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OCCASIONAL PAPERS
OF THE

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California Academy of Sciences

No. 114^ 27 pages, 72 figures

A REVISION OF THE DELTOCEPHALINE

LEAFHOPPER GENUS GIPRUS OMAN

(HOMOPTERA: CICADELLIDAE: DELTOCEPHALINAE)

By
Jane R. Sawbridge

SAN FRANCISCO
PUBLISHED BY THE ACADEMY

January 16, 1975

COMMITTEE ON PUBLICATION
George E. Lindsay, Chairman
Diana R. Young, Editor

OCCASIONAL PAPERS

OF THE

California Academy of Sciences

No. 114, 27 pages, 72 figures.

A REVISION OF THE DELTOCEPHALINE

LEAFHOPPER GENUS GIPRUS OMAN

(HOMOPTERA: CICADELLIDAE: DELTOCEPHALINAE)

By

Jane R. Sawbridge

Department of Entomology,
Oregon State University,
Corvallis, Oregon 97331

Abstract: Members of the deltocephaline leafhopper
genus Giprus Oman live on grasses, primarily in the
desert scrub, chapparal, and marine sand dune
communities of western North America. Six species
are redescribed and ten new species are described.
Recharacterization of the genus results in the
exclusion of Laevicephalus ineongruus Oman.
Differentiation of Giprus species is primarily based
on the male genital structures; characterization of
species is through illustration, verbal description,
and a diagnostic key. The fifth instar nymph is
characterized. Significant inf raspecif ic variation
occurs in the size and shape of the accessory
aedeagal processes of G. siskiyou (Oman) .

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Acknowledgments

I would like to thank Dr. Paul Oman for suggesting the
project, helping to collect specimens, and offering advice
and constructive criticism throughout the study. Dr. Paul
0. Ritcher kindly read and commented on the manuscript. I
acknowledge the loan of material by the United States
National Museum of Natural History, California Academy of
Sciences, University of Kansas, University of Arizona, and
University of California at Davis. This study was supported
primarily by Grants GB-7706 and GB-28292 "Phylogeny and
Distribution of Nearctic and Palearctic Leaf hoppers" from
the National Science Foundation to Dr. Paul Oman. Addi-
tional support by the Department of Entomology, Oregon State
University is acknowledged.

Introduction

Giprus Oman is a member of that subgroup of the Tribe
Deltocephalini characterized by a linear connective and
articulation between the connective and the aedeagus (Oman,
1949). Members of the genus are small to medium in size,
cinereous to amber in color, and usually with fuscous
markings on the wings and head. The known species inhabit
western North America, occurring in climatic zones ranging
from coastal to continental at latitudes from 32°N. to 47°N.
and from sea level to about 2000 m. elevation. They are
associated with various species of grasses, usually in
chaparral, desert scrub, or sand dune communities. Giprus
is most closely related to Laevicephalus DeLong but may be
distinguished from it by the structure of the aedeagus.

The first known species, described by Van Duzee (1892)
as Deltocephalus cinerosus , was subsequently placed in the
subgenus Laevicephalus by DeLong (1926). When Laevicephalus
was elevated to generic status in 1929 by DeLong and Slees-
man, it was noted that the genus contained species with a
great diversity of genitalic structures. In 19 37, primarily j
on the basis of the male genitalia, Oman recognized and
characterized a "cinerosus group" of Laevicephalus which
included L. cinerosus and five additional species: L.
angelus Oman, L. pacificus Oman, L. joaquinus Oman, L.
Siskiyou Oman, and L. incongruus Oman. In 1949 Oman removed
the "cinerosus group" and L. cartwrighti , which he had des-
cribed in 19 32, from Laevicephalus and erected the genus
Giprus to contain them.

The need for a comprehensive review of the genus became
apparent when it was discovered that samples of several
populations encountered in Oregon and California could not
be referred to described species. The study reported here
endeavors to summarize knowledge of a group that is now
known to be taxonomically much more complex than was pre-
viously suspected.

No. 114] SAWBRIDGE: GIPRUS

Materials and Hethods

Over 3000 specimens of Giprus , including one holotype
and 87 paratypes of seven species, were examined. Most were
collected during the course of this study. Borrowed mater-
ial served primarily to confirm the identity of previously
described species.

Drawings were prepared with the aid of a micrometer disc
ruled in 0.1 mm. squares placed in the ocular of a stereo-
scopic microscope, and drawing paper with a 1/2 inch square
grid. Anatomical details were checked with a compound
microscope at 150x. All drawings of homologous structures
are to the same scale. Drawings of the internal male
genitalia show a dorsal view of the connective, aedeagus,
and left style; the right style is omitted. In addition, a
lateral view of the aedeagus from the left side is shown.

Measurements were made with a filar micrometer. Total
body length was measured as the distance from the apex of
the crown to the wing tips. Median length of the head and
its width immediately anterior to the eye were measured and
the ratio of width (W) to length (L) computed for each
individual in a sample. The mean of the W/L values for a
sample (usually 10 of each species) was computed and
assigned as the W/L value for the species.

Comparative morphology was used as the primary basis for
classification and species discrimination; field observation
of populations was used as additional evidence to supplement
and reinforce conclusions.

Field collection of specimens was undertaken with four
main objectives: to obtain large samples (50 or more
specimens) at each collection site for the study of infra-
population variation as a means of understanding infra-
specific variation; to examine and compare habitats; to
determine host associations; and to determine whether G.
Siskiyou is a single variable species or a group of sibling
species. In order to determine whether variation in G.
siskiyou could be considered topoclinal or ecoclinal in
nature, two transects were made through the region where the
species is most commonly found. The north-south transect
(170 km.; 12 sites) extended between Mt. Ashland, Oregon,
and 57.4 km. north of Redding, California. The east-west
transect (233 km.; 14 sites) extended between Cedarville,
California, and Klamath River, California.

The aedeagi of specimens from each site where G.
siskiyou was collected were compared in a study of infra-
specific variation. In order to determine the amount of
variation present in any one area at a given time, individ-
uals collected at a single site on the same day were
compared.

Host associations were determined by selective
sweeping of grass inflorescences.

Holotypes of new species will be deposited in the United
States National Museum of Natural History (NMNH) except for
that of Giprus decurvatus which is deposited in the Cali-
fornia Academy of Sciences. Paratypes of all new species

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

described in this paper, except G. vastitatus , were
collected at the same locality as the holotype.

Characterization of the Genus and Species Groups

Head approximately twice as long medially as the
shortest distance between the eyes, giving the head a
rather pointed appearance which is accentuated by oblique
fuscous dashes on each anterolateral submargin of the crown
(fig. 5). Pronotum brown with parallel rows of longitudinal
pale stripes. Clypeus usually dark colored with horizontal
rows of thin pale lines and a pale median line. Ratio of
the head width immediately anterior to the eyes to the median
length of the head between 1.20:1 and 1.30:1 in most species.

Significant forewing characters are the small appendix,
inner anteapical cell almost always closed basally, and the
central anteapical cell usually constricted (fig. 3) .
Forewings transparent with ground color cinereous to amber
in all species except G. cartwvighti which has a greenish-
gold ground color. In most species at least some individ-
uals have fuscous markings concentrated around the
periphery of the wing cells.

Most specimens are between 3.50 mm. and 4.50 mm. long.
Males tend to be slightly smaller than females although
frequently the smallest female in a sample may be smaller
than the largest male.

Important characters of the genitalia include heavily
setose pygofers in both sexes, broad male plates, large
triangular valve, aedeagus with a basal submembranous hood-
like extension for attachment to the tenth segment, and
aedeagal shaft arising from a point dorsad of the point of
articulation of the phallobase and the connective (fig. 14) .
The latter condition is the main difference between Giprus
and its nearest relative, Laevioephatus , as the articu-
lation of the connective and the phallobase in Laevioephatus
is directly in line with the aedeagal shaft (fig. 15).
Length, curvature, and thickness of the aedeagal shaft, and
position and number of paired aedeagal processes show
considerable diversity among members of the genus, and
therefore species may be grouped by these characteristics
(fig. 1). Group I includes only G. cavtwvighti , placed
alone because of its elongate, sinuate aedeagus unlike any
other in Giprus; Group II includes those with a long slender
aedeagus with short apical processes; Group III includes

those with a short stout aedeagus which is enlarged
apically; and Group IV includes those with a pair of
elongate processes at some point on the aedeagus. Genital
plates of males of Giprus, except for those of G. cart-
wrighti which are elongate and gradually tapering (fig. 13) ,
may be similar for several species with markedly different
internal structures. Three types of plates are present:
plates entirely contiguous with apices acute (fig. 10);
plates broadly rounded, basally contiguous but diverging
posteriorly and abruptly narrowing in some to acute apices
(fig. 11) ; or an intermediate condition with plates

No. 114] SAWBRIDGE: GIPRUS

divergent posteriorly and gradually tapering to acute apices
(fig. 12) . Groupings according to plate type do not
coincide with groupings by aedeagal structure.

The posterior margin of the seventh sternum of the
female is characterized by two pairs of median dentate pro-
jections (figs. 6, 7, 8). The dorsal extension of the
seventh sternum mentioned by Readio (1922) in connection
with his work on the ovipositors of leafhoppers, is readily
visible in cleared specimens of Giprus and appears as either
a single sclerotized structure in some species (figs. 7, 8)
or as a pair of sclerotized lobes connected by a membrane
in others (fig. 6) . The dorsal extension arises from the
seventh sternum posteromedially between the outer pair of
dentate projections. The posterior portion of the ventral
part of the seventh sternum is unpigmented; the heavily
sclerotized dorsal extension gives the posterior margin of
the seventh sternum its characteristic color pattern.

Bluntness or pointedness of the head varies among the
species. Mean W/L values range from 1.20, indicating a very
pointed head, to 1.46, indicating a very blunt head. It is
notable that the species with blunt heads all occupy
coastal habitats, but head shape, while perhaps useful as
an indicator of habitat, is not a reliable character for
species discrimination because there is too much overlap of
values between the species (fig. 2).

Fifth instar nymphs of Giprus are cream colored with
extensive light brown to fuscous markings on the head,
pronotum, wing pads, and abdominal dorsum (fig. 4). The
abdominal venter has fewer dark markings. The eyes, head,
and body of nymphs frequently have a reddish cast.

Laevioephalus inoongruus , placed in Giprus by Oman
(1949), is atypical of Giprus in several characteristics.
It is conspicuously larger than other members of the genus;
the posterior margin of the seventh sternum of the female
is emarginate; and most significantly, the structure of the
aedeagus is unlike that of typical Giprus species. The
aedeagus of L. inoongruus is more like that of Laevioephalus
than that of Giprus in that the point of articulation of the
connective and the phallobase is directly opposite the shaft
of the aedeagus (fig. 16). For these reasons, I remove L.
inoongruus from Giprus and return it to Laevioephalus . On
the basis of the structure of the aedeagus, it would appear
that L. inoongruus would fit most closely into the "parvulus
subgroup" as characterized by Ross and Hamilton (1972).

Several characteristics of G. oartwrighti suggest that
it is not closely related to other species in the genus.
The almost semicircular seventh sternum of the female, the
elongate, tapered genital plates of the male, and the
sinuate aedeagus are reminiscent of Laevioephalus . However,
the position of the aedeagal shaft relative to the con-
nective places the species in Giprus. Structurally, G.
oartwrighti seems to be intermediate between the two genera,
and it is of interest that the only instances where two
species of Giprus occur together are those where G. oart-
wrighti is one of the sympatric pair. This suggests that

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

G. oartwrighti and other Giprus species do not compete
ecologically to such an extent as to allow only one species
to exist at a given locality. Also, G. oartwrighti is most
frequently encountered in open grassland habitats such as
those favored by species of Laevioephalus .

Key to the Species of Giprus

1 Aedeagus approximately 3 times the length

of the connective, slender, and sinuate,

with minute apical processes (fig. 36);

male genital plates attenuate with

diverging apices (fig. 13) ; 7th sternum

of female nearly semicircular with small

median incision (fig. 9) G. oartwrighti

1' Aedeagus no more than 2 times the length of
the connective; male plates short and
stout; 7th sternum of female incised
posterolaterally (figs. 6, 7, 8) 2

2(1') Aedeagus with at least one pair of elongate

retrorse processes that diverge obliquely

from the longitudinal axis of the shaft;

if the longest pair of processes is less

than 1/3 the length of the aedeagal shaft,

then they originate considerably basad

of the apex 3

2' Aedeagus without elongate retrorse processes;
aedeagal armature consisting of stout
retrorse dentate processes, lateral
flanges, or slightly elongate processes
that originate near the apex 8

3(2) Aedeagus with one pair of straight or sinuate

lateral or lateroventral elongate processes

arising at or near the apex and no more

than one other pair of processes 4

3' Aedeagus with 3 pairs of processes, or if

only 2, then the slightly elongate pair

originates considerably basad of the

apex 5

4(3) Aedeagus with a single pair of elongate,

sinuate, lateroventral apical processes
about 3/4 the length of the aedeagal
shaft; no ventral processes (fig. 29)
G. vastitatus

4' Two pairs of aedeagal processes: short
ventral processes originating at the
base of the elongate lateral processes
and curving mesad (figs. 30, 31);
lateral processes either straight or
sinuate in lateral view G. oalifornious

No. 114] SAWBRIDGE: GIPRUS

5(3') Aedeagus with 3 pairs of processes: one

pair short and ventral, one pair elongate

and lateral, and one pair very short

and dorsal 6

5' Aedeagus with fewer than 3 pairs of

processes 7

6(5) All pairs of processes apparently origi-
nating at the same point on the
aedeagal shaft; ventral processes not
visible in dorsal view (fig. 35) ... G. cinevosus

6' Elongate processes clearly originating

basad of the other processes; ventral
processes extending slightly laterad and
therefore visible in dorsal view
(fig. 34) G. desertovum

7(5') Aedeagus with dorsal and lateral processes

only; shaft stout and straight (fig. 33)

, G. oajon

T Aedeagus with lateral and lateroventral pro-
cesses only; shaft somewhat sinuate
(fig. 32) G. pacifieus

8(2') Aedeagus with a single pair of lateroventral

processes 9

8' Aedeagus with more than one pair of processes

or processes not as above 10

9(8) Aedeagal shaft slender, slightly decurved,
and with acute apex in dorsal view;
processes pointing posteriorly in
lateral view (fig. 22) G. arenarius

9' Aedeagal shaft not overly slender; processes
pointing anteriorly in lateral view
(fig. 21) G. hidentatus

10(8') Aedeagus with a single pair of dorsal

processes combined with a pair of more

or less well developed (and at times

asymmetrical) lateral flanges (figs.

17-20) G. siskiyou

10' Aedeagus with more than one pair of

processes ; no lateral flanges 11

11(10') Aedeagus with 2 pairs of processes 12

11' Aedeagus with 3 pairs of processes 14

12(11') Aedeagal shaft strongly decurved with
dorsal and lateral processes well
developed; shaft fairly robust (fig.
24) G. decurvatus

12' Aedeagal shaft not strongly decurved; if
sli<^tly decurved then the processes
very small 13

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

13(12') Aedeagus with dorsolateral and ventro-
lateral processes very small (fig. 23) ;
female 7th sternum much longer medially
than laterally (fig. 8) G. joaquinus

13' Aedeagus with dorsal and lateroventral

processes well developed (fig. 25)... G. angelus

14(11') Aedeagus with a pair of short, thin,

lateral processes arising distinctly

basad of the bulbous apex (fig. 21).. G. orarius

14' Aedeagus without lateral processes as

above 15

15(14') In dorsal view aedeagus 4 times as long as
greatest width of apical processes; in
lateral view apical processes oriented
obliquely to the longitudinal axis of
the aedeagal shaft (fig. 28) G. agrestis

15' In dorsal view aedeagus twice as long as
the greatest width of the apical pro-
cesses; in lateral view apical processes
essentially transverse to the longi-
tudinal axis of the aedeagal shaft
(fig. 26) G. arassus

Descriptions of Species

Giprus oavtwvighti (Oman) .

Laevioephalus oartwrighti Oman 1932, p. 90.

LENGTH. Female: 4.01 mm. - 4.54 mm.; male: 3.87 mm. -
4.19 mm.

COLOR. Pale, pattern indistinct, ground color greenish
gold.

HEAD SHAPE. W/L = 1.20.

FEMALE 7TH STERNUM. Dorsal extensions separate and
posterior margin semicircular; dentate projections on pos-
terior margin equal in length and projecting only slightly
(fig. 13).

MALE GENITALIA. Plates elongate and gradually tapering
to acute diverging apices. Aedeagus whiplike, sinuate, and
with minute apical processes. Styles flared near tip (fig.
36) .

HOLOTYPE. Male, Sloughhouse, California, 19 May 19 31,
Cartwright.

DISTRIBUTION. CALIFORNIA: Butte County: Oroville;
Fresno County: Coalinga; Kern County: Tehachapi Pass;
Monterey County: Bryson; San Bernardino County: 3.2 km. S.
Rock Camp, Lake Arrowhead; San Joaquin County: Locke ford;
Santa Clara County: Los Gates Canyon; Siskiyou County:
25.7 km. E. Klamath River, 18.5 km. N. Weed, 25.1 km. N.
Yreka, 0.8 km. E. Yreka. OREGON: Jackson County: 5.1 km.

No. 114] SAWBRIDGE: GIPRUS

S. Ruch, Agate Desert, Agate Lake, Upper Table Rock N. of
Medford; Josephine County: 8 km. SW. Cave Junction (April -
December) .

HABITAT AND HOSTS. Usually collected in mixed grass-
land, host unknown.

Giipvus stskiyou (Oman) .

Laevioephalus siskiyou Oman 1937, p. 478.

LENGTH. Female: 3.76 mm. - 4.5 8 mm.; male: 3.5 0 mm. -
4. 32 mm.

COLOR. Dark, pattern distinct.

HEAD SHAPE. W/L = 1.27.

FEMALE 7TH STERNUM. Dorsal extensions separate; inner
pair of dentate projections on the posterior margin shorter
than outer pair.

MALE GENITALIA. Genital plates entirely contiguous with
acute apices (fig. 10) . Aedeagus with one pair of dorsal
processes near apex. Lateral flanges basad of processes
very variable, may be more or less well developed, and are
often asymmetrical. Style tips small and blunt. Aedeagal
apex notched in lateral view (figs. 17-20 & 37-72).

HOLOTYPE. Male, Mt. Shasta, California, 29 June 1935,
Oman.

DISTRIBUTION. CALIFORNIA: Modoc County: 24.1 km. NE.
Alturas , 11.3 km. NE. Alturas , Fandango Pass 1829 m. ;
Shasta County: 20.9 km. S. Castella, 3.2 km. S. Castella;
Siskiyou County: Bray, 26.1 km. N. Yreka, 21.6 km. N.
Yreka, 0.8 km. E. Yreka, 25.7 km. E. Klamath River, 18.5 km.
N. Weed, N. of Weed, 4.8 km. S. Weed, Mt. Shasta City, S.
of Mt. Shasta City, 24 km. SE. Mt. Shasta, Dunsmuir. IDAHO:
Bonneville County: 7 km. SE. Irwin. NEVADA: Ormsby
County: Carson City. OREGON; Jackson County: Mt. Ashland
1402 m. and 1250 m. , Siskiyou Summit, Pinehurst; Klamath
County: 30.6 km. W. Klamath Falls, Klamath Falls, 35.4 km.
E. Klamath Falls, 48.3 km. E. Klamath Falls, 57.9 km. E.
Klamath Falls, 9.6 km. E. Bly; Harney County: 17.7 km. E.
Frenchglen. WASHINGTON: Kittitas County: Cliff dell;
Pierce County: Mt. Rainier (Cottonwood Flats); Yakima
County: Naches , 40.2 km. S. Toppenish, 57.9 km. S. Toppenish
(May - July) .

HABITAT AND HOSTS. Chaparral and desert scrub on Stipa
lemmonii , Poa species, Bromus species (one locality),
Sitanion jubatum (one locality) .

Giprus arenarius Sawbridge, new species.

LENGTH. Female: 3.68 mm. - 3.96 mm.; male: 3.30 mm. -
3. 86 mm.

COLOR. Very pale, little pigment deposited around
periphery of wing cells.

HEAD SHAPE. W/L = 1.43.

FEMALE 7TH STERNUM. Dorsal extensions separate; both
pairs of dentate projections on posterior margin of equal

10 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

length.

MALE GENITALIA. Genital plates divergent posteriorly
and gradually tapering to acute apices (fig. 12) . Aedeagal
shaft slightly decurved and supporting a single pair of
lateroventrally projecting processes. Apex of aedeagus
acute. Styles broad at tips (fig. 22).

HOLOTYPE. Male, 4.0 km. N. Waldport, Oregon, 3 July
19 70, Oman.

PARATYPES. 21 females, 22 males, 3 July 1970, Oman;
13 females, 15 males, 1 July 1971, Sawbridge.

DISTRIBUTION. OREGON: Coos County: 8 km. S. Bandon;
Curry County: 9.6 km. S. Brookings, 12.8 km. N. Gold
Beach; Lane County: 13.7 km. N. Florence; Lincoln County:
4.0 km. N. Waldport (June - September).

HABITAT AND HOSTS: Poa macvantha and Festuca rubra on
sand dunes.

Giprus hidentatus Sawbridge, new species.

LENGTH. Female: 4.26 mm. - 4.49 mm.; male: 4.12 mm.

COLOR. Medium brown.

HEAD SHAPE. W/L = 1.27.

FEMALE 7TH STERNUM. Dorsal extensions separate; inner
and outer pairs of dentate projections on posterior margin
of equal length.

MALE GENITALIA. Genital plates as in G. arenarius .
Aedeagus with one pair of short, lateroventrally projecting
apical processes which point anteriorly in lateral view
(fig. 21) . Style tips slender and elongate.

HOLOTYPE. Male, 8 km. SW. Cave Junction, Oregon,
15 June 19 72, Oman.

PARATYPES. 11 females, 6 males, 15 June 19 72, Oman;
1 female, 3 males, 13 May 19 72, Sawbridge.

DISTRIBUTION. CALIFORNIA: Del Norte County: 3.2 km.
E. Gasquet. OREGON: Josephine County: 8 km. SW. Cave
Junction (May - July) .

HABITAT AND HOST. In chaparral, host unknown.

Giprus joaquinus (Oman) .

Laeviaephalus joaquinus Oman 1937, p. 478.

LENGTH. Female: 4.12 mm.; male: 3.95 mm.

COLOR. Light brown.

HEAD SHAPE. W/L = 1.23.

FEMALE 7TH STERNUM. Elongate medially, about three
times the length at the lateral margin. Dorsal extensions
fused (fig. 8) .

MALE GENITALIA. Genital plates as in G. siskiyou.
Aedeagus long and slightly decurved with two pairs of small
apical processes, one pair dorsal and the other ventral.
Style tips broad and flat (fig. 23).

HOLOTYPE. Male, Califa, California, 12 June 1935, Oman.

DISTRIBUTION. CALIFORNIA: Madera County: Califa
(June) •

No. 114] SAWBRIDGE: GIPRUS 11

HABITAT AND HOST. Unknown.
Giprus deourvatus Sawbridge, new species.

LENGTH. Female: 4.82 nun.; male: 4.08 mm. - 4.36 mm.

COLOR. Light brown.

HEAD SHAPE. W/L = 1.25.

FEMALE 7TH STERNUM. No record; only female has abdomen
missing.

MALE GENITALIA. Genital plates as in G. desertorum new
species. Aedeagus strongly decurved with two pairs of
short, narrow, apical processes, one pair originating
dorsally and the other laterally. Styles with blunt tips
(fig. 24).

HOLOTYPE. Male, Grass Valley, California, 18 May 1930,
Van Duzee.

PARATYPES. 1 female, 1 male, 18 May 19 30, Van Duzee.

DISTRIBUTION. CALIFORNIA: Nevada County: Grass
Valley; Placer County: Dutch Flat (May - June).

HABITAT AND HOST. Unknown.

Giprus angelus (Oman).

Laevicephalus angelus Oman 1937, p. 478.

LENGTH. Female: 3.59 mm. - 4.04 mm.; male: 3.79 mm. -
4. 05 nrni.

COLOR. Light brown.

HEAD SHAPE. W/L = 1.26.

FEMALE 7TH STERNUM. Dorsal extension fused; inner pair
of dentate projections on posterior margin shorter than
outer pair.

MALE GENITALIA. Genital plates as in G. siskiyou.
Aedeagus with two pairs of apical processes, one pair dorsal
and the other pair lateroventral . Style tips blunt (fig.
25) .

HOLOTYPE. Male, above Mint Canyon, California, 8 June
1935, Oman.

DISTRIBUTION. CALIFORNIA: Los Angeles County: Mint
Canyon; Monterey County: Salinas (June).

HABITAT AND HOST. Unknown.

Giprus crassus Sawbridge, new species.

LENGTH. Female: 4.00 mm. - 4.2 3 ma.; male: 3.77 mm. -
4. 07 mm.

COLOR. Medium brown.

HEAD SHAPE. W/L = 1.23.

FEMALE 7TH STERNUM. Dorsal extension fused; inner pair
of dentate projections on posterior margin much shorter
than outer pair.

MALE GENITALIA. Genital plates as in G. desevtorum
except apices are subacute. Aedeagus short, stout, broadest
at apex, and with three pairs of short apical processes:
one pair lateroventral, one pair lateral, and one pair

1

'■
12 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers !

dorsal. Styles short and broad with blunt tips (fig. 26).

HOLOTYPE. Male, Sams Valley, Oregon, 15 May 19 70, Oman.

PARATYPES. 14 females, 16 males, 15 May 19 70, Oman;
3 females, 12 males, 29 May 19 71, Oman.

DISTRIBUTION. OREGON: Jackson County: Sams Valley
(May - June) .

HABITAT AND HOSTS. In chaparral on Stipa lemmonii , Poa
scabre lla.

Giprus ovavius Sawbridge, new species.

LENGTH: Female: 3.66 mm. - 3.96 mm.; male: 3.84 mm. -
3.96 mm.

COLOR. Very pale, little pigment deposited around
periphery of wing cells.

HEAD SHAPE. W/L = 1.46.

FEMALE 7TH STERNUM. Dorsal extensions separate; inner
pair of dentate projections on posterior margin shorter than
outer pair.

MALE GENITALIA. Genital plates as in G. siskiyou.
Aedeagus with a bulbous apex produced into a pair of j

lateroventral processes. A pair of thin laterodorsally j
projecting processes originate basad of the bulbous apex. I
Style tips broad (fig. 27) . '

HOLOTYPE. Male, 16.1 km. (10 mi.) N. Pacific City, :l
Oregon, 2 0 July 19 71, Sawbridge. \\

PARATYPES. 18 females, 13 males, 20 July 1971, Saw- |
bridge; 47 females, 22 males, 20 July 1971, Viraktamath. li

DISTRIBUTION. OREGON: Coos County: 17.7 km. S. j

Reedsport; Douglas County: 17.7 km. S. Florence; Lincoln
County: 1.6 km. S. Newport; Tillamook County: Nehalem j
Bay State Park, 16.1 km. N. Pacific City, 1.6 km. N. Pacific
City (July - August) .

HABITAT AND HOSTS. On sand dunes on Poa maorantha and
Festuca rubra.

Giprus agrestis Sawbridge, new species.

LENGTH. Female: 3.91 mm. - 4.41 mm.; male: 3.71 mm. -
4. 07 mm.

COLOR. Dark brown.

HEAD SHAPE. W/L = 1.20.

FEMALE 7TH STERNUM. Dorsal extension fused; inner pair
of dentate projections on posterior margin almost as long
as outer pair (fig. 7) .

MALE GENITALIA. Genital plates as in G. siskiyou.
Aedeagus short with three pairs of apical processes: one
pair dorsal, one pair lateral, and one pair ventral.
Aedeagus broadly rounded apically. Styles with small flared
tips (fig. 28).

HOLOTYPE. Male, Merlin, Oregon, 27 May 1970, Oman.

PARATYPES. 16 females, 15 males, 27 May 19 70, Oman;
13 females, 10 males, 13 May 1972, Sawbridge.

DISTRIBUTION. OREGON: Josephine County: Merlin (May -
June) .

No. 114] SAWBRIDGE: GIPRUS 13

HABITAT AND HOST. In chaparral on Danthonia oatifornioa
var. amerioana.

Giprus oinerosus (Van Duzee) .

Dettocephalus oinerosus Van Duzee 1892, p. 305.

Laevicephalus oinerosus (Van Duzee) ; DeLong and Sleesman
1929, p. 93.

Giprus oinerosus Oman 1949, p. 164.

LENGTH. Female: 4.30 mm. - 4. 31 nun. ; male: 3.91 mm. -
4. 17 ram.

COLOR. Light brown.

HEAD SHAPE. W/L = 1.27.

FEMALE 7TH STERNUM. Lateral margins short; dorsal
extensions separate; inner pair of dentate projections on
posterior margin slightly shorter than outer pair.

MALE GENITALIA. Genital plates as in G. desertorum.
Aedeagus with three pairs of apical processes all origi-
nating at the same point on the shaft: a short ventral
pair, an elongate lateral pair, and a short dorsal pair
(fig. 35). Style tips broad.

LECTOTYPE. Male, California, Coquillett.

DISTRIBUTION. CALIFORNIA: Los Angeles County; Orange
County: Newport Beach; Riverside County: Perris; San Diego
County: El Cajon, Pala, Witch Creek (February - May).

HABITAT AND HOST. Unknown.

Giprus desertorum Sawbridge, new species.

LENGTH. Female: 3.59 mm. - 3.88 mm.; male: 3.29 mm. -

3. 50 mm.

COLOR. Light brown.

HEAD SHAPE. W/L = 1.28.

FEMALE 7TH STERNUM. Dorsal extensions separate; inner
pair of dentate projections on the posterior margin longer
than outer pair.

MALE GENITALIA. Genital plates broadly rounded,
anteriorly contiguous, but diverging posteriorly and
abruptly narrowing to acute apices (fig. 11) . Aedeagus with
three pairs of processes: a short ventral pair, an elongate
lateral pair arising basad of the ventral pair, and a very
short dorsal pair. Style tips broad (fig. 34) .

HOLOTYPE. Male, 4.8 km. (3 mi.) N. Rosamond, Cali-
fornia, 2 3 March 19 71, Sawbridge.

PARATYPES. 10 females, 20 males, 2 3 March 1971, Saw-
bridge; 8 females, 14 males, 23 March 1971, Oman.

DISTRIBUTION. CALIFORNIA: Los Angeles County: 4.8 km.
S. Lancaster, 4.8 km. N. Rosamond (March - April).

HABITAT AND HOSTS. Amongst desert scrub on Stipa
speoiosa and Orysopsis species.

Giprus oalifornicus Sawbridge, new species.

LENGTH. Female: 3.95 mm. - 4.21 mm.; male: 3.65 mm. -

4. 20 mm.

14 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

COLOR. Medium brown.

HEAD SHAPE. W/L = 1.28.

FEMALE 7TH STERNUM. Dorsal extensions separate; dentate
projections on posterior margin of equal length.

MALE GENITALIA. Genital plates as in G. siskiyou,
Aedeagus with a pair of straight or sinuate laterally pro-
jecting processes from which arise a pair of ventral
processes which curve toward the midline (figs. 30, 31).
Curvature of lateral processes varies between populations.

HOLOTYPE. Male, 43.5 km. (27 mi.) SE. Bakersfield,
California, 23 March 1971, Sawbridge.

PARATYPES. 9 females, 12 males, 2 3 March 19 71, Saw-
bridge; 5 females, 5 males, 22 April 1971, Oman.

DISTRIBUTION. CALIFORNIA: Kern County: 43.5 km. SE.
Bakersfield, Tehachapi Pass 579 m. and 823 m. ; Marin County:
Mill Valley; Napa County: 3.2 km. N. St. Helena, Bothe-
Napa State Park, 4.2 km. W. Oakville; Santa Cruz County:
Highland District (March - June) .

HABITAT AND HOST. In chaparral, host unknown.

Giprus vastitatus Sawbridge, new species.

LENGTH. Female: 3.73 mm. - 4.12 mm.; male: 3.51 mm. -
4. 13 mm.

COLOR. Medium brown.

HEAD SHAPE. W/L =1.29. 1

FEMALE 7TH STERNUM. Dorsal extensions separate; inner 1
pair of dentate projections on posterior margin shorter \
than outer pair.

MALE GENITALIA. Genital plates as in G. siskiyou.
Aedeagus short and slightly sinuate with one pair of sinuate
processes originating lateroventrally and extending basally
approximately 3/4 the length of the aedeagal shaft (fig.
29) .

HOLOTYPE. Male, 16 km. (10 mi.) N. Winnemucca, Nevada,
2 3 June 19 71, Sawbridge.

PARATYPES. 2 females, 1 male, 6.4 km. (4 mi.) NW.
Winnemucca, Nevada, 2 3 June 19 71, Oman.

DISTRIBUTION. NEVADA: Humboldt County: Winnemucca,
6.4 km. NW. Winnemucca, 16 km. N. Winnemucca. UTAH:
Washington County: St. George. March (Utah) - June
(Nevada) .

HABITAT AND HOST. In desert scrub, host unknown.

Giprus oajon Sawbridge, new species.

LENGTH. Female: 3.79 mm. - 4.03 mm.; male: 3.53 mm. -
3.93 mm.

COLOR. Medium brown.

HEAD SHAPE. W/L = 1.26.

FEMALE 7TH STERNUM. Dorsal extensions separate; inner
pair of dentate projections on posterior margin shorter
than outer pair.

MALE GENITALIA. Genital plates as in G. desevtovum.
Aedeagus with somewhat elongate dorsal processes arising

No. 114] SAWBRIDGE: GIPRUS 15

more than 1/3 of the distance from the apex to the base and
with lateroventral processes arising near the apex.
Aedeagal shaft somewhat stout. Style tips broad (fig. 33) .
Considerable variation.

HOLOTYPE. Male, 1.6 km. (1 mi.) NW. top of Cajon Pass,
California, 23 April 1971, Oman.

PARATYPES. 11 females, 5 males, 23 April 1971, Oman.

DISTRIBUTION. CALIFORNIA: San Bernardino County: 1.6
km. NW. top of Cajon Pass (April) .

HABITAT AND HOST. In chaparral on Stipa speciosa.

Giprus paoificus (Oman) .

Laevieephalus paoificus Oman 1937, p. 477.

LENGTH. Female: 3.2 3 mm. - 3.26 mm.; male: 2.81 mm. -
3. 01 mm.

COLOR. Pale to light brown.

HEAD SHAPE. W/L = 1.33.

FEMALE 7TH STERNUM. Dorsal extensions separate; inner
pair of dentate projections on posterior margin as long as
outer pair; median incision short.

MALE GENITALIA. Genital plates as in G. siskiyou,
Aedeagus sinuate with two pairs of processes: a somewhat
elongate lateral pair arising at a point about 1/3 of the
distance from the apex of the aedeagus to the base and a
short ventral pair arising near the apex. Style tips broad
(fig. 32).

HOLOTYPE. Male, Montara, California, 13 June 1935,
Oman.

DISTRIBUTION. CALIFORNIA: Alameda County: Bay Farm;
San Mateo County: Montara, Pillar Point, (?) Half Moon
Bay (May - July) .

HABITAT AND HOST. On coastal bluffs, host unknown.

Infraspecific Variation

Inf raspecif ic variation in Giprus is evident in the
overall size, pattern and degree of pigmentation of the
head and wings, shape and pigmentation of the female
seventh sternum, wing venation, and structure of the aede-
agus. The latter is the most taxonomically significant
type of variation as species can be readily distinguished
only on the basis of aedeagal structure. Aedeagal vari-
ation is evident in five species: G. oajon, G. paoificus ,
G. catifornicus , G. vastitatus , and G. siskiyou.

The most obvious and frequently encountered infra-
specific variation is found in G. siskiyou. While the
dorsal processes and apical notch are more or less constant
in shape and size, noticeable variation occurs in the
degree of development of the lateral flanges. In the
original description of the species Oman (1937) made no
reference to the presence of lateral flanges, and the
individual illustrated in his paper was probably an extreme

16 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

case showing little or no flange development.

While most specimens from northern California and
southern Oregon and the single specimen from Carson City,
Nevada, have an aedeagus with moderately developed lateral
flanges (fig. 17), specimens from localities in Washington,
northeastern California, and Steens Mountain, Oregon, show
much greater development of the lateral flanges (figs. 18-
20) .

Two samples of populations, one from south of Weed,
California, in the north-south transect and the other from
58 km. east of Klamath Falls, Oregon, in the east-west
transect were chosen for dissection to determine the extent
of aedeagal variation present at any one time in the core
area of G. siskiyou distribution. Flange development in
these samples ranges from slight lateral protrusions which
merely suggest the presence of flanges (figs. 52, 53, 72)
to well developed structures (figs. 37, 54, 55). Figure 54
illustrates extreme development of the lateral flanges.
The lateral flanges exhibit greater asymmetry than is
usually found among aedeagal structures of Giprus , an
extreme condition being that illustrated in figure 49.
Variation also occurs in the position of the lateral flanges
on the shaft relative to the dorsal processes. Figures 37,
38, and 47 show specimens which have the lateral flanges
projecting only slightly basad of the dorsal processes
while figures 39 and 55 show individuals with lateral
flanges extended considerably basad of the dorsal processes.

The variation shown in these samples is highly signif-
icant because it indicates that structures such as those
seen in specimens from southcentral Washington, Steens
Mountain, Oregon, and northeastern California fall within
the demonstrable range of variation of G. siskiyou. A
single specimen (fig. 54) from 58 km. east of Klamath Falls,
Oregon, shows greater development of the lateral flanges
than is shown in any of the individuals from isolated or
peripheral areas.

Although the aedeagal structure of only five G. cajon
specimens was examined, considerable infraspecific vari-
ation was noted. One specimen had two very small dorsal
processes originating at the bases of the dorsal elongate
processes. Another was markedly asymmetrical with the
processes on the right arising from the aedeagal shaft basad
of those on the left. The angle between the elongate dorsal
processes and the shaft is also variable. Variation in the
angle formed by the lateral processes and the aedeagal
shaft occurs in G. pacificus.

Structural variation between populations of the same
species also occurs. Specimens of G. calif ovnicus from
Tehachapi Pass (fig. 30) , Napa Valley, and the Highland
District of Santa Cruz County, California, have straight
elongate processes and ventral processes with acute apices.
Specimens of G. oatifornicus from Mill Valley, California,
(fig. 31) have distinctly sinuate elongate processes and
ventral processes with acute apices. Minor differences
also occur between specimens of G. vastitatus collected in

No. 114] SAWBRIDGE: GIPRUS 17

southern Utah and northern Nevada. Specimens from inter-
vening localities may show such differences to be part of
the normal range of variation in this species.

Habitat

Species of Giprus have been found in five types of
habitats: chaparral, desert scrub, marine sand dunes,
coastal bluffs, and grassland. While all the members of
the genus are found on grasses, Giprus does not exhibit any
obvious host specificity within that plant group. Grasses
from which Giprus species have been taken are not confined
to any group of grasses recognized in Gould's (1968)
classification of genera. In several instances a species
of Giprus has been collected from more than one genus of
grass and a single species of grass has been observed to.
host more than one species of Giprus, although only at
different localities.

Chaparral habitats (Cooper, 1922; Detling, 1961) are
occupied by G. hidentatus , G. crassus , G. agrestis , G.
californicus , G. siskiyou , and G. oajon. All but the latter
inhabit the type of chaparral described by Detling (1961)
as northern California - southern Oregon chaparral, domi-
nated by Ceanothus ouneatus . Other common plants include
Arotostaphylos visoida , A. oanescens , Querous species.
Arbutus menziesii , and Purshia tridentata. Grass hosts
determined included Stipa Zemmonii , Poa soabrella , and
Danthonia oalifornica var. americana. Giprus oajon occurs
in a southern California chaparral at the transition between
Yuoea brevi folia and Juniperus species zones. Specimens
were collected from Stipa speeiosa amongst Eriogonum
fasciculatum and Salvia dorii subspecies carnosa.

Desert scrub habitats are occupied by G. siskiyou , G.
desertorum , and G. vastitatus . East of the Cascade Range
G. siskiyou inhabits an Artemisia tridentata dominated
shrub-steppe as described by Franklin and Dyrness (1969)
and is found on Stipa lemmonii and Poa species. Giprus
vastitatus is found in the even drier sagebrush - grass zone
of the Great Basin described by Billings (1951) . Giprus
desertorum lives on Stipa speeiosa and Oryzopsis species in
the Mojave Desert amongst Yucca brevifolia, Grayia spinosa ,
Atriplex species, Tetradymia canes cens , Coreopsis species,
and Larrea divaricata.

Giprus arenarius and G. orarius are found on both
Festuca rubra and Poa macrantha on the sand dunes of the
Oregon Coast. The part of the dunes on which these grasses
and leafhoppers occur is still very unstable with few if
any shrubs present. Other plants which may occur along
with the host grasses are Lathyrus species, Tanecetum
camphoratum , Lupinus species, and Glehnia leiocarpa. Giprus
arenarius and G. orarius are almost always found on the lee
side of foredunes or on the slopes of higher dunes farther
inland; they have never been found sympatrically .

18

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Giprus paoificus has been collected from bluffs at
Pillar Point, California, a habitat with considerably more
vegetation than the Oregon dunes. However, female speci-
mens which may also be G. paoificus were taken from grasses
on unstable dunes about a mile south of the bluff site.

Giprus oavtwrighti is typically found in flat, shrub-
less, open areas with mixed short grasses and f orbs .
Occasionally it is found in the chaparral association.

Literature Cited

BILLINGS,
1951.

COOPER, W.
1922.

DELONG, D,
1926.

DELONG, D.
1929.

DETLING,
1961

FRANKLIN,
1969.

GOULD, F.
1968.

OMAN , P .
1932

W. D.

Vegetational zonation in the Great Basin of
North America. In'. Les bases ecologiques de
la regeneration de la vegetation des zones
arides. International Union of Biological
Sciences Serie B, Colloquia, no. 9, 149 pp.

S.

The broad-sclerophyll vegetation of California.
Carnegie Institution of Washington Publi-
cations, no. 319, pp. 1-124.

M.

A monographic study of the North American

species of the genus Deltooephalus . The Ohio
State University Studies, vol. 2, no. 13,
Contributions in Zoology and Entomology no. 3,
129 pp.

M. , AND J. P. SLEESMAN.

New genera and subgenera from the genus Delto-
oephalus. A study of the internal male
genitalia of the American species and their
bearing upon taxonomy. Annals of the Entomo-
logical Society of America, vol. 22, pp. 81-114,
E.

The chaparral formation of southwestern Oregon
with considerations of its postglacial history.
Ecology, vol. 42, pp. 348-357.

1

F . , AND C ,

DYRNESS

Vegetation of Oregon and Washington. Portland.
216 pp. (USDA Forest Service Research Paper
PNW-80) .

W.

Grass systematics
382 pp.

McGraw-Hill, New York, xi +

W.

1937

1949

Two new species of Laevicephalus from California.
Proceedings of the Entomological Society of
Washington, vol. 34, no. 6, pp. 90-92.

The cinerosus group of the genus Laevicephalus
(Homoptera: Cicadellidae) . Journal of the
Washington Academy of Sciences, vol. 27,
no. 11, pp. 474-479.

The Nearctic leafhoppers, a generic classifi-
cation and checklist. Memoirs of the

No. 114]

SAWBRIDGE:

GIPRUS

19

Entomological Society of Washington, no,
25 3 pp.

3,

READIO, P
1922.

ROSS, H. H
1972.

VAN DUZEE,
1892.

A.

Ovipositors of Cicadellidae. The Kansas Univer-
sity Science Bulletin, vol. 14, no. 8, pp. 217-
298.

, AND K. G. A. HAMILTON.

A review of the North American leafhopper genus
Laevicephalus (Hemiptera: Cicadellidae).
Annals of the Entomological Society of America,
vol. 65, no. 4, pp. 929-942.

E. P.

A synoptical arrangement of the genera of the
North American Jassidae, with descriptions of
some new species. American Entomological
Society Transactions, vol. 19, pp. 295-307.

Group IV

/ Group III

B'n? aO'

' SH a^\

\

1

I

/

^ilftss^ ^^J

FIGURE 1. Species groups of Gipvus based on the struc-
ture of the aedeagus. Group I: A, G. oartwrighti . Group
II: B, G. arenavius , C, G. bidentatus , D, G. siskiyou , E,
G. siskiyou , F, G. joaquinus , G, G. decurvatus . Group III:
H, G. angelus , I, G. arassus , J, G. orarius , K, G. agvestis .
Group IV: L, G. vastitatus , M, G. aalifornious , N, G. cali-
forniaus , 0, G. aajon, P, G. paaificus , Q, G. oinerosus , R,
G. desertorum .

2 0 CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

Mean ^L
m -Range

G. orarius

G. arenar i us
G. pacrf icus

G. vastitatus
G. desertorum
■^^BB G. calKornicus

G. bidentatus
G. cinerosus
G. Siskiyou
^^m G.ongelus

^^m G. cajon

G. decurvatus
G. joaquinus

1^ G. crassus

G. agresf is
G. cartwr ight i

1.12 1.16 1.20 1.24 1.28 1.32 1.36 1.40 1.44 1.48 1.52

FIGURE 2. Ratio of head width (W) to head length (L)
for all species of Giprus .

No. 114]

SAWBRIDGE: GIPRUS

21

FIGURES 3-8. FIGURE 3. Giprus siskiyou: forewing;
FIGURE 4. G, agrestis: fifth instar nymph; FIGURE 5. G.
siskiyou: head; FIGURE 6. G. cajon: female 7th sternum;
FIGURE 7. G. agrestis: female 7th sternum; FIGURE 8: G.
joaquinus : female 7th sternum.

22

CALIFORNIA ACADEMY OF SCIENCES

[Occ. Papers

FIGURES 9-17. FIGURE 9. Giprus cartwrighti: female
7th sternum; FIGURE 10. G. siskiyou: male genital seg-
ments; FIGURE 11. G. desertorum: male genital segments;
FIGURE 12. G. avenarius: male genital segments; FIGURE
13. G. cartwrighti : male genital segments; FIGURE 14. G.
cinerosus : connective and aedeagus ; FIGURE 15. Laevi-
cephalus sylvestris (Osborn & Ball): connective and aedea-
gus; FIGURE 16. Laevioephalus incongruus : connective,
style, and aedeagus; FIGURE 17. G. siskiyou (Mt. Shasta,
California): connective, style, and aedeagus.

J

No. 114]

SAWBRIDGE : GIPRUS

23

FIGURES
FIGURE 18.
FIGURE 19.
G. Siskiyou

hidentatus ;
quinus .

18-23. Connective, style, and aedeagus of:
Giprus siskiyou (24 km. NE. Alturas, California)
G. siskiyou (Cliffdell, Washington); FIGURE 20.
(18 km. E. Frenchglen, Oregon) ; FIGURE 21. G.
FIGURE 22. G. arenarius ; FIGURE 23. G. joa-

24

CALIFORNIA ACADEMY OF SCIENCES

[Occ. Papers

t^

26

FIGURES 2 4-29. Connective, style, and aedeagus of:
FIGURE 24. Giprus deourvatus ; FIGURE 25. G. angelus;
FIGURE 26. G. arassusj FIGURE 27. G. ovavius ', FIGURE 28,
G. agrestis ; FIGURE 29. G. vastitatus .

No. 114]

SAWBRIDGE: GIPRUS

25

FIGURES 30-36. Connective, style, and aedeagus of:
FIGURE 30. Giprus californicus (Tehachapi Pass, California);
FIGURE 31. G. californious (Mill Valley, California);
FIGURE 32. G. pacifiaus) FIGURE 33. G. aajonj FIGURE 34.
G. desertorum; FIGURE 35. G. ainerosus ; FIGURE 36. G.
cartwrighti .

26

CALIFORNIA ACADEMY OF SCIENCES [Occ. Papers

nJl M

37

42

47

U

52

l5

38

43

0«

48

53

39

iUi M

U

44
49

40

45

0"

50

M^l

?

41

46

//

51

u

?i

54

U "

59
64

1;^*'

55

60
65

56

i'Wn

U

61

^i

66

[il

\J

0

57

U

62

67

58

63

^ V A)

1/

68

69

70

71

72

No. 114] SAWBRIDGE: GIPRUS 27

^ FIGURES 37-72. FIGURES 37-53. Giprus siskiyou (5 km.
S. Weed, California): apex of aedeagus; FIGURES 54-72. G.
siskiyou (58 km. E. Klamath Falls, Oregon): apex of
aedeagus.

I

Mlii " II' 'I 11

1 I l\ \ IV I

UH nCl X