GENERA OF AMERICAN CICADAS NORTH OF MEXICO
MAXINE SHOEMAKER HEATH
A DISSERTATION PRESENTED TO THE GRADUATE COUNCIL OF
THE UNIVERSITY OF FLORIDA
IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE
DEGREE OF DOCTOR OF PHILOSOPHY
UNIVERSITY OF FLORIDA
To my family,
Cindy, Pam, Jessica
Many people have generously contributed time, advice, and information
useful in the preparation of this dissertation. I would like to thank
Dr. Thomas Moore of the University of Michigan, Dr. George Byers, Dr.
Peter Ashlock, and Dr. Charles Michner of the University of Kansas, Dr.
Frank Hasbrouck and Mr. Martin Kolner of Arizona State University, Dr. and
Mme. Michel Boulard of the Mus4um National d'Histoire Naturelle, Dr. T.
Knight and Dr. Peter Broomfield of the British Museum (Natural History),
Mr. Donald Webb of the Illinois Natural History Survey, Dr. Frank Mead of
the Florida Division of Plant Industry, and Dr. Jack Cranford of Virginia
Polytechnic Institute for their interest, information, access to col-
lections, and loans and gifts of cicada specimens. Mrs. Alice Prickett
assisted me in the planning of the illustrations. I thank my committee,
Professors Reece Sailer, Harvey Cromroy, Lewis Berner, Ellis Matheny, and
Frederick King, for continued advice and encouragement. Most of all I
would like to thank my family for assistance and sacrifices, especially
my husband, James E. Heath, who introduced me to the study of cicadas,
and whose continued help and encouragement made this dissertation possible.
TABLE OF CONTENTS
ACKNOWLEDGEMENTS. . . . . . .
ABSTRACT . . . . . . .
I INTRODUCTION . . . . .
Life Cycle . . . . .
The Problem. . . . . .
II MATERIALS AND METHODS. . .
III CLASSIFICATION: HIGHER TAXA .
Order . . . . . .
Suborder . . . . . .
Superfamily . . . .
Family Cicadidae . . . .
Subfamilies . . . .
Tribes . . . . . .
IV CLASSIFICATION: GENERA .
V GENERAL MORPHOLOGY . . .
General Body Proportions
Head . . . . .
Thorax . . . . .
Legs . . . . .
Wings . . . .
Abdomen . . . .
Sexual Characteristics .
Internal Structure Reveal
. . . . . . iii
. . . . . . .. viii
. . . . . .
. . . . . .
. . . . . .
. . . . . .
. . . . . .
. . . . . .
. . . . . .
ed by Radiographs .
VI KEYS TO GENERA . . . . . . . . ... ... 61
Males. . . . . . . . . ... . . . 61
Females. . . . . . . . . ... . . . 63
VII Becaeria . . . . . . . . ... . . . 65
Taxonomy . . . . . . . . ... . . . 63
Description. . . . . . . . . ... . . 66
Distribution . . . . . . . . . . . 70
Discussion . . . . . . . . . . 70
VIII Cacama . . . . . . . ... . . . 71
Taxonomy . . . . . . . .... . . . 71
Description. . . . . . . . . . . ... 72
Distribution . . . . . . . . . . . 76
Discussion . . . . . . . . . . . 76
IX Cicadetta . . . . . . . . . . .. 79
Taxonomy. . . . . . . . .. ..... . 79
Description . . . . . . . .... . ... 81
Distribution . . . . . . . . ... . 86
Discussion . . . . . . . . . . . 87
X Clidoph eps. . . . . . . . . . .. 39
Taxonomy. . . . . . . . .. . . . 89
Description. . . . . . . . . . . ... 90
Distribution . . . . . . . .. . . 94
Discussion . . . . . . . . . . . 95
XI Cornuptura . . . . . . . . . . . 97
Taxonomy . . . . . . . .. . . . 97
Description. . . . . . . . . . . . 98
Distribution . . . . . . . . . . 102
Discussion . . . . . . . . . . . 102
XII Diceroprocta . . . . . . . . . . 104
Taxonomy . . . . . . . . ... . . . 104
Description. . . . . . . . . ... . . 105
Distribution . . . . . . . . . . . .10
Discussion . . . . . . . . . . . 111
XIII Magicicada . . . . . . . . . . . 114
Taxonomy . . . . . . . .. . . . 114
Description. . . . . . . . . .. . . 115
Distribution .... . . . . . . . 119
Discussion . . . . . . . . . . . 120
XIV Neocicada. .... . . . . . . . 125
Taxonomy . . . . . . . . . . . 125
Description. . . . . . . . . . . . 126
Distribution . . . . . . . .... . .. 131
Discussion . . . . . . . . . . . 131
XV Neoplatypedia. . . . . . . . . . .. 134
Taxonomy . . . . . . . . . . . .. 134
Description. . . . . . . . . . . .. 135
Distribution . . . . . . . . . . .. 137
Discussion . . . . . . . . . . .. 140
XVI Okanag . . . . . . . . . . .. 142
Taxonomy . . . . . . . . . . . .. 142
Description. . . . . . . . . . . .. 144
Distribution . . . . . . . . . . .. 149
Discussion . . . . . . . . . . .. 149
XVII Okanagodes . . . . . . . . . . .. 152
Taxonomy . . . . . . . . . . . .. 152
Description. . . . . . . . . . . .. 153
Distribution . . . . . . . . . . .. 157
Discussion . . . . . . . .. . . . 137
XVIII Pacar~na . . . . . . . . . . . .. 159
Taxonomy . . . . . . . . . . . .. 159
Description. . . . . . . . . . . .. 159
Distribution . . . . . . . . . . . 164
Discussion . . . . . . . .. .. . . 164
XIX Platypedia . . . . . . . . ... . .165
Taxonomy . . . . . . . . . . .... 165
Description . . . . . . . . . 166
Distribution . . . . . . . . . . .. 172
Discussion . . . . . . . . . . . 172
XX Q tesada . . . . . . . . . ...... 176
Taxonomy . . . . . . . . . . . .. 176
Description. . . . . . . . . . . .. 177
Distribution . . . . . . . . . . .. 181
Discussion . . . . . . . . . . .. 181
XXI Tibicen . . . . . . . . . . . 133
Taxonomy . . . . . . . .. . . . 183
Description. . . . . . . . . .. . . 186
Distribution . . . . . . . . . . .. 190
Discussion . . . . . . . .. .. . . 192
XXII T2ici noide . . . . . . . . . . . 195
Taxonomy . . . . . . . . . . . 195
Description. . . . . . . . . .. . . 196
Distribution . . . . .... . . . . . 200
Discussion . . . . . . . .. . . . 200
XXIII DISCUSSION AND CONCLUSIONS . . . . . . ... 202
Origins and Dispersal. . . . . . . . .. 202
Ecology and Distribution . . . . . . . .. 207
Evolution of Higher Taxa . . . . . . . . 209
Evolution and Morphology . . . . . . . .. 210
Summary. . . . . . ... . . . . . 217
LITERATURE CITED. . . . . . . . . . . . ... 219
BIOGRAPHICAL SKETCH . . . . . .... . . . . 231
Abstract of Dissertation Presented to the Graduate Council
of the University of Florida in Partial Fulfillment of the Requirements
for the Degree of Doctor of Philosophy
GENERA OF AMERICAN CICADAS NORTH OF MEXICO
Maxine Shoemaker Heath
Chairman: Reece I. Sailer
Co-chairman: Harvey L. Cromroy
Major Department: Entomology and Nematology
Cicadas are large, vocal insects, often heard, but seldom seen.
Sixteen endemic genera occur in the Nearctic biogeographic region north
of Mexico. The genera are Beameria, Cacama, Cicadetta, Clidophleps,
Cornuptura, Diceroprocta, Magicicada, Neocicada, Neoplatypedia, Okanagana,
Okanagodes, Pacarina, Platypedia, Quesada, Tibicen,and Tibicinoides.
Each of these genera is redescribed in a consistent format, thereby
facilitating comparisons of common characters. Keys to males and females
are included, and diagnostic characters are given for each genus.
Radiographs, made with soft x-rays, are used as a taxonomic technique to
elucidate internal structures. The following revisions are recognized:
Okanagana rubrocaudata, a former variety of 0. bella is raised to species
rank; Okanagana rotundifrons is transferred to the genus Clidophleps;
Tibicinoides hesperia is moved back to Okanagana; Cicada chisos is placed
in the genus Neocicada. A new tribe, Okanaganini, is proposed for six
western genera with the possibility of including two palearctic genera.
This tribe has two subtribes, Platypedaria and Okanaganaria.
The cicada fauna in North America is old. Earliest fossils date
from the Paleocene. Current distribution is dependent on origin,
dispersal patterns, habitat, and past geologic history. Two major
radiations of genera have occurred on the North American continent,
one allied to Tibicen, the other to Okanagana. Ancestral and derived
characters are proposed and related to the North American genera.
Cicadas undoubtedly intrigued the imagination of man before recorded
history. A frequently related ancient observation noted how lucky the
male cicada is because his mate is mute. The following pages will show
that even among cicadas some females have managed to develop a "voice."
Myers (1929), in his book Insect Singers, reviewed both the classical and
modern literature, summarizing what was known about cicada biology at
The first written references to cicadas, "tettix" in Greek, can be
found in those sections of the Iliad attributed to Homer and written
around 1,000 B.C. Another poet from the same era, Hesiod, noted that the
sound comes from beneath the wings, an observation made and lost many
times before sound production organs in cicadas were identified and de-
scribed. The ancient Greeks apparently found the song of the cicada
pleasing and musical. In Greek mythology, cicadas were formerly men who
were so enchanted when the Muses brought music to the earth that they
became absorbed by the music, neglected to eat and sleep, and eventually
wasted away. Their voices may still be heard in the singing of cicadas.
In contrast the ancient Romans found the sound of the cicada unpleasant
and raucous. Roman mythology tells of the goddess Aurora, who fell in
love with a mortal. She asked the gods to grant her lover the gift of
immortality, but in her enthusiasm she neglected to ask for eternal youth
for him. As the years passed Tithonus, her lover, became toothless and
infirm and prayed for death. However, the gift of the gods could not be
returned, so Aurora changed her toothless lover into "the ever complaining
cicada." I prefer to think that these differences in the perception of
cicada song relate to a taxonomic difference in cicadas rather than a
difference in aural perception of ancient Greeks and Romans.
From the fables of Aesop, the stories of ancient India, and the folk-
lore of the Maori tribes of New Zealand come amazingly similar tales
contrasting the industrious ant at the foot of a tree with the lazy
cicada singing in the sun at the top of its crown. When winter came the
ant was well fed and survived, while the cicada begged for food and
eventually starved. The Zuni Indians of the American Soutwest attribute
more cunning to the cicada. They tell the tale of a coyote who wished to
learn the song of a cicada singing from the boughs of a pinyon pine. The
coyote had difficulty learning the song and twice forgot it on the way
home. When he returned for the second time, the cicada, distrustful of
the coyote, divested himself of his old skin, filled it with a quartz
pebble, sealed the vent with pitch, and flew to the safety of another tree.
The coyote became frustrated when this cicada image did not respond,
siezed it and splintered his teeth on the stone, driving some into his
jaw and leaving others protruding as tusks. His descendants inherited
these broken teeth. The cicada of the pinyon pine is either Tibicen
duryi or Okanagana ."agniF-ica. In the Orient cicada symbolism was de-
veloped to its greatest heights by the Taoists. They believe that the
soul of the departed escapes the body and is carried aloft by a cicada.
Early burials often included a cicada-shaped image, while in later
burials a cicada amulet was placed in the mouth of the corpse to facili-
tate the soul leaving the body.
As the centers of western civilization moved northward to areas
where cicadas are less prevalent, literary allusions to cicadas became
scarcer. Keats and Browning mentioned them in their poetry, but they had
lived in the southern Mediterranean regions of Europe. In the United
States nearly all cultural allusions to cicadas center around the genus
Magicicada, misnamed "seventeen-year locusts" because their periodical
abundance reminded early colonists of the biblical locusts.
The earliest scientific work on cicadas was that of Aristotle. He
described the morphology of the cicada mouth, their reproductive behavior,
and life history. Taxonomically, he made the distinction between the
large ones that sing and the smaller ones that do not sing. This latter
observation probably discriminated between our modern family Cicadidae
and other auchenorryncous Homoptera, although we now know many of these
produce sounds that are seldom audible to the human ear. Although
Aristotle was often quoted, cicada biology remained essentially in sus-
pended animation until Reamur described the sound producing organs and
made observations on the nests and eggs in the eighteenth century.
Linnaeus established the genus Cicada in 1758 in Systemma Naturae. In
the nineteenth century Fabre recorded the habits of some Mediterranean
Cicadidae. Since that time numerous other writers have described species
and various aspects of cicada morphology and biology. These are sum-
marized by Myers (1929). Distant, in a series of papers entitled
Rhynochotal Notes appearing in the Annals and Magazine of Natural History
(1904-1905),established our current system of cicada taxonomy.
The earliest attempt to treat the classification of North American
cicadas comprehensively was Uhler's Preliminary Survey of the Cicadaea
of the United States, Antilles, and Mexico, published in two parts in
Entomologica Americana in 1888. Van Duzee published a checklist of
genera and species occurring north of Mexico in 1916 and followed this
with a catalog of the same genera and species the subsequent year.
Davis treated the distribution of the genera and species in 1930.
Metcalf's (1963) world catalog is the most recent work to treat North
American genera comprehensively. Other papers dealing with several North
American genera are Lawson (1920), Beamer (1928), Beamer and Beamer
(1930), Froeschner (1952), Simons (1953, 1954), Moore (1966), and Alexan-
der, Pace, and Otte (1972). The writers primarily concerned with each
genus are reviewed under the individual genera in the following chapters.
William T. Davis made the most significant contributions to our
knowledge of western North American Cicadidae. For Davis the study of
cicadas was a second vocation. What began as a hobby during his years
as a successful industrialist became, after an early retirement, his
primary preoccupation until his death in 1944. Davis described many of
our genera and species. He was dependent for most of his specimens on
the collections of other people, but in his writings he included whatever
biological information was sent to him in letters accompanying the
specimens. Since Davis' death most work in North America has centered
around the periodical cicadas of the genus Magicicada.
Cicada eggs are laid in nests along the stems and branches of various
host plants. The nests may or may not be arranged in a definite order.
The female pierces the host with the ovipositor gradually driving into
the host tissue with an in-and-out sawing motion of the lateral valves.
Some species simply lay the eggs in the dry nest. Other species fill the
ends of the nest or the entire nest with a glue-like secretion, and
some, e.g., Tibicen lyricen, press the torn tissues of the plant back
into place before moving on to the next nest. Beamer (1928) recorded
from as few as three to as many as twenty eggs in the nests of Kansas
Cicadidae, while Myers (1929) cited estimates of egg production for a
single female that range from three hundred to seven hundred. All de-
scribed eggs resemble-each other closely in structure, varying only in
size. Some species overwinter as eggs. The eggs of other species hatch
the same summer they are deposited.
The hatching embryo ruptures the anterior end of the egg capsule
with its egg burster and makes its way to the nest opening, where it
sheds the postnatal membrane and falls to the ground. Upon reaching
the ground, the young nymph searches for a crevice in the earth to
enter. A quick retreat underground is especially important in dry
habitats as the newly hatched nymphs appear to be particularly vulnerable
Once in the ground, the nymph continues its downward movement until
it locates a suitable spot for a cell. The cell apparently is constructed
before food is located (Beamer, 1928). The cell protects the nymphal
cicada from both flooding and desiccation and probably provides some
protection from predators. The cell consists of walls of hard packed
earth. It is cylindrical and large enough to permit the nymph to do
a half somersault, his method of turning around. Once the cell is built,
the nymph looks for food. If food lies at the edge of the cell, he
begins to eat. If there is no food on the periphery of the cell, he
searches for food by laboriously moving his cell with him. Digging and
packing of the earth is done with special adaptations of the prothoracic
legs. Much of the above was verified by Beamer (1928) from observations
made on cicada nymphs in glass-sided cages.
Like most Homoptera, cicadas have five nymphal instars. The shortest
life cycle known is four years, a fact established by Beamer (1928) for
Cicadetta calliope. The longest cycle is the seventeen year cycle of
three species of Magicicada.
In the last month before emergence, the fifth nymphal instar extends
its cell to within an inch or two of the surface. J.E. Heath (1968)
showed that emergence from the ground of seventeen-year cicadas is
dependent on soil temperature. Temperature is probably the environmental
cue for emergence of other cicadas also. Upon emergence the nymph
climbs a nearby stem or trunk before the final molt to an adult.
The primary function of the adult stage is mating and renewal of the
life cycle. Adult cicadas live approximately six weeks to two months.
In some species, females appear to outlive the males by several days.
The two sexes are attracted to each other by a mating call or song
usually performed by the male. In Magicicada the song appears to have
a congregating function, bringing together both males and females rather
than uniting specific pairs. In Platypedia and Neoplatypedia both sexes
signal in a two-way communication. Several species produce three dif-
ferent sounds: the mating call, a sound used during courtship and mating
of specific pairs, and an alarm squawk emitted by males when they are
disturbed. Wing fluttering is also used as a signaling device between
individual cicadas. After mating, the female lays her eggs and the
cycle begins again.
In June, 1968, J.E. Heath, P.J. Wilkin, and I began a study of the
adaptations to climate of cicadas in Arizona. When we tried to identify
our experimental animals, we became frustrated by the lack of taxonomic
keys. No key to the genera of North American cicadas existed. Published
descriptions of genera were often vague and incomplete. There was
usually no way of comparing similar characters in one genus with another.
Many of the descriptions were of a comparative nature, but they required
identified specimens of the other genus or genera in hand in order to
make the comparisons. In some cases a lone female was unidentifiable
because only male diagnostic characters had been used.
We were faced with two alternatives. We could compile a list and
original descriptions of all species previously reported from the general
region and hope that what we had collected was not a range extension or a
new species, or we could travel to a large museum collection, and hope
that our species were available for comparative identifications. In
either case we would be working backward from species to genus or from
a larger group to a smaller group. Everyone wishing to identify indi-
vidual cicada specimens to genus could hardly follow these procedures.
Clearly a comprehensive taxonomic study of the genera of American cicadas
north of Mexico was needed. This paper presents such a study.
The aims of this dissertation are
1. To provide thorough, consistent descriptions of adults of the
genera of cicadas occurring in North America north of Mexico.
2. To provide keys and diagnostic characters for identification of
individual specimens to genus.
3. To indicate the general distribution of North American genera
north of Mexico.
4. To summarize the known biology of cicada genera found north of
5. To indicate evolutionary relationships of the North American
genera north of Mexico.
MATERIALS AND METHODS
The personal collection of cicadas used in this study dates from
1964. Most of the specimens were collected under the auspices of the
University of Illinois and arrangements have been made to deposit these
specimens in the Illinois Natural History Survey collection. Several
broods of Magicicada are included in the collection. Extensive col-
lections of western genera were made in Arizona in June and early July
of 1968, 1969, 1970, 1972, and 1973. Additional specimens were collected
in Colorado, Illinois, and Florida. Notes were made on location, time of
day, and habitat of each collection. Live weights and temperature data
were frequently recorded. Specimens were preserved in alcohol and pinned
and examined at a later date.
Several museum collections were examined in part or whole. These
included the Museum of Zoology, University of Michigan, Ann Arbor;
Arizona State University, Tempe; Illinois Natural History Survey, Urbana;
the Snow Collection, University of Kansas, Lawrence; the British Museum
(Natural History), London; and the Mus4um National d'Histoire Naturelle,
Paris. Numerous friends and colleagues contributed both specimens and
Measurements were made using the following tools; a Wild M5 stereo-
scopic dissecting microscope, calipers, and a plastic rule read easily
to the nearest half millimeter. Measurements were made from both
specimens and photographs. The photographic plates found in the col-
lected works of Davis (1921a, 1942) as well as my personal photographs
were utilized for proportional measurements. Descriptions were written
using the stereoscopic microscope and notes made while using the facili-
ties of the museums visited. Radiographs were made with a Faxitron 804
field emission x-ray unit and Polaroid positive-negative 55 film. Details
of exposure are given in the general morphology chapter. Additional
prints and enlargements were made by the photographic services of the
University of Florida.
Illustrations were drawn with a camera lucida. Original sketches
were compared with the descriptions and later reduced on a reducing
Xerox prior to the final inking. Three illustrations were made directly
from enlarged photographs. Color patterns are indicated, but the decision
was made to leave black insects uncolored so that morphological features
would not be obscured by the color. Several excellent drawings of male
external genitalic segments are found in the literature. Some of these
were redrawn from the originals after a comparison was made with specimens
Unless otherwise indicated in the text, the terminology used in
this paper conforms with that found in Torre-Bueno, A Glossary of
CLASSIFICATION: HIGHER TAXA
Cicadas are usually placed in the order Homoptera. This order is
comprised of hemipteroid insects with a sucking beak arising from the back
of the head. The proboscis contains four piercing stylets consisting of
two mandibles and two maxillae. In winged members, the forewings are
usually of a uniform texture throughout, either membranous or slightly
thickened. The hind wings are membranous. Winged forms hold the wings
tentlike over the body when at rest. Members of this order are plant
Cicadas belong to the suborder Auchenorrhyncha. Insects in this
suborder have three-segmented tarsi and short, bristle-like antennae. The
beak arises from the back of the head and not between the fore coxae as in
the Sternorrhyncha. Members of the Auchenorrhyncha are described as active
Cicadas are classified in the superfamily Cicadoidea. The designa-
tion Cicadoidea, however, means different things to different authors.
Those writers who divide cicadas into two or more families (cf. Kato,
Boulard, and Metcalf) define the Cicadoidea as those insects comprising
the former family Cicadidae. Authors who still maintain the single family
Cicadidae as representing all cicadas use the designation Cicadoidea to
separate the families Cicadellidae, Cercopidae, Membracidae, and Cicadidae
from the Fulgoroidea. This definition of the Cicadoidea is characterized
by antennae which arise on the front of the head between or anterior to
the eyes; middle coxae short and contiguous; and no "Y" vein in the anal
area of the wing (Borror and DeLong, 1971,and Borror and White, 1970).
Lamarck (1801) described the genus Cicada and named Cicada orni Linn.
as the type, thereby establishing the validity of the genus. Essig (1942)
gave the following history of the family: "Latrielle grouped these in-
sects as Cicadariae in 1802, and Leach erected the tribe Cicadides and
the family Cicadida in 1815. The later was changed to Cicadidae by
Samouelle in 1819 and has been generally accepted for this homogeneous
group of remarkable insects." Westwood (1840) and Metcalf (1963) attrib-
uted the name to Leach, but Leach's description of the tribe Cicadides
reads, "Antennae three-jointed. Ocelli two. Tarsi with three joints"
(Leach, 1832, First American edition, after Leach,1815). This description
specifically excludes the genus Cicada as established by Lamarck.
Lamarck's genus falls under Leach's tribe Tettigonides. Leach's family
Cicadida also excludes Lamarck's genus Cicada. Samouelle's (1819) work
follows the classification of Leach. According to Article l0e and
Article 63 of the "International Code of Zoological Nomenclature" (1961),
a family name must be based on a valid generic name. Latreille's (1802)
"Cicadariae" was roughly equivalent to Linnaeus' genus Cicada. It in-
cluded two subgroups, "Cicadae verae" and "Cicadellae." The genus Cicada
made up the "Cicadae verae." There were five genera in the "Cicadellae."
I believe this example is equivalent to the example given for Tipulidae
on page 11 of the 1961 International Code of Zoological Nomenclature and
the family Cicadidae, therefore, should be attributed to Latreille.
Westwood (1840) was the first author to limit the term "Cicadidae" to
this family of homopterous insects as we now recognize it. He equated
Cicadidae to "Cicadae maniferae" of Linnaeus, the "Chanteuses" of Latreille,
and the "Stridulantia" of Burmeister. Distant (1912), Myers (1929), and
Kato (1961) gave credit to Westwood for establishing the family. Metcalf
omitted Westwood from his "Bibliography of the Cicadoidea" (1962).
Westwood followed his designation by a detailed description of the family.
Other descriptions I consulted were Amyot and Serville (1843), Ashmead
(1888), Edwards (1896), Distant (1912), Lawson (1920), Metcalf (1951--
cohors Cicadida), Kato (1961--superfamily Cicadoidea), and Borrer and
The head is short and broad with an enlarged, cross-striated clypeus.
Antennae are short and setaceous, arising between the eyes and beneath
the supraantennal plates. There are three ocelli arranged triangularly
on the epicranium. The head terminates beneath in an elongated, three-
The thorax is large and broad. The pronotum is short except in
Tettigarcta where it covers most of the mesonotum. There are two diagonal
grooves on each side of the pronotum. The mesonotum is large and ter-
minates in a raised X or cruciform elevation ("in Tettigarcta forming a
scutellum," Kato, 1961). The metanotum is short and in some genera only
visible laterally. Ventrally the thoracic segments are subequal.
The abdomen consists of eleven segments. The first two are united
ventrally and the ninth and tenth segments form the external genitalia.
The eleventh segment bears the anal style. The third to seventh spiracles
are in the sternites and not the pleura.
The anterior femora are enlarged and spinose in all the genera of
North America north of Mexico. Tibiae are slender and subcylindrical.
There are no pulvilli or empodia between the tarsal claws. The meta-
thoracic mera are well developed and extend posteriorly. The legs are
not adapted for jumping, as in other auchenorrynchous Homoptera.
Wings may be hyaline, opaque, or both. Venation is distinct. An
ambient vein encloses the marginal cells. Forewings contain a basal cell.
A nodal line running from the nodus of the costal margin to the anal
margin may be indicated by auxiliary bumps and breaks in the venation.
Wings are coupled in flight by a recurved flange on the dorsal surface
of the costal border of the hind wing which hooks with a similar flange
on the ventral surface of the anal border of the forewing.
The ninth segment in both sexes is the pygofer, a more or less
cylindrical, strongly chitinized case surrounding the genitalia. The
aedeagus is further protected by a projection of the tenth segment known
as the uncus. The shape of the uncus has been used as the basis for
species determination in many species of North American Cicadidae. The
female external genitalia consist of three pairs of appendages making up
the ovipositor and its sheaths. The central pair is fused distally. The
apices of the middle pair are highly chitinized and possess cutting teeth
or edges. The outer pair forms the sheath.
Because methods of sound production among cicadas vary, the following
sound "producing" structures cannot be considered characteristic of the
entire family. Sound production has been so closely linked with cicadas,
however, that a description of the family would not be complete without
some mention of the most common structures associated with sound produc-
tion and reception. Sounds are most commonly produced by the contraction
and expansion of a tymbal organ located lateral to the first abdominal
segment of the males. The tymbals may be either partially or totally
covered from above by a forward projection of the second abdominal ter-
gite or by a posterior projection of the mesonotum (Boulard, 1976a).
Males with functional tymbal organs usually possess posterior metasternal
projections called opercula. The hearing or chordotonal organ is located
in the second abdominal segment. It may be indicated externally by a
lateral bulge on the sternite. Many cicadas have accessory stridulating
mechanisms and some produce sound by beating their wings together or
against the substrate on which they are sitting.
Current Taxonomic Status of the Family
At least three authors use the superfamilial name Cicadoidea to
encompass up to five families of cicadas. All of these divisions are
based on sound production mechanisms and/or the coverings associated
Kato (1961) lists four families. The two families Cicadidae and
Tettigadidae have tymbal organs. The Tettigadidae have an accessory
stridulating organ. The two families Platypediidae and Tettigarctidae
lack a functional tymbal organ. Platypediidae produce clicking sounds
with the forewings and have a chordotonal mirror. Tettigarctidae lack
the usual chordotonal mirror.
Metcalf (1951, 1963) lists two families of cicadas, Cicadidae and
Tibicinidae. Cicadidae possess an anterior projection of the second
abdominal tergite covering or partially covering the tymbal organ.
Tibicinidae lack this projection.
Boulard (1976a) divides cicadas into five families. The Tettigarc-
tidae are separated out by their very long pronotum covering most of the
mesonotum. Platypediidae lack a functional tymbal organ. A new family,
Plautillidae, is characterized by tymbal coverings emanating from the
metanotum and a reduction of the vannal region of the wings. Cicadidae
and Tibicinidac include the remaining genera, with Cicadidae possessing
an anterior projection of the second abdominal tergite in the males and
Tibicinidae lacking such a projection.
A definite difference exists in the philosophy and usefulness of the
family taxon between those who place cicadas in a single family, Cicadidae,
and the above three authors. If we accept Smith's definition of family
as reiterated by Torre-Bueno (1962), we can analyze the above divisions:
Family, a division or classification including a number
of genera agreeing in one or a set of characters and so
closely related that they are apparently descended from
Fossil evidence is sparse, but modern characteristics seem to in-
dicate that cicadas evolved from one stem. Almost all authors working
in cicada systematics eventually point out the basic similarity of all
cicadas and bemoan the lack of truly separable characters. Most mor-
phological characteristics of cicadas represent a continuum rather than
a "have" or "have not" status. Some authors have attempted to explain
this basic similarity on the basis of a homogeneous, long, subterranean
life shared by the nymphal forms. To me this explanation seems fallacious,
as edaphic characteristics of soil and differences in root characteristics
of host plants can surely be as varied as the external environment of the
images. The long life of the nymphs has resulted, however, in fewer
generations of cicadas responding to evolutionary pressures than those
insects with one or more generation per year. In North America north of
Mexico, the shortest confirmed life cycle is the four year cycle of
Cicadetta calliope (Beamer, 1928). The long life cycle could be a con-
tributing factor to the lack of diversity in cicadan morphology. It seems
evident, therefore, that the basic similarity of all cicadas verifies
their descent from a single phylogenetic stem.
The family Tettigarctidae used by both Kato and Boulard consists
of one genus. Members of this genus are generally considered the most
primitive of living cicadas. Although Tettigarcta apparently differs
considerably from other cicadas, separation of a single genus from its
nearest relatives obscures that relationship. A subfamilial ranking
seems more logical.
Boulard (1975a, 1976a), himself, recognized that the family Platy-
pediidae is not a natural grouping. The loss of the tymbal organ has
occurred at least three times, in North America, in Africa, and in China.
In both the African and North American genera, other characteristics
relate them more closely to other genera in their respective geographical
areas than to each other. Platypedia and Neoplatypedia have several
characteristics in common with the okanaganoid genera and would be better
placed as a subdivision of the same taxa in which these genera are placed.
The family as delineated by Kato and Boulard is characterized by a single
negative character. If one of the functions of systematics is to clarify
phylogenetic relationships rather than obscure them, this is surely not
a valid family.
In Kato's family, Tettigadidae, both males and females possess an
accessory stridulating surface on the mesonotum. A similar structure is
found in four genera of North American cicadas not generally considered
to be closely related to the Tettigadidae. In addition Boulard (1976a)
has described stridulating files on the posterior margin of the mesonotum
in the genus Moana. Because of the unavailability of specimens, I have
not had an opportunity to examine the stridulating mechanisms of the
family Tettigadidae as recognized by Kato. However, the mere presence
of accessory stridulating mechanisms now appears to be more widespread
among cicadas than previously thought. Therefore, Kato's family,Tetti-
gadidae,as originally circumscribed is no longer valid.
The three remaining families are all diagnosed by the presence or
absence and/or derivation of tymbal coverings. Only males have tymbal
organs, and hence, only males can be identified by these structures.
Even among males, only Boulard's Plautillidae appear definitely to form
a phylogenetic group. I think it is safe to assume that the tymbal organ
itself developed prior to any accessory protective devices. This struc-
ture functions to bring the two sexes into proximity for mating. Because
of its importance and relative fragility, the development of strengthening
or protective devices seems like the next evolutionary step. If the lack
of a projection of the second abdominal tergite indicates a primitive
state, then other characters must be drawn upon to show any special
phylogenetic relationship within the Tibicinidae. Myers (1929) was aware
of this when he stated, regarding Distant's subfamily Tibicininae from
which the family Tibicinidae is drawn, that the "Tibicininae . was
nothing but a dumping-ground for a miscellany of unrelated forms charac-
terized by a single negative quality." Some cicadas have increased the
number of ribs and the degree of rib chitinization within the tymbal
itself, and others have lost functional tymbals and replaced them with other
sound-producing mechanisms, perhaps as a response to tymbal fragility.
Even among the Cicadidae of Metcalf and Boulard, there is such a
variation in second abdominal tergal projections that it is not clear
whether this character developed once or many times. In our North
American cicadas it ranges from the mere upturned ribbon-like acrotergite
of Quesada gigas with its small lateral projections to the enormous
saddlebags of Diceroprocta arizona.
If we consider only males there is at least one other obvious
characteristic that may be equally important or more so from an evolu-
tionary standpoint than the presence of tymbal covers. That character
is the ability of some males to withdraw some genitalic structures as
contrasted with those that do not possess this ability. Thus, even con-
sidering only males, it is unclear whether the proposed groupings should
rank at the family level. Boulard's Plautillidae may represent a
phylogenetic grouping of subfamilial status.
In my opinion, the family taxon in insect systematics serves a
utilitarian function. It is the lowest level to which a nonexpert in a
group can easily identify a given specimen. While admittedly smaller
groupings may be easier to work with in some cases, I see no justifica-
tion for the splitting of phylogenetically valid families unless such
splitting further elucidates evolutionary and zoogeographic relationships
and increases the utilitarian function of the taxon. Any division that
obscures the identification of large numbers of a group at the familial
level fails to meet these criteria. The proposed family groupings de-
scribed by Kato, Metcalf, and Boulard are based primarily on male secondary
sexual characteristics. At least fifty per cent of the world's cicadas
are females. These proposals obscure the identification to family of
half of the possible specimens. I, therefore, propose we maintain the
family Cicadidae as a single unit until such time as more definitive and
useful criteria for familial designations are discovered.
Several attempts have been made to divide the Cicadidae into sub-
families. Amyot and Serville (1843) erected two subfamilies based on the
presence or absence of a reticulate condition of the veins of the fore-
wing. Ashmead (1888) added a third subfamily based on the opaqueness of
the wings. Wing characteristics have proven unsuitable at the subfamilial
level. The reticulate condition of the veins has apparently arisen at
least three times in the family, and several modern genera contain species
with both completely membranous, semiopaque, or partially opaque wings
(e.g., Okanagana). More recent attempts to divide the family(ies) into
subfamilies have been based on sound-producing mechanisms and their
attendant coverings. Distant (1889) proposed two subfamilies: Cicadinae
for those forms with the tymbals completely covered by a forward pro-
jection of the second abdominal tergite and Tibicininae for those forms
lacking a projection of the second abdominal segment. Distant's sub-
familial divisions proved inadequate. He found many intermediate forms
with the tymbals incompletely covered. For these he erected the sub-
family Gaeaninae in 1905. Jacobi (1907) erected the Tettigadinae.
Handlirsch (1925) attempted to erect a phylogenetic classification and
established the subfamily Platypleurinae for those forms with totally
covered tymbals. He united forms without tymbal covers or with the
tymbals partially covered under the subfamily Cicadinae. Myers (1929)
raised Tettigarcta to subfamilial level. Kato's family Platypediidae was
reduced to subfamilial level by Metcalf (1963). Metcalf (1963) used a
combination of tymbal cover characteristics and geographic distribution
to delineate subfamilies. His Tibiceninae includes Handlirsch's
Platypleurinae and geographically related forms with partially covered
tymbals with the exception of the genus Neocicada which he placed in the
Cicadinae. Boulard (1976a) added Moaninae for those Cicadidae with a
stridulatory surface on the posterior border of the mesonotum, Plautil-
linae to encompass his family Plautillidae, and Ydiellinae as a subfamily
of his Platypediidae with a stridulatory surface on the costal margin of
the hind wing.
Metcalf (1963) places seven of our North American genera in the
Tibiceninae (Cacama, Cornuptura, Diceroprocta, Tibicen, Becmeria,
Pacarina, and Quesada), one in the Cicadinae (Neocicada), six in the
Tibicininae (Okanagodes, Okanagana, Tibicinoides, Clidophleps, Magicicada,
and Cicadetta), and two in the Platypediinae (Platypedia and Neoplaty-
pedia). The last eight genera belong to the Tibicinidae of Metcalf,
while in Boulard's groupings, the last two are in the family Platypediidae.
Their subfamily designations, however, remain the same. The first eight
belong in the family Cicadidae under both arrangements; however, here
the resemblance ends. In Boulard's arrangement, by definition, Tibicen,
Diceroprocta, CornulpZra, and Caoama belong in the Platypleurinae because
the tymbals are completely covered, while Quesada, Pacarina, and Beameria
would be moved to the Cicadinae because the tymbal coverings are incom-
plete. The tymbal covers on Neocicada, on the other hand, are far more
extensive that they are on Cicada orni, the type for the genus Cicada from
which Neocicada was removed by Kato in 1932. Individuals following.
Boulard's key could conceivably place specimens of Neocicada heirogLyphica
in the Platypleurinae without representative samples of other Platypleur-
inae genera present for comparison. A comparison of the subfamilial and
familial arrangements of Metcalf and Boulard is given in Table 1.
Obviously subfamily designations are dependent on family designations.
The problem is that the same terms have now been used with so many varia-
tions in meaning that a complete overhauling on a world-wide basis seems
desirable. This is, of course, beyond the scope of this paper.
Distant was responsible for much of the tribal classification of
the Cicadidae in a series of papers in the "Annals and Magazine of Natural
History" from 1904 to 1905. These are summarized in his synoptic catalog
(Distant, 1906). His work was based on the vast collection of the British
Museum. Though his output was tremendous, his techniques have been
severely criticized. Myers (1929) sums up the problem thusly:
If any of the [referring to Distant] divisions or
tribes prove to be real morphological entities, it will
be largely a happy chance, since most of them appear to
be cabinet associations of superficially similar genera,
for the elucidation of which a study of his collection is
the first desideratum.
A comparison of Metcalf's and Boulard's family and subfamily groupings of
the genera of American cicadas north of Mexico.
Metcalf's "General Catalogue of the Homoptera, Fascicle VII,
Cicadoidea" (1963) gives the most recent listing of tribal affiliation.
He placed the North American genera in the following tribes:
There are no tribal designations for the two genera Playpedia and Neo-
piatypedia which comprise Metcalf's subfamily Platypediinae. Myers (1929)
felt that the tribe Tibicinnini represented a "miscellaneous collection
of all those forms which will not go into any of the other gymnotympanate
divisions." No improvement seems to have been made since that time.
Tribal classification has not been treated in recent papers on the
Cicadidae. Those characteristics which are applicable will be treated
in the discussion of individual genera in this paper. The tribal taxon
needs to be reworked on a cosmopolitan level.
Linnaeus (1758) separated the genus Cicada into five subdivisions.
The subdivision Manniferae became the "non saltantes" or true Cicadae.
Fabricius (1775) in Systema Entomologiae introduced the genus Tettigonia
for most of what we now consider cicadas, but the two names were frequently
synonymized. The genus Cicada remained the only generic designation of
the group for a long time. Latrielle founded the genus Tibicen in 1825.
The name was simply listed without description but with the naming of
C. plebeja Scop. as type. Some authors tried to use the name with various
interpretations, but most ignored it. Several species had been named as
types of the genus Cicada, among them the species Cicada tibicen. In
1914, Van Duzee firmly stated he felt Lamarck (1801) intended to name
Cicada orni as the type of the genus Cicada, thus freeing the name
"tibicen" to be used for another genus. He accepted plebeja as the type
of the genus Tibicen following the "International Code" and two years
later established many of our North American species as members of this
genus in his Check List of the Hemiptera of America, North of Mexico.
Among our American genera north of Mexico, the next genus to be
named was Cicadetta, Kolenati (1857). Diceroprocta appeared in a key
in Stal's Hemiptera insularum Philippinarum in 1870 but was not widely
used until Davis (1928) moved several North American species from the
genus Tibi-en to Diceroprocta. Uhler named Platypedia in 1888. Cacama
was named by Distant in 1904, and the same author named Pacarina, Quesada,
and Okanagana in 1905. Distant added Tibicinoides in 1914 and Van Duzee
named Clidophleps in 1915. All but one of the remaining genera were
named by William T. Davis. Those named by Davis include Okanagodes, 1919a;
Neoplatypedia, 1920; Magicicada, 1925; Beameria, 1934; and Cornuplura,
1944. In 1932, Kato transferred our remaining North American representa-
tives of the genus Cicada to their own genus, Neocicada.
The first attempt to list specifically the genera of America north
of Mexico was Van Duzee's checklist of 1916 followed the next year by his
catalogue of the same genera and species. Although Van Duzee listed six-
teen genera, the same number we have now, only eight of the names remain
the same. Metampsalta has been synonomized with Cicadetta. Our repre-
sentatives of Proarna have become Beameria; Tibicina have become Magicicada;
and Cicada have become Neocicada. Fidicina, Odopoea, Tettigades, and
Herrera were apparently erroneously attributed to this geographical re-
gion by early writers.
The only other paper to deal with the genera as a whole is Davis'
1930 distributional study. He also lists sixteen genera with only twelve
that correspond to our current list. Davis omits Fidicina, Odopoea, and
Herrara and adds Neoplatypedia, Diceroprocta, and Okanagodes. He replaces
Tibicina with Magicicada. Although Davis retains Tettigades as a United
States genus, based on Cochise County records, I have never seen specimens
of the genus collected north of Mexico. I have collected extensively in
Cochise County, Arizona, where Davis reports they occur. I have also
consulted with Dr. Thomas Moore of the University of Michigan, who has
collected in both the United States and Mexico, and he assures me that
the northern distributional limits of Tettigades lie far south of the
United States border (Moore, 1977, personal communication). Beameria,
Neocicada, and Cornuplura were not yet named when Davis wrote his 1930
paper, and the synonomy of Melampsalta with Cicadetta was not widely
The most recent listing of cicadas is the world wide catalogue of
Metcalf (1963). Metcalf occasionally indicates what he thinks is an
erroneous location, but he usually reiterates locations given by authors
without comment. His generic nomenclature is up to date. Early writers
were often obscure in listing their localities, thus one can extract
twenty-four possible genera for North American cicadas north of Mexico
from Metcalf's list. Errors in Metcalf's catalogue contribute additional
genera, for example, Neocicada chisos is still listed under the genus
The current list of American genera from the region north of Mexico
is as follows: 1. Beameria, 2. Cacama, 3. Cicadetta, 4. Clidophleps,
5. Conuplura, 6. Diceroprocta, 7. Magicicada, 8. Neocicada,
9. Neoplatypedia, 10. Okanagana, 11. Okanagodes, 12. Pacarina,
13. Platypedia, 14. Quesada, 15. Tibicen, 16. Tibicinoides.
The largest Homoptera are found in the family Cicadidae. These
insects are generally characterized by a rather short, stout body with
moderately long wings. The head is short, but wide, with prominent eyes
and three conspicuous ocelli. In the genera north of Mexico. the meso-
thorax is the largest thoracic segment. Males of most genera contain a
sound producing tymbal organ lateral to the first abdominal segment. A
chordotonal organ is indicated by a lateral bulge on the second abdominal
segment in both sexes.
Although several writers have written on specific aspects of both
the internal and external anatomy of cicadas, Myers' study (1928) remains
the most comprehensive treatment of cicada morphology. A number of
specialized terms have been used in the past to refer to features of
cicada anatomy. This paper incorporates several of these terms into the
descriptions and keys in order to provide continuity with past work. The
following discussion is presented to clarify terms used in the keys and
generic descriptions. General characteristics of cicada morphology are
illustrated by diagrammatic views of the dorsal and ventral surfaces
A, dorsal view of a cicada. The following structures are
labeled: clypeus (clp), epicranium (ep), pronotum (pr),
pronotal collar (pc), prescutum (psc), scutum (sc), scutellum
or "cruciform elevation" (scl), metathorax (mt), tymbal cover
(tc), pygofer (p). The left tymbal cover is cut away to
reveal the tymbal organ (tym). In this example the mesothorax
covers most of the metathorax. The arrow points to the loca-
tion of the lateral carina when viewed laterally. B, ventral
view. Legs have been cut away below the coxae (cx). The
following structures are labeled: clypeus, operculum (op),
propleuron (prp), meron (me), hypandrium (h), epimeron of
the metathorax (mep). The right operculum is cut away to
reveal the folded membrane (fm) and the mirror or tympanum
(m). Roman numerals in both figures indicate abdominal
General Body Proportions
Two measurements are given for each genus to provide an indication
of general range of size of members of the genus. Length of body is
measured from the apex of the clypeus to the tip of the abdomen. Expanse
of wings is the greatest expanse of the forewings. This measurement is
the only wing measurement consistently given in species descriptions.
The relationship of abdomen length to overall length of the body is also
indicated. This relationship usually is characteristic of a genus. The
variability in size range is greater, of course, in specimens of the
larger genera. While size ranges overlap considerably, from a practical
standpoint, size can often narrow the number of genera under considera-
tion when identifying an individual specimen. Thus, a very large cicada
would not fall in the range of Tibicinoides, Beameria, or Pacarina, and
a very small cicada is unlikely to be a Quesada, Tibicen, or Cornuptura.
The cicada head contains several characteristic features. Tradi-
tional measurements and nomenclature of the head are in some cases
obscure and in others a point of controversy in the literature. A
characteristic measurement used by Distant and others is a comparison
of the width of the head, including the eyes, versus the width of the
"base" of the mesonotum. Torre-Bueno (1962) defines the base of a
thoracic structure as "that part nearest the abdomen"; however, this is
a meaningless measurement in cicadas. Previous writers obviously meant
the visible anterior edge of the mesonotum. Since the real anterior
edge of the mesonotum is overlapped by the collar of the pronotum, a
sometimes curved structure, this measurement is also awkward. Therefore,
the width of the head, including the eyes, in the following descriptions
is compared with the width of the mesonotum measured at its broadest
point between the forewings, excluding the axillary sclerites.
The eyes in cicadas are prominent and may be narrower, in a line
with, or extend laterally beyond the anterior angles of the pronotum.
This relationship is usually obvious. However, a straight edge may be
lined up with the lateral edge of the eye to determine its relative posi-
tion with the anterior lateral angles of the pronotum.
The definitions of the head regions are controversial, also.
Snodgrass (1935) defined the clypeus as the "region of the cranial wall
on which the dorsal dilator muscles of the . cibarium take their
origin." The frons was interpreted as a triangular region which "bears
the median ocellus on its upper part." When a frontoclypeal or epistomal
suture is present, it separates these two regions. The anterior tentorial
pits are normally associated with the lateral ends of this sulcus and
mark the point of articulation with the anterior arms of the tentorium.
In some insects the pits have migrated medially on the sulcus. When a
frontoclypeal suture is lacking, a line drawn between the anterior
tentorial pits is used to delineate the frontal and clypeal regions of
In the cicada the median facial plate is bordered both dorsally and
laterally by a continuous suture. The anterior tentorial pits are
located in the upper half of the lateral portion of this sulcus. DuPorte
(1962) interpreted the transverse portion of the sulcus as a "transfrontal
ridge such as found in mantids." The lateral regions between the trans-
verse portion and the anterior tentorial pits, he called "frontogenal
ridges," and those distal to the pits, he interpreted as "clypeogenal
ridges." He supported this view by stating that the anterior mandibular
articulation would then correspond to that typically found in Hymenoptera.
He further stated that the entire head evolved from a hypothetical
psocid-like ancestor. In his interpretation, the frons would consist of
the region anterior and dorsal to a hypothetical line drawn between the
tentorial pits, and the post clypeus would be the region posterior to this
line. In reality this interpretation is a return in part to the de-
scriptions of earlier morphologists and taxonomists who called the entire
median facial plate the "front" or frons and the remainder of the epi-
cranium the "disk of the vertex." This terminology was used prior to
Snodgrass' detailed morphological studies.
DuPorte's interpretation is interesting but requires several assump-
tions: 1. The frontoclypeal sulcus has disappeared or never existed;
2. the muscles of the cibarium have invaded the frons; 3. The modern
cicada head has developed or evolved from both specialized orthopteran
and hymenopteran characteristics; 4. The entire head, and by extension,
the rest of the organism developed from a psocid-like ancestor. Both
Snodgrass and Kramer (1950) cited evidence of a "Y" shaped frontal suture
encompassing the median ocellus. They interpreted the arms of this suture
as the demarcation of the sides of the modern frons. Kramer further
used these ridges as part of his phylogentic evidence that ancestral
cicadas branched from a common ancestoral stem with cercopids, cicadellids,
and membracids much earlier than the separation of the three from each
In an attempt to verify DuPorte, I felt that if even part of the
lateral edges of the median facial sclerite split during ecdysis, it would
be evidence in favor of his theory. A careful examination of exuviae in
my collection indicated that the ecdysial split terminates at the trans-
verse sulcus. This is not, however, inconsistent with other writings of
The dorsum of the cicada head contains three distinct regions, a
horizontal vertex, a sloping, sometimes vertical, middle region anterior
to the vertex containing the median ocellus, and an anterior horizontal
region containing the dorsum of the median facial plate flanked by the
supraantennal plates. If we accept DuPorte's ideas, what should we call
the central section? If we accept Snodgrass' interpretation, the length
of this segment is equivalent to the length of the frons.
There is a much simpler explanation for the dorsal structures of the
cicada head than DuPorte's theory. The clypeus has obviously expanded to
accommodate the dilator muscles of the cibarium. If it expanded upward as
well as outward, the frontoclypeal sulcus would lengthen around the
sclerite. Without moving the tentorial pits, the sulcus would attain the
current broad "U" shape. The frons would be subsequently shortened as
the clypeus expanded. This interpretation does not require the loss of
one sulcus and the acquisition of another, the invasion of the frons by
the muscles of the cibarium, or other specialized characteristics except
those associated with the more efficient acquisition of food.
I have no evidence either in support of or against DuPorte's theory.
DuPorte worked with an unidentified Tibicen. Until I see additional
studies of closely related Homoptera, including Tettigarcta, which
apparently retains a vestigial subgenal suture (Kramer, 1950), I prefer
the simpler explanation. Therefore, in the following descriptions, the
length of the frons will refer to the more or less vertical portion of
the head, as distinct from the vertex, and will be compared with the
dorsum of the clypeus. The epicranium will be assumed to end with the
epistomal or frontoclypeal suture (see Figure 2).
Cicadas have three ocelli. The two lateral ocelli lie on the ver-
tex of the head, the middle ocellus is located on the frons. The rela-
tionship of the distance between the two lateral ocelli versus the
distance between a lateral ocellus and the corresponding eye usually is
characteristic of a genus. Measurements were made from the interior
margin of the eye to the exterior edge of the ocellus and between the
medial edges of the two lateral ocelli.
The supraantennal plates lie above the insertion of the antennae.
The anterior edge of these plates may form a more or less continuous line
with the apex of the clypeus or the clypeus may protrude substantially
beyond the margins of the supraantennal plates. The dorsal shape of the
clypeus and the extent of its protrusion are generically characteristic.
The clypeus is flanked laterally on the "face" by the lora (singular,
lorum) sometimes referred to as the "cheeks."
The rostrum or proboscis is an extended mouth piece representing
the sucking beak in cicadas. The rostrum is comprised of an elongated
labium forming 'the outer sheath and containing the mandibular and max-
illary bristles. The maxillary bristles interlock centrally forming two
primary ducts, one for the egestion of salivary fluid and one for the
ingestion of food. The length of che rostrum may be species specific
within a genus; however, a prescribed range of lengths is usually
characteristic of a genus. Thus, Okanagana have relatively short rostra,
extending within the range of the middle coxae; whereas, Cacama have long
rostra extending to or beyond the hind coxae.
The cicada head. A, face or ventral side showing position
of eyes (e), ocelli (oc), frons (fr), supraantennal plates
(sap), lora (1), postclypeus (pclp), anteclypeus (aclp),
labrum (1m), rostrum (ros). B, dorsum showing the eyes,
ocelli, frons, supraantennal plates, the dorsum of the
clypeus (clp), and the frontoclypeal or epistomal suture
(fcs). The lateral circles just media to the eyes mark the
location of an unidentified sensory organ. C, cross section
of the rostrum showing the relative positions of the man-
dibular (md) and maxillary (mx) setae. The large central
lumen is the food channel. The smaller lumen to the right
is the salivary channel. Four smaller luma, one in each of
the setae form passageways for nerves and tracheae. The
labium (lab) forms the outer sheath.
The pronotum is characterized by two pairs of diagonal grooves. The
posterior margin is broad and ends with the dilated lateral posterior
angles. The posterior margin is frequently referred to as the pronotal
collar. The lateral margins may be ampliated or dilated. Laterally the
pronotum joins the propleuron. If the pronotum is ampliated, when viewed
laterally, this juncture forms a continuous keel or carina extending the
length of the pronotum. The extent of the lateral amplification deter-
mines the extent of the carina. The lateral edges of the pronotum are
usually smooth in the genera found north of Mexico. They are irregularly
toothed or jagged in Quesada and in some species of Okanagana. Occasion-
ally the lateral edges will bear one or two spines.
The mesothorax is the largest thoracic segment in the North American
genera of cicadas. The anterior part of the notum projects beneath the
pronotum forming an anterior phragma. A "U" shaped prescutum is located
in the center of the anterior portion of the notum. The scutum occupies
the greater part of the notum. The scutellum forms a raised "X" which
has been termed the "cruciform elevation" by taxonomists. In some genera
the mesonotum is ridged where the forewing sits at rest. A portion of
this ridge may project medially on the anterior part of the notum near
what is sometimes called the "shoulder." TWhen this occurs, the projec-
tion is finely striated and used as an accessory sound producing mechanism
with the wing acting as the plectrum. Sometimes the ridged portion
bears fine, even striations (see Figure 3).
The metathorax is very short dorsally. In half of the genera found
north of Mexico, the scutellum of the mesothorax overlies and conceals
A portion of the thorax of an okanaganoid cicada showing
striations on the mesonotum. Large arrow points to the
stridulitrum. Small arrow points to the presumed plectrum
on the forewing. Terminology follows Ashlock and Lattin
(1963). The stridulitrum is frequently located in a pale
area on a dark background. This area is indicated on the
figure. Drawn from Clidophleps blaisdelli.
the central portion of the metathoracic tergum. Ventrally, the epimera
in males are usually produced caudally forming the opercula, or lower
covers of the tymbal organs.
The legs of cicadas are normally segmented. They are not modified
for jumping as in some of their closest relatives. There are three
tarsal segments, but no pulvilli between the tarsal claws. The foreleg
of the nymph is fossorial. In the adult the fore femora retain a
lessened fossorial expansion and usually bear spines on the ventral
margin (Figure 4). The tibia are subcylindrical. The metathoracic mera
are elongated and form projecting triangular structures. These projec-
tions are the meracanthi of some older works, but more recent publications
use mera (singular, meron), e.g., Kramer (1950), Bullough (1950).
Wing notation is by no means fixed for the family. The most common
deviation is the proximal portion of the radius. Some authors combine
this portion of the radius with the base of the media where it emerges
from the basal cell or the arculus. This of course modifies the notation
of the veins at the apex of the wing and also the notation of the inter-
vening cells. The following descriptions follow the modified Comstock-
Needham interpretation of venation accepted by Myers (1928, 1929),
Cooper (1941), Kramer (1950), and others. This is the interpretation
most commonly given in modern general textbooks (e.g., Borrer and DeLong,
1971). In this interpretation the radius is united with the subcosta
Femur of a cicada foreleg showing expanded configuration
and spines on the ventral surface. Drawn from Okanaganc
proximal to the body. This interpretation is used for most original
descriptions of North American genera and species. The following remarks,
therefore, are consistent with prior descriptions. The modified
Comstock-Needham systemwhich I am following, is shown in Figure 5.
Cell notation in the Comstock-Needham system is complex. The cells
or areoles in cicada wings are often referred to by groups, for example,
the eight apical areas or marginal cells or the ulnar areas. This is the
system of cell notation used and diagrammed by Simons (1954). Simons'
notation is simpler and lends itself to use in keys and the more
generalized descriptions at a generic level. Simons' system of cell
notation is shown in Figure 5 and is utilized in the following text.
A special feature of the forewings is the presence of the nodal line.
This line runs from the nodus or node of the costal margin across the
width of the wing. Its presence is marked by minute breaks in the
sclerotization of some of the principal vein branches. The function of
this line became apparent to me during a sequence of a movie showing
Magicicada septendecim in the stages of the final molt to an imago (Moore
and Kausch, 1975). The wings are apparently folded along this line in
the last instar nymph prior to emergence as an adult. The line is well
marked in some members of the genus Tibicinoides by a rudimentary vein.
The line apparently is also present in the Cicadelloid family Hylicidae
The hindwings of some genera are susceptible to individual variation.
Therefore, hindwing characteristics have not been utilized in this
Cicada forewing showing two systems of notation in current
usage. A, cell notation used by Simons (1954). Abbrevia-
tions are as follows: A, anal cell; arc, arculus; Ba, basal
cell; C, costa; Cu, cubital cell; Ml, M2, etc., marginal
cells; Me, medial cell; N, nodus; UI, U2, etc., ulnar cells;
1st and 2nd equal the first and second crossveins. B,
Comstock-Needham system used in current textbooks of entomology.
Abbreviations used in the notation are longitudinal veins--C,
costa; Sc, subcosta; R, radius; M, media; Cu, cubitus; A,
anal. Crossveins--m, medial; m-cu, medio-cubital; r-m, radio-
medial. Cells are named according to the longitudinal vein on
the anterior side of the cell. The general position of the
nodal line is indicated by dots in both A and B.
The first two abdominal segments are modified for the production and
reception of sound. Pringle (1954, 1957) gives detailed accounts of the
morphology of this region. The exterior cuticle is depressed and
thickened forming a wide vertical groove. This groove "is divided into
dorso-lateral and ventro-lateral portions by a horizontal ridge projecting
forwards from the lateral angle of the abdomen" (Pringle, 1957). Ven-
trally the groove is bounded by two membranes, an anterior "folded
membrane" that permits the abdomen to move relative to the thorax and a
posterior chordotonal "tympanum" sometimes termed the "mirror." At the
base of the groove is a heavily sclerotized skeletal structure that forms
the basal attachment of the tymbal muscles and projects dorso-laterally
to form the chitinouss V." Dorsally the groove is bounded posteriorly
by a strongly sclerotized surface, while the anterior surface is the
sound producing "tymbal organ" or simply "tymbal" (Figure 6). The tymbal
organ itself is a ribbed membrane whose buckling produces the sound
vibrations associated with cicadas. An "auditory capsule" is indicated
exteriorly by a lateral bulge on the second segment. Interiorly this
capsule contains a number of chordotonal sensilla connected to the
thoracic ganglion by an auditory nerve running forward through the base
of the chitinous "V" to the thoracic ganglion. In Neocicada and Cicada
this capsule has migrated ventrally. Pringle (1954, 1957) and Vasvary
(1966) detail the musculature and nervous connections of this area in
Platypleura capitata and Tibicen chioromera, respectively.
Abdominal segments III through VI are not specialized. The terga
form lateral angles and project across the lateral areas of the ventral
Structures associated with the first and second abdominal
segments in male cicadas. A, vertical groove, viewed
laterally with left opercula and tymbal cover removed.
Structures shown are tymbal organ (tym), auditory capsule (a),
horizontal ridge (hr), folded membrane (fm), chitinous "V"
(V), tympanum or mirror (m), opercula (op), posterior coxae
(cx), meron (me). The direction of the thorax (tho) and
abdomen (abd) are indicated. B, posterior view of the same
area. The tymbal cover (tc) and tymbal organ (tym) are re-
moved on the right to reveal the sclerotized posterior
surface. Tymbal muscles (t mus) are shown. Drawn from
side of the abdomen. The terminal segments of the abdomen are discussed
under the heading "sexual characteristics."
One of the primary functions of this study is to enable the reader to
identify easily specimens "in hand" to genus. For that reason only ex-
ternal portions of the segments bearing genitalia have been described.
These are the parts that are readily seen or easily extracted without
complicated dissection. Numerous other genitalic structures are available
for use at the specific level. The terminal segments of the abdomen are
modified by the genitalic appendages. The ninth and tenth segments form
the external genitalia. The eleventh segment is small and bears the anal
style. The ninth tergite in both sexes forms the pygofer. The pygofer in
cicadas is a simplified structure forming nearly a complete sclerotized
ring. Ventrally the ends of the tergite may meet, but they remain un-
connected in females to allow for the extrusion of the ovipositor. The
posterior portion of the ventral region in males is invaginated into the
segment to form the genital chamber that holds the phallus. "Genital
plates and genital valve as well as styles and connective are absent"
(Tuxen, 1970). The tenth segment in males bears an interesting structure
apparently functioning to protect, and in some cases, hold the phallus.
This structure may be united or simple as in Tibicen and the okanaganoid
genera, or it may be bifurcated, consisting of a pair of structures as
in Cicadetta, Magicicada, and Diceroprocta. It has been called by a
variety of names, e.g., "copulatory claspers" (Myers, 1928, 1929),
"lateral lobes" (Snodgrass, 1935), "anal hooks" (Tuxen, 1970), "ventral
processes" (DuPorte, 1977). Taxonomists call it the "uncus," and it
will so be named in this study. The uncus has formed the basis for both
generic and specific separation especially in the cicadas found in North
America north of Mexico. Six western genera are characterized by an
elongated uncus lying dorsally over the phallus. The hypandrium or
eighth sternite, a "boat-shaped" sclerite, is correspondingly elongated
and protects the phallus from below. The uncus in these genera does not
retract into the abdomen. In the remaining genera the pygofer may be
substantially withdrawn into the eighth segment. The uncus maintains a
ventral position in the noncopulating male. It is hidden ventrally by
the correspondingly shorter hypandrium (see Figure 7).
In females the seventh sternite forms the subgenital plate. Its
basic shape is frequently characteristic of a genus. The eighth sternite
is not visible. It is greatly reduced in size and consists of two lateral
sclerites lying under the seventh sternite. In some Tibioen and
Conuplura the eighth sternite may project into hooks medially. The
female possesses three pairs of appendages, or "valves" that make up
the ovipositor and its sheaths. The central pair is fused distally. It
is united with the middle pair by a series of interlocking grooves that
prevent lateral movement but allow the lateral pieces to slide longi-
tudinally relative to the central piece. The middle pair of valves are
highly chitinized and possess cutting teeth or edges. The outer pair of
valves form the sheath of the ovipositor (Figure 7).
Internal Structure Revealed by Radiographs
Systematists have recognized for a long time that characteristics
other than external anatomical features could be useful in identification
and in correlating relationships of various taxcnomic units. Numerous
External genital bearing segments of cicadas. A through E
are males. A, lateral view of retractable form with the
pygofer (p) and uncus (u) pulled out; B, same as above in a
normal resting position; C, caudal view. D, lateral view of
nonretractable form with the uncus raised; E, same in a normal
resting position with the uncus lying in the trough of the
hypandrium (h). F through H show female characters. F,
caudal view showing hooks of sternite VIII lying under the
subgenital plate (sp). This character occurs in Cornuplura
and some Tibicen. G, ventral view showing the pygofer (p)
surrounding the ovipositor. H, female ovipositor showing
saw valves (vl), middle piece (v2), sheaths (v3). Terminology
after Myers (1929).
approaches have been taken ranging from simple internal dissections to
complicated chemical extractions. While most of these techniques yield
valuable information, they may be impractical for the average entomologist
to use. They frequently require fresh specimens or at least freshly
preserved specimens for best results. Many techniques are extremely ex-
pensive. They all require destruction of part or all of the specimen,
and,therefore, cannot be used on museum types. Professor H.L. Cromroy of
the University of Florida suggested I explore the use of low energy
x-rays (soft x-rays) to elucidate internal characteristics. The results
are only preliminary, but a few remarks can be made regarding the technique.
The model [Faxitron 804] used is small and fits on a table-top. It is a
relatively inexpensive piece of scientific equipment and is as easy to
operate as a kitchen oven. It in no way alters or damages the appearance
of the specimen and, therefore, should be safe to use even on type
specimens. Two series of radiographs were taken. Dorsal views were
made at 50 KVP (kilovolt potential) with exposures ranging from 45 to 60
seconds, and ventral views at 25 KVP with an exposure time of 60 seconds.
Polaroid P/N 55 film was used, and the resulting radiographs were en-
larged for analysis. At these exposures and times, elements of dried
internal musculature are the most obvious features revealed. The median
dorsal indirect flight muscles show clearly in both sexes. The other
thoracic musculature is less distinct. Elements of the reproductive
musculature are also seen in some of the pictures. The most significant
muscular feature of taxonomic value revealed is the generalized shape of
the male tymbal muscles.
Pringle (1954) defines the tymbal muscles as, "The main muscles of
the sound-producing apparatus, attached basally to the base of the
chitinous V and distally to a thin approximately circular disk from the
centre of which a flat apodeme runs to the tymbal." While the muscle runs
dorso-ventrally through the insect, a characteristic pattern is shown by
x-rays made of a dorsal or ventral view. Thus the four North American
genera having complete tymbal covers and belonging to the tribe Tibicenini
have a characteristic deep V shape to the x-rayed tymbal muscle. The V
is united posteriorly forming a Y in both Tibicen and Cornuplura. This
pattern holds for both the parallel sided and gently tapering forms of
Tibicen discussed later in the text. Kato (1961) has also taken
radiographs of cicadas. He pictures two genera of Tibicenini found in
Japan (Kato, 1961, Plate XIX). The x-rayed tymbal muscle in Tibicen
bihamatus (Motschulsky) illustrated by Kato agrees perfectly with the
pattern seen in North American Tibicen, while Cryptotympana japonensis
Kato lacks the united posterior extension of the V. Cacama also lacks
the posterior extension, its V shape being broader and shallower than
Tibicen. The pattern in Diceroprocta shows the greatest deviation from
the other Tibicenini. Although the central posterior V is present, the
lateral connections are broader. Additional partial anterior muscles
extend upward, almost forming a hexagonal pattern. The pattern in
Diceroprocta is very similar to that found in Magicicada which lacks
tymbal covers completely. Pacarina has a similar pattern, but the arms
do not meet posteriorly. In Neocicada the distal connections are narrow.
The four okanagancid genera, Okanagana, Tibicinoides, Okanagodes, and
Clidophleps exhibit a rather flat latitudinal pattern indicating a more
vertical structure within the insects. This pattern appears broadest in
Okanagana. Quesada shows a broad latitudinal pattern and Beameria falls
between Okanagana and Quesada. Cicadetta has a narrow pattern projecting
slightly anteriorly. X-rays of Platypedia and Neoplatypedia reveal no
sign of tymbal muscles. While all the patterns of tymbal muscles
generated by North American genera occurring north of Mexico appear to
be deviations of two basic patterns, they do not group the genera com-
pletely around traditional lines. The most notable exception is the
pattern exhibited by Magicicada, resembling more closely the muscular
pattern in the group containing tymbal covers rather than the group
lacking tymbal covers. The tymbal covers in Beameria and Quesada are
minimal. Handlirsh (1925) felt that cicadas with partial tymbal covers
should be considered more closely related evolutionarily to those lacking
covers than to those with complete covers. The two genera with extensive,
but not complete covers, Pacarina and Neocicada, are more similar to
those genera with tymbal covers than they are to those without. Although
variation in the tymbal musculature patterns found in the North American
genera are not as striking as one might hope for, quite different pat-
terns are illustrated by Kato for the genera Tanna and Graptopsaltria
from Japan, indicating that both this character and the technique that
reveals it are worth pursuing further. Figure 8 shows examples of the
Other morphological features exhibited by all or several of the
x-ray radiographs are the shape and position of the frontoclypeal suture,
the insertion of the forelegs, and the extent of ovarian development.
All the specimens x-rayed were dry, pinned specimens. Therefore,
care must be used in interpretation of the radiographs. Comparative
studies need to be made between fresh specimens and dry ones to see the
extent and variation of change with time. Various exposures and times
may reveal different structures. For these reasons, the above analysis
Radiographs of cicadas. A though H are males. Arrows
point to the location of the tymbal muscles. A, Tibicen,
and B, Cornuplura, show a Y configuration. C, Cacama, shows
a broad V. D, Diceroprocta, and E, Magicicada, are almost
hexagonal. F, Quesada, and G, Tibicinoides, are transverse.
H, Platypedia, lacks a tymbal muscle. I through L are
females. Large arrows point to the ovaries. Small arrows
on I, Quesada, point to the frontoclypeal sulcus and the
insertion of the foreleg. The small arrow on J, Tibicen,
points to the muscles surrounding the ovipositor. K and L
are Magicicada and Cacama, respectively. E, G, and H are
enlarged. The remainder approximate the actual size.
is regarded as both speculative and preliminary. The technique does show
promise, however, not only in taxonomy but in ecology as well. Struc-
ture and utilization of nests, nymphal development, analysis of parasitism,
and infestations within plants are but a few of the possibilities. The
greatest advantage of low energy x-ray radiography in taxonomy is that
it reveals additional information without damaging the source.
KEYS TO THE GENERA
1 Posterior margin of metanotum partially covered by mesonotum; lateral
anterior projections of 2nd abdominal tergite covering or partially
covering the tymbal organ (Fig. 1). . . . . . . . . 2
1' Posterior margin of metanotum completely visible behind mesonotum;
lateral anterior projections of 2nd abdominal tergite absent
(Fig. 12). . . . . . . . . . . . . . 10
2 (1) Tymbal covers complete dorsally; opercula meeting or nearly
meeting (Fig. 2). . . . . . . . . . . . 6
2' Tymbal covers incomplete dorsally; opercula may meet (Fig. 20) . 3
3 (2') Tymbal covers triangular and concave medially; opercula meeting
or nearly meeting (Fig. 20). . . . . . . .Pacarina
3' Tymbal covers not triangular or concave medially; opercula separated
(Fig. 9) . . . . . . . . . . . . . . 4
4 (3') 1st and 2nd crossveins of forewings parallel (Fig. 5). . . 5
4' 1st and 2nd crossveins of forewings not parallel (Fig. 9). .Beameria
5 (4) Lateral edges of pronotum smooth (Fig. 16) ....... eocicada
5' Lateral edges of pronotum jagged or irregularly toothed
(Fig. 22). . . . . . . . . . . . . Quesada
6 (2) Ninth abdominal tergite produced posteriorly to a central spine
or point (Fig. 6) . . . . . . . . . . . 7
6' Ninth abdominal tergite without a central spine (Fig. 14). . . 9
Diceroprocta and Tibicen in part
7 (6) Width of head including eyes much narrower than the width of the
mesonotum; length of body less than 2 1/2 times the width of the
mesonotum; abdomen broadly rounded caudally (Fig. 10) . .Cacama
7' Width of head including eyes about as wide or wider than mesonotum;
length of body at least 2 1/2 times the width of the mesonotum;
abdomen tapering caudally (Fig. 1) . . . . . . . . 8
8 (7') Ninth abdominal tergite with a pair of spines protruding backward
from the ventral edge (Fig. 13). . . . . . .Cornuplura
8' Ninth abdominal tergite lacking ventral spines (Fig. 23 and
Fig. 7). . . . . . . . . . . . ... .Tibicen
9 (6') Length of body greater than 35 mm; uncus simple
(Fig. 23). . . . . . . . . . .. ... Tibicen
9' Length of body less than 35 mm; uncus bifurcated (Fig. 14) Diceroprocta
10 (1') Uncus retractable (Fig. 7). . . . . . . . .. 11
10' Uncus exposed; hypandrium elongated (Fig. 7). . . . . . .12
11 (10') Medial and cubital veins joined when emerging from basal cell
of forewing (Fig. 11) . . . . . . .... Cicadetta
11' Medial and cubital veins emerging separately from basal cell of
forewing (Fig. 15). . . . . . . . ... .Magicicada
12 (10') Tymbal organs present (Fig. 1) . . . . . .... .14
12' Tymbal organs absent (Fig. 21). . . . . . . . . .. 13
13 (12') Eight marginal cells in forewing (Fig. 21) .... .Patypedia
13' Seven marginal cells in forewing (Fig. 17). . . ... Neoplatypedia
14 (12) Pronotum flared laterally forming longitudinal ridge or carina
when viewed from the side; ocelli variously colored ... .15
14' Pronotum constricted, rounding laterally; ocelli ruby red
(Fig. 19) . . . . . . . . .. . Okanagodes
15 (14) Marginal cells of forewing about 1/2 the length of ulnar cells
Ul and U2 (Fig. 24) . . . . . .... .Tibicenoides
15' Marginal cells of forewing at least 2/3 the length of ulnar cells
Ul and U2 (Fig. 5) . . . . . . . . . . . 16
16 (15') Costal margin of radial cell at least 2 1/2 times longer than
the medial vein anterior to the first branch
(Fig. 18). . . . . . . . . . ... .Okanagana
16' Costal margin of radial cell less than 2 1/2 times longer than the
unbranched segment of the medial vein (Fig. 12) . .. Clidophleps
1 Posterior margin of metanotum partially covered by mesonotum
(Fig. 1) . . . . . . . . . . . . . . 2
1' Posterior margin of metanotum completely visible behind mesonotum
(Fig. 12) . . . . . . . . . . . . . .11
2 (1) Width of head including eyes about as wide or wider than width
of mesonotum (Fig. 1) . . . . . . . . . . 3
2' Width of head including eyes much narrower than the width of the
mesonotum (Fig. 10) . . . . . . . . . .. Ca ama
3 (2') First and second crossveins of forewing parallel or nearly
parallel (Fig. 5). . . . . . . . . . . . 4
3' First and second crossveins of forewing not parallel (Fig. 9). . 9
4 (3) Longitudinal veins with infuscated dots near the ambient vein
(Fig. 16). . . . . . . . . . . Neocicada
4' Longitudinal veins without infuscated dots near the ambient vein
(Fig. 5) . . . . . . . . . . . . . . 5
5 (4') Lateral edges of pronotum jagged or irregularly toothed
(Fig. 22) . . . . . . . . . . .. . Quesada
5' Lateral edges of pronotum smooth (Fig. 1). . . . . . . 6
6 (5') Subgenital plate sinuous and/or length of R4+5 between the first
and 2nd crossveins equal to or greater than length of R4+5
from first crossvein to the ambient vein in the forewing
(Fig. 14) . . . . . . . . ... . .Diceroprocta
6' Subgenital plate produced into two symmetrical, smooth lobes; length
of R4+5 between the first and second crossveins less than the
length of R4+5 from the first crossvein to the ambient vein in
the forewing (Fig. 23) . . . . . . . . ... . . 7
7 (6') Eighth abdominal sternite visible as 2 hook-like projections
beneath the subgenital plate when viewed caudally (Fig. 7) . 8
Cornuplura and Tibicen in part
7' Eighth abdominal sternite not visible beneath subgenital plate or
or lacking hook-like projections (Fig. 23). . . . .. Tibicen
8 (7) Basal cell of forewing green or brown (Fig. 23) . . .. .Tibicen
8' Basal cell of forewing predominantly black (Fig. 13) . . Cornuplura
9 (3') Length of 7th marginal areole more than 2 times the width
(Fig. 14). . . . . . .. . . . .Diceroprocta
9' Length of 7th marginal areole not greater than 2 times the width
(Fig. 9). . . . . . . . . . . . . .. .10
10 (9') Eyes extending laterally away from pronotum; abrupt color change
on face (Fig. 20). . . . . . . . .... .Pacarina
10' Eyes not extending laterally away from anterior edge of pronotum;
no abrupt color change on face (Fig. 9). . . . . .Beameria
11 (1') Medial and cubital veins joined when emerging from basal cell
of forewing (Fig. 11). . . . . . . ... Cicadetta
11' Medial and cubital veins emerging separately from basal cell of
forewing (Fig. 5). . . . . . . . ... ..... .12
12 (11') Nodus of forewing located centrally on the costal margin
(Fig. 5). . . . . . . . . . ....... .14
12' Nodus of forewing located on the distal third of the costal margin
(Fig. 21). . . . . . . . . . . . ... .13
13 (12') Eight marginal cells in forewing (Fig. 21). . .FPatypedia
13' Seven marginal cells in forewing (Fig. 17) ..... .Neoplatypedia
14 (12') Pronotum expanded laterally forming a longitudinal carina for
its entire length . . . . . . . . ... . 15
14' Pronotum rounded laterally, lacking a complete longitudinal
carina . . . . . . . . ... . . . . . .17
15 (14) Marginal cells of forewing about 1/2 the length of ulnar cells
Ul and U2 (Fig. 24). . . . . . . . .Tibicenoides
15' Marginal cells of forewing at least 2/3 the length of ulnar cells
Ul and U2 (Fig. 5) . . . . . . . .... .. ... .16
16 (15') Costal margin of radial cell at least 2 1/2 times longer than
the medial vein anterior to the first branch
(Fig. 18) . . . . . . . . ... .. .Okanagana
16' Costal margin of radial cell less than 2 1/2 times longer than the
unbranched segment of the medial vein (Fig. 12). . Clidophieps
17 (14') First crossvein of forewing oblique (Fig. 15) . .Magicicada
17 First crossvein of forewing perpendicular (Fig. 19) .. . Okanagodes
Beameria Davis (1934, p. 47)
Beameria venosa (Uhler) (Davis, 1934)
= Pruanasis venosa Uhler
= Proarna venosa (Uhler)
There are two species in this genus, Beameria venosa (Uhler) and
B. wheeleri Davis. Both occur in North America north of Mexico.
Davis' remarks are less a description than a comparison of the new
genus with the two genera in which Beameria venosa was previously placed.
Neither Pruanasis nor Proarna occur in North America north of Mexico so
these comparisons are of little value in distinguishing this genus from
other American genera north of Mexico.
General Body Proportions
Length of body 11.0 to 17.0 mm
Expanse of forewings 31.0 to 40.0 mm
The abdomen represents approximately one-half the body length.
Width of head including eyes about the same as the width of the meso-
notum; eyes not prominently projecting laterally; epicranium in a con-
tinuous gentle slope with dorsum of clypeus; width between ocelli about
the same as width between lateral ocelli and eyes; clypeus rounded and
prominently produced beyond the supraantennal plates; dorsum of clypeus
longer than frons; rostrum reaching posterior coxae. Dorsum and venter
of head similarly colored.
Pronotum adpressed laterally except at posterior angles, lacking a
continuous lateral carina. Length of pronotum shorter than length of
mesonotum; mesonotum obscuring central portion of metanotum. Opercula
in males large, broadly rounded, separated medially by a third, centrally
Anterior femora with two prominent spines beneath, sometimes with
a minute third spine distally. Posterior mera triangular, frequently
Forewing venation is illustrated in Figure 9. Both wings hyaline.
First crossvein in forewing perpendicular, second crossvein slightly
oblique, first and second crossveins not parallel; medial and cubital
veins arising separately from the arculus; basal cell clear; nodus cen-
tral; costal margin of forewings minutely spinose.
First abdominal segment in males containing tymbal organs; central
dorsal sclerite modified to resemble a "bow tie." Second abdominal seg-
ment produced anteriorly on each side into a small tymbal covering that
leaves approximately two-thirds to three-fourths of the tymbal organ
exposed; sternal sclerite produced into lateral bulges.
Ninth dorsal segment in males not prominently produced centrally,
lateral margins produced into two upturned points; uncus withdrawable,
notched both caudally and laterally (see Figure 9). Subgenital plate
of female shallowly notched (Figure 9).
The adpressed pronotal margins and non parallel first and second
crossveins in the forewings are primary diagnostic characters. Second-
arily, small size, basically light coloration, and the relatively small
eyes are helpful. Males are easily identified by the shape and extent of
the tymbal covers and the central sclerite separating the opercula
Figure 9. Beameria. A, dorsal view of male; B, ventral view of male
abdomen; C, ventral view of terminal segments of female
abdomen; D, caudal view of male abdomen showing uncus
(after Davis, 1934); E, map showing distribution of the
genus by state. Arrows point to diagnostic characters of
the genus. Species illustrated is Beameria venosa.
Beameria is apparently confined to America north of Mexico. It has
been recorded in the literature from Texas, New Mexico, Arizona, Okla-
homa, Kansas, Colorado, and Nebraska.
Changes in morphology of the nymphs of four of the five nymphal in-
stars of B. venosa are described in Beamer (1928). Beameria venosa adults
frequently inhabit high, dry, rocky hillsides or hilltops where sparse
grass and occasional yucca make up the only vegetation (Beamer, 1928).
Davis (1921b) also reports B. venosa as a grassland species living in
desert grass. It has been reported singing from thistle (Davis, 1921b).
I have personally captured only one specimen of this genus, a B. venosa
singing in Juniperus sp. just south of Sedona, Arizona. There was sparse
grass in the area. It is not unusual for grass or weed inhabiting cicadas
to utilize a higher perch for singing if one is available. Altitudes
recorded for specimens taken in Arizona fall in the range of 1,500 to
2,500 meters. The song of B. venosa is shrill but difficult to hear
beyond a few meters. The light coloration of B. venosa makes it incon-
spicuous among the dry grass it inhabits.
Beamer (1928) records B. venosa as sluggish and not easily frightened
into flight. He describes the nests as arranged serially in a single row
along a stem. Each nest contains three to four white eggs and there may
be a dozen or more nests in a series.
Very little is known about Beameria wheeleri. It is also presumed
to be a grassland species. Dates of capture on specimens of this genus
I have examined range from June 9 to July 14.
Cacama Distant (1904b, p. 429)
Cacama maura (Distant) (Distant, 1904b)
= Proarna maura Distant
The five species of Cacama reported from North America north of
C. californica Davis
C. crepitans (Van Duzee)
C. dissimiZis (Distant)
C. valvata (Uhler)
C. variegata Davis
The genus Cacama was described more completely than most North
American genera. I have added additional features in the following de-
scription, and Distant's remarks are referred to in the appropriate
sections. In 1919, Davis considered the genus as a whole, adding four
species to the list of Cacama and providing a key to Cacaina identifica-
tion. No new species have been described since that time.
General Body Proportions
Length of body 21.0 to 30.0 mm
Expanse of forewings 64.0 to 83.0 mm
Cacama are robust insects with the abdomen representing approximately
one-half the body length. The length of the abdomen is less than two and
one-half times the width of the mesonotum.
Width of head, including eyes, much narrower than the width of the
mesonotum; eyes in a line with or extending slightly beyond the anterior
edge of the pronotum; vertex of epicranium horizontal, frons sloping or
vertical, forming an obtuse to right angle with the dorsum of the clypeus;
distance between lateral ocelli about the same to slightly less than dis-
tance between lateral ocelli and eyes; clypeus moderately truncate, ex-
tending only slightly beyond the supraantennal plates and giving the head
a blocky appearance; dorsum of clypeus shorter than frons; "rostrum
reaching or passing the posterior coxae" (Distant, 1904b).
Pronotum ampliated posteriorly, strongly arched anteriorly,
lateral carinae complete but minimal on the anterior half, lateral edges
smooth; length of pronotum shorter than mesonotum; "Mesonotum somewhat
convexly gibbous" (Distant, 1904b); mesonotum obscuring central portion
of metanotum; opercula in males large, "about half the length of abdomen
above, broad, their apical margins convexly rounded, their lateral margins
almost straight" (Distant, 1904b), overlapping medially.
Anterior femora with two prominent spines beneath; posterior mera
short, sometimes blunt, usually equilateral.
Forewing venation is illustrated in Figure 10. Both "wings (ex-
cepting base) hyaline . apical areas [of forewings] eight in number,
the two lowermost small, subquadrangular" (Distant, 1904b); first and
second crossveins may or may not be parallel; medial and cubital veins
arising separately from the arculus; nodus central; costal margin of
forewings not noticeably spinose, may have a few tiny spines proximal
"Abdomen short, broad, convex above" (Distant, 1904b); first ab-
dominal segment in males containing tymbal organs, central dorsal sclerite
with two rounded lateral lobes (Figure 10); second dorsal segment in
males produced anteriorly on each side into a complete tymbal cover;
"lateral margins of the tympanal coverings subparallel to the abdominal
margins" (Distant, 1904b); lateral bulge inconspicuous.
Figure 10. Cacama. A, dorsal view of male; B, ventral view of male
abdomen; C, ventral view of terminal segments of female
abdomen; D, caudal view of male abdomen showing uncus (after
Davis, 1919b); E, map showing distribution of genus by state.
Arrows point to diagnostic characters of the genus. Species
illustrated is Cacamna valvata.
Ninth dorsal segment of males rounded laterally, produced into a
central spine dorsally; uncus withdrawable, simple (Figure 10). Sub-
genital plate of female shallowly notched centrally, smooth, not sinuous
Cacama is easily recognized by its stout gestalt. The combination
of the mesonotum overlapping the metanotum and the narrow, blocky head
will identify both males and females. In addition the abdomen is broadly
Cacama extends from Yucatan, Mexico, northward to the southwestern
United States. Within the United States it has been captured in Texas,
Oklahoma, New Mexico, Colorado, Arizona, Utah, Nevada, and Southern
Cacama is the only cicada genus found north of Mexico that might be
considered a mimic. Their black or nearly black, robust bodies super-
ficially resemble bumble bees. They initiate flight by hovering away
from a perch rather than jumping into flight. The behavior of Cacama
when captured in a net alSp is reminiscent of a bumble bee. Live Cacama
are marked with a light colored pruinosity usually not seen in museum
specimens. Heath, Wilkin, and Heath (1972) showed that C. valvata from
Arizona extended activity in the sun by orienting the reflective surface
of the white, pruinose venter toward the sun's rays thereby diminishing
the rate of heat gain. I have consistently found peak activity in
C. vatvata and C. dissimilis to range between 8:00 AM and 1:00 PM mountain
standard time. Beamer and Beamer (1930) reported a similar activity
period for C. crepitans. No data are available on the other two American
species found north of Mexico. In Arizona I have captured Cacama from
the desert plant formations to the pinyon-juniper plant formation. Al-
though Beamer and Beamer (1930) reported capturing C. vatvata from a
yucca, sagebrush, and cedar habitat, and again, from a "habitat of small
cedars" and C. dissimilis from palo verde, I have found the same two
species almost always associated with some species of Opuntia. All
other reports in the literature are from Opuntia. Cacama will sing from
higher perches when these are available. I have observed them on barbed-
wire fences and telephone poles. I cannot help but wonder if some
Opuntia was not present in the habitats reported by Beamer and Beamer.
Cacama valvata was observed to spend the night feeding near the base of
clusters of the flat pads of prickley pear by Heath, Wilkin, and Heath
(1972). Beamer and Beamer (1930) found that wetting nests of C. valvata
laid in cane cactus or exposing the nests to a cold morning rain approxi-
mately three months after egg laying precipitated the emergence of the
nymphs. They concluded that moisture, and consequently reduced soil
temperatures, would increase the survival of nymphs from arid, hot
habitats. Cacama has been given the popular name "cactus dodger." The
song of Cacama resembles the sound of a chain saw cutting logs.
Cacama dissimiiis was described by Distant (1881) from Mexico. I
have seen the type specimen in the British Museum and believe it to be a
different species from that commonly identified as C. dissimilis in the
southwestern United States. Dr. Thomas Moore of the University of
Michigan has a manuscript in preparation naming the U.S. species. Remarks
on C. dissimitis given above will be pertinent to this new species when
Dr. Moore's description appears.
Cicadetta Kolenati (1857, p. 417)
Cicadetta montana (Scopoli) (Kolenati, 1857
= Cicada montana Scopoli
= Cicada orni (nec Linne)
= Tettigonia montana (Scopoli)
= Cicada pygmea Olivier
= Cicada tibialis Latreille (nec Panzer)
= Cicada angelica Samouelle
= Cicada haematodes Linnd (nec Scopoli)
= Cicadetta Amyot (nonbinomial)
= Tettigonia haematodes (Linne) (nec Sco
= Tettigonia dimidiata Fabricius
= Cicada schaefferi Gmelin
= Cicada parvuZa Walker
= etcampsalta montana (Scopoli)
= Cicada saxonica Hartwig
= Tettigia montana (Scopoli)
Four species and one named variety occur in North America north of
Mexico. The species are
C. calliope (Walker)
C. camerona (Davis)
C. kansa (Davis)
C. texana (Davis)
Alexander (unpublished) considers the geographical variety Cicadetta
calliope floridensis (Davis) a distinct species. Cicadetta calliope
floridensis is distinguishable by color and patterning.
At least six generic names have been used for all or part of the
genus Cicadetta. Metcalf (1963) catalogs the names and synonomies while
Chen (1943) gives a brief history of the status of the names. They each
come to different conclusions regarding the proper name. References to
North American species prior to Metcalf (1963) use the generic name
Melampsalta. In 1847, Amyot published descriptions of Melampsalta,
Cicadetta, and Tettigetta in that order. Although he included the range
of body lengths, the major part of the descriptions deals with color
patterning and not anatomical features. In this sense each description
resembles a species description and not a generic description. Amyot
apparently intended these names as part of a monomial system of nomen-
clature and meant to refer only to what we now consider single species.
Kolenati (1857) picked up Amyot's names and gave them subgeneric status
using the species described by Amyot as the types and attributing the
names to Amyot. In Kolenati's work the names appear in the following
order: Cicadetta, Tettigetta, Melampsalta. Kolenati's descriptions are
morphological. According to Metcalf (1963) Stal was the first to synonomize
the three in 1861. He, however, recognized Amyot as the author and used
the name Melampsalta. Since that time various authors have recognized
the same synonomy. The favored name depends on whether Amyot or Kolenati
is recognized as author. Other authors have attempted to separate por-
tions of the genus into new genera, but most of these names have not been
generally accepted. The current status rests on the argument that Amyot
did not intend to describe part of a binomial name. The species de-
scribed by Amyot already had valid specific names. According to article
llc of the International Code of Zoological Nomenclature, a name does not
become available unless the author consistently applied the principles of
binomial nomenclature in the work in which the name is published. Since
Amyot was proposing monomial names, the names cannot be attributed to
him. Kolenati, therefore, becomes the author. The subgenera described
by him constitute the establishment of the genus and Cicadetta becomes
the valid name. This is apparently the view taken by Metcalf and now
accepted on a world-wide basis. I have consulted the descriptions of
Amyot (1847), Kolenati (1857), Ortega (1957), and Kato (1961). The
following description is intended to encompass only those species found
in North America north of Mexico.
General Body Proportions
Length of body 12.0 to 20.0 mm
Expanse of forewings 25.5 to 40.0 mm
The abdomen represents more than one-half the body length.
Width of head, including eyes, subequal with the width of the meso-
notum; eyes prominent, may or may not project laterally beyond the
anterior angles of the pronotum; vertex of epicranium horizontal; frons
sloping, forming a broad obtuse angle with the dorsum of the clypeus;
vertex and clypeus centrally sulcate; dorsum of clypeus somewhat centrally
depressed; distance between lateral ocelli about the same or narrower
than distance between lateral ocelli and eyes; clypeus and supraantennal
plates forming the more or less continuous sides of a broad triangle with
the sides notched where the clypeus and supraantennal plates meet; dorsum
of clypeus shorter than frons; rostrum reaching to or barely beyond the
Pronotum strongly arched laterally, not strongly ampliated, but
usually having continuous carinae where the notal and pleural regions
meet, lateral margins smooth, posterior angles lobately dilated; length
of pronotum shorter than mesonotum; posterior margins of metanotum com-
pletely visible behind mesonotum; opercula in males well developed,
Anterior femora with at least three distinct spines; a fourth, less
developed, distal spine is usually present. Posterior mera narrowly
drawn out at the tip.
Forewing venation is illustrated in Figure 11. Wings hyaline.
First and second crossveins in the forewings perpendicular and usually
parallel; medial and cubital veins coalescent where they arise from the
arculus; nodus more or less central, may be slightly further from the
base of the wing than from the apex; costal margin of forewing may have
several very fine spines.
First abdominal segment in males containing tymbal organs, posterior
margin of central dorsal sclerite concave centrally; anterior margin of
second dorsal sclerite curved smoothly entad laterally behind the tymbal
organs; lateral bulge on sternite well developed in both sexes; tymbal
The ninth dorsal segment of the males may or may not project into
an acute point dorsally, posterior margin with a caudally projecting,
ventral lobe, a third lobe may be present along the ventral margin; uncus
retractable, short, bifurcated with the two parts diverging outward from
a caudal view and the apices subacute, curved inwardly from a lateral
view (Figure 11). Subgenital plate of female with a large simple,
triangular notch extending nearly to the base of the sternite (Figure
Figure 11. Cicadetta. A, dorsal view of male; B, ventral view of male
abdomen; C, ventral view of terminal segments of female
abdomen; D, lateral view of extended male pygofer, caudal
view of uncus (after Davis, 1920); E, map showing distribu-
tion of the genus by state. Open circle is a questionable
report. Arrow points to the coalesced media and cubitus
emerging together from the arculus. Species illustrated is
The coalesced medial and cubital veins in the forewings emerging
together from the apex of the basal cell will separate Cicadetta from all
other genera found in North America north of Mexico. In the other genera
in this region the medius and cubitus leave the arculus separately.
Cicadetta is widely distributed throughout all the world's biogeo-
graphic regions except the Neotropical. Within the Nearctic region it is
found throughout the southeastern United States, as far west as eastern
Colorado, and north to Nebraska, Iowa, Illinois, Ohio, and Virginia.
Cicadetta camerona and C. texana have thus far been reported only from
Texas. Cicadetta caZliope fZoridensis is found in Florida and Georgia.
Davis (1930) records C. kansa from Kansas, Colorado, Oklahoma, and Texas.
Cicadetta calliope calliope is the most widely distributed species. It
is found in all of the above states except Oklahoma. Additional state
records are from Missouri, Arkansas, Louisiana, Mississippi, Alabama,
Tennessee, and North Carolina. There is a doubtful report from New
Jersey. There are no records from Indiana, Kentucky, South Carolina, or
West Virginia, but part or all of these states fall within a more
generalized distributional pattern, particularly if the Ohio designation
is accurate. These four states represent likely extensions of the
Cicadetta is the most widespread genus in the family Cicadidae.
Several factors may influence the widespread dispersal of Cicadetta,
among which are the age of the genus, size of the insects, and the period
of development. There are, at present, no known fossil Cicadetta to give
us a clue to the age of the genus. If, however, Cicadetta does represent
a single phylogenetic line, the wide distribution would seem to infer
that it is a very old genus. The majority of Cicadetta species are small.
Beamer (1928) remarks on the effects of wind in lengthening the flight of
C. calliope in Kansas. Because of the small size of the adults, wind may
well be a factor in Cicadetta dispersal. Myers (1929) suggests that the
entire New Zealand cicada fauna derives from a single immigrant species
of Cicadetta. Whether this invasion resulted from rafting, wind, or a
combination is difficult to say. Most cicadas spend a lengthy immature
period underground. Although some cicadas have received popular names
such as "annual cicada" and a two year life cycle has been proposed
frequently, the shortest documented life cycle of a cicada is the four
year cycle of Cicadetta calliope (Beamer, 1928). If a relatively short
life cycle is characteristic of the genus, more generations of Cicadetta
have been available for dispersal than have been available among those
cicada genera of equal age with a longer subterranean existence.
The probable habitat of the species of Cicadetta in North America is
perennial grasses and weeds. Like other cicadas, they will utilize the
higher perches of shrubs or trees when singing. Beamer (1928) studied
C. calliope calliope in detail in Kansas. This species is abundant
frequently in meadowlands as long as fires or serious flooding do not
prevail. Females oviposit in live material and the eggs wither if the
branch dies. Nests are placed in a single row on pithy stems. Matura-
tion of the eggs takes about two months. The song of the males is
audible for only a distance of a few meters. Adults are preyed upon
extensively by both robber flies (Asilidae) and spiders. Eggs are
parasitized by chalcids of the genus Syntomaspis. Cicadetta in North
America are active as adults from late April to August. Beamer (1928)
also observed the behavior of the nymphs in glass sided cages. Cells are
generally about one inch long. If the host plant dies the cell may be
extended for several inches as the nymph searches for a new host. Loose
dirt is dug from the front of the cell with the forelegs. It is then
packed into a ball between the forelegs and the postclypeus and sub-
sequently carried to the rear of the cell. Beamer found nymphs in the
field usually six to ten inches beneath the surface. A disturbed nymph
will rear and prepare to fight the offender with its fore femora. Beamer
described the morphology of the five nymphal instars of C. calliope
Clidophleps Van Duzee (1915, p. 31)
CZiodphZeps distant (Van Duzee) (Van Duzee, 1915)
= Okanagana distant Van Duzee
= Okanagana pallida Van Duzee
= Okanagana paZlidus Van Duzee
= Clidophleps pallida (Van Duzee)
Seven species and one named variety of Clidophleps occur in North
America north of Mexico. The species are
C. beameri Davis
C. bZaisdeZli (Uhler)
C. distant (Van Duzee)
C. rotundifrons (Davis)
C. tenuis Davis
C. vagans Davis
C. wright Davis
There is one named variety of ClidophZlps distant.
The original description of Van Duzee (1915) compares features of
Clidophleps with similar features in Okanagana and Platypedia. A few
diagnostic features are given. Simons (1954) keys and illustrates a
generalized wing and the terminal abdominal segments of the males of the
California species. No other general work on the genus has appeared, and
no complete description has been written.
General Body Proportions
Length of body 19.0 to 26.5 mm
Expanse of forewings 44.0 to 71.0 mm
The abdomen represents approximately one-half the body length. In
addition the long, slender body has a "bullet or torpedo-shaped"
Width of head, including eyes, narrower than the mesonotum; eyes in
a line with or slightly wider than the anterior angles of the pronotum;
vertex of epicranium horizontal, centrally sulcate; frons steeply slanted
to vertical, forming an obtuse to right angle with the dorsum of the
clypeus; distance between lateral ocelli greater than distance between
lateral ocelli and eyes; clypeus broadly rounded, extending beyond the
supraantennal plates, centrally sulcate apically and ventrally; dorsum of
clypeus about the same to slightly longer than frons; rostrum reaching
Pronotum steeply arched laterally; lateral carinae continuous but
varying between minimal and ampliated on a species specific basis;
lateral edges usually smooth, posterior angles dilated; length of pro-
notum shorter than mesonotum; mesonotum with a raised striated surface
located next to the base of the forewing when folded against the body;
posterior margin of metanotum completely visible behind mesonotum;
opercula in both sexes small, transverse, rounded, widely separated,
not extending beyond the base of the abdomen.
Anterior femora with two prominent spines; posterior mera broadly
Forewing venation is illustrated in Figure 12. Wings hyaline, basal
cell clear; first and second crossveins in forewings usually oblique and
parallel; third marginal cell at least two-thirds the length of second
ulnar cell; costal margin of the radial cell less than two and one-half
times longer than the medial vein anterior to the first branch; radial
cell of forewing frequently expanded; the transverse crossvein between
M8 and Cu usually thickened; medial and cubital veins arising separately
from the arculus; the cubital vein with at least a slight arch; nodus
central; costal margin of forewing with a few very minute spines.