormenis pruinosa (Say)

Lateral view of head of Ormenis pruinosa

Frontal view of head of Epitera septentrionalis Say

Frontal view of head of Oliarus aridus Ball, with maxillary lobes displaced
Frontal view of head of Scolops pungens

CORA OO aa

Lateral view of head of Scolops pungens
8a. Lateral view of head of Scudderia sp. (Orthoptera)

80

81

PLATE II

Sagittal section of lower part of head of Scolops pungens, showing at-
tachment of hypopharynx to the head capsule

Anterior views of hypopharynx of Scolops pungens

Posterior view of hypopharynx, maxillary plate and posterior tentorial
bridge of Oliarus aridus

Muscles and articulation of right mandibular stylet of Scolops pungens
Sagittal section of the head of Scolops pungens

Median section of labium of Scolops pungens

Muscles and articulation of right maxillary stylet of Scolops pungens
Posterior view of head of Scolops pungens, with labium removed
Posterior view of labium of Scolops pungens

a
ae SOOO

DpHowwpwp wv
oo 19

bo
be

b bp bo
Sa Sh

a

PLATE III

Tibicina septendecim
Sagittal section of head
Anterior view of head
Posterior view of hypopharyngeal wing plate
Lateral view of labium
Median section of basal segments of labium
Muscles of the maxillary stylet

Cross section of head through hypopharynx, sucking pump, and post-
clypeus

Posteroventral view of tentorium and hypopharyngeal wing plate of up-
per right half of head

Dorsal view of head
Posterior view of head, with labium removed
Muscles of the mandibular stylet

84

29.
30.
31.
32.
33.
34.
35.

36.

37.

PLATE IV

Lepyronia quadrangularis

Anterior view of head
Sagittal section of head
Dorsal view of head
Posterior view of head

Basal portion of right maxillary stylet
Muscles and articulation of right mandibular stylet
Anterior view of left half of head showing the connection of the

hypopharynx with the lorum

Cross-section of head through the hypopharynx, sucking pump, and

postclypeus
Median section of the labium

86

Nv

87

38.
39.
40,
41.
42.
43.
44,
45.

46.

PLATE V

Aulacizes irrorata
Anterior view of head
Dorsal view of head
Sagittal section of head
Posterior view of head
Muscles and articulation of the maxillary stylet
Lateral view of labium
Median section of labium

Cross-section of head through hypopharynx, sucking pump and _ post-
clypeus

Posterior view of internal section of head showing the articulation and
muscles of the mandibular stylet

88

i

| ly 1 |

s
=
=

89

A7.
48.
49.
50.
le
D2:
oe
54.

PLATE VI

Ceresa bubalus
Ventral view of head
Anterior view of head
Sagittal section of head
Posterior view of head
Lateral view of labium
Median section of labium
Articulation and muscles of maxillary stylet

Sagittal section of head, showing articulation and muscles of mandibular
stylet

90

91

PLATE VII

Lateral view of thorax of Scolops pungens
Dorsal view of thorax of Scolops pungens
Dorsal view of thorax of Tibicina septendecim
Lateral view of thorax of Tibicina septendecim

Dorsal section through intersegmental area of pro- and mesothoracic
segments of Tibicina septendecim

Ventral view of thorax of Scolops pungens
Ventral view of thorax of Tibicina septendecim

Posterior view of phragma between meso- and metathoracic segments of
Tibicina septendecim

Posterior view of prothoracic endoskeleton of Scolops pungens

92

PLATE VIII

Lateral view of thorax of Aulacizes irrorata
Dorsal view of thorax of Aulacizes irrorata

Dorsal view of thorax of Lepyronia quadrangularis
Lateral view of thorax of Lepyronia quadrangularis
Ventral view of thorax of Lepyronia quadrangularis
Ventral view of thorax of Aulacizes irrorata

94

95

PLATE IX

Ceresa bubalus
Dorsal view of meso- and metathoracic segments
Ventral view of meso- and metathoracic segments
Posterior view of prothorax
Posterior view of metathorax
Ventral view of thoracic segments
Anterior view of prothorax
Anterior view of metathorax
Posterior view of endoskeleton of mesothorax
Anterior view of mesothorax

Yy /,
Uy
Yj Yi A
Vy

VHD -=:
Ss PCX2

97

Tes),
80.
oul
82.
83.
84.
85.
86.
87.
88.

PLATE X

Forewing of Scolops pungens
Hindwing of Scolops pungens
Forewing of Tibicina septendecim
Hindwing of Tibicina septendecim
Forewing of Lepyronia quadrangularis
Hindwing of Lepyronia quadrangularis
Forewing of Aulacizes irrorata
Hindwing of Aulacizes irrorata
Forewing of Ceresa bubalus

Hindwing of Ceresa bubalus

98

99

89.
90.
wil,
92.
93.
94.
95.
96.
oie
98.
99.
100.

101.
102.
103.
104.
105.

PLATE XI

Prothoracic leg of Scolops pungens

Mesothoracic leg of Scolops pungens

Metathoracic leg of Scolops pungens

Tarsal segments and claws of mesothoracic leg of Scolops pungens
Prothoracic leg of Tibicina septendecim

Mesothoracic leg of Tibicina septendecim

Metathoracic leg of Tibicina septendecim

Tarsal segments and claws of metathoracic leg of Tibicina septendecim
Prothoracic leg of Lepyronia quadrangularis

Mesothoracic of Lepyronia quadrangularis

Metathoracic leg of Lepyronia quadrangularis

Tarsal segments and claws of mesothoracic leg of Lepyronia quadrang-
ularis

Prothoracic leg of Aulacizes irrorata

Mesothoracic leg of Aulacizes irrorata

Metathoracic leg of Aulacizes irrorata

Tarsal segments and claws of prothoracic leg of Aulacizes irrorata
Lateral view of claw of prothoracic leg of Aulacizes irrorata

100

101

106.
107.
108.

109.
110.
iT
112.
113.
114.
115.
116.
117.

PLATE XII

Ceresa bubalus
Median sagittal section of head and thorax
Lateral sagittal section of meso- and metathoracic segments

Lateral sagittal section of meso- and metathoracic segments, showing
latermost muscles

Lateral sagittal section of mesothoracic segment
Posterior view of prothoracic muscles

Prothoracic leg

Mesothoracic leg, showing coxal muscles
Metathoracic leg

Trochanteral and femoral muscles of mesothoracic leg
Tibial muscles of mesothoracic leg

Tarsal and posttarsal muscles of mesothoracic leg
Internal view of metathoracic pleuron and muscles

102

Y=
54c & !

‘Se

103

118.
11S:
120.
ile
122.
123.
124.
125.
126.
127.

PLATE XIII

Ventral view of basal abdominal segments of Scolops pungens

Lateral view of female abdomen of Scolops pungens

Ventral view of basal abdominal segments of Tibicina septendecim
Lateral view of female abdomen of Tibicina septendecim

Ventral view of basal abdominal segment of Lepyronia quadrangularis
Lateral view of female abdomen of Lepyronia quadrangularis

Ventral view of basal abdominal segments of Aulacizes irrorata

Lateral view of female abdomen of Aulacizes irrorata

Ventral view of basal abdominal segments of Ceresa bubalus

Lateral view of female abdomen of Ceresa bubalus

104

128.
129.
130.
131.
132.
133.
134.
135.
136.
137.

138.
139.
140.
141.
142.

PLATE XIV

Lateral view of distended parts of ovipositor of Lepyronia quadrangularis
Basal plate, aedeagus and paramere of Lepyronia quadrangularis
Terminal abdominal segments of male, Tibicina septendecim

Lateral view of genitalia of female, Scolops pungens

Ventral view of genitalia of female, Scolops pungens

Lateral view of genitalia of male, Scolops pungens

Lateral view of genitalia of female, Tibicina septendecim

Ventral view of genitalia of female, Tibicina septendecim

Lateral view of male genitalia, Tibicina septendecim

Lateral view of female terminal abdominal segments, Tibicina septen-
decim

Caudoventral view of ninth segment and aedeagus, Tibicina septendecim
Lateral view of female genitalia, Lepyronia quadrangularis

Lateral view of male genitalia, Lepyronia quadrangularis

Lateral view of female genitalia, Aulacizes irrorata

Lateral view of male genitalia, Aulacizes irrorata

106

139 Wit DViE

107

143.
144.
145.
146.
147.
148.
149.

PLATE XV

Ceresa bubalus
Sagittal section of visceral abdominal segments
Ventral view of female genitalia
Terminal abdominal segments
Muscles of the ovipositor
Second valvifer with second and third valvulae
Internal view of valvifers, valvulae, and rami of ovipositor
Lateral view of ovipositor

108

Ay,

——1

Tih

4
NP) Sor ce

a

IIT shall Mr 4)
ae: :
Mit

109

PLATE XVI

Ceresa bubalus
150. Caudal view of male genitalia
151. Lateral view of male genitalia
152. Muscles of the male genitalia, lateral view

153. Dissection showing paramere, basal plate, aedeagus, tenth segment, and
musculature

154. Ventral view of subgenital plate
155. Caudal view of aedeagus

110

ALL

.

-

n

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