JAMES F. GREE A Revision of the

w Nearctic Species of the

Genus Psammotettix

(Homoptera: Cicadellidae)

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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 74

James F. Greene A Revision of the

Nearctic Species of the

Genus Psammotettix

(Homoptera: Cicadellidae)

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

»97>
ABSTRACT

Greene, James F. A Revision of the Nearctic Species of the Genus Psammotettix.

Smithsonian Contributions to Zoology, number 74, 40 pages. 1971.—Nineteen

Nearctic Psammottetix species were treated. Four of the species are Holarctic.

Fifteen species were redescribed and four new species were described. The limits

of the genus were redefined.

Interspecific, intraspecific, and individual variations were discussed. The first

complete written keys to males and females of Nearctic species were prepared.

Important diagnostic characters for each species were illustrated. New specific

taxonomic characters in the genus, lobes at the base of the first valvula, were

employed and illustrated.

Possible avenues for future investigation, host-plant data, disease vector

relationships, and parasite data were included in the text.

Official publication date is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year.

UNITED STATES GOVERNMENT PRINTING OFFICE

WASHINGTON : 1971

For sale by the Superintendent of Documents. U.S. Government Printing Office

Washington. DC. 20402 - Price 50 cents (paper cover*
James F. Greene A Revision of the

Nearctic Species of the

Genus Psammotettix

(Homoptera: Cicadellidae)

Introduction

Since there has been a growing interest, both economically and taxonomically, in the genus Psammotettix Haupt in the last decade, the purpose of this

paper is to revise the Nearctic species of this genus.

Workers have experienced difficulty in resolving a

species complex known to European workers as the

striatus group. They were finding individual variation in the aedeagal characters which had been used

as primary species diagnostic characters (Moravskaia, 1956).

When this study began there was a need for a

closer comparison of Nearctic and Holarctic specimens of Psammotettix in order to identify objectively Nearctic specimens of Holarctic species that

had been described from Europe. The fact that

some of the Holarctic species were members of the

unresolved P. striatus complex added to the problem. In addition there was a need for a revision of

the Nearctic species of the genus to complement a

revision of the Palaearctic species by Ribaut (1938).

The present work has resulted in the preparation

of keys to male and female species of Nearctic

Psammotettix. Fifteen species were redescribed, four

new species were described and named, and it was

determined that Psammotettix striatus (Linnaeus)

sensu Ribaut did not occur in the Nearctic region.

Concerning the difficult striatus complex mentioned

above, that portion of the problem related to the

Nearctic species has been resolved if Ribaut's interpretation of P. striatus (Linnaeus) is accepted. Four

James F. Greene, Xorlh Carolina Department of Agriculture,

Raleigh, Xorth Carolina 27602.

Nearctic species groups or complexes and one heterogeneous group were discovered and illustrated.

Intraspecific variation in characters previously frequently interpreted as specific have been discovered

and illustrated in this work. Female genitalic characters hitherto unknown were discovered. These

characters were very helpful in establishing species

relationships, but they have posed problems in determining relationship when they have not paralleled the aedeagal characters of the male.

The work on Nearctic Psammotettix is not yet

complete. Additional distribution and host-plant

data are needed. The most important work that remains to be done is an intensive study of Psammotettix on a Holarctic basis, utilizing die newly discovered female genitalic characters as well as the

other characters employed in this paper.

Psammotettix species are extensively listed in the

literature as feeding on grasses. The type of the

genus, Psammotettix maritimus (Penis), feeds on

Convolvulus (Haupt, 1929). Psammotettix lividellus

(Zetterstedt) in North Carolina appears to be host

specific for Poa pratensis Linnaeus. Moravskaia

(1956) lists P. striatus (Linnaeus) as being adapted

to young crops and P. confinis (Dahlbom) as polyphagous. In addition to previous records, species of

Psammotettix have been collected on celery, corn,

and sugar beet, reported here for the first time.

Psammotettix species have been reported as being

vectors of virus diseases of sugar beet (Muraviov,

1930), wheat mosaic (Moravskaia, 1956), and a virus

disease of potatoes (von Archimowitsch, 1952).

Haupt (1929) described the genus Psammotettix

for Athysanus maritimus Perris, 1857 (type locality,

I
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

southern France by the sea, on Convolvulus soldonella). Zachvatkin (1933) described the genus Ribautiellus for Deltocephalus striatus (Linnaeus),

1758 (type locality, Europe). Ribaut (1938) synonymized Psammotettix Haupt, 1929, and Ribautiellus

Zachvatkin, 1933.

European workers such as Ossiannilsson (1937,

1938, 1941), Ribaut (1938), Wagner (1939), Dlabola

(1945, 1954), and Remane (1965) added species until

a total of 46 Palearctic species are included in the

genus (Remane, 1965). The most comprehensive

works on the Palearctic species of the genus are

those of Wagner (1939) and Ribaut (1938).

American workers Osborn (1891), Ash mead

(1904), Oman (1931), and DeLong and Davidson

(1934, 1935) described a number of species in Deltacephalus and Laevicephalus, which Oman subsequently (1947, 1949) placed in Psammotettix

Haupt.

There was some confusion among American

workers as to the proper identity of Nearctic specimens of Holarctic species. DeLong and Davidson

(1935) illustrated Psammotettix lividellus (Zetterstedt) as Laevicephalus striatus (Linnaeus). True

P. lividellus (Zetterstedt) was also treated as Deltocephalus affinis Gillette and Baker (Beirne, 1954),

and true Psammotettix alienus (Dahlbom) was

described as Laevicephalus latipex DeLong and

Davidson (1935). Oman (1949) synonymized latipex with Psammotettix striatus (Linnaeus) and

listed ten Nearctic species of Psammotettix. Knull

(1954) named and described three new species of

Nearctic Psammotettix: P. obesus, P. revae, and P.

dentatus. Beirne (1954) redescribed Psammotettix

lapponicus (Ossiannilsson) and recorded it from

Chesterfield, Northwest Territories. He redescribed

Psammotettix con finis (Dahlbom) and recorded it

as a Nearctic species. Deltocephalus harrimani Ashmead (Ashmead, 1904), one of the ten species

listed by Oman (1949), was synonymized with confinis by Beirne. Beirne (1954) redescribed Psammotettix alienus (Dahlbom) and synonymized it with

Deltocephalus latipex DeLong and Davidson and

with Psammotettix striatus DeLong, 1948 (not Linnaeus). Beirne (1954) redescribed Psammotettix

lividellus (Zetterstedt) and synonymized it with

Deltocephalus affinis Gillette and Baker and with

Psammotettix striatus DeLong, 1948 (not Linnaeus). McClure (1943) reported Psammotettix

asper from Manitoba.

Beirne (1956) redescribed Psammotettix cephalotes (Herrich-Schaeffer) and synonymized it with

Psammotettix obesus Knull.

Combining Oman's (1954) checklist, Knull's

(1954) three new species, McClure's (1943) report

from Manitoba, and accounting for Beirne's (1954,

1956) synonymies, there were 15 Nearctic species of

Psammotettix recorded in the literature in 1956, not

including what then were considered to be synonyms. This was the situation when this study was

started.

Dried, pinned specimens of Psammotettix from

the United States, Canada, Europe, and Asiatic

Russia were used in this study. Living specimens

of Psammotettix lividellus (Zetterstedt) from North

Carolina were examined, but all descriptions arc

based on thied or cleared specimens.

The methods of Young (1952) were followed in

preparing tiie specimens with the following exceptions. The genital capsule was treated in 17 percent

K.OH at room temperature for two hours rather

than in 10 percent KOH heated to the boiling point.

The specimens were not stained with acid fuchsin,

and the glycerine in which they were observed was

not acidulated.

The genitalic parts were observed under transmitted light of varying intensities with both dissecting and compound microscopes. In the males, drawings of genitalic structures were made at a magnification of 160x with use of a camera lucida. A

wide-angled ocular was used in order to get the

complete structure to be drawn in the field of view.

When making drawings of the aedeagus in caudodorsal aspect, the aedeagal shaft was placed as

nearly perpendicular to the axis of the barrel of the

microscope as possible. This was difficult in specimens with curved aedeagal shafts. Drawings were

made of the aedeagus in the lateral aspect, which

give a more accurate measure of the shaft length.

The aedeagal shaft apex was studied in cross section by making "end-on" observations of the longitudinal axis of the shaft. Only a few such observations are incorporated in this paper. This technique

gives a more complete picture of the aedeagal shaft.

Cross-sectioning the aedeagal shaft might yield additional taxonomic information. In the female subtle

differences can regularly be observed between

species after clearing. The seventh abdominal stern-
NU.MBKR 74

urn of the female was drawn while attached to the

abdomen, after being cleared in 17 percent KOH.

In dried, unrelaxecl specimens it is often not possible to see constant differences between species. In

order to get a constant orientation under the microscope, for drawing purposes, the sternum was oriented in such a way that the median part of the

posterior and the anterior edge were in sharp focus

at the same setting of the microscope. These drawings were made with camera lucida at 160x magnification.

After clearing the female genital capsule in KOH,

the capsule was removed from the abdomen for

observation and illustration. To do this it was

necessary to rupture the membrane connecting the

posterior edge of the seventh abdominal sternum

and the base of the first valvulae. This was accomplished by probing gently with a fine point under

and to the base of the seventh sternum and the first

valvulae. The membrane was thus torn from its

connection with the base of the first valvulae. The

genital capsule was separated from the eighth

tergum with a needle.

be made accurately for subbrachypterous forms

because of the tendency of the abdominal apex to

bend dorsally in dried specimens.

All length measurements of parts of the specimens

were made parallel to the median longitudinal axis

of the specimen in the dorsal aspect. The length of

crown was measured medially and also tangentially

to the mesal border of the eye. The width of the

head, including eyes, was measured between the

outermost parts of the eyes.

As is true with many leafhopper species, it is

usually possible to see differences in the head shape

between species in this group, but it is hard to

document or describe them. The curvature of the

anterior edge of the crown, when viewed in dorsal

aspect, seems to be the most variable character.

There is much intraspecific variation in this character.

Unusual variations were studied for possible unnatural causes. In several specimens the capsule in

which the genitalia were stored had lost the preserving fluid. This dry storage resulted in some

distortions that could be misinterpreted as being

The bases of the first valvulae were illustrated NUMBKR 74iracters of taxonomic importance.

ventral aspect from the intact genital capsule. The

bases of the first valvulae were oriented as nearly

perpendicular to the axis of the barrel of the microscope as possible. The basidorsal median lobes and

the articulating lobes were drawn or observed.

Drawings of the head, pronotum, and scutellum

were made in the dorsal aspect with a stereoscopic

microscope at 160x magnification and a camera

lucida. In order to get a constant attitude, the head

was arranged so the ocelli were barely visible at

the front edge of the crown. Observations, drawings,

and measurements were made in this position under

reflected light. Note that longitudinal head measurements in this paper were made while the specimen was in this position. Measurements made with

the long axis of the specimen perpendicular to the

line of sight of the microscope would have a different value. Measurements of total length and greatest

width were made with long axis of the specimen

perpendicular to the line of sight. Measurements

of length of macropterous forms were made from

apex of crown to apex of fore wings. Total length

measurements were made from apex of crown to tip

of hindmost setae of pygofer in subbrachypterous

forms. Very precise length measurements could not

Differences of possible taxonomic importance

were noted in the setal pattern on the caudal apex

of the style, but not incorporated in this paper.

The general morphology of Psamrnotettix is not

discussed in this paper, which is concerned for the

most part with characters of taxonomic importance.

The morphology and terminology used in this paper

are from Oman (1949), Young (1952), and Snodgrass (1935).

The anterior edge of the crown in dorsal aspect

varies from blunt as in Psammotettix attenuens

(DeLong and Davidson) (Figure 1A) to pointed

as in Psammotettix dcntatus Knull (Figure 1B) .

This character varies both interspecifically and intraspecifically. A median longitudinal sulcus, the

base of the epicranial suture, extends from the

posterior edge of the crown anteriorly for a distance about equal to the longitudinal length of

the crown next to the mesal border of the eye.

The area of the pronotum varies relative to the

area of the crown. The relative median longitudinal

lengths of the crown and pronotum give an index

of their relative area. The anterior edge of the pronotum is raised or ridged dorsally (Figure 1). The
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

posterior edge of this raised portion outlines the

anterior edge of the scutum of the alinotum which

lies beneath. The raised portion is above the cervix.

The area above the alinotum is covered with circular pits. The pits are often found to be confined

to the anterior edge of this area.

The wings of Psammotettix vary in length. Psammotettix species are macropterous, subbrachypterous, or brachypterous. The distance the hind wings

extend posteriorly varies as compared to the distance the fore wings extend posteriorly. This character is used in the key to the females.

XROWN

PRONOTUM

CIRCULAR PITS.._/ _ Z

ANTERIOR EDGE

SCUTELLUM

BASIDORSAL MEDIAN LOBE

ARTICULATING LOBE

FIRST VALVULA__

FIRST VALVIFER

FIGURE 1.—Head and thorax of Psammotettix in dorsal aspect and lust valvulac and fust valvifcr

in ventral aspect: A, head and thorax of P. attenuens (Ik*Long and Davidson) ; B, head and

thorax of P. dentetus Knull: c. first valvulac and first valvifer of P. totalus iPcLong and

Davidson).
NUMBER 74

The female genitalic characters do not delimit

species as precisely as do the aedeagal characters

of the male. The basal lobes of the first valvula

vary in relative position, length, and degree of

sclerotization and can be used in conjunction with

wing length and shape of the posterior edge of the

seventh abdominal sternum to separate species.

These two lobes (Figure lc) have not been previously named. One lobe that is at the base of the

first valvula in a dorsal position is called the

basidorsal median lobe; the other that is laterad

of the basidorsal median lobe and appears to articulate with the first valvifer is called the articulating

lobe.

The most reliable taxonomic characters in Psammotettix are on the male aedeagus. The aedeagal

shaft varies from linear (Figure 11A) to expanded

at apex (Figure 17c), from entire (Figure lie) to

dentate (Figure 18c), and from symmetrical to

asymmetrical (Figure 3c) in caudodorsal aspect. In

lateral aspect the shaft varies from straight (Figure

4D) to curved (Figure 10D) to bisinuate in lateral

aspect. The connective varies interspecifically from

linear with basal arms contiguous (Figure 7c) to

Y-shaped with basal arms separated (Figure 2A) .

The caudal end of the shaft of the connective varies

from truncate (Figure 7c) to bifid (Figure 20c) in

dorsal aspect.

Acknowledgments

The writer is indebted to many people who have

directly or indirectly made completion of this paper

possible. Acknowledgment and gratitude are extended to the following who have been especially

helpful.

Dr. David A. Young, chairman of the author's

graduate committee, initiated the project, made preliminary studies, and obtained specimens; he gave

direction, advice, criticism, and encouragement. The

other members of the author's graduate committee

at North Carolina State University, Dr. M. H.

Farrier of the Department of Entomology and Dr.

J. VV. Hardin of the Department of Botany, helped

the author through teaching, advice, encouragement, and review of the work. The staffs of the

Department of Botany and the Department of Entomology made suggestions and provided equipments as needed. The writer's colleagues in the

North Carolina Department of Agriculture were

patient and helpful. Mr. J. A. Harris was particularly understanding in the arrangement of work

schedules and Dr. D. L. Wray gave advice and

provided equipment.

The following people loaned specimens from

their respective collections: Dr. D. M. DeLong of

Ohio State University, specimens from his personal

collection; Dr. Paul H. Freytag and Dr. C. A.

Triplehorn, specimens from the Ohio State University collection; Dr. J. P. Kramer, specimens from

the National Museum of Natural History; Dr. C. D.

Michener, specimens from the Snow Entomological

Museum at the University of Kansas; Dr. W. R.

Richards, specimens from the Canadian National

collection; Dr. Edward S. Ross, specimens from the

California Academy of Science collection: Dr. A.

Soos, specimens from the Hungarian Natural History Museum, Budapest; Dr. D. A. Young, specimens

from the North Carolina State University collection; Dr. VV. Wagner, specimens from the Zoological

Institute in Hamburg.

Genus Psammotettix Haupt

Psammoteltix Haupt, 1929. pp. 173-286. [Type by original

designation, Athysatius mnritimus Pcrris, 1857.]

Ribauticllus Zachvatkin, 1933. pp. 262-272. [Type by original

designation, D lellocephalus) slrintus Linnaeus.]

From elongate, slender to elliptical in dorsal aspect.

Length 2.80 to 4.20 mm, greatest width 0.86 to

1.30 mm. Head, pronotum, scutellum, and abdomen

with ground color creamy, occasional amber or

cinereous. Head, pronotum, and scutellum with or

without brown markings. Abdomen marked with

black, brown, orange, or red. Wings transparent,

subhyaline, or opaque; with brown to smoky markings. Hind wing veins clear or darkened.

Head including eyes slightly wider than pronotum. Crown produced, with median length

greater than length tangent to niesal border of eye;

with median sulcus extending anteriorly 0.56 to 0.90

length from pronotum to apex. Crown with anterior

edge a median blunt curve with radius of curvature

varying from equal to one-fifth the radius of curvature of the pronotum in dorsal aspect. Crown granulate along front margin, glabrous discally; center

of disc depressed in dried specimens. Clypeus with

horizontal brown stripes connected longitudinally

at or near median line in cephalic aspect. Clypellus
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

nearly parallel-sided, gradually tapering to a

rounded point distally, longer than lorum; transclypeal suture distinct.

Pronotum almost as large as crown, including

eyes; from 0.8 to 1.6 as long as crown measured

longitudinally in dorsal aspect, with anterior portion slightly raised beginning at anterior edge of

scutum. With or without six longitudinal brown

stripes. Scutellum almost covered by pronotum, exposed portion triangular, with straight, crescentiform, or undulate transverse suture. Wings macropterous, subbrachypterous, or brachypterous. Hind

wings terminating much before or nearly at apex

of fore wings when in resting position. Wings varying in length, attaining or not, or exceeding apex

of abdomen when in resting position. Outer anteapical cell not always stalked.

Female abdomen with seventh sternum wider anteriorly; anterior edge truncate; posterior edge truncate, emarginate, or with median projection. Ventral

intersegmental membrane connecting posterior edge

of seventh sternum with base of first valvulae rigid

posteriorly, appearing as a closely appressed flap

of the sternum, visible as wrinkles through cleared

sternum (Figure 4B) . Female pygofer setiferous,

cephalodorsal edge with well-developed, horseshoeshaped, internal apodeme. First valvula with a basidorsal median lobe flanked by (usually) sclerotized

folds or articulating points where the first valvifer

articulates (Figure lc).

Male valve wider anteriorly, large compared to

exposed tips of plates. Pygofer with group of setae

near point of attachment to tenth tergum which is

a narrow C-shaped sclerite with arms of C attached

to pygofer. Male plates thickened, each with mesal

groove which receives apex of style and with lateral

row of setae; distal third of plate twice as wide

as proximal two-thirds. Connective linear (Figure

7c) rarely Y-shaped (Figure 2A) , length equal to

or three times as long as aedeagus, divided anteriorly

in two, usually contiguous, arms. Style attached to

anterior third of connective, with well-developed

preapical lobe; apex curved, forming a hook, with

numerous small setae on inner surface. Aedeagus

not fused to connective, with well-developed dorsal

apodemes. Shaft of aedeagus tubular or flattened,

curved or Insinuate, with apex flanged, dentate or

entire in caudodorsal aspect. Gonoduct median or

asymmetrical. Gonopore subterminal, opening into

a spatulate or spoonlike apex.

The linear connective, the articulation between

aedeagus and connective, the male plates with

spinelike setae, the length of the connective (at least

twice as long as aedeagus), and the small male plates

set Psammotettix apart from other Deltocephaline

genera (Oman, 1949). The connective of Psammotettix ferratus (DeLong and Davidson) differs from

other species in the genus in that it is Y-shaped

anteriorly, the arms are not contiguous, and it is

not twice as long as the aedeagus. The connective in

all other Nearctic species is divided anteriorly and

would be Y-shaped if the anterior arms were

separated. The genus is classed with those Deltocephalinae having a linear connective because the contiguous anterior arms give a linear appearance.

The genus appears to be close to the genus

Laevicephahis DeLong. Psammotettix Haupt and

Lacvicephalus are similar in many external characters, differing in characters associated with the external genitalia. The female seventh abdominal

sternum of Laevicephahis has a very pronounced

median projection on the posterior margin. This

margin is usually emarginate in Psammotettix, being blunt and rounded in those species with a

median projection. The sclerite that is appressed

to the female seventh abdominal sternum is weakly

sclerotized or membranous in Psammotettix, heavily

sclerotized in Laevicephalus. This sclerite is possibly

associated with the eighth abdominal sternum. The

basidorsal median lobe does not extend dorsad and

is expanded anteriorly in Laevicephalus. The basidorsal median lobe extends dorsad and is not

expanded anteriorly in Psammotettix. The basidorsal median lobe of Laevicephalus is flanged and

covered dorsally by two flaplike projections that are

not present in Psammotettix. The dorsal aedeagal

apodemes are large in Psammotettix, small in

Laevicephalus. The exposed portion of each male

plate of Psammotettix is small; comprising about

one-third the length of the plate, while that of

Laevicephalus comprises about one-half the total

length of the plate.

The aedeagal shaft of Laevicephalus DeLong is

long, slender, and undulate in lateral aspect, and

with a terminal gonopore. The aedeagus of Psammotettix varies from short to long, from slender

to expanded at apex, and the gonopore is sub-
NL'MBER 74

terminal. The connective is shorter than the

aedeagus in Laevicephalus but longer than the

aedeagus in Psammotettix.

Psammotettix species possess many discordant

characters, making the determination of species relationships difficult. The species can be grouped in

at least five different ways, with no two groups the

same, by grouping them on the basis of different

single characters. Males grouped on the basis of

aedeagal characters fall into different groups from

females grouped on the basis of ovipositor characters. This, per se, is a strong argument against onecharacter taxonomy.

Probably, however, some of the characters should

be weighted. For example, the Y-shaped connective

is held to be a phyletic character in Deltocephalinae; hence, Psammotettix ferratus (DeLong and

Davidson), the only species in Psammotettix with

this character, can be grouped to itself, based on

the weight of this one character, which is probably

secondary in this genus. Psammotettix totalus

(DeLong and Davidson) has an asymmetrical gonoduct, and might, therefore, be grouped alone since

asymmetry is generally accepted as an advanced

character in the Metazoa.

The present arrangement of species in Psammotettix is based on geographic and distributional

data in combination with morphological characters. Each of the species differs from the others in

at least two characters. Usually these are aedegal

characters, comparative wing lengths, characters at

base of the first valvula, and at posterior edge of the

female seventh sternum. Some of the differences

found are clinal.

Psammatettix amplus (DeLong and Davidson),

P. cahuillus (Van Duzee), P. lividellus (Zetterstedt),

and P. alienus (Dahlbom) are species with much

intraspecific variation. Some of these variations are

illustrated (Figures 7, 8, and 9). Further study of

these species, as indicated in the text, would be

profitable.

Psammotettix is divided into the following

Nearctic species groups:

1. The lapponicus group, consisting of P. lividellus (Zetterstedt), P. lapponicus (Ossiannilsson), P.

alexanderi, new species, and P. revae Knull. Psammotettix alienus (Dahlbom), a member of the

Paleartic striatus group of European authors, can

also be grouped with the lapponicus group.

2. The cahuillus group consisting of P. cahuillus

(Van Duzee), a species with much intraspecific variation, and P. attenuens (DeLong and Davidson).

3. The excavatus group consisting of P. excavatus

(Oman), P. shoshone (DeLong and Davidson), and

P. emarginatus, new species.

4. The amplus group consisting of P. ampins

(DeLong and Davidson), a species with much intraspecific variation, and P. asper (Ribaut).

5. A heterogeneous group with each species

actually constituting a group to itself: P. confinis

(Dahlbom), another member of the Palearctic striatus group of European authors, P. ferratus (DeLong

and Davidson), P. totalus (DeLong and Davidson),

P. knullae, new species, and P. dentatus Knull.

Key to Species of Psammotettix

MALES

1. Connective Y-shaped (Figure 2A) . the basal arms not contiguous

ferratus (DeLong and Davidson)

Connective linear (Figure 7c), the basal arms contiguous 2

2(1). Aedeagus with gonoduct symmetrical in caudodorsal aspect 3

Aedeagus with gonoduct asymmetrical in caudodorsal aspect (Figure 3c)

totalus (DeLong and Davidson)

3 (2). Aedeagus with apex of shaft not expanded or not more than one-eighth greater than

narrowest width of shaft in caudodorsal aspect (Figure 6c) 4

Aedeagus with apex of shaft expanded to more than one-fourth greater than narrowest

width of shaft in caudodorsal aspect (Figure 17c) 13

4 (3). Aedeagus with apical two-thirds of shaft straight in lateral aspect (Figure 4D)

knullae, new species

Aedeagus with apical two-thirds of shaft curved in lateral aspect (Figure 10D) 5

5 (4). Aedeagus with apical two-thirds of shaft bisinuate in lateral aspect (Figure 6D) 6

Aedeagus with apical two-thirds of shaft not bisinuate in lateral aspect (Figure 1 ID) . . 7
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

6 (5). Aedeagus with apex abruptly narrowed to one-half greatest width well before tip;

resembling a nipple in caudodorsal aspect (Figure 5E) eahuiUua (Van Duzee)

Aedeagus with apex gradually narrowed to a sharp or blunt point in caudodorsal

aspect (Figure 7c) attenuent (DeLong and Davidson)

7 (5). Aedeagus with apical one-third of shaft gradually tapering to a blunt point in caudorsal aspect 8

Aedeagus with apical one-third of shaft not gradually tapering to a blunt point in

caudodorsal aspect 9

8 (7). Aedeagus with shaft regularly tapering to a point at apex in lateral aspect (Figure 8E)

amplus (DeLong and Davidson)

Aedeagus with shaft not regularly tapering to a point at apex in lateral aspect (Figure 10c) atper (Ribaut)

9(7). Aedeagus with lateral teeth or lobes on apical portion of shaft in caudodorsal aspect

(Figure 16c) 10

Aedeagus without lateral teeth and lobes on apical portion of shaft in caudodorsal

aspect (Figure HE) lividellus (Zetterstedt)

10(9). Aedeagus with apical portion of shaft bearing more than eight lateral dentations per

side in caudodorsal aspect (Figure 13c) dentatu* Knull

Aedeagus with apical portion of shaft bearing less than six lateral dentations per side

in caudodorsal aspect (Figure 15c) 11

11 (10). Aedeagus with lobe of toothed portion at apex in caudodorsal aspect (Figure 14c)

emarginatus, new species

Aedeagus with lobe of toothed portion preapical in caudodorsal aspect (Figure 15c) .. . 12

12(11). Aedeagus with greatest width of Iobcd portion more than five times width of tip in

caudodorsal aspect (Figure 15c) shonhone (DeLong and Davidson)

Aedeagus with greatest width of Iobcd region less than four times width of tip in

caudodorsal aspect (Figure 16c) excavatu* (Oman)

13(3). Aedeagal shaft with apical emargination in caudodorsal aspect (Figure 17c)

alienut (Dahlbom)

Aedeagal shaft without apical emargination in caudodorsal aspect 14

14(13). Aedeagal shaft with apex wide, truncate in caudodorsal aspect (Figure 18c)

confinis (Dahlbom)

Aedeagal shaft with apex rounded, not wide, truncate in caudodorsal aspect 15

15 (14). Aedeagal shaft with widest portion of apex equal to or greater than distance from

gonopore opening to extreme tip in caudodorsal aspect (Figure 19c) . . obetut Knull

Aedeagal shaft with widest portion of apex less than distance from gonopore opening

to extreme tip in caudodorsal aspect (Figure 21D) 16

16(15). Aedeagal shaft with greatest width of apex equal to five-eighths or more distance from

gonopore opening to extreme tip in caudodorsal aspect (Figure 20o) 17

Aedeagal shaft with greatest width of apex equal to one-half or less the distance from

gonopore opening to extreme tip in caudodorsal aspect (Figure 22D) 18

17 (16). Connective stem with posterior end clearly bifid at point of articulation with aedeagus

in caudodorsal aspect beirnei, new species

Connective stem with caudal end not clearly bifid at point of articulation with

aedeagus in caudodorsal aspect alexanderi, new species

18 (16). Hind wings exceeding apex of abdomen revae Knull

Hind wings not reaching apex of abdomen when at rest position

lapponicu* (Ossiannilsson)

FEMALES

I. Hind wings not reaching apex of abdomen when at rest 2

Hind wings exceeding apex of abdomen when at rest 12

2(1). Fore wings not or barely reaching apex of abdomen when at rest 3

Fore wings exceeding apex of abdomen when at rest 7

3 (2). Fore wings not reaching apex of abdomen when at rest alexanderi, new species

Fore wings barely reaching apex of abdomen when at rest 4

4 (3). Seventh abdominal sternum with median projection on posterior margin (Figure 5B)

cahuillu* (Van Duzee)

Seventh abdominal sternum without median projection on posterior margin 5
NUMBER 74

5 (4). First valvula with median basidorsal lobe extending farther cephalad than articulating

lobe (Figure 4A) knullae, new species

First valvula with median basidorsal lobe not extending farther cephalad than articulating lobe 6

6 (5). First valvula with articulating lobe sclerotized or darkened laterally in ventral aspect

obesu* Knull

First valvula with articulating lobe not sclerotized or darkened laterally in ventral

aspect lapponicu* Ossiannilsson

7(2). Seventh abdominal sternum with posterior margin emarginate medially 8

Seventh abdominal sternum with posterior margin not emarginate medially

cahuillut (Van Duzee)

8 (7). Seventh abdominal sternum with depth of median emargination one-fourth or more

median exposed length of sternum 9

Seventh abdominal sternum with depth of median emargination one-eight or less median exposed length of sternum 10

9 (8). Seventh abdominal sternum with depth of median emargination one-fourth to one-half

median exposed length of sternum emarginatua, new species and exeavatu* (Oman)

Seventh abdominal sternum with depth of median emargination two-thirds median

exposed length of sternum thothone (DcLong and Davidson)

10(8). Seventh abdominal sternum produced posteriorly (Figure 21B) 11

Seventh abdominal sternum not produced posteriorly (Figure 22B)

lapponicug (Ossiannilsson)

11 (10) . Seventh abdominal sternum with posterior margin uniformly sinuate; with convexities

and concavities approximately equal (Figure 19B) obeuu Knull

Seventh abdominal sternum with posterior margin irregularly sinuate; with convexities

narrower than concavities (Figure 20B) beirnei, new species

12(1). Ground color two-toned; head, pronotum, and scutellum dark amber; fore wings white

atper (Ribaut)

Ground color uniform IS

13(12). Hind wings extending no farther than midlength of inner apical cell of fore wings

at rest 14

Hind wings extending to distal end of inner apical cell of fore wings at rest 15

14(13). First valvula with articulating lobe sclerotized or darkened laterally (Figure 23A)

revae Knull

First valvula with articulating lobe not sclerotized or darkened laterally

knullae, new species

15(13). Seventh abdominal sternum with posterior margin clearly emarginate (Figure 3B) . 16

Seventh abdominal sternum with posterior margin truncate, weakly emarginate, or

with weak median projection (Figure 7B) 19

16 (15). Seventh abdominal sternum with posterior margin produced posteriorly (Figure 6B)

attenuen» (DcLong and Davidson)

Seventh abdominal sternum with posterior margin not produced posteriorly (Figure

13B) 17

17(16). First valvula with articulating lobe not appearing contiguous to median basidorsal

lobe in ventral aspect (Figure 3A) totalut (DeLong and Davidson)

First valvula with articulating lobe appearing contiguous to median basidorsal lobe in

ventral aspect 18

18(17). Hind wing veins darkened distally, sclerotized part of articulating lobe not broadly

triangular in ventral aspect (Figure 13A) dentatu* Knull

Hind wing veins not darkened distally, sclerotized part of articulating lobe broadly

triangular in ventral aspect (Figure 12A) lividellu* (Zetterstedt)

19(15). First valvula with basidorsal median lobe reduced, appearing concave anteriorly in

ventral aspect 20

First valvula with basidorsal median lobe not reduced, appearing convex anteriorly

in ventral aspect am plus (DeLong and Davidson)

20(19). Basidorsal median lobes not sclerotized, not well developed alienus (Dahlbom)

Basidorsal median lobes sclerotized, deveolped into thin vertical plates

confinu (Dahlbom)
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

B

FIGURE 2.—Male genitalia of Psammotettix ferratus (DcLong and Davidson): A, connective and

style in dorsal aspect; B, aedeagus in caudodorsal aspect; c, same in lateral aspect.

Psammotettix ferratus (DeLong and Davidson)

FIGURE 2

Laevicephalus ferratus DeLong and Davidson, 1935, p. 170,

pi. 7.

Psammotettix ferratus.—Oman, 1947, p. 62.

Length of male 3.16 mm; width of head including

eyes 0.94 mm. Crown 1.26 times longer medially

than next to eye, with median longitudinal suture

extending 0.62 median length from pronotum to

apex. Ground color of head, pronotum, scutellum,

and fore wings creamy; brown markings very faint.

Fore wings with apical cells smoky. Hind wings

transparent; veins not darkened.

Pronotum with round pits confined to area overlying anterior edge of scutum. Scutellum with

sinuate transverse suture. Macropterous form with

fore and hind wings exceeding abdomen when at

rest.

Aedeagal shaft tubular, not expanded at apex in

caudodorsal aspect. Aedeagus with dorsal apodemes

short, less than one-fifth length of shaft. Connective

not much longer than aedeagus, Y-shaped. Style

with arms broadly triangular. Plates fused to valve.

The type series consists of male holotype, allotype, and female paratype, Presque Island, Pennsylvania, 21 July 1920 (DeLong) in the DeLong

collection.

This description is based on the male holotype.

The Y-shaped connective, short with respect to the

length of the aedeagus, and the short dorsal

apodemes set Psammotettix ferratus (DeLong and

Davidson) well apart from other species in Psammotettix. Oman (1949) separates deltocephaline

genera on the basis of the connective being Y-shaped

or linear and sets Psammotettix apart by the connective being twice as long as the aedeagus, and

the arm of the style being long and slender.

Psammotettix ferratus differs from other species of

Psammotettix in all of these respects. Psammotettix

ferratus (DeLong and Davidson) is retained in

Psammotettix became of the small number of available specimens, and because it is similar to other

species of Psammotettix in many respects. Occasionally, speciments of Psammotettix lividellus

(Zetterstedt) have been observed with a Y-shaped

connective. In these specimens it was obvious that

the Y-shaped condition was caused by distortion.

The connective of P. ferratus (DeLong and Davidson) appears to be normal, very flat, and evenly

formed without distortion. No internal parasites

were found. The only unusual condition was that

the aedeagus was pliable and the style appeared in-
NUMBER 74 II

flated. These structures are normally sclerotized

and hard. This pliability would indicate that the

specimen is either teneral or has been softened in

KOH or some other desclerotizing agent. This specimen could be a teneral form. The aedeagus of a

specimen of a species of Psammotettix left in 17

percent KOH remained for 36 hours at room temperature before becoming softened. The styles

softened before the aedeagus. This would indicate

that the softening was not due to a desclerotizing

agent and that the specimen is probably teneral.

Psammotettix totalus (DeLong and Davidson)

FIGURE 3

Laevicephalus totalus DeLong and Davidson, 1935, p. 171.

Psammotettix totalus.—Oman, 1947, p. 62.

Length of male 3.5 to 3.6 mm, of female 3.9 to

4.1 mm; width of head, including eyes, of male 1.0

mm, of female 1.1 mm. Crown with median length

1.3 to 1.4 length next to eye, with median suture

extending anteriorly from 0.66 to 0.69 median

length. Ground color of head, pronotum, scutellum,

A

B

FIGURE 3.—Male and female genitalia and female seventh abdominal sternum of Psammotettix

totalus (DeLong and Davidson): A, first valvulae and first valvifers in ventral aspect; B, seventh

sternum in ventral aspect; c, aedeagus in caudodorsal aspect; D, same in lateral aspect.
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and wings light cream, lightly to heavily marked

with brown. Crown unmarked or with two triangular anterolateral areas, two longitudinal posterolateral stripes, and a spot cephalad of each eye,

brown. Pronotum unmarked or with six longitudinal brown stripes. Fore wing with veins unmarked

or bordered with brown in claval, brachial, central,

and inner anteapical and apical cells. Hind wings

and veins colorless.

Pronotum with round pits randomly distributed

over area above scutum. Scutellum with crescentiform transverse suture. Macropterous form with

fore and hind wings exceeding apex of abdomen,

hind wings reaching distal end of inner apical cell

of front wings when at rest.

Aedeagus with shaft flattened dorsoventrally, with

decurved and serrated lateral flanges; gonoduct

asymmetrical in caudodorsal aspect.

Female seventh abdominal sternum with posterior margin scarcely produced, with a broad median

shallow emargination. First valvula with basidorsal

median lobe well developed and sclerotized, with

articulating lobe arising laterad of median basidorsal lobe at a distance of about one-third width of

latter, in ventral aspect; median basidorsal lobe

extending only slightly farther cephalad than articulating lobe.

The male holotype and the allotype are in the

DeLong collection. The paratypes are in the DeLong and Osborn collections. The type habitat is

not known. The type locality is Murtaugh, Idaho,

and Cane Tree and Cattail Spring, Wyoming. The

distribution is in the Northwestern States (Oman,

1949).

This is one of the larger species of Psammotettix.

It resembles P. confinis (Dahlbom), P. alienus

(Dahlbom), and P. lividellus (Zetterstedt) externally. This species is difficult to relate closely to

any one species in Psammotettix. The asymmetrical

gonoduct and gonopore set it apart. The shape of

the posterior edge of the female seventh abdominal

sternum would place it close to P. obesus Knull

and P. attenuens (DeLong and Davidson). Based on

wing lengths it is in a group with P. confinis

(Dahlbom), dentatus Knull, alienus (Dahlbom), and

lividellus (Zetterstedt). Based on characters of the

first valvulae, it is more closely related to P. amplus

(DeLong and Davidson), lapponicus (Ossiannilsson), and alexanderi, new species.

Psammotettix knullae, new species

FIGURE 4

Psammotettix ferratus.—KmM, 1954, p. 56, figs. 3, 4.

Length of male 3.20 to 3.64 mm, of female 3.13 to

3.58 mm; width of head, including eyes, of male

0.97 to 1.03 mm, of female 0.99 to 1.10 mm; crown

with median longitudinal sulcus extending 0.60 to

0.70 median length from pronotum to apex. Ground

color of head, pronotum, scutellum, and abdomen

creamy. Crown unmarked or marked with two

broad longitudinal stripes laterally. Pronotum unmarked or with six longitudinal brown stripes.

Scutellum unmarked or with two longitudinal

stripes laterally. Fore wings subhyaline, unmarked,

or marked with brown along commisural border;

hind wings transparent and unmarked.

Pronotum with round pits as in P. totalus (DeLong and Davidson). Scutellum witli crescentiform

transverse suture. Macropterous form with fore

wings barely reaching or exceeding abdominal

apex when at rest, hind wings shorter than abdomen, reaching proximal end of inner apical cell of

fore wing when at rest.

Aedeagus with shaft tubular throughout length,

apex with internal sclerite resembling a "y" in

caudodorsal aspect, shaft with apical two-thirds

straight in lateral aspect. Connective with stem

greatly widened caudally, with small notch at point

of articulation with aedeagus in ventral aspect.

Female seventh abdominal sternum with notchlike median emargination. Intersegmental membrane lightly sclerotized, with wrinkles visible

through cleared seventh sternum (Figure 42). First

valvula (Figure 4A) with median basidorsal lobe

well developed, extending farther cephalad than

articulating lobe which is not well defined medially.

Holotype male, allotype, two male and three

female paratypes. Toppenish, Washington, 8 July,

1935 (Beamer); paratypes from Rocky Ford, Colorado; Hot Lake, Oregon; Snowville, Utah; Gardner,

Montana; Burley and Twin Falls, Idaho; Saskatoon

and Elbow, Saskatchewan; McMurrary, Foremost,

and Manyberries, Alberta. Holotype in the National

Museum of Natural History.

This species is named in honor of D. J. Knull.

It was misidentified and illustrated as Psammotettix

ferratus (DeLong and Davidson) by Knull. The
NUMBER 74 13

f\

B

FIGURE 4.—Male and female genitalia and female seventh abdominal sternum of Psammotettix

knullae, new species: A, first valvulae and first valvifer in ventral aspect; B, seventh sternum in

ventral aspect; c, aedeagus in caudodorsal aspect; D, same in lateral aspect.

type of P. ferratus, described elsewhere in this

paper, is entirely different in aedeagal characters

from those illustrated by Knull. This species stands

apart from other Nearctic Psammotettix species in

that the apical two-thirds of aedeagal shaft is

straight in lateral aspect. The relative wing lengths

are close to P. revae Knull. The internal sclerotization of the aedeagal apex is close to an illustration

by Remane (1965, fig. 2c) which is either Psammotettix koeleriae Zachvatkin or Psammotettix

angulatus.

A specimen from Rocky Ford, Colorado, was

collected from sugar beet.

Psammotettix cahuillus (Van Duzee)

FIGURE 5

Deltocephalus cahuilla Van Duzee, 1925, p. 417.

Latalus cahuilla.—DeLong and Caldwell, 1937.

Laevicephalus cahuillus.—DeLong and Knull, 1946, p. 28.

Psammotettix cahuillus.-Oman, 1949, p. 162.

Length of male 3.00 mm, of female 2.70 to 3.25

mm; width of head, including eyes, of male 0.95 to

1.00 mm, of female 0.93 to 1.30 mm. Crown with

median length 1.13 to 1.29 length next to eye, with

median sulcus extending from 0.56 to 0.86 median

length. Ground color of head, pronotum, fore

wings, and scutellum varying from creamy to cinereous, and from unmarked to heavily marked with

brown. Crown with four spots on anterior margin,
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

D

FIGURE 5.—Male and female genitulia and female seventh abdominal sternum of Psammotettix

cahuillus (Van Duzec) : A, first valvulae and first first vulvifer in ventral aspect; B, seventh sternum

in ventral aspect; c, acdeagus in caudodor>al aspect. Riverside, California; D, same in lateral

aspect: E, aedcagus in caudodorsal aspect, Keen Camp. California: F, same in lateral aspect.

two mediolateral triangular spots, and four stripes

along caudal edge, brown. Pronotum with six longitudinal brown stripes with superimposed brown

splotches. Fore wing veins bordered with brown;

with eight spots, two anteriorly, four in transverse

row at midlength, and two in inner anteapical cells,

brown. Scutellum with two longitudinal brown

stripes.

Pronotum with round pits absent or confined to

area just above anterior edge of scutum. Scutellum
NUMBER 74 15

with transverse suture crescentiform. Macropterous

form with fore wings reaching or exceeding apex of

abdomen, with hind wings shorter than abdomen,

reaching midlength of inner anteapical cell of fore

wings when at rest.

Aedeagus with shaft tubular, apical two-thirds

bisinuate in lateral aspect, with apical portion

abruptly narrowed to one-half greatest width well

before tip, resembling a nipple in caudodorsal

aspect; without lateral dentations on shaft; connective weakly concave in ventral aspect at articulation with aedeagus.

Female seventh abdominal sternum with relatively strong median projection on posterior margin. First valvula with basidorsal median lobe well

developed and sclerotized; articulating lobe arising

one-half its width laterad of basidorsal median lobe

in ventral aspect; median basidorsal lobe and articulating lobe extending about equally cephalad.

The type locality is Keen Camp, California. The

type habitat is grass. The distribution is California

(Van Duzee, 1925).

Psammotettix cahuillus (Van Duzee) shows a wide

color variation. The bisinuate shaft of the aedeagus

would place it close to P. attenuens (DeLong and

Davidson), but other characters such as head shape

and wing length indicate a more distant relationship. The median projection of the posterior edge

of the female seventh abdominal sternum is similar

to P. alienus (Dahlbom). The wing lengths of P.

B

FIGURE 6.—Male and female genitalia and female seventh abdominal sternum of Psammotettix

attenuens (DeLong and Davidson) : A, first valvulac and first valvifers in ventral aspect;

B, seventh sternum in ventral aspect; c, aedeagus in caudodorsal aspect: D. same in lateral aspect.
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

B

FIGURE 7.—Male and female genitalia and female seventh abdominal sternum of Psammotettix

ampins (DeLong and Davidson): A, first valvulac and first valvifer in ventral aspect; B, seventh

sternum in ventral aspect; c, connective and style in dorsal aspect; D, aedeagus in lateral aspect,

Monterey, California; E, same in caudodorsal aspect.

cahuillus are close to those of P. alexanderi, new

species, P. lapponicus (Ossiannilsson), P. excavatus

(Oman), P. ampins (DeLong and Davidson), P.

obesus Knull, and P. shoshone (DeLong and Davidson). In the characters at the base of the first valvula

cahuillus is close to amplus.

Psammotettix cahuillus (Van Duzee) shows a

greater range of color variation than do most of the

Psammotettix species. There is also variation in the

aedeagus (Figures 5 c, D, E, and F). More information is necessary to verify the infraspecific taxonomic status of these forms.

Psammotettix attenuens (DeLong and Davidson)

FIGURE 6

Laevicephalus attenuens DeLong and Davidson, 1935, p. 170.

Psammotettix attenuens.—Oman, 1947, p. 62.

Length of male 3.50 mm, of female 3.46 to 3.60

mm; width of head, including eyes, of male 0.96

to 1.00 mm, of female 0.96 to 1.00 mm; crown 1.10

to 1.33 times longer medially than next to eye, with

median longitudinal suture extending 0.64 to 0.98

median length from pronotum to apex. Pronotum

1.21 to 1.42 times longer medially than length of

crown. Ground color of head, pronotum, scutellum,

and wings creamy, lightly to heavily marked with

brown. Dorsal horizontal brown stripe of face visible from above, causing crown to appear more

pointed than it actually is. Crown with two anterolateral triangles, mediolateral blotches, and four

spots along posterior edge, brown. Pronotum with

or without six longitudinal stripes, brown; without

numerous circular pits. Scutellum with or without

four brown spots or two inner brown and two outer
NUMBER 74 17

orange spots, under posterior edge of pronotum;

with transverse suture straight to crescentiform.

Macropterous form with front and hind wings

longer than abdomen; similar to Psammotettix

totalus (DeLong and Davidson).

Shaft of aedeagus bisinuate in lateral aspect;

gradually tapering to a blunt point in caudodorsal

aspect; without lateral wings or projections.

Seventh sternum of female with posterior edge

strongly produced caudad and emarginate. First

valvula with basidorsal median lobe well developed; with articulating lobe poorly developed, represented by a basal fold of the first valvula

(Figure 6A). Articulating lobe or fold appearing

contiguous to median basidorsal lobe in ventral

aspect. Articulating lobe and median basidorsal

lobe extending about equally cephalad.

The type series is from Yellowstone Park, Wyoming, 4 July 1930 (D. M. DeLong). The type habitat

is not known.

In aedeagal characters, Psammotettix attenuens

is close to Psammotettix cahuillus (Van Duzee). In

the blunt apex of the crown, attenuens is not close

to other species of Psammotettix.

A female specimen of Psammotettix attenuens

(DeLong and Davidson) from Kalispell, Montana,

was stylopized between the fifth and sixth abdominal segments.

B

FIGURE 8.—Aedeagus of Psammotettix ampins (DeLong and Davidson) : A, aedeagus in caudodorsal aspect, Cuyama Ranch, California; B, same in lateral aspect; c, aedeagus in caudodorsal

aspect, Tehachapi, California; D, same in lateral aspect; E, aedeagus in caudodorsal aspect,

Monlith, California; F, same in lateral aspect.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

B

FIGURE 9.—Aedeagus of Psammotettix amplus (DeLong and Davidson): A, aedeagus in caudodorsal aspect, San Margarita, California; B, same in lateral aspect; c, aedeagus in caudodorsal

aspect, Visalia, California; D, same in lateral aspect; E, aedeagus in caudodorsal aspect, Fort

Lewis, Washington; F, same in lateral aspect.

Psammotettix amplus (DeLong and Davidson)

FIGURES 7. 8, 9

Laevicephalus amplus DeLong and Davidson, 1935, p. 171.

Psammotettix amplus.—Oman, 1947, p. 62.

Length of male 2.90 mm to 3.97 mm, of female

2.97 mm to 3.80 mm; width of head, including eyes,

of male 0.90 mm to 1.20 mm, of female 0.90 mm

to 1.00 mm. Color varying as in Psammotettix

totalus (DeLong and Davidson).

Pronotum with round pits on area overlying

anterior edge of scutum. Scutellum with straight to

weakly crescentiform transverse suture. Macropterous form with fore and hind wings longer than

abdomen when at rest; hind wings at rest reaching

distal end of inner apical cell of fore wings.

Aedeagus with shaft changing from tubular at

gonopore to flattened dorsoventrally at apex; with

apex of shaft varying in attitude from curved ventrally and anteriorly to curved dorsally in lateral

aspect; lateral edge of apical one-half of shaft varying from entire to dentate with up to five teeth on

each side in caudodorsal aspect, shaft without lateral flanges, with apical one-third gradually taper-
NUMBER 74 19

ing to a blunt point in caudodorsal aspect.

Female seventh abdominal sternum with posterior margin slightly produced, weakly emarginate,

or almost truncate. First valvula with basidorsal

median lobe well developed and sclerotized, almost

reaching base of valvula which with its complement

is convex in ventral aspect; with articulating lobe

arising laterad of median basidorsal lobe at a distance of less than one-third width of latter in ventral aspect and extending slightly farther cephalad

than basidorsal median lobe.

The male holotype and the allotype are in the

Osborn collection. There are no paratypes. The

type locality is Bakersfield, California. The distribution is California (Oman, 1949) and Fort Lewis,

Washington.

P. ampins (DeLong and Davidson) is one of the

more variable species of Psammotettix. There is a

morphological continuum from an aedeagal apex

that is turned dorsally to one that is turned ventrally and anteriorly in the lateral aspect. The

aedeagal shaft varies from un toothed to toothed

laterally in caudodorsal aspect. Both characters vary

geographically according to available data. The

ventral and anteriorly directed apex occurs in

higher altitudes; the coiled dorsally directed apex

occurs in coastal areas. The untoothed aedeagal

shaft occurs in the south and teeth increase in number to the north. In addition to the clinal aedeagal

variation in this species, there is much individual

variation in size and coloration. Our knowledge of

P. ampins would benefit from an intensive geographic and host-plant study.

Psammotettix ampins (DeLong and Davidson)

appears to be close to Psammotettix asper (Ribaut).

The tip of the aedeagal shaft of asper is developed

more than that of amplus, and it has a slight lateral

flange in caudodorsal aspect.

Psammotettix asper (Ribaut)

FIGURE 10

Deltocephalus asper Ribaut, 1925, p. 16.

Psammotettix asper.—Ribaut, 1938, p. 168.

Length of male 2.90 to 3.05 mm, of female 2.90 to

3.00 mm; width of head, including eyes, of male

0.88 to 0.92 mm. Crown with median sulcus extending anteriorly about 0.7 median length. Ground

color of head, pronotum, and scutellum creamy to

amber, marked with brown or orange. Ground color

of wings ivory; claval, brachial, inner discal, inner

anteapical, and apical cells marked with brown.

Brown markings on crown as in Psammotettix

totalus (DeLong and Davidson), occasionally with

median longitudinal stripe orange. Pronotum unmarked or with two orange longitudinal stripes.

Pronotum with round pits as in Psammotettix

totalus (DeLong and Davidson). Scutellum with

transverse suture straight or undulate. Macropterous form with wing lengths as in totalus.

Aedeagus tubular, with widest point of shaft in

caudodorsal aspect about midway between gonopore opening and apex; aedeagal shaft not widened

at apex, with series of serrations on cephaloventral

surface between dorsal apodemes and gonopore

opening; shaft with weak lateral flanges.

Female seventh abdominal sternum with posterior margin not produced, sinuate, with convexities

and concavities approximately equal; median emargination not as broad as and deeper than that of

Psammotettix totalus (DeLong and Davidson). First

valvula with basidorsal median lobe well developed

and sclerotized, with articulating lobe extending

farther cephalad than basidorsal median lobe, with

articulating lobe appearing to arise contiguous to

the median basidorsal lobe in ventral aspect.

No holotype has been designated. The type

locality is Sarkeresztur, Hungary. This description

is based on specimens obtained from W. Wagner.

The Nearctic specimens of Psammotettix asper

(Ribaut) have not been examined. The Nearctic occurrence of asper is based on a Manitoba record

(McClure, 1943). The distribution is Hungary

(Ribaut, 1925), Bohemia, Moravia (Dlabola, 1945),

Slovakia (Dlabola, 1954), and Manitoba.

Psammotettix asper (Ribaut) appears to be close

to Psammotettix amplus (DeLong and Davidson).

This is the only Nearctic species of Psammotettix

with a two-toned ground color. Psammotettix asper

also stands apart from other Nearctic species by the

row of serrations dorsad on the aedeagal shaft.

Psammotettix lividellus (Zetterstedt)

FIGURES 11, 12

Cicada lividella Zetterstedt, 1840, p. 290.

lassus lividellus.-W'alkcr, 1851, p. 882.

Deltocephalus lividellus.—Sa hi berg, 1871, p. 37.

Deltocephalus affinis Gillette and Baker, 1895, p. 84.
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Deltocephalus excisus Matsumura, 1906, p. 79.

Deltocephalus exisus (sic) Matsumura, 1908, p. 2.

Deltocephalus incisus (sic) Matsumura, 1908, p. 2.

Deltocephalus striatus.—Sanders and DeLong, 1920, p. 8.

Deltocephalus striatus lividellus.—Blole, 1927, p. 220.

Laevicephalus striatus.—DeLong and Davidson not (Linnaeus,)

1935, p. 168.

Psammotettix lividellus.—Ribzut, 1938, p. 166.

Psammotettix striatus.—DeLong (not Linnaeus), 1948, p. 258.

Length of male 3.15 to 3.70 mm, of female 3.13 to

3.80 mm, width of head, including eyes, of male

0.86 to 1.01 mm, of female 0.95 to 1.05 mm. Crown

with median sulcus varying in length. Ground color

of head, pronotum, scutellum, and wings light

cream to yellowish amber, very lightly marked with

brown. Crown unmarked or with paired anterolateral brown lines forming crescentiform arcs from

apex to ocelli. Pronotum unmarked. Fore wings unmarked or with apical cells bordered with brown.

Hind wings transparent, veins not darkened. Pronotum with pits as in P. totalus (DeLong and

Davidson). Scutellum with transverse suture almost

straight, with weak median undulate impression.

Macropterous form with fore and hind wings exceeding apex of abdomen; hind wings extending to

midlength or distal end of inner apical cell of fore

wings at rest.

Aedeagus tubular with shaft of nearly uniform

diameter throughout length, not appreciably narrowed or expanded in any aspect; with gonoduct

symmetrical in caudodorsal aspect; varying in size;

lateral edge of shaft entire or with minute dentations; slightly narrowed or expanded distad of

gonopore in caudodorsal aspect (Figures 11A, C, E).

Female seventh abdominal sternum with median

emarginations as in P. totalus (DeLong and Davidson); lateral emarginations shallower than in

B

FIGURE 10.—Male and female genitalia and female seventh abdominal sternum of Psammotettix

asper (Ribaut): A, first valvulae and first valvifers in ventral aspect; B, seventh sternum in

ventral aspect; c, aedeagus in caudodorsal aspect; D, same in lateral aspect.
NUMBER 74

FIGURE 11.—Aedeagus of Psammoteltix lividcllus (Zcttcrstcdt): A, acdeagus in caudodorsal aspect,

Indian House Lake, Quebec; B, same in lateral aspect; c, acdeagus in caudodorsal aspect, Soda

Creek, British Columbia; D, same in lateral aspect; E, aedeagus in caudodorsal aspect, Hebron,

Labrador; F, same in lateral aspect.

totalus. First valvulae with articulating lobe arising laterad of median basidorsal lobe, the two appearing contiguous in ventral aspect. Median basidorsal lobe and articulating lobe extending about

the same distance cephalad.

Described from Greenland on the basis of external characters. The labeled types are in Zetterstedt's collection. Their abdomens are missing

(Beirne, 1954). No holotype was designated. Ossiannilsson (1938) designated a neotype (invalid

because original types are not lost) and illustrated

the aedeagus.

Psammotettix lividellus (Zetterstedt) shows variation in size, internal sclerotization, and lateral ornamentation of the aedeagal shaft. It has not been

determined whether these variations are individual

or geographic or a combination of the two. This

species is most closely related to a group of Psammotettix species referred to herein as the lapponicus

group. Nearctic species in this group are lapponicus, alexanderi, new species, revae Knull, beirnei,

new species, and obesus Knull. This group is characterized by a lateral expansion of the aedeagal

shaft apex, giving it an oval appearance in the
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 12.—Male and female genitalia and female seventh abdominal sternum of Psammotcttix

lividellus (Zetterstedt): A, first valvulae and first valvifer in ventral aspect; B, seventh sternum

in ventral aspect; c, aedeagus in caudodorsal aspect, Lusk, Wyoming; D, same in lateral aspect.

caudodorsal aspect. Some specimens of lividellus

show a tendency toward a similar expansion. A

morphological gradation for this character can be

demonstrated through the increasingly oval shaft

apex of lapponicus (Ossiannilsson), revae Knull,

alexanderi, new species, to the almost round apex

of obesus Knull. This gradation could not be clinal

as the morphological changes do not parallel the

geographic distribution of these species. The apparent geographic variation in P. amplus yields

intraspecific aedeagal character differences as great

as the interspecific character differences in the

lapponicus group. Supporting characters such as

relative length of wings in rest position, shape of

posterior edge of female seventh abdominal sternum, and characters at the base of the first valvulae

confirm the specific status of the taxa in the

lapponicus group.

Psammotettix lividellus (Zetterstedt) is closely related to P. revae, based on characters of the first

valvulae. This species has been reported as P.

striatus (Linnaeus) by earlier North American

workers. This is not the striatus of European

workers. A lectotype for P. striatus (Linnaeus) has

not yet been designated. The erroneous reporting

of lividellus as striatus is confusing. It will not be

possible to resolve the problem until a lectotype or

neotype for striatus (Linnaeus) has been designated.

Ribaut (1938) redescribed striatus (Linnaeus) but

did not designate a type. If Ribaut's redescription

of striatus (Linnaeus) is accepted, lividellus (Zetterstedt) is a separate and distinct species, and records
NUMBER 74 23

of Nearctic striatus are actually records of lividellus

(Zetterstedt). The writer has not found a Nearctic

specimen of P. striatus (Linnaeus) sensu Ribaut. Dr.

W. J. Knight has examined the male type of

"striata" in the Linnean Society collection and finds

it to be Aphrodes flavostriatus (Donovan, 1799). He

is not fully convinced that he has the true Cicada

striata Linnaeus since it is difficult to imagine that

past authors, even without dissection, could have

considered his specimen even remotely similar to

a species of Psammotettix.

Psammotettix dentatus Knull

FIGURE 13

Psammotettix dentatus Knull, 1954, p. 55.

Length of male 3.4 to 3.6 mm, of female 3.5 to 3.8

mm; width of head, including eyes, of male 0.9 mm

to 1.0 mm; of female 1.0 mm; crown with median

length 1.2 to 1.4 length next to eye, with median

longitudinal sulcus extending anteriorly 0.6 to 0.8

median length. Ground color of head, pronotum,

scutellum, and wings cream, lightly to heavily

marked with brown. Crown and pronotum with

color as in P. totalus (DeLong and Davidson).

FIGURE 13.—Male and female genitalia and female seventh abdominal sternum of Psammotettix

dentatus Knull: A, first valvulae and first valvifer in ventral aspect; B, seventh sternum in ventral

aspect; c, aedeagus in caudodorsal aspect; D, same in lateral aspect.
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 14.—Male and female genitalia and female seventh abdominal sternum of Psammotettix

emarginatus, new species: A, first valvulae and first valvifer in ventral aspect; B, seventh sternum

in ventral aspect; c, aedeagus in caudodorsal aspect; D, aedcagus in lateral aspect.

Scutellum unmarked or with four spots behind

pronotum, and with two spots behind transverse

suture, brown. Fore wing veins unmarked to bordered with brown in all cells. Hind wing veins

darkened distally or colorless.

Crown with area cephalad of ocelli triangular in

shape in dorsal aspect, its apex sharp. Pronotum

with round pits as in P. totalus (DeLong and Davidson). Scutellum with transverse suture crescentiform

to straight. Macropterous form with wing length

ratios as in totalus.

Aedeagus with shaft long, slender, changing from

round in cross section at gonopore to flattened

dorsoventrally at apex with decurved and serrated

lateral flanges in caudodorsal aspect, with more

than twelve dentations each side.

Female seventh abdominal sternum with posterior

margin not produced, with median emargination as

in P. lividellus (Zetterstedt), lateral emarginations

weak (Figure 13B); intersegmental membrane sclerotized with wrinkles visible through cleared female

seventh sternum. First valvula with basidorsal median lobe well developed and scletorized; with articulating lobe arising contiguous to the basidorsal

median lobe in ventral aspect. Basidorsal median

lobe extending farther cephalad than articulating

lobe.

The male holotype, allotype, and paratypes are

in the D. J. and J. W. Knull collection. The type

locality and locality of allotype and paratypes is

Quincy, California. The type habitat is not known.

The distribution is northern California (Knull,

1954), Siskiyou National Forest, California, North

Farmington, Utah, and British Columbia.

Psammotettix dentatus Knull appears to stand

well apart from other species of Psammotettix. The

shape of the crown in dorsal aspect is close to P.

ampins (De Long and Davidson), P. cahuillus (Van
NUMBER 74 25

Duzee), P. lapponicus (Ossiannilsson), and P. alexanderi, new species. The long slender aedeagal shaft

sets it apart from the other Nearctic species of

Psammotettix. Psammotettix lividellus (Zetterstedt),

P. ferratus (DeLong and Davidson), P. kniillae, newspecies, P. attenuens (DeLong and Davidson), and

P. cahuillus (Van Duzee) have slender aedeagal

shafts, but they are not as long in comparison as

that of dcntatus. Psammotettix alboniger (Lethierry)

(Linnavuori, 1953), a Palearctic species, has a long

slender undulating aedeagal shaft which is compartively longer than that of dentatus. The form

of the aedeagal shaft of alboniger resembles the

aedeagal shaft of the species of the genus Laevicephalus. Psammotettix dentattis is of the typical

striata color. The relative positions of the median

basidorsal lobe and the articulating lobe place this

species close to lividellus, revae, asper, and amplus.

Psammotettix dentatus is set apart from most other

Nearctic species of Psammotettix by the darkening

of some of the veins of the hind wing.

A specimen of this species was collected from

celery at North Farmington, Utah.

Psammotettix emarginatus, new species

FK.LRF 14

Length of male 3.00 to 3.40 mm, of female 2.95 to

3.40 mm, width of head, including eyes, of male

0.94 to 0.98 mm, of female 0.92 to 1.00 mm; crown

with median sulcus extending anteriorly 0.61 to

0.79 median length. Color varying as in P. totalus

(DeLong and Davidson).

Pronotum with round pits confined to area overlying anterior edge of scutum. Scutellum with transverse suture crescentiform to straight. Macropterous

form with fore wings at rest exceeding abdominal

apex, hind wings at rest shorter than abdomen,

reaching distal end of inner apical cell of fore wings.

Aedeagus with shaft tubular at gonopore, dorsoventrally flattened distally, with less than six lateral

dentations and with large lateral projection on

each side at apex in caudodorsal aspect. Connective

divided at point of articulation with aedeagus;

stem one-half length of arms.

Female seventh abdominal sternum with posterior margin strongly emarginate, with median emargination extending one-third the distance from posterior margin to base (Figure 14B). Intersegmental

membrane lightly sclerotized, with wrinkles visible

through cleared seventh sternum (Figure 14B). First

valvula with median basidorsal lobe well developed,

anteriorly resembling a hook that overlaps articulating lobe at base in ventral aspect (Figure 14A).

Holotype male, allotype, and paratypes, Klamath,

Oregon, 1 July 1935 (Oman) in the National Museum of Natural History.

Psammotettix emarginatus, new species, is closely

related to P. excavatus (Oman) and P. shoshone

(DeLong and Davidson). The chief distinguishing

character is the degree of emargination of the posterior margin of the female seventh abdominal

sternum. The emargination of emarginatus extends

about one-third, of excavatus about one-half, and

of shoshone about two-thirds the distance from the

hindmost point to the hindmost point of the sixth

sternum. In the aedeagus, excavatus and shoshone

are very close, but the difference in the posterior

margin of the female seventh abdominal sternum

is distinct. In the aedeagus, excavatus and emarginatus are clearly different, but there seems to be

some overlap in characters at the posterior margin

of the seventh abdominal sternum. Further study

of this group is desirable.

Psammotettix shoshone (DeLong and Davidson)

FIGURE 15

Liievicephalus shoshone DeLong and Davidson, 1934, p. 222.

Psammotettix shoshone.—Oman, 1947, p. 62.

Length of male 3.4 mm; width of head, including

eyes, of male 0.92 to 0.96 mm, of female 1.0 mm;

crown with median length 1.3 to 1.5 length next

to eye, with median sulcus extending anteriorly

0.7 to 0.8 median length from pronotum to apex.

Color varying as in P. totalus (DeLong and Davidson).

Pronotum with round pits as in Psammotettix

totalus (DeLong and Davidson). Scutellum with

transverse suture as in totalus. Macropterous form

with fore wings barely exceeding apex of abdomen,

hind wings at rest shorter than abdomen.

Aedeagus with shaft changing from tubular at

gonopore to dorsovcntrally flattened at apex; apex

of shaft with lateral flanges bearing four dentations

on each side, with greatest width of toothed region

more than five times width of tip in caudodorsal

aspect. Connective as in Psammotettix emarginatus,

new species.
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FICURE 15.—Male and female genitalia and female seventh abdominal sternum of Psammotettix

shoshone (DeLong and Davidson): A, first valvulae and first valvifer in ventral aspect; B,

seventh sternum in ventral aspect; c, aedeagus in caudodorsal aspect; D, same in lateral aspect.

Female seventh abdominal sternum with posterior margin strongly emarginate; with median

emargination extending one-half the distance from

posterior margin to base; extending two-thirds the

distance from posterior margin to posterior margin

of sixth abdominal sternum. Intersegmental membrane with wrinkles visible through cleared seventh

sternum. First valvula with median basidorsal lobe

well developed, with anterior hook ending at base

of articulating lobe in ventral aspect (Figure 15A).

The female holotype and twelve female paratypes

are in the DeLong collection. The type locality is

Shoshone Basin, Idaho. The type habitat is grasses.

The distribution is Shoshone Basin, Idaho, and

Contact, Nevada.

Psammotettix shoshone (DeLong and Davidson)

is close to P. excavatus (Oman). Psammotettix shoshone differs from excavatus in the relative width

of the aedeagal apex in caudodorsal aspect; the

greatest width of the toothed region being more

than five times the width of the apex in shoshone,

less than four times the width of the apex in excavatus. Psammotettix shoshone differs from excavatus in the relative distance the median emargination of the female seventh abdominal sternum

extends from the posterior edge of the seventh

abdominal sternum to the posterior edge of the

sixth abdominal sternum, extending about twothirds the distance in shoshone, about one-half the

distance in excavatus.
NUMBER 74 27

Psammotettix excavatus (Oman), P. shoshone

(DeLong and Davidson), and P. emarginatus, new

species, are in a closely related group which is hereafter referred to as the excavatus group. The bifid

stem of the connective in the excavatus group is

present in some species of the lapponicus group.

Psammotettix excavatus (Oman)

FIGURE 16

Laevicephalus excavatus Oman, 1931, p. 430.

Psammotettix excavatus.—Oman, 1947, p. 62.

Length of female 3.05 to 3.20 mm; width of head

including eyes 0.97 to 0.99 mm. Crown with median

sulcus extending anteriorly 0.73 to 0.80 median

length. Ground color of head, pronotum, scutellum,

and fore wings light cream, unmarked or lightly

marked with brown as in P. totalus (DeLong and

Davidson). Hind wings transparent, veins colorless.

Pronotum with round pits confined to area above

anterior edge of scutum. Scutellum with transverse

suture nearly straight. Macropterous form with fore

wings exceeding apex of abdomen, with length of

hind wings at rest equal to or shorter than abdoFICURE 16.—Male and female genitalia and female seventh abdominal sternum of Psammotettix

excavatus (Oman): A, first valvulae and first valvifer in ventral aspect; B, seventh sternum in

ventral aspect; c, aedeagus in caudodorsal aspect; D, same in lateral aspect.
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

men. Hind wings at rest extending only to midlength or distal part of inner anteapical cell of fore

wings.

Aedeagus with shaft flattened distally and toothed

as in Psammotettix emarginatus, new species, with

largest lateral lobe anteapical in caudodorsal aspect;

with greatest width of toothed region less than four

times width of apex in caudodorsal aspect (Figure

16c). Connective as in emarginatus.

Female seventh abdominal sternum strongly

emarginate, with median emargination extending

one-half distance from posterior margin to base

(Figure 16B). Intersegmental membrane as in P.

emarginatus, new species. First valvula with median

basidorsal lobe well developed, anteriorly resembling a hook that terminates at base of articulating

lobe in ventral aspect.

The type locality is Gazelle, California. Other

specimens in type series from the type locality,

Siskiyou, Oregon, and Lake Tahoe, California. The

holotype is catalog number 43584 USNM. The type

habitat is not known. The distribution is in the

northwestern states (Oman, 1949).

This species is closely related to P. emarginatus,

new species, and P. shoshone (DeLong and Davidson).

Psammotettix alienus (Dahlbom)

FIGURE 17

Thamnolettix aliena Dahlbom, 1850, p. 187.

Athysanus alienus— Kirschbaum, 1858, p. 3.

Deltocephalus breviceps Sahlberg, 1871, p. 342.

Thamnolettix alinea.— (sic) Melichor, 18%.

Laexicephalus latipex Beirne, 1954, p. 496.

Deltocephalus striatus.—\zx. breviceps Ossiannilsson, 1937,

p. 133.

Psammotettix allienus.— (sic) Farsky, 1952, p. 75

Length of male 3.8 to 4.2 mm, of female 3.9 to 4.1

mm; width of head, including eyes, of both sexes

1.0 to 1.1 mm. Crown with median length 1.1 to

1.5 length next to eye, with median sulcus extending anteriorly from 0.7 to 0.8 median length. Color

varying as in P. totalus (DeLong and Davidson).

Scutellum with or without one median longitudinal

brown stripe and two lateral orange stripes. Hind

wing veins mostly colorless; some specimens with

apical veins partially darkened.

Pronotum with median length 1.3 to 1.6 median

length of crown, with round pits as in P. totalus.

Aedeagus with shaft tubular, expanding into a

keeled flanged spoonlike apex distad of the gonopore, with apex of shaft distinctly emarginate in

caudodorsal aspect, without lateral teeth or dentations (Figures 17c and F).

Female seventh abdominal sternum with posterior margin slightly produced or almost truncate

(Figure 17B). Intersegmental membrane visible as

wrinkles through cleared seventh sternum. First

valvula (Figure 17A) with basidorsal median lobe

absent or weakly developed; base of valvula with

its complement concave in ventral aspect. Articulating lobe much farther cephalad than basidorsal

median lobe if present.

The type locality is Gotland Island. The original

description was based on external characters. The

aedeagus of the holotype was illustrated by Ossiannilsson (1947). The distribution is Holarctic.

Psammotettix alienus (Dahlbom) is close to P.

stria tits (Linnaeus) sensu Ribaut. This relationship

has been discussed in the introduction. This species

appears to be the same as P. provincialis Ribaut,

based on the shape of the aedeagus. Ribaut's (1938)

illustration of the aedeagus of alienus differs from

Ossiannilsson's (1947) illustration of the type of alienus. Figures 17c and F show some of the variation

in the aedeagal base.

In the characters at the base of the first valvula

this species is close to P. confinis (Dahlbom). The

shape of the posterior edge of the female seventh

abdominal sternum of P. alienus (Dahlbom) is close

to P. cahuillus (Van Duzee).

Von Archimowitsch (1952) reports P. alienus

(Dahlbom) as a vector of a virus disease of potato

in Spain.

A male from Fort Yukon, Alaska, is stylopized

between the second and third abdominal segments.

There are no apparent morphological changes in

the genitalia as a result of this parasitism.

A female was collected from corn at Caldwell,

Idaho.

Psammotettix confinis (Dahlbom)

FIGURE 18

Deltocephalus confinis Dahlbom, 1850, p. 193.

fassus striatus.—Kirschbaum (not Linnaeus), 1868, p. 132.

Thomson, 1869, p. 70.

Deltocephalus heydeni Fieber, 1872, p. 3.

Deltocephalus thenii Edwards, 1915, p. 208.

Deltocephalus spat Infer Ribaut, 1925, p. 19.

Deltocephalus striatus var. thenii Blote, 1927, p. 57.

Ribautiellus thenii.-China, 1938, p. 195.
NUMBER 74

Psammotettix thenii.—Ribaut, 1938, p. 166.

Psammotettix spalhifer Ribaut, 1938, p. 166.

Psammoteltix confmis.—Ribaut, 1938, p. 166.

Psammotettix striattts.—Wagner (not Linnaeus), 1939, p. 162.

29

Ribautiellus confinis.—Roche, 1944, p. 84.

Deltocephalus harrimani Bcirne, 1954, p. 495.

Length of male 3.2 to 3.8 mm, of female 3.5 to 3.6

B

FIGURE 17.—Male and female genitalia and female seventh abdominal sternum of Psammotettix

alienus (Dahlbom): A, first valvulae and first valvifer in ventral aspect; B, seventh sternum in

ventral aspect; c, aedeagus in caudodorsal aspect, Pingre Park, Colorado; D, same in lateral

aspect; E, aedeagus in lateral aspect, Sigurd, Utah; F, same in caudodorsal aspect.
so

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

B

FIGURE 18.—Male and female genitalia and female seventh abdominal sternum of Psammotettix

confinis (Dahlbom): A, first valvulae and first valvifers in ventral aspect; B, seventh sternum in

ventral aspect; c, aedeagus in caudodorsal aspect; D, same in lateral aspect.

mm; width of head, including eyes, of male 9.9 to

1.0 mm, of female 0.96 to 1.0 mm. Crown with

median length 1.2 to 1.4 length next to eye; with

median sulcus extending anteriorly 0.7 to 0.9 median length. Color varying as in P. alienus (Dahlbom). Hind wings transparent, veins colorless.

Pronotum with median length 1.14 to 1.41 times

length of crown; with round pits as in P. totalus

(DeLong and Davidson). Scutellum with crescentiform transverse suture. Macropterous form with

wings as in totalus.

Aedeagus with shaft expanded into a flat-bottomed scooplike apex distad of gonopore, sides of

apex joining bottom at approximately a right angle,

apex truncate in caudodorsal aspect, greatest width

two-thirds greatest distance between dorsal apodemes in caudodorsal aspect (Figure 18c).

Female seventh abdominal sternum with posterior margin broadly and shallowly emarginate, median emargination broader than the two lateral

emarginations combined (Figure 18B). First valvula

(Figure 18A) with basidorsal median lobe developed

into thin vertical plates that do not extend as far

cephalad as the articulating lobe; base of valvula
NUMBER 74 31

with its complement appearing concave in ventral

aspect, with articulating lobe arising laterad of median basidorsal lobe, the two appearing contiguous

in ventral aspect (Figure 18A).

The holotype was not designated in the original

description which was based on external characters.

Ossiannilsson (1937) listed the type as a male and

illustrated the aedeagus of the "type." The type

locality is Carices I Rohne, Sweden. The distribution is Holoarctic.

Psammotettix con finis (Dahlbom) is close to P.

nodosus (Ribaut) in the form of the aedeagus. It

belongs to the striatus complex. In external characters it is close to P. alienus (Dahlbom), P. totalus

(DeLong and Davidson), and P. attenuens (DeLong

and Davidson) in the Nearctic region. The shape

of the posterior edge of the female seventh abdominal sternum of confinis is close to P. alexanderi,

new species.

Remane (1965) described Psammotettix viridiconfinis Remane. The aedeagus of the Nearctic P. confinis (Dahlbom) (Figure 18c) differs from Remane's

illustration of confinis as much as does viridiconfinis. Further study may warrant recognizing specific or subspecific standing in the Nearctic form.

Psammotettix harrimani (Ashmead) would be the

type of this new taxon if Beirne's (1954) interpretation of the type were confirmed.

Kontkanen (1950) reports dryinid and pipinculid

parasites attacking this species in Finland.

Moravskaia (1956) reports this species as being

polyphagous. It is found on grasses and herbaceous

plants of the Compositae, Juncaceae, and Cyperaceae.

Psammotettix obesus Knull

1-K.I RE 19

Psammotettix obesus Knull, 1954, p. 57.

Length of male 2.9 to 3.3 mm, of female 3.0 to

3.6 mm; width of head, including eyes, of male 0.9

to 1.0 mm, of female 1.0 to 1.1 mm. Crown with

median length 1.1 to 1.4 length next to eye, with median sulcus extending anteriorly from 0.6 to 0.9

median length. Color varying as in P. totalus (DeLong and Davidson). Scutellum marked with median stripe, or with two spots at posterior edge of

pronotum and two lateral stripes, brown.

Pronotum with round pits as in P. totalus (DeLong and Davidson). Scutellum with transverse

suture that is almost straight medially with lateral

ends angled anteriorly. Macropterous form with

fore wings reaching or exceeding abdominal apex,

with hind wings shorter than, equal to, or longer

than abdomen, reaching distal end of first apical

cell of fore wings at rest position.

Aedeagus with shaft tubular throughout length

to gonopore; apex of shaft expanded with lateral

flanges in caudodorsal aspect (Figure 19D). Median

longitudinal distance from gonopore to apex less

than or equal to greatest width of apex of shaft in

caudodorsal aspect. Apex of shaft almost round in

caudodorsal aspect.

Female seventh abdominal sternum slightly produced posteriorly, with a shallow median emargination (Figure 19B). First valvula with basidorsal

median lobe well developed and sclerotized, not

extending as far cephalad as the articulating lobe,

which is strongly folded laterad, its form resembling an interrogation point, its mesal border not

well defined in ventral aspect (Figure 19A).

The holotype, allotype, and two male paratypes

are in the Knull collection at the Ohio State University. The type series is from Grand Teton National Park and San Francisco Mountains, Arizona.

The allotype and two male paratypes are from the

type locality.

Psammotettix obesus Knull is one of the shorter

species of Psammotettix. Its front wings at rest

reach or slightly exceed the apex of the abdomen.

The short wings of P. obesus Knull are like those

of P. alexanderi, new species, P. cahuillus (Van

Duzee), P. lapponicus (Ossiannilsson), and P. beirnei, new species, in the Nearctic region. The shape

of the aedeagus of obesus is close to beirnei. The

shape of the seventh abdominal sternum and the

characters at the base of the first valvula of the

female are close to P. amplus (DeLong and Davidson). A specimen from Yellowstone Park, Fry Pan

Lake, appears to be intermediate between obesus

and beirnei, the aedeagus resembling beirnei in

lateral aspect but resembling obesus in caudodorsal

aspect.

Psammotettix beirnei, new species

FIGURE 20

Psammotettix cephalotes.—Bcirne, 1956, p. 103.

Psammotettix obesus— Beirne (not Knull). 1956, p. 103.
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

B

FIGURE 19.—Male and female genitalia and female seventh abdominal sternum of Psammotettix

obesus Knull: A, first valvulae and first valvifer in ventral aspect; B, seventh sternum in ventral

aspect; c, aedeagus in caudodorsal aspect; D, same in lateral aspect.

Length of male 2.80 to 3.10 mm, of female 3.24 mm,

width of head, including eyes, of male 0.93 to 1.00

mm. Crown with median length 1.20 to 1.41 length

next to eye, with median sulcus extending anteriorly

from 0.64 to 0.73 median length. Ground color of

head, pronotum, wings, and scutellum varying

from light cream to cinereous. Crown and pronotum

marked with brown as in P. totalus (DeLong and

Davidson). Almost all cells of front wings marked

with some brown. Hind wings transparent, some

veins darkened. Scutellum with two anterior median

spots partly overlaid by pronotum and triangular spot behind transverse suture, brown. Abdomen

with all terga black, often with transverse creamy

band on posterior edge of each segment.

Pronotum with round pits as in P. totalus (DeLong and Davidson). Scutellum with transverse

suture undulate. Macropterous form with fore wings

at rest exceeding apex of abdomen. Hind wings at

rest shorter than abdomen, not reaching distal end

of central anteapical cell of fore wings.

Aedeagus similar to that of P. alexanderi, new

species, differing as discussed above. Median longitudinal distance from gonopore to apex greater

than greatest width of apex of shaft in caudodorsal

aspect. Apex appearing elongate in caudodorsal

aspect. Posterior end of connective strongly sclerotized and bifid at articulation with aedeagus (Figure

20c).

Female seventh abdominal sternum relatively
NUMBER 74 33

FIGURE 20.—Male and female genitalia and female seventh abdominal sternum of Psammotettix

beirnei, new species, and aedeagus of P. cephalotes (Herrich-Schaeffer): A, first valvulae and

first valvifer in ventral aspect; B, seventh sternum in ventral aspect; c, connective and style in

dorsal aspect; D, aedeagus of P. beirnei, new species, in caudodorsal aspect; E, same in lateral

aspect; F, aedeagus of P. cephalotes (Herrich-Schaeffer) in caudodorsal aspect; c, same in lateral

aspect.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

B

FIGURE 21.—Male and female genitalia and female seventh abdominal sternum of Psammotettix

alexanderi, new species: A, first valvulae and first valvifers in ventral aspect; B, seventh sternum

in ventral aspect; c, connective and style in dorsal aspect; D. aedeagus in caudodorsal aspect;

E, same in lateral aspect.

strongly produced posteriorly with shallow concave

median emargination (Figure 20B). First valvula

with basidorsal median lobe well developed and

sclerotized, not extending as far cephalad as the

articulating lobe, which is well defined mesally and

laterally, appearing to arise and diverge from median basidorsal lobe (Figue 20A).

Holotype male and allotype from First Summit

Mt. Harry, N.W.T., 7600' 20 August 1952 (G. J.

Spencer) in Canadian National collection.

Psammotettix beirnei is named in honor of B. P.

Beirne. It was determined as P. cephalotes (HerrichSchaeffer) by Beirne (1956). This species differs

from specimens of P. cephalotes, from Germany, in

color, body form, and shape of aedeagus. The European specimens are elongate with both fore and

hind wings exceeding the apex of abdomen when

at rest position; the Canadian specimens are short

with fore wings exceeding the apex of abdomen

and hind wings shorter than abdomen when at rest

position. The European specimens are a uniform

creamy ivory color with few markings; the Canadian specimens vary from a ground color of creamy

to cinereous heavily marked with brown. There is

a definite difference in size and in the lateral profile

of the aedeagus between the European and Canadian specimens (Figures 2c, E, and c). The stem of

the connective is bifid posteriorly in the Canadian specimens, not bifid in the European specimens. The published illustrations of the aedeagus
NUMBKR 74 35

of P. cephalotes (Herrich-Schaeffer) differ from the

Canadian specimens, and in a personal communication Dr. R. Remane, Marburg, Germany, expressed

the opinion that these were two different species.

In view of the above differences the Canadian form

is being treated as a new species.

Psammotcttix beirnei, new species, is closely related to the lapponicus group, possessing the characteristic elliptic-ally expanded or flanged aedeagal

shaft apex. This species is very closely related to

P. alexanderi, new species, as has been previously

discussed.

The shape of the posterior edge of the female

seventh abdominal sternum of P. beirnei is close to

P. attcmiens (DeLong and Davidson). The wing

lengths of P. beirnei, new species, are close to those

of P. alexanderi, new species, P. obesus Knull, P.

lapponicus (Ossiannilsson), P. excavatus (Oman),

P. shoshone (DeLong and Davidson), and P.

cahuillus (Van Duzee). In the characters at the

base of the first valvula, P. beirnei, new species, is

close to P. ampins (DeLong and Davidson), P. lapponicus (Ossiannilsson), and P. alexanderi, new

species. In its darkened veins in the hind wing, this

species is closely related to P. lapponicus (Ossiannilsson) and P. den tat us Knull.

Psammotettix alexanderi, new species

FIGURE 21

Length of male 2.6 to 2.84 mm, of female 2.85 to

3.25 mm; width of head, including eyes, of male

0.93 to 0.95 mm, of female 1.00 to 1.08 mm. Crown

with median length 1.23 to 1.50 length next to

eye, with median sulcus extending anteriorly from

0.56 to 0.65 median length.

Color of crown and pronotum varying as in P.

totalus (DeLong and Davidson). Ground color of

crown, pronotum, scutellum, wings, and abdomen

as in totalus. Wings unmarked or with apical cells

very lightly marked with brown. Scutellum unmarked. All abdominal terga heavily marked with

black medially except eighth.

Pronotum with round pits as in P. totalus (DeLong and Davidson). Scutellum with transverse suture straight. Males macropterous with fore wings

exceeding apex of abdomen, hind wings at rest

shorter than abdomen, not exceeding tip of brachial

cell of fore wings. Females subbrachypterous with

front wings at rest not attaining apex of abdomen,

hind wings as in male.

Aedeagus with shaft tubular throughout length

to gonopore; apex of shaft expanded into lateral

flanges in caudodorsal aspect (Figure 2ID). Greatest

width of tip in caudodorsal aspect five-eighths or

more the longitudinal distance from gonopore opening to tip (Figures 21 D and E). Some specimens with

tubercles ventrad of flanges. Connective weakly concave in ventral aspect at articulation with aedeagus.

Female seventh abdominal sternum almost truncate apically, with weak median and two lateral

emarginations of approximately equal size (Figure

21B). First valvula with median basidorsal lobe and

articulating lobe as in P. obesus Knull; the articulating lobe not as strongly folded laterad as in

obesus and with its mesal border distinct cephalad.

Holotype male, White Mts., New Hampshire,

U.S.A., 23 August 1935 (C. P. Alexander); allotype,

Kings Ravine, White Mts., New Hampshire, U.S.A.,

26 August 1935 (C. P. Alexander). These and

eighteen paratypes are in the National Museum of

Natural History.

This species is named in honor of the collector,

C. P. Alexander. Psammotettix alexanderi, new

species, is close to other members of the previously

mentioned lapponicus group. The male is very

close, in external characters, to P. beirnei, new

species. The female can usually be distinguished

from beirnei, new species, and Psammotettix lapponicus (Ossiannilsson) by the subbrachypterous

wings.

The aedeagus of the male is very similar to the

aedeagus of P. beirnei, new species. The tip of the

aedeagal shaft of P. cephalotes (Herrich-Schaeffer)

is curved anteriorly at an angle to the axis of the

shaft in lateral aspect. The tip of the aedeagal shaft

of P. alexanderi, new species, is not curved at any

appreciable angle to the axis of the shaft in lateral

aspect. The connective of P. alexanderi is not bifid

at articulation with aedeagus as is that of P. beirnei.

Based on the posterior edge of the female seventh

abdominal sternum, P. alexanderi is closely related

to P. lapponicus (Ossiannilsson).

Psammotettix lapponicus (Ossiannilsson)

FIGURE 22

Deltocephalus lapponicus Ossiannilsson, 1938, p. 6.

Psammotettix lapponicus.—Ribaut, 1938, p. 166.
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

B

FIGURE 22.—Male and female genitalia and female seventh abdominal sternum of Psammotettix

lapponicus (Ossiannilsson): A, first valvulae and first valvifers in ventral aspect; B, seventh

sternum in ventral aspect; c, connective and style in dorsal aspect; D, aedeagus in caudodorsal

aspect; K, same in lateral aspect.

Length of male 2,.5 to 2.8 mm, of female 2.7 mm;

width of head, including eyes, of male 0.90 to 0.92

mm, of female 0.97 mm. Crown with median length

1.31 to 1.44 length next to eye, with median sulcus

extending anteriorly from 0.64 to 0.73 median

length. Ground color of head, pronotum, and scutellum cream. Crown with upper horizontal stripes

of front visible from above, with two anterior

stripes paralleling stripes of front, with two broad

lateral longitudinal stripes and two spots anterior

to eyes, brown. Pronotum with six longitudinal

brown stripes. Scutellum marked as in P. beirnei,

new species. Wings transparent, almost all cells

marked with some brown. Abdomen with all terga

black. Hind wings transparent, some veins darkened.

Pronotum with round pits as in Psammotettix

totalus (DeLong and Davidson). Scutellum with

transverse suture crescentiform to straight. Macropterous form with wings as in P. beirnei, new species.

Aedeagal shaft with apex in the form of a very

elongate ellipse, length from gonopore to apex 2.6

times the greatest width in caudodorsal aspect

(Figure 22D). Posterior end of connective truncate,

not strongly bifid at articulation with aedeagus.

Female seventh abdominal sternum not produced

posteriorly, with median and two lateral emarginations of about equal size. First valvula with base as

in Psammotettix beirnei, new species.

The holotype is in the Ossiannilsson collection.

The type locality is Arvidjous, Sweden. The distribution is Sweden (Ossiannilsson, 1938), and Chesterfield, Northwest Territories (Beirne, 1954).
NUMBKR 74 37

The specimens from the Northwest Territories

(Figure 22D) have an aedeagal shaft with an apex

that is more elongate and slender than is the apex

of a paratype from the type locality. In lateral aspect

the Northwest Territory specimens are closer to

Ossiannilssons illustration of the type than to the

paratype. These specimens are similar to the paratype in external characters.

Psammotettix revae Knull

FIGURE 23

Psammotettix revae Knull, 1954, p. 57.

Length of male 2.90 mm, of female 3.20 mm; width

of head, including eyes, of male 0.99 mm, of female

0.98 mm. Crown with median length 1.21 to 1.28

length next to eye, with median sulcus extending

anteriorly from 0.69 to 0.73 median length. Ground

color of head, pronotum, and scutellum light cream.

Crown with two apical triangles, and markings

around each ocellus brown, the usual striate markings missing or very faint on other areas. Wings

creamy subhy aline, very sparsely and faintly marked

with brown. Hind wings transparent; veins not

darkened.

Pronotum with round pits as in P. totalus (DeLong and Davidson). Scutellum with transverse

suture straight. Macropterous form with fore wings

exceeding apex of abdomen. Hind wings at rest

extending only to midlength of apical cells of front

wings.

Aedeagus with apex in the form of an ellipse;

length from gonopore to apex 1.88 times greatest

width in the caudodorsal aspect (Figure 23c, D).

Posterior end of connective bifid at articulation

with aedeagus, as in Psammotettix beirnei, new

species.

Female seventh abdominal sternum with posterior

margin as in P. lividellus (Zetterstedt), with median

FIGURE 23.—Male and female genitalia and female seventh abdominal sternum of Psammotettix

revae Knull: A, first valvulae and first valvifer in ventral aspect; B, seventh sternum in ventral

aspect; c, aedeagus in caudodorsal aspect; D, same in lateral aspect.
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

emargination flanked by two smaller lateral emarginations (Figure 23B). First valvula with basidorsal

median lobe well developed and sclerotized, extending about equally as far cephalad as the articulating lobe which appears to be contiguous with

the basidorsal median lobe (Figure 23A).

The type series from Reva Alkali Flats, Reva,

Springfield, and Buffaloe, South Dakota, are in the

Knull collection, Ohio State University.

Psammotettix revae Knull is very close to P.

cephalotes (Herrich-Schaeffer), the chief distinguishing character being the notched or bifid posterior

end of the connective in revae which is unnotched

in cephalotes. Psammotettix revae appears to be

closer to cephalotes (Herrich-Schaeffer) than beirnei,

new species. This form is possibly a subspecies of

cephalotes (Herrich-Schaeffer).

Psammotettix revae is a member of the lapponicus

group. Based on the posterior edge of the female

seventh abdominal segment and on characters at

the base of the first valvula, it is close to P. lividellus

(Zetterstedt) and lapponicus (Ossiannilsson). Based

on wing lengths this species is close to P. knullae.

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Index to Species

(Synonyms are in italics. Page numbers of principal accounts are in boldface.)

affinis, Deltocephalus, 2, 19

alboniger, Psammotettix, 25

alexanderi, Psammotettix, 8, 16, 20, 22, 25, 31, 34, 35

alienus, Psammotettix, 2,8, 9, 15, 28. 29, 31

allienus, Psammotettix, 28

amplus, Psammotettix, 8, 9, 16, 17, 18, 19, 22, 24. 25, 31, 35

angulatus, Psammotettix, 13

aspci, Psammotettix, 2, 9, 19. 20, 25

attcnuens, Psammotettix, 8, 9, 15, 16. 25, 31, 35

beirnei, Psammotettix, 8, 9, 20, 31, 33, 34, 35, 36, 37, 38

brcviceps, Deltocephalus, 28

cahuilla, Deltocephalus. 13

cahuilla, La talus, 13

cahuillus, Psammotettix, 8, 9, 13. 14, 15, 16, 17, 24, 25, 31, 35

cahuillus, Laevicephalus, 13

cephalotes, Psammotettix, 2, 33, 34, 35, 38

cephalotes, Psammotettix. 31

confinis, Psammotettix, 1, 2, 8, 9, 28. 30, 31

dentatus, Psammotettix, 2, 8, 9, 23. 24, 25, 35

emarginatus, Psammotettix, 8, 9, 24, 25, 27, 28

excavatus, Psammotettix, 8. 9, 16, 25, 26, 27, 35

exeisus, Deltocephalus, 20

exisus, Deltocephalus, 20

ferratus, Psammotettix, 6, 7, 10. 13, 25

Havostriatus, Aphrodes, 23

harrimani, Deltocephalus, 2. 31

heydeni, Deltocephalus, 28

knullae, Psammotettix. 7. 9. 12, 13. 25. 38

koelcriac, Psammotetlix, 13

lapponicus. Psammotettix, 2, 8. 9, 16, 20, 22. 25, 31. 35. 36, 38

latipex, Deltocephalus, 2

lividcllus, Psammotettix, 1, 2. 8, 9. 19. 20. 21, 22, 23, 25. 37, 38

maritimus, Athysanus, 1, 5

maiiiiinus, Psammotetlix, I

nodesus. Psammotettix, 31

obesus, Psammotettix, 2, 8. 9. 16, 20, 22. 31. 32, 35

obesm, Psammotettix, 31

Psammotettix, 5

revae, Psammotettix, 2, 8, 9, 22, 25. 37, 38

shoshone, Psammotettix, 8.9, 16, 25. 26, 27, 28, 35

spathifer, Psammotettix, 29

striatus, Psammotettix. 2, 20

striatus, Psammotettix. 1, 22

striatus, Jassus, 28

striatus var. thenii, Deltocephalus. 28

striatus, Psammotettix, 29

thenii, Psammotettix, 28

totalus, Psammotettix. 7, 9, 11, 17. 18, 19, 24, 25, 31, 32, 35,

36,37

viiidiconfinis, Psammotettix, 31

40 * U. S. GOVERNMENT PRINTINC OFFICE : 1971 O - 403-48S


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titles will carry a final line indicating the higher categories to which the taxon is referable:

"(Hymenoptera: Sphecidae)." Include an abstract as an introductory part of the text. Identify

the author on the first page of text with an unnumbered footnote that includes his professional

mailing address. A table of contents is optional. An index, if required, may be supplied by the

author when he returns page proof.

Two headings are used: (1) text heads (boldface in print) for major sections and chapters

and (2) paragraph sideheads (caps and small caps in print) for subdivisions. Further headings

may be worked out with the editor.

In taxonomic keys, number only the first item of each couplet; if there is only one couplet,

omit the number. For easy reference, number also the taxa and their corresponding headings

throughout the text; do not incorporate page references in the key.

In synonymy, use the short form (taxon, author, date, page) with a full reference at the

end of the paper under "Literature Cited." Begin each taxon at the left margin with subsequent lines indented about three spaces. Within a taxon, use a period-dash (.—) to separate

each entry. Enclose with square brackets any annotation in or at the end of the taxon. For

synonymy and references within the text, use the author-date system: "(Jones 1910)." Use

the colon system for page references: "(Jones 1910:122)," and abbreviate further data:

"(Jones 1910:122, fig. 3, pi. 5: fig. 1)."

Simple tabulations in the text (e.g., columns of data) may carry headings or not, but they

should not contain rules. Formal tables must be submitted as pages separate from the text, and

each table, no matter how large, should be pasted up as a single sheet of copy.

Use the metric system instead of (or in addition to) the English system.

Illustrations (line drawings, maps, photographs, shaded drawings) usually can be intermixed throughout the printed text. They will be termed Figures and should be numbered

consecutively; however, if a group of figures is treated as a single figure, the individual components should be indicated by lowercase italic letters on the illustration, in the legend, and

in text references: "Figure 9b." Submit all legends on pages separate from the text and not

attached to the artwork. An instruction sheet for the preparation of illustrations is available

from the Press on request.

In the bibliography (usually called "Literature Cited"), spell out book, journal, and article

titles, using initial caps with all words except minor terms such as "and, of, the." (For capitalization of titles in foreign languages, follow the national practice of each language.) Underscore (for italics) book and journal titles. Use the colon-parentheses system for volume number

and page citations: "10(2) :5-9." Spell out such words as "figures," "plates," pages."

For free copies of his own paper, a Smithsonian author should indicate his requirements on

"Form 36" (submitted to the Press with the manuscript). A non-Smithsonian author will

receive fifty free copies; order forms for quantities above this amount, with instructions for

payment, will be supplied when page proof is forwarded.