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Journal

of the

New York

ENTOMOLOGICAL SOCIETY

Devoted to Entomology in General

VOLUME LXXIV

Published by the Society
New York, N. Y.

ALLEN PRESS, INC.
Lawrence, Kansas

INDEX OF AUTHORS

ALEXANDER, CHARLES P. Undescribed Species of Crane Flies from the Himalaya
Mountains (Diptera: Tipulidae), XII 66

ALEXANDER, CHARLES P. Undescribed Species of Crane Flies from the Himalaya
Mountains (Diptera: Tipulidae), XIII 180

BAHADUR, J. and B. B. L. SRIVASTAVA The Nerves of the Thoracic Segments of
the Larva of Prodenia litura (Lepidoptera: Noctuidae) 168

BENNETT, FREDERICK D. Notes on the Biology of Stelis ( Odontostelis ) biline olata
(Spinola), a Parasite of Euglossa cordata (Linnaeus) (Hymenoptera: Apoidea:

Megachilidae) 72

BROWN, F. MARTIN David Bruce (1833-1903) and Other Entomological Collectors
in Colorado 126

dos PASSOS, CYRIL F. Pieris narina oleracera (Harris) in New Jersey (Lepidoptera:
Pieridae) 222

FREDRICKSON, RICHARD W. An Apparent Association of Mites (Acarina) with
the Rock Barnacle Balanus 101

GUPTA, A. P. Further Studies on the Internal Anatomy of the Meloidae (Coleoptera) .
II. The Digestive and Reproductive Systems of the S. A. Blister Beetle, Picnoseus

nitidipennis Fairmaire and Germain (Coleoptera: Meloidae) 72

HUNG, AKEY C. F. and WILLIAM L. BROWN, Jr. Structure of Gastric Apex as
a Subfamily Character of the Formicinae (Hymenoptera: Formicidae) 198

IVIE, WILTON Two North American Spiders (Araneae: Linyphiidae) 224

KLOTS, ALEXANDER B. Melanism in Connecticut Panthea fur cilia (Packard) (Lep-
idoptera: Noctuidae) 95

KLOTS, ALEXANDER B. Life History Notes on Lagoa laceyi (Barnes and McDun-
nough) (Lepidoptera: Maegalpygidae) 140

KLOTS, ALEXANDER B. The Larva of Amblyscirtes samoset (Scudder) (Lepidop-
tera: Hesperiidae) 185

LUDWIG, DANIEL and MARGARET R. GALLAGHER Vitamin Synthesis by the
Symbionts in the Fat Body of the Cockroach, Periplaneta americana (L.) 134

MANISCHEWITZ, JACK R. Studies on Parasitic Mites of New Jersey 189

O’BRIEN, JAMES F. Origin and Structural Function of the Basal Cells of the Larval
Midgut in the Mosquito, Aedes aegypti Linnaeus 59

ROZEN, JEROME G., Jr. Taxonomic Descriptions of the Immature Stages of the
Parasitic Bee Stelis ( Odontostelis ) biline olata (Spinola) (Hymenoptera: Apoidea:

Megachilidae) 84

ROZEN, JEROME G., Jr. and BARBARA L. ROZEN Mature Larvae of the Old
World Bee Genus Panurgus (Hymenoptera: Apoidea) 92

iii

TREAT, ASHER E. A New Blattisocius (Acarina: Mesostigmata) from Noctuid

Moths 143

VASVARY, LOUIS M. Musculature and Nervous System of the Thorax, of the Sound
Mechanism, and of a Typical Pregenital Abdominal Segment of the Male of the Annual
Cicada, Tibicen chloromera (Walker) (Homoptera: Cicadidae) 2

VOGEL, BEATRICE R. Spiders from Powdermill Nature Reserve 55

WOOLLEY, TYLER A. and HAROLD G. HIGGINS Xenillidae, a New Family of
Oribatid Mites (Acari: Cryptostigmata) 201

NOTES

HOPF, ALICE L. Help for Ailing Caterpillars? 111

dos PASSOS, CYRIL F. The Discovery of Additional Journals of Frank E. Watson .... 188

BOOK REVIEWS

KLOTS, ELSIE B. Monarch Butterflies by Alice L. Hopf 64

and

Fireflies in Nature and the Laboratory by Lynn and Gray Poole 64

FREDRICKSON, RICHARD W. The Tarantula by William J. Baerg 109

ARNETT, ROSS H., Jr. The Beetles of the Pacific Northwest by Melville H. Hatch __ 109

MILLER, DAVID C. Wandering Through Winter by Edwin W. Teale 110

QUEDNAU, F. W. The Callaphidini of Canada by W. R. Richards 228

BROWN, F. M. A History of Entomology by O. E. Essig 229

VASVARY, L. M. Plant Galls and Gall Makers by E. P. Felt 230

RECENT PUBLICATIONS 58, 116, 164

PROCEEDINGS of the NEW YORK ENTOMOLOGICAL SOCIETY 117, 160

BYLAWS of the NEW YORK ENTOMOLOGICAL SOCIETY 103

MEMBERSHIP of the NEW YORK ENTOMOLOGICAL SOCIETY 112

NECROLOGY 122

gf fjT 10 0 7 3

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Vol. LXXIY

MARCH 1966

No. 1

Devoted to Entomology in General

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New York Entomological Society

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Reincorporated February 17, 1943

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The meetings of the Society are held on the first and third Tuesday of each month (except

June, July, August and September) at 8 p.m., in the American Museum of Natural

History, 79th St., & Central Park W., New York 24, N. Y.

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Members of the Society will please remit their annual dues, payable in January, to the

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Officers for the Year 1966

President, Dr. Richard Fredrickson 1 >. . f V A

College of the City of New York 1003 1(

Fzce President, Dr. Kumar Krishna a

American Museum of Natural History, New York 10024

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Secretary, Mrs. Lucy Heineman 115 Central Park West, New York 10023

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Assistant Secretary, Mrs. Albert Poelzl

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230 E. 78th Street, New York 10021

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Treasurer,* Mr. Raymond Brush

American Museum of Natural History^ New York 10024

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Assistant Treasurer, Mrs. Patricia Vaurie

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1 Year Term

Dr. Alexander Br Klots

2 Year Term

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Dr. Jerome Rozen, Jr.

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The Journal of the New York Entomological Society is published quarterly for the Society by Allen Press

Inc., 1041 New Hampshire, Lawrence, Kansas. Second class postage paid at Lawrence, Kansas.

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Journal of the

New York Entomological Society

Volume LXXIV March 31, 1966

No. 1

EDITORIAL BOARD

Editor Emeritus Harry B. Weiss

Editor Lucy W. Clausen
Columbia University College of Pharmacy
115 West 68th Street, New York, N. Y. 10023

Associate Editor James Forbes
Fordham University, New York, N.Y. 10458

Publication Committee

Dr. Pedro Wygodzinsky Dr. Asher Treat

Dr. David Miller

CONTENTS

Musculature and Nervous System of the Thorax, of the Sound Mechanism, and
of a Typical Pregenital Abdominal Segment of the Male of the Annual
Cicada, Tibicen ehloromera (Walker) (Homoptera: Cicadidae)

Louis M. Vasvary 2

Spiders from Powdermill Nature Reserve Beatrice R. Vogel 55

Origin and Structural Function of the Basal Cells of the Larval Midgut in the

Mosquito, Aedes aegypti Linnaeus James F. O’Brien 59

Recent Publications 58

Book Reviews 64

New York Entomological Society

[Vol. LXXIV

Musculature and Nervous System of the Thorax, of the Sound
Mechanism, anti of a Typical Pregenital Abdominal Segment of
the Male of the Annual Cicada, Tibicen chloromera
( W alker ) ( Homoptera : Cicadidae ) 1

Louis M. Vasvary

Rutgers — The State University, New Brunswick, N. J.

Abstract: The musculature and innervation of the thorax, sound mechanism, and the

fourth abdominal segment of the male annual cicada, Tibicen chloromera (Walker) are
described.

The ventral nerve cord consists of a subesophageal ganglion, prothoracic ganglion, and a
thoracic-abdominal ganglionic mass. There are no ganglia present in any of the abdominal
segments. The prothoracic ganglion supplies innervation to some of the muscles of the cervical
area and the muscles of the prothorax. The thoracic-abdominal ganglionic mass provides in-
nervation to the posterior tergo-sternal muscles of the prothorax, the muscles of the pro-
thorax, the muscles of the mesothorax, metathorax, and all of the abdominal segments. The
abdominal segments are innervated by lateral nerve branches which arise from a pair of
nerves that originate from the posterior portion of the thoracic-abdominal ganglionic mass
located in the mesothorax. No median nerves are visible between the subesophageal ganglion,
prothoracic ganglion, and the thoracic-abdominal ganglionic mass. The median nerves are
probably included within the interganglionic connectives.

The members of the family Cicadidae are among the largest insects classified
in the order Homoptera. Their periodic occurrences in large numbers and the
shrill “song” produced by the males have probably aroused the curiosity of man
since the beginning of time. Despite their large size and the interest they have
received by virtue of their sound-producing apparatus, cicadas have been some-
what neglected by morphologists. This study was undertaken as a contribution
to our knowledge of the musculature and innervation of the thorax, of the sound
mechanism, and of a typical pregenital abdominal segment of the male of the
annual cicada, Tibicen chloromera (Walker).

A study of the nerve patterns in insects may be approached with at least two
different objectives in mind. From a physiological or histological standpoint, a
knowledge of nerve and muscle arrangements is a necessary prerequisite for pre-
cise investigations. From a morphological standpoint, a knowledge of the
hexapod nervous system is essential in establishing nerve and muscle homologies
and thereby provide additional information on the course of phylogenetic devel-
opment. This paper is an attempt in the latter direction with the full understand-
ing that detailed investigations of many more forms are necessary in order to
establish the course of phylogenetic development.

1 Paper of the Jour. Series, N. J. Agric. Expt. Station, Rutgers-The State University of
New Jersey, Dept. Ent. and Econ. Zool.

March, 1966]

Vasvary: Morphology of Annual Cicada

Figure 1A

Fig. 1A. Lateral view of the subesophageal ganglion of the male annual cicada Tibicen
chloromera (Walker) .

The concept of an underlying homology of segmental musculature has pro-
vided important evidence on the evolution of the insect thorax and appendages.
This concept is based on the assumption that at sometime in the past history of
the Hexapoda, the abdominal somites, as leg-bearing segments, had essentially
the same structure as the primitive thoracic and gnathal segments. If we assume
that the innervation pattern as well as the musculature was homologous in each
ancestral segment then the nerve configuration manifested in insects today is
a variation of the ancestral pattern. Moreover, since the inherent purpose of
the nervous system is to transmit nerve impulses, selective pressure on the
nerve pattern would be less than on the structures innervated (Schmitt, 1959).
This assumption should not be interpreted to imply that the nervous systems of
insects have remained static in the course of phylogenetic development, but
rather that through investigations of the segmental innervation patterns of
insects and by establishing criteria of homology of nerves through the utilization
of primitive muscle groups and nerve junctions, a knowledge of the course of
the phylogenetic development of the nervous system should be possible.

Unfortunately, only a very few comparative morphological investigations
have been presented in the literature concerning the establishment of nerve
homologies in insects. The writer hopes that this paper will be a significant addi-
tion to the existing studies and serve to cultivate further interest regarding the
concept of a basic plan of segmental innervation.

New York Entomological Society

IVol. LXXIV

Fig. IB. Dorsal view of ventral muscles that cover the prothoracic ganglion and anterior
portion of the thoracic-abdominal ganglionic mass of the male annual cicada Tibicen
chloromera (Walker).

OBJECTIVES

1. Determine the musculature of the thorax of the male of the annual cicada,
Tibicen chloromera (Walker) and compare this musculature to that of
Hnechys sanguinea var. philaemata as described by Maki (1938) and to
that of Cicada (= Tibicen ) plebeia as described by Berlese (1909).

2. Describe the ventral nerve cord of Tibicen and compare its configuration
to the ventral nerve cords previously described in the family Cicadidae.

3. Determine and describe the cervicothoracic nervous system of the male of
Tibicen chloromera (Walker) and, if feasible, to establish criteria of
homology.

4. Determine the musculature of the first abdominal segment of Tibicen
which contains the sound mechanism and compare this musculature to
that described by Maki (1938) for Huechys and to that of Cicada
(= Tibicen) plebeia as described by Berlese (1909).

5. Determine the innervation of the first abdominal segment of the male of
T ibicen chloromera ( W alker ) .

6. Determine the musculature of a typical pregenital abdominal segment of

March, 1966]

Vasvary: Morphology of Annual Cicada

Table 1. Ventral muscles covering the prothoracic and thoracic-abdominal ganglia of

Tibicen chloromera (Walker).

Muscle number

Origin

Insertion

Pleural arm of
prothorax

Zygomatic with muscles 2 and 3
over the prothoracic ganglion
and the anterior portion of
thoracic-abdominal ganglionic
mass.

Anterior margin of
episternum ventral
to tergo-pleural 40

Zygomatic with muscles 1 and 3
over the prothoracic ganglion
and the anterior portion of thoracic-
abdominal ganglionic mass.

Anterior mesofurcal
arm

Zygomatic with muscles 1 and 2
over the prothoracic ganglion and
the anterior portion of thoracic-
abdominal ganglionic mass.

Tibicen and compare this musculature to that described by Maki (1938)
for Huechys.

7. Determine the innervation of a typical pregenital abdominal segment of
Tibicen and, if feasible, to establish criteria of homology.

REVIEW OF LITERATURE

The literature will be reviewed under five major headings corresponding to
their order of presentation in this paper.

1 . THE VENTRAL NERVE CORD

Comparatively little is known concerning the general nerve configuration in
the families of the order Homoptera. The principal writers reporting on the
ventral nerve cord of cicadas are: Binet (1894), Dufour ( 1833), Hilton (1939),
and Myers (1928). It may be stated that within the family Cicadidae a high
degree of specialization has taken place as far as the nervous system is con-
cerned (Myers, 1928). The chief evidence of this specialization is the fact that
all abdominal ganglia have become consolidated within the large thoracic-ab-
dominal ganglionic mass located in the mesothorax.

Binet (1894) described the subintestinal nervous system of Cicada orni. By
microscopic sections of the thoracic-abdominal ganglionic mass, Binet was able
to distinguish the abdominal ganglia by the absence of crural lobes correlated
with the absence of legs in corresponding segments (Myers, 1928).

Dufour (1833), in an earlier publication, described the ventral nerve cord in
Cicada orni as having a cephalic ganglion and two thoracic ganglia. The thoracic
ganglia are nearly fused, forming one oblong body which is covered dorsally by
a mass of muscles which occupy the lower wall of the thorax. Dufour states
that the anterior thoracic ganglion gives rise to four pairs of principal nerves,
while the posterior ganglion gives rise to six pairs of nerves. The nerve cords

New York Entomological Society

LVol. LXXIV

Fig. 2. First stage dissection showing muscles of the cervix, thorax, and first abdominal
segment and the ventral nerve cord in longitudinal section in the male annual cicada Tibicen
chloromera (Walker).

March, 1966]

Vasvary: Morphology of Annual Cicada

which innervate the abdominal segments are adherent at their origin but separate
before finally dividing in the abdominal cavity. Dufour did not describe the
subesophageal ganglion which, according to Myers (1928), may have been
mistaken for the brain.

Hilton (1939) described the central nervous system for both the immature
and adult stages of a cicada. Unfortunately, no mention is made of the species
studied. There were two ganglia in both the immature and mature cicada other
than the superesophageal ganglion. Hilton further stated that there are many
large, long nerves issuing from the caudal portion of the large thoracic-abdominal
ganglion.

Myers (1928) described the brain and ventral nerve cord of Melampsalta
sericea. The round subesophageal ganglion is connected to the first ganglionic
mass by a pair of long, stout, well-separated interganglionic connectives. The
first ganglionic mass lies largely in the prothorax. Two short, very stout inter-
ganglionic connectives join the first ganglionic mass to the second thoracic mass
which lies wholly within the mesothorax. Myers states that the second thoracic
mass is much longer than broad and displays signs of a two fold origin. However,
from the standpoint of gross anatomy, the abdominal ganglia cannot be dis-
tinguished. Nerves that innervate typical abdominal segments superficially ap-
pear to arise as a single cord as they leave the second thoracic mass. Later the
single cord splits into two nerves as it enters the abdomen.

2. THORACIC MUSCULATURE

The thoracic musculature of two species of cicadas have been described by
Berlese (1909) and Maki (1938). Berlese (1909) described in some detail
the thoracic musculature of Cicada ( — Tibicen ) plebeia . Muscles are identified
in figures by Roman or Arabic numerals while descriptions of muscle origins and
insertions are included in the text. An attempt is made to homologize the thoracic
musculature of several species of insects. Unfortunately, with respect to Cicada
(= Tibicen) plebeia , it appears many of the muscles that originate on the furcal
and pleural arms and attach to the coxae and trochantine are omitted.

Maki (1938) presents a very detailed description of the thoracic muscles of
Huechys sanguinea var. philaemata. Muscles are identified by their position and
function; however, in tables and figures, Arabic numerals are utilized for muscle
numbers. Muscle origins and insertions are described in the text.

In his study of Hemiptera, Maki presents the thoracic musculature of Eurostus
validus , Sigara substriata , Cicadella ferruginea , Macrohomotoma gladiatum , and
Huechys sanguinea var. philaemata . Maki includes in his tables the musculature
of Nezara viridula by Malouf (1933), Cicada plebeia by Berlese (1909), and
Psylla mali by Weber (1929).

Snodgrass ( 1927 and 1935), illustrates a portion of the thoracic musculature
of Tibicina (= Magicicada) septendecim as an example of indirect wing muscles.

New York Entomological Society

I Vol. LXXIV

Table 2. Prothoracic musculature of Tibicen chloromera (Walker).

Muscle Muscle 0rigin

number (or attachment)

Insertion
(or attachment)

Dorsal muscles

Median dorsal

Posterior edge of head

First phragma

Median dorsal

Dorsolaterally on middle
of tergum

First phragma

Lateral dorsal

Dorsolaterally on middle
of tergum

Anterolateral region of first
phragma

Lateral dorsal

Dorsolaterally on middle
of tergum

Anterior edge of first phragma

Anterior dorsal

Posterior edge of head

Dorsolateral midportion of
tergum

Ventral muscles

Internal ventral

Posterior tentorial arm

Sternal apophyses

External ventral

Posterior end of cervical
sclerite

Pleural arm of prothorax

Tergo-sternal muscles

Anterior intersegmental

Posterior edge of head

Ventrolateral cervical sclerite

Anterior intersegmental

Posterior edge of head

Ventrolateral cervical sclerite

Anterior intersegmental

Anterior dorsolateral region
of tergum

Posterior tentorial arm

Anterior intersegmental

Dorsolateral region of
tergum

Posterior tentorial arm

Anterior intersegmental

Anterior dorsolateral region
of tergum

Ventrolateral cervical sclerite

Anterior intersegmental

Dorsolateral region of
tergum

Base of tentorium

Posterior tergo-sternal

Anterolateral portion of
mesotergum

Pleural arm of prothorax

Tergo-pleural muscles

Anterior tergo-pleural

Dorsolateral portion of pos-
terior edge of head

Base of prothoracic pleural arm

Anterior tergo-pleural

Dorsolateral portion of pos-
terior edge of head

Base of prothoracic pleural arm

Ordinary tergo-pleural

Middle of lateral region of
tergum

Pleural arm of prothorax

Coxal muscles

Tergal promotor

Dorsolateral region of
tergum

Anterior rim of coxa

Tergal promotor

Lateral region of tergum

Apodeme of trochantin

Sternal promotor

Prof urea

Anterior basal rim of coxa

Tergal remotor

Middle dorsolateral region
of tergum

Remotor apodeme of coxa

Tergal remotor

Oblique ridge at middle of
lateral region of tergum

Remotor apodeme of coxa

Tergal remotor

Tergum external to 25

Posterior basal rim of coxa

Tergal remotor

Lateral region of tergum
beneath 26

Posterior basal rim of coxa

Sternal remotor

Profurca

| j 1

Posterior basal rim of coxa

Tergal abductor

Midportion of dorsolateral
region of tergum

Apodeme anterolateral basal
rim of coxa

Pleural abductor

Pleural arm of prothorax

Anterior basal rim of coxa

Pleural abductor

Pleural arm of prothorax

Anterior basal rim of coxa

Trochanteral muscles

Tergal depressor

Midlateral region of tergum

Depressor apodeme of trochanter

Pleural depressor

Pleural arm of prothorax

Depressor apodeme of trochanter

March, 1966]

Vasvary: Morphology of Annual Cicada

The muscles illustrated in the mesothorax are the longitudinal dorsal, oblique
dorsal, anterior tergo-sternal, and posterior tergo-sternal. The metathoracic de-
pressor muscles of the trochanter and the coxal part of the depressor muscle of
the trochanter are also included.

The above muscles are homologous to those of Cicada (= Tibicen ) plebeia
(Berlese, 1909), Huechys sanguinea var. philaemata (Maki, 1938), and Tibicen
chloromera with respect to their origins and insertions.

3. THE CERVICOTHORACIC NERVOUS SYSTEM

Detailed descriptions of the thoracic nervous system have not appeared in
the literature for any member of the family Cicadidae nor for any insect in the
order Homoptera. Moreover, the literature contains only a relatively few studies
regarding the thoracic nervous systems of insects. One reason for this lack of
information is due to the time-consuming nature and patience necessary for
such research. Therefore, the majority of nerve studies have been restricted to
anatomical facts and descriptions of nerve cord configurations. The principal
writers who have contributed detailed information on thoracic nervous systems of
insects are: Holste (1910) on Dytiscus marginalis , Johansson (1957) on

Oncopeltus fasciatus , Maki (1936) on Chauliodes formosanus, Marquardt
(1939) on Carausius morosus , Matsuda (1956) on Agulla adnixa and Blattella
germanica , Niiesch (1957) on Telea polyphemus , Pipa and Cook (1959) on
Periplaneta americana , Schmitt (1959) on Dissosteira Carolina , and Wittig
(1955) on Perla abdominalis.

Schmitt (1962 ) states that an additional reason for the lag of nerve topography
studies in insects is due to the difficulty in relating the findings on one group to
those on another group. Furthermore, Maki (1936) and Pipa and Cook (1959)
state that there exists a remarkable degree of variability in nerve distribution
patterns of different individuals of the same insect species. However, Pipa
and Cook (1959) also state that the existence of a fundamental plan in the
peripheral distribution of thoracic nerves in widely separated insects is evident.
Wittig (1955) describes the innervation pattern in the thorax of the larva and
adult of Perla abdominalis . She presents a comparison of the innervation fields
of the thoracic nerves of Perla abdominalis with those of Chauliodes formosanus
as reported by Maki (1936), Carausius morosus as reported by Marquardt
(1939), and Dytiscus marginalis as reported by Holste (1910) and establishes
the existence of nerve homologies in these widely separated insects.

Pipa and Cook (1959) state that the pattern of nerve distribution in Peri-
planeta americana essentially agrees with that found in other insects which have
been investigated. A similar indication in Periplaneta americana was made by
Nijenhuis and Dresden (1955).

Schmitt (1959) describes the cervicothoracic nervous system of Dissosteira
Carolina and presents several areas of nerve homology with respect to Chauliodes

trx

New York Entomological Society

[Vol. LXXIV

Fig. 3. Second stage dissection showing muscles of the cervix, thorax, and first abdominal
segment in longitudinal section in the male annual cicada Tibicen chloromera (Walker).

March, 1966]

Vasvary: Morphology of Annual Cicada

formosanus as reported by Maki (1936). However, he states that the ptero-
thoracic dorsal nerves pass beneath the ventral longitudinal muscles and differ
in this respect from the prothoracic dorsal nerves of Dissosteira and all the
thoracic dorsal nerves of Chauliodes. Schmitt compared the nerves of the pro-
thoracic muscles of Dissosteira with those of the pterothorax and concluded that
there is evidence of a loss of anterior prothoracic musculature as a result of the
evolution of the cervix. The cephalic muscles of the cervical sclerites and the
ventral lateral neck muscles are derived from this anterior prothoracic muscula-
ture. Schmitt also includes a comparative study of the anterior ganglionic con-
nectives of the dorsal nerves of Dissosteira , Periplaneta , and Orchelimum and
indicates that the anterior ganglionic connectives of the dorsal nerves may have
a wider distribution than in Orthoptera but are not recognizable because of
juxtaposition with the connectives of the ventral nerve cord. Schmitt describes
the median nerves and the innervation of the spiracular muscles in Dissosteira
and mentions that the transverse nerves, dorsal nerves, and the innervation of
the spiracular muscles of Chauliodes as described by Maki (1936) present a
pattern identical with that in Dissosteira. There appears to be no essential dif-
ferences in the innervation pattern of the thoracic spiracles as compared with
the innervation pattern of the abdominal spiracles in both Chauliodes and
Dissosteira. Schmitt concludes that the nerves to the thoracic spiracles agree
sufficiently with the nerve pattern of the abdominal spiracles to indicate that
the thoracic spiracles may be homologous with the abdominal spiracles.

Schmitt (1962), in a later paper, despite unfortunate differences in nomen-
clature applied by different workers, presents additional information establishing
the presence of nerve homologies in several insects. Schmitt utilizes the dorsal
longitudinal muscles as a starting point since these muscles are homologous both
in the thorax and abdomen of insects. Usually, from a descriptive standpoint it
is quite simple to identify the dorsal nerves to these muscles. Schmitt arranges in
tabular form the names and designations used by various authors for the nerves
to the thoracic dorsal longitudinal muscles, designations of the anterior ganglionic
connectives, designations of the subesophageal nerves to the protergal muscles,
and a comparison of thoracic nerve designations used by various authors with
those utilized by Maki for Chauliodes. The wing nerves, median and transverse
nerves, innervation of the ventral muscles and spinosternal musculature, and a
discussion of the prothoracic nervous system in various insects is also presented.

4. THE MUSCULATURE AND INNERVATION OF THE SOUND MECHANISM

The majority of investigations appearing in the literature concerning the
sound mechanism of cicadas describes the construction of the sound apparatus
and the mechanics of sound production. Myers (1928) presents a summary of
the studies pertaining to the sound-producing apparatus as well as including his

New York Entomological Society

[Vol. LXXIV

Table 3. Mesothoracic musculature of Tibicen chloromera (Walker).

Muscle

Origin

Insertion

number

(or attachment)

Dorsal muscles

Median dorsal

Anterior median portion of
tergum

Median area of second phragma

Lateral dorsal

Middle of dorsolateral por-
tion of tergum

Lateral portion of second
phragma

Ventral muscles

Longitudinal ventral

Profurcal arm

xMiterior mesofurcal arm

Tergo-Sternal muscles

Anterior tergo-sternal

Anterior portion of dorso-
lateral region of tergum

Ventrolateral sternal region

Posterior tergo-sternal

Ventral portion of second
phragma

Posterior mesofurcal arm

Tergo-Pleural muscles

Tergo-pleural

Anterolateral margin of
tergum

Anterior margin of episternum

Tergo-pleural

Lateral margin of tergum

Anterior margin of episternum

Tergo-pleural

Lateral margin of tergum

Mesothoracic pleural arm

Tergo-pleural

Lateral margin of tergum

Prothoracic pleural arm

Tergo-pleural

Lateral margin of tergum

Wing process

Tergo-pleural

Lateral margin of tergum

Base of mesothoracic pleural arm

Pleural-axillary

Episternum

Third axillary sclerite

Pleural-axillary

Episternum

Third axillary sclerite

Pleuro-subalar

Posterior margin of
epimeron

Subalar sclerite

Sterno-Pleural muscles

Sterno-basalar

Anterodorsal portion of epi-
sternum

Ventrolateral sternal region

Furco-entopleural

Furcal arm of mesothorax

Pleural arm of mesothorax

Coxal muscles

Tergal promotor

Anterolateral region of
tergum

Trochantin

Trochantino-basalar

Laterodorsal margin of epi-
sternum

Trochantin

Trochantino-basalar

Anterolateral margin of epi-
sternum

Trochantin

Sternal promotor

Base of mesofurcal arm

Anterior basal rim of coxa

Tergal remotor

Anterolateral region of
tergum

Posterior basal rim of coxa

Tergal remotor

Posterior dorsolateral region
of tergum

By a tendon to posterior basal
rim of coxa

Coxo-subalar

Posterior basal rim of coxa

Subalar sclerite

Sternal remotor

Posterior mesofurcal arm

Posterior basal rim of coxa

Sternal remotor

Mesofurcal arm

Posterior basal rim of coxa

Sternal adductor

Mesofurca

Mesal basal edge of coxa

Coxo-basalar

Dorsal margin of episternum Anterolateral basal rim of coxa

Troehanteral muscles

Tergal depressor

Anterolateral portion of
tergum

Depressor apodeme of trochanter

Trochantero-basalar

Dorsal margin of episternum

Depressor apodeme of trochanter

Sternal depressor

Mesofurcal arm

Depressor apodeme of trochanter

Muscles of the spiracle

Occlusor

Subspiracularum

Ventral portion of atrial chamber

March, 19661

Vasvary: Morphology of Annual Cicada

own findings based on M damp salt a sericea and M damp salt a muta , two species
of cicadas found in New Zealand.

Complete studies regarding the musculature of the first abdominal segment
which contains the sound-producing apparatus have been described for Cicada
(— Tibicen) plebeia by Berlese (1909) and for Huechys sanguinea var. philae-
mata by Maki (1938). Berlese utilizes both Roman and Arabic numerals for
muscle identification in his figures while descriptions of muscle attachments are
included in the text. Berlese (1909) shows the structure of the sound mechanism
in his figures 879 to 882. Berlese considers the sclerotized V-shaped structure,
yAd2 in his figure 880, as the furca of the second abdominal sternite. However,
Carlet (1876), Vogel (1923) and Myers (1928) who have given this structure
the most attention, ascribe it to the first abdominal segment. Maki (1938)
shows the musculature of the sound mechanism in his figure 24 and utilizes
Arabic numerals for muscle numbers. Maki presents in tabular form the muscles
of the first six abdominal segments with their muscle numbers. Descriptions
of the muscle attachments are not included in the text.

A complete presentation of the innervation pattern of the first abdominal seg-
ment of cicadas has not appeared in the literature. However, the auditory or
tymbal nerves which innervate the large tymbal muscles have been mentioned by
various writers since Binet (1894). Swinton (1880) traced the auditory nerve
from the thoracic ganglionic mass, presumably in the mesothorax, to the abdomen
and around the tymbal muscle. The auditory nerve then forms a ganglion which
enters a groove. According to Vogel (1923) the auditory nerve arises in the
ventral nerve strands and rises, running parallel with the body wall, in a
sclerotized groove and passes dorsally to the sense organ, where its fibers run
into the base of each sense cell. Myers (1928), in poorly preserved material,
found a distinct nerve emerging on each side of the last thoracic-ganglionic mass
and running parallel to a sclerotized ridge leading up to the auditory capsule.
Myers (1928) states that it is very improbable that the auditory nerve should
arise from the abdominal strands, as Vogel ( 1923) states.

Investigations utilizing electric stimulation of the auditory or tymbal nerve
and the sympathetic nerve have appeared in the literature. Pringle (1954)
concluded that the frequency of tymbal movements resulting from the contrac-
tions of the tymbal muscle exceeds the rhythm of tymbal nerve stimulation.
Pringle also reported that an isolated tymbal muscle does not give multiplied
rhythmic reactions when stimulated but functions the same as a common skeletal
muscle. Hagiwara and Watanabe (1956) found that at a certain intensity and
frequency of nerve stimulation, repetitive potentials up to ten or more resulted
from each stimulus in the tymbal muscle, tymbal nerve, and motor neuron.
Voskresenskaya and Svidersky (1960) investigated the electrical activity of
the tymbal muscle, the tymbal nerve, and the sympathetic nerve during and

New York Entomological Society

[Vol. LXXIV

Fig. 4. Third stage dissection showing muscles of cervix, thorax, and first abdominal seg-
ment in longitudinal section in the male annual cicada Tibicen chloromera (Walker).

March, 1966]

Vasvary: Morphology or Annual Cicada

after electric stimulation and concluded that the sympathetic nervous system is
essential to normal sound production in cicadas.

5. THE MUSCULATURE AND INNERVATION OF THE FOURTH ABDOMINAL SEGMENT

The musculature of the pregenital abdominal segments of male cicadas have
been described by Maki (1938) for Huechys sanguined var. philaemata and by
Berlese (1909) for Cicada (= Tibicen) plebeia. Maki in his figure 24 shows the
musculature of the first three abdominal segments and utilizes Arabic numerals
for muscle numbers. Maki presents the muscles of the first six abdominal seg-
ments and their muscle numbers in a table on page 168 where he compares the
musculature of Erostus validus , Sigara sub striata , Huechys sanguined var.
philaemata, Cicadella ferruginea, and Macrohomotoma gladiatum. Maki does
not describe the muscle attachments for Huechys in his text; however, they are
clearly shown in his figure 24. Berlese (1909) describes the musculature of the
first three abdominal segments in Cicada (= Tibicen) plebeia and utilizes both
Roman and Arabic numerals for muscle identification. Descriptions of the
muscle attachments are included in the text.

No studies dealing with the innervation of a pregenital abdominal segment of
a male cicada have been found in the literature. Moreover, the literature con-
tains only a few studies on the abdominal nervous system of insects.

In recent years some interest has been shown regarding the establishment of
basic segmental nerve pattern within the Hexapoda. Schmitt ( 1954) describes
the nervous system of the pregenital abdominal segments of Dissosteira Carolina ,
Acheta assimilis, Periplaneta americana , and Diapheromera femorata. Schmitt
utilizes various points of nerve homology or “landmarks” in presenting the in-
nervation pattern of the above insects. The innervation of the ventral diaphragm
in Dissosteira is also described. Libby (1959) describes the musculature and
innervation of the second and third abdominal segments of the cecropia larva
and concludes that the dorsal, ventral, and transverse nerve roots arising from
each segmental ganglion of the cecropia larva seem homologous with those de-
scribed by Schmitt (1954) for the pregenital segments of certain Orthoptera.
Libby concludes, by utilizing the points of nerve homology set forth by Schmitt,
that the homogeneity of the innervation pattern in such widely separated orders
as Orthoptera and Lepidoptera lend further support to the concept of a basic
segmental nerve pattern within the Hexapoda. Libby (1961) describes the mus-
culature and innervation in the fourth abdominal segment of the adult male
cecropia moth Hyalophora cecropia and compares his finding with the pregenital
abdominal segments of Chauliodes formosanus , as described by Maki (1936),
Acheta assimilis , as described by Schmitt (1954), and the larva of Hyalophora ,
as described by Libby ( 1959).

Schmitt (1963) describes the abdominal nervous system in the nymph of
Pteronarcys proteus and the adult of Pteronarcys calif ornica and concludes that

New York Entomological Society

[Vol. LXXIV

Fig. 5. Fourth stage dissection showing muscles of the cervix, thorax, and first abdominal
segment in longitudinal section in the male annual cicada Tibicen chloromera (Walker).

March, 19661

Vasvary: Morphology or Annual Cicada

the ganglia of segments 3 and 4 have coalesced and only the first three segments
contain both dorsal and ventral nerves. The transverse nerves of segments 4,
5, and 6 arise from the ganglia of the immediately following segments. No
occlusor or dilator muscles of the spiracles could be found in the two above-men-
tioned species of Pteronarcys. Schmitt also describes the muscles and nerves of
the genital segments.

Schmitt (1964) describes the nerve pattern of the pregenital abdominal seg-
ments of N eoconocephaiis exiliscanorus and Cento philus gracilipes gracilipes,
two Orthoptera classified in the family Tettigoniidae. The segmental nerve
patterns of these two insects were comparable and conformed to the patterns
described in the Acrididae, the Gryllidae, and the Blattidae, as described by
Schmitt (1954), and in Carausius (Phasmidae) as described by Marquardt
(1939). Similarities in the nerve patterns to Hyalophora cecropia as described
by Libby (1959 and 1961) and by Beckel (1958) and in some degree to the
Plecoptera and the Megaloptera were noted. No innervation to the alary muscles
could be found in N eoconocephalus or Ceutophilus.

Schmitt (1965) presents a comparative study on the transverse nerves of
the pregenital abdominal segments of insects. By comparing the segmental
innervation patterns of Peri planet a, N eoconocephalus, Hyalophora, Chauliodes ,
Pteronarycs, Acroneurai, Apis, and Tibicen, Schmitt concludes that, in those
insects which apparently lack median and transverse nerves, these nerves are in-
corporated in the longitudinal connectives and lateral segmental nerves.

MATERIALS AND METHODS

Insect Material Used in the This Study. — The male of the annual cicada, Tibicen
chloromera (Walker), was selected for this study in order to provide information
concerning the musculature and nervous system of the thorax, sound mechanism,
and a typical pregenital abdominal segment. The annual cicada’s large size and
ready availability make them especially attractive subjects for such investigation.
N omenclature. — Nomenclature used in this study involve primarily the muscula-
ture and nervous system. Various methods of nomenclature have been devised
for each of these organ systems.

Nomenclature of the musculature is based on the general outline set forth by
Maki (1938) in his work on Huechys sanguinea var. philaemata. Muscles are
named according to their position, attachment, or function and are assigned
Arabic numerals which serve as muscle numbers in figures.

Effective nerve nomenclature requires not only that it describe the nerves in
question, but also that it can be applied or adapted to as many nervous systems
as possible in order to demonstrate nerve homologies. However, before a stan-
dard terminology can be devised, it is essential to have a relatively thorough
knowledge of the musculature and nervous systems of manj^ different insect

New York Entomological Society

[Vol. LXXIV

Table 4. Metathoracic Musculature of Tibicen chloromera (Walker)

| Muscle Origin

Number (or attachment)

Insertion
(or attachment)

Dorsal muscles

Median dorsal

Dorsal portion of second
phragma

Dorsal portion of third phragma

Ventral muscles

Longitudinal ventral

Posterior mesothoracic
furcal arm

Metafurcal arm

Tergo-Sternal muscles

Anterior tergo-sternal

Anterior dorsolateral region
of tergum

Ventrolateral sternal region

Posterior tergo-sternal

Anterolateral edge of first
abdominal tergum

Metafurcal arm

Tergo-Pleural muscles

Tergo-pleural

Lateral portion of tergum

Pleural arm of metathorax

Tergo-pleural

Lateral portion of tergum

Dorsal border of episternum

Pleuro-axillary

Pleural ridge

Third axillary sclerite

Pleuro-axillary

Pleural ridge

Third axillary sclerite

Sterno-Pleural muscles

Sterno-pleural

Mesofurcal arm

Anterior end of metathoracic
episternum

Furco-entopleural

Metafurcal arm

Pleural arm of metathorax

Coxal muscles

Tergal promotor

Anterior dorsolateral region
of tergum

Trochantin

Pleural promotor

Anterior portion of epi-
sternum

Anterior basal rim of coxa

Sternal promotor

Metafurcal arm

Anterior basal rim of coxa

Tergal remotor

Mid dorsolateral region of
tergum

Posterior basal rim of coxa

Tergal remotor

Posterior dorsolateral region
of tergum

Posterior basal rim of coxa by a
tendon

Coxo-subalar

Lateral basal rim of coxa

Subalare

Sternal remotor

Metafurcal arm

Posterior basal rim of coxa

Sternal remotor

Metafurcal arm

Posterior basal rim of coxa

Pleural abductor

Anterior region of epi-
sternum lateral to 75

Anterolateral basal rim of coxa

Trochanteral muscles

Tergal depressor

Second phragma

Depressor apodeme of trochanter

Tergal depressor

Anterior portion of dorso-
lateral region of tergum

Depressor apodeme of trochanter

Pleural depressor

Episternum

Depressor apodeme of trochanter

Sternal depressor

Metafurcal arm

Depressor apodeme of trochanter

Muscles of the spiracle

Occlusor

Ridge between mesothorax
and metathorax

Ventral end of spiracle

species. Several systematic methods of nerve terminology have been devised and
each have their advantages and disadvantages.

The method of nerve designation utilized in this paper is similar to that used
by Whittig (1955) in her work on Perla abdominalis Burm. Ganglia, except for
the subesophageal ganglion, are assigned Roman numerals. Nerve roots arising
from each ganglion are designated by the Roman numeral of the ganglion followed

March, 1966]

Vasvary: Morphology op Annual Cicada

by the letter N and an Arabic numeral. Lower case letters following Arabic
numerals are used to identify nerve branches. Prime (') and double prime (")
designations are utilized where it appears necessary for better understanding of
nerve branch description.

Methods of Illustration. — Illustrations in this paper representing nerves and
muscles are of two types. One type, the semiperspective illustration, is an at-
tempt to represent as clearly as possible the various stages of dissection. Each
stage is illustrated separately and in series beginning with the median muscle
groups and progressing to the body wall. In illustrations that combine two
consecutive stages of dissection, the lower half of the figure represents the earlier
stage.

The second type of illustration used in this study are diagrams indicating the
spatial relationships of nerves. The right side of the insect is illustrated and
viewed in a laterad aspect. Muscle innervations are designated by Arabic nu-
merals which represent muscle numbers. Where two nerves cross, the laterad
nerve is interrupted. Nerves which terminate in the integument are indicated by
a short line drawn across the nerve.

An explanation of abbreviated designations may be found under “Abbrevia-
tions used in the Figures” at the conclusion of this paper.

RESULTS AND DISCUSSION
1. THE VENTRAL NERVE CORD

General: The ventral nerve cord of insects is the postcephalic portion of the
nervous system which lies beneath the alimentary canal and extends posteriorly
through the thorax and abdomen. This portion of the central nervous system
contains the subesophageal ganglion, thoracic ganglia, and abdominal ganglia
arranged metamerically and joined by paired longitudinal connectives. However,
modifications of the above generalized ventral nerve cord exists in a number of
insect orders and is evidenced by the reduction in number or complete absence
of ganglia in abdominal segments. Snodgrass (1935) states that there is a
tendency for the ganglia of the ventral nerve cord to migrate anteriorly and
unite with each other. This process is referred to as condensation. The forward
migration and fusion of ganglia results in the shortening and external disap-
pearance of connectives and commissures.

A dorsal view of the ventral nerve cord in the male cicada, Tibicen chloromera
(Walker) is illustrated in Fig. 9 and consists of a subesophageal ganglion, pro-
thoracic ganglion, and a thoracic-abdominal ganglionic mass. There are no
ganglia in any of the abdominal segments. All abdominal segments are innervated
by nerves originating from the posterior portion of the thoracic-abdominal
ganglionic mass located in the mesothorax.

The subesophageal ganglion is the anterior ganglion of the ventral nerve cord.

New York Entomological Society

[Vol. LXXIV

Table 5. Comparison of prothoracic musculature of Tibicen chloromera , Huechys
sanguined var. philaemata (Maki, 1938), and Cicada (— Tibicen) plebeia (Berlese, 1909).

Muscle groups

Tibicen

chloromera

Huechys sanguined
var. philaemata
(Maki, 1938)

Cicada (= Tibicen)
plebeia

(Berlese, 1909)

Dorsal muscles

Median dorsal

140

Median dorsal

CIX

Lateral dorsal

110

Lateral dorsal

—

CXII

Anterior dorsal

CXXXVI

Anterior dorsal

cxxxv

Ventral muscles

Internal ventral

136

External ventral

CXXXI

Tergo-Sternal muscles

Anterior intersegmental

147

Anterior internal tergo-sternal

—

Anterior internal tergo-sternal

CXXXV

Anterior internal tergo-sternal

—

Anterior internal tergo-sternal

CXXXVa

Anterior internal tergo-sternal

—

144

Anterior internal tergo-sternal

—

Posterior tergo-sternal

112

Tergo-Pleural muscles

Anterior tergo-pleural

Ordinary tergo-pleural

Coxal muscles

Tergal promotor

113

Tergal promotor

Sternal promotor

Tergal remotor

116

Tergal remotor

—

Sternal remotor

—

Tergal abductor

Pleural abductor

—

Pleural abductor

Troehanteral muscles

Tergal depressor

115

Pleural depressor

—

In Tibicen chloromera eight pairs of nerves arise from the ganglion and innervate
the salivary glands and lateral salivary gland ducts, muscles associated with'
the feeding apparatus, and some muscles of the cervical area.

The prothoracic ganglion and the anterior portion of the thoracic-abdominal
ganglionic mass are covered dorsally by ventral muscles (Fig. IB). Dufour
( 1833) mentions similar ventral muscles in Cicada orni.

An invagination of the first abdominal sternite serves as a muscle attachment
for the large tympanal muscles. A sternal canal is located within this invagina-
tion. Two pairs of nerves, IIN8 and IIN9, pass through the sternal canal.

March, 19661

Vasvary: Morphology of Annual Cicada

Fig. 6. Fifth stage dissection showing muscles of the cervix, thorax, and first abdominal
segment in longitudinal section in the male annual cicada Tibicen chloromera (Walker).

New York Entomological Society

[ Vol. LXXIV

One pair of nerves, IIN8, innervates the posterior muscles of the first abdominal
segment while the other pair of nerves, IIN9, innervates the remaining abdominal
segments.

No median nerve is visible between the subesophageal ganglion, prothoracic
ganglion, and thoracic-abdominal ganglionic mass. However, the median nerve
is probably included within the interganglionic connectives.

Spiracular muscles in the thoracic segments are innervated by nerves which
arise from the dorsolateral portion of the prothoracic ganglion and thoracic-
abdominal ganglionic mass. Spiracular muscles in pregenital abdominal seg-
ments are innervated by a nerve branch from the dorsal nerve.

The ventral nerve cord of the male Tibicen chloromera (Walker) is not re-
stricted to a definitive positional relationship in the thorax by spinae or muscles
that attach to these structures. Schmitt (1959) described an opposite situation
in the thorax of Dissosteira , where possible future evolution of the ventral nerve
cord towards condensation will require drastic skeletal and muscle system
changes.

Subesophageal Ganglion: The anterior portion of the subesophageal gan-
glion is covered by the tentorial bridge (TB, Fig. 9). A pair of short, stout cir-
cumesophageal connectives link the subesophageal ganglion to the brain.

A lateral view of the subesophageal ganglion is shown in Fig. 1A. Eight pairs
of nerves arise from the ganglion, five pairs of nerves from the lateroventral
surface, and three pairs from the ventral area.

The first pair of nerves, SN1, arise from the anterior medioventral surface of
the ganglion in close association with the ventral portions of the circumesopha-
geal connectives. SN1 nerves divide into labral nerves (LmNv) and nerves which
innervate the protractor muscles of the mandibular bristles (pmdb).

The second pair of nerves, SN2, are mandibular nerves and arise from the
anterior lateroventral surface of the ganglion. The SN2 nerve divides soon
after leaving the ganglion into a dorsal branch that innervates the retractor
muscle of the mandibular bristle (rmdb) and a ventral branch that innervates the
protractor muscles of the mandibular bristles.

The third pair of nerves, SN3, are maxillary nerves and arise from the latero-
ventral surface of the ganglion. The SN3 nerve bifurcates into anterior and
posterior nerve branches. The anterior branches innervate the internal (rmxbi)
and the external (rmxb2) retractor muscles of the maxillary bristles. Posterior
nerve branches innervate both internal and external retractor muscles of the
maxillary bristles, protractor muscles of the maxillary bristles (lpmxb and
2pmxb), and provide nerve branches which enter the base of the maxillary
bristles, mxb (Fig. 1A).

The fourth, SN4, and fifth, SN5, pairs of nerves arise from the mediolateral
and posterolateral areas, respectively, of the subesophageal ganglion and co-

March, 1966]

Vasvary: Morphology of Annual Cicada

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alesce to form the first cervical nerve SN4 + SN5. The SN4 + SN5 nerves ex-
tend dorsally and innervate the anterior internal tergo-sternal muscle 13.

The sixth pair of nerves, SN6, arises from the posterior lateroventral surface
of the ganglion and innervates the salivary glands (S1G1) and the anterior in-
ternal tergo-sternal muscle 16. The latter nerves are the second cervical nerves.
A nerve branch from SN6, and SN6a, proceeds in a posterior direction and
coalesces with the SN9 + INI nerve.

The seventh pair of nerves, SN7, are the labial nerves and arise from the
posterior medioventral surface of the ganglion ventrad to SN6. Nerve SN7 pro-

New York Entomological Society

[Vol. LXXIV

vides nerve branches to the dilator muscles of the salivary syringe (dlSyr) and
the lateral muscles of the sclerotized rod (mr) before entering the labium.
Johansson ( 1957) shows a similar innervation pattern for the labial nerve in
the milkweed bug One o pelt us fasciatus (Dallas). The labial nerves innervate
the dilator muscles of the salivary syringe before entering the base of the
labium.

The eighth pair of nerves, SN8, arises from the posterior medioventral sur-
faces of the ganglion in close association with the interganglionic connectives and
innervates the salivary ducts (SID).

A pair of large nerves, SN9, arises laterally from each interganglionic connec-
tive and coalesce with the IN 1 nerves which arise from the anterior surface of the
prothoracic ganglion. INI nerves can easily be separated from the intergangli-
onic connectives to their origin on the prothoracic ganglion.

A pair of long, sturdy, well-separated interganglionic connectives link the
subesophageal ganglion to the prothoracic ganglion. The interganglionic con-
nectives pass laterally around the muscles of the sclerotized rod. The sclerotized
rod is an extension of the labium and appears to have a stronger association with
the prothorax than with the head since it hangs freely from the cervical mem-
brane.

Prothoracic Ganglion: The prothoracic ganglion lies wholly within the pro-
thorax and is situated between the sternal apophyses. Three pairs of ventral
muscles (1, 2, and 3) cover the entire ganglion dorsally (Fig. IB).

Four pairs of nerves arise from the anterior portion of the prothoracic ganglion
(Fig. 9). The anterior pair of nerves (INI) proceed anteriorly and join nerves
SN9 which branch from the interganglionic connectives. Nerves IN2, IN3, and
IN4 pass under the internal ventral longitudinal muscles (9) and innervate
muscles of the cervix, prothorax, and prothoracic leg muscles. (A detailed de-
scription of the innervation pattern is presented under the section entitled “The
Cervicothoracic Nervous System.”)

A pair of large nerves, IIN1, issues from the interganglionic connectives be-
tween the prothoracic ganglion and thoracic-abdominal ganglionic mass (Fig.
9). The IIN1 nerves pass over the IIN2 nerves originating from the thoracic-
abdominal ganglionic mass and then pass under the longitudinal ventral muscles
36 of the mesothorax. Nerve IINl innervates the longitudinal ventral muscles
36, median dorsal muscles 34, and lateral dorsal muscles 35 of the mesothorax.

A pair of fine, short nerves, IN5, arise on each side of the middorsal portion
of the prothoracic ganglion and innervates the ventral muscles 1 which cover the
ganglion.

Two pairs of fine nerves, IN6 and IN 7, arise from the middorsal area of the
prothoracic ganglion and coalesce with nerve INS arising from the dorsal surface
of the interganglionic connective between the prothoracic ganglion and thoracic-
abdominal ganglionic mass.

March, 1966 1

Vasvary: Morphology of Annual Cicada

A pair of very short, stout interganglionic connectives links the prothoracic
ganglion to the large thoracic-abdominal ganglionic mass.

Thoracic— Abdominal Ganglionic Mass: The thoracic-abdominal gangli-

onic mass is the terminal ganglion of the ventral nerve cord and is located above
the basisternum of the mesothorax. With the exception of the XIN2a nerves
which innervate the posterior tergo-sternal muscles 17 of the prothorax, nerves
originating from the thoracic-abdominal ganglionic mass innervate muscles of
the mesothorax, metathorax, sound mechanism, and abdominal segments.

Eight pairs of lateral nerve roots arise from the thoracic-abdominal ganglionic
mass: one pair anteriorly, IIN2; two pairs laterally, TIN3 and IIN4; and five
pairs posteriorly, IIN5, 1 1X6. IIN7, IIN8, and TIN9 (Fig. 9). Nerves IIN2,
IIN3, and IIN4 pass under the longitudinal ventral muscles 36 while the remain-
ing nerve roots extend posteriorly and pass over the mesofurca. IIN2 is the
anterior wing nerve while IIN3 and IIN4 innervate muscles in the mesothorax.
Nerves IIN5 and IIN6 pass under the posterior arms of the mesofurca and inner-
vate muscles of the metathoracic segment with the exception of nerve branch
IIN6a' which innervates the posterior tergo-pleural muscle 38 of the mesothorax.
The 1 1 N 5 nerve is the dorsal nerve since it innervates the dorsal muscles 65.
Ventral muscles are innervated by a nerve branch from IIN10 + IIN11. Nerve
IIN6 provides a nerve branch IIN6a which is the posterior wing nerve. Nerves
IIN7 supply innervation to the muscles located in the anterior portion of the
first abdominal segment and the membrane forming the large abdominal air
chamber. Nerves IIN8 provide nerve branches IIN8a to the large tympanal
muscles before passing through the sternal canal to innervate the muscles located
in the posterior portion of the first abdominal segment. The IIN9 nerves pass
through the sternal canal and innervate muscles of the remaining pregenital
abdominal segments by providing a lateral nerve branch to each consecutive
segment. Two pairs of fine nerves (IIN10 and IIN11) arise dorsolaterally from
the thoracic-abdominal ganglionic mass (Fig. 9). The IIN11 nerve divides
soon after leaving the ganglion and provides a fine nerve branch IINlla which
coalesces with nerve IIN5. Nerves IIN10 and XIN11 are connected by a fine
nerve designated as XIN10 + IIN11. It appears that both the IIN10 and IIN11
nerves are responsible for innervation of the occlusor muscle (88) of the meta-
thoracic spiracle and ventral muscle 3.

Discussion: Unfortunately, only the gross anatomy of the central nervous sys-
tem of cicadas has been described in the literature. Therefore, comparisons of
ventral nerve cords in order to establish areas of homology are limited to their
general configuration.

Hilton (1939) in his Figure 190-1 presents an unlabeled drawing of the cen-
tral nervous system of an unnamed adult cicada showing the brain and two gan-
glia of the ventral nerve cord. If it is assumed that the anterior ganglion is the

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rVoL. LXXIV

Table 6. Comparison of mesothoracic musculature of Tibicen chloromera, Huechys san-
guined var. philaemata (Maki, 1938), and Cicada {— Tibicen ) plebeia (Berlese, 1909).

Muscle groups

Tibicen

chloromera

Huechys sanguined
var. philaemata
(Maki, 1938)

Cicada (— Tibicen )
plebeia

(Berlese, 1909)

Dorsal muscles

Median dorsal

—

Lateral dorsal

Ventral muscles

Longitudinal ventral

105 + 106

Spino-furcal ventrals

104

Tergo-Sternal muscles

Anterior tergo-sternal

LXXVIII

Posterior tergo-sternal

Tergo-PIeural muscles

Tergo-pleural

XCI

Tergo -pleural

Tergo-pleural

Pleuro-axillary

XC1II

Pleuro-axillary

XCII

Pleuro-subalar

Sterno-Pleural muscles

Sterno-basalar

Furco-entopleural

100

Coxal muscles

Tergal promotor

74?

Trochantino-basalar

79 + SO

Trochantino-basalar

Sternal promotor

Tergal remotor

LXXXII

Tergal remotor

Coxo-subalar

Sternal remotor

Sternal adductor

Pleural abductor

Coxo-basalar

Trocha literal muscles

Tergal depressor

Trochantero-basalar

Sternal depressor

Muscles of the spiracle

Occlusor

—

subesophageal ganglion, then the remaining ganglionic mass contains all of
the thoracic and abdominal ganglia.

Dufour (1833), describing the ventral nerve cord in Cicada orni , states that
the central nervous system consists of a cephalic ganglion and two thoracic
ganglia. No mention is made of the subesophageal ganglion, which, according
to Myers (1928), Dufour may have confused with the brain. Dufour does

March, 1966]

Vasvary: Morphology of Annual Cicada

mention that the cephalic ganglion is produced by a fusion of two hemispheroid
lobes and the cleft which separates the two lobes is only superficial. Dufour
continues by describing the thoracic ganglia as not being separate and distinct
but nearly fused into one. However, with difficulty, a light demarcation of an
anterior ganglion can be observed.

Myers (1928) states that the ventral nerve cord in Melampsalta sericea
consists of a subesophageal ganglion, prothoracic ganglion, and thoracic-ab-
dominal ganglionic mass, each separated by visible interganglionic connectives.

Berlese (1909), in his Figure 697, presents a diagram of the brain and the
subesophageal ganglion of Cicada (= Tibicen) plebeia , and shows that the
subesophageal ganglion is separated from the brain by a pair of stout circum-
esophageal connectives. The remainder of the ventral nerve cord is not described.

Snodgrass (1935), in his Figure 237, presents a longitudinal section of
Tibicina (— Magicicada) septendecim showing two thoracic ganglia, one in the
prothorax and the other in the mesothorax. The subesophageal ganglion is
not illustrated.

If the above investigations are correct, then there appears to be some diversity
in the family Cicadidae regarding the number of ganglia in the ventral nerve
cord. Cicada orni is the most specialized with a central nervous system composed
of a cephalic ganglion and two very closely associated thoracic ganglia while in
Melampsalta sericea and Tibicen chloromera there is a subesophageal ganglion
and two separate thoracic ganglia.

There also appears to be a diversity in the number of principal lateral nerve
roots arising from the thoracic ganglia. Dufour (1833) mentions that the an-
terior thoracic ganglion in Cicada orni gives rise to four pairs of principal nerves
while six pairs of nerves issue from the posterior thoracic ganglion. Hilton
(1939), in his Figure 190-1, of the central nervous systems of an unnamed
species of cicada, shows three principal nerves arising from the anterior lobe
of the thoracic-abdominal ganglionic mass while the posterior lobe possesses
three pairs of lateral nerves and a single caudal nerve.

Tibicen chloromera has three pairs of principal lateral nerve roots (not count-
ing the INI nerve which adheres to the interganglionic connective) arising from
the prothoracic ganglion. One nerve, IIN1, appears to arise from the intergan-
glionic connective between the prothoracic ganglion and thoracic-abdominal
ganglionic mass, and eight principal pairs of nerves arise from the thoracic-ab-
dominal ganglionic mass.

2. THORACIC MUSCULATURE

General: The thoracic musculature of the male cicada, Tibicen chloromera
(Walker) is illustrated in Figs. IB to 8. Figs. 2 to 8 represent stage dissections
which proceed from the interior muscle groups to the exterior muscle groups on
the body wall. Arabic numerals are utilized for muscle numbers. Thoracic

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| V ol. LXXIV

Table 7. Comparison of metathoracic musculature of Tibicen chloromera, Huechys san-
guined var. philaemata (Maki, 1938), and Cicada {— Tibicen) plebeia (Berlese, 1909).

Muscle groups

Tibicen

chloromera

Huechys sanguined
var. philaemata
(Maki, 1938)

Cicada (= Tibicen)
plebeia

(Berlese, 1909)

Dorsal muscles

Median dorsal

Ventral muscles

Longitudinal ventral

Tergo-Sterual muscles

Anterior tergo-sternal

XXXVI

Posterior tergo-sternal

XXXVII

Tergo-Pleural muscles

Tergo-pleural

Pleuro-axillarv

Pleuro-axillary

Sterno-Pleural muscles

Sterno-pleural

Furco-entopleural

Coxal muscles

Tergal promotor

42?

Pleural promotor

48 + 49

Sternal promotor

Tergal remotor

Coxo-subalar

XLIX

Sternal remotor

—

Pleural abductor

Trochauteral muscles

Tergal depressor

XLV

Tergal depressor

Pleural depressor

Sternal depressor

Muscles of the spiracle

Occlusor

—

muscles are listed with their muscle numbers, origins, and insertions in Tables 1
to 4. A comparison of the thoracic musculature of Tibicen chloromera , Huechys
sanguinea var. philaemata described by Maki ( 1938) and Cicada (= Tibicen)
plebeia described by Berlese (1909) is presented in Tables 5 to 7.

Ventral Muscles Which Cover the Thoracic Ganglia: The prothoracic

ganglion and the anterior portion of the thoracic-abdominal ganglionic mass
are covered dorsally by three pairs of muscles (Fig. IB). The muscle numbers,
origins, and insertions of the three muscles groups are described in Table 1.
Ventral muscles 1,2, and 3 are mutually joined by zygomatic connections. Mus-
cles 1 and 3 are quite sturdy, while muscle 2 is broad at its zygomatic junction
and compressed dorsoventrally. Muscle 3 is joined laterally to ventral longi-
tudinal muscle 36 for a portion of its length.

Prothoracic Musculature: The prothoracic muscles in Tibicen chloromera
are fundamentally homologous to Huechys sanguined var. philaemata (Table 5).
The lateral dorsal 7, anterior internal tergo-sternals 12 and 14, anterior tergo-
pleural 19, and sternal remotor 28 muscles in Tibicen chloromera were not re-
ported in Huechys sanguinea var. philaemata .

The anterior internal tergo-sternal muscle 10 reported by Maki (1938) and
muscle 144 by Berlese (1909) are absent in Tibicen chloromera. This muscle
arises on the tergum and attaches to the ventrolateral cervical sclerite. However,
the anterior tergo-sternal muscles 12 in Tibicen chloromera , which has its origin

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[Vol. LXXIV

on the posterior end of the head and attaches to the ventrolateral cervical
sclerite, is probably the homologue.

Berlese (1909) did not report the presence of tergo-pleural, sternal promotor,
sternal remotor, tergal abductor, pleural abductor or pleural depressor muscles
in the prothorax of Cicada { — Tibicen) plebeia. The anterior internal tergo-
sternal 14, anterior tergo-pleural 19, and sternal remotor 28 muscles in Tibicen
chloromera do not have counterparts in the other two species of cicadas.

Mesothoracic Musculature: The mesothorax is the largest division of the
thorax and necessarily so, since it contains the very large dorsal longitudinal
(34) and oblique dorsal (35) muscles. The dorsal longitudinal muscles serve as
depressors of the wings while the oblique dorsal muscles are probably wing ele-
vators (Snodgrass, 1927 and 1935).

The tergo-pleurals 42, 43, and 44, trochantino-basalar 52, sternal remotor 58,
and sternal adductor 59 muscles in Tibicen chloromera have not been reported
in Huechys sanguinea var. philaemata by Maki ( 1938) or in Cicada (= Tibicen)
plebeia by Berlese (1909). The trochantino-basalar muscle 51 in Tibicen
chloromera is present in Cicada {—Tibicen) plebeia (79 + 80) but not in
Huechys sanguinea var. philaemata. The pleural abductor of the coxa, Maki’s
muscle number 46 in Huechys sanguinea var. philaemata , was not described by
Berlese (1909) in Cicada {—Tibicen) plebeia nor is it present in Tibicen
chloromera.

Berlese (1909) includes median dorsal 69 and spino-furcal ventral 104 muscles
in Cicada { — Tibicen) plebeia. Both of the above muscles are not present in
the two other species of cicadas (Table 6). Berlese (1909) did not report the
presence of pleural-subalar, sternal promotor, sternal remotor, sternal adductor,
sternal depressor, or spiracular muscles.

Metathoracic Musculature: The metathorax is extremely short, especially

dorsally, where the entire notum is reduced to a narrow band behind the scutel-
lum of the mesonotum.

The metathoracic musculature in Tibicen chloromera is homologous to that of
Huechys sanguinea var. philaemata , with the exception of the tergo-pleural
muscle 70 and the sternal remotor muscle 82. Berlese (1909) did not report the
presence of tergo-pleural, sternal-pleural, sternal promotor, pleural abductor,
pleural depressor, sternal depressor, and spiracular muscles. However, all of
the above mentioned muscles were reported by Maki (1938) in Huechys san-
guinea var. philaemata and are present in Tibicen chloromera.

3. THE CERVICOTHORACIC NERVOUS SYSTEM

General: A dorsal view of the ventral nerve cord in the male of Tibicen chloro-
mera (Walker) is shown in Fig. 9. A general description of the thoracic nervous
system is presented under the section entitled “The Ventral Nerve Cord.”

March, 1966] Vasvary: Morphology of Annual Cicada 31

Fig. 9. Dorsal view of the ventral nerve cord of the male annual cicada Tibicen chloromera
(Walker) from the head to the first abdominal segment.

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[Vol. LXXIV

The Cervix and the Prothorax: The narrowed membranous region between
the head and prothorax of insects is called the cervix or neck and is pre-
sumably derived from portions of both the labial and prothoracic segments.
Muscles contained within the cervical region are believed to have evolved from
both the labial and prothoracic segments. Therefore, the concept that the
muscles of a segment are innervated from the ganglion of that segment suggests
that each muscle of the cervix can be assigned either to the labial or prothoracic
segment by determining the segment of innervation.

Three pairs of nerves from the subesophageal ganglion innervate muscles
located in the cervical region: SN4 + SN5, SN6 (Fig. 9), and SN7 (Fig. 1A).
Nerve SN4 + SN5 innervates the anterior intersegmental muscle 13 and
nerve SN6 innervates the anterior intersegmental muscle 16. The SN7 nerve
innervates the muscles of the sclerotized rod, mr, and muscles within the
labium, mlb (Fig. 1A). The sclerotized rod is an extension of the labium and
hangs freely from the cervical membrane.

A nerve branch from SN6, designated as SN6a in figs. 1A and 9, may be
associated with the innervation of the anterior intersegmental muscle 15 and
possibly other muscles in the cervicoprothoracic area. A precise determination
could not be made since the SN6a nerve joins with a nerve formed by the
coalescence of nerves SN9 and INI. The resulting nerve, INI + SN9 + SN6a,
then coalesces with the IN2 nerve to form nerve IN2 + INI +SN9 +SN6a
which innervates muscles associated with the cervical sclerites, dorsal muscles,
and muscles located in the anterior portion of the prothorax.

The SN9 nerves issue from the interganglionic connectives between the
subesophageal ganglion and prothoracic ganglion. Nerve branch SN9a inner-
vates the internal ventral muscle 9 and external ventral muscle 10 before joining
the IIN10 + IIN11 nerve originating from the thoracic-abdominal ganglionic
mass (Fig. 9). Nerve SN9 then coalesces with nerve INI and later receives
the SN6a nerve before joining nerve IN2 originating from the prothoracic
ganglion.

The INI nerves arise from the anterior portions of the prothoracic ganglion
adjacent to the interganglionic connectives. Nerves INI proceed anteriorly in
close association with the interganglionic connectives before coalescing with the
SN9 nerves.

The IN2 nerves issue from the anterolateral area of the prothoracic ganglion
and pass under the internal ventral muscles 9. The first nerve branch, IIN2a,
provides two sensory nerve branches to the integument, then passes around the
tergal promotor muscle of the coxa 21 and over the anterior basal rim of the
prothoracic coxa into the leg. After IN2 coalesces with nerve INI + SN9 +
SN6a to produce nerve IN2 + INI + SN9 + SN6a, a nerve branch is formed
which combines with nerve branches from nerve IIN10 + IIN11 to innervate
the anterior intersegmental muscle 15. Nerve IN2 + INI + SN9 + SN6a then

March, 1966]

Vasvary: Morphology of Annual Cicada

Fig. 10. Dorsal view of the ventral nerve cord of the male annual cicada, Tibicen chloro-
mera (Walker), from nerve root IIN3 of the thoracic-abdominal ganglionic mass to the
second abdominal segment and showing the innervation pattern of the first abdominal
segment.

ramifies into three nerve branches. One nerve branch proceeds anteriorly and
innervates the anterior intersegmental muscles 11 and 12, the ordinary tergo-
pleural muscle 20, and the anterior tergo-pleural muscles 18 and 19. The
lateral nerve branch bifurcates into a ventral branch which innervates the pleural
abductors 30 and 31 and a dorsal branch which innervates the tergal abductor

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I Vol. LXXIV

muscle 29 and the tergal promotor muscles 21 and 22. The remaining nerve
branch proceeds dorsally and innervates the anterior intersegmental muscle 14,
anterior dorsal muscle 8, median dorsal muscle 5, lateral dorsal muscles 6 and 7,
and median dorsal muscle 4.

The IN3 nerves arise from the anterolateral portion of the prothoracic gan-
glion, pass under the anterior edge of the prothoracic pleural apophysis, and
innervate the sternal promotor 23, pleural depressor 33, and sternal remotor 28
muscles.

Nerve IN4 arises from the anterolateral portion of the prothoracic ganglion
posterior to IN3 and proceeds in a lateral direction passing under the prothoracic
pleural apophysis. IN4 provides a nerve branch into the leg before dividing
into nerve branches IN4a and IN4b. IN4a is a sensory nerve and provides
nerve branches to the posterolateral protergal area. Nerve IN4a innervates
tergal remotor muscles 24, 25, 26, and 27 and the tergal depressor muscle 32.

Nerve IN5 is very short and issues from the mediodorsal area of the pro-
thoracic ganglion and innervates ventral muscle 1.

Nerves IN6 and IN 7 arise from the mediodorsal portion of the ganglion pos-
terior to nerve IN5 and coalesce with nerve IN8 which arises from the dorsal
area of the interganglionic connective between the prothoracic ganglion and
thoracic-abdominal ganglionic mass (Fig. 9). It appears that nerves IN6, IN7,
and INS are responsible for the innervation of ventral muscle 2 and the occlusor
muscle 64 of the mesothorax.

The posterior tergo-sternal muscle 17 of the prothorax is innervated by
nerve branch IIN2a which arises from the anterolateral area of the thoracic-
abdominal ganglionic mass. Nerve IIN2a passes under nerve IINl and proceeds
lateral to the longitudinal ventral muscle 36 and along the posterior edge of the
prothoracic pleural arm to the posterior tergo-sternal muscle 17.

Mesothorax: The mesothorax is the largest thoracic segment in the male of
the annual cicada, Tibicen chloromera (Walker). Innervation of the meso-
thoracic segment, with the exception of the posterior tergo-sternal muscle 38
and the occlusor muscle 64, is achieved by five pairs of nerves: IINl, IIN2,
IIN3, IIN4, and IIN10 + IIN11.

The large IINl nerves arise from the short interganglionic connectives between
the prothoracic ganglion and thoracic-abdominal ganglionic mass GII (Fig. 9).
Nerve IINl passes over nerve IIN2 and provides nerve branch IINl a to the
longitudinal ventral muscle 36.

Nerve IINla bifurcates into two nerve branches. One nerve branch enters
muscle 36 along its mesal surface while the remaining nerve branch enters the
lateral surface of the muscle 36. The IIN2 nerve passes laterad to the longi-
tudinal ventral muscle 36 and ramifies into six nerve branches along the ventral
edge of the median dorsal muscle 34. The large median dorsal muscle 34
is innervated by three nerve branches while the lateral dorsal muscle 35 is in-

March, 1966]

Vasvary: Morphology of Annual Cicada

Fig. 11. Posterolateral view of the nerves and muscles of the right side of the fourth
abdominal segment of the male of Tibicen chloromera (Walker). First stage of dissection.

New York Entomological Society

[Vol. LXXIV

nervated by a single nerve branch which passes between the median dorsal
muscle 35 and the anterior tergo-sternal muscle 37. The remaining nerve
branches (IINlb, IINlc, and IINld) are sensory nerves and terminate in the
integument of the mesotergum (Fig. 9). Nerve branch IINlb passes obliquely
between the anterior tergo-sternal muscle 37 and the median dorsal muscle 34
and continues nresad to the furco-entopleural muscle 49 and tergal remotor
muscle 54. Nerve IINlb provides a nerve branch to the trachea between
muscle 54 and 55 before terminating in the integument in the posterior portion
of the mesotergum. IINlc passes mesad to median dorsal muscle 34 and
divides into two nerve branches. One nerve branch continues dorsally and
terminates in the middorsal region of the mesotergum while the other nerve
branch passes between the latter nerve and muscle 34 and terminates in the
integument in the anterodorsal region of the mesotergum. Nerve IINld passes
obliquely over the anterior tergo-sternal muscle 37 then proceeds along the
posterior edge of muscle 37 and terminates in the integument of the middorsal
region of the mesotergum.

Nerve IIN2 arises from the anterolateral surface of the thoracic-abdominal
ganglionic mass. Two nerve branches, IIN2a and IIN2b, issue from the base
of nerve IIN2. The IIN2 nerve passes under the ventral longitudinal muscle 36,
and provides a nerve branch IIN2c to nerve IIN3 (Fig. 9). Nerve IIN2 pro-
ceeds anterodorsally along the posterior edge of the profurcal arm and then
passes laterad to the anterior tergo-sternal muscle 37. The IIN2 nerve divides
into three nerve branches, IIN2d, IIN2e, and IIN2f, prior to entering the
forewing. Nerve branch IIN2d enters the integument below the tegula of the
mesothorax. Nerve branch IIN2e terminates in the integument in the region
of the third axillary sclerites and nerve branch IIN2f enters the mesothoracic
tegula. Nerve IIN2 is the anterior wing nerve (AWN, Fig. 9) and enters the
base of the mesothoracic wing. Nerve branch IIN2a innervates the posterior
tergo-sternal muscle 17 of the prothorax. Nerve branch IIN2b passes under
the ventral longitudinal muscle 36, proceeds around the posterior edge of the
anterior tergo-sternal muscle 37, and innervates the furco-entopleural muscle
49 and the tergal remotor muscle 54, before terminating in the integument along
the lateral edge of the mesotergum.

Nerve IIN3 is a large nerve and arises from the lateral surface of the thoracic-
abdominal ganglionic mass. The first nerve branch passes under the nerve IIN4
and bifurcates into two nerve branches. One nerve branch enters the integument
while the remaining nerve branch passes over the anterior basal aim of the
mesothoracic coxa and enters the leg. Nerve branch IIN3a innervates the sternal
promotor of the coxa 53, sternal remotor of the coxa 58, sternal adductor of the
coxa 59, tergal depressor 61, trochantero-basalar 62, and the tergal promotor of
the coxa 50. Nerve IIN3 is then connected to IIN2 by way of nerve branch
IIN2c. The next nerve branch, IIN3b, innervates the sterno-basalar muscle 48,

March, 1966]

Vasvary: Morphology of Annual Cicada

the trochantero-basalar muscle 62, the trochantino-basalar muscles 51 and 52,
and the coxo-basalar muscle 60.

Nerve IIN3 then ramifies into three nerve branches, one innervating the
anterior tergo-sternal muscle 37, another which innervates the tergal promotor
muscle 50, and a nerve branch designated as IIN3c (Fig. 9). Nerve branch
IIN3c innervates the anterior tergo-sternal muscle 37 and the tergo-pleural
muscles 39, 40, 42, and 43.

Nerve IIN4 issues from the lateral surface of the thoracic-abdominal gangli-
onic mass posterior to IIN3 and proceeds posteriorly and ramifies into three
nerve branches (Fig. 9). One nerve branch innervates the sternal depressor
muscle of the coxa 63 before passing over the posterior basal rim of the coxa
and into the mesothoracic leg. Nerve branch IIN4a innervates the furco-ento-
pleural muscle 49 and the sternal remotor muscle 57. Nerve IIN4 passes
around the posterior edge of the sternal remotor muscle 57 and provides a nerve
branch to the coxo-subalar muscle 56. Nerve IIN4 continues dorsally and
innervates the tergo-pleural muscles 41 and 44, the pleuro-axillary muscles 45
and 46, and the tergal remotor muscles 54 and 55.

The posterior tergo-sternal muscle 38 of the mesothorax is innervated by
nerve branch IIN6a'. Nerve branch IIN6a is the posterior wing nerve (PWN,
Fig. 9). It is noteworthy that the posterior tergo-sternal muscle 17 of the
prothorax is innervated by a nerve branch IIN2a of the anterior wing nerve
IIN2.

The pleuro-subalar muscle 47 is innervated by a nerve branch formed by the
coalescence of IIN10 + IIN11 and nerve branch IIN6a//r.

Ventral muscle 3 is innervated by a nerve branch from the IIN10 + IIN11
nerve (Fig. 9).

Metathorax: The metathorax is the shortest thoracic segment in the male of
the annual cicada, Tibicen chloromera (Walker). The entire notum is reduced
to a narrow band behind the scutellum of the mesonotum (Fig. 2). Innervation
of the metathoracic segment is achieved by three pairs of nerves: IIN5, IIN6,
and IIN1Q + IIN11 (Fig. 9).

Nerve IIN5 arises from the lateroposterior surface of the thoracic-abdominal
ganglionic mass and passes mesad to the mesofurca. After receiving nerve
branch IINlla, nerve IIN5 continues posteriorly and ramifies into five nerve
branches (Fig. 9). The anterior nerve branch, IIN5, is the dorsal nerve and
innervates the tergal promotor muscle 75, the anterior tergo-sternal muscle 67,
and terminates in the median dorsal muscles 65. The next nerve branch origi-
nates at the base of IIN5 and bifurcates into a nerve branch which enters the
integument and nerve branch IIN5a which innervates the pleural promotor
muscle 76 and the pleural abductor muscle 83. A nerve branch originating
between IIN5 and IIN5b provides a nerve to the integument before passing
over the anterior basal rim of the metathoracic coxa and into the leg. Nerve

New York Entomological Society

[Vol. LXXIV

IN9

Fig. 12. Posterolateral view of the nerves and muscles of the right side of the fourth
abdominal segment of the male of Tibicen chloromera (Walker). Second stage of dissection.

March, 1966]

Vasvary: Morphology or Annual Cicada

branch IIN5b passes under the posterior mesof ureal arm and provides a nerve
branch to the pleural depressor muscle 86 before innervating the tergal depressor
muscles 84 and 85. The remaining nerve branch innervates the sternal promotor
muscle 77, the sternal depressor muscle 87, and the sternal remotor muscle 81.

Nerve IIN6 passes under the posterior mesofurcal arm and forms nerve
branch IIN6a which passes around the posterior tergo-sternal muscle 38 of the
mesothorax providing the nerve branch, IIN6a' to muscle 38. Nerve branch
IIN6a" passes around the tergal depressor muscle 84 and proceeds along the
posterior edge of the second phragma (2 Ph) and provides a nerve branch to
the membranous sac of the abdominal air chamber (MS) before terminating in
the integument along the posterolateral edge of the metatergum (Fig. 9).
Nerve branch IIN6ar// coalesces with nerve IIN10 + IIN11. Nerve IIN6a is
the posterior wing nerve and passes along the anterior edge of the second
phragma and enters the base of the metathoracic wing. Nerve IIN6 passes under
the posterior metafurcal arm and proceeds dorsally along the posterior edge of
the tergal remotor muscle 79. Nerve IIN6 provides a nerve branch which enters
the integument of the epimeron before innervating the following muscles: coxo-
subalar 80, posterior tergo-sternal 68, tergal remotors 78 and 79, tergo-pleural
69 and 70, and the pleural-axillary muscles 71 and 72. Nerve IIN6b provides a
nerve branch to the metathoracic leg before innervating the sternal remotor
muscles 82 and the coxo-subalar muscle 80. Nerve branch IIN6c innervates
the furco-entopleural muscle 74.

Nerves IIN10 and IIN11 arise from the posterior dorsolateral surface of the
thoracic abdominal ganglionic mass and coalesce to form nerve IIN10 + IIN11.
The anterior branch of nerve I1N10 + IIN11 passes mesad to the ventral muscle
3 and provides a nerve branch to muscle 3 prior to passing mesad to the pro-
furcal arm and the posterior tergo-sternal muscle 17 of the prothorax. Nerve
IIN10 + IIN11 continues anteriorly passing over the large trachea of the
mesothoracic spiracle and provides two nerve branches which coalesce with a
nerve branch from IN 2 + INI + SN9 + SN6a that innervates the anterior
internal tergo-sternal muscle 15 of the prothorax. A nerve branch from SN9a
also joins nerve IIN10 + IIN11. The pattern of axon distribution resulting
from this fusion requires histological clarification. However, it appears that
the anterior intersegmental muscle 15, which attaches to the anterior dorso-
lateral region of the protergum and the ventrolateral cervical sclerite, does, in
part, receive its innervation from nerve IIN10 + IIN11. The IIN10 + IIN11
nerve continues anteriorly and passes under the tentorial bridge.

Nerve IIN11 provides a short nerve branch, IINlla, that coalesces with nerve
IIN5.

The posterior branch of IIN10 + IIN11 passes dorsally over the caudal
portion of the thoracic-abdominal ganglionic mass and bifurcates into a
dorsal branch and ventral branch. The dorsal branch divides forming two

New York Entomological Society

[Vol. LXXIV

nerve branches. One nerve branch passes mesad to IIN6a' and enters
the large trachea originating from the metathoracic spiracle. The second
nerve branch passes laterad to IIN6a' and coalesces with IIN6a/r/ to innervate
the sterno-pleural muscle 7 3, the pleuro-subalar muscle 47 of the meso thorax,
and the occlusor muscle 88 of the metathoracic spiracle. The ventral branch,
IIN10 + IIN11, continues posteriorly and innervates the longitudinal ventral
muscle 66, then forms a loop which proceeds anteriorly and provides a nerve
branch which coalesces with nerve IIN7. The IIN10 + IIN11 nerve proceeds
in a posterior direction and enters the first abdominal segment which contains
the sound mechanism. Further description of the innervation pattern of the
IIN10 + IIN11 nerve is presented under the section entitled “The Musculature
and Innervation of the Sound Mechanism.”

Nerve IIN7 innervates the muscles located in the anterior portion of the
first abdominal segment while nerve IIN8 innervates the muscles located in
the posterior portion of the first abdominal segment. The IIN9 nerves innervate
the remaining pregenital abdominal segments by providing a pair of lateral
nerve roots to each consecutive segment.

Discussion: The thoracic nervous system in the male of the annual cicada,
Tibicen chloromera (Walker), presents a perplexing enigma regarding the
determination of nerve homologies. This problem is due to the coalescence of
the mesothoracic, metathoracic, and abdominal ganglia into a single ganglionic
mass located in the mesothorax. Condensation of the ventral nerve cord has
presumably resulted in the coalescence of lateral nerve branches thereby produc-
ing apparent variations in the nerve distribution pattern in Tibicen when com-
pared to nerve patterns described in other insects.

Schmitt (1962) suggests utilization of the dorsal longitudinal muscles as a
starting point in establishing nerve homologies. The dorsal longitudinal muscles
are innervated by the dorsal nerves of each consecutive segment. Therefore,
from a descriptive standpoint, it is usually easy to identify the dorsal nerve
as it issues from its ganglion. However, Niiesch (1954) has shown that some
of the axons which supply the dorsal longitudinal muscles also originate from
the immediately anterior ganglion.

The dorsal muscles of the prothorax in Tibicen is innervated by nerve IN2 +
INI + SN9 + SN6a. Nerve SN9 may be the anterior ganglionic connective of
the dorsal nerve and has adhered to the interganglionic connective. However, it
cannot be determined without recourse to histological examination if nerve INI
or nerve IN2 is the prothoracic dorsal nerve.

The dorsal longitudinal muscles of the mesothorax are innervated by the
I INI nerves which arise from the very short interganglionic connectives between
the prothoracic ganglion and the thoracic-abdominal ganglion mass. Anterior
ganglionic connectives are not visible in Tibicen. Niiesch (1954) has demon-
strated in Telea that motor axons from the prothoracic ganglion pass through

March, 1966] Vasvary: Morphology of Annual Cicada 41

the anterior ganglionic connectives to the mesothoracic dorsal nerve. Anterior
ganglionic connectives are not visible in Chauliodes , as described by Maki (1936),
nor in Agulla , as described by Matsuda (1956). However, Schmitt (1962)
proposes that if the findings of Niiesch regarding the innervation of the dorsal
longitudinal muscles are applicable to other Neopterygota, it is probable that the
fibers of the anterior ganglionic connectives are also present in Chauliodes and
Agulla , but are incorporated in the interganglionic connectives to their connec-
tion with the dorsal nerves.

Nerve IIN5 in Tibicen is the dorsal nerve of the metathorax and provides in-
nervation to muscles located in the anterior portion of this segment and the
dorsal longitudinal muscles.

In the Neopterygota, wing nerves may also be useful in establishing nerve
homologies. The wing nerve enters the wing cavity and is associated with
sensory structures at the base of the wing. In Tibicen there are two wing nerves.
The anterior wing nerve IIN2 arises from the anterolateral surface of the
thoracic-abdominal ganglionic mass. Two nerve branches arise from the base
of the anterior wing nerve (IIN2a and IIN2b) and a third nerve branch IIN2c
coalesces with nerve IIN3 (Fig. 9). Nerve IIN2 then continues as a completely
independent nerve providing sensory nerve branches to the integument below
the tegula, the tegula, and the integument in the region of the third axillary
sclerites before entering the wing cavity. Maki (1936) describes a similar
condition in Chauliodes , where a separate nerve which he labeled “fourth root”
arises from the mesothoracic ganglion and passes directly into the wing. In
Dissosteira , Schmitt (1959) found that in addition to innervating the dorsal
muscles of the mesothorax, the dorsal nerve also provides an anterior nerve
branch which enters the tegmen anteriorly and a posterior branch, entering
the same wing posteriorly. In Agulla , Matsuda (1956) also found that a
branch of the dorsal nerve enters the wing. It appears that the wing nerves of
Tibicen and Chauliodes are homologous to the wing nerves of Dissosteira and
Agulla despite their association with dorsal nerves in the latter two insects.
The posterior wing nerve in Tibicen arises as a nerve branch, IIN6a, from nerve
IIN6 (Fig. 9). Nerve IIN6a provides three nerve branches, IIN6a', IIN6a",
IIN6a"', before continuing as a separate nerve and passing directly into the
metathoracic wing cavity.

It is interesting to note that the thoracic legs receive their innervation from
two pairs of nerves, one entering the coxae anteriorly and the other posteriorly.
The prothoracic legs are innervated by nerve branches from nerves IN2 and
IN4 and the mesothoracic legs by nerve branches from IIN3 and IIN4 while
nerve branches from IIN5 and IIN6 enter the metathoracic legs.

In Tibicen no median nerves are visible between the subesophageal ganglion
and the prothoracic ganglion nor between the prothoracic ganglion and the
thoracic-abdominal ganglionic mass. However, the median nerves may be

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[Vol. LXXIV

included within the interganglionic connectives. The transverse or lateral nerves
to the occlusor muscles of the mesothoracic spiracles arise from the dorsal
surface of the prothoracic ganglion. The mesothoracic occlusor muscle 64 is
innervated by a nerve resulting from the coalescence of nerves IN6, IN 7, and
IN8 (Fig. 9). Case ( 1957) has shown that in the cockroach the axons to muscles
of a thoracic spiracle leave the anterior ganglion by way of the median nerve,
passing to the transverse nerve and then to the muscles. Hoyle (1959) has
reported a similar axon path in the thorax of Schistocerca gregaria. It appears
then that the nerve formed by the coalescence of IN6, IN7, and INS is in part
the transverse nerve since it terminates in the occlusor muscle of the mesothoracic
spiracle. In many instances the transverse nerves from the prothoracic ganglion
coalesce with the mesothoracic dorsal nerves. This was found to be true in
Agulla and Blattella by Matsuda, in Carausiaus by Marquardt (1939), in
Chauliodes by Maki (1936), in Dissosteira by Schmitt (1959), in Periplaneta
by Pipa and Cook (1959), in Perla by Wittig ( 1955), and in Telea by Niiesch
(1957). Schmitt (1962) mentions that no explanation has been offered for
the coalescence of dorsal nerves to the transverse nerves and presumably the
transverse nerves provide other functions in addition to exercising control
over the spiracles.

The IIN10 + IIN11 nerves may contain axons of a transverse nerve, since a
nerve branch from IIN10 + IIN11 coalesces with nerve branch IIN6a//r and
the resulting nerve terminates in the occlusor muscle 88 of the metathoracic
spiracle.

4. THE MUSCULATURE AND INNERVATION OF THE SOUND MECHANISM

General: The musculature of the sound mechanism of Tibicen chloromera
(Walker) is shown in Figs. 2 to 4 and a list of these muscles with their muscle
numbers and attachments is presented in Table 8. Fig. 10 shows that the
innervation of the first abdominal segment is achieved by nerves IIN7, IIN8, and
IIN10 + IIN11. Nerve branch IIN8a is the auditory or tymbal nerve since it
innervates the large tergo-sternal muscle 94 or tymbal muscle of the sound
mechanism. Nerve IIN10 + IIN11 provides nerve branches to IIN8a and the
tymbal muscle.

The Musculature of the Sound Mechanism: The musculature of the sound
mechanism of Tibicen chloromera (Walker) is contained within the first ab-
dominal segment. The tergo-sternal muscle 94 (Fig. 2), or tymbal muscle, is the
largest muscle of the sound mechanism and has its attachments on the basal
portion of the first abdominal sternum and a sclerotized terminal plate which
attaches to the tymbal by a tendon. The tymbal muscles and the tymbals are
the essential elements of the sound-producing apparatus (Myers, 1928).

The first abdominal sternite has become modified into a sclerotized V-shaped
structure which provides attachments and support for the large tymbal muscles.

March, 1966]

Vasvary: Morphology of Annual Cicada

dim

Fig. 13. Diagram of the nerve pattern of the right side of the fourth abdominal segment
of the male of Tibicen chloromera (Walker) viewed mesally.

Carlet (1876), Vogel (1923), and Myers (1928) have established that this
sclerotized V-shaped structure is a modification of the first abdominal sternum.
A sternal canal is present within the base of this structure and provides a
passageway for two pairs of nerves, IIN8 and IIN9. The “wings” or “arms”

New York Entomological Society

I Vol. LXXIV

Table 8. Musculature of the sound mechanism of Tibicen chloromera (Walker).

Muscle

Muscle Origin

number (or attachment)

Insertion
(or attachment)

Dorsal muscle

Anterior intersegmental fold Tergal ridge

Dorsal muscle

Tergal ridge

Posterior intersegmental fold

Ventral muscle

Metafurca

Anterior edge of first abdominal
sternite

Ventral muscle

Metafurca

Lateral apodemal arm of anterior
edge of sternum

Ventral muscle

Posterior edge of first ab-
dominal sternum

Posterior intersegmental fold

Tergo-sternal muscle
(Tvmbal muscles)

Basal portion of first ab-
dominal sternum

Terminal plate which attaches to
tymbal by a tendon

Tergo-sternal muscle

Anterolateral edge of ter-
gum

Lateral apodemal arm of ante-
rior edge of sternum

Tergo-sternal muscle

Anterolateral edge of ter-
gum ventrad to 95

Lateral apodemal arm of ante-
rior edge of sternum

Tergo-sternal muscle

Anterolateral edge of ter-
gum ventrad to 96

Lateral apodemal arm of ante-
rior edge of sternum

Tergo-sternal muscle

Anteroventral edge of tym-
bal

Lateral apodemal arm of ante-
rior edge of sternum

Tergo-sternal muscle

Tergum along lateral edge
of mirror

Lateral apodemal arm of poste-
rior edge of sternum

Tergo-sternal muscle

100

Posterior intersegmental
fold

Lateral apodemal arm of poste-
rior edge of sternum

Occlusor of spiracle

101

Lateral apodemal arm of
posterior edge of sternum

Ventral end of spiracle

Occlusor of spiracle

102

Lateral apodemal arm of
posterior edge of sternum

Base of spiracle

of the sclerotized V-shaped structure attach to the tergal ridge of the first
abdominal segment.

A comparison of the musculature of the sound mechanism of Tibicen chloro-
mera with Huechys sanguinea var. philaemata described by Maki (1938) and
Cicada {— Tibicen ) plebeia described by Berlese (1909) is presented in Table
9. The musculature of Tibicen differs from that found in the two other species
of cicadas compared in Table 9 by the presence of tergo-sternal muscles 96, 97,
and 98. Maki’s ventral muscle 76 in Huechys was not described by Berlese
(1909) in Cicada (= Tibicen) plebeia ; however, it is present in Tibicen chloro-
mera. In Tibicen , the dorsal muscles 89 and 90 have attachments on the tergal
ridge (tr) of the first abdominal segment (Fig. 2). Berlese (1909) shows in his
figure 542 similar points of attachments for the dorsal muscles 37 and 28-29 in
Cicada (= Tibicen) plebeia. However, the dorsal muscle 74 of Huechys san-
guinea var. philaemata , as shown by Maki (1938) in his figure 24, has its
attachment on the anterior and posterior intersegmental folds of the first
abdominal segment.

Innervation of the Sound Mechanism: The innervation of the sound mech-
anism is shown in Fig. 10. Nerves IIN7, IIN8, and IIN10 + IIN11 provide
innervation to the muscles of the first abdominal segment which contains the

March, 19661

Vasvary: Morphology or Annual Cicada

sound mechanism. Nerve IIN7 issues from the posterior portion of the thoracic-
abdominal ganglionic mass and proceeds posteriorly and passes over the meso-
furca. Nerve IIN7 receives a nerve branch from the IIN10 + IIN11 nerve
before forming the nerve branch IIN7a. Branch IIN7a provides innervation to
the ventral muscle 92, the membranous sac of the abdominal air chamber, and
occlusor muscle of the first abdominal spiracle 101 before coalescing with the
dorsal nerve IIN7. Nerve IIN7 is the anterior dorsal nerve of the first abdomi-
nal segment since it terminates in the dorsal muscles 89. Nerve IIN7 provides a
nerve branch to the ventral muscle 91 and passes under muscle 91, around the
posterior arm of the metafurca, and continues along the anterior edge of the
first abdominal segment. Nerve IIN7 provides a sensory nerve branch to the
integument and coalesces with nerve IIN10 + IIN11 before innervating the
tergo-sternal muscles 95, 96, and 97. Nerve IIN7 then provides a nerve branch
to the membranous sac surrounding the tymbal muscle 94 and coalesces with
nerve branch IIN7a before providing a nerve branch to the tergo-sternal muscle
98 and dorsal muscle 89 (Fig. 10).

Nerve IIN8 issues from the posterior portion of the thoracic-abdominal gan-
glionic mass and proceeds posteriorly over the mesofurca. Nerve IIN8 then
divides into a dorsal nerve branch IIN8a which terminates in the tergo-sternal
muscle 94, and a ventral nerve IIN8 which passes between the ventral muscles
91 and enters the sternal canal. Nerve branch IIN8a is the auditory nerve and
proceeds dorsally and for a portion of its length adheres to the IIN8a nerve
from the opposite side. Nerve branch IIN8a then receives a nerve branch from
the IIN10 + IIN11 nerve and proceeds in a dorso-oblique path over the mesal
surface of the large tergo-sternal muscle 94. Nerve IIN8a then passes around
the dorso-posterior edge of muscle 94 and enters this muscle along its lateral
surface. Nerve IIN8 passes through the sternal canal and provides nerve
branches to the integument and ventral muscle 93. Nerve IIN8 then passes
along the posterior edge of the first abdominal segment and provides nerve
branches to the occlusor muscle of the spiracle 102, tergo-sternal muscles 99 and
100, and the dorsal muscle 90. Nerve IIN8 is the posterior dorsal nerve since
it terminates in the dorsal muscle 90 located in the posterior portion of the first
abdominal segment.

The IIN10 + IIN11 nerve enters the first abdominal segment after supplying
a nerve branch to nerve IIN7 and proceeds posteriorly in a dorso-oblique path
and provides a nerve branch which coalesces with nerve IIN8a. The IIN10 +
IIN11 nerve continues dorsolaterally and provides a short nerve branch to
the tymbal muscle 94 before it loops in an anterior direction and coalesces with
nerve IIN7 (Fig. 10). The IIN10 + IIN11 nerve is the sympathetic nerve of
Voskresenskaya and Svidersky (1960), who report that without the innervation
of the sympathetic nerve the sound-producing system cannot function normally.
Therefore, both the auditory nerve, IIN8a, and the sympathetic unpaired nerve,

New York Entomological Society

[Vol. LXXIV

Table 9. Comparison of the musculature of the sound mechanism of Tibicen chloromera,
with Huechys sanguined var. philaemata (Maki, 1938) and Cicada (— Tibicen) plebeia

(Berlese, 1909) .

Muscles

Tibicen

chloromera

Huechys
sanguined
var. philaemata
(Maki, 1938)

Cicada
( = Tibicen )
plebeia

(Berlese, 1909)

Dorsal muscles

—

28-29

Dorsal muscle

—

Ventral muscle

—

Ventral muscle

—

14 + 15

Tergo-sternal muscle

XXVI

Tergo-sternal muscle

—

Tergo-sternal muscle

—

Tergo-sternal muscle

—

Tergo-sternal muscle

—

Tergo-sternal muscle

—

XVII

Tergo-sternal muscle

100

XVII

First interpleural muscle

—

LIII

Occlusor of spiracle

101

—

Occlusor of spiracle

102

IIN10 + IIN11, are necessary for the rhythmic “singing” of the cicada. It is
interesting to note that Hagiwara and Watanabe (1956) concluded that the
paired tymbal muscles receive alternate impulses from the ganglion, and this
alternate activity of the two tymbals may give a double sound vibration
frequency.

5. THE MUSCULATURE AND INNERVATION OF THE FOURTH ABDOMINAL SEGMENT

General: The abdominal musculature of adult and larval insects conforms to
a simple fundamental pattern which is repeated with only minor variations in
each of the pregenital segments (Snodgrass, 1935). The major groups of ab-
dominal muscles found in insects are the dorsal muscles, ventral muscles, lateral
muscles, transverse muscles, and spiracular muscles. The dorsal and ventral
muscles in most insects occur in two layers and thereby form dorsal internal and
external muscles and ventral internal and external muscles. In the male of
Tibicen chloromera (Walker) the dorsal external and ventral external muscles
are absent. The writer observed a similar condition in all of the typical pre-
genital abdominal segments. Another common form of diversification affecting
dorsal and ventral muscles includes a more or less distinguishable grouping of
the muscles into median and lateral sets (Snodgrass, 1935). In Tibicen the
dorsal muscles can be classified into dorsal internal median and dorsal internal
lateral muscle groups; however, the ventral internal muscles cannot be classified
into median and lateral sets since there are no ventral transverse muscles or a
wide separation between the ventral internal muscles.

March, 1966]

Vasvary: Morphology of Annual Cicada

The musculature of the fourth abdominal segment of the male of the annual
cicada, Tibicen chloromera (Walker) is shown in Figs. 11 and 12 and a list
of the muscles with their muscle numbers and attachments is presented in
Table 10.

The innervation of the abdominal musculature with the exception of the
muscles of the first abdominal segment is achieved by nerves IIN9. The IIN9
nerves provide a pair of lateral nerve branches to each consecutive pregenital
abdominal segment. Figs. 11 and 12 show the innervation of the fourth abdomi-
nal segment.

The Musculature of the Fourth Abdominal Segment: The musculature
of the fourth abdominal segment of the male of Tibicen chloromera (Walker)
can be classified into dorsal, ventral, lateral, and spiracular muscles (Table 10).
The dorsal muscles are subdivided into dorsal internal median (103) and dorsal
internal lateral (104) muscles, dorsal (105) and ventral (106) muscles of the
apodeme, and dorsal transverse muscles (107). The dorsal internal median and
dorsal internal lateral muscles have their attachments on the anterior and pos-
terior intersegmental folds while the dorsal and ventral muscles of the apodeme
have their attachments on the anterior edge of the apodeme and the posterior in-
tersegmental fold. It is interesting to note that there is a complete absence
of dorsal external muscles in the pregenital abdominal segments. The usual
location of the dorsal external muscles is the posterior portion of the abdominal
segment with their attachments on the posterior margin of the tergunr and the
posterior intersegmental fold. In this position the dorsal external muscles serve
as protractors of the abdomen. It appears that the protraction of the abdomen
in Tibicen is achieved by the contraction of the dorsal and ventral muscles of the
tergal apodeme. The dorsal transverse muscle, 107, has its attachments along
the lateral edge of the dorsal vessel and the anterolateral intersegmental fold
of the tergum.

Eight closely associated sets of ventral internal muscles, 108, are present in
Tibicen. The ventral internal muscles cannot be grouped into specific median
and lateral muscle sets.

Four pairs of lateral muscles are present in Tibicen : lateral internal 109,
lateral external 110, lateral intrasegmental 111, and the dilator of the abdomen
112 (Figs. 11 and 12). The lateral internal and external muscles are tergo-sternal
muscles and have their attachments on the anterior intersegmental fold of the
tergum and the internal surface of the anterior sternal apodeme. The lateral
intrasegmental muscle is tergo-sternal in its attachments and is located in the
posterior portion of the segment. The dilator of the abdomen is attached to the
anterolateral edge of the tergum and the external surface of the anterior sternal
apodeme.

The spiracle of the fourth abdominal segment is located in the anterolateral
corner of the sternum. The occlusor muscle of the spiracle has its attachments

New York Entomological Society

[Vol. LXXIV

on the anterolateral portion of the sternum adjacent to the sternal apodeme and
to the ventral edge of the spiracle.

Maki (1938) in his figure 24 shows the musculature of the third abdominal
segment of Huechys sanguined var. philaemata. The musculature of the fourth
abdominal segment of Tibicen chloromera (Walker) is homologous to the third
segment of Huechys sanguinea var. philaemata with the exception of the dorsal
internal lateral muscles 104, the lateral intrasegmental muscle 111, and the
dilator of the abdomen 112 of Tibicen (Table 11).

The Innervation of the Fourth Abdominal Segment: The innervation of
the fourth abdominal segment is achieved by a pair of lateral nerve branches
which arise from the IIN9 nerves. The IIN9 nerves issue from the posterior end
of the thoracic-abdominal ganglionic mass and pass over the mesofurca and meta-
furca and between the ventral muscles into the sternal canal. After the IIN9
nerves pass through the sternal canal they provide a pair of lateral nerve
branches to the second and remaining pregenital abdominal segments. The
lateral nerve branch from nerve IIN9 divides into a dorsal nerve and ventral
nerve prior to passing under the ventral internal muscles of the fourth abdominal
segment. The innervation of the fourth abdominal segment is shown in Figs. 1 1
and 12. The ventral nerve (VNv) passes under the dorsal nerve (DNv) and
terminates in the integument beneath the ventral internal muscles 108 (Fig. 12).
The dorsal nerve provides a nerve branch to the ventral internal muscles which
are innervated along their external surface.

The dorsal nerve proceeds laterally and provides a nerve (TNv) to the
occlusor muscle of the spiracle 113 (Fig. 13). Case (1957) presents experi-
mental evidence that the median and transverse nerves provide a neural pathway
connecting the spiracular mechanism with the central nervous system. Schmitt
(1965) presents a comparative morphological study on the transverse nerves
in the abdominal nervous system of insects and concludes that, in insects which
apparently lack median and transverse nerves, these nerves have become in-
corporated in the longitudinal connectives and lateral segmental nerves. In the
majority of insects reviewed by Schmitt, the transverse nerve also innervates the
alary muscles. In Tibicen chloromera (Walker) the writer could not determine
the innervation of the alary muscles; however, it appears reasonable to con-
clude that the innervation of the occlusor muscle of the spiracle of Tibicen is
accomplished by fibers of the transverse nerve which have become incorporated
in the dorsal nerve.

After providing a nerve branch to the occlusor muscle of the spiracle, the
dorsal nerve ramifies into three nerve branches. The anterior nerve branch in-
nervates the lateral internal muscle 109, lateral external muscle 110, and the
dilator of the abdomen 112. The posterior nerve branch divides into a sensory
nerve which terminates in the integument and a nerve branch which innervates
the lateral intrasegmental muscle 111 (Figs. 11 and 12).

March, 1966]

Vasvary: Morphology of Annual Cicada

Table 10. The musculature of the fourth abdominal segment of Tibicen chloromera

(Walker).

Muscle

number

Origin

(or attachment)

Insertion
(or attachment)

Dorsal muscles

Dorsal internal median muscles

103

Anterior intersegmental
fold

Posterior intersegmental fold

Dorsal internal lateral muscles

104

Anterior intersegmental
fold

Posterior intersegmental fold

Dorsal muscle of apodeme

105

Anterior edge of tergal
apodeme

Posterior intersegmental fold

Ventral muscle of apodeme

106

Anterior edge of tergal
apodeme

Posterior intersegmental fold

Dorsal transverse muscle

107

Anterior intersegmental
fold

Lateral edge of the dorsal
vessel

Ventral muscles

Ventral internal muscles

108

Anterior intersegmental
fold

Posterior intersegmental fold

Lateral muscles

Lateral internal muscle

109

Anterior intersegmental
fold of tergum

Internal surface of sternal
apodeme

Lateral external muscle

110

Anterior intersegmental
fold of tergum

Internal surface of sternal
apodeme

Lateral intrasegmental muscle

111

Posterolateral portion
of tergum

Lateral edge of sternum

Dilator of the abdomen

112

Lateral edge of tergum

External surface of sternal
apodeme

Muscles of the spiracle

Occlusor

113

Anterolateral portion of
sternum adjacent to the
sternal apodeme.

Ventral edge of spiracle

The dorsal nerve proceeds dorsally in an oblique-posterior direction over the
tergal apodeme and supplies a nerve branch to the dorsal and ventral muscles
(104 and 105) of the apodeme (Figs. 11 and 12). The dorsal nerve continues
dorsally along the posterior portion of the tergum and passes over the dorsal
internal lateral muscles 104 and provides nerve branches to these muscles. The
dorsal nerve then divides into three nerve branches; two nerve branches pass
laterally under the dorsal internal median muscles and terminate in the integu-
ment while the dorsal nerve passes mesally over the dorsal internal median
muscles supplying these muscles with nerve branches. The dorsal nerve termi-
nates in the first set of dorsal internal median muscles (Fig. 11).

The segmental nerve pattern in the male cicada, Tibicen chloromera (Walker),
is notably abbreviated when compared to the innervation pattern of some
families of Orthoptera, as described by Schmitt (1954), Chauliodes jormosanus
as described by Maki (1936), the larva and adult of Hyalophora cecropa as
described by Libby (1959 and 1961), and in Pteronarchys as described by
Schmitt (1963). The abbreviated nerve pattern in Tibicen is largely due to
the absence of the dorsal and ventral external muscles and by the condensation

New York Entomological Society

[Vol. LXXIV

Table 11. A comparison of the musculature of the fourth abdominal segment of Tibicen
chloromera (Walker) to the musculature of the third abdominal segment of Huechys
sanguined var. philaemata described by Maki (1938).

Muscle groups

Tibicen

chloromera

Huechys
sanguinea
var. philaemata
(Maki, 1938)

Dorsal muscles

Dorsal internal median muscles

103

Dorsal internal lateral muscles

104

—

Dorsal muscle of apodeme

105

Ventral muscle of apodeme

106

Dorsal transverse muscle

107

Ventral muscles

Ventral internal muscles

108

Lateral muscles

Lateral internal muscle

109

Lateral external muscle

110

Lateral intrasegmental muscle

111

—

Dilator of the abdomen

112

Muscles of the spiracle

Occlusors of the spiracle

113

of the ventral nerve cord which has resulted in the formation of a thoracic-
abdominal ganglionic mass located in the mesothorax. With the condensation of
the ventral nerve cord, the motor axons which supply the innervation to the
typical pregenital abdominal segments have become incorporated within a
single pair of nerves, IIN9. The IIN9 nerves supply a pair of lateral nerve
branches to each consecutive abdominal segment after which there are no nerve
connections between segments.

The innervation pattern of the fourth abdominal segment of Tibicen is shown
in Fig. 13. The dorsal nerve supplies innervation to the dorsal and ventral
internal longitudinal muscles which are considered primitive muscle groups of
the segmental musculature and are therefore useful in the establishment of a
criteria of nerve homology. Schmitt (1954) shows that the dorsal and ventral
internal muscles of Dissosteira , Acheta , and Periplaneta are innervated by nerve
branches from the dorsal nerve. Libby (1959 and 1961) shows a similar innerva-
tion of the same muscle groups in the larva and adult of Hyalophora.

Further investigations of insects possessing a thoracic-abdominal ganglionic
mass located in the thorax must be conducted before significant comparisons
can be made with the segmental nerve pattern of Tibicen.

SUMMARY AND CONCLUSIONS

The musculature and innervation of the thorax, of the sound mechanism, and
of a typical pregenital abdominal segment of the male of the annual cicada,
Tibicen chloromera (Walker) are described. The musculature of the thorax and

March, 1966]

Vasvary: Morphology of Annual Cicada

abdominal segments of Tibicen is essentially homologous to the musculature of
the male cicada Heuchys sanguined var. philaemata as described by Maki
(1938).

The ventral nerve cord consists of a subesophageal ganglion, prothoracic
ganglion, and a thoracic-abdominal ganglionic mass. There are no ganglia pres-
ent in any of the abdominal segments. The abdominal segments are innervated
by lateral nerve branches arising from a pair of nerves that originate from the
posterior portion of the thoracic-abdominal ganglionic mass located in the meso-
thorax. Eight pairs of nerves arise from the subesophageal ganglion and supply
innervation to the muscles associated with the feeding apparatus, the salivary
glands, the lateral ducts of the salivary glands, and some of the muscles of the
cervical area.

The prothoracic ganglion and the anterior portion of the thoracic-abdominal
ganglionic mass are covered dorsally by ventral muscles. The prothoracic
ganglion supplies innervation to some of the muscles of the cervical area and
the muscles of the prothorax. The thoracic-abdominal ganglionic mass provides
innervation to the posterior tergo-sternal muscles of the prothorax, the muscles
of the mesothorax, metathorax, and all the abdominal segments. No median
nerves are visible between the subesophageal ganglion, prothoracic ganglion, and
the thoracic-abdominal ganglionic mass. However, the median nerves are prob-
ably included within the interganglionic connectives. Spiracular muscles of the
thoracic segments are innervated by nerves which arise from the dorsolateral area
of the prothoracic ganglion and the thoracic-abdominal ganglionic mass. The
nerves to the spiracular muscles are apparently the transverse nerves of the
“ventral sympathetic nervous system.”

The sound mechanism is contained within the first abdominal segment. An
invagination of the first abdominal sternite serves as an area for attachment
and support for the large tymbal muscles. A sternal canal is located within the
sternal invagination and permits the passage of two pairs of nerves. One pair
of nerves innervates the muscles in the posterior portion of the first abdominal
segment while the remaining pair of nerves provides innervation to the remain-
ing abdominal segments.

Each typical pregenital abdominal segment is innervated by a pair of lateral
nerve branches which arises from a single pair of nerves originating from the
posterior end of the thoracic-abdominal ganglionic mass and pass through the
sternal canal. There are no nerve connections between the typical pregenital
abdominal segments once the lateral nerves enter their respective segments. A
single nerve branch from the dorsal nerve innervates the occlusor muscle of the
spiracle of the fourth abdominal segment. It appears that the innervation of
the occlusor muscle of the spiracle is achieved by fibers of the transverse nerve
which have become incorporated in the lateral nerve branches to the abdominal
segments.

New York Entomological Society

[Vol. LXXIV

Acknowledgments

I wish to express my gratitude to Dr. J. B. Schmitt of the Department of Entomology
and Economic Zoology, Rutgers-The State University, for his assistance and guidance in
the selection and suggestions for carrying out this morphological study. This paper is a
portion of a thesis submitted to the Graduate School of Rutgers-The State University in
partial fulfillment of requirements for the degree of Doctor of Philosophy.

Literature Cited

Beckel, W. E. 1958. The morphology, histology and physiology of the spiracular regu-
lating apparatus of Hyalophora cecropia (L.) Proc. Intern. Congr. Entomol. 10th
Meeting, Montreal, Que. 1956, 2: 87-115.

Berlese, A. 1909. Gli Insetti. Vol. I. Milan.

Binet, A. 1894. Contribution a l’etude du systeme nerveux sous intestinal des Insects.

Journ. de l’Amat. de la Phys. 30 Ann., Nr. 5: 449-543.

Brandt, E. 1878. Vergleichend-anatomische Untersuchungen fiber das Nervensystem der
Hemipteren. Horae Soc. Ent. Ross., Tom. 14: 496-505.

Carlet, G. 1876. Sur Panatome de l’appareil musical de la Cigale. C. R. Acad. d. Sc., T.

86.

Case, J. F. 1957. The median nerves and cockroach spiracular function. J. Insect Physiol.,
1 : 85-94.

Dufour, L. 1833. Recherches anatomiques et physiologiques sur les Hemipteres. Mem. d.

savant, etrang. a l’Acad. d. Sc. Tom. 4: 129-462.

Hagiwara, S., and A. Watanabe. 1956. Discharges in motoneurons of Cicada. Journ. Cell.
Comp. Physiol., 47: 415-428.

Hilton, W. A. 1939. The nervous system of Homoptera and Hemiptera. Journ. Ent. and
Zool., 31 : 36-38.

Holste, G. 1910. Das Nervensystem von Dytiscus marginalis. Z. Wiss. Zool., 96: 419-
476.

Hoyle, G. 1959. The neuromuscular mechanism of an insect spiracular muscle. J. Insect
Physiol., 3: 378-394.

Johansson, A. S. 1957. The nervous system of the milkweed bug, Oncopeltus jasciatus
(Dallas) (Heteroptera, Lygaeidae). Trans. Amer. Ent. Soc., 83: 119-183.

Libby, J. L. 1959. The nervous system of certain abdominal segments in the cecropia larva.
Ann. Ent. Soc. Amer., 52: 469-480.

— . 1961. The nervous system of certain abdominal segments and the innervation of

the male reproductive system and genitalia in Hyalophora cecropia. Ann. Ent. Soc.
Amer., 54: 887-896.

Maki, T. 1936. Studies on the skeletal structure, musculature and nervous system of
the alder fly, Chauliodes formosanus Peterson. Mem. Fac. Sci. and Agric., Taihoku
Imp. Univ., 16 (3): 117-253.

— . 1938. Studies on the thoracic musculature of insects. Mem. Faculty Sci. and Agric.,

Taihoku Imp. Univ., Formosa, Vol. 24, No. 1. Entomol. No. 10.

Malouf, N. S. R. 1933. The skeletal motor mechanism of the “Stink bug”, Nezara
viridula L. Bull. Soc. ent Egypte, Cairo, xvi (1932). 161-203.

Marquardt, F. 1939. Beitrage zur Anatomie der Muskulatur und peripheren Nerven von
Carausius ( Dixippus ) morosns Br. Zool. Jahrb. Anat., 66: 63-128.

Matsuda, R. 1956. The comparative morphology of the thorax of two species of insects.
Microentomology, 21: 1-63.

Myers, J. G. 1928. Morphology of the Cicadidae (Homoptera). Proc. Zool. Soc. London,
25: 365-472, 74 figs.

March, 1966]

Vasvary: Morphology of Annual Cicada

Nijenhuis, E. D., and D. Dresden. 1955. On the topographical anatomy of the nervous
system of the mesothoracic leg of the American cockroach ( Periplaneta americana) .
Konikl. Ned. Akad. Wetenschap. Proc. Ser. C., 58: 121-136.

Nuescil, H. 1954. Segmentierung und Muskelinnervation bei Telea polyphemus Cr. Rev.
Suisse Zook, 61 : 420-428.

— . 1957. Die Morphologie des Thorax von Telea polyphemus Cr. II. Nervensystem.

Zool. Jahrb. Anat., 75: 615-642.

Pipa, R. L., and E. F. Cook. 1959. Studies on the hexapod nervous system. I. The
peripheral distribution of the thoracic nerves of the adult cockroach, Periplaneta
americana. Ann. Entomol. Soc. Am., 52: 695-710.

Pringel, J. W. S. 1954. A physiological analysis of Cicada song. Journ. Exp. Biol., 31:
255-560.

Schmitt, J. B. 1954. The nervous system of the pregenital abdominal segments of some
Orthoptera. Ann. Ent. Soc. Amer., 47: 677-682.

. 1959. The cervicothoracic nervous system of a grasshopper. Smithsonian Inst.

Pubis. Misc. Collections, 137: 307-329.

. 1962. The comparative anatomy of the insect nervous system. Ann. Rev. of

Entomol., 7: 137-156.

. 1963. The abdominal nervous system in Pteronarcys. Jour. N. Y. Entomol. Soc.,

71: 202-217.

. 1965. Variations in the transverse nerve in the abdominal nervous system of

insects. Jour. N. Y. Entomol. Soc. 73: 144-150.

Snodgrass, R. E. 1927. Morphology and mechanism of the insect thorax. Smithsonian
Misc. Coll., 80: No. 1, 108 pp., 44 figs.

. 1935. Principles of Insect Morphology. McGraw-Hill Book Co., New York, N. Y.

Swinton, A. H. 1880. Insect Variety: its propagation, and distribution, treating of

odours, dances, colours, and music in all grasshoppers, cicadae and moths. London.
Vogel, R. 1923. Uber ein tympanales Sinnesorgan, das mutmalsliche Hororgan der
Singzikaden. Zeitschr f. wissensch. Zook, 120: 190-231.

Voskresenskaya, A. K., and V. L. Svidersky. 1960. The role of the central and sympa-
thetic nervous system in the function of the tymbal muscles of cicadas. Journ. Ins.
Physiol., 6: 26-35.

Weber, H. 1929. Kopf and Thorax von Psylla Mali Schmids. (Hemiptera-Homoptera) .
Z. Morph. Oekol. Tiere, xiv, 59-165.

Wittig, G. 1955. Untersuchungen am Thorax von Perla abdominalis Burm. (Larve und
Imago). Zook Jahrb. Anat., 74: 491-570.

Received for publication September 20, 1965

ABBREVIATIONS USED ON THE FIGURES

AT — Anterior tentorial arm
AWN — Anterior wing nerve
Ba:i — Metathoracic basalare
bp — Bristle plate

CoeCon — Circumesophageal connective

Con — Connective

Cv — Cervix

cv — Cervical sclerite

Cxi — Coxa of the prothoracic leg

Cx^ — Coxa of the mesothoracic leg

Cx:; — Coxa of the metathoracic leg

New York Entomological Society

[Vol. LXXIV

dil — Dorsal internal lateral muscle

dim — Dorsal internal median muscle

dlra — Dilator muscle of the abdomen

dlSyr — Dilator muscle of the salivary syringe

dma — Dorsal muscle of the tergal apodeme

DNv — Dorsal nerve

Ful — Mesofurca

F u3 — Metafurca

GI — Prothoracic ganglion

GII — Thoracic-abdominal ganglionic mass

L— Leg

Lb — Labium

le — Lateral external muscle

li — Lateral internal muscle

lis — Lateral intrasegmental muscle

LmNv — Labral nerve

M — Mirror of the sound mechanism

mlb — Muscles of labium

mr — Muscles of the rod

MS — Membranous sac of the sound mechanism
mxb — Maxillary bristle
Op — Operculum

osp — Occlusor muscle of the spiracle

pi — Tymbal muscle plate

PlAi — Prothoracic pleural arm

PlAs — Mesothoracic pleural arm

PlAs — Metathoracic pleural arm

pmdb — Protractor muscle of the mandibular bristle

PT — Posterior tentorial arm

PWN — Posterior wing nerve

r — Rod

rmdb — Retractor muscle of the mandibular bristle

rmxbi — Internal retractor muscle of the maxillary bristle

rmxbs — External retractor muscle of the maxillary bristle

Sai — Subalare of the mesothorax

Sa;! — Subalare of the metathorax

SLD — Salivary duct

SLGL — Salivary gland

SoeGng — Subesophageal ganglion

Sp2 — Mesothoracic spiracle

Spy — Metathoracic spiracle

sp — Fourth abdominal spiracle

Ti — Prothoracic tergum

To — Mesothoracic tergum

Ty — Metathoracic tergum

tapd — Tergal apodeme

TB — Tentorial bridge

tg — Tegula

tn — Tendon between tymbal plate and tymbal
TNv — Transverse nerve

March, 1966 |

Vogel: Spiders from Pennsylvania

Tr — Trachea

tr — Tergal ridge

TYM — Tymbal

Vi — Ventral internal muscles

Vma — -Ventral muscle of the tergal apodeme

Vnv — Ventral nerve

wg — Wing of sclerotized V-shaped structure of first abdominal segment
lPh — First phragma
2Ph — Second phragma

I — First abdominal segment
ISp — First abdominal spiracle

II — Second abdominal segment

lpmxb — Protractor muscle of the maxillary bristle
2pmxb — Protractor muscle of the maxillary bristle

Spiders from Powdermill Nature Reserve

Beatrice R. Vogel

Biology Department, Yale University

Abstract: This paper is a list of 150 species of spiders collected during a 2 -week study of
the fauna of Powdermill Nature Reserve in Pennsylvania.

Local and regional faunal lists are the backbone of ecological and zoogeograph-
ical studies. Such lists provide the detailed information on local faunas necessary
for syntheses of a broader scope. The spider fauna of Pennsylvania has been
sadly neglected. Distribution maps in recent taxonomic revisions almost invari-
ably show a lack of locality records from this state. There has been only one
list of Pennsylvania spiders published during the last half century (Truman,
1942): a list of spiders from Presque Isle, Erie County. This present paper is
also a local faunal list.

The Powdermill Nature Reserve of Carnegie Museum is an area of about
1,500 acres in the Ligonier Valley of Westmoreland County, Pennsylvania. The
reserve includes a variety of woodland and open (chiefly old field) habitats.
This collection was primarily made during a 2 -week study in June and July,
1965, with a few additional specimens collected at other times of the year. This
list must, of necessity, be regarded as preliminary. With the exception of
duplicates retained by the author, the specimens are deposited in Carnegie
Museum.

The collection consists of over 1,000 specimens of mature spiders, representing
about 150 species. The list from Presque Isle also includes about 150 species,
but the two lists have only half their species in common. These lists, along with
scattered reports, bring the published number of Pennsylvania species between
200 and 250. Judging by the fauna of New York state, there should eventually
be more than 500 species of Pennsylvania spiders.

New York Entomological Society

LVol. LXXIV

I wish to thank C. J. Goodnight, of Western Michigan University, for identifi-
cation of the opilionids; W. Ivie, of the American Museum of Natural History,
for identification of the erigonids; and W. J. Gertsch, Curator of Spiders at the
American Museum of Natural History, for identification of Clubiona and for
help with some of the difficult species. I am also indebted to M. Graham
Netting, Director of Carnegie Museum, for making my fieldwork at Powdermill
Nature Reserve possible.

ORDER ARANEIDA

Suborder MY GALOMORPHAE

ANTRODIAETIDAE
Antrodiaetus unicolor (Hentz)

Suborder ARANEAMORPHAE
AMAUROBIIDAE

Amaurobius bennetti (Blackwall)

DICTYNIDAE
Dictyna cruciata Emerton
Dictyna joliacea (Hentz)

Dictyna frondea (Hentz)

Dictyna sublata (Hentz)

Lathy s foxi Marx

ULOBORIDAE
Hyptiotes sp. (immature)

SEGESTERIIDAE
Ariadna bicolor (Hentz)

TETRAGNATHIDAE

Leucauge venusta (Walckenaer)
Mimognatha foxi (McCook)
Pachygnatha autumnalis Keyserling
T etragnatha elongata Walckenaer
T etragnatha straminea Emerton
T etragnatha versicolor Walckenaer

THERIDIIDAE

Achaearanea globosum (Hentz)
Achaearanea tepidariorum (Koch)
Ancylorrhanis hirsutum (Emerton)
Asagena americana Emerton
Crustulina altera Gertsch and Archer
Ctenium frontata (Banks)

Ctenium pumilis (Emerton)

Dipoena nigra (Emerton)
Enoplognatha tecta (Keyserling)
Euryopis funebris (Hentz)

Steatoda borealis (Hentz)

Teutana triangulosa (Walckenaer)
Theridion albidum Banks
Theridion differens Emerton
Theridion frondeum Emerton
Theridion lyricum Walckenaer
Theridion murarium Emerton
Theridion sexpunctatum Emerton
Theridion spirale Emerton
Theridula opulent a Walckenaer

ERIGONIDAE
Ceraticelus bulbosus (Emerton)
Ceraticelus fissiceps (Cambridge)
Ceraticelus laetabilis (Cambridge)
Ceratinopsidis formosa (Banks)
Ceratinopsis inter pres (Cambridge)
Ceratinopsis nigriceps Emerton
Collinsia oxypaederotipus (Crosby)
Cornicularia minuatus Emerton
Cornicularia vigilax (Blackwall)
Eperigone macndat a (Banks)

Erigone autumnalis Emerton
Grammonota trivittata ? Banks
Hypselisthes florens (Cambridge)
Maso sarcocuom (Crosby and Bishop)
Maso sundevalli (Westring)

Origanates rostratus (Emerton)
Scylaceus pallida (Emerton)

LINYPHIIDAE

Bathyphantes albiventris (Banks)
Lepthy phantes subalpina (Emerton)
Linyphia waldea Chamberlin and Ivie
Meioneta fabra (Keyserling)

Pity ohy phantes costatus Hentz
Tennesseellum formica (Emerton)

ARANEIDAE

Araneinae

Acacesia hamata (Hentz)

Acanthepeira stellata (Walckenaer)

March, 1966

Vogel: Spiders from Pennsylvania

Araneus attestor Petrunkevitch
Araneus cornutus Clerck
Araneus marmoreus Clerck
Araneus solitarius (Emerton)

Araneus trifolium (Hentz)

Araniella displicata (Hentz)

Cyclosa conica (Pallos)

Cyclosa turbinata (Walckenaer)

East ala anastera (Walckenaer)

M angora gibber osa (Hentz)

Mastophora bisaccata ? (Emerton) (imma-
ture)

Neoscona arabesca (Walckenaer)

Singa praetensis Emerton
Singa sp. aff. variabilis

Argiopinae
Argiope trifasciata (Forskal)

Gea heptagon (Hentz)

Gasteracanthinae
Micrathena gracilis (Walckenaer)

Micrathena sagittata (Walckenaer)

Theridiosomatinae
Theridiosomma radiosum McCook

AGELENIDAE

Agelenopsis pennsylvanica (Koch)

Cicurina arcuata Keyserling
Cicurina brevis (Emerton)

Cicurina idahoana Chamberlin
Cicurina pallida Keyserling
Coras medicinalis (Hentz)

Wadotes sp. (immature)

HAHNIIDAE

Antistea brunnea ? (Emerton) (immature)
Hahnia cinerea Emerton

OXYOPIDAE
Oxyopes salticus (Hentz)

PISAURIDAE
Dolomedes scriptus Hentz
Dolomedes tenebrosus Hentz
Dolomedes triton sexpunctatus Hentz
Dolomedes urinator Walckenaer
Dolomedes vittatus Walckenaer
Pisaurina brevipes (Emerton)

Pisaurina mira (Walckenaer)

Pisaurina mira var. subinflata

LYCOSIDAE

Arc.tosa virgo (Chamberlin)

Lycosa frondicola Emerton
Lycosa gulosa Walckenaer
Lycosa helluo Walckenaer
Lycosa rabida Walckenaer
Pardosa distinct a (Blackwall)

Pardosa lapidicina Emerton
Pardosa milvina Hentz
Pardosa moesta Banks
Pardosa saxatilis (Hentz)

Pirata insularis Emerton
Pirata maculatus Emerton
Pirata minutus Emerton
Pirata montanus Emerton
Schizocosa avida (Walckenaer)
Schizocosa crassipes (Walckenaer)
Schizocosa saltatrix (Hentz)

GNAPHOSIDAE
Drassyllus fallens Chamberlin
Sosticus insularis (Banks)

Zelotes duplex Chamberlin

ANYPHAENIDAE
Any phaenella saltabunda (Hentz)

CLUBIONIDAE
Castianeira sp. (immature)
Chiracanthium inc.lusum (Hentz)
Clubiona abboti Koch
Clubiona kastoni Gertsch
Clubiona obesa Hentz
Clubionoides pallens (Hentz)

THOMISIDAE

Misuminae

Misumenoides formosipes (Walckenaer)
Misumenops asperatus (Hentz)
Misumenops oblongus (Keyserling)

X ysticus elegans Keyserling
Xysticus ferox (Hentz)

X ysticus f rat emus Banks
Xysticus funestus (Keyserling)

Xysticus triguttatus Keyserling

Philodrominae
Philodromus placidus Banks
Philodromus rufus Walckenaer
Thanatus sp. (immature)

New York Entomological Society

[Vol. LXXIV

Tibellus maritimus (Menge)

Tibellus oblongus (Walckenaer)

SALTICIDAE
Evarcha hoyi (Peckham)
Habrocestum pulex (Hentz)
Habronattus decorus (Balckwall)
Hasarius adansoni (Audouin)
lcius harti Emerton
Maevia incle mens (Walckenaer)
Marpissa lineata (Koch)

Marpissa undata (DeGeer)
Metaphidippus galathea (Walckenaer)
Neon nelli Peckham

Paraphidippus marginata (Walckenaer)
Peckhamia scorpiona (Hentz)
Phidippus clams Keyserling
Phi dip pus prince ps (Peckham)

Phlegra fasciata (Hahn)

Salticus scenicus (Linnaeus)

Sittacus jloridanus Gertsch and Mulaik
Synemosyna formica Hentz
Zygoballus bettini Peckham

ORDER OPILIONIDA
Leiobunum nigropalpi Wood
Leiobunum ventricosum Wood
Leiobunum verrucosum Wood

Literature Cited

Included is a brief list of works for studying Pennsylvania spiders. Many of these con-
tain bibliographies of more specialized papers. Kaston is probably the most useful single
work for identifying species.

Bonnet, P. 1945-1959. Bibliographia Araneorum. Toulouse, 1, 2: i-xvi, 1-832; 1-5058.
Crosby, C. R., and S. C. Bishop. 1928. Araneae. In A list of the Insects of New York.

Cornell Univ. Agr. Exper. Sta., Mem. 101 : 1034-1074.

Kaston, B. J. 1948. Spiders of Connecticut. State Geological and Natural History Survey,
Bull. 70: 1-874.

Levi, H. W. 1957. The spider Genera Enoplognatha , Theridion and Paidisca in America
north of Mexico (Araneae:Theridiidae) . Bull. Amer. Mus. Nat. Hist., 112 (1): 1-123.
Truman, L. C. 1942. A list of spiders collected in western Pennsylvania. Proc. Penn.
Acad. Sci., 16: 25-28.

Received for Publication November 29, 1965

Recent Publications

The Natural History of Mosquitoes. Marston Bates, Harper and Row, New York, $2.45
(paper) 378 pp., 1965.

A Systematic Revision of the Amenidae (Diptera: Calliphoridae) , R. W. Crosskey,

Bull. Brit. Mus. (Nat. Hist.), Entomology, 16: 2, about $5.00, 107 pp., 1965.

The Culicoides of New York State (Diptera: Ceratopogonidae) , Bull. # 399. Hugo Jamn-
back, New York State Museum and Science, $1.00, 154 pp., 24 plates, 1965.

Mierolepidoptera of Juan Fernandez Island. J. F. Gates Clarke, Proc. U. S. Nat. Mu-
seum 117 No. 3508, Smithsonian Institution, Washington, D. C., 106 pp., 1965.

A Revision of the Nodini and A Key to the Genera of Eumolpidae of Africa (Cole-
optera: Eumolidae), B. J. Selman, Bull. Brit. Mus. (Nat. Hist.), Entomology, 16,

No. 2, about $1.90 (paper), 31 pp., 1965.

Review of the Genus Cerceris in America North of Mexico (Hymenoptera: Sphecidae),
Herman A. Scullen, Proc. U. S. Nat. Museum, 116, No. 3506, Smithsonian Institution,
Washington, D. C., 2121 pp., 1965.

Defensive Secretion of a Caterpillar (Papilio). Thomas Eisener, and Yvonne C. Mein-
wald, Science, 150, Dec. 1965, pp. 1733-1738, illus.

March, 1966]

O’Brien: Aedes aegypti

Origin and Structural Function of the Basal Cells of the Larval Midgut
in the Mosquito, Aedes aegypti Linnaeus1

James F. O’Brien2

Biological Laboratories, Fordham University, Bronx, New York 10458

Abstract: This study of a series of midgut whole mounts of larval and pupal Aedes aegypti

shows that basal or regenerative cells first appear as a distinct cell type in the mosquito
midgut at about the ninth hour of larval life. These cells seldom take part in forming the
epithelial lining of the larval midgut. After their appearance, frequent mitotic divisions
occur in the basal cells throughout the larval instars resulting in the presence of a large
number of these cells in the prepupal midgut. During metamorphosis in the pupal stage,
the basal cells remain to form the epithelial layer of the imaginal midgut.

Relatively little is known about the cytological development of the midgut in
the mosquito, Aedes aegypti Linnaeus. Christophers’ (1960) description of the
Aedes digestive tract indicates that the midgut has received little attention from
cytologists. Among the three types of cells comprising the larval midgut of Aedes ,
the regenerative or basal cells remained somewhat of a mystery as to their origin.
Christophers stated that the origin of the basal cells is unknown. Berger (1938)
reported finding regenerative cells in the larval midgut of the mosquito Culex
pipiens but offered no explanation as to their origin. The fairly constant size,
active divisions, and increasingly larger numbers of these cells during the larval
stages indicate that they must perform some function other than to replace
epithelial cells in the larval midgut. The question regarding the origin of the
basal cells as well as the fact that such a large number of these cells is present
in the later larval instars indicated the need for further cytological study of the
midgut of A. aegypti. While the investigation is principally concerned with the
larval midgut, pupal and adult midguts were also studied to determine the
origin and structural function of the basal or regenerative cells.

MATERIALS AND METHODS

The Aedes larvae used in this study were obtained from the colony main-
tained in this laboratory (O’Brien, 1965). Beginning about 6 hours after hatch-
ing and at intervals of from 1 to 4 hours throughout the larval and pupal stages,
the midguts were dissected from the specimens. They were then prepared as
whole mounts, stained with the Feulgen reaction and counterstained with Orange
G. The dissections, the fixation, and the staining procedure were performed on
depression slides to eliminate loss or damage to the tissue (O’Brien, 1965).

1 A portion of the author’s dissertation submitted in partial fulfillment of the require-
ments for the degree of Doctor of Philosophy at Fordham University. The author wishes
to acknowledge the assistance and encouragement given him during this study by Prof.
C. A. Berger, S.J., of the Fordham Biological Laboratory. This work was supported in
part by an Educational Assistance Grant from the Arthur J. Schmitt Foundation.

-Present address: Regis College, Willowdale, Ontario, Canada.

New York Entomological Society

I Vol. LXXIV

Fig. 1. Photomicrograph of portion of stomach area of 17-hour larva showing three
potential regenerative cells (arrows). X 1,290.

RESULTS

In 6-hour larval midguts, only two cell types are present, the longitudinal
and circular rows of muscle cells and larger cells forming the epithelial lining
of the midgut. At about the twelfth hour of larval life, growth of the midgut
has resulted in an increase in size of the epithelial cells, making the epithelial
cells easily distinguishable from the smaller regenerative cells which have ap-
peared by this time. Examination of whole mounts of midguts between 6 and
12 hours old shows that at about 9 hours, some of the original midgut epithelial
cells are undergoing mitotic division. Such a division gives rise to two cells
that are smaller than the neighboring cells. These smaller cells are regenerative
cells. Up to this time, the midgut wall is only two cell layers thick, the outer
cells being the rows of muscle cells and the inner layer the epithelial cells. The
division of some of the initial epithelial cells results in the formation of the
smaller cells that lie on the basement membrane, at the bases of the epithelial
cells — hence the term “basal” cells.

Study of the cells of the midguts obtained from larvae between 6 and 9 hours
old reveals the presence of large, uniformly sized epithelial cells resting on the
basement membrane. A few of these cells exhibit nuclei that appear to be in
early prophase of mitotic division, while the neighboring cells contain normal
“resting” nuclei. Since all the cells are of about the same size, the cells appear-
ing to be in early prophase must be the potential basal cells (Fig. 1). The origin
of the basal cells from epithelial cells was confirmed when mitosis was observed

March, 1966]

O’Brien: Aedes ciegypti

Fig. 2. Photomicrograph of portion of a pouch of gastric ceca of 24-hour larva showing
one of the primordial epithelial cells in mitotic prophase. X 1,290.

in epithelial cells of the gastric ceca (Fig. 2) where regenerative cells appear at
a later stage and in fewer numbers than in the stomach area of the midgut.

All divisions of the basal cells are normal mitotic divisions, exhibiting the
somatic pairing of homologous chromosomes characteristic of dipteran cells
(Figs. 2, 3, 4).

After the basal cells appear in the midgut, their number increases rapidly by
repeated divisions. These cells lie at the bases of the large primordial epithelial
cells which continue to grow larger during the larval instars and never divide
after about 24 hours of larval life. By the fourth instar, the regenerative cells
form almost a complete layer of cells, intermediate in size between the large
primordial epithelial cells and the smaller muscle cells, against the basement
membrane. The number of basal cells found in the gastric ceca is considerably
smaller than in the stomach area of the midgut.

Examination of the pupal midgut shows that the regenerative cells form the
new epithelial lining of the imaginal midgut. During the larval instars, few
of the basal cells help to form the epithelial layer of the midgut. But, after the
onset of pupation, the primordial epithelial cells quickly separate from the
basement membrane, are sloughed off into the lumen of the midgut, and begin
to disintegrate. The basal cells that have been increasing in number throughout
larval life continue their divisions and soon form the epithelial lining of the
imaginal midgut. Since the adult midgut contains no structure similar to the
pouches of the larval gastric ceca, the basal cells that formed in the region of

New York Entomological Society

[ Vol. LXXIV

Fig. 3. Photomicrograph of portion of gastric ceca of 17-hour larva showing a large
epithelial cell in mitotic prophase. X 1,290.

the gastric ceca during the larval instars combine with the cells of the cardiac
region and those of the anterior portion of the stomach area to form the epithe-
lium of the anterior region of the adult midgut.

DISCUSSION

The results of this study indicate the need for revising some statements based
upon earlier findings. Berger ( 1938) reported that the cells comprising the
epithelial lining of the larval mosquito midgut (the “primordial” epithelial cells)
never undergo mitotic division but rather only increase in size during larval life.
The basal cells were thought to function primarily as replacement cells for the
worn-out epithelial cells in the larval midgut. But the findings here presented
show that the early first-instar midgut contains only muscle cells and primordial
epithelial cells and that very few of the basal cells function as replacement cells
in the larval instars. Therefore, it seems that some of the primordial epithelial
cells in the young larva become potential basal or regenerative cells soon after
hatching. Once larval feeding and growth begin, these potential basal cells
cease functioning as epithelial cells, undergo mitotic division, and become basal
cells. In the process of this transformation, their places in the epithelial layer
are taken by the nearby primordial epithelial cells which do not divide, but
rather enlarge to fill the space left in the epithelial lining. Thus, some of the
primordial epithelial cells do undergo division, but only during the early hours
of larval life. The factor determining the time of transformation from primordial
epithelial cells to potential basal cells is not known.

Since so few of the basal cells in the larval midgut function as replacement

March, 1966]

O’Brien: Aedes aegypti

Fig. 4. Photomicrograph of portion of stomach area of 24-hour larva showing a basal
cell in mitotic prophase. X 1,290.

cells in the epithelial coat, the main role of these cells must be to form the
epithelial lining of the imaginal midgut. Therefore, the formation of the adult
midgut does not take place principally in the pupa. Both the muscular coat
for the midgut, basically that present in the prepupa (O’Brien, 1965), and the
epithelial lining of the imaginal midgut, derived from the basal cells of the
larval midgut, have been steadily developing throughout the larval stages.

SUMMARY

Regenerative or basal cells in the larval midgut of A. aegypti first appear
about 9 hours after hatching.

The basal cells generally take no part in forming the epithelial lining of the
larval midgut.

After their appearance in the early larval midgut, the basal cells undergo
frequent mitotic divisions, resulting in the presence of a large number of basal
cells in the prepupa.

Early in the pupal stage, the primordial epithelial cells of the larval midgut
are sloughed off into the midgut lumen and the basal cells remain to form the
epithelial lining of the imaginal midgut.

Literature Cited

Berger, C. A. 1938. Multiplication and reduction of somatic chromosome groups as a
regular developmental process in the mosquito, Culex pipiens. Pub. 496. Carnegie
Institution of Washington.

Christophers, S. R. 1960. Aedes aegypti (L.). Cambridge Univ. Press.

O’Brien, J. F. 1965. Development of the muscular network of the midgut in the larval
stages of the mosquito Aedes aegypti Linnaeus. Jour. N. Y. Ent. Soc. 73(4): 226-231.

New York Entomological Society

[Vol. LXXIV

BOOK REVIEWS

TWO BOOKS FOR YOUNG NATURALISTS

Monarch Butterflies. Alice L. Hopf. Illustrated by Peter Burchard. Thomas Y. Crowell
Company, 1965, 135 pp., price $3.75.

Fireflies in Nature and the Laboratory. Lynn and Gray Poole. Illustrated by Christine
Sapieha. Thomas Y. Crowell Company, 1965, 149 pp., price $3.95.

These two little books are valuable additions to the young naturalist’s library. Mrs. Hopf’s
book will have greater appeal, probably, since it is based on personal experiences and trans-
mits the author’s enthusiasm for field work and dedication to the Monarch Butterfly. The
Poole book brings together a large amount of information about luminescence which might
be difficult for the young person to ferret out by himself. It is of broader scope than its
title implies. Both books will arouse the reader’s interest in getting outdoors and “swinging a
net.”

Monarch Butterflies describes the work that has been done in tagging butterflies for
the purpose of gathering information on flight and migration and offers practical suggestions
for the young collector who would like to cooperate in this scientific study. It discusses
the collecting, rearing, and photographing of Monarchs and gives detailed and practical
suggestions which are applicable to many other species of insects. In touching briefly upon
the studies that have been made to test the Monarch’s protection against predation it
shows the reader how he, too, may make observations which might be of value.

Fireflies includes a discussion of the vocabulary of luminescence and describes luminous
dinoflagellates, annelid worms, molluscs, mycetophilids, bacteria, and fungi. The chapters
about the early research on luminous organisms, and the various authors’ accounts of them,
may not hold the readers’ interest, but those on recent and current work and on the
methods of collecting fireflies, shipping them to scientists, and exchanging them with other
young collectors for natural history specimens from their particular area will certainly elicit
an enthusiastic response. The black and white illustrations are enchanting. Unfortunately
the book is marred by poor editing: scientific names are sometimes italicized, sometimes not;
genus names are sometimes capitalized, sometimes not; and the arachnid daddylonglegs is
called an insect.

The price of these books seems high for text that is scarcely more than magazine article
length. But the books are attractive; they are printed in large, clear type; and they seem to
have been written not just to give information but to encourage young people to be active.

Elsie B. Klots

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JUNE 1966

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> The

New York Entomological Society

Organized June 29, 1892- — Incorporated February 25, 1893

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Reincorporated February 17, 1943

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The meetings of the Society are held on the first and third Tuesday of each month (except
June, July, August and September) at 8 p.m., in the American Museum of Natural
History, 79th St., & Central Park W., New York 24, N. Y.

Annual dues for Active Members, $4.00; including subscription to the Journal, $9.00.

Members of the Society will please remit their annual dues, payable in January, to the
Treasurer. V , ,v 77

7 '7

Officers for the Year 1966

President , Dr. Richard Fredrickson

College of the City of New York 10031

Vice President, Dr. Kumar Krishna

American Museum of Natural History, New York 10024

Secretary, Mrs. Lucy Heineman 115 Central Park West, New York 10023

Assistant Secretary, Mr. Albert Poelzl

230 E. 78th Street, New York 10021

Treasurer , Mr. Raymond Brush

American Museum of Natural History, New York 10024

Assistant Treasurer , Mrs. Patricia Vaurie

4- z'-' r - £

American Museum of Natural History, New York 10024

Trustees

A- X ■ y v* /Xj . N\; .. ' y i/7 j- Iv \ ■ " X/ • »‘i(y ✓ / \

1 Year Term

Dr. Alexander B. Klots Dr. John B. Schmitt

2 Year Term

Mr. Robert Buckbee 1

Dr. Jerome Rozen, Jr.

'-V -V- S'

Mailed June 29, 1966

The Journal of the New York Entomological Society is published quarterly for the Society by Allen Press
Inc., 1041 New Hampshire, Lawrence, Kansas. Second class postage paid at Lawrence, /Kansas.

\ -

7 X

Journal of the

New York Entomological Society

Volume LXXIV June 29, 1966

No. 2

EDITORIAL BOARD

Editor Emeritus Harry B. Weiss

Editor Lucy W. Clausen
Columbia University College of Pharmacy
115 West 68th Street, New York, N. Y. 10023

Associate Editor James Forbes
Fordham University, New York, N.Y. 10458

Publication Committee

Dr. Pedro Wygodzinsky Dr. Asher Treat

Dr. David Miller

CONTENTS

Undescribed Species of Crane Flies from the Himalaya Mountains (Diptera:
Tipulidae), XII Charles P. Alexander 66

Notes on the Biology of Stelis ( Odontostelis ) bilineolata (Spinola), a Parasite
of Euglossa cordata (Linnaeus) (Hymenoptera : Apoidea: Megachilidae)

Frederick D. Bennett 72

Further Studies on the Internal Anatomy of the Meloidae (Coleoptera) . II.

The Digestive and Reproductive Systems of the S.A. Blister Beetle, Picnoseus
nitidipennis Fairmaire and Germain (Coleoptera: Meloidae) A. P. Gupta 80

Taxonomic Descriptions of the Immature Stages of the Parasitic Bee Stelis
( Odontostelis ) bilineolata (Spinola) (Hymenoptera: Apoidea: Megachilidae)

Jerome G. Rozen, Jr. 84

Mature Larvae of the Old World Bee Genus Panurgus (Hymenoptera: Apoidea)

Jerome G. Rozen, Jr. and Barbara L. Rozen 92

Melanism in Connecticut Panthea furcilla (Packard) (Lepidoptera : Noctuidae)

Alexander B. Klots 95

An Apparent Association of Mites (Acarina) with the Rock Barnacle Balanus

Richard W. Fredrickson 101

Bylaws of the New York Entomological Society

Book Reviews

Notes — Help for Ailing Caterpillars?

Membership of the New York Entomological Society

Recent Publications

Proceedings

Necrology

Invitation to Membership

103

109

Alice L. Hopf 111

112

116

117

122

123

New York Entomological Society

I Vol. LXXIV

Undescribetl Species of Crane Flies From the Himalaya Mountains

(Diptera: Tipulidae) , XII1

Charles P. Alexander
Amherst, Massachusetts

Abstract: Six new species of the Eriopterine genera Ormosia and Erioptera are described,
including Ormosia ( Oreophila ) licina n. sp., from Kashmir and Kumaon, and Ormosia
( Par ormosia ) atrotibialis n. sp., Ormosia ( Ormosia ) subpulehra n. sp., and 0. (0.) um-
bripennis n. sp., from Sikkim: Erioptera ( llisia ) diadexia n. sp. and E. (I.) epicharis
n. sp., from Sikkim.

Part XI of this series of papers was published in the Journal of the New York
Entomological Society, 73: 163-167, 1965. The materials upon which the new
species are based were collected by Dr. Fernand Schmid, of Ottawa, to whom
1 express my deepest appreciation for this outstanding series of Asiatic Tipulidae.

Doctor Schmid collected insect specimens in India and adjoining countries
between 1953 and 1961 as a member of the Swiss Zoological Expedition. His
insect collections were restricted to certain groups, where they proved to be of
paramount importance in making known the exceedingly rich fauna of the
region. A summary of the stations visited, as they pertain to the crane flies, is
given in a paper by the writer (Philippine Jour. Sci., 90: 163; 1961), covering
the period between 1953 and 1960. Between February and October, 1961,
still further collections were made by Doctor Schmid in the Kameng Frontier
Division of the North East Frontier Agency (NEFA), Assam.

In the crane fly materials several hundred new species were included that
have been discussed and presently are being described in a long series of papers
that are summarized herewith in order to assist other students of the subject:

Philippine Jour. Sci. (chiefly Tipulinae and Limoniini)

Ann. and Mag. Nat. Hist. (London) (chiefly Tipulinae and Eriopterini)

Proc. Royal Ent. Soc. London (Pediciini)

Trans. Royal Ent. Soc. London (Hexatomini; Phyllolabis)

Bull. Brooklyn Ent. Soc. (Tanyderidae, Ptychopteridae, Trichoceridae)

Jour. N. Y. Ent. Soc. (chiefly Eriopterini)

Ent. News (Hexatomini)

Trans. Amer. Ent. Soc. (Eriopterini)

Ormosia ( Oreophila ) licina n. sp.

General coloration of mesonotum light brown, sparsely pruinose, pleura yellow; antennae
moderately long; wings yellowed, very restrictedly patterned with pale brown, vein 2nd A
sinuous; male hypopygium with the outer dististyle black, coarsely spinulose; lateral margins
of gonapophyses produced into two or three acute points.

1 Contribution from the Entomological Laboratory, University of Massachusetts.

June, 1966 I

Alexander: Himalayan Crane Flies, XII

male: Length about 4.8-5 mm; wing 5.8-6 mm; antenna about 1.5-1 .6 mm.
female: Length about 5 mm; wing 6 mm.

Rostrum light yellow; palpi pale brown. Antennae of male moderately long, scape obscure
yellow, the remainder black; flagellar segments oval to long-oval, shorter than their verticils.
Head light gray.

Thorax light brown, sparsely pruinose, pronotum more yellowed, pretergites clear yellow.
Pleura and lateral prescutal borders light yellow. Halteres yellow. Legs with coxae and
trochanters yellow; remainder of legs yellowish brown, femora more yellowed basally,
tarsi brownish black. Wings yellowed, very restrictedly patterned with pale brown, includ-
ing the stigma, cord, outer end of cell 1st M2 , and small spots at Sc2 and origin of Rs. Vena-
tion: Sci ending nearly opposite the fork of R2+ s+i, Sc2 far retracted, about opposite one-third
to two-fifths Rs ; vein R» close to fork of R2+-m', cell 1st M» elongate, subequal to distal
section of M ]+2; vein 2nd A sinuous. One wing of the holotype has cell M2 open by atrophy
of the basal section of Ms.

Abdominal tergites brown, basal sternites paler. Male hypopygium with the dististyles
slightly subterminal, broadly united basally; outer style blackened, relatively short and
stout, the outer half with numerous strong spinules; inner style pale, relatively short, the
outer margin with a strong lobe before midlength. Phallosome with lateral margins of
outer apophyses produced into two or three acute points; aedeagus short, black.

holotype 3, Dakwani, Pauri Garhwal, Kumaon, 9,300-11,000 feet, August 5,
1958 (Schmid). Allotopotype, 9, pinned with type. Paratypes, 8 9, Gangrea,
Pauri Garhwal, 7,500-10,000 feet, June 12, 1958; 13, Tales, Kashmir, August
13, 1954 (Schmid).

Ormosia ( Oreophila ) licina is most similar to O. (O.) hutchinsonae Alex-
ander, which differs in the coloration, venation, as the short straight vein 2nd A,
and in the structure of the hypopygium, especially the phallosome and the
elongate dististyle.

Ormosia ( Parormosia ) atrotibialis n. sp.

Generally similar and closely allied to Ormosia ( Parormosia ) leucoplagia Alexander,
differing in the coloration of the legs in the male.
male: Length about 4.5-5 mm; wing 5. 2-5. 6 mm.
female: Length about 5 mm; wing 5.8 mm.

Antennae light yellow, in cases with the intermediate flagellar segments bicolored, their
bases narrowly dark brown, the outer two-thirds to three-fourths yellow. Mesonotal
prescutum obscure yellow with a more or less distinct capillary brownish black median line ;
scutum brown, scutellum and postnotum darker. Legs black in both sexes, the tips of the
femora narrowly yellow, including about the outer tenth of segment, extreme tibial bases
more narrowly yellowed. In leucoplagia the tibiae and basitarsi of male yellow, of the
female black, as in the present fly.

holotype 8, Lachen, Sikkim, 8,900 feet, June 13, 1959 (Schmid). Allotopotype,
9 . Paratopotypes, 48 9; paratypes, 1 8, 2 9 9 , Lachung Sikkim, 8,610 feet, July
2, 1959 (Schmid).

Ormosia ( Ormosia ) subpulolira n. sp.

Allied to puchra ; general coloration of thorax gray, prescutum with a broad brown

New York Entomological Society

[Vol. LXXIV

central stripe, humeral region yellowed; femora yellow with two subequal broad brownish
black rings, the outer one nearly apical ; wings whitened, with conspicuous pale brown
clouds; male hypopygium with both dististvles extended into acute blackened points; gona-
pophvses appearing as a massive black triangular head, its outer margin with three strong
spines.

male: Length about 4.5 mm; wing 5.3 mm.

Head broken. Pronotal scutum dark brownish gray, scutellum testaceous yellow. Mesonotal
prescutum gray, with a broad brown central stripe that is narrowly darker medially ; humeral
region, including the pseudosutural foveae, yellow, tuberculate pits very reduced; posterior
sclerites of notum dark brown, sparsely pruinose. Pleura dark gray, the yellow setae of the
posterior pteropleurite very long. Halteres broken. Legs with coxae dark brown; trochanters
obscure yellow; femora yellow, each with two subequal broad brownish black rings that
are about equal to the pale base or intervening interspace, the tip narrowly yellow; re-
mainder of legs light brown. Wings with the ground color whitened, with conspicuous pale
brown clouds chiefly in the outer three-fourths, stigma darker; whitened marginal spots in
cells R>, R,, and Ri, less evident in cells R5 and 2nd Mz, larger in cells Ms, Cu and the anals;
cells basad of cord more extensively whitened; veins brown, prearcular field and Sc, R, and
Cu more yellowed. Venation: R2 at fork of R2+ 3+4; cell 2nd M2 square at base; vein 2nd A
strongly sinuous, close to border on outer end.

Abdomen, including hypopygium, brownish black. Male hypopygium with apical end
of tergite short and broad, the lobes low. Both dististyles extended into acute blackened
points. Gonapophysis appearing as a massive blackened triangular head, the basal stem
relatively slender, outer margin with three strong spines, with a further series of about four
microscopic denticles on lower margin near the stem.

holotype a broken S , mounted on microscope slide, Zema, Sikkim, 9,100 feet,
June 14, 1959 (Schmid).

Ormosia ( Ormosia ) siibpulchra is related to O. (O.) kashrniri Alexander and
O. (O.) pulchra (Brunetti), all three species differing among themselves chiefly
in important characters of the male hypopygium.

Ormosia ( Ormosia ) umbripennis n. sp.

General coloration of head and thorax brownish black; palpi, antennae, halteres, and legs
black; wings strongly infuscated; Sc2 beyond midlength of Rs, cell 1st M2 shorter than vein
Mi, vein 2nd A gently sinuous.

female: Length about 6 mm; wing 6.5 mm; antenna about 1.6 mm.

Rostrum and palpi black. Antennae black throughout ; flagellar segments long-oval, with
dense white setae, the verticils longer than the segments. Head brownish black.

Thorax uniformly very dark brown to brownish black, the surface of mesonotum subniti-
dous; prescutum and scutellum with a few long setae. Halteres brownish black, base of
stem obscure yellow. Legs black. Wings strongly infuscated, especially the prearcular and
costal fields and the stigma; veins brown. Venation: ending opposite the oblique R2,

Sc2 moderately retracted, about opposite three-fifths the long Rs; R2+ 3+i shorter than basal
section of R:,; cell 1st M-> shorter than vein Ah; m-cu at fork of M, perpendicular and
slightly sinuous; vein 2nd A gently wavy.

Abdomen brown, the outer segments more blackened. Ovipositor with cerci horn yellow,
long and slender, gently upcurved to the acute tips.

June, 1966]

Alexander: Himalayan Crane Flies, XII

holotype 9, Namnasa, Sikkim, 10,000 feet, July 1, 1959 (Schmid).

The only other generally similar regional species is Ormosia ( Ormosia ) nyc-
topoda Alexander, of Pakistan, which similarly has the legs black but with the
wings pale and having the venational details distinct.

Erioptera ( Ilisia ) diadexia n. sp.

Allied to asymmetrical general coloration of thorax gray, the prescutum with two diffuse
brown stripes; antennae black; femora brownish yellow, tips brownish black; wings brownish
yellow, conspicuously patterned with brown spots and dots, the latter on all veins excepting
Sc and Cu ; male hypopygium with the outer dististyle bilobed, inner style broad, yellow,
and the tip very obtuse; gonapophyses with the two arms virtually identical, appearing as
straight blackened rods, the tip microscopically toothed.
male: Length about 5-6.5 mm; wing 5.8-8 mm.

Rostrum gray, palpi black. Antennae relatively long, black; flagellar segments long-oval
to fusiform, basal segments with long verticils, all with further dense pale setulae. Head
brownish gray.

Prothorax brownish gray; anterior pretergites obscure yellow. Mesonotal prescutum
brownish gray, the interspaces more infuscated to form two diffuse stripes; posterior
sclerites of notum brownish gray, central area of scutum narrowly brown. Pleura brownish
gray. Halteres with stem yellow, knob weakly infuscated. Legs with coxae brownish gray ;
trochanters brownish yellow ; femoral and tibiae brownish yellow, tips brownish black, the
tibiae slightly enlarged and darkened beyond bases; basitarsi light brown, remainder of
tarsi black. Wings brownish yellow, conspicuously patterned with brown spots and dots,
the former including about five costal areas, the second over Sc2, the third largest, over tip
of Sci and R2, fourth area at tip of i?i+2; smaller marginal spots at ends of all longitudinal
veins; narrow brown seams over cord, m, arculus, and at near midlength of Cu\ ; paler brown
spots on all longitudinal veins excepting Sc and Cu, those basad of cord paler; veins yellow
in the ground areas, brown in the patterned markings. Venation: R>+ 3+<t about twice

R ’+s, Ri+2 nearly as long as Rs ; m-cu far before fork of M ; vein 2nd A straight.

Abdomen brownish black. Male hypopygium with inner apical angle of basistyle pro-
duced, with very long setae. Dististyles subterminal, the outer style blackened, bilobed, the
outer lobe a slender paddlelike blade, its tip obtuse, outer end with delicate setae, inner arm
a shorter blade that is dilated outwardly, apex broadly obtuse to truncate ; inner style pale,
broadly flattened, apex very obtuse to bluntly triangular, surface with long yellow setae.
Gonapophysis with the two arms virtually identical in length and diameter, appearing as
straight blackened rods, the tips microscopically toothed.

holotype $ , Chachu, Sikkim, 11,500 feet, June 29, 1959 (Schmid). Paratype,
S , Darkot, Kashmir, 8,900 feet, August 17, 1954 (Schmid).

The most similar described regional species is Erioptera ( Ilisia ) jansta
Alexander, which is generally similar in coloration of the body and wings, dif-
fering most evidently in the hypopygial structure, including the trilobed outer
dististyle, slender arcuated inner style, and the unequal arms of the gonapophysis.
The paratype from Kashmir is much smaller (the smallest measurements given)
but the hypopygium is so similar to that of the type that I regard it as being
conspecific.

New York Entomological Society

[ Vol. LXXIV

Erioptera ( Ilisia ) epicharis n. sp.

Allied to a symmetrica] general coloration of thorax brownish gray, the prescutum faintly
patterned with darker; halteres yellow; femora darkened, tips brownish black, preceded
by a yellow ring; wings whitish yellow with a conspicuous brown pattern, including large
costal darkenings and paler brown areas in the anal field, the discal areas restricted; male
hypopygium with the outer dististyle trilobed, the inner oval blade with a blackened spur
at base, inner dististyle extended into a point at apex; gonapophysis unequally bifid.
male: Length about 6.5 mm; wing 7.5 mm.
female: Length about 6.5 mm; wing 8 mm.

Rostrum brownish gray; palpi black. Antennae relatively long, brownish black, the bases
of proximal flagellar segments narrowly paler; segments elongate, a little shorter than the
verticils. Head brownish gray.

Pronotum brownish gray, scutum darker laterally, sides of scutellum yellow. Mesonotum
brownish gray, the prescutum with a poorly indicated pale brown stripe, narrowly darkened
in front and on the sides behind; pseudosutural foveae and tuberculate pits black, shiny.
Pleura gray. Halteres pale yellow. Legs with coxae and trochanters pale brown; femora
light brown to brownish black, tips broadly brownish black, preceded by a broad yellow
ring; tibiae and tarsi pale brown. Wings very pale whitish yellow, clearer yellow in the
costal interspaces; a conspicuous brown pattern that is chiefly marginal in distribution,
including six darker costal areas that are more extensive than the interspaces, the larger
markings at origin of Rs and over tip of Sci, the last area at the wing tip; cubital and anal
fields with comparable large paler brown markings, most extensive in the anal cells, small
brown marginal spots on veins M» through Mi narrow darker brown seams over cord, outer
end of cell 1st M2, and isolated at near midlength of vein Cu, with small dots along vein R5]
veins light yellow in the ground, dark brown in the patterned areas, in cases including series
of four or five dashes. Venation: Sci ending shortly beyond R2] m transverse, about one-
third to one-half the basal section of Ms] m-cu before fork of M; vein 2nd A nearly
straight.

Abdomen dark brown, including the hypopygium. Ovipositor with valves light horn yellow.
Male hypopygium with posterior border of tergite nearly truncate, at midregion with two
small paired darkened lobes, separated by a tiny V-shaped emargination, densely set with
microscopic spicules. Apex of basistyle produced beyond insertion of the dististyles. Outer
dististyle trilobed, the outer blade longer, pale yellow, with abundant very delicate pale
setae, those at apex longer; inner arm including an oval to subcircular darkened blade with
a blackened spur at its base; inner style yellow, the lower apical angle produced into a
point, surface with pale setae, some of the outer ones very long. Phallosome including
bifid gonapophyses, the lateral arm an erect blackened rod, its margin nearly smooth in the
holotype, microscopically roughened in the Kumaon paratype; inner arm much smaller,
at apex dilated into a triangular head ; aedeagus appearing as two short slightly divergent
spines.

holotype 3, Yagtang, Sikkim, in Rhododendron association, 11,200 feet, May
28, 1959 (Schmid). Allotopotype, 9. Paratopotype, 9, with the types, 11,600
feet, June 17, 1959; paratypes, 8 9, Chachu, Sikkim, 11,500 feet, June 29, 1959;
Chamiteng, Sikkim, 9,900 feet, August 24, 1959; Gey, Sikkim, in Rhododendron
association, May 18, 1959; Lachung, 8,610 feet, July 10, 1959; Namnasa,
Sikkim, 9,500 feet, July 13, 1959; Talam, Sikkim, in Rhododendron association,
11,300 feet, June 16, 1959; Tsomgo, Sikkim, in Rhododendron association,

June, 1966]

Alexander: Himalayan Crane Flies, XII

12,500 feet, August 26, 1959; Dakwani, Pauri GarhVval, Kumaon, 9,300-11,000
feet, August 5, 1959; Kanol, Pauri Garhwal, 8,530 feet, August 19, 1958;
Kulara, Pauri Garhwal, 12,000 feet, August 4, 1958 (Schmid).

Erioptera ( Ilisia ) epicharis is quite distinct from the other known regional
species of the subgenus, including E. (/.) asymmetrica Alexander ( indica Senior-
White), E. (/.) diadexia n. sp., and E. (I.) jausta Alexander, especially in the
wing pattern and hypopygial structure. One male paratype from Tsomgo is
smaller (length about 5 mm; wing 5.2 mm) and has the femora almost uniformly
darkened but from the wing pattern and hypopygium evidently pertains to
this species.

Received for Publication October 7, 1965

New York Entomological Society

[Vol. LXXIV

Notes on the Biology of Stelis ( Odontostelis ) bilineolata
(Spinola), a Parasite of Euglossa cordata (Linnaeus)
(Hymenoptera: Apoidea: Megacliilidae)

Frederick D. Bennett1

Abstract: The activities of the parasitic bee Stelis ( Odontostelis ) bilineolata (Spinola) in
specially constructed box nests of its host Euglossa cordata (Linnaeus) are reported. The
female enters the nest, forces the attendant Euglossa female to abandon the nest, and remains
in the nest for several days. She opens those cells containing eggs or small larvae, seals, re-
moves, and destroys them and after depositing her own egg reseals the cell. Cells with older
stages of Euglossa are not opened but the larva or pupa contained therein is killed. Feeding
behavior of the small and large larvae and construction of the cocoon are described.

During studies on the biology of Euglossa spp. in Trinidad some of the obser-
vation nests were invaded by the parasitic bee Stelis ( Odontostelis ) bilineolata
(Spinola). It is planned to publish the results of the Euglossa studies separately
when they are completed but it seems advisable at this time to publish a note
on the activities of its parasite as a companion paper to one on the morphology
of the immature stages by Rozen (1966).

nesting habits of Euglossa cordata

To explain the behavior of S. bilineolata it is necessary to describe briefly the
nesting habits of Euglossa. Two species of this genus, E. cordata and E. vari-
abilis (Friese),2 which are solitary species, have been induced to nest in small
wooden boxes (inside dimensions 10 X 6 X 4.5 cm) with a 10-mm circular
entrance hole on one side. Once a nest is occupied and cell construction started
the wooden top can be replaced by a pane of glass through which activities within
the nest can be readily observed.

The bee cements the glass lid to the wood with a brown resinlike plant material
which is used to seal all cracks and joints on the inside of the box and also
to close the entrance hole. When leaving the nest the female opens a smaller
circular hole in the resin just large enough for passage. She never closes the
entrance when leaving the nest, even when leaving for the last time, but always
seals it in the evening and frequently during the day while working inside. Cells
of the same resinlike material are constructed either on the floor of the box or
on the side, usually each cell being provisioned and sealed before another is
started.

1 Entomologist-in-Charge, W. I. Station, Commonwealth Institute of Biological Control,
Curepe, Trinidad, West Indies.

2 Although all observations were in nests of E. cordata the parasite has also been reared
from nests of E. variabilis .

June, 1966]

Bennett: Stelis and Euglossa

OBSERVATIONS ON THE BEHAVIOR OF Stelis

When a Stelis adult was first noted in observation box No. 33 on July 12, 1964
neither its identity nor the significance of its presence was immediately appre-
ciated. This nest which had been first occupied by Euglossa in April contained
nine cells, some with immature stages, others from which adults had emerged,
and a tenth cell only partially constructed and partially provisioned. Work on
this cell stopped about July 4. It is probable that the bee was attacked by a
conopid and, although still capable of flight, her ovaries had ceased to function.
She was present in the nest with the intruder and on the bottom of the box there
was also a small Euglossa larva. The Stelis frequently approached and nudged
the Euglossa female with her mandibles; the latter kept turning away and
finally retreated to a corner of the box. During the next 20 minutes the Stelis
female spent most of the time examining the cell mass.

At 6:45 A.M. the following morning both adults were in the nest, the Euglossa
in a corner and the Stelis on the cell mass. By 7:45 A.M. the Euglossa went
out and the Stelis began to close the entrance from the inside. The larva on
the floor was dead and partially covered with wax.

The box was examined from time to time on succeeding days and the presence
or absence of the two bees noted. This information is summarized in Table 1.
Although the Stelis female was frequently on the cell mass she did not open any
of the cells during the periods of observation. The walls of two cells from
which Euglossa adults had emerged a few days earlier were partially broken
down by the Euglossa while the Stelis was either resting on the the side of the
box or absent from the nest.

No further activity occurred in this nest when (due to my absence from
Trinidad) observations were suspended on October 3. When I next examined the
nest on December 6 a Stelis adult had emerged from one cell. The other cells
when opened later contained dead pupae and mature larvae of Euglossa.

When observations on the box nests were continued in December it was evident
that a number of other nests had been attacked, i.e., sealed cells present, no
attendant Euglossa adult but the entrance sealed. This was confirmed later
when Stelis adults emerged from one or more cells in these boxes.

On December 6, nest 12, which was started on September 30 but still retained
its wooden cover, contained a dead Euglossa adult (parasitized by a conopid) ;
a Stelis adult; three sealed cells with somewhat flattened walls, and a fourth
partially completed cell. When examined on December 8 the Stelis was absent
(an adult, possibly the same one, was present in nest 13) but was in again on
December 9 and 10. On December 11 the wooden top was replaced by a sheet of
glass and by December 12 the Stelis utilizing bits of resin present in the box
had sealed the glass in place in a manner similar to Euglossa. Although out

The boxes were numbered serially as they were occupied by Euglossa.

New York Entomological Society

[Vol. LXXIV

Table 1. Records of the presence of Euglossa and Stelis in the nest during periods of

observation.

Date and time of
observation

Euglossa

Stelis

Entrance hole

July 12

7:30

A.M.

Closed

July 13

6:45

7:45

Out

Being closed

12:10

P.M.

Out

Closed

1:20