THE MORPHOLOGY 05 ‘fHE
GENSTALIA AN!) A$$OClATED ABDOMINAL

SEGMENTS OF
MONOCHAMUS FAROLINENSIS (OLIVIER)

(COLEOPTERA: CERAMBYCIDAE)

 

 

Thesis {or Hm Degree of M. S
MEEHIGAN SMTE Ufii‘s’ERSlTY
Shu Chen Chang
1963

THESIS

LIBRARY

Michigan State
University

 

 

THE MORPHOLOGY OF THE GENITALIA AND ASSOCIATED ABDOMINAL

SEGMENTS 0F MONOCHAMUS CAROLINENSIS (OLIVIER)

 

(COLEOPTERA: CERAMBYCIDAE)

BY

Shu Chen Chang

A THESIS

Submitted to
Michigan State University
in partial fulfillment of the requirements
for the degree of

MASTER OF SCIENCE

Department of Entomology

1963

ACKNOWLEDGEMENTS

The author wishes to eXpress his sincere thanks to Professor
Roland L. Fischer of the Department of Entomology, under whose kind
guidance, rigorous criticism, and constant supervision this research
work was undertaken.

The author is greatly indebted to Professor Gorden E. Guyer,
Chairman of the Department of Entomology, for his stimulating advice
and frequent encouragement.

Appreciation is extended to Mr. Steven Ilnitzky for his aid
in obtaining material from Gull Lake.

The staff of the Department of Entomology provided valuable aid
during the course of the work. Mr. Lan-Lin wu of the University of
Alberta, Canada, provided important references, Mr. Chia-Moo King of
Ta Shu Shan Forestry Corporation of Taiwan, and Mr. Chung Tu Yang of
Chung Hsing university sent relative material. This aid is acknow-

ledged gratefully.

ii

TABLE OF CONTENTS

Page
ACKNOWLEDGEMENTS........................ ii
I.INTRODUCTION....................... 1
II.REVIEWOFLITERATURE................... 2
III. THE GENERAL MORPHOLOGY OF THE ABDOMEN. . . . . . . . . . . 4
IV. THE ASSOCIATED ABDOMINAL SEGMENTS......... . .. . 6
V.THEGENITALIA....................... 9
VI. DISCUSSION . . . . . . . . . . . . . . . . . . . . . . . . 11
VII. CONCLUSION . . . . . . . . . . . . . . . . . . . . . . . . l6
REFERENCESCITED........................ 18

EXme IONS OF PIAT E S O O O O O O O O O O O O O O O O O O O I O 2 0

iii

I. INTRODUCTION

The present paper deals with the structure of the genitalia and
associated abdominal segments of one of the North American Species of

long-horned beetles, MOnochamus carolinensis (Olivier). Those who have

 

previously figured the genitalia of certain groups of Coleoptera have
usually made no attempt to homologize the structures with those of the
related groups. Workers have unfortunately used their own Special terms
without regard to previous terminology. This has resulted in a system
of terminology which is complex and confused. An attempt is made to
elucidate concepts pertaining to the structure of the genitalia of
coleopterous insects.

Little has been published concerning either the morphological or
ecological aSpects of the long-horned beetles. The structure of the
genitalia is important in both the fields of morphology and taxonomy.
Representatives of the subfamilies and tribes should be studied in
detail throughout the family Cerambycidae. AS time was limited only

this Species was studied, the results of which are presented.

II. REVIEW OF LITERATURE

In the literature there is limited information on the muscular
mechanism of the abdominal segments of the cerambycids and only a few
important papers on the genitalia of beetles can be found. In addition,
most of the papers were published in the second half of last century or
early in the 20th century. Their points of view are usually beyond the
SCOpe of homology and phylogeny and are inadequate for the present.

The most outstanding paper of these is that of Tanner (1927),
who has done considerable work on the morphology of female genitalia of
cole0pterous insects. He has advanced our knowledge of the female
genital apparatus considerably, but like many others has failed to
homologize the Structures and was mislead as to the segmentation and
interpretation of the Structure of genitalia. He has treated the whole
structure that retracted into the body including the genital and post-
genital segments as the female genitalia.

Another remarkable paper is that of Sharp and Muir (1912). They
divided the male genitalia of coleOpterous insects into two main portions,
the zygotic and azygotic. The azygotic portion was further subdivided
into the eurazygotic and stenazygotic portions. As they explained these
terms, the azygotic portion, or azygos, comprises all the unpaired
portion of the tube from the body wall to the divergence of the ends of
seminal ducts; the zygotic portion, or zygos, is formed by the two
seminal ducts preceeding from the testes; the eurazygotic portion, or

eurazygos, is the enlarged portion of the genital tube; and the

2

3
stenazygotic portion or stenazygos, is the slender portion of the
azygos.

It is very clear that the zygotic portion (the zygos), is the
seminal duct or the vasa deferentia; the stenazygotic portion (the
stenazygos) is the ejaculatory duct, or the ductus ejaculatorius; and
the eurazygotic portion (the eurazygos), is the phallus with internal
sac. The whole apparatus is rather simple in structure. To divide
the genital apparatus into so many portions and establish so many
Special terms seems unnecessary and merely adds to the confusion in

terminology.

III. THE GENERAL MORPHOLOGY OF THE ABDOMEN

A consideration of the functional aSpects of the insect genitalia
must be based upon adequate understanding of the morphology of the

abdomen. Thus, a brief description of the abdomen of Monochamus

 

carolinensis (Olivier) (which this study deals with) follows.
In general, the insect abdomen may be divided into three divisions:
First, the pregenital or visceral segments, including the first
to seventh segments, are usually of simple form and structure and differ
but little from one another.
Second, the genital division, including the eighth and ninth
segments, usually bears the genital aperture and genital appendages.
Third, the postgenital division includes segments ten and eleven.
This region usually bears the anus and terminal appendages, such as the
cerci and the telson, and in certain forms the median caudal filament.

In Monochamus carolinensis (Olivier) as well as other long-horned

 

beetles, the abdomen is rather Simple. Only seven segments are visible
on the dorsal surface (Figure 2) and five on the ventral (Figure 1).

All the terga, except the last one, are similar in size, Shape and weakly
sclerotized, as they are covered by the hardened elytra. Some forms

with narrow elytra as Thranius spp., with short elytra as MO1orchuS $22.,

 

and StenoEterous Spp., and certain others are exceptions. The last

 

visible tergum, the seventh, is larger than the preceding and unlike
others is strongly sclerotized, as it is usually beyond the cover of the

elytra, SSpecially in the female. The sterna are five in number, as

4

5
the first two are concealed and overlapped by the hind coxae. All of
the sterna are strongly sclerotized. Only the last visible sternum,
the seventh, differs in shape from the preceding.
The eighth, the ninth, and the post-genital segments are all re-
tracted into the abdomen and concealed by the seventh tergum and

sternum.

IV. THE ASSOCIATED ABDOMINAL SEGMENTS

The abdominal segments associated with the genitalia are the
seventh segment (the last visceral segment), the eighth segment (the
first genital segment), the ninth segment (the second genital segment),

and the tenth segment (the periproct).

The male

The seventh segment.--The seventh tergum is much more sclerotized

 

than the preceding, setiferous along the posterior margin, and with a
broad projection on the anterior margin which projects forward into the
body cavity under the preceding tergum. This projection functions as an
apodeme and increases the area of attachment for muscles. This tergum
is termed the pygidium in taxonomic literature, visible externally and
usually extending beyond the cover of the elytra.

The sternum.of the seventh segment is taxonomically known as the
subgenital plate. Its posterior margin is much narrower than its
anterior margin. Along the posterior margin there are many bristles
and a thin triangular region demarked by darker color and larger hairs.

The eighth Segment.--This segment, as well as the ninth, is re-

 

tracted within the body in its usual condition. It is much reduced in
size. The tergum of the eighth is complete, with the same shape as the
seventh tergum only much smaller in size (Figure 15). The sternum
(Figure 16) is much reduced and present as a broad band-like sclerite
with a thin Y-Shaped and strongly sclerotized apodeme at the base

6

7
(the anterior portion). This apodeme projects forward and functions for
support of muscles which evert and retract this segment (Figure 11).

The ninth segment.--The structure of this segment (Figure 17) is
greatly modified. The entire tergum and sternum are pliable and toughly
membranous. When the ninth segment retracts inwardly it successively
ensheathes the genitalia in its usual state, and forms the so called
genital chamber. On the ventral basal margin there is a slightly
larger Y-shaped apodeme similar to that of the eighth sternum.

The tenth segment.--This segment has been called the periproct which

 

bears the epiproct and paraprocts surrounding the anus. But in this in-
sect, it appears only as a small projection from the dorsal wall of the

entrance chamber.

The female

 

The seventh segment.--The subgenital plate (Figure 3) and tergum

 

or the pygidium (Figure 4) are Similar to those of the male. The
posterior margins are Slightly broader when compared with the anterior.

The eighth segment.--This segment (Figures 7, 8, 9) is extremely

 

modified in shape and structure. Its tergum is complete but much re-
duced, with long hairs on its posterior angles, and its lateral margins
extended to the ventral aSpect involving the lateral portions of the
sternum. The sternum is membranous, in great part, with a wing-like
and slightly sclerotized plate at the basal portion and a large and
strongly sclerotized plate mesally. A very long and strongly sclero-
tized apodeme arises from the mesal plate and projects forward through
the abdominal cavity with its tip into the metathorax. This extremely

long apodeme serves for the attachment of muscles which evert the eighth,

8

and the following segments, and genitalia at the time of COpulation.
Muscles are connected to the tip of this apodeme and the anterior margin
Of the seventh sternum ventrally. When contracted, these muscles re-
sult in the eversion of the eighth segment, the following segments, and
genitalia. In the dorsum, there are muscles arising from the tip of
the apodeme which connect to the oviductus communis directly. When
contracted these same muscles indirectly pull the ninth segment within
the abdomen prOper. Thus they also function as retractors of the ninth
segment and the genitalia.

In its usual condition, the eighth segment as well as those
posterior to it are retracted within the body.

The ninth segment.--This Segment is notable by its entirely com-

 

plicated folding membranous condition with the dorsal anus and the
external genitalia ventrally at its terminal end. In the usual con-
dition, the terminal-half retracts into the basal-half with the apex

of the tube forming a double-walled tube (Figure 9). The cavity of the
double-walled tube is the genital chamber. The retracted ninth segment
with the genitalia in its cavity retracts into the eighth segment. It
everts only during the period of COpulation or egg-oviposition.

The tenth segment.--This segment is markedly reduced, only the

 

proctiger, a portion of the tenth tergite, is visible. It bears the
anus which is located on the dorsal base of the genitalia and beneath

the proctiger. The epiproct and paraprocts are lacking.

V. THE GENITALIA

The male

The male genitalia (Figures 11-14) of this Species consists only
of the phallic structures, the aedeagus, and the phallobase. All the
structures are derived from the appendages of the ninth abdominal
segment.

The aedeagus has develOped from the second gonapOphysis and con-
tains the phallus and the internal sac. The two united gonapOphyseS
form the phallus prOper, which invests the internal sac. The bases of
two parameres connect to the phallobase with membrane. These continue
anteriorly to form the struts. At the terminal end of the phallus there
is an Opening, the median orifice, along the edge of which it connects
with the membrane of the internal sac, and through which the internal
sac may evert during COpulation.

The internal sac (terminal vesica of Snodgrass) connects to the
median orificedat its basal end, to the ejaculatory duct with its apical
end. It is retracted into the ninth abdominal segment in the usual
condition and everted through the median orifice during COpulation.
When it is evaginated the Opening at its terminal end is the genital
aperture or gonOpore.

The phallobase (tegmen of Sharp and Muir) is develOped from the
coxites of the ninth abdominal segment and well differentiated from the
aedeagus. It bears a pair of valves called the parameres or lateral
lobes and a median dorsal lobe called the epimere or the basal piece.

9

10
From its base, an apodeme called the terminal strut projects into the
body cavity and serves as the attachment of muscles.

In coition, only the aedeagus is the intromittent organ, namely
only the phallus with the internal sac inserts into the female genitalia
and the latter when everted is the only functional part. The phallobase
is left outside of the female genitalia but in the genital chamber with
its terminal end against the ventral chamber wall and its struts against
the dorsal chamber wall. It maintains the position of the aedeagus in
the female genitalia and is similar to that of the clasPers in other
male insects. Thus it functions for the aedeagus during copulation not

only as a support but as a guide and also as a sustaining apparatus.

The female

 

The female genitalia (Figures 5-9) of this insect consists of two
main parts, the genitalia prOper and the genitalia valvae or egg guides.
The former is presumed to have develOped from the ninth gonapOPhyseS
which united together to form a completely membranous tube-like structure.
It bears the anus dorsally at its base, and the valvae on its sides.

The latter are presumed to have develOped from the ninth gono-
coxites. Apically each valva bears a small Stylus which internally is
supported by the baculum, a rod-like sclerotized structure. This
functions as a stiffening structure for the valva and a support for the
genitalia. The valvae serve as a guide for COpulation as well as for

oviposition.

VI. DISCUSSION

An attempt should be made to establish a common understanding re-
garding the morphological concepts of pterygotan insect genitalia. For

the sake of convenience, the interpretation of Michener (1944) is

 

quoted:
Origin Female Male
Coxites of eighth abdominal First valvifer I Absent in Pterygota
segment
Gonapophyses of eighth First valvae Absent in Pterygota

abdominal segment

Coxites of ninth Second valvifer Gonocoxites
abdominal segment ‘

Gonap0physes of ninth Second valvae Penis valves (2)
abdominal segment

Styli of ninth Third valvae Gonostyli
abdominal segment

This table shows that the higher female insectan genitalia is com-
posed of the coxites and the gonapOphyses of the eighth abdominal
segment, and the coxites, the gonapophyses, and styli of the ninth
abdominal segment. In the male, only the coxites, gonapophyses, and
styli of the ninth abdominal segment are involved.

Gustafson (1949) Stated that, "The appendage rudiments develOp
into the coxites and their styli; while the ampullae from the gonap0physes
of both male and female and the phallus of the male. The genitalic
patterns of all insects are derived from a common plan." He states

11

 

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a
a __ ,n 7
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12
further that, “Evolutionary tendencies of the ovipositor are demon-
strated in the complete loss of the Structure, in the reduction in Size
of the inner valvae (gonopophyses of the ninth segment) with the con-
current development of gonocoxites of segment nine to form a pair of
valves greatly resembling the gonopOphyses of segment eight, and in the
general reductiOn of all component structures. The latter condition is
best demonstrated in the Coleoptera.

"The males show a great stability in the retention of the gono-
Coxites of the ninth abdominal segment and to a lesser degree the styli
of that segment. There is a strong tendency for the gonapOphyses
(Parameres) of the male to fuse with the phallus.

"The primitive phallus of the insects is primarily a paired
structure.

“The genitalia appendages of all insects may be directly compared
with the basic pattern as demonstrated in the Thysanura.

"No completely new structures have arisen, all variations from
the basic pattern occurring by the modifications of existing appendages
and sclerites or by loss".

In fact, there is a common tendency in the evolution of the in-
sectan abdomen to gradually Simplify it from the abdomen of lower
groups to those of the higher groups.

The number of the abdominal segments is gradually reduced,,as in
the Order Protura where the number of abdominal segments is twelve.

The Order Collembola is an extreme exception, for the abdomen of this
group has only six segments, even in embryonic stages. In the lower
groups of ptergotan insects, such as the orthopteroid and hemipteroid

orders, the visible abdominal segments are always less than twelve,

13
usually nine or ten in number. In higher groups, the neuropteroid
orders, the number of visible abdominal segments is even more reduced.
It is not rare that six or seven abdominal segments may be found in
Cole0ptera, five in HymenOptera, and three in the Diptera.

Abdominal appendages borne on the pregenital segments (one to
seven) are found only in the apterygotan orders, such as Protura,
Collembola, Diplura, and Thysanura, and have never been found in the
pterygotan orders. As for those appendages borne on the genital or
postgenital segments, though they may be found throughout the orders,
they have disappeared or are concealed in the abdomen in higher
groups.

Because the insect abdomen and its appendages are reduced in sum
and form in the higher groups, there is no reason to think of the
genitalia which developed from the appendages of the eighth, and ninth
abdominal segments as being more complex than those of the lower in-
sects.

In this Species as well as its allies and even in the whole order
of Coleoptera, the female genitalia consists only of the appendages of
the ninth abdominal segment. However, former workers have used various
terminologies reflecting their Opinions concerning the Structures of the
female genitalia of coleopterous insects. Stein (1847) considered the
structures of the female genitalia Of cole0pterous insects as the
divided ninth sternum. Verhoeff (1894) considered them as the tenth
sternum. Wheeler (1893) considered them as rudimentary abdominal
appendages and designated them as ninth valvae which consist of the
valvifers, coxites, and styli. Peytoureau (1895) thought that they

were the eighth sternum in the Dytiscidae and the seventh in the

l4

HydrOphilidae. Wendollek (1905) considered them as the tergum or sternum
of ninth segment. MacGillivray (1923) contended that the "OctOpOphyses"
have migrated caudad from the eight segment to their present position,
the posterior margin of the ninth segment. Crampton (1925) considered
them as rudimentary abdominal appendages and designated them as ninth
valvae, including the valvifers, coxites, and Styli, the same as that of
Wheeler, and suggested that the paraproct of the beetles may be a part
of the ninth tergite called the surstylus in other insects, as in
TrichOptera. Tanner (1927) considered them as the coxites and styli of
the ninth segment, and that the valvifer are a sclerotized portion of
the adjacent end of the ninth sternite which bears the cOxites and styli,
and some sclerotized remnants of the tenth sternite in some Species,
and proposed to call the rod like structures of the paraprocts, valvifers
and coxites, the baculi which are lying in the integument and serving
as stiffening structures for the long membranous expanses of the ovi-
positor.

From all the evidence obtainable, there seems to be no reason for
considering parts of the female genitalia of coleOpterous insects as
the structures of the eighth or tenth segment. Some aSpects in Tanner's
paper (1927) are worthy of considerable attention. His statement that
"The genitalia consist of the eighth, ninth, and tenth abdominal
segments and their appendages" is not true in the Cerambycidae. The
associated eighth, ninth, and tenth abdominal segments are not included
in the composition of the genitalia. The genitalia are developed from
the appendages of the ninth abdominal segment only. In addition, the
appendages of the tenth segment, the cerci, which have functions re-

lated to reproduction are only found in Embioptera or other very rare

15
cases. So the statement, "the genitalia consist of eighth, and tenth
segments and their appendages" is to inclusive. The statements that
"The proctiger is a part of the tenth tergite. The paraprocts are
probably parts of the ninth tergite.”‘is not true in any case. The
paraprocts are always accompanied with the epiproct, if they are
present, and the epiproct is always located on the proctiger. Since
the proctiger is a portion Of the tenth tergite, the paraprocts should
also belong to the tenth segment. To think of the proctiger as an
upper genital valvae is also a misleading concept, because it is always

located above the anus.

VII. CONCLUSION

From a morphological study of the structure of the genitalia and

abdomen of Monochamus carolinensis (Olivier), it has been shown that

 

seven visible abdominal terga and five sterna are present. The
structure of the external genitalia is much simpler than that of the
lower pterygotan insects. The male genitalia consists only of the
phallus and phallobase, but the former with an internal sac and the
latter with the modified valvae. Both of these structures are develOped
from the appendages of ninth abdominal segment. The phallobase functions
similarly to those of the claSpers of other insects, but serves to
prevent its forward movement instead of holding. The female external
genitalia consists of the genitalia prOper and the genital valvae.

Both of these structures are develOped from the appendages of the ninth
abdominal segment. The modified genital valvae with styli on or near
their terminal end have the functions of support for the genitalia and
guide for COpulation and oviposition. The seventh abdominal tergum,

the dorsogenital plate, is well develOped, with one (in male) or two

(in female) expansions (the apodeme) along the anterior margin and long
bristles along the posterior margin, the bristles at the angle being
much longer. The seventh abdominal sternum, the subgenital plate, is
also well develOped, with long bristles along the posterior margin and
conceals the genital and postgenital segments within the body. The
eighth abdominal segment is much reduced in size and retracted within

the body in its usual condition. In the ventral area there is a rod-like

l6

 

I
-_ A
A

 

17
structure, the apodeme, which serves as an attachment for muscles and
has the functions of supporting, everting, and retracting this segment
and the others succeeding to it. This rod-like apodeme in the female is
eSpecially long and retracted through the abdominal cavity into that of
the metathorax. The ninth abdominal segment is much modified in shape
and much reduced in size. The tergum and sternum of this segment are
entirely membranous. When its terminal half retracts inwardly it
successively ensheathes the genitalia and forms the genital chamber.
In the male, there is a similar apodeme to that of the eighth sternum,
in the ventral region. The tenth abdominal segment, which bears the
anus, is located dorsally on the base of the external genitalia and
behind the ninth segment, It is very reduced, but in the female its
epiproct and paraprocts seem to be sclerotized. The external genitalia
both of the male and female is develOped only from the appendages of
the ninth abdominal segment. The terminal abdominal segments are not
included in the composition of the external genitalia but merely

associated with it.

REFERENCES CITED

Crampton, G. C. 1920. Remarks on the basic plan of the terminal
abdominal structures of the males of winged insects. Canad.
Entom. 53:178-183.

. 1921. Correction of a statement concerning the terminal
abdominal structures of male insects. Canad. Entom. 53:75.

. 1925. .EE Tanner (1927) fide below.

. 1929. The terminal abdominal Structures of female insects
compared throughout the orders from the standpoint of phylogeny.
Jour. New YOrk Ent. Soc. 33:453-490.

. 1938. The structures called parameres in male insects.
Bul. Brooklyn Ent. Soc. 33:16-24.

Gustafson, J. F. 1949. The origin and evolution of the genitalia of
the insecta. Stanford Univ. Bul. 24:74-76.

1950. The origin and evolution of the genitalia of the
insecta. Mflcroentomology 15(2):35-67.

Lindroth, C. H. 1957. The principal terms used for male and female
genitalia in Coleoptera. Opuscula Ent. 22(2/3):241-256.

MacGillivray, Alex D. 1923. External insect anatomy. Urbana, 111.,
Scarab Co. 388 pp.

Michener, C. 1944. A comparative study of the appendages of the eighth
and ninth abdominal segments of insects. Ann. Ent. Soc. Amer.
37:336-351.

Muir, F. 1924. The male genitalia of Cupes concolor westw. (ColeOptera).
Jour. New YOrk Ent. Soc. 32:167-169.

 

Peytoureau, A. 1895. Contribution a l'etude de la Morphologie de
1 Armure Genitale des Insectes. Bordeaux. 248 pp.

Sharp, D. and F. Muir. 1912. The comparative anatomy of the male
genital tube in ColeOptra. Trans. Ent. Soc. London 1912:477-461.

Singh-Pruthi, Hem. 1929. Homologies of the genitalia of insects.
Ent. Monthly Mag. 65:189-201.

18

19

Snodgrass, R. E. 1931. Morphology of insect abdomen. Part I.
General structure of the abdomen and its appendages. Smithsonian
Misc. Collections 85(6):l-128.

. 1933. Morphology of the insect abdomen. Part II. The
genital ducts and the ovipositor. Smithsonian Misc. Collections
89(8):1-l48.

. 1936. MOrphology of the abdomen. Part III. The male
genitalia (with arthrOpods other than insects). Smithsonian.Misc.
Collections 95(14):1-96.

. 1957. A revised interpretation of the external reproductive
organs of male insects. Smithsonian Misc. Collections 135(6):1-60.

Stein, F. 1847. Vergleichende Anatomie und Physiologie der Insecten.
Erste Mbnographie: Die weiblichen Geschlechtsorgane der Kafer.
Berlin, Dunker und Humblot. 139 pp.

Tanner, F. M. 1927. A preliminary study of the genitalia of female
ColeOptera. Trans. Amer. Ent. Soc. 53:5-50.

Verhoeff, Carl. 1894. Zur Kenntniss der vergleichenden MOrphologie
des Abdomens der Weiblichen ColeOptera. Deutsche Entomol.
Zeitschr. 1894:177-188.

wandolleck, Benno. 1905. Zur vergleichenden Morphologie des Abdomens
der weiblichen Kfifer. Zool. Jahrb. 22:477-576.

Wheeler, Wm. M. 1893. A contribution to insect embryology. JOur.
Morphology 8:1-160.

EXPLANAT IONS 0F PLATES

Figure
Figure
Figure

Figure

21

Ventral aSpect of the female abdomen.
Dorsal aSpect of the female abdomen.
Ventral aSpect of the female seventh sternum.

Dorsal aSpect of the female seventh tergum.

 

 

 

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25

Figure 7. Dorsal view of the eighth and ninth segments showing
the relation of the female genitalia.

Figure 8. Ventral view of the eighth and ninth segments showing
the relation of the female genitalia.

Figure 9. Dorsal view of the eighth and ninth segments showing
the female genitalia protruded.

Figure 10. Ventral view of the eighth and ninth segments showing
the female genitalia protruded.

'— ”VIII TE RGUM

    
  
  
   
  
  
   

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27

Figure 11. Dorsal view of the eighth and ninth segments showing
the relation of the male genitalia.

Figure 12. Ventral view of the eighth and ninth segments showing
the relation of the male genitalia.

Figure 13. Enlarged dorso-lateral View of the external male
genitalia.

Figure 14. Enlarged latero-ventral view of the external male
genitalia.

Figure 15. Eighth tergum of male.
Figure 16. Eighth sternum of male.

Figure 17. Ninth segment of male.

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. 3 1293 03046 1911