Title: Genera of American cicadas north of Mexico
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Title: Genera of American cicadas north of Mexico
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Language: English
Creator: Heath, Maxine Shoemaker, 1935-
Copyright Date: 1978
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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.





ABSTRACT . . . . . . .



Life Cycle . . . . .
The Problem. . . . . .



Order . . . . . .
Suborder . . . . . .
Superfamily . . . .
Family Cicadidae . . . .
Subfamilies . . . .
Tribes . . . . . .



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


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



Maxine Shoemaker Heath

August, 1978

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

that time.

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.

Life Cycle

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

to desiccation.

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.


The Problem

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.



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

collection data.

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

in hand.

Unless otherwise indicated in the text, the terminology used in

this paper conforms with that found in Torre-Bueno, A Glossary of

Entomology, 1962.




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).

Family Cicadidae


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

DeLong (1971).


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-

jointed rostrum.


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

with them.


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
one stem.

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.


Table 1

A comparison of Metcalf's and Boulard's family and subfamily groupings of
the genera of American cicadas north of Mexico.






Cornup ura

















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:


Subtribe Tibicenaria


Cornup lura


















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

(Figure 1).


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

Figure 1.



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

the head.

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

DuPorte (1977).

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.

Figure 2.




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.

Figure 3.



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

Figure 4.



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

(Kramer, 1950).

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.

Figure 5.




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
Tibicen duLri.

Figure 6.



side of the abdomen. The terminal segments of the abdomen are discussed

under the heading "sexual characteristics."

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).

Figure 7.





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.

Figure 8.








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.




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)

Type Species

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.

Descriptive History

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

projecting schlerite.


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.

Sexual Characteristics

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).

Diagnostic Characters

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).

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)

Type Species

Cacama maura (Distant) (Distant, 1904b)

= Proarna maura Distant


The five species of Cacama reported from North America north of

Mexico are

C. californica Davis

C. crepitans (Van Duzee)

C. dissimiZis (Distant)

C. valvata (Uhler)

C. variegata Davis

Discriptive History

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

to body.


"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.





Sexual Characteristics

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

(Figure 10).

Diagnostic Characters

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

rounded caudally.


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)

Type Species

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.

Descriptive History

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

intermediate coxae.


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,

separated, rounded.


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

organs exposed.

Sexual Characteristics

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
C. kansa.




Diagnostic Characters

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

reported distribution.



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

in detail.


Clidoph eps


Clidophleps Van Duzee (1915, p. 31)

Type Species

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.



Descriptive History

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

intermediate coxae.



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.

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