HARVARD UNIVERSITY

SO.

LIBRARY

OF THE

Museum of Comparative Zoology

The Great Basin Naturalist

VOLUME XXIX, 1969

Editor: Vasco M. Tanner
Associate Editor: Stephen L. Wood

Published at Brigham Young University, by
Brigham Young University

Volume XXIX December 31, 1969 Nos. 1-4

TABLE OF CONTENTS
Volume XXIX

Number 1 — March 28, 1969

Undescribed Species of Nearctic Tipulidae (Diptera), X.

Charles P. Alexander 1

The Buprestidae and Cleridae of the Nevada Test Site

(Coleoptera). Illustrated. William F. Barr 11

Bees of the Nevada Test Site. Dorald M. Allred 20

A Note on Geranium richardsonii Fisch. & Trautv. Illus-
trated. John W. Van Cott 25

New Combinations and a New Species of Perennial Cryp-

tantha. Illustrated. Larry C. Higgins 28

New Records and Distributional Notes for Reptiles of the

Nevada Test Site. Wilmer W. Tanner 31

The Efficacy of Haloxon and Thiabendazole as Anthel-
mintics Against Gastro-intestinal Nematodes in Sheep.
Ferron L. Andersen, Keith H. Hoopes, and J. Carl Fox 35

NOTE: Lepidoptera of the Nevada Test Site. Dorald M.

Allred 42

NOTE: Wislouchiella planctonica Skvortz (Chlorophyta.
Volvocales), A New Algal Record for Nevada. James
L. Reveal 43

NOTE: The Infection of Immature Aquatic Insects by

Larval Paragordius (Nematomorpha). David A. White .... 44

Number 2 — July 31, 1969

The Ecology of the Western Spotted Frog, Rana pretiosa
pretiosa Baird and Girard, a Life History Study. Illus-
trated. Ronald L. Morris and Wilmer W. Tanner 45

Fossil Chelonians. Chrysemys and Clemmys, from the

upper Pliocene of Idaho. G. R. Zug 82

Notes on the Distribution and Bionomics of Some Idaho
Cerambycidae (Coleoptera). William F. Barr and
Richard L. Penrose 88

Overwintering Phytoseiid Mites in Central Utah Apple
Orchards. Illustrated. John W. Leetham and Clive D.
Jorgensen 96

Spiders of the National Reactor Testing Station. Dorald M.

Allred 105

NOTE: Individual Rodent Movements in Upper Sonoran

Desert of Central Nevada. Jon Ghisehn 109

ii

NOTE: An Albino Pacific Tree Frog, Hyla regilla, from

Death Valley, California. Fenton R. Kay Ill

Number 3 — September 30, 1969

New Synonymy and Records of Platypodidae and Scoly-

tidae (Coleoptera). Stephen L. Wood 113

Zoogeographic and Systematic Notes on the Lace Bug Tribe

Litadeini, with the Description of the New Genus

Stragulotingis (Hemiptera: Tingidae). Illustrated.

Richard C. Froeschner 129

A New Species of Oppia from Colorado (Cryptostigmata;

Oppiidae). Illustrated. Tyler A. Woolley 133

A New Genus and Species of Oribatid Mite (Acari: Lia-

caroidea, Metrioppiidae) . Illustrated. Tyler A. Woolley .... 137
A New^ Species of Linear us from a Wood Rat Nest (Aca-

rina: Cryptostigmata, Liacaridae). Illustrated. Harold

G. Higgins and Tyler A. Woolley 140

Invasion of a "Trapped-out" Southern Nevada Habitat by

Perognathus longimembris . Lester D. Flake and Clive

D. Jorgensen 143

List of the Insect Type Specimens in the Entomological

Collection of the Brigham Young University, Provo.

Utah, No. III. Vasco M. Tanner and Don R. Harris 150

Studies on Raptor Mortality in Western Utah. David H.

Ellis, Dwight G. Smith, and Joseph R. Murphy 165

Number 4 — December 31, 1969

Qualitative Behavior of a Pupfish {Cyprinodon atrorus)
in Differing Environments. Illustrated. Murray Itz-
kowitz and W. L. Minckley 169

Does Vireo gilvus swainsonii Occur in Utah? Gary L.

Worthen 181

List of Insect Type Specimens in the Entomological Collec-
tion of the Brigham Young University, Provo, Utah,
No. IV. Vasco M. Tanner and Don R. Harris 183

Index 206

lU

5- Qd-j^^'^

Volume XXIX, No. 1
March 28, 1969

The

5-171

Great Basin

iiirruiJiiiiST

Published by
Brigham Young University

GREAT BASIN NATURALIST

Editor: Vasco M. Tanner, Department of Zoology and Entomology
Brigham Young University, Provo, Utah

Associate Editor: Stephen L. Wood, Department of Zoology and
Entomology, Brigham Young University, Provo, Utah

Members of the Editorial Board:

Ferron L. Andersen (5), Zoology

Jay V. Beck (3), Bacteriology

Robert W. Gardner (1), Animal Science

Joseph R. Murdock (4), Botany

WiLMER W. Tanner (2), Zoology, Chairman of the Board

Stanley L. Welsh ( 1 ) , Botany

Ex officio Members:

A. Lester Allen, Acting Dean, College of Biological and
Agricultural Sciences

Ernest L, Olson, Chairman, University Publications, Uni-
versity Editor

The Great Basin Naturalist

Published at Provo, Utah by
Brigham Young University

Volume XXIX March 28, 1969 No. 1

UNDESCRIBED SPECIES OF NEARCTIC TIPULIDAE
(DIPTERA), X

Charles P. Alexander'

At this time I am describing various new species of crane flies
from western North America, including Alaska and British Colum-
bia, collected by Dr. Marion E. Smith and David L. Carson; Nevada
by Dr. Donald (j. Denning; and Utah, by Dr. George W. Knowlton.
Dr. C. Dennis Hynes, and Dr. Donald W. Davis. 1 am greatly
indebted to the persons listed for permission to retain the types of
the novelties in my personal collection of World Tij)ulidae.

Genus Tipula Linnaeus
Subgenus Arctotipula Alexander

Tipula {Arctotipula) Alexander; Philippine Jour. Sci., 52:410; 1933.

Several species of Arctotipula now are known from Alaska and
adjoining territories and a brief discussion of certain of the previously
described forms seems to be advisable. The references cited are listed
at the conclusion of the paper.

Lackschewitz (1936:290) has placed Tipula {Arctotipula) aleu-
tica Alexander (1923) in the synonymy of T. {A.) hesselsi Osten
Sacken, certainly in error. The outer wing veins of aleutica have
abundant macrotrichia while the hypopygial structure is quite
distinct from that of besselsi and certain other allied species. In
the same paper, Lackschewitz (1936:288) places T.{A) alascaensis
Alexander (1923), described from the Pribilof islands, in the sy-
nonymy of ciliata Lundstrom (1915), a course that was followed by
Savchenko (1961:325). It may be affirmed that the two flies are
quite distinct in hypo{)ygial characters, very definitely in the struc-
ture of the inner dististyle. In alascaensis the beak of the latter is
short and obtusely rounded to appear almost suboval in outline, ex-
tending only a short distance beyond the very stout outer spine of the
style. T. {A.) 0klandi Alexander (1921), described from Nova
Zembla. likewise was considered by both Lackschewitz and Sav-
chenko as being identical with ciliata but paratype specimens in my

'Amherst, Massachusetts.

The Great Basin Naturalist
2 CHARLES P. ALEXANDER Vol. XXIX. No. 1

collection have the conformation of the inner dististyle different from
that shown by Savchenko (1961 :326, fig. 3) for ciliata, including the
beak and the outer spine.

Tipula (Arctotipula) denali, n.sp.

Size medium (wing of male 14.5 mm.); flagellar verticils short;
general coloration of body dark gray, vestiture very reduced to
nearly absent; wings yellowish white, restrictedly patterned with
brown; Rs long, about two and one-half times m-cu; male hypopygi-
um with posterior border of tergite triemarginate, the intermediate
lobes acute, their mesal borders microscopically toothed; outer disti-
style small, vestiture short.

Male. — Length about 16 mm.; wing 14.5 mm.; antenna about
4 mm.

Frontal prolongation of head about three-fourths the remain-
der, light gray above, more plumbeous on sides, nasus stout; vestiture
reduced to very sparse short stout black setae on outer half of pro-
longation; palpi black. Antennae black; flagellar segments without
basal enlargements, longer than the verticils, beyond the first with
dense short yellow setulae. Head uniformly gray with a vague
more darkened line on posterior vertex; vertical tubercle low. de-
pressed medially; posterior half of head with sparse short black
setae, longer and porrect on genae and beneath, very reduced in
number and size on vertex, lacking on the midportion.

Cervical region and pronotum blackened, heavily gray pruinose.
Mesonotal praescutum light gray with four scarcely indicated darker
gray stripes that are very narrowly bordered by darker; scutum gray,
each lobe with two confluent darker areas, the anterior one smaller;
posterior sclerites of notum light gray, parascutella pale; notum
virtually glabrous, praescutal interspaces with microscopic punctures
that bear tiny yellow setulae. Pleura light gray, vaguely patterned
with darker gray areas; dorsopleural region buffy yellow. Halteres
brown. Legs with coxae light gray, vestiture reduced to sparse
microscopic yellow setulae; trochanters dark gray; femora and tibiae
reddish brown, tips darker; tarsi dark brown; claws of male with a
single slender tooth. Wings yellowish white, with an extensive pale
brown clouded pattern that is subequal in extent to the ground,
stigma darker brown, oval; more extensive brown clouds include a
continuous seam along Rs. two large areas in cell M, most cells be-
yond cord, and outer ends of cubital and anal cells; the major ground
areas include the prearcular and costal fields, cell R and inter-spaces
of cells M and Cu, and the broad central part of cell 1st A; outer
radial field chiefly of the ground, including cell Sc-., most of /?.., and
a subbasal brightening in cell /?3; veins brown, obliterative sections
very extensive, especially in the veins that enclose cell 1st Mo. Veins
posterior to R glabrous. Venation: Rs very long, about two and one-
half times m-cu; petiole of cell M] shorter than m.

Abdomen dark plumbeous gray, basal tergite paler gray; poster-
ior borders of both tergites and sternites very narrowly obscure yel-

March 28, 1969 species of nearctic tiphlidaf, 3

low. Male hypopygiuni with tergite relatively nan'ow. transverse,
posterior border with three U-shaped emarginations to produce two
narrow acute intermediate lobes and broader lateral ones, the mesal
borders of the intermediate pair microscopically toothed, of the later
al pair with a pale plate; vestiture of tergite relatively abundant but
very short and inconspicuous. Ninth sternite with lobes elongate-
oval, with long yellow setae. Basistyle oval, entire. Outer d'stistyle
small, long-oval, with weak delicate setae; inner style with beak
flattened, pale, tip obtuse; lower beak blackened, obtuse, posterior
crest conspicuous, with darkened parallel lines at base; region of
outer basal lobe a small plate that narrows into an acute black spine.
Aedeagus long and very stout. Eighth sternite transverse, posterior
border convexly rounded, medially with a triangular membranous
area; setae very small and sparse, with a group of longer and more
conspicuous yellow setae adjoining the membranous area.

Habitat. — Alaska.

HoLOTYPE, Muldrow Glacier, McKinley National Park, July 19,
1952 (D. L. Carson).

The specific name 'denali is that of the native name for Mount
McKinley, Denali, 'Home of the Sun'. I am referring this fly to the
subgenus Arctotipula despite the differences in hypopygial structure
from the subgenotype and allied species. The virtually glabrous body
similarly is discordant in this subgenus. The wing pattern is ap-
proached by that of Tipula {Arctotipula) twogwoteeana Alexander
of the higher Rocky Mountains of Colorado and Wyoming which
has the hypopygial structure quite different, being more suggestive
of species in the subgenus Sinotipula Alexander.

Tipula {Arctotipula) epios, n.sp.

Allied to thulensis, differing chiefly in hypopygial characters,
especially the acutely pointed lobes of the ninth tergite and the
broad outer dististyle.

Male. — Length about 14 mm.; wdng 15 mm.; antenna about 4
mm.

Frontal prolongation of head relatively short, about one-half the
remainder, light gray; dorsal surface with long black setae, nasus
long and stout; palpi black. Antenna black throughout; flagellar
segment subequal to their longest verticils, basal enlargements poorly
to scarcely developed. Head gray, front and orbits light gray; vertex
with abundant long erect black setae, lacking on midarea; eyes rela-
tively small, anterior vertex correspondingly broad.

Pronotal scutum gray, with long erect setae, scutellum brownish
gray, the angles more reddened. Mesonotal praescutum light gray
with four darker gray stripes, the intermediate pair narrowly sepa-
rated by a central ground stripe without setae, remainder of ground
areas with abundant long erect setae; scutum light gray, each lobe
with two separate darker gray areas, vestiture short and very sparse;
scutellum dark gray with very long setae, parascutella glabrous;
postnotum uniformly gray, mediotergite with very long setae, pleuro-

The Great Basin Naturalist
4 CHARLES P. ALEXANDER Vol. XXIX. No. 1

tergite glabrous. Pleura gray, doropleural region buffy yellow;
sternopleurite and metapleura with long setae, lacking on meron.
Halteres with stem brownish yellow, knob clearer yellow, base weak-
ly darkened. Legs with coxae gray, setae very long; trochanters
blackened, gray pruinose; femora brownish yellow, tips blackened;
tibiae yellowish brown, darker outwardly; tarsi black; claw of male
with a small triangular tooth at near midlength. Wings subhyaline
to weakly tinted, cell Sc more yellowed; stigma oval, brown; veins
light brown. Veins posterior to R chiefly glabrous, with trichia on
R2+,. distal section of /?,+-, and restrictedly near outer end of 2nd A.
Venation: Petiole of cell M, shorter than w; M^+4 short, about one-
half the basal section of Mi+j or less.

Abdomen dark plumbeous gray, extreme borders of the segments
yellowed; hypopygium blackened. Male hypopygium with tergal
lobes acute, tips pointed; median emargination broadly U-shaped,
its diameter subequal to that of the lateral lobe. Outer dististyle very
broad, the greatest width about three-fourths the length; inner style
with lateral spine blackened, slender, gently curved.

Habitat. — British Columbia.

HoLOTYPE. cf , Alaska Highway. Mile 316, along 107 Creek, June
26, 1952 (M. E. Smith).

Among the numerous members of the subgenus the present fly
is closest to Tipula {Arctotipula) thulensis Alexander, still known
only from Baffin Island, Canadian Northwest Territories. It differs
especially in details of hypopygial structure, as described. None of
the northern Palaearctic members of the subgenus has the tergal
lobes acute as in the present fly.

Tipula {Arctotipula) mckinleyana, n.sp.

General coloration of head and thorax gray, praescutum with four
poorly differentiated darker gray stripes; antenna of male relatively
short; wings faintly darkened, the long-oval stigma solidly darkened;
male hypopygium with beak of inner dististyle relatively slender,
on side before apex with a low blackened flange, margin at base of
beak with a powerful broad-based lobe that narrows into a strong
black spine, margin of style with one or two more basal spinous
points.

Male. — Length about 14 mm.; wing 15.5 mm.; antenna about
3.3 mm.

Frontal prolongation of head gray above, more brownish black
beneath; nasus long and stout, with long black setae; palpi brownish
black. Antennae relatively short; scape dark brown, sparsely prui-
nose, pedicel brown; flagellum black, segments without basal enlarge-
ments, slightly exceeding their longest verticils. Head with front
and the broad orbits clear light gray, posterior vertex darker gray,
with numerous relatively short black setae, midregion glabrous, light
brown; ventral surface of head with very long pale setae.

Pronotal scutum dark gray, scutellum yellowed. Mesonotal prae-
scutum light gray with four poorly differentiated darker gray stripes;

March 28, 1969 species of nearctic tipulidae 5

vestiture of interspaces long, erect; scutum almost uniformly dark
gray, vestiture restricted to sparse long setae on the posterior callo-
sities; scutellum brownish gray, with long setae. j)arascutella paler
gray; mediotergite gray, with sparse long setae, pleurotergite gray.
Pleura gray, glabrous; dorsopleural membrane buffy yellow. Hal-
teres with stem pale brown, yellowed at base, knob darker, apex
obscure yellow. Legs with coxae gray, with long j)ale setae; tro-
chanters dark gray; femora yellow, tips conspicuously blackened;
tibiae obscure yellow, darker outwardly; tarsi dark, claws of male
toothed. Wings faintly darkened, stigma long-oval, dark brown; veins
brown, those in costal and prearcular fields light brown. Veins pos-
terior to R essentially glabrous, with three or four trichia on distal
section of /?,+:, and still more sparsely on 2nd A. Venation: Rs about
two and one-half times m-cu; cell ist M. long, nearly parallel-sided.

Abdomen dark brownish gray, posterior borders of tergites ob-
scure yellow, of the sternites more narrowly so; hypopygium brown-
ish black, tergal lobes and styli paler. Male hypopygium with tergite
longer than broad, posterior third with two broad lobes that are
separated by a very narrow emargination, inner edges of lobes and
outer half of tergite with abundant black setae. Outer dististyle
broadest at near midlength, the greatest width at near two-thirds
the length; inner style with beak relatively slender, with a small
blackened lateral flange beyond midlength; posterior margin of style
with one or two basal spines and an outer very powerful broad-based
lobe that bears the sensory area at its base, thence narrowed into a
strong blackened spine that is directed outwardly. Eighth sternite
transverse, posterior border nearly truncate, narrowly membranous,
outer half of plate with numerous moderately long setae.

Habitat. — Alaska.

HoLOTYPE. Sable Pass. McKinley National Park. July 20, 1952
(M. E. Smith).

Tipula (Arctotipula) mckinleyana differs from other generally
similar species chiefly in details of structure of the male hypopygi-
um. particularly the tergite and inner dististyle. The most similar
species is T. {A) smithiae Alexander which differs in the wing
coloration and in the hypopygium, especially the quite different in-
ner dististyle.

Tipula (Platytipula) knowltoniana, n.sp.

Size small (wing about 10.5 mm.); mesonotal praescutum brown-
ish gray with four darker brown stripes. i)leura light gray; wings
tinged with brown, costal border darker brown; abdominal segments
with a darkened longitudinal central line; male hypopygium with
tergal lobes appearing as broad circular blades that are bordered
by black, at base of each on lower surface with a slender blackened
rod; inner style with beak stout, disk with a conspicuous blackened
bilobed structure.

Male. — Length about 13 mm.; wing 10.5 mm.

Female. — Length about 14 mm.; wing 10.5 mm.

The Great Basin Naturalist
6 CHARLES P. ALEXANDER Vol. XXIX, No. 1

Frontal prolongation of head light gray above, brownish yellow
laterally; palpi brownish black. Antennae with scape and pedicel
brownish yellow; flagelluni broken. Head gray, orbits and sides of
vertical tubercle more yellowed; a vague central darkening from
summit of tubercle caudad almost to occiput.

Cervical region yellow. Pronotal scutum restrictedly yellowed,
with three extensive gray areas, scutellum yellowed medially, sides
gray. Mesonotal praescutum brownish gray with four darker brown
stripes, the intermediate pair confluent at anterior ends, divided at
near midlength, not reaching the suture behind; posterior sclerites
of notum gray, each scutal lobe with a darkened spot laterally be-
hind the suture, parascutella more yellowed. Pleura light gray, dor-
sopleural membrane yellow. Halteres with stem light brown, knob
dark brown. Legs with coxae light gray; trochanters yellowed; fe-
mora brown, more yellowed basally, tibiae brown, tarsi passing into
brownish black; claws of male small, simple; vestiture of legs of fe-
male long and erect, much as in perhirtipes. Wings tinged with
brown, cells C and Sc conspicuously darker brown; stigmal region
chiefly obscure yellow, with a more darkened cloud at either end,
proximally including 5co, distally involving R^ and free tip of ^Sc^;
veins brown. Sparse macrotrichia on veins R.u /?4+5. M, and M . in
male, in female more sparse and restricted to outer ends of veins
/?3 to Ml. Venation: Rs and /?o+:i subequal; petiole of cell Mi from
about one-third to one-half the length of the cell.

Abdominal tergites yellowish brown, sternites paler, both with
a narrow central dark brown longitudinal stripe that becomes ob-
solete at about the seventh segment. Male hypopygium with tergal
lobes appearing as broad circular pale brown blades with narrowly
blackened margins, the intervening emargination subrectangular;
each lobe at base of lower margin with a slender blackened rod. Outer
dististyle relatively small, about three times as long as broad, tip
obtuse; inner style with beak stout, on disk of style with a conspicu-
ous blackened structure, its outer part unequally bilobed.

Habitat. — Utah.

HoLOTYPE. d", Farmington, Davis Co., September 23, 1967 (G.
F. Knowlton). Allotopotype, 9 , pinned with type.

I take great pleasure in naming this fly for my longtime friend
and colleague. Dr. George F. Knowlton. to whom much of our present
knowledge of the crane flies of Utah is due.

The fly is generally similar to the economically important Smoky
Crane fly, Tipula {Platytipula) cunctans Say, differing evidently
in hypopygial structure. A few further American members of the
subgenus bear a resemblance to these, the one most similar to the
present fly being T. (P.) perhirtipes Alexander which still is known
only from the female type taken in Kennebec County. Maine, differ-
ing in its larger size and in slight details of coloration, wing vena-
tion and trichiation of the wing veins. It is believed that the discov-
ery of the male sex of the eastern species will provide further char-
acters for the separation of the two flies.

March 28, 1969 species of nearctic tipulidae 7

Tipula (Sinotipulo) denningi, n.sj).

Belongs to the commiscibilis group, allied to josephus, differing
evidently in hypopygial structure, especially the tergite and disti-
styles.

Male. — Length about 20-21 mm.; wing 19-20 mm.; antenna
about 3.0-3.5 mm.

Female. — Length about 18 mm.; wing 22 mm.

Frontal prolongation of head dark brown, base yellowed; nasus
long and slender; palpi brown. Antennae with scape and pedicel
yellow, flagellum black; flagellar segments shorter than their verti-
cils. Head light grayish brown, orbits and lateral parts of anterior
vertex slightly more yellowed; vertex with a narrow dark brown
median vitta.

Pronotal scutum light gray, with three brown areas; scutellum
and pretergites light yellow. Mesonotal praescutum brownish yellow
with four gray stripes that are narrowly margined with black, lateral
borders broadly blackened; each scutal lobe with two darkened areas,
the major inner one oval, gray, bordered with darker, central area
obscure yellow; scutellum light brown with a narrow central dark-
ened line, parascutella pearly yellow; mediotergite gray with a
darkened central area, pleurotergite obscure yellow, patterned with
brown. Pleura obscure yellow, mesepisternum variegated with gray;
dorsopleural membrane light yellow. Halteres with stern yellow,
base of knob brown, apex broadly light yellow. Legs with coxae
yellow, patterned with brown; trochanters yellow; femora and tibiae
obscure yellow, tips blackened; tarsi light brown; claws of male
small, simple, nearly straight. Wings brownish gray, very incon-
spicuously patterned with pale brown and yellow; costal field yel-
lowed; stigma oval, brown; disk with pale brown and yellow mark-
ings, the latter near outer end of cell M adjoining vein cu, and in
cell Cu at either end of a darkened area at near one-third the length
of cell; beyond cord, with conspicuous pale yellow seams along
veins and marginally at centers of cells; veins brown. Costal fringe
short and dense; trichia of veins Sc and R abundant, /?.,^r. with
numerous trichia on more than outer half. Venation: Ra about one-
third longer than /?•_.+;!•

Abdominal tergites obscure orange yellow with a broad brown
central stripe and narrow darker brown sublateral lines, extensively
interrupted at bases and apices of segments, lateral borders narrowly
yellow; sternites yellowish brown, posterior margins of intermediate
segments yellow, narrower on outer sternites. Male hypopygium
with lateral tergal arms unequally bilobed, with dense black setae,
median area pale. Dististyles very. unequal, outer style very large,
conspicuously bilobed. provided with long black setae, upper arm
narrower, terminating in a microscopic black jioint. near base with
two blackened lobes, one glabrous, the other with abundant black
setae; inner style nmch smaller, body with long black setae, rostrum
long and slender. Phallosome a strong blackened horn, with a smaller

The Great Basin Naturalist
8 CHARLES P. ALEXANDER Vol. XXIX. No. 1

basal point. Eighth sternite with posterior border truncate, unmodi-
fied.

Habitat. — Nevada.

HoLOTYPE, (S , South Fork of Humboldt River, Elko County.
August 27. 1967 (D. G. Denning). Allotopotype. 9. with type.
Paratopotypes, 2 cT cT .

This interesting crane fly is dedicated to the collector, Dr. Donald
G. Denning, outstanding student of the North American Trichoptera.
The nearest relative is Tipula (Sinotipula) josephus Alexander, of
eastern Oregon, readily distinguished by the structure of the hypo
pygium. particularly the tergite and dististyles. T. (S.) commiscibilis
Doane is more distantly allied.

Tipula (Eremotipula) eurystyla, n.sp.

Size medium (wing of male 14 mm.); general coloration of
thorax light gray, praescutum with narrow intermediate pale brown
stripes; antennal flagellum black, segments incised; wings pale
brownish gray, prearcular and costal fields yellowed, stigma light
brown, veins brownish black; male hypopygium with tergal canthi
small, tips obtuse; outer dististyle very broad, especially outwardly,
diameter at apex nearly equal to the length; posterior margin of
eighth sternite with long yellow setae.

Male. — Length about 14.5 mm.; wing 14 mm.

Frontal prolongation of head stout, slightly exceeding the re-
mainder; nasus virtually lacking, represented by a small tubercle
that bears a few long setae; prolongation brown, gray pruinose above,
polished beneath. Antennae with scape and pedicel light yellow,
flagellum abruptly black, segments with basal enlargements well-
developed to present an incised appearance. Front and anterior
vertex yellow; posterior vertex more grayish yellow with indications
of a capillary darker median line.

Prothorax light yellow, including the scutellum and dorsopleural
membrane. Mesonotal praescutum light gray with intermediate pale
brown stripes that are narrower than the broad ground interspaces,
lateral stripes very pale; pseudosutural foveae reduced to small
shallow pits; posterior sclerites gray, each scutal lobe with two pale
brown areas, the anterior one very small; scutellum and medio-
tergite with a poorly indicated darkened median line, parascutella
more yellowed. Pleura gray. Halteres with stem yellow, knob
brown. Legs with coxae gray; trochanters yellow; femora obscure
yellow, tips darker; tibiae yellowish brown, darker outwardly, tarsi
passing into black; claws of male with a strong tooth. Wings pale
brownish gray, prearcular field, together with cells C and Sc yellow,
stigma light browTi, cell Sci above it more yellowed; outer ends of
radial cells and extensive clouds in both anal cells very slightly
darker than the ground; obliterative area at cord whitened, incon-
spicuous; veins brownish black, much paler in the brightened costal
field. Vein 1st A without trichia, 2nd A with these numerous and
well distributed, extending basad almost to the arculus. Venation:

March 28, 1969 species of nearctic tipulidae 9

Petiole of cell Mi shorter than the oblique m; m-cu shortly before
fork of A/:,+4.

Basal abdominal tergites light brownish gray, remaining tergites
obscure yellow, the more proximal ones with indications of three
brownish gray stripes, outer segments more uniformly yellow; ster-
nites yellow, with abundant small erect dark setae. Male hypojiygi-
um with subtergal })rocess of tergite elongate, the slender outer half
pale and more membranous; canthi small, outer j)art narrow, apex
obtuse. Outer dististyle distinctive, very broad, widened outwardly,
apex nearly truncate; the diameter across apex nearly equal to the
length of the style; inner style with dorsal crest strongly rounded,
blackened; beak obtuse, lower beak very stout; outer basal lobe sub-
oval, in size and outline not greatly different from the outer style.
Eighth sternite with caudal margin fringed with very long yellow
setae, none conspicuously enlarged. In the unique type the sternite
is damaged but appears to have posterior lateral lobes that bear
unusually long setae, with a further development of long setae be-
tween these groups.

Habitat. — Utah.

HoLOTYPE, cT. Dixie State Park, Washington County, April 24,
1968 (G. F. Knowlton and D. W. Davis).

The most similar species appears to be Tipula {Eremotipula)
mitrata Dietz, of New Mexico, which is known to me only from the
original description. From this the species appears to differ in colora-
tion and structure of the antennae, coloration of the wings, and in
details of hypopygial structure. The outer dististyle of the hypo-
pygium in the present fly shows the maximum breadth of any
species in the subgenus.

Gonomyia (Idiocera) per simplex, n.sp.

In general appearance most as in coloradica and gothicana,
especially in the unpatterned wings; male hypopygium with all three
dististyles simple.

Male. — Length about 6 mm.; wing 5 mm.

Described from alcoholic material. Rostrum and palpi brown.
Antennae black, proximal half of scape brownish yellow; flagellar
segments oval, the outer ones more elongate. 1 lead dark brown,
anterior vertex, sides of vertical tubercle and the genae yellowish

gray-

Pronotal scutum yellow, patterned with dark brown, scutellum
yellowed. Mesonotal praescutum dark brown, lateral borders and an
isolated area in humeral region light yellow; scutal lobes dark brown;
scutellum brown. i)osterior border yellow; parascutella. pleurotergite
and lateral borders of mediotergite light yellow. Pleura dark brown,
dorsopleural region and a broad longitudinal stripe whitened. 1 lal-
teres with stem j)ale. knob weakly darkened. Legs with coxae yel-
lowed, fore pair slightly more infuscated; trochanters yellow; femora
light brown, slightly darker near tips; remainder of legs brown.
Wings subhyaline, unpatterned except for the small pale brown

The Great Basin Naturalist
10 CHARLES P. ALEXANDER Vol. XXIX. No. 1

stignia, prearcular and costal fields slightly more yellowed; veins
pale brown. Longitudinal veins beyond cord with trichia except /?,,.
also occurring on Rs, extreme outer end of basal section of Cui and
the distal third of ist A. Venation: Veins /?uj and R^ contiguous at
margin; m-cu more than its own length before fork of M.

Abdomen dark brown, pleural membrane yellowed. Male hypo-
pygium with outer lobe of basistyle broadly obtuse, with abundant
long setae. All three dististyles simple; outer style longest, appearing
as a slender pale spine that narrows to the long terminal point; inter-
mediate style blackened on outer part, similar in shape to the outer
style but shorter; inner style very pale, widened basally. outer end
slender, blackened. Aedeagus of unique type broken. The inter-
mediate styles of the two sides are unequal, one being only about
one-half the size of the other, this presumably representing an ab-
normal condition.

Habitat. — Utah.

HoLOTYPE, alcoholic cT- Fish Creek, Sevier County, June 21.
1967 (C. D. Hynes).

The most similar regional species include Gonomyia (Idiocera)
color adica Alexander an,d G. (/.) gothicana Alexander, both quite
different in hypopygial structure, especially the dististyles, as
described.

References

Lackschewitz, Paul. Das genus Tipula (Diptera Nematocera) in der Arktis

und dem borealen Waldgebiet Eurasiens. Trav. Inst. Zool. Acad. Sci. U R S S,

4: 246-312, 34 figs, (with numerous subfigs); 1936.
Savtshenko, Eugen N. (Savchenko) Fauna USSR, Diptera II, no. 3. Fam.

Tipulidae, Subfam. Tipulinae, Genus Tipula Linn, (part l)-in Russian.

Zool. Inst. Akad. Nauk, S S S R (Moscow), N. S., No. 79: 1 - 486, 295

figs.; 1961.

Tl IE BUPRESTIDAE AND CLERIDAE OF THE
NEVADA TEST SITE (COLEOPTERA) '

William F. Barr"

The collecting phases of the comprehensive ecological investiga-
tions conducted under the auspices of Brigham Young University at
the Nevada Test Site in southern Nye County, Nevada, as reported
by AUred. Beck and Jorgensen (1963a) yielded relatively few
specimens of the beetle families Buprestidae and Cleridae. This is
somewhat surprising inasmuch as these groups are usually considered
to be major components of the Coleopterous fauna occurring in the
desert and adjacent mountainous areas of the southwestern United
States. However, several of the collections that were made do provide
new or otherwise significant distributional or bionomical information
for some of the species encountered. Therefore, a preliminary listing
of the species of these two families taken at the Test Site along with
a presentation of the available collection information seems war-
ranted at this time.

For the exact location of collection sites and areas as listed herein,
reference must be made to the papers by Allred. Beck and Jorgensen
(1963b) and Knight (1968).

Appreciation is expressed to Drs. D. M. Allred and the late
D. E. Beck of Brigham Young University for allowing me the oppor-
tunity of studying their material. This material was collected under
Contracts AT( 11-1) 786 and AT(11-1)1326 between the Atomic
Ejiergy Commission and Brigham Young University.

Buprestidae
Hippomelas {Gyascutus) near obliterata LeConte

Specific determination of the 14 specimens taken at the Test Site
is not possible at this time. Perhaps they represent one of the several
species described by Casey (1909) from southwestern Utah which
have been relegated to synonymy in the most recent catalogs. Un-
fortunately. Casey's descriptions are not sufficient in their charac-
terization of important features to allow identification of the Test
Site material and I have not had opportunity to make comparisons
with his type material.

The Test Site collections were made between July 2 and August
2 in 1960, 1961. and 1962, at 9 and 9.5 miles N of Well 3B in a
Coleogyne habitat, in the environs of Well 3B in a Grayia-Lycium

'Published with the approval of tlie Direttor of the Idalio Agi-itullural F,xpcrin\pnt Station
js Research Paper No. 780.

'University of Idaho, Moscow.

11

The Great Basin Naturalist
12 WILLIAM F. BARR Vol. XXIX. No. 1

community, at 1 0 miles NW of Mercury and in the environs of Well
5B in a Larrea-Franseria community.

Melanophila consputa LeConte

Three large specimens, ranging in length from 12 to 14 mm.,
were taken in the euAdrons of Mercury on Larrea divaricata (DC.),
at 10 miles NW of Mercury and at Ground Zero in a can pit- trap
in a Grayia-Lycium community. Collection dates were July 8, 1960.
and July 22, 1962, and 1965.

M. consputa is known to develop only in coniferous trees, conse-
quently the occurrence of specimens in desert areas of the Test Site
some distance from a coniferous habitat may seem unusual. How-
ever, these specimens probably were attracted to smoke or fumes
that were present in the desert areas. Such materials are known to
be highly attractive to M. consputa and related species.

Melanophila pini-edulis Burke

This uncommon species is known from the intermountain West
and adjacent regions, but has not been reported specifically from
Nevada. One specimen was collected in Area 18 on June 20, 1965,
and was associated wdth Pinus monophylla Torrey and Fremont.

Anthaxia {Haplanthaxia) deleta deleta LeConte

One specimen of this common and wide-ranging subspecies was
taken in Area 12 on June 11, 1965, from Cowania stansburiana
Torrey in a Juniper-Pinyon community.

Chrysobothris cuprascens LeConte

Six specimens were collected at Rainier Mesa on June 15, 1964,
from Pinus monophylla Torrey and Fremont and Juniperus osteo-
sperma (Torrey). They are assigned to C. cuprascens rather than
to the closely related C. semisculpta LeConte on the basis of host
plan.t association and geographic distribution. According to Fisher
(1942) these two species are doubtfully distinct.

Chrysobothris arizonica Chamberlin

A single specimen, tentatively assigned to this species, was col-
lected along the Basalt Area road, 12 miles NW of Tippipah Spring
in a Grayia-Lycium community on July 13. 1964. from Baileya
pleniradiata Harvey and Gray. The finding of C. arizonica on this
plant probably constitutes a visitation rather than a host record.

Chrysobothris platti Cazier

This species has not been recorded previously from Nevada. One
female was collected in the environs of Tippipah Spring on July

March 28, 1969 buprestidae & cleridaf. in nkvada 13

23. 1965. from Ephedra viridis (yoville which uncloubtcdly is a host
of this beetle.

Acmaeodera lanata Horn

A single specimen of this well-known southwestern species was
collected in Area 16 on June 11, 1965. from Dalea polyadenia Torrey.
The larva of this species is known to bore in the roots and lower
stems of Ephedra spp.

Acmaeodera purshiae Fisher

Five specimens of this attractive species were collected. Three
were taken in the environs of Tippipah Spring on June 14. 1965,
from Fallugia paradoxa (D. Don), one was found 10 miles NW of
Mercury on June 20, 1965, on Purshia glandulosa Curran and one
was taken in the Mid Valley Area on July 28. 1962, and was
associated with P. tridentata (Pursh). This species has been recorded
previously from several areas of California and Oregon and from
the Reno area of Nevada. Its only known host plant is P. tridentata.

Acmaeodera diffusa Barr, n. sp.

(Fig. 1)

Acmaeodera variegata, auctorum

Male. — Medium-sized, moderately robust, black; pronotum
blackish-bronze with a small yellow spot at sides immediately be-
hind middle; each elytron with four transverse irregular straw-
colored spots extending from lateral margin to near suture, first spot
in front of basal fourth, second spot in front of middle and anteriorly
oblique on disk, third spot in front of apical fourth and anteriorly
oblique on disk, fourth spot in front of apex, small and oblique; upper
surface moderately clothed with rather long erect and suberect brown
hairs; ventral surface and legs moderately clothed with suberect
whitish hairs.

Head densely punctured, punctures small and deep; vertex with-
out a median carina; front convex except upper portion obliquely
flattened; clypeus with front margin broadly, subtriangularly emar-
ginate; antenna shining, extending to about hind margin of proster-
num, serrate from fifth segment, outer segments slightly wider than
long.

Pronotum slightly less than twice as wide as long, irregularly
convex with a shallow transverse depression behind front margin,
a small transverse median depression in front of base, a very faintly
indicated median longitudinal depression in front of middle and a
broad shallow depression surrounding the conspicuous subbasal pits;
sides arcuate, widest at about basal third; lateral margins very
slightly reflexed. visible from above on anterior half; front margin
very broadly and feebly triangularly lobed at middle; surface more

The Great Basin Naturalist
14 WILLIAM F. BARR Vol. XXIX, No. 1

coarsely and densely punctured than on head, becoming sub-reticu-
late towards sides.

Elytra subequal in width to pronotum; disk irregularly de-
pressed and flattened except for slightly elevated area near sides at
basal fourth and for the narrowly elevated suture on apical three-
fourths; sides nearly vertical, when viewed from above, feebly nar-
rowing at basal fourth, slightly expanded at middle, and gradually
evenly narrowing to conjointly rounded apices; front angles rectan-
gular when viewed from the side; lateral margins broadly bisinuate
when viewed from the side, most deeply emarginate at basal fourth,
coarsely serrate on apical half; surface deeply punctured, slightly
roughened at extreme base, strial punctures about subequal to those
on thorax becoming smaller apically, interstrial spaces flattened
with inconspicuous tiny punctures, third interstrial space feebly
elevated, ninth interstrial space elevated at extreme base.

Ventral surface. — Thorax coarsely, densely punctured; front
margin of prosternum retracted, subtruncate, very slightly reflexed
on either side of middle. Abdomen rather finely densely punctured
except for first sternite which is more coarsely punctured at middle
and along front margin; last sternite short and broad, feebly convex
at middle and vsdthout a subapical plate or elevation, margins nar-
rowly reflexed, lateral margin straight, oblique, hind margin very
broadly rounded.

Length. — 8.4 mm.; width, 2.9 mm.

Ho lo type, male (California Academy of Sciences) from Highland
Range, above Mendha, Lincoln County, Nevada, July 9, 1965 (W.
F. Barr) . Two hundred and twenty- three paratypes from type locali-
ty July 9, 1965 (W. F. Barr and R. L. Westcott), and June 25, 1966
(E. J. Allen, W. F. Barr, D. S. Horning, Jr., R. L. Westcott and R. L.
Penrose), most specimens collected from the flowers of Opuntia sp.
and Sphaeralcea sp., others on the flowers of Chaenactis sp., Garrea
sp. and Calochortus sp. One paratype from Nevada Test Site, Mer-
cury, Nevada, June 11, 1965. on Viguiera multiflora (Nutt.) Blake.
Ten paratypes from Pine Valley, Washington County, Utah. June
2. 1960 (Verity and Raven), and from the same locality. 31 para-
types, June 12 and 14, 1961 (D. W. Davis and B. Haws), one para-
type June 25, 1948, and one paratype June 27, 1933 (J. T. Howell).
Paratypes in the collections of the American Museum of Natural
History, California Academy of Sciences, IjOs Angeles County
Museum, U. S. National Museum, Brigham Young University, Uni-
versity of Idaho, Utah State University; W. F. Barr, F. M. Beer,
D. S. Horning. Jr., J. N. Knull, G. H. Nelson, R. L. Penrose, D. S.
Verity, and R. L. Westcott.

Many additional specimens have been examined which have not
been designated as paratypes. Distributional information from these
specimens is as follows: Arizona: Betatakin Cyn.; Navajo National
Monument; Grand Canyon; Jacob L^ake; Kayenta, Navajo County;
Navajo Mt., Navajo County; Prescott; and near Williams. Colo-

March 28, 1969 buprestidae & cleridae in Nevada

15

Fig. 1. Acmaeodera diffusa, new species, dorsal view of holotype.

RADO: Durango; Glenwood Springs; Meeker; and Mesa Verde.
Utah: American Fork Canyon; Aspen Grove. Timpanogos; Beaver
Canyon; Beaver Cr., Kamas; Bryce Canyon; Cainville; City Cr.
Canyon; Dugway; Emigration Canyon, Salt Lake County; Escalante;
Farmington, Davis County; Fort Douglas; Fruita; (ilendale. Tlanks-
ville; Indianola; Jordan River; Lehi; Little Cottonwood Canyon,
Wasatch Mountains; Mount Dell Cr.. Salt Lake City; Mt. Nebo.
Juab County; Mt. Timpanogos; North Fork of Provo Canyon; Oak
Creek Canyon; Ogden; Park City; Parley's Canyon; The Pass. Table
Cliff Mountain. Garfield County; Pay son Canyon, Utah County;
Provo; Provo Canyon; Red Butte; Rock Canyon, Provo; St. George;

The Great Basin Naturalist
16 WILLIAM F. BARR Vol. XXIX. No. 1

Salt Lake; Salt Lake City; Spanish Fork; Stockton; Vivian Park;
Wasatch Mountains; Zion Lodge; Zion National Park.

A. diffusa is a well-known species that has a wide distribution
range in the western United States. It is especially abundant in the
Intermountain Regions. In most of the literature and in most collec-
tions it is referred to as A. variegata LeConte, but that name must be
utilized for another distantly related species that occurs in New-
Mexico and Arizona.

This species may be separated from other known species by the
lack of a subapical plate or elevation on the shortened and broadly
rounded last abdominal sternite, by the irregularly flattened elytral
disk and by the coarsely and densely, nearly cribrately punctured
pronotmn which usually is of a dark bronzy color. The yellowish
elytral markings, consisting of three pairs of fasciae and a pair of
subapical spots, although distinctive, are variable. These markings
usually are irregular and interrupted at the suture, but may be
broken, reduced or greatly expanded. The pronotal spots also are of
a variable nature. They exhibit considerable difference in size, may
be present on one side of the pronotum or may be entirely absent.
In the case of the type series these spots are present on approximate-
ly 70 percent of the specimens.

Acmaeodera sp.

Four specimens belonging to the retifer species complex were
taken in an Artemisia community 4.4 miles S of Tippipah Spring
and at Tippipah Spring on June 13, 1964. They were found on
Argemone corymnosa (Greene) and Eriogonum fasciculatum Benth.

The retifer complex is in need of a thorough taxonomic study
before names can be assigned to its many populations that occur in
the Pacific Coast and intermountain states.

Acmaeodera immaculata Horn

One specimen of this widespread Great Basin and Mohave Desert
inhabitant was collected 4.4 miles S of Tippipah Spring in an
Artemisia community. It was found on Baileya pleniradiata Harvey
and Gray. In southern Idaho A. immaculata is known to develop
only in the roots of Eurotia lanata (Pursh), but adults are frequent-
ly encountered on the flowers of a variety of plants including the
Compositae.

Agrilus pubifrons Fisher

This species, previously known from southern Idaho and eastern
Oregon, is recorded from Nevada for the first time. Four specimens
were taken at the Test Site in Area 12 and in Area 18 on June 11,
July 27 and August 7. 1965, on Chrysothamnus visidiflorus (Hook.),
C. parryi (A. Gray) and Grayia spinosa (Hook.). C. viscidiflorus
is its only known host plant. .

March 28. 19()9 buphkstidak & clkkidak in nkvada 17

Agrilus felix Horn

One specimen, tentatively identified as this species by J. N. Knull,
was collected at 4.5 miles SE of Well 5B on luric 11. l<Hi4. from
Sphaeralcea ambigua.

Cleridae
Cymatodera unijormis Schaeffer

Collections of this s|)ecies were made at Yucca Flat in a (irayia-
Lycium community, at Rainier Mesa in a Juniper-Pinyon com-
munity and in Area 12 in a Juniper-Pinyon community. The five
specimens collected on August 13 and 14. 1964. were attracted to
incandescent and black light.

C. uniformis has been recorded previously from southern Arizona
and southern California where it has been reared from Juniperus
spp.

Cymatodera latefascia Schaffer

This distinctive species has not been known from Nevada. Two
specimens were collected at Rainier Mesa on August 9, 1964. in an
Artemisia community and in Area 18 on July 25, 1965. In both
collections the specimens were attracted to incandescent light traps.

Cymatodera oblita Horn

Four specimens were found at Rainier Mesa in a Juniper-Pinyon
community and in the Area 12 residence area. They were attracted
to black light and incandescent light on August 8 and 11, 1964.

Cymatodera fuchsii Schaeffer

This was the most commonly encountered clerid species of the
Test Site collections. Forty-seven specimens were collected at the
following locations: 10 miles NW of Mercury; 9.3 miles W of
Mercury; environs of Mercury; Area 5 in a Grayia-Lycium com-
munity; Well 5B; W of Frenchman Playa. in Larrea and Lycium
communities; Yucca Flats. 5.5 miles NW of Well 3B, in a Grayia-
Lycium community; and Jackass Flats, W of Cane Springs. Dates
of the collections ranged from July 15 to August 24. The specimens,
which w^ere taken in can ])it-traps. at blacklight and at incan-
descent light, are remarkedly uniform in the coloration of the
elytra. All have the elytral fascia moderately distinct. The great
majority are of a pale reddish-brown color, with few specimens
showing a darker coloration, but none exhibit the usual tan color of
specimens from other localities.

Phyllobaenus (juadrimaculatus (Van Dyke)

A single female specimen was found at Cane Springs on June 13.
1965. on Stanleya pinnata (Pursh). This specimen appears to fit the

The Great Basin Naturalist
18 WILLIAM F. BARR Vol. XXIX. No. 1

original description of P. quandrimaculatus except that the dark por-
tions of the elytra lack a greenish luster and the elytral markings are
more extensive. In addition the legs of the Cane Springs specimens
are yellowish except for the darkened apices of the mesofemora and
the predominately darkened metafemora.

Phyllobaenus pygmaeus (Wolcott)

One specimen, tentatively assigned to this species, was collected
W of Cane Springs on June 18, 1965, from Artemisia sp. Positive
identification of many of the described species of North American
Phyllobaenus is not possible because of the poor taxonomic state of
the genus.

Phyllobaenus subfasciatus (LeConte)

A specimen collected W of Cane Springs on June 20. 1965, from
Atriplex canescens (Pursh) falls within the currently recognized
concept of this widespread and very variable species.

Trichodes ornatus Say

This well-known species was not commonly found at the Test
Site. Only 13 specimens were taken, but two variants are represented
in the series. Twelve of the specimens have reddish elytra with the
black fasciae showing varying degrees of reduction, especially on the
basal half. This variation is characteristic of populations occurring
in the Great Basin and immediately adjacent areas. The other speci-
men represents one of the Sonoran Desert variations which has the
elytra predominately yellow in color with narrow black fasciae.

Collections were made at Tippipah Spring, Area 17, Mercury.
Yucca Flat and Mid- Valley on May 18, 1960, June 12-17, 1965, and
July 21, 1962. The specimens were found probably on the flowers of
Sphaeralcea sp., Tetradymia glabrata Gray, Chrysothamnus sp. and
Malacothrix glabrata Gray.

Serriger reichei Spinola

This uncommon species was collected at Yucca Flat in a Grayia-
Lycium community, at Cane Springs, W of Cane Springs and at Area
9. Seven specimens were taken between June 10 and 18, 1965, from
Sphaeralcea sp., Tetradymia glabrata Gray, Lepidium fremontii S.
Wats, and Salazaria mexicana Torrey. These collections constitute
a new state record for Nevada. Previously, S. reichei has been known
in the literature only from single localities in southeastern California
and southwestern Idaho.

Literature Cited

Allred, D. M., D E. Beck and C. D. Jorgensen. 1963a. Biotic Communities
of the Nevada Test Site. Brigham Young Univ. Sci. Bull., Biol. Sen, 2(2) :1-
52.

March 28, 1969 buprestidae & clkridae in nkvada 19

1963b. Nevada Test Site Study Areas and Specimen Depositories. Brig-

ham Young Univ. Sci. Bull., Biol. Ser., 2(4): 1-15.

C.\SEY, Thom.'^s L. 1909. Studies in the American Buprestidae. Proc. Washing-
ton Acad. Sci., 11 (2) : 47- 178.

Fisher, W. S. 1942. A Revision of the North American Species of Buprestid
Beetles Belonging to the Tribe Chrysobothrini. USDA Misc. Publ. No. 470.
pp. 1-274.

Knight, H. H. 1968. Taxonomic Reviews: Miridae of the Nevada Test Site
and the Western United States. Brigham Young Univ. Sci. Bull., Biol. Ser.,
9(3): 1-282.

BEES OF THE NEVADA FEST SITE^

Dorald M. Allred

During the years from 1959 to 1965, many bees were collected
at the Nevada Test Site as part of the ecological studies conducted
by the Department of Zoology and Entomology of Brigham Young
University, under contract with the U. S. Atomic Energy Commis-
sion. Most of these were recently identified by Dr. G. E. Bohart,
Entomology Research Division, Department of Agriculture. Logan,
Utah, and Mr. P. II. I'imberlake. Riverside, California. The identifi-
cations form the basis of this report on the host relationships, sea-
sonal occurrence, and geographic distribution at the test site.

Further studies specifically directed toward the bees, particularly
during the blooming periods of the desert plants, undoubtedly would
yield additional records. Genera and species are listed alphabeti-
cally for convenience. Areas of collection refer to divisions deline-
ated by Allred. Beck, and Jorgensen (1963a and b). Methods of
collection used were hand capture, aerial and sweep nets, and can
pit-traps ( AWred et al., 1963a).

Species And Plant Associations

Agapostemon cockerelli: 2$ $ ; areas 17. 18; Malacothrix glabrata;
June. July.

A. texana: 2cf cT 3 ? ? ; areas 5, 12, CB. Asclepias erosa, Petradoria
pumila, Stanleya pinnata (most common host); June. Aug.

Andrena sp.: cf ; area CE; host unknown; March.

Anthidium dammersi: d" 9 ; areas 1. 18; Erigeron pumilus; May;
June.

Anthophora californica: (S 2 ; area 5; Stanleya pinnata., June.

A. porterae: cf ; area 4; host unknown; March.

A. urbana: 8crcf 24? 5 ; areas 5, 12. 17, 18. CB, TM; Asclepias
erosa. Astragalus lentiginosus, Gayophytum ramosissima, Mala-
cothrix glabrata, Oenothera californica, Petradoria pumila, Stan-
leya pinnata; most common host was S. pinnata, with A. erosa
and P. pumila also frequently represented; most were taken in
June, about half as many in July, and few in August.

Anthophora new species: ScT d" 1 2 ; area CE; host unknown; March.

Ashmeadiella aridula: ? ; area 2; Pectis papposa, Aug.

A. australis: 9 9 2?, area 12. Penstemon palmeri; July.

A. bigeloviae: 9 ; area 10; Salsola kali; Sept.

A. inyoensis: 4cf cf ; area 5; host unknown; May.

A. opuntiae: 2<S d 3 9 9 ; areas 12, 13. CB; Echinocereus engelman-
nii (most common host). Opuntia erinacea; May. July.

Bombus morrisoni: 7 9 9; areas 5, 12; Petradoria pumila (most
common host) , Stanleya pinnata, Viguiera multi flora; July, Aug.

^BYU-AEC publication no. COO- 1355 19.

20

March 28, 1969 bees of the Nevada test site 21

Centris rhodopus: \9d d 6 9 9 ; areas 5, 6, CB, 401. Cleome lutea,
Eriogonum inf latum, Stanleya pinnata (most common host),
Tamarix pcntandra; June-Sept., mostly July.

Ceratina nanuhr. ?; area 17; Malacothrix glabrata; June.

Chelostomoides lohatifrons: cf ; area CB; Asclepias erosa; June.

Dialictus alhohirtus: SScTcT 39 9; areas 5, 6. 12. CB, EM, T;
Artemisia tridentata. Asclepias erosa (most common host),
Chrysothamnus paniculatus, Eriogonum inflatum, Eriogonum
fasiculatum, Larrea divaricata, Petradoria pumila. Salix good-
dingii, Stanleya pinnata, Tamarix pentandra; June (mostly),
July.

D. hyalinus: d H ? ? ; areas 1, 5, 6, 12, 17, CB. EC; Acamptopap-
pus shockleyi, Asclepias erosa, Calochortus flexuosus, Malacothrix
glabrata, Penstemon palmeri, Stanleya pinnata, Viguiera multi-
flora; May-Aug.

D. incompletus: 129 9 ; areas 12. FM; Bromus rubens. Calochortus
flexuosus, Penstemon bridgesii, Penstemon jloridus, Penstemon
palmeri (most common host) ; July.

D. microlepoides: W d d 79 9; areas 12. CB. EM; Artemisia tri-
dentata, Asclepias erosa (most common host), Eriogonum fasicu-
latum, Penstemon bridgesii, Tamarix pentandra, Viguiera multi-
flora; June (mostly). July.

D. nevadensis: 29 9 area 12; Penstemon palmeri; July.

D. pruinosum: cT ; area CB; Asclepias erosa; June.

Dialictus sp. 1, ? new: 29 9; area 12; Castilleja linariae folia,
Chrysothamnus sp.; July. Aug.

Dialictus sp. 2, ? new: d" ; area CB; Asclepias erosa; June.

Dialictus sp. 3, ? new: 9 ; area 410; Eriogonum fasiculatum; July.

Colletes eulophi: d 9 ; areas 5. 17; Sphaeralcea sp.. Stanleya pinnata;
June.

Colletes sp., ? new: 9 ; area 17; Sphaeralcea sp.; June.

Diadasia australis: 2cf cf; areas 12. 17; Opuntia erinacea; June. July.

D. diminuta: 4cf d" 3 9 9 ; areas 5, 17; Sphaeralcea ambigua (most
common host) , Stanleya pinnata; June.

D. lutzi: cT 9 1?; areas 5. 17; Sphaeralcea sp.. Stanleya pinnata;
June.

Dianthidium pudicum: d ? ; areas 16, 19; Chrysothamnus viscidi-
florus; June. Aug.

D. subparvum: cT 9 ; area 12; Chrysothamnus viscidiflorus , Petra-
doria pumila; July. Aug.

D. ulkei: 2(f cT; areas 12. FC; Artemisia tridentata; July.

Dioxys productus: <S ; area 5; host unknown; March.

Dufourea new species 1: 9 ; area JA; host unknown; April.

Dufourea new species 2: 9 9; area 12; Phacelia peirsoniana- June.

Epeolus minimus: 9 ; area 5; host unknown; May.

Evylaeus ruficornis: 9 ; area 19; host unknown; June.

Halictus hyalinus: 9 ; area 401 ; Chrysothamnus parryi; Aug.

H. tri parti t us: 79 9; areas 12. CB, FM; Calochortus flexuosus,
Eriogonum fasiculatum, Lupinus palmeri, Penstemon sp., Petra-
doria pumila, Viguiera multiflora; June, mostly July, Aug.

The Great Basin Naturalist
22 DORALD M. ALLRED Vol. XXIX. No. 1

Heriades timberlakei: 6<S d 4? $ ; areas 12. 13; Opuntia erinacea.
Viguiera multiflora; July.

Hesperapis wilmattae: ? ; area 401; Malacothrix glabrata; June.

Hylaeus asininus: 2^ d ; area 12; Chrysothamnus viscidijlorus;
Aug.

Hypomacrotera subalpinus: 4j'cr; area 5; Baileya multiradiata,
Sphaeralcea ambigua, Stanleya pinnata; June.

Lasioglossum sisymbrium: 3 9 9 5?; areas 5, 12, 18. 19; Chryso-
thamnus viscidijlorus, Erigeron pum.il is, Hymenoxys cooper i,
Penstemon sp., Stanleya pinnata, Viguiera multiflora; June-Aug.

Lithurgus apicalis: 3d d 19; areas 12. 13; Opuntia erinacea; July.

Melectamorpha californica-. 2d d 29 9; areas 5, CB; Asclepias
erosa, Stanleya pinnata; June. July.

Melissodes subagilis: 79 9; areas 5, 12. 17; Baileya multiradiata,
Chrysothamnus viscidijlorus. Petradoria pumila, Sphaeralcea
ambigua, Stanleya pinnata; June-Aug.

M. tristis: 9 ; area 12; Petradoria pumila; July.

Micranthophora hololeuca: 2d d 29 9; area TA; Sphaeralcea
munroana; June.

M. phenax: d ; area 1; host unknown; April.

Nomia tetrazonata: d ; area CE; Berula erecta; July.

Osmia titusi: d ; area 5; host unknown; March.

Osmia sp., ? new: d 9 ; areas 5. 17; Astragalus lentijinosus. Descu-
rainia pinnata; May, June.

Perdita arcuata: cf 2 9 9 ; area 5; Sphaeralcea ambigua; June.

P. callicerata: d '-, area 12; Chrysothamnus paniculatus; June.

P. chloris: 9 ; area 16; Eriogonum deflexum; Aug.

P. jallugia: 5 d d '-, areas 12. 18; Cowania sp. (most common host),
Erigeron pumilus, Oenothera calijornica; June.

P. nasuta: 2 9 9 7?; area 6; Eriogonum injlatum; July.

P. thermophila: d 9 ; areas 6. 410; Eriogonum injlatum; July.

Perdita new species: 89 9; areas 16. 401; Dalea polyadenia, Erio-
gonum dejlexum (most common host) ; Aug.

Sphecodes eustictus: 9 ; area 12; Eriogonum dejlexum; Aug.

Stelis new species: 9 ; area 10; host unknown; April.

Tetralonia quadricincta-. d ; area 4; host unknown; March.

Tetralonia new species "al": cT 2 9 9 ; areas 4, 5; Stanleya pinnata;
March. June.

Tetralonia new species "an": d '-, area 4; host unknown; March.

Tetralonia new species "mo": 9 ; area 4; host unknown; March.

Tetralonia new species "pr": 4cr d 32?; areas 4, 5; host unknown;
March.

Triepeolus helianthi: d; area 5; Sphaeralcea ambigua; June.

Xylocopa calijornica: 9 7?; areas 5, M; Stanleya pinnata; June
(mostly), July.

Plant-Bee Associations

Acamptopappus shockleyi Dialictus albohirtus

Dialictus hyalinus D. microlepoides

Artemisia tridentata Dianthridium ulkei

March 28, 1969 bees of the Nevada test site

23

Asclepias erosa

Agapostemon texana

Anthophora urbana

Chelostornoides lobatifrons

Dialictus albohirtus

D. hyalinus

D. microlepoides

D. pruinosum

Dialictus sp. 2

Melectaniorpha calif or nica
Astragalus lentiginosus

Anthophora urbana

Osmia sp.
Baileya multiradiata

Hypomacratera subalpinus

Melissodes subagilis
Berula erecta

Nomia tetrazonata
Bromus rubens

Dialictus incompletus
Calochortus flexuosus

Dialictus hyalinus

D. incompletus

Halictus tripartitus
Castilleja linariaefolia

Dialictus sp. 1
Chrysothamnus paniculatus

Dialictus albohirtus

Perdita callicerata
Chrysothamnus parryi

Halictus hyalinus
Chrysothamnus viscidiflorus

Dianthidium pudicum

D. subparvum

Hylaeus asininus

Lasioglossum sisymbrium

Melissodes subagilis
Chrysothamnus sp.

Dialictus sp. 1
Cleome lutea

Centris rhodopus
Cowania sp.

Perdita fallugia
Dalea polyadenia

Perdita sp.
Descurainia pinnata

Osmia sp.
Echinocereus engelmanni

Ashmeadiella opuntiae
Erigeron pumilis

Anthidium dammersi

Lasioglossum sisymbrium

Perdita fallugia
Eriogonum defleiuni

Perdita chloris

Perdita sp.

Sphecodes eustictus
Eriogonum fasiculatum

Dialictus albohirtus

D. microlepoides

Dialictus sp. 3

Halictus tripartitus

Eriogonum inflatum

Centris rhodopus

Dialictus albohirtus

Perdita nasuta

P. thermophila
Gayophytum ramosissima

Anthophora urbana
Hymenoxys cooperi

Lasioglossum sisymbrium
Larrea divaricata

Dialictus albohirtus
Lupinus palmeri

Halictus tripartitus
Malacothrix glabrata

Agapostemon cockerelli

Anthophora urbana

Ceratina nanula

Dialictus hyalinus

Hesperapis wilmattae
Oenothera californica

Anthophora urbana

Perdita fallugia
Opuntia erinacea

Ashmeadiella opuntiae

Diadasia australis

Heriades timberlakei

Lithurgus apicalis
Pectis papposa

Ashmeadiella aridula
Penstemon bridgesii

Dialictus incompletus

D. microlepoides
Pensteman floridus

Dialictus incompletus
Penstemon palmeri

Ashmeadiella australis

Dialictus hyalinus

D. incompletus

D. nevadensis
Penstemon sp.

Halictus tripartitus

Lasioglossum sisymbrium
Petradoria pumila

Agapostemon texana

Anthophora urbana

Bombus morrisoni

Dialictus albohirtus

Dianthidium subparvum

Halictus tripartitus

Melissodes subagilis

M. tristis
Phacelia peirsoniana

Dufourea sp. 2
Salii gooddingii

Dialictus albohirtus
Salsoli kali

Ashmeadiella bigcloviae
Sphaeralcea ambigua

Diadasia diminuta

Hypomacrotera subalpinus

Melissodes subagilis

Perdita arcuata

24

DORALD M. ALLRED

The Great Basin Naturalist

Vol. XXIX. No. 1

Triepeolus helianthi
Sphaeralcea rnunroana

Micranthophora hololeuca
Sphaeralcea sp.

Colletes eulophi

Colletes sp.

Diadasia lutzi
Stanleya pinnata

Agapostemon texana

Anthophora calif or nica

A. urbana

Bombus morrisoni

Centris rhodopus

Dialictus albohirtus

C. hyalinus
Colletes eulophi
Diadasia diminuta

D. lutzi

Hypornacrotera subalpinus
Lasioglossum sisymbrium
Meleclamorpha californica
Melissodes subagilis
Tetralonia sp.
Xylocopa californica

Tamarii pentandra
Centris rhodopus
Dialictus albohirtus
D. microlepoides

Viguiera multiflora
Bombus morrisoni
Dialictus hyalinus
D. microlepoides
Halictus tripartitus
Heriades timberlakei
Lasioglossum sisymbrium

Summary

Between 1959 and 1965, bees representing 71 species of 35 genera
were collected from plants of more than 40 species at the Nevada
Test Site. Bees of nine of the species are new to science, and six
others possibly are new. Most specimens were taken during June
and July, but some were found as early as March and others as late
as September. Bees of greatest abundance, as indicated by those
collected, were Dialictus albohirtus, Tetralonia n. sp. "pr," Antho-
phora urbana, and Centris rhodopus. Those with the widest geo-
graphic distribution at the test site were Dialictus hyalinus, D. albo-
hirtus, and Anthophora urbana. The species found on the greatest
variety of plants was Dilictus albohirtus. A greater variety of bees
was found on Stanleya pinnata, Asclepias erosa, and Petradoria
pumila than on other plants.

References

Allred, D. M.. D E. Beck, and C. D. Jorgensen. 1963a. Biotic communities

of the Nevada Test Site. Brigham Young Univ. Sci. Bull., Biol. Ser..

2 (2): 1-52.
Allred, D. M., D E. Beck, and C. D. Jorgensen. 1963b. Nevada Test Site

study areas and specimen depositories. Brigham Young Univ. Sci. Bull.,

Biol. Ser., 2(4): 1-15.

A NOTE ON GERANIUM RICHARDSONII FISCH. & TRAU TV.

John W. Van Cott'

While examining specimens of the genus Geranium it became
apparent that there were discrepancies between published botanical
descriptions based on herbarium specimens and plants as they occur
naturally. It is always difficult to collect a large specimen and to
arrange it to fit a herbarium sheet. Often, the specimen is separated
into individual stems, or portions are discarded before pressing.
When such an alteration is necessary and when incomplete or altered
specimens are placed in collections, they form the basis upon which
descriptions are made. Such circumscriptions have led to confusion of
students attempting to understand taxonomic entities.

Descriptions of Geranium richardsonii are cited as an example.
In 1907, Hanks and Small (p. 18) described G. richardsonii as
"caudex often slightly branched." Jones and Jones (1943. p. 32)
described it wdth "the usually simple caudex" whereas Harrington
(1954, p. 352) cited that species as "plants . . . erect from a simple
caudex." These three statements are slightly misleading because
the Richardson Geranium is often definitely rhizomatous (Figs. 2,
3, 4) even though a great many specimens appear to have only a
simple caudex (see Fig. 1). Plants such as those appearing in Figures
2. 3, and 4. are often broken apart in such a manner as to appear
like the specimen in Figure 1.

Shaw (1952, p. 302), who did extensive field work on G. richard-
sonii, described it more accurately as having the "caudex often
branched."

It seems probable that the more misleading descriptions cited
above might well have been derived from specimens which lack com-
plete caudices. Of course, there are specimens in herbaria which do
represent the situation as it occurs in nature, but there are a large
number of inadequate specimens and it is these which prompt this
note. Thus, after an examination of some three hundred herbarium
sheets and numerous plants in the field. I present herein a descrip-
tion of the caudex of G. richardsonii.

Caudex clothed with scale-like, brownish, remnants of petioles
and stipules, simple or branched; rhizomes, when present. 3-17 cm
long, one to several from a central, stout, woody. [)erennial rootstock
with a more or less prominent, simple or branched taproot.

Representative Specimens. Arizona: Apache Co., White Mt.
Scout Camp, Deaver 4660 (asc). Coconino Co.. Weatherford Road,
San Francisco Peaks, Carson 268 (asc); road to Weimer Springs. N
of Mormon Mts., Carson 49 (asc) ; NW side of Mormon Mt.. Kynoch
332 (bry). CALIFORNIA: Alpine Co., head of E^st Carson River,

1. Department of Botany, Brigham Young University, Provo, Utah.

25

26

JOHN W. VAN COTT

The Great Basin Naturalist

Vol. XXIX, No. 1

Fig. 1. UTAH. Sanpete Co.: Head of Lake Fork, T19S, R4E, ca. 10,000'. Van Cott

943 (bry).
Figs. 2, 4. UTAH. Garfield Co.: Escalante Canyon. 5 mi E of Widstoe. T34S,

RIW, ca. 8,000'. Van Cott 981a and 981b (bry).

Fig. 3. UTAH. Garfield Co.: 9 mi N of Boulder on Deer Creek, T32S, R5E,

7,500'. Van Cott 987 (bry).

March 28, 1969 note on geranium richardsoni 27

Johnson 213 (usu). Colorado: Boulder Co., y^ mi above Science
Lodge, Murdoch 542 (bry). idaho: Bear Lake Co., Emigration Can-
yon Forest Camp. Shaw 69 (usu); Caribou Co. N of Pruess and
Montpelier Creek Divide, S23, TUS, R45E. Caribou Natl
For, Major 20 (usu). Montana: Beaverhead Co.. U.S. Sheep Exp
Station. Odell Creek, Calvert 102 (bry). utah: Kane Co., left fork
of headwater of Virgin River 15 mi NW of Orderville, Maguire
18823 (bry),- Summit Co., near Lost Lake, Uinta Mts., Patrick 150
(bry); 3 mi west of Holiday Park, Welsh et al. 6305 (bry); San
Juan Co., shady places along Indian Creek 13 mi west of Monti-
cello, Conquist & Holmgren, 9403 (usu).

Literature Cited

Hanks, L. T. & J. K. Small. 1907. Geraniaceae. N. Am. Flora 25(1):3-21.
Harrington, H. D. 1954. Manual of the Plants of Colorado. Sage Books,

Denver. 351-353.
Jones, G. N. & F. F. Jones. 1943. A revision of the perennial species of

Geranium of the United States and Canada. Rhodora 45:5-53.
Shaw, R. J. 1952. A cytotaxonomic studj^ of the genus Geranium in the

Wasatch Region of Idaho and Utah. Madrono 11:297-304.

NEW COMBINATIONS AND A NEW SPECIES OF PERENNIAL

CRYPTANTHA

Larry C. Higgins'

The preparation of a revision of the perennial Cryptantha, sub-
genus Oreocarya of North America, has led to the recognition of
an undescribed species, and to adjustments in nomenclature. Ihe
conclusions presented here are the results of three years work with
this group. Numerous field trips resulting in the collection of several
thousand herbarium sheets has partially filled in some of the gaps
in the distribution and understanding of this subgenus.

The writer wishes to express his thanks to the Society of Sigma
Xi for their grant-in-aid of research, which made it possible to do
the necessary field work for this paper.

1. Cryptantha shackletteana sp. nov.

A C. spiculifera (Piper) Payson. differt foliis longior et angustior,
pilis pustulatis nullis. inflorescentia subcapitata, nuculis lineari-
lanceolatis longior proprietaibus evidentibus minoribus.

Herbae perennes caespitosae. 1-3 dm altae; caules graciles. in-
firmi, 1-plures 0.7-1.8 dm longi, strigosi et setosi effusi pilis gracili-
bus infirmisque; folia lineari 2-13 cm longa. 0.1-0.5 cm lata, strigosa
supra et infra, pilis pustulatis inconspicuis paucis dorsaliter; inflore-
scentia angusta. 0.2-0.8 dm longa; segmenta calycum linearia vel
anguste lanceolata, sub anthesi 3-5 mm longa, sub fructu 7-10 mm
longa. setis flavidis divaricatis; pedicelli 0.5-1 mm longi; corolla
alba, tubo 3-3.3 mm longo cristis ad basi tubi conspicuis, fornicibus
flavis emarginatis 0.5 mm longis, limbo 5-6 mm lato; stylus fructo
exceedens ab 1.3-1.6 mm; nuculae lanceolatae 3.3-3.6 mm longae,
1.6-2 mm latae, pleurmque totae quatuor evolutar, marginibus
acutis, contiguis pagina dorsalis muricata et rugulosa cristis humili-
bus inconspicuisque pagina ventralis similaris sed proprietaibus
evidentibus minoribus; sulcus aperta. subulatus, marginibus sine
elevatibus.

Caespitose perennial herbs. 1-3 dm tall; stems slender, weak,
1-several. 0.7-1.8 dm long, strigose and spreading setose with slender
weak hairs; leaves linear, 0.1-0.5 cm wide. 2-13 cm long, strigose
on both surfaces, and with a few inconspicuous pustulate hairs on
the dorsal surface; inflorescence narrow, nearly capitate. 0.2-0.8
dm long; calyx segments linear or narrowly lanceolate, in anthesis
3-5 mm long, in fruit becoming 7-10 mm long, with yellowish
spreading hairs; pedicels 0.5-1 mm long; corolla white, the tube
3-3.3 mm long, crests at base of tube conspicuous, fornices yellow,
emarginate, 0.5 mm long, limb 5-6 mm broad; style exceeding ma-
ture fruit 1.3-1.6 mm; nutlets lanceolate, 3.3-3.6 mm long, 1.6-2

1. Department of Botany, Brigham Young University. Provo. Utali.

28

March 28, 1 969 new species of cryptantha

29

Figure 1. Habit sketch of Cryptantha shackle tleana showing the narrow
leaves, capitate inflorescence, and both surfaces of a Single Nutlet.

The Great Basin Naturalist
30 LARRY C. HIGGINS Vol. XXIX, No. 1

mm wide, usually all four maturing, the margins acute, in contact,
dorsal surface muricate and rugulose with the ridges low and incon-
spicuous, the ventral surface similar but the markings much less
evident, scar open, subulate, and without an elevated margin.

Type. Locality; Alaska: On steep, southfacing slope of Eagle
Bluff, about one mile northwest of Eagle. Growing with Artemisia
frigida and Agropyron smithii, 26 July, 1968, Stanley L. Welsh and
Glen Moore 8629. Holotype deposited at BRY.

Distribution. Apparently confined to the type locality near Eagle,
Alaska. Growing on serpentine talus slopes, 1,000 to 1,500 feet.
Flowering from June to July.

Specimens Examined. Alaska, USA, on dry slope of ultrabasic
talus, Mission Bluff, Eagle area, 1,300 feet, June 26, 1960. Hansford
T. Shacklette 6183 (US).

Cryptantha shackletteana is probably closely related to C.
spiculifera but differs in the longer and narrower leaves with only
inconspicuous pustulate hairs, the more capitate inflorescence,
longer and narrower nutlets with less evident markings, and weaker
stems.

This species is named in honor of Hansford T. Shacklette, who is
a student of Alaskan Botany and Geology, and who was the first
to collect this distinctive species.

2. Cryptantha subgenus Oreocarya {Greene) stat. nov.

(Based on Oreocarya Greene, Pitt. 1:57-58. 1887.)

Payson included all the species in this subgenus under the section
Oreocarya, but if a sectional name is to be used it cannot be Oreo-
carya, but must be Pseudomyosotis according to the rules. Rather
than use a sectional name all the perennial species have been ele-
vated to the rank of subgenus.

Eritrichium section Pseudomyosotis A. DC. Prod. 10:129. 1846.
Krynitzkia section Pterygium Gray, Proc. Am. Acad. 20:276.

1885 in part.
Krynitzkia section Pseudokrynitzkia Gray, Proc. Am. Acad.

20:276-280. 1885 in large part.
Hemisphaerocarya Brand, Fedde, Rep. Spec. Nov. 24:59. 1927.
Cryptantha section Oreocarya (Greene) Payson, Ann. Mo. Bot.
Gard. 14:237. 1927.

3. Cryptantha fulvocanescens (Gray) Payson var. echinoides (M.E.
Jones) Higgins stat. nov. (Based on: Krynitzkia echinoides M. E.
Jones, Proc. Cahf. Acad. Sci. II 5:709. 1895.

References

Payson, E. B., 1927. A monograph of the section Oreocarya of Cryptantha, Ann.
Mo. Bot. Gard. 14:211-359.

NEW RECORDS AND DIS IRIBU IIONAL NOTES FOR
REPTILES OF THE NEVADA TES I SITE'

Wilmer W. Tanner

In 1963 Tanner and Jorgensen published their study, "Reptiles of
the Nevada Test Site." and listed twenty-eight sj)ecies for the Test
Site. Since then four summer of intensive ecological activity in sev-
eral of the habitat areas of the Test Site have added three additional
species and increased our understanding of others. Two of the added
species were listed as probable species in 1963. but the third was
quite unexpected.

Dipsosaurus dorsalis dorsalis Baird and Ciirard

Three additional specimens have been taken, two from the
Larrea-Franseria community in Frenchman Flat (can traps) and
one IX)R in western Jackass Flat. Although more widely distributed
than originally reported, this species still remains one of those less
commonly observed on the Test Site.

Sceloporus graciosus graciosus Baird and Girard

The northern Sagebrush Lizard reaches its most southern limits
in the uplands of southern Nevada. Records from Pahute Mesa are
now available. In this area we find the Pinyon-Juniper forests with
open areas of desert shrubs of which Artemisia tridentata is usually
jiresent.

.\n ecological point of interest is the occurrence of Sceloporus
occidentalis at the summit of Rainier Mesa well above the distribu-
tion of S. g. graciosus. The three species of Sceloporus at NTS have
little overlapping of their habitat areas. Sceloporus magister is found
in the flats and particularly on the upper bajadas and foothills that
surround the broad valleys. Sceloporus graciosus has been found in
the lower limits of the Pinyon-Juniper Community and Sceloporus
occidentalis seems to occur primarily in the Pinyon-Juniper Com-
munity with the greatest concentrations in the higher elevations such
as Rainier Mesa. In a narrow area along the eastern base of
Rainier Mesa both magister and occidentalis have been seen on
boulders which line the road. There may be other areas where their
habitats overlap; however, it is quite obvious that all three exist for
the most part as single species populations within their own habitat
iiiche.

The comi)lete range of Sceloporus graciosus for the Test Site is
not yet known. Its occurrence in the Pinyon-Juniper-Artemisia

'This study was supported by Research Grant (AT(1 11 ) IWli willi the United States .\toniic
F.nergy Commission.

-Department of Zoology and Entomology. Brighani Young I'niversity, Provo, Utah.

31

The Great Basin Naturalist
32 WILMER W. TANNER Vol. XXIX, No. 1

plant associations would indicate its occurrence wherever these
plants predominate. This would place the range in the northern
parts of NTS and extending northward into central Nevada.

Morphological (meristic) characters of the small series available
does not indicate any important differences from those series seen
from central Nevada and southwestern Utah.

Leptotyphlops humilis Baird and Girard

In spite of extensive field work and can traps placed in what
appears to be favorable habitat, only one specimen has been found.
On May 19, 1964, one specimen (BYU 23929) was taken in Rock
Valley (NTS) by A. P. Aschwanden. It has a total length of 282 mm
and a snout- vent length of 270 mm. There are 14 rows of dorsal
and 282 scales in the dorsal row; head scales are normal except
that the fifth dorsal is noticeably wider than is usual in humilis.

The nearness of NTS to southwestern Utah and the fact that a
specimen taken at Indian Springs (Clark Coun,ty) was identified
as utahensis (La Rivers, 1942) leads one to suspect that the sub-
species utahensis should be at the test site. An examination of the
scale counts as listed above, however, places this specimen as an
intergrade humilis x utahensis. The dorsals fall between these two
subspecies {humilis 265-280; and utahensis 289-308) and the fourth
dorsal is not divided but the fifth is much wider than the sixth.
Such intermediate characters indicate that one might expect to find
at NTS individuals which would key to either subspecies or be
intermediate.

Diadophis punctatus regalis Baird and Girard

An adult female (BYU 31287) was taken on July 14, 1968,
along Holmes Road one mile from its junction with Stockade Wash
Road, Rainier Mesa, by James M. Hopkin and Wilmer W. Tanner.
Its peculiar behavior of coiling its tail so as to show the bright
Carnelian Red color (Ridge way 1912) on the caudals led to its
capture. The snake was actually on the shoulder of the road; how-
ever, as we approached, the vibration apparently initiated the above
behavior permitting Mr. Hopkin to see it as we passed.

The color and color pattern does not vary from the few live
specimens seen from southwestern Utah and is the same as pre-
viously reported by Tanner (1952) for a specimen from Washing-
ton County, Utah. The ventral scale count of 235 is noticeably
higher than has been observed in previous specimens from Nevada
and Utah, however, the caudal count is lower thus producing a
ventral-caudal total of 301 scutes. This is comparable to other
specimens seen from the range of regalis.

This record extends the range for this species approximately 250
miles southwest from its previous record near Caliente. Nevada
and for at least 300 miles west of the nearest locality in north-
western Arizona.

March 28, 1969 hi.ptii.i-.s oi- thk nkvaua tkst site 33

The apparent extensive range of this species seems to bo re-
stricted to those habitats associated with the scrub oak, l^inyon-
Juniper or some combination of these plant comnmnities. In
western Utah and adjoining eastern Nevada these plant communi-
ties occur jjrimarily between 4.500 to 7,000 feet elevation. At
the Nevada Test Site such plant communities occur at higher ele-
vations with the Pinyon-.Iuniper at a near climax on the higher
mountains and mesas.

This specimen was found on the south sloj)e of Rainier Mesa
at an elevation of approximately 6500 feet and well within the
Pinyon-Juniper community. On the basis of its occurrence on
Rainier Mesa, I would expect it to be found or at least to inhabit
other similar adjoining areas, such as Pahute Mesa, and to extend
north and northeast into central and eastern Nevada.

Masticophis taeniatus taeniatus Hallowell

In 1963 only three specimens of this species were recorded for
the Test Site. At that time it was indicated that the scarcity of
records was undoubtedly due to the lack of collecting in the more
desirable areas. During the period from 1965 to 1968 (3^2 sum-
mers) only three additional s[)ecimens have been taken and two
seen DOR.

All records are from areas in or near the Juniper-Pinyon pine
habitat. Our records would indicate that this species (subspecies)
is at its southern limits in the higher elevations of the Nevada Test
Site and we do not expect to find it in the deseret valleys (French-
man. Yucca, or Jackass Flats). Although this species is not what
we might call a rare species, it is not a common one and may not
be seen daily even though we traverse its habitat area.

Phyllorhynchus decurtatus parkinsi Klauber

We reported the first record of this species for Nye County.
Nevada, in 1963 and indicated on the basis of the few specimens
available (four) that it was perhaps one of the less common, species
on the Test Site. In the last few years, however, we have found
this species to be one of the more common, particularly in the lower
valleys. Certain areas in Frenchman, Mercury, and .Jackass Flats
have produced large series of s[)ecimens. We have noted that our
greatest number of observations have occurred in those habitats
where Larrea occurs as a part of the vegetative cover. It is now
possible for us to provide a range of variation for some of the more
common scale characters and these are as follows: Ventrals 9 9 .
10(178-190)183.5; caudals. 10(26-30)28.0. Ventals dd- 16(165-
176) 170.9; caudals, 16(33-40)37.5. In both the males and the
females it will be noted that the above scale counts average less
than those reported by Klauber (1935). The color pattern is es-
sentially as reported for NTS specimens in 1963.

The Great Basin Naturalist
34 WILMER W. TANNER Vol. XXIX, No. 1

Hypsiglena torquata deserticola Tanner

In 1963 a single specimen was reported for the Test Site. Since
1965 a series of this species has been found by driving the roads
in the evenings. Present records show it not to be a rare species,
particularly in the lower foothill and bajada areas of the Test Site.
The scale counts and color pattern, especially the dorsal spots and
the nape spots, are essentially as reported by Tanner (1946).

Trimorphodon lambda Cope

A second specimen of this species was taken approximately one
mile north of the junction between the NLRD road and the con-
necting road from Mercury. It is a male with approximately the
same scale counts as reported previously. We have made a de-
termined effort to find additional specimens of this species using
methods which have been successful in other areas by other workers.
We, therefore, must conclude that it really is a rare species at the
Test Site and that it perhaps is very near its most northern dis-
tribution limits.

Tantilla p. utahensis (Blanchard)

A third specimen of this species was taken at Cane Springs in
a can trap, 31 August, 1968. It is an adult male specimen with
scale patterns that fit nicely into the averages as presented by Tanner
in 1966. We are still not certain as to the abundance of this species
but suspect that it is more common than our collecting records
would indicate, particularly in the rocky foothills and gravelly
bajadas. This specimen was taken on a rocky northeast slope.

Literature Cited

Klauber, L. M. 1935. Phylorhynchus, the leaf-nosed snake. Bull. Zool. Soc.

of San Diego, No. 12:5-31.
LaRivers, Ira. 1942. Some new amphibian and reptile records for Nevada.

Jour. Entomol. and Zool., 34(3):53-68.
RiDGWAY, Robert. 1912. Color standards and color nomenclature. Washington,

D. C. (published by author), 43 pp., 53 pis.
Tanner, W. W. 1946. A taxonomic study of the genus Hypsiglena. Great

Basin Nat. 5:25-92.
Tanner, W. W. 1952. Diadophis regalis regalis (Baird and Girard) found in

Nevada. Great Basin Nat., 12(1-4) : 63-64.
Tanner, W. W. 1966. A reevaluation of the genus Tantilla in the south-
western United States and northwestern Mexico. Herp. 22(2): 134-150.
Tanner, W. W. and Clive D. Jorgensen. 1963. Reptiles of the Nevada Test

Site. Brigham Young University Sci. Bull., 3 (2): 1-35.

THE EFFICACY OF IIALOXON AND THIABENDAZOLE

AS ANTHELMINTICS AGAINS I' GASTROINTESTINAL

NEMATODES IN SHEEP

Ferrou L. Andersen^, Keith H. Hoopes-, and J. Cail Fox'

Introduction

Since many species of parasitic nematodes are picked up by
the host grazing on contaminated pastures, control measures are
generally dir€K-ted towards proper range or pasture management as
well as tow^ards an appropriate animal husbandry program. In
most areas where severe parasitism occurs, livestock owners must
treat their animals several times each year with anthelmintic
drugs in order to keep the animals as free of parasites as possible.
Some of the problems which relate to drug treatment are not only
the cost of the drugs and of handling the livestock several times each
year, but also the fact that drug resistant strains of parasites fre-
quently develop to routinely-used drugs.

Examination, in the fall of 1967, of the sheep kept at the BYU
Research Farm at Provo. Utah, indicated that gastro-intestinal
parasitism by certain nematodes could represent a serious problem.
One yearling lamb died as a result of natural infection of the large
stomach worm of sheep, Haemonchus contortus, on 22 September.
1967. This lamb had a count of 5,200 eggs per gram (e.p.g.) of
feces, and exhibited other symptoms of haemonchosis. including
edema of the head and thoracic and abdominal cavities. Other
lambs killed on different experiments at this same approximate
time also had several hundred H. contortus adults in the abomasum.

Since all of the sheep at the farm had been treated routinely
during the past eight years with low levels of phenothiazine mixed
in the feed, we decided to test the anthelmintic efficacy of Thiaben-
dazole* and Haloxon**. two drugs which had not been used pre-
viously at the BYU Research Farm.

Procedure and Results

On 30 September, 1967, we obtained fecal samples from 107
sheep maintained at the farm. Some of these sheep had been on

'As'iix iiite l'rf)fessor, Department of Zoolopv and Entomology, Brighani Young Univcrsilv.
I'lovo. rt;,l.

-.\s-()< uitc I'rofessor, D.V.M., Dei>artnient of Animal Science, Brigliani Yoimg University,
Provo. Ii.iIk

■'(iiiiiliialc Sliideiil. Dopiirtment of '/.oolonv ami Knloinologv. Urigliam doling I'mversilv.
I'rovo, llali

"Tliiahenda/.ole i-i an imidazole derivative with the chemical foniuila of J-(4"-thia/olyl )■
hen/imidazole, and a i>rodurt of Merck and Company. Hahway. New Jei-sey.

"Haloxon is an organophosphate compound with the chemical fonuula of O.Ocli( Jchloroethyl)-
0-( i-chloro-4-methylcoumarin-7-yl) phosphate, and a product of William Cooper and Nephews,
Inc., (Chicago. Illinois.

This research was supported in part liv a grant-in-aid from William Cooper & Nephews.
Inc., Chicago, Illinois, through Dr. Paul V. Christofferson, D.V.M., Veterinary Director.

35

The Great Basin Naturalist
36 ANDERSON, HOOPES, FOX Vol. XXIX. No. 1

irrigated grass pasture during the entire summer, while others had
been used on nutrition experiments, and had been held on drylot
only. Those on drylot had only small numbers of eggs or none in
the feces. Of those held on pasture, we were able to identify 25
adult sheep that had egg counts varying from 200 to 3500 tricho-
strongylid eggs per gram of feces. For the purpose of the experi-
ment, these sheep were divided into three groups on the basis of the
egg count. Group A. consisting of 8 sheep, had an average e.p.g.
of 1028 (range: 200 to 2100) and on 19 October, 1967, each animal
was given, by dose syringe. 1 fluid ounce of Thiabendazole mixture,
containing 2 gm Thiabendazole per fluid ounce. Group B, consisting
of 9 sheep, had an average e.p.g. of 1230 (range: 200 to 3500) and
on 19 October, 1967, each animal received 1 fluid ounce of Haloxon
containing 1.4 gm Haloxon per fluid ounce. Drug dosages were
based on manufacturers' recommendations. Group C, consisting of
8 sheep, which had an average e.p.g. of 1300 (range: 200 to 3400)
served as the control group. Fecal samples were collected again
12 days after treatment was given. At that time the average e.p.g.
in Group A (Thiabendazole-treated) had dropped to 8 (range: 0 to
100), a 99% reduction in total egg count. Group B (Haloxon -treated)
had dropped to 19 (range: 0 to 100), a 98% reduction, and the
control sheep had also dropped to an average e.p.g. of 275 (range:
0 to 900). a 79% reduction. The average e.p.g. counts for all three
groups of sheep prior to and 12 days after treatment are listed in
Table 1.

In order to determine which genera of nematodes were affected
by the drugs, larval cultures were made from all fecal samples
from the 25 sheep prior to and after treatment. Table II shows the
numbers of larvae examined and their assignment to various genera,
based on the tables prepared for total length and sheath tail length
by Dikmans and Andrews (1933) and Keith (1953). Our ranges
of total and sheath tail measurements differed slightly from the
tables we were using, but the identification of Haemonchus,
Chabertia. Ostertagia, Trichostrongylus, and Nematodirus was con-
sidered valid.

From the results we obtained, it was noted that Thiabendazole
was highly effective against Trichostrongylus and Ostertagia, since
no larvae of these genera were detected after treatment. The per-
cent of Haemonchus larvae constituted 67% of the larvae identified
before treatment and 27% of those identified after treatment. The
percentage of Chabertia larvae increased from 4% prior to treat-
ment to 47% after treatment, and the percentage of Cooperia rose
from 7% to 20%. Haloxon appeared completely effective in
eliminating Haemonchus, Cooperia and Ostertagia, but not as effec-
tive against Trichostrongylus as was Thiabendazole. The percentage
of Trichostrongylus in the Haloxon-treated group increased from
14% prior to treatment to 21% after treatment. Oesophagostomum
rose from 2% to 5%, and 68% of all larvae identified after treat-
ment from the Haloxon-treated group were identified as Chabertia,

March 28, 1969

NEMATODES IN SHEEP

37

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The Great Basin Naturalist
38 ANDERSON, HOOPES, FOX Vol. XXIX. No. 1

whereas none had been identified in that group prior to treatment.
The most significant change in the percentages of the various
genera in the control sheep was a decrease in Haemonchus from
68%, prior to the time the two groups were treated, to 39% when
the control sheep were sampled afterwards. The percentages of
larvae identified as Trichostrongylus and Cooperia showed corres-
ponding increases during this period.

Discussion and Conclusions

Although the variation in nematode egg counts was considerably-
greater than would have existed had we infected the sheep experi-
mentally with equal numbers of infective larvae, this project,
nevertheless, showed that both Thiabendazole and Haloxon are
effective anthelmintics against the majority of gastro-intestinal
nematodes encountered. It was unfortunate, however, that the
problem was identified and started so late in the pasture season,
since the marked drop in egg counts in the control sheep undoubted-
ly related to decreased numbers of infective larvae being picked
up by the grazing sheep at that time of the year. Haemonchus
contortus larvae in particular are highly susceptible to colder tem-
peratures as demonstrated by the seasonal e.p.g. decline in the
control animals, and generally do not develop from the egg to the
infective larval stage at mean monthl}^ temperatures of less than
65° F, or when the total monthly rainfall is less than 2 inches per
month (Levine. 1963; Kates, 1965). The mean monthly tempera-
tures in a standard weather shelter in Provo for June. July. August.
September, and October, 1967, were 62.3, 73.3, 72.3. 62.6. and 47.9 F
respectively. The total precipitation for those five months was
1.26, 0.96, 0.16. 0.77. and 0.55 inches, respectively. Therefore, the
conditions for the development and survival of the free-living
stages of Haemonchus, as far as available moisture was concerned,
would appear to be considerably less than optimum for these months.
The precipitation for these months in 1967 did not differ markedly
from the long-term averages for Provo for the four main pasture
months of June, July. August, and September, since these are all
less than 1 inch per month. During this same period, however,
farmers in this area put as much water on their lands through irri-
gation as that comparable to 20 inches of rainfall. Some of the
lowland pastures are frequently completely saturated by irrigation
water that may be allowed to run for several days in any one
pasture location. These conditions of moisture, coupled wdth the fact
that the soil temperature measurements may actually be 15-20° F
higher than that recorded in a standard weather shelter (Andersen
and Levine, 1967) certainly are capable of producing micro-
environmental conditions optimum for the development and survival
of these nematode larvae. Nevertheless, when the project herein
reported was conducted during the late pasture season, the number
of larvae available on the pasture would naturally^ be decreasing

March 28. 1969 nematodes in sheep 39

duo to low lenipenituros. On iho 1 ^tli and 14th of Sopteniher the
minimum daily temperatures in the weather shelter were only 29
and 32° F, respectively, and 6 of the 16 remaining days in Septem-
ber had minimum temi)eratures less than 40° F. These low tem
peratures. undoubtedly, would decrease the numbers of viable
larvae on the pasture, thereby resulting in fewer new infections in
the grazing sheep. Had the total egg counts from the control sheep
remained at the same approximate level as that before treatment, the
efficacy of the two anthelmintics tested would have been much
clearer.

Since the numbers of larvae recovered from fecal samples fol-
lowing treatment was very limited in the two treated grou{)S. the
comparison of the effect of the drugs on the specific genera as
determined by larval identification must be interpreted provisional-
ly. No more than 100 eggs per gram of feces was noted in any of
the treated sheep. With such small numbers, the recovery of suf-
ficient numbers of larvae for identification after laboratory culture
is very difficult, and only a total of 54 infective larvae were iden-
tified from these two treated groups. Nevertheless, as mentioned
above, the identification of most of the genera was considered valid.
Infective Ilaemonchus contortus larvae are particularly easy to
identify and the fact that none of these were found in any fecal
samples taken from the Haloxon-treated sheep seemed quite signifi-
cant.

As stated above, one of the main difficulties in using continual
chemotherapy as a control measure in parasitism is the frequent
development of strains of the parasites resistant to the chemicals
being used. Phenothiazine, for example, was first introduced as an
anthelmintic in swine by Harwood, Jerstad and Swanson in 1938,
and then by a number of authors in 1939 (See Gibson, 1965) as
an anthelmintic in sheep. Early rej)orts showed this drug to be
75-90% effective against Haemonchus when used in crude prepara-
tions, and later up to 100% effective when used in a micronized-
powder form. Strains of Haemonchus resistant to phenothiazine
were first reported in 1957 in Kentucky by Drudge, Leland, and
Wyant, and later noted by such authors as Enzie et al. (1960) in
Maine, and Levine and Garrigus (1961) in Illinois. In the latter
case, phenothiazine had been used routinely at the sheep division
at the University of Illinois for 19 years before the resistant strain
was identified.

Thiabendazole was first introduced as an anthelmintic in sheep
by Brown et al. in 1961. This drug proved to be an extremely
effective broad-spectrum anthelmintic and has been used widely
in many geographical locations for treatment of parasitic helminths.
Efficacy against Haemonchus in sheep ranged as high as 96-100%
(Hebden, 1961; Gordon. 1961). Since that time, however, strain
resistance has appeared against this drug also. Smeal ct al. (1968)
recently reviewed accounts of resistance to Thiabendazole that have
been noted in the United States and Brazil, as well as the work

The Great Basin Naturalist
40 ANDERSON, HOOPES, FOX Vol. XXIX, No. 1

they reported on for Australia. Since 27% of the larvae recovered
after treatment in the Thiabendazole-treated animals in our project
were Haemonchus^ there is at least some suggestion that the
Haemonchus in this geographical area are also somewhat resistant
to this drug. Since the drug has not been used at the BYU Research
Farm, however, such a conclusion would imply that resistant
strains have been brought to this area by sheep that have been
purchased elsewhere.

The efficacy of Thiabendazole against parasitic helminths has
been reported in several hundred scientific articles, and annotated
bibliographies are available on this subject from Merck and Com-
pany, Inc., Rahway, New Jersey. Haloxon has not been tested as
widely in the United States, but its efficacy against several genera
of gastro-intestinal nematodes in sheep as well-documented by such
authors as Armour, Brown and Sloan (1962), Harbour (1963),
Barnett, Berger, and Rodrigues (1964). Turk, Galvin and Bell
(1965), and Baker and Douglas (1965). In general, these workers
found Haloxon to be a highly efficacious anthelmintic against most
of the same genera of parasitic nematodes as was Thiabendazole,
with the exception that it was not as effective against such genera
as Chabertia, the large-mouthed bowel-worm, or Oesophagostomum,
the large intestinal nodular worm. Haloxon is apparently hydro-
lyzed before it reaches the large intestine, and is therefore more
effective against nematodes which occur in the abomasum and
small intestine. We also found that Haloxon was not effective
against Chabertia and Oesophagostomum^ although as stated earlier,
the small numbers of larvae identified suggests that this conclusion
be interpreted provisionally.

Although comparative studies on the efficacy of phenothiazine
were not included in this project, the fact that the animals had
been treated routinely with phenothiazine for the past eight years,
and yet the fact that severe haemonchosis can occur here, also
suggests that strains resistant to this drug exist in this area. Experi-
ments involving critical testing where worm counts and speciations
can be made at necropsy after treatment with Thiabendazole,
Haloxon, and phenothiazine should be conducted, before the problem
of possible resistant strains in this area can be resolved.

References

Andersen, F. L., and N. D. Levine. 1967. Methods and problems in micro-
environmental measurements. 111. Vet. 10:10-17.

Armour, J.. P. R. M. Brown, and J. E. N. Sloan. 1962. The anthelmintic
efficacy of haloxon against the adult and immature stages of gastro-intes-
tinal nematodes. Vet. Rec. 74:1454-1457.

Barnett, S. F., J. Berger, and C. Rodrigues. 1964. A small controlled trial
of the anthelmintics haloxon and thiabendazole in sheep. Afr. Agr. For. J.
29:195-198.

Baker, N. F., and J. R. Douglas. 1965. Anthelmintic activity of haloxon in
lambs. Am. J. Vet. Res. 26:651-653.

March 28, 1969 nematodes in sheep 41

Brown. H. D.. A. R. Mat/uk, 1. R. Ilves, I,. H. Peterson, S. A. Harris, L. H.
Sarett, J. R. EfiERTON. J. J. Yakstis, W. C. Campbem,, and a. C. Cuckler.
1961. Aiitipiuasitic drugs. IV. 2-(4'-thia7,ol3'l)-benziniidazole, a new an-
thelmintic. J. Am. Chem. Soc. 83:1764-1765.

Dikmans. G. and J. S. Andrews. 1933. A compxarative morphological study of
the infective larsae of the common nematodes parasitic in the alimentary
tract of sheep. Tr. Am. Micr. Soc. 52:1-25.

Drudge, J. H., S. E. Lei-and, Jr., and '/. N. Wyant. 1957. Strain variation
in the response of sheep nematodes to action of phenotfiiazine. II. Studies
on pure infections of Haenionchus contortus. Am. J. Vet. Res. 18:517-325.

Enzie, F. D.. M. L. Colglazier, G. E. Wiiitmore, and D. E. Thompson. 1960.
Effectiveness of tliree management systems on parasitism in lambs. III. Re-
sistance of Haenionchus contortus to phenothiazine. J. Parasit. 46 (suppl.):
41.

Gibson. T. E. 1965. Veterinary Anthelmintic Medication. Technical Communi-
cation No. 33 of the Commonwealth Bureau of Helminthology. St. Albans,
Herts. 206 pages.

Gordon, H. McL. 1961. Thiabendazole; a highly effective anthelmintic for
sheep. Nature 191:1409-1410.

Harbour, H. E. 1963. Expeirence with organo-phosphorus compounds as ant-
helmintics. Internat. Vet. Cong. pp. 751-756.

Hl^RwooD, P. D., A. C. JERST.^D, AND L. E. Swanson. 1938. The efficacy of
phenothiazine for the removal of ascarids and nodular worms from swine.
J. Parasit. 24:16-17.

Hebden, S. p. 1961. The anthelmintic activity of thiabendazole (MK-360).
Austral. Vet. J. 37:264-269.

K.\TES. K. C. 1965. Ecological aspects of helminth transmission in domesticated
animals. Am. Zool. 5:95-130.

Keith. R. K. 1953. The differentiation of the infective larvae of some common
nematode parasites of cattle. Austral. J. Zool.. 1:223-235.

Levine, N. D. 1963. Weather, climate, and the bionomics of ruminant nema-
tode larvae. Adv. Vet. Sci. 8:215-261.

Levine, N. D., and U. S. Garrigus. 1961. The effect of Cu-Nic and organc^-
phosphorus anthelmintics on phenothiazine-resistant Haenionchus in sheep.
Am. J. Vet. Res. 23:489-493.

Sme.^l, M. G.. p. a. Gough. a. R. Jackson, and I. K. Hotson. 1968. The
occurrence of strains of Haenionchus contortus resistant to thiabendazole.
Austral. Vet. J. 44:108-109.

Turk, R. D.. T. J. Galvin. and R. R. Beli,. 1965. Comparative tests of the
anthelmintics haloxon, phenothiazine and thiabendazole in sheep and goats.
Southwest. Vet. 19:44-47.

NOTES

LEPIDOPTERA OF THE NEVADA TEST SITE^

During ecological investigations by Brighani Young University at the Nevada
Test Site between 1959 and 1965, many moths and butterflies were taken, mainly
by incandescent and ultraviolet light-traps. Many of these were submitted to Dr.
Jerry A. Powell, University of California at Berkeley, who kindly made some
general as well as specific identifications as a basis for further studies with this
difficult group of insects. These are reported herein. All listings are alphabetically
arranged for convenience.

Arctiidae

Arachnis prob. picta

Pygaractia murina
Coleophoridae

Heliodines nr sexpunctella

Scythris (12 spp.)
Cosmopterygidae

Inga concolorella
Geometridae

Caripeta sp.

Glaucina sp.

Phaeoura sp.

Semiothisa nr colorata

Stenospilates sp.
Lasiocampidae

Gloveria arizonensis
Noctuidae

Agrotis ypsilon

Conochares nr arizonae

Conochares nr hutsoni

Grotella sp.

Oxycnemis nr gtacillima

Phobolosia anfracta

Synedoida sp.

Triocnemis sp.
Papilionidae

Cercyonis sp.

Pieris protodice
Plutellidae

Abebaea sp.

Cerostoma nr angelicella

Cerostoma nr delicatella

Cerostoma nr- flavistrigella

Plutella maculipennis
Pyralidae

Dichozoma parvipicta

—Donald M. Allred'

'BYU-AEC publication no. COOl 355-20.

Dioryctria nr gulosella
Etiella zinckenella
Eumysia mysiella
Heterographis morrisonella
Hulstia undulatella
Jocara sp.

Loxostege albiceralis
Nephopteryx bifasciella
Ommatopteryx texana
Passadena flavidorsella
Salebriacus odiosellus
Sosipatra rileyeUa
Staudingeria albipenella

Sphingidae

Celerio lineata
Sphinx dollii

Tineidae

Acrolophus laticapitanus
Acrolophus variabilis
Acrolophus (4 spp.)
Dystospasta yumaeella
Myrmecozela nr obliquella
Tinea sp.

Tortricidae

Decodes fragarianus
Eucosma bobana
Eucosma nr bolanderana
Eucosma rorana
Ofatulena duodecemstriata
Phaneta indagatricana
Phaneta setonana
Platynota labiosana
Platynota nr yumana

Unplaced

Adela punctijerella

42

March 28, 1969 43

WISLOUCHIELLA PLANCTONICA SKVORTZ. (CHLOROPHYTA.
VOLVOCALES), A NEW ALGAL RECORD FOR NEVADA

III 1925, Skvort/.uvv described a iiioiiotyi)ii genus of Infldf^ellated green
algae as W islouchiella planctonica, and according to SniitJi (IQ'jO). the species
is known only from California and a few sites in the states of the Mississippi
River Valley. The genus is quickly distinguished by its strongly compressed lorica
witli broad wing-like expansions on both sides of the massive cup-shaped chloro-
plast. Pour projections may be seen; two arise at the proximal end and extend
upwards and for-ward slightly beyond the apex of the lorica while the two distal
jirojections extend upwards and ba( k from an insertion point near the base of the
protoplasm. It is these projections and the wing-like expansions of the compressed
lorica that characterize the genus W islouchiella.

The Nevada record is based on Reveal 1570, collected at The Reservoir, about
0.5 miles nortli of the Buckboard Mesa Road and about 2 airline miles northwest
of VABM Tippipah at an elevation of 5.200 feet, north of Shoshone Mountain.
Nye Co., Nevada, 16 July 1968. and deposited in the algal collection at Brigham
Young University. This site is a recent man-made reservoir and the collection
was made during a dense algal bloom; only the one species was found. — James
L. Reveal '•'

References

Skvortzow, B. B. 1925. W islouchielle planctonica nov. gen. et spec, of Vol.

vocales. Proc. Sungari River Biol. Stat. 1:27-29.
Smith, G. M. 1950. The freshwater algae of the United States. McGraw-Hill

Co., New York, pp. 89-90.

'Laboratory of Nuclear Mediiiiie and Kadialioii Biology, University of California. I.os Angeles.

California 90024. Present address: Department of Botany, Brigham Young University, I'rovo, Utah
84«01 .

-Work perfonned under Contract No. AT (04-1) Gen-12 between the Universitj' of (California

and the Division of Biology and Medicine. United States Atomic Energy Commission.

44

Vol. XXIX. No. 1

THE INFECTION OF IMMATURE AQUATIC INSECTS
BY LARVAL PARAGORDIUS ( NEMATOMORPH A )

Most of the hosts lecorded for immature Pragordius sp. have been terrestrial
insects (Carvalho, 1942; Hyman, 1951). The discovery of an infection of larval
Paragordius in the mayfly Baetis sp. (White, 1966) led me to e.xamine more
closely the other aquatic insects collected from Lawrence Creek, Adams Co.,
Wisconsin.

Fifty bottom samples, V^m', collected each month during 1966 supplied the
aquatic insects. Aquatic insects found to be infected with larval Paragordius are
listed in the following table:

Table I

List of aquatic insects from Lawrence Creek, Wisconsin,

infected with larval Paragordius sp, 1966.

% Collection

Family

Genera

Months

Infected

Baetidae

Baetis spp.

IV,

V

2.1

Leptophlebiidae

Leptophlebia sp.

IV,

V

2.3

Ephemerellidae

Ephemerella spp.

IV,

V. VI

4.1

Corixidae

Sigara sp.

IV

1.5

Brachycentridae

Brachycentrus sp.

IV,

V, VI

1.2 no pupation

Simuliidae

Simulium spp.

IV,

V, VI, VII

2.4 no pupation

Of the 83 genera of aquatic insects examined, 6 genera of 4 orders were
found to contain lai-val Paragordius. The larval neniatomorphs all appeared to
be similar. The characteristics of larval nematomorphs are not well known but
only Paragordius varius have been collected from Lawrence Creek.

Individuals that were parasitized showed several morphological variations
from the non-parasitized individuals of the same genera. The wing pads of the
Ephemeroptera nymphs were smaller than the average while parasitized. Simu-
liidae larvae were larger than average during June and July. The parasitized
Simulliidae and Brachycentridae could not be induced to pupate in the laboratory
while the non-parasitized larvae pupated readily.

From these observations I conclude that larval Paragordius can infect several
different orders of immature aquatic insects of both holometabolous and hemi-
metabolous development. — David A. White, Assistant Professor of Zoology and
Entomology, Brigham Young University, Provo, Utah.

References

Carvalho, J. C. M. 1942. Studies on some Gordiacea of Noi-th and South

America. J. Parasit., 28:212-222.
Hyman, L. H. 1951. The Invertebrates. Vol. III. Acanthocephala Aschelmin-

thes and Endoprocta. McGraw-Hill, New York, pp. 455-473.
White, D. A. 1966. A new host record for Paragordius varius (Nematomor-

pha). Trans. Amer. Microsc. Soc. 85:579.

(or7au o

The

Volume XXIX, No. 2
July 31, 1969

MUS. COMP. ZOOLL
LIBRARY.

MAR 11971

HARVARD
UNIVERSITY

Great Basin

mmmim

Published by
Brigham Young University

GREAT BASIN NATURALIST

Editor: Vasco M. Tanner, Department of Zoology and Entomology
Brigham Young University, Provo, Utah

Associate Editor: Stephen L. Wood, Department of Zoology and
Entomology, Brigham Young University, Provo, Utah

Members of the Editorial Board:

Ferron L. Andersen (5), Zoology

Jay V. Beck (3) , Bacteriology

Robert W. Gardner (1), Animal Science

Joseph R. Murdock (4) , Botany

WiLMER W. Tanner (2) , Zoology, Chairman of the Board

Stanley L. Welsh ( 1 ) , Botany

Ex officio Members:

A. Lester Allen, Acting Dean, College of Biological and
Agricultural Sciences

Ernest L. Olson, Chairman, University PubUcations, Uni-
versity Editor

The Great Basin Naturalist

Published at Provo, Utah by
Brigham Young University

Volume XXIX July 31, 1969 No. 2

TTTE ECOLOGY OF THE WESTERN SPO ITED FROG,

RAN A PRETIOSA PRETIOSA BAIRD AND GIRARD,

A LIFE HISTORY STUDY

Ronald L. Morris and Wilmer W. Tanner^

Introduction

Numerous individuals have assisted in the preparation and com-
pletion of this study. We are indebted to Mr. Lawson Hamblin for
allowing free access to his property where the main portion of this
study was conducted, and to Dr. Frederick B. Turner for his sugges-
tions and review of the manuscript.

We also thank Mr. David F. Avery for his help and for the
data he furnished for the years 1962 and 1963. and also for photo-
graphs taken by him. Thanks also to the other students and staff
who have helped in data analysis.

Since the turn of the century, the study of amphibian life history
has received more attention than before; however, the natural his-
tory of many amphibians is still only partially known. Much of our
knowledge is based upon laboratory studies where observations have
been made on eggs obtained directly from the female as described
by Rugh (1948) and as used by Johnson (1965). or directly from
the pond as reported by Skousen (1952, unpublished). These types
of studies provide for contn)lled conditions thereby giving a low
variance to the results. They can also give an actual growth curve
rather than an estimated one since the same individuals can be
studied continually throughout their larval development. 1 lowever,
as stated by Bragg (1940a), to understand the reactions and behavior
of an animal within its complex environment, one must still of nec-
essity go directly to nature. Therefore, this study is based on an
em{)irical approach.

Review ok Literature. Since the description of Rana prctiosa
Baird and Girard. only limited life history accounts have appeared

'Departnicnl of /oolnpv and Kntoinology, Biigliaiii Young University, I'rovo. Ulali.

45

The Great Basin Naturalist
46 MORRIS AND TANNER Vol. XXIX. No. 2

in the literature as compared to most other species of this genus in
North America. Most references are small notes relating to distribu-
tion, habitat, or stomach content, and refer to populations occurring
in Oregon, Washington. British Columbia, Wyoming, Idaho, and
Utah. A brief summary is given by Stebbins (1951 and 1954).

Several accounts of R. pretiosa have appeared since the reports
by Stebbins. Turner (1957) did a four year study from 1953-1956
on the ecology and morphology of Rana pretiosa pretiosa at Yellow-
stone Park in Wyoming, and Johnson (1965, unpublished) worked
on the early development, embryonic temperature tolerance, and
rate of development of Rana pretiosa luteiventris Thompson, from
central Oregon. Rana pretiosa has also been included by Dunlap
(1955) and Dumas (1966) in excellent works on the genus Rana.
The influence of nerves in limb regeneration was studied by Thorn-
ton (1956) on pretiosa tadpoles taken at Moran, Wyoming.

Other earlier papers include Svihla's (1935) brief report on the
eggs and tadpoles of pretiosa in Washington; Middendorf's (1957)
observations on the frog's early spring activities in Montana; Car-
penter's (1953a) brief ecological notes on pretiosa in the Grand
Teton-Jackson Hole area of Wyoming, and his (1953b) notes on the
aggregation behavior of the tadpoles. In a later study on amphibian
movement. Carpenter (1954) found that there is a tendency for in-
dividuals to return to their original point of capture. Dunlap (1959)
described briefly some morphological characteristics of pretiosa
found in Deschutes County. Oregon.

The first published report of amphibians in Utah was by Yarrow
(1875); however, no comprehensive work was done on the popula-
tions of Utah Rana pretiosa until Tanner (1931) included them in
his work entitled, "A Synoptical Study of Utah Amphibia." The only
other reports to include this species are those of Van Denburgh and
Slevin (1915), Slevin (1928) who provides a description, and Skou-
sen (1952) in which he characterized the eggs and larvae of Utah
amphibians.

Taxonomic Status

Rana pretiosa was first described from Puget Sound, Washington,
in 1853 by Baird and Girard. Rana pretiosa luteiventris was later
described by Thompson (1913) from Anne Creek, Elko County, Nev-
ada. Although R. pretiosa has been recognized as a valid species since
its description, the status of the subspecies luteiventris has been ques-
tioned by several writers. Slevin (1928), for example, did not separ-
ate R. p. luteiventris from R. p. pretiosa. He based this decision upon
the work of Van Denburgh and Slevin (1915) which stated that it
was not possible to find constant differences in plantar or palmar
tubercles between those specimens thought to be R. p. luteiventris
from Utah and those R. p. pretiosa from Fort Klamath, Oregon, and
Mount Rainer, Washington. Storer (1925) also questioned the valid-
ity of the subspecies luteiventris as did Stebbins (1951) when he
stated, "it appears to be no more than a slightly differentiated sub-

July 31, 1969 ecology of r. pretiosa in utah 47

species of possibly very local occurrence." He did, however, recognize
the subspecies luteiventris at this time. In a later work Stebbins
(1954) did not recognize luteiventris. and in his latest work (1966)
makes no mention of it.

Dunlap (1955) did recognize the two subspecies in his work on
the variation within the genus Rana and states:

R. p. luteiventris may be distinguished from R. p. pretiosa by the dif-
ference in coloration and by the foot tubercles. The bright color on the
ventral surface is orange-yellow in adult R. p. luteiventris, and bright
salmon-red in adult R. p. pretiosa. R. p. luteiventris, furthermore, lacks
the tubercle at the base of the fourth toe, which is characteristic of
R. p. pretiosa.

Dumas (1966) in his study on the Rana species complex also
recognized the validity of the two subspecies. Livezey and Wright
(1947) in their work on salientian eggs said that the eggs are en-
tirely different between the two subspecies.

In Utah the subspecies has been recognized as R. p. pretiosa by
most writers (V. M. Tanner 1927, Stejneger and Barbour 1943, W.
W. Tanner 1940, Stebbins 1951, Schmidt 1953); however, certain
variations have made its identification confusing. Skousen (1952)
stated that the larvae of R. p. subsp. of Utah do not fit the description
of R. p. pretiosa by Svihla (1935), but rather the description of R.
p. luteiventris by Thompson (1913) and Svihla (1935). He also
stated that the eggs are smaller than those of R. p. pretiosa of eastern
Washington, and. therefore, suggested that the Utah population
along the Wasatch front be called R. p. luteiventris. In a recent
work Dumas (1966) summarized the distribution of R. pretiosa sub-
species as follows:

Pretiosa (sic) is found from northern British Columbia and south-
western Alberta southward through northern Idaho, western Montana,
Washington, northern and western Oregon, and extreme northern Cali-
fornia. Luteiventris (sic) occurs in southern Oregon, southern Idaho,
western Wyoming, and in isolated pockets in northern Utah and northern
Nevada.

Much of the confusion over the Utah subspecies is well founded
and the decision one reaches in classification is largely determined by
the stage of the life history which is studied. We have experienced
similar difficulties in an attempt to classify Utah pretiosa by the
diagnostic features given in the different taxonomic references. Live-
zey and Wright (1947) published a classification of anuran eggs of
the United States and differentiated the pretiosa subspecies mainly
upon: (1) the number of gelatinous envelopes surrounding each
egg, and (2) the size of the eggs and number of the eggs per clutch.
When special care is used in the observation of the jelly layers. Utah
pretiosa key out to subspecies pretiosa becavise of their size and pre-
sence of the inner memljrane. The differentiation of larvae is based
upon the number and size of labial teeth rows. Tadpoles in this case,
as pointed out by Skousen (1952). have two upper rows of teeth and
fit the description of luteiventris by Thompson (1913) rather than

The Great Basin Naturalist
48 MORRIS AND TANNER Vol. XXIX, No. 2

that of pretiosa by Svihla (1935). The diagnostic characteristics used
in the separation of adults are based upon coloration and the presence
or absence of tubercles on the inner surface of the feet. The ventral
coloration of live specimens range from orange to red which would
place them as pretiosa, whereas tubercles are reduced and indistinct
which is characteristic of luteiventris. Based upon the above informa-
tion and the fact that some authors question the validity of luteivent-
ris^ we consider the Utah population to be R. p. pretiosa with a few
aberrant characteristics which, as pointed out by Skousen (1952),
may be the result of intergradation between the two subspecies. This
possibility exists since both subspecies are present in the northern
regions of the Great Basin and are approximately equal distances
from the Utah population. In either case more work is needed on
the taxonomy of the species, and the lines of distribution need to be
established.

Life History and Ecology

Amphibians are used extensively in different experimental fields
of biology and many times this is done without an understanding of
the activities and life history of these animals in nature. This lack
of information may also have a bearing upon other studies and their
results. For example, Johnson (1965) studied temperature tolerance
and developmental rate of R. p. luteiventris in the laboratory, but
was unable to relate these to its distributional pattern based upon the
breeding biology because of lack of information on R. p. pretiosa and
other western species of ranids. In this study we are concerned with
life history and breeding biology of Rana pretiosa pretiosa in Utah.

Study involves data which have been collected intermittently from

1962 to the spring of 1968 with the majority of the data collected
regularly during spring and summer of 1966. This was done by fol-
lowing the activities of frogs from the time they emerged in the spring
until offspring metamorphosed and hibernated in the fall. Observa-
tions and collections were made at regular intervals throughout this
period of activity to provide a continuous record of growth and
activity.

The study actually began in the spring of 1962 as a research pro-
ject by David Avery, but was discontinued when eggs under obser-
vation were destroyed by children. The same study was resumed in

1963 except that all observations and measurements were obtained
from eggs which were removed from the pond and taken to the lab-
oratory where embryonic and larval development were observed in
a 44 liter aquarium until metamorphosis was completed.

To gain a more complete picture of the life history of the west-
ern spotted frog, we obtained permission to study the frogs inhabiting
the ponds on the private property of Mr. Lawson Hamblin (study
area II) during 1966. This proved to be an ideal site, not only be-
cause of protected surroundings, but also because R. p. pretiosa is the
only frog to use these ponds for breeding. As a result no difficulty
was encountered in keeping separate the eggs and tadpoles from

July 31, 1969 ecology of r. pretiosa in utah 49

those of R. pipiens bracliycephala which normally ocx uj)ies the same
waters in this area.

Extensive observations were made and specimens collected at
these ponds during the spring and summer of 1966. This study
is supported primarily by these data, supplemented by additional
observations made by us and others both before and after 1966.

Study Areas

Observations on the western spotted frog were made primarily in
the vicinity of Provo. Utah, along drainage areas of the Provo River
at an elevation of approximately 4550 feet. A population of R. pret-
iosa along the San Pitch River three miles north of Fairview\ San-
pete County, and one near Mona Reservoir in Juab G)unty (collec-
tion records of adults only) are also included. The area used for this
study is at the extreme southeastern extension of the range for R.
pretiosa (See Stebbins, 1966) and includes those populations which
have extended south from Snake River Basin in Idaho into the water
courses along eastern edge of the Great Basin and the western front
of the Wasatch mountains.

All study area were located after much field investigation, and
can be characterized by several features which they have in common.
Each site at which R. pretiosa has been observed was located, in
relation to the local topography, in a low. swampy situation with
some type of spring water supply nearby. Except for study area IV
all sites were near the base of Provo Bench where water collects as
a result of its relative position to the water table. Each observed
site is a small permanent pond of water which has a continual source
of water. Because of their low level and seep springs inflow, the
ponds seldom have an external outlet; therefore, very little move-
ment results because of flow of water through them. As a result each
pond is made up of standing water with a deep silt or muck bottom
in which frogs presumably hibernate during the winter.

Stonewort. Chara sp., makes up the dominate aquatic vegetation
and forms a complete mat covering over the bottom of the pond. Cat-
tails. Typha sp., are present in the deeper parts of each pond, and
provide a cool, moist place for adult frogs to feed during the warmer
summer months. By the end of June Spirogyra sp. is usually com-
mon in water providing an excellent place for hiding and a source
of food for developing tadpoles, which can normally be found within
or beneath floating vegetation.

Study area I was located near the old Provo Brick and Tile Com-
pany in a swampy area resulting from several small sj)rings and
seepage from higher irrigated lands. The three ponds used as breed-
ing sites were all small ponds of standing water with seepage as the
only inlet (Fig. 1). Several other jwnds which were interconne<-ted
by small streams were present, but none of these were used by the
frogs. Study area I has since been filled as a result of construction.

50

MORRIS AND TANNER

The Great Basin Naturalist

Vol. XXIX. No. 2

:<^

Fig. 1. Main breeding pond at study area I. Arrow indicates location of
egg deposition.

Study area II is located at 2160 N. 750 W., 1 mile northwest of
area I on property of Mr. Lawson Hamblin. This site consists of
four man-made ponds, only two of which were used as breeding
ponds. Pond A (Fig. 2) is the only one continually spring fed and
therefore has a temperature several degrees lower than other ponds
in this specific area.

Pond A measures approximately 5 by 8 meters, and has the
smallest surface area. However, it is the deepest pond used for breed-
ing at this location. The depth of the water at the north end is ap-
proximately 30 cm. and has a rocky bottom. The south end of the
pond, where most of the adult frogs are concentrated, is approximate-
ly one meter deep and has a muck bottom. A thick growth of cattails

July 31, 1969 ecology of r. pretiosa in utah

51

Fig. 2. Pond A at study area II. Arrow indicates location of egg deposition.

is present at this end. All the ponds are at the base of a 16 to 20
meter embankment (lake terrace) which extends along the west
side. Willows growing thickly along the bank hang over the water
for approximately one meter. A field with several species of grasses
and weeds borders the ponds from the east. Pond B is located 30
meters north of pond A and is much larger, measuring approximately
12 by 21 meters. Chara sp. completely covers the bottom of the pond
with only a thin layer of water appearing above it. The water is 1 5 to
45 cm deep with a muck bottom 60 to 90 cm dee{).

Study area III is a drainage area from higher irrigated lands and
is located west of Provo along the D. & R.G.W. Railroad tracks at
approximately 1000 N. and 2100 W. (Fig. 3).

Study area IV (Fig. 4) is the highest elevation 5350 ft., at which
observations were made in the Provo River drainage area. It is
located on the south fork of Provo River approximately one and one-
half miles east of Vivian Park, Utah County. These are permanent
spring-fed ponds which have been impounded by man for a water
storage area.

Several observations were made on a population of Rana pretiosa
along the San Pitch River in Sanpete County. This population as
reported by Tanner (1940) is the farthest south that this species has
been collected. The elevation is approximately 6140 feet and because
of the cooler temperature, eggs are laid about two weeks later than
they are near Provo.

All ponds exhibit a basic reaction as shown by the pH readings
in Table 1.

52

The Great Basin Naturalist
MORRIS AND TANNER Vol. XXIX, No. 2

Fig. 3. Habitat of study area III. Arrow indicates location of breeding
activities.

Fig. 4. Hatitat of study area IV.

July 31, 1969 ecology of k. pretiosa in utah 53

Table 1. The pH readings taken from study areas.

Location pH Reading

Study area I 7.45

Study area II pond A 7.50

Study area II pond B 8.48

Study area III 8.52

Study area IV 8.42

Methods and Procedures

Intensive field work has been carried out for the past three years
beginning in the early spring months during 1965 to 1967 for the
piu-pose of locating suitable habitats and noting the first dates of
emergence. Temperature data taken from the ponds were later com-
pared with temperature data obtained from United States Depart-
ment of Commerce Weather Bureau Climatogical Data Reports for
Provo and Geneva. Utah, to determine the physical conditions of
the environment which released frogs from their hibernacula.

Daily observations were made on the breeding and egg laying
habits after their spring appearance. Photographs were taken of eggs
and mating frogs, and recordings made of breeding call of males in
the field with a model 301 Martel Tape Recorder.

All egg clutches were measured in the field and left in their
natural surroundings. Clutch dimensions were measured to the near-
est centimeter by means of a metric ruler. Volume was determined
by water displacement in a 500 ml graduated cylinder. Several eggs
were taken from each clutch, preserved in 10 percent formalin, and
removed to the laboratory where each egg was examined and mea-
sured individually to the nearest 0.1 mm with a 10 power magnifica-
tion of a dissecting microscope equipped with an ocular micrometer.

I'o facilitate viewing of the inner membrane of the eggs, several
techniques were employed. One such method, useful in the field, was
to float a few eggs over the surface of the water in a small, white
enamel pan. and then to observe the shadow cast by each membrane
on the bottom of the pan when placed in direct sunlight. Although
this method was successful for fresh eggs, it was not satisfactory for
preserved eggs because of irregular shadows resulting from distortions
in the outer surface. Another technique which proved very useful
was staining. 1 his was accomplished by first embedding individual
eggs in dental impressive material, 1 Ty dtocolloid Algenate, according
to directions on the label. After approximately four minutes the
capsule was cut with a razor blade as close to the center of egg as
possible. A drop of either Congo Red or Giesman stain was then
placed over the exposed surface for one or two minutes and then
removed. The resulting ring or rings within the jelly could then be
observed and measured under a dissecting microscope.

Tadpoles were collected randomly at regular intervals and pre-
served in 10 percent formalin, and all measurements were taken

The Great Basin Naturalist
54 MORRIS AND TANNER Vol. XXIX, No. 2

to the nearest 0.1 mm as follows: head-body length, from tip of
snout to the midpoint of cloacal opening; tail length, from midpoint
of cloacal opening to tip of the tail. Total length was derived from
the sum of the body and tail lengths; widths were taken at widest
part of the body. Weights were taken on a Mettler electric balance
and read to the nearest 0.1 mg and estimated to the nearest 0.01 mg;
tadpoles were placed on blotter paper for 15 seconds after removal
from the formalin before weighing.

Egg numbers were obtained by three different methods: (1) by
counting eggs in each clutch after ovulation, (2) by counting the
number of eggs in a given volume and then measuring the volume
of the whole clutch, and (3) by actual count of eggs dissected out of
mature females collected at intervals during late summer of 1966
and early spring of 1967.

Some difficulty was encountered during 1965 in locating tadpoles
because of the dense growth of vegetation. To facilitate their collec-
tion, a small mesh wire screen was placed around the eggs at study
area II on March 26, 1966 (Fig. 4 and 5). The screen was approxi-
mately 8 meters long and made a semi-circle in the water. The open
end of the screen was approximately 3 meters long. The water level
at pond A increased progressively during the summer so that it
provided no barrier late in the summer. The effect of the screen at
pond B is discussed in conjunction with larval development.

Analysis of dissolved minerals in water taken from the ponds
was obtained through a measurement of conductivity with a salt
bridge. Results are expressed in parts per million.

Hydrogen-ion concentrations were determined with an electric
pH meter in the laboratory.

Discussion

The first appearance. The western spotted frog emerges from
hibernation normally during the middle of March when the air
temperature has risen to 13-16° C for several days or after a rain
storm which has warmed the water sufficiently (Fig. 5-8). Spotted
frogs appear about one week after the chorus frog Pseudacris nigrita
triseriata which has been the first anuran to appear in the spring
within the time covered by this study. The emergence of the leopard
frog R. pipiens normally follows R. pretiosa by 7-10 days so that by
the end of March the chorus frog and both ranids are present in the
ponds often in large number. The peaks of the spawning seasons for
these species, however, normally do not overlap in a given habitat
at the same general elevation.

During the course of this study, earliest record of appearance for
R. pretiosa was March 6. 1967. The seasonal temperature was unusu-
ally warm during this time. More important, however, than the air
temperature itself is the temperature of the water under which the
frogs hibernate. A temperature of 10-11° C seems to be a critical
point in their activity. Not only must the water reach this tempera-

July 31, 1969 ecology of r. pretiosa in utah

55

1962

DAILY PRECIPITATION (cm)

'L^JL^JWAJiJLJ'LJJJ

30

-2^-

-30

DAILY MAX. & MIN. TEMP, {t.)

Fig. 5. Climatic data for the spring of 1962. Arrow indicates approximate
date of emergence; X indicates approximate date hatching began.

56

MORRIS AND TANNER

1963

The Great Basin Naturalist

Vol. XXIX, No. 2

3-
2-

1

DAILY PRECIPITATION (cm.)

u

3(fr

o

25

DAILY MAX. & MIH. TEMP.Cc)

JANUARY

FEBRUARY

MARCH

APRIL

Fig. 6. Climatic data for the spring of 1963. Arrow indicates approximate
date of emergence; X indicates approximate date hatching began.

July 31, 1969 ecology of r. pretiosa in utah

1965

57

3p

2-

1 -

DAILY PRECIPITATION (cm.)

_>^-A.w^_AJLiJ(L--_

i^aOL.

Au)L_L.

35
30'

DAILY MAX. & MIN. TEMRTcI

JANUARY

FEBRUARY

MARCH

APRIL

Fig. 7. Climatic data for the spring of 1965. Arrow indicates approximate
date of emergence; X indicates approximate date hatching began.

58

MORRIS AND TANNER

The Great Basin Naturalist

Vol. XXIX, No. 2

1966

DAILY PRECIPITATION (cm.)

aJV__A.

j\ Av

FCBRUARY

Fig. 8. Climatic data for the spring of 1966. Arrow indicates approximate
date of emergence; X indicates approximate date hatching began.

July 31. 1969

ECOLOGY OF R. PRETIOSA IN UTAH

59

ture to release them from hibernation, but when early morning tem-
j)eratures fall below 10° C, or during a cold period causing a similar
drop, frogs are not found at the surface of the water. Dredging of
the bottom of the ponds reveals their presence in a semidormant state.
As the temperature of the water again warms to 10° C they can be
seen resting in vegetation just under the surface of the water. As the
temj>erature warms to 1 1 ° C they will come to the surface.

There are undoubtedly other factors involved in stimulating their
release from hibernation. The effect of moisture has been mentioned
by both Vernberg (1953) and Martof (1953) as being important in
releasing amphibians from hibernation.

During the spring of 1966 and 1967 it was noticed at two separ-
ate locations that frogs were appearing later in those ponds where the
water level had dropped so that in the more shallow portions of the
jx)nds the bottom was exposed. There may have been several factors
involved in their retarded arrival, two of which may have been a
lower temperature or a greater fluctuation in temperature because
of a smaller quantity of water. This is an area of study open to
further investigation. However, one item was noted, namely that
these receding ponds showed a greater concentration of soluble salts,
approximately 100 ppm higher than the others. This may possibly
provide a type of physiological dryness to the frogs causing a delay
in their appearance.

The males are first to appear and are present three or four days
before the larger females emerge. The smaller immature frogs do not
appear for two or three weeks after the females or until the breeding
season is completed.

A summary of measurements taken on 36 frogs collected and re-
leased at study area II, pond A, on March 27. 1966, can be found in
Table 2.

Table 2. Measurements of postbreeding frogs taken March 27, 1966, from
study area II, pond A.

Sex

Number of
specimens

Snout-vent length in mm
Range Mean

Female

Male

Immature

5

25

6

78.6-61.6
64.2-47.2
42.6-31.0

70.0
49.9
37.8

As males emerge they do not exhibit much activity and can be
found either calling from the pond with only their heads above water,
or buried in vegetation during a cold period.

Vocality. The call itself closely resembles the sound made by re-
peatedly clicking the tongue against the top of one's mouth. The calls
come in a series of clicks ranging in number from six to approxi-
mately 50. rhe rate of the notes varies depending upon the tempera-
ture. An examination of several calls made at 16° C showed a var-

60

MORRIS AND TANNER

The Great Basin Naturalist

Vol. XXIX. No. 2

iance in rate of from 300 notes per minute to 480 notes per minute
by different males. The duration of an individual call is from 4-10
seconds and repeated at a rate of as many as 10 calls per minute.
With each note the floor of the mouth is depressed approximately
2 or 3 mm. The call can be given either above or below the surface
of the water with the call from below the surface being somewhat
louder. The call is weak and is muffled when other anuran species
are calling from the same pond. The maximum distance for hearing
the call is somewhat less than 25 meters, the average being 7 to 10
meters. The call of one male has been observed to stimulate others
to call. This was noted after the frogs had been disturbed and many
moved to the bottom of the pond. As one came near the surface and
called, other frogs came from under the vegetation and swam to the
top as if responding to a signal; soon all were at the surface and
calling again. Normally at this time they pay little attention to any
movement around the pond, even to the point that many can actually
be touched before they attempt to escape.

Mating. Mating takes place immediately upon the arrival of the
female as evidenced by the following: ( 1 ) at no time was an adult
female observed to be present which was not in amplexus until after
ovulation, and (2) the aggressiveness with which males attempt
amplexus would require only a few minutes before a receptive gravid
female would be clasped. The sexual drive is very strong in males of
this species, so much so that not only do they attempt amplexus with
others of the same sex. but at times two and three males have been
observed embracing one female (Fig. 9) ; however, this multiple

Fig. 9. Multiple embrace of two males with one female. Note typical posi-
tion of calling males at breeding sites.

July 31. 1969 ecology of h. prktiosa in utah 61

embrace lasts for only a few iuiiiutes. The drive is so strong that
attempted amplexus has been observed with other objects in the
water, even to the point that one male, rather than escaping as it
was reached for, swam closer and clasped onto my finger. The tenaci-
ty to mate is no doubt correlated with competition exerted by the
large male to female ratio, and the fact that this species is very gre-
garious with all of the breeding activity limited to a small area with-
in the pond. Of 30 adult frogs collected on March 27. 1966. there
was a 1 to 5 female to male ratio (Table 2).

The difference in size between the larger females and smaller
males (Table 2) places the vent of the males directly above that of
the females during amplexus. Amplexus is axillary with the male
embracing from above.

Ovulation. The females are apparently ready for ovulation as
soon as they emerge and several clutches of eggs have been observed
the same day. The eggs are normally laid at a water temperature of
approximately 14° C and may be laid during the day or night. In
nature, egg-laying has been observed to be interrupted for several
days during a cold period, but would proceed as soon as the water
again warmed to the approximate temperature. The embrace is
maintained during the entire period. Experimentally males have
maintained the clasp in the laboratory during a lowering of tempera-
ture almost to the point of freezing at which time the clasp was
slowly released. Throughout the entire period of amplexus. the male's
main physical activity seems to be to maintain the clasp until ovula-
tion. All activities, such as protection and movement into deeper
water during cold periods, are dependent upon the action of the
female. The male remains passive during the entire period.

Fertile eggs have not been taken from a clasping pair which have
been removed to the laboratory. This artificial condition seems to
inhibit the females from laying even though the males have main-
tained their embrace for as long as two weeks.

The actual process of ovulation has not been observed in nature,
although eggs have been found before the first cleavage has taken
place. An examination of a clutch of eggs collected March 29, 1966,
at study area III. showed that some eggs were attached together by a
small gelatinous chord 2 to 3 mm in diameter as reported by Turner
(1958), whereas the surfaces of adjacent eggs were attached directly
by 3 to 5 mm of their outer membrane. A close examination of a
typical egg revealed that it was connected to five other eggs. Two
of these adjacent eggs were attached by means of the gelatinous
chord, while the other three were attached directly by their outer
membranes. As eggs are laid the adhesive character of the mem-
branes causes them to become attached to one another and form one
large, irregular clump. They also weakly adhere to vegetation below
them as they come in contact with it.

Oviposition sites. The sites used for egg laying have several
characteristics in common: (1) they are normally in the part of the

62

MORRIS AND TANNER

The Great Basin Naturalist

Vol. XXIX, No. 2

pond which has warmest temperature resuhing from solar radiation,
and in most instances this is on the west side where the eggs catch
the morning sun; (2) the eggs were always laid in an open area in
clear water and never in among the cattails or in floating Spirogyra
which was oftentimes only a few feet way; (3) eggs are usually in,
the shallow portion of the pond 10 to 20 cm under the surface of the
water and attached to Chara sp. This attachment is rather weak,
however, and within a week the clutch usually breaks loose and
floats on the surface where the eggs become more of a flattened mass
and are scattered by wind and water movements.

As one clutch of eggs is laid, there apparently is a stimulation for
other females to lay their eggs in the same area. This behavior was
best shown at study area IV where approximately 50 clutches of eggs
were laid within an area 75 cm in diameter, even though other
apparently ideal locations were available. This large assemblage of
eggs was weakly attached, one to another and to the vegetation (Fig.
10). In this large pond area only a few clutches were found away
from the large mass of eggs.

Eggs. Durin,g the spring of 1965, 10 clutches of eggs were mea-
sured and counted in the field (Table 3). The average number of
eggs per clutch was 605. The largest clutch contained 725 eggs and
the smallest 430 eggs.

In 1966 eggs in 21 different clutches were counted and measured
(Table 4), and the average number per clutch for this year was
746. The maximum was 1160 and minimum 147 eggs per clutch.

Fig. 10. Oviposition site at study area IV. Note the closeness of egg
masses (approximately 50 clutches).

July 31, 1969 ecology ok r. pretiosa in utah 63

Table 3. Comparison of clutch size and number of eggs collected from study
area I on March 30, 1965.

Total no. Eggs Clutch size Diameter of eggs

per clutch in cm in mm

541
625
695
715
615
635
650
725
514
430

Egg

Outer Mem.

9-8-5

2.5

10.0

9-7-5

2.3

10.1

11-8-7

2.4

10.5

10-8-7

2.5

13.0

12-9-9

2.5

10.0

11-9-7

2.4

10.0

13-9-7

2.5

13.0

11-6-6

2.2

10.7

6-5-3

2.1

12.0

5-4-2

2.3

11.0

Table 4. Relationship of clutch volume and number of eggs per clutch.

Total no.

Clutch

eggs per

volume

Eggs per cc

Locality

Date

clutch

in cc

of clutch

Study area II

24 March 66

940

320

2.62

pond A

660

110

6.00

500

105

5.24

900

215

4.37

950

400

2.38

29 March 66

600

4O0

1.50

840

415

2.02

Stury areall

29 March 66

716

285

2.58

pond B

900

475

1.72

148

130

1.13

1160

550

1.11

30 March 66

980

500

1.96

31 March 66

655
625

325

2.01

Study area III

23 March 66

900

285

3.17

925

230

3.96

380

115

3.30

460

100

4.60

Study area IV

5 April 66

990

250

3.96

716

235

3.05

730

375

1.98

Because of the great variation, in egg numbers and because the
numbers were consistently lower than those reported by such workers
as Svihla (1935). Livezey and Wright (1947), Wright and Wright
(1949). and Stebbins (1951), it was suspected that perhaps these
frogs were laying a small number of eggs at one location and then
producing others later, as was reported by Bragg (1944) for the
common leopard frog in Oklahoma. However, in the dissection of
gravid females of R. p. pretiosa from Utah County, this same varia-
tion in numbers has been observed before any eggs had been laid
(Table 5). Therefore, it is reasonable to assume that Utah popula-
tions produce a smaller number of eggs for each complete clutch.

The Great Basin Naturalist
64 MORRIS AND TANNER Vol. XXIX. No. 2

Table 5. Comparison of egg numbers and size of females collected from
study area II, pond A.

Total no.

Clutch

eggs per

volume

Eggs per cc

Total length

Date

clutch

m cc

of clutch

of female

27 July 66

902

80

11.3

74

19 August 66

763

50

15.2

73

13 September 16

168

67

7 October 66

860

60

14.4

70

8 October 66

1060

72

14.7

64

15 October 66

393

40

11.2

59

We are unable to explain the reason for variation in the number of
eggs per clutch. The size of females does not reliably explain the
variation in clutch size.

Freshly deposited eggs soon increase in volume by absorption of
water, so that within several hours after laying they may range
from 110 cc to 550 cc with the average of those measured being 291
cc. Clutches of the same age are variable as to size and number of
eggs per cc as is indicated in Table 3.

Several days after being deposited, a clutch usually appears as
an irregular, oval mass just under the surface of the water. Within
about a week egg mass becomes less coherent, and breaks loose to
float at the surface in more of a plinth shape. By the time the eggs
are ready to hatch they are almost undiscemible from above, because
of the accumulation of dirt and debris on the exposed surface and the
breaking up and amalgamation of the adjoining ielly masses. There
is also a dry, crusty appearance to the mass resulting from destruction
of 10 to 20 percent of surface eggs. This high mortality rate among
the eggs is a result of several factors. First, the eggs are laid early
in the spring so that there are many nights in which they become
encrusted in ice which destroys some of them. Also, because of the
long period of embryonic development they may later be forced out
of the water by the new growth of submerged vegetation, which ex-
poses the top eggs to desiccation.

Each egg is enclosed in two gelatinous envelopes separated by an
indistinct inner membrane. The total diameter of the outer capsule
normally varies within a range of from 9.0 mm to 13.0 mm although
variations as great as 8.0 mm to 21.0 mm have been measured. The
average diameter of those eggs measured (Tables 3, 6-9) was 10.0
mm. The diameter of the indistinct inner membrane has a normal
variation of from 3.8 mm to 6.1 mm and a mean of 5.0 mm. This
membrane is difficult to see in the fresh eggs without the techniques
employed as described previously, but can be seen even in unferti-
lized eggs if these methods are used. As development progresses this
inner membrane becomes more obvious because of the infestation of
algae from the outside which stops at the inner membrane. If eggs
fail to develop the ovum deteriorates clouding the inner jelly coat
out as far as the inner membrane.

July 31, 1969 ecology of r. pretiosa in utah

65

Table 6. Measurements in mm of eggs
collected March 29, 1965, at study area I.

Egg Diameter

Capsule
Diameter

Inner

Outer

2.7

5.0

10.0

2.8

6.0

13.0

2.7

4.9

12.0

2.6

5.0

10.0

2.0

4.7

11.0

2.5

4.9

11.0

2.4

3.8

11.0

2.1

4.5

10.0

3.0*

8.0*

21.0*

*Extremely large egg

Table 7. Measurements in mm of un-
fertilized eggs laid in the laboratory by a
female collected March 27, 1967, at study
area III.

Egg Diameter

Capsule
Diameter

Irmer

Outer

2.5

4.7

9.0

2.4

4.7

9.0

2.6

4.9

8.0

2.7

4.8

10.0

3.0

5.0

11.0

2.8

5.0

12.0

2.8

5.0

12.0

Table 8. Measurements in
collected March 29, 1966, at

mm of eggs
study area III.

Egg Diameter

Inner

Capsule

Diameter

Outer

2.6 6.0
2.5 6.1
2.4 5.2
2.4 5.0

11.0

10.1

10.0

8.0

Table 9. Measurements in
collected April 5, 1966, at

mm of eggs
study area IV.

Egg Diameter

Inner

Capsule

Diameter

Outer

2.6

5.0

10.6

2.6

5.0

10.5

2.6

3.0

10.0

2.7

5.1

10.3

2.6

5.0

10.3

2.5

4.5

10.2

2.4

4.9

10.1

The Great Basin Naturalist
66 MORRIS AND TANNER Vol. XXIX, No. 2

The ovum is normally 2.5 mm in diameter, but does vary within
a range of 2.1 to 2.9 mm. Preservation can cause some distortion
in size if not done properly and may account for some of the varia-
tion shown in Tables 3, 6-9. The color of an egg is dark brown to
black above, and pale yellow or light tan below. The vitelline mem-
brane is closely applied to the ovum at time of hatching, which allows
an enlargement of the fluid-filled chamber for embryonic develop-
ment and movements.

As the embryonic development progresses, the embryo becomes
progressively lor^ger imtil at the time of hatching it is 8 to 10 mm
in length. An attempt to analyze the physical factors involved in the
actual hatching process has not been made. Discussions of these
factors are given by Noble (1954) and Bragg (1940a, 1940b). A
change in temperature was observed to have a noticeable influence
on the hatching process. This was demonstrated when eggs placed
in a natural spring did not hatch even though embryonic develop-
ment seemed to be complete. The temperature of the spring water
remained constant, at approximately 1 1 ° C, with only minor fluctua-
tions. Samples of these eggs when removed from the cold environ-
ment would hatch in approximately one hour while those left in the
spring hatched 7-10 days later. Eggs which were brought into the
laboratory hatched at room temperature in about seven days, whereas
those in nature and subiect to the lower temperatures required from
13 to 23 days to hatch. Under normal pond temperature where the
eggs were laid, the majority hatched about two weeks after ovulation.
However, length of time required for hatching varies from year to
year depending upon the fluctuations of atmospheric temperatures
and the amount of cloud cover which reduces solar radiation.

Larval Period. The hatching activities cover a period of several
days, with most eggs hatching within three weeks after ovulation.
The tadpoles remain attached to the gelatinous material by their oral
suckers for two or three days following hatching. As they break
loose from the jelly they sink to the bottom where only an occasional
swimming movement is made. The results of ciliary movement along
the body surface keeps a fresh supply of oxygenated water flowing
from anterior to posterior past their external gills, as observed under
a dissection microscope (30x mag.). The mouth and anal openings
do not develop until one or two days following hatching. Actual
feeding begins in conjunction with the first swimming activities.
Associated with this free feeding stage, an operculum develops over
the external gills and water is taken in through the mouth and out
the spiracle on the left side of the body. The food eaten is mostly
secured by scraping or rasping off the loose outer surface of decom-
posed plant material, a function for which the teeth are well adapted.

The rows of larval teeth are two upper and three lower as des-
cribed by Skousen (1952) except that in a high percentage of the
specimens measured, the first lower row was continuous rather than
divided medially. For a more complete discussion on the development
of larval mouth parts see Johnson (1965). An examination of the

July 31, 1969 ecology of r. pretiosa in utah

67

digestive tracts of several larvae show mostly decomposed material
and some green algae. Tadpoles in the laboratory have been raised
on live Spirogyra which gives a green color to the digestive tract.
Burke (1933) was able to raise Rana pretiosa tadpoles through
metamorphosis on a 24 hour mixed culture of bacteria. He concluded
that the common water bacteria contain all the food factors necessary
for their normal growth. The scavenger feeding of tadpoles advanced
beyond stage 26 (Table 10) was further observed as they ate the
remains of dead pretiosa tadpoles.

Stage 1
Stage 2
Stage 3
Stage 4:
Stage 5
Stage 6
Stage 7
Stage 8
Stage 9
Stage 10
Stage 11
Stage 12
Stage 13
Stage 14
Stage 15
Stage 16
Stage 17
Stage 18
Stage 19
Stage 20
Stage 21
Stage 22
Stage 23
Stage 24
Stage 25
Stage 26
Stage 27
Stage 28
Stage 29

Stage 30
Stage 31
Stage 32
Stage 33
Stage 34
Stage 35
Stage 36
Stage 37
Stage 38
Stage 39

Stage 40
Stage 41
Stage 42
Stage 43
Stage 44
Stage 45
Stage 46

Table 10. Summary of developmental stages.

Egg fertilization

Appearance of the Gray Crescent

Two cells (first cleavage)

Four cells

Eight cells

16 cells

32 cells

Mid cleavage

Late cleavage

Beginning of gastioilation

Involution at dorsal lip

Blastopore complete

Neural plate develop

Neural folds and groove formed

Beginning of ciliary rotation, closing of neural fold

Neural tube formed, gill plates discernible

Development of tail bud

Muscular movement

Heart beat

Gill circulation

Cornea becomes transparent, mouth opens

Tail fin circulation

Opercular fold formed

Operculunr covers right gills

Operculum covers left gills

Hind limb bud appears

Limb bud length equal to or greater than one-half the diameter

Limb bud length equal to or greater than diameter

Limb bud length equal to or greater than one and one-half the

diameter

Limb bud length equal to two times the diameter

"Foot" becomes paddle shaped

Indentation formed for fourth and fifth toes

Indentation formed between third and fourth toes

Indentation formed between second and third toes

Indentation formed between first and second toes

Beginning of toe separation

All toes separated

Appearance of metatarsal tubercles

Subarticular tubercles appear as light patches on the inner surface

of the foot

Subarticular tubercles fully developed

Loss of cloacal tail piece, "skin window" appears for forelimb

Forelimbs free

Angle of mouth midway between nostril and eye

Angle of mouth below midpoint of eye

Angle of mouth posterior to eye, tail stub remains

Metamorphosis complete

The Great Basin Naturalist
68 MORRIS AND TANNER Vol. XXIX, No. 2

The actual growth of tadpoles, like that of the unhatched em-
bryos is greatly influenced by different temperatures (King, 1903;
Moore, 1938; Brattastrom, 1963; and Johnson, 1965). Tadpole act-
ivities at the early stages indicate that they employ to some extent
a biological control of their temperature by their position within the
pond. The young tadpoles prefer to stay close to the bottom and do
so in the shallower areas where water temperature during the day
is warmer. Aggregations of tadpoles have been noted at various
times throughout the summer and as suggested by Carpenter (1953b)
and Brattstrom (1962) these close aggregation may contribute, be-
cause of their melanistic color, to the warming of their immediate
surroundings by absorption of solar radiation which in effect speeds
up development leading towards metamorphosis. On overcast and
rainy days tadpoles move to deeper water and are usually within
the vegetation. By stage 30 (Table 10) they become more active,
continually swimming and feeding throughout the pond. This activi-
ty continues until metamorphosis at which time a modification of the
digestive tract permits them to feed closer to the shore on small arth-
ropods, and to rest in the vegetation.

Summary of Developmental Stages. To facilitate a description
of the developmental process taking place in embryonic (prefeeding
stage) and larval (free feeding stage) frogs, staging tables have been
used. They are of value in a life history study since the external
morphology of each stage of development is described and illustrated.
These staging tables have been used for some time by authors in
descriptive and taxonomic works. Two systems of numbering the
stages were in use until Gosner (1960) presented a table which
simplified them. Johnson (1965) described and illustrated the stages
as they applied to the spotted frog, Rana pretiosa luteiventris.

Table 10 is a smnmarization from Gosner (1960) and Johnson
(1965), and is used as a standard of comparison since growth rates
are too variable from one locality to another to be used for this pur-
pose.

Growth Rate. The embryonic growth up through stage 27
(transformation stage from embryo to larvae) is dependent upon food
stored in the yolk. All growth beyond this stage is dependent upon
the tadpole's own feeding. For this reason no constan,t measurements
of growth were taken, until this stage. Those tadpoles which developed
at warmer temepratures reached this stage in a shorter period of time
and were 2 to 3 mm longer in total length. Study site II. pond B,
continually maintained a 4-8 C warmer temperature than pond A
until the middle of September when they became equivalent. An
average of temperature readings taken at different hours of the day
throughout the larval period shows pond A with 13.6 C and pond
B with 18.7° C. The more rapid development resulting from the
warmer temperature at pond B was, however, counteracted by the
crowded conditions imposed upon the tadpoles when a screen, (Fig.
2) was placed around some of the eggs. The effects of the screen were
evident by the middle of May when it was obvious that the growth

July 31, 1969 kcology of r. pretiosa in i tah 69

of tlie tadpoles at j)oiid A exceeded that of the now stunted tadpoles
of pond B (Fig. 11). Another factor which may have had an effect
upon the growth of the tad[)oles at j)ond B was the high concentration
of dissolved salt in the water. The concentration was fi22j)pni at pond
B as comj)ared with a 380 ppni reading at pond A. Pond B also had
a j)H of 8.9 compared with a 1 J at pond A. A detailed study of dis-
solved minerals was not made; however, their effects prove to be
very similar to those causing the stunted condition in plants grown
in the same concentration of minerals because of the checked absorp-
tion of water.

Cameron (1940), in a summary of his work, stated that for Rami
pipiens:

\ . The nature and amount of dissolved substances and their relative
proportions exert separate and unlike influences upon the rate of de-
velopment and the stage attainefi at hatching of normal frog embryos.
Eggs kept in well water containing fluorine hatch earlier but at a less
advanced stage of development than those kept in i)ond or distilled
water.

2. Flourine in concentrations as low as 1 ppm is able to exert a
constant and measurable retardation on the rate of development and
stage at hatching.

The tadpoles of pond B showed their greatest rate of growth from
April 10 to May 23. At the end of this time their total length varied
within a range of from 33 to 47 mm (Fig. 11). This growth rate
was a maximum of approximately 2.7 mm per day and a minimum
of 1.5 mm per day since stage 26. The development stage at the end
of this growth period varied from 33 to 37 (Fig. 12) indicating that
the greatest period of growth was from the first appearance of the
hind limb bud until the full development of the hind foot. There was
no appreciable growth from May 23 to July 27. The ontogenetic
development, however, had advanced to a stage where several of
the larger tadpoles were transforming into frogs (Fig. 12). This
transformation of larger tadpoles seemed to release some of the
growth retarding effects on smaller ones (Fig. 16-21). Thus, there
followed another period of increased growth so that by the end of
August their total length varied from 40 to 57 mm. Bose (1960)
stated that when stunted tadpoles were separated from their larger
siblings normal growth again took place.

The tadpoles of pond A continued their growth until stage 40
when metamorphosis began without any noticeable decrease in the
growth rate (Fig. 13); therefore, tadpoles at this pond had mostly
transformed into immature frogs by the end of August. Although
the development of larvae at Pond A was approximately two weeks
later than at i)ond B, they developed at the same rate until the first
of June at which time the tadpoles of pond B became stunted (Fig.

12.)

The individual size of each tadpole showed signs of variability
shortly before the hatching stage and these differences became more
exaggerated with their development. Cameron (1940) suggested that

70

The Great Basin Naturalist
MORRIS AND TANNER Vol. XXIX, No. 2

+ -t- 4+4-1- + 4-

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TOTAL LENGTH (m.m.

Fig. 11. Scatter diagram showing total length of specimens collected ran-
domly at study area II on days indicated. Dots are pond A and crosses are
pond B.

July 31, 1969 ecology of r. pretiosa in utah

71

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STAGES OF DEVELOPMENT

Fig. 12. Scatter diagram showing stages of development of specimens col-
lected randomly at study area II on days indicated. Dots are pond A and crosses
pond B.

72

MORRIS AND TANNER

The Great Basin Naturalist

Vol. XXIX, No. 2

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domly at study area II on days indicated. Dots are pond A and crosses pond B.

there is a genetic variation of as much as 10 percent in the size of
eggs reaching a given stage. This difference in size was more appar-
ent at pond A where the total mass of the individuals, as shown by
the weights in Fig. 1 1, on August 3, had a range of 1 .69 to 6.12 gm.

An examination of Figs. 11-13 shows that absolute body dimen-
sions are so variable throughout the larval development that the use
of these measurements as key characters in species identification
would be most difficult. The use of body ratios as suggested by Lim-
baugh and Volpe (1957) is also questionable since there is a variance
from one individual to another even in those of the same age (Fig.
14). Data show that the larger tadpoles have a smaller tail to head
ratio, and when the head length is greater than 24 mm the tail can
have a variance of approximately 23 mm (21 to 44 mm) (Fig. 16).

July 31, 1969 ecology of r. pretiosa in utah

73

.80r
.60 -
.40 -
.20 -

a.00-
.80-
.60-
.40-
.20-

5.00-
.80-
.60-
.40-
.20-

4.00-
.80-.
.60-
.40 -
.20-

loo-

.80-
.60-
.40-
.20-

2.00-
.80-
.60-
.40-
.20 -

1.00 -
.80-
.60-
.40-
.20-

■s^

^

.^

+ + +

_i I I I i_

Fig. 14.

5 10 15 20 25 30 35 40 4 5 50 55 60 65

TOTAL LENGTH (mm.)

rig. 1^. Scatter diagram showing comparison of length and weights of
specimens collected randomly at study area II. Dots indicate specimens up
through stage 45 at pond A, crosses indicate specimens up through stage 45 at
pond B, open stars indicate specimens past stage 45 at pond A, and closed stars
indicate si>ecimens past stage 45 at pond B.

Tadpoles at pond A normally began metamorphosis, stage 41.
once they reached a maximum total length of 50-55 mm (Fig. 11, 12
and 15). Some tadpoles, however, continue to grow even after others
of approximately equal size have begun transformation, so that they
reached a maximum size of up to 70 mm two or three weeks later.
There is a tendency for these large tadpoles to remain in an arrested
state of development between stages 36 and 40 once they have grown
past the normal size for transformation. They remained as large tad-
])oles up until the last of August before any started to transform

74

MORRIS AND TANNER

The Great Basin Naturalist

Vol. XXIX, No. 2

and then all were able to complete metamorphosis by September
15, 1966.

The tadpoles at pond B were not only retarded in size (as a
group), but also retarded in their ontological development (Fig. 12
and 15). The first transformation was on August 3, 1966, two weeks
later than the first tadpoles from pond A (Fig. 12). Transformation
was a slow process, requiring until October 15, 1966, for the remain-
der of the population to complete metamorphosis (Fig. 12). There-
fore, the time required for metamorphosis to occur at this locality
varied from 122 days to a maximum of 209 days after egg laying.

The maximum and minimum total lengths of tadpoles at pond B
reaching stage 40, were 57 to 47 mm respectively (Fig. 15), com-
pared wdth the same maximum and minimum of 70 and 60 mm at
pond A. An examination of Fig. 14 gives another picture of the
smallness of transforming frogs from pond B. When comparing their
weights with total length, the pond B tadpoles are consistently smaller
at the time of metamorphosis, with the smallest weighing 0.52 gm
and a total length (snout- vent) of 19.5 mm. This would indicate
that in some tadpoles very little growth would have taken place
since the middle of May (when the stunted condition developed) be-
cause they were approximately the same head-body (snout- vent)
length and total weight as the tadpoles on May 23 (Fig. 13). It soon

•h+ ~ •• •••

+

4 •

....

4

4-1.
4 -t-

• • -t- ■«— ^

-44t.

+ •

■»++-H-

+f4. -M- +■•■

4 +

+ ^+4

4

•

■*-

+ •*•+

+++

-f •

t

-M-

• + ■•-

■H-++

4 ■»■

• •• «•

4- ++ ■»+«••"

+<••»■•-

• *■••

•

30 3S 40 45 SO SS 60 6S

TOTAL IINGTM (mm.)

Fig. 15. Scatter diagram showing comparison of total length and stage of
development of specimens collected at study area II. Dots indicate pond A and
crosses pond B.

July 31, 1969 ecology of r. pretiosa in utah

75

becomes apparent that size is not one of the stimulating factors in
metamorphosis (Fig. 14), but rather metamorphosis is dependent
on an internal reaction.

Once metamorphosis begins (stage 41) the time required to com-
plete the transformation (stage 46) is comparatively short. A series
of 10 tadpoles at stage 40 were examined to determine the time re-
quired for the appearance of the front legs. The front leg is the first
to appear and the joint of the elbow can be seen appearing out of the
spircular opening for 3 to 4 hours before the full leg becomes appar-
ent. Only an occasional three legged frog will be observed in nature
because of the brevity of this stage. Of the 10 specimens examined
after the left leg appeared, the time varied from 4 to 8 hours before
the right leg appeared. Detailed descriptions of the tadpoles are given
by Stebbins (1951), Skousen (1952), Turner (1958), and Johnson
(1965). and it is for this reason that we have not included detailed
descriptions.

Hibernation. The ponds at study area II were checked at iner-
vals during the fall of 1966 to determine the data of disappearance
for hibernation. Both frogs and tadpoles were seen until the first of
October, but by the 15th all of the tadpoles had metamorphosed and
were basking at the edge of the pond. The adult frogs were seen in
the deeper water among the cattails. The next observation was Octo-
ber 27 at five o'clock p.m. The water temperature at this time was 9°
C at pond A and 14° C at pond B; no frogs were evident. The air
temperature had been down to freezing for several nights since the
last visit, but it had taken about two weeks for the water to cool to
or below the critical 10° C level (Fig. 16). The frogs had been ob-
served in about the same location of the pond throughout the sum-

40

■

3S

-

30

■

.

S5

-

*. *+ . : ;

» • *

• *•

ao

IS
10

-

■t +

+
+

... ^**'

5

-

■f
1 1

1 1 1-

1 1 _J .

T Al L lENOT H ( mm.)

Fig. 16. Scatter diagram showing comparison of tail and head-body lengths
up through stage 40 of specimens collected at study area II. Dots are pond A
and crosses pond B.

The Great Bsisin Naturalist
76 MORRIS AND TANNER Vol. XXIX. No. 2

mer and right up until the time of hibernation. It is assumed, there-
fore, that these frogs hibernate in the muck under the same water
in which they breed.

Conclusions and Summary

This study was in progress with some interruptions from 1962
to 1967 for the purpose of investigating the breeding biology and life
history of Rana pretiosa pretiosa Baird and Girard in central Utah.
Some effects of various environmental factors upon the growth and
habits of this species were considered.

Collections were made from the time the adults emerged from
hibernation in the spring until the tadpoles metamorphosed and
frogs hibernated in the fall. Data gathered at the ponds and samples
of the life history stages were taken to the laboratory where they
were analyzed and studied.

The western spotted frogs emerged from hibernation normally
during the middle of March as a result of the warming of the en-
vironment. This same species does not emerge until May at Yellow-
stone National Park, Wyoming (Turner 1958), but is usually present
by late February in Washington (Svihla 1935). A few individuals
were reported by Dickerson (1906) to be out sunning themselves
throughout the winter at Puget Sound.

In Utah this species prefers small ponds of standing water grown
thick with stonewart and possessing a deep muck bottom from which
cattails emerge. Tanner (1931) in speaking of the Utah population
states, "it is always found near springs, small streams, and swamps."
Turner (1958) recorded them as visitors of "pools of stagnant water
..." for egg deposition. Stebbins (1966). however, reported the
habitat as follows: "A highly aquatic species found in the vicinity
of cold, permanent water — streams, rivers, marshes, springs, pools,
and small lakes. Seems not to occur in warm stagnant ponds grown
to cattails."

The adult male frogs appear in large numbers at the beginning
of the breeding season, which begins immediately following the
emergence of the larger females. The emergent male frogs seem to
congregate in small areas of the ponds as breeding choruses. The
breeding season usually lasts for a short period and all eggs are laid
within a week or two depending on prevailing temperatures. The
males outnumber the females by a ratio of 5 to 1, thus there is strong
competition for a mate. The strong sexual drive in the males may be
the result of this strong competition.

The voice of the male is characterized by a low clicking sound
which can be reproduced by clicking ones tongue against the top of
his mouth. The call is very weak and can normally be heard for only
20-30 feet. The distance compares favorably with that reported by
Turner (1958), but is considerably less than the quarter of a mile
reported by Svihla (1935). Stebbins (1966) stated that the call was
unknown in Rana pretiosa.

July 31, 1969 ecology of r. pretiosa in utah 11

J he eggs are laid shortly after the arrival of females and vary in
number from 147 to 1 160 per clutch, the average being approximately
750 per clutch. The range of eggs per clutch is lower than the 1 100 to
1500 reported by Svihla (1935). It is. however, higher than the 206
to 802 reported by Turner (1958). Turner (1958) rej)orted that his
eggs numbers were based upon counts of eggs in clusters 1 to 3 days
old; Svihla's (1935) reported that the number is an estimation based
upon two egg masses which measured 1500 cc and 1100 cc. His
determinations were made as follows: "Since each egg measured
more than 1 cc the number of eggs in these masses would approxi-
mate 1500 and 1100 respectively." Because these same 1100 and
1500 figures are reported by Livezey and Wright (1947). Wright
and Wright (1949), and Stebbins (1951). and on the basis of data
gathered for this study, it appears that no actual counts were made
by these authors and that their numbers are. at least for the Utah
population, too high.

The eggs average 2.5 mm in diameter and are surrounded by a
distinct outer gelatinous envelope and an indistinct inner one, aver-
aging 10.0 and 5.0 mm respectively. Following the key prepared by
Livezey and Wright (1947). these populations are R. p. pretiosa.

The eggs have required from one week to hatch in the laboratory
to 13-23 days in nature, the majority hatching about two weeks after
oviposition. This two week incubation period is the same as that re-
quired by the frogs in Wyoming as reported by Turner (1958), but
longer than the four days given by Carl (1943) for those in British
Columbia. Johnson (1965) reported the eggs as taking 72 hours to
hatch at 25° C but longer at cooler temperatures.

Several factors were noted which affect the frogs' growth and be-
havior. These were temperature, crowding, and perhaps dissolved
minerals in the water.

Temperature has an effect upon all stages of the life history of
the frog. The adult activity seems to be affected by temperatures
approximately 10-11° C. Temperatures below this point bring on a
quiescence in their activities. The eggs are laid at temperatures above
14° C. These temperatures are higher than the 5° C which Midden-
dorf (1957) reported as being critical in the frogs' activity. The eggs
and tadpoles are subject to a wide range of temperature but growth
is more rapid at the warmer temperatures. Johnson (1965) found
6° and 28° C to be the minimum and maximum temperatures limit-
ing normal embryonic development for this species. Other studies
on temperature tolerance and rates of develojmient are by Moore
(1938. 1939. and 1942).

A high concentration of soluable salts in the water was suspected
of having retarding effects both upon the spring emergence of the
adults and the growth of the larvae. Cameron (1940) observed that
low concentrations of flourine retarded development. Gosner and
Black (1957) studied the effects of acidity on the development and
hatching of frogs, while Merwin and Allee (1943) noted the re-
tarding effects of carbon dioxide on the cleavage rates of frog eggs.

78

MORRIS AND TANNER

The Great Basin Naturalist

Vol. XXIX, No. 2

1966

DAILY MAX. & MIN. TEMP.l'C.^

SEPTEMBER

NOVEMBER

Fig. 17. Air temperature for the fall of 1966. Stars indicate approximate
date of hibernation.

July 31, 1969 ecology of r. pretiosa in utah 79

Crowding of tadpoles was observed to have a retarding effect
upon size and development. This same phenomonon has been ob-
served and studied by Lyrm and Edelman (1936), Rose (1960).
Adolph (1929), Rugh (1934). and Richards (1958) in other species
of Anura.

Metamorphosis was observed to begin on July 27. 1966; this was
approximately 122 days following ovulation. Some tadpoles required
up to 209 days to transform. It may therefore be concluded that at
higher elevations or more northern latitudes having a shorter summer
that tadpoles may require more than one summer to complete their
growth and to transform as reported by Logier (1932) in British Col-
umbia and Turner (1958) in Wyoming.

An examination of the larval mouth parts reveals a tooth row
formula of two rows and three lower rows which according to Svihla
(1935) would place them as R. p. luteiventris.

Hibernation began during the middle of October approximately
two weeks after the first freezing temperatures, and presumably
under the same water where their summer atcivities were concen-
trated.

The adults are variable in key characteristics and normally are
classified as R. p. pretiosa. The ventral coloration is variable between
orange and red. The following color description given by Stebbins
(1966) does not fit the Utah population: "Populations in Nevada.
Utah, Idaho south of Salmon River, and SE Oregon usually have
yellowish ventral color; elsewhere red or salmon predominates."

The Utah pretiosa do not completely fit present keys to the sub-
species and have been identified in various studies as either pretiosa
or luteiventris. The taxonomic characters examined in this study show
an intergradation of sub-specific types. Based upon the eggs and adult
coloration we consider the Utah population to be Rana p. pretiosa.
The susceptibility of the larvae to different water conditions may be
the factor which limits the distribution of the subspecies rather than
the differences in the adults which seem to be ecologically similar.
Wright and Wright (1949) in giving the distribution of the pretiosa
subspecies state that luteiventris occupies the tertiary volcanic areas,
whereas pretiosa prefer the continental deposit areas. The key to the
taxonomy and distribution of this species may very well be the dif-
ferences in ecological tolerance of the immature frog rather than the
adults.

Literature Cited

Adolph, Edward F. 1931. The size of the body and the size of the environ

ment in the growth of the tadpoles. Biological Bulletin, 61:350-375.
Baird, S. F., and C. Girard. 1853. Communication describing Rana preliosa

and Bufo columbiensis . Proc. Acad. Nat. Sci. Phila.. 6:378-379.
Bragg, A. N. 1940a. Observations on the ecology and natural history of Anura.

II. Amer. Midi. Nat., 24:306-321.
. 1940b. Observations on the ecology and natural historj' of Anura. V.

The process of hatching in several species. Proc. Oklahoma Acad. Sci..

20:71-74.
. 1944. Egg laying in leopard frogs. Proc. Oklahoma Acad. Sci.. 24.

The Great Basin Naturalist
80 MORRIS AND TANNER Vol. XXIX. No. 2

Brattstrom, B. H. 1963. A preliminary review of the thermal requirements

of amphibians. Ecology, 44:238-255.
Burke, V. 1933. Bacteria as food for vertebrates. Science, 78:194-195.
Cameron, John A. 1940. Effects of fluorine on hatching and hatching stage in

Rana pipiens. Ecology, 21:288-292.
Carpenter, C. C. 1953a. An ecological survey of the herpetofauna of the

Grand Teton-Jackson Hole area of Wyoming. Copeia, 1953(3) : 1 70-174.

. 1953b. Aggregation behavior of tadpoles of Rana p. pretiosa. Herpe-

tologica, 9:77-78.

. 1954. A study of amphibian movement in the Jackson Hole vsild life

park. Copeia, 1954:197-200.
Dickerson, Mary C. 1906. The frog book. Doubleday Co., New York, xvii-

253.
Dumas, Philip C. 1966. Studies of the Rana species complex of the Pacific

northwest. Copeia, 1966:60-74.
Dunlap, D. G. 1955. Inter- and intraspecific variation in Oregon frogs of the

genus Rana. Amer. Midi. Nat., 54:314-331.
. 1959. Notes on the amphibians and reptiles of Deschutes County,

Oregon. Herpetologica, 15:173-177.
GosNER, Kenneth L., and Irving H. Black. 1959. The effects of acidity on

the development and hatching of New Jersey frogs. Ecology, 38:256-262.
GosNER, Kenneth L. 1960. A simplified table for staging anuran embryos

and larva with notes on identification. Herpetologica. 16:183-190.
Johnson, Oliver W. 1965. Early development, embryonic temperature toler-
ance and rate of development in Rana pretiosa luteiventris Thompson. Un-
published Ph.D. dissertation, Department of Zoology, Oregon State Univer-
sity, Corvallis, Oregon.
King, H. D. 1903. The effects of heat on the development of the toad's egg.

Biol. Bull., 5:218-232.
Limbaugh, B. A., AND E. P. VoLPE. 1957. Early development of the Gulf Coast

toad, Bufo valliceps Wiegmann. Amer. Mus. Novitates, 1842:1-32.
LivEZEY, R. L., AND A. H. Wright. 1947. A synoptic key to the salientian

eggs of the United States. Amer. Midi. Nat., 37:179-222.
Martof, Bernard. 1953. Home range and movement of the green from Rana

clamitans. Ecology, 34:529-543.
Merwin, Ruth M., and W. C. Allee. 1943. The effect of low concentration

of carbon dioxide on the cleavage rates of frogs eggs. Ecology, 24:61-69.
Middendorf, L. J. 1957. Observations on the early spring activities of the

western spotted frog (Rana pretiosa pretiosa) in Gallatin County, Montana.

Proc. Montana Acad. Sci., 17:55-56.
MooRE, John A. 1938. Temperature tolerance and rates of development of the

eggs of amphibians. Anat. Rec, 72(Supp. 1):96.
. 1939. Temperature tolerance and rates of development in the eggs of

amphibia. Ecology, 20:459-476.

1940. Adaptative differences in the egg membranes of frogs. Amer.

Nat., 74:89-93.

Noble, G. K. 1954. The biology of the amphibia. Dover Publications, New
York. 577 front., illus., diagrs.

Richards, Christina M. 1958. The inhibition of growth in crowded Rana
pipiens tadpoles. Physiological Zoology. 31:138-151.

Rose, S. M. 1960. A feedback mechanism of growth control in tadpoles. Ecol-
ogy, 41:188-199.

RuGH, Roberts. 1934. The space factor in the growth rate of tadpoles. Ecology,
15:407-411.

. 1948. Experimental embryology. A manual of techniques and pro-
cedures. Burgess Publishing Co., Minneapolis. 480.

Schmidt, Karl P. 1953. A checklist of North American amphibians and
reptiles. 6th Ed. American Society of Ichthyologists and Herpetologists,
Chicago.

Skousen, Don B. 1952. A taxonomic survey of the eggs and larvae of some
species of Utah amphibia. Unpublished master's thesis. Department of Zo-
ology, Brigham Young University, Provo. Utah.

July 31. 1969 ecology of r. pretiosa in utah 81

Slevin, J. R. 1928. The iimphibia of western North America. Occ. Papers

Calif. Acad. Sci., \&.\i6.
Stebbins, R. C. 1951. Ampliibians of western North America. University of

California Press, Berkeley and Los Angeles. Calif.
. 1954. Amphibians and reptiles of western North America. McGraw

and Hill Book Co.. Inc.. New York.

1966. A field guide to western reptiles and amphibians. Houghton

Mifflin Co., Boston.
Storer. T. I. 1925. A snyopsis of the amphibia of California. Univ. Calif.

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Stejneger, L.. and T. Barbour. 1943. A checklist of North American am-
phibians and reptiles. Bull. Mus. Comp. Zool., 93(l):xix, 260.
SviHLA, Arthur. 1935. Notes on the western spotted frog, Rana p. pretiosa.

Copeia, 1935 (3): 119-122.
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159-198.
Tanner, Wilmer W. 1940. Herpetological specimens added to the Brigham

Young University collection. Great Basin Naturalist. Vo. 1:138-146.
Thompson, H. B. 1913. Description of a new subspecies of Rana pretiosa from

Nevada. Proc. Biol. Soc. Wash., 26:53-56.
Thornton, C. S. 1956. Epidermal modifications in regenerating and in non-
regenerating limbs of anuran larvae. J. Exp. Zoo., 131:373-393, pi. 1. 2.
Turner. Frederick B. 1957. The ecology and morphology of Rana pretiosa

pretiosa in Yellowstone Park, Wyoming. Ph.D. dissertation. University of

California. Berkeley.
. 1958. Life-history of the westeiTi spotted frog in Yellowstone National

Park. Herpetologica, 14:96-100.
Vernberg, John F. 1953. Hibernation studies of two species of salamanders,

Plethodon cinerius cinerius and Eurycea bislineata bislineata. Ecology,

34:55-61.
Wright, A. H., and A. A. Wright. 1949. Handbook of frogs and toads of the

United States and Canada. Comstock Publ. Co.. Ithaca, New York.
Yarrow. H. C. 1875. Report upon the collection of batrachians and reptiles

made in ix)rtions of Nevada. Utah. California, Colorado, New Mexico, and

Arizona, during the years 1871. 1872. 1873, 1874. Wheeler Survey. Vol.

V: 509-633.

FOSSIL CHELONIANS, CHRYSEMYS AND CLEMMYS,
FROM THE UPPER PLIOCENE OF IDAHO

G. R. Zugi

The extant reptilian fauna of the Columbian Plateau and the
northern half of the Basin and Range Province lacks turtles. Their
absence appears to be of relatively recent occurrence, for turtles per-
sist in the surrounding areas and have been found in Plio-Pleistocene
deposits, Glenn Ferry Formation, of this region. The latest faunal
list for the Glenn Ferry Formation (Taylor. 1966. p. 75) includes
Kinosternon sp. and Pseudemy idahoensis. Through the effort of
Dr. C. W. Hibbard and his field parties, a large collection of chelo-
nian shell fragments has been amassed from the vicinity of Hager-
man, Twin Falls County, Idaho. Unless indicated otherwise, the
specimens reported herein are from the Hagerman area.

Family Kinosternidae
Kinosternon Spix

Remarks. There are no elements that can be assigned to this
taxon. Unfortunately, it seems that Taylor (1966) misread Hibbard's
(1959), p. 11) statement. Hibbard indicated only that the shell
fragments may represent "Kinosternon, Chrysemys, or Pseudemy sJ'
This error was compounded by the limited distribution of a faunal
compilation and as such has appeared elsewhere (Holman, 1968).

Family Testudinidae
Chrysemys idahoensis (Gilmore)

Material. Peripherals, UMMP V45276A, V52778; Hyoplas-
tron, UMMP V53417; Xiphiplastra, UMMP V53417, V54129.

Remarks. Of the two turtles represented in this collection, very
few elements can be assigned to C. idahoensis. All Chrysemys ele-
ments were compared with Gilmore's type series and match the
morphology therein. A humeropectoral sulcus extends across the
entire width of the hyoplastron and distally forms an anterior loop
(Gilmore, 1933, PI. 2). Both xiphiplastra possess a strong step later-
ally between the femoral and anal scutes and a shallow dorsal groove
medial of the scutes. These features are characteristic of C. idaho-
ensis.

An anterior half of a plastron (UMMP V42609) has been recov-
ered from the Late Hemphilian Wolf Canyon area of Meade County.
Kansas. The humeropectoral sulcus of this plastron forms an an-
terior loop laterally on each side, indicating that this fossil is also
C. idahoensis. Thus temporally, C. idahoensis extends from approxi-

^United States National Museum, Washington, DC.

82

July 31, 1969 fossil chelorians in idaho 83

mately 3.5 million BP to about 5 million BP and spatially from
south-central Idaho to southwestern Kansas.

In the original description, Gilmore (1933) suggested that the
affinities of this species were with the rubriventris group and not
with Trnchemys, i.e.. the scripta group. The reverse, however, seems
to be true. In the rubriventris group, the skull characteristically
possesses a beak bordered by cusps, a high, serrated, median ridge
with a pair of anteromedial cusps on the dorsal alveolar surface, a
shallow posteromedial indentation of the dorsal alveolar surface, and
a small pro-otic tuberosity. C. idahoensis and the scripta group have
no cusp on the beak, a low. smooth, median ridge with no cusps, a
deep posteromedial indentation of the dorsal alveolar surface, and a
well developed pro-otic tuberosity. Unlike the skulls of the rubriven-
tris or the scripta group, the anterior edge of the inferior process of
the parietal is buttress-like in C. idahoensis, similar to the condition
seen in Malaclemys geographica. This buttress-like process and the
broad alveolar surfaces suggest that this species feed on mollusks.

Clemmys owyheensis Brattstrom and Sturri

Material. Nuchals, UMMP V34444, V49940, V50201, V53128.
V53527,V53556, V53731. V53773; Peripherals, UMMP V45276 (2).
V49956, V50349 (14), V52778C-E. V53128 (7), V53264 (6).
V53445 (7), V53555, V54595 (5).V55477 (2), V55537; Pvgals.
UMMP V34444 (4). V49558 (2). V50349 (3), V50553 (2). V50938.
V51132. V51277, V51737. V53128 (2). V53264. V53731 (2).
V54706, V56076 (3), V56261. V56267; Epiplastra. UMMP V34444.
V50349 (3), V53264 (2), V53445, V53553 (2). V53755; Endoplas-
tra, UMMP V34444, V50091. V50349. V53128, V53264 (3). V53445
(2). V53731, V53772 (2), V56327; Hyoplastra, UMMP V53128.
V53528 (2); Hypoplastra, UMMP V52778, V53264, V53421.
V53555; Xiphiplastra, UMMP V53264 (2), V53445 (3), V53553.
V53731.

Remarks. Brattstrom and Sturn (1959) described Clemmys
owyheensis on an endoplastron from a Hemphillian (Middle Plio-
cene) locality in Oregon. C. owyheensis is characterized by each
gulahumeral sulcus of the endoplastron forming a 33-degree angle
or less with the midline, a prominent anterior knob on the endojjlas-
tron. and a humeropectoral sulcus crossing the endoplastron. 1 he
last character occurs in both Terrapene and Clemmys. The endo-
plastron of Terrapene is diamond-shaped and in adult specimens
inseparably fused with the epiplastra and hyoplastra. The range of
variation of the gulahumeral angle is 24 to 37 degrees for the endo-
plastra from the Hagerman and 31 to 47 degrees for nuchals from
seven extant C. marmorata. Some recent C. marmorata possess
prominent anterior knobs on the end()[)lastra. Thus on the basis of
endoplastral characters, the fossil Clemmys from the Pliocene cannot
be separated from the extant C. marmorata. However, the variation
in the morphology of the other shell elements from Hagerman. Idaho,
suggests that the fossil remains represent a distinct population. Since

The Great Basin Naturalist

84 G. R. zuG Vol. XXIX. No. 2

the Hagerman endoplastra match the diagnosis of Brattstrom and
Sturn, Clemmys owyheensis is recognized, at least until more mater-
ial is available, as a distinct species. The conspecificity of the Hager-
man fossil Clemmys and C. oivyheensis is strengthened by their oc-
currence in the same drainage system, the Idaho Lake system. Tem-
porally, these two Clemmys may be separated by as much as 5.5
million years.

The nuchals of C. owyheensis differ from those of Malaclemys
and the Chrysemys floridana group by an extensive nuchal scute
underlay and from all Chrysemys by a proportionately narrower
nuchal scute. The Clemmys nuchal can be distinguished from that
of Emydoidea, because in the latter the anterolateral border of the
first vertebral scute is rarely impressed on the bone. If the sulcus of
vertebral scute is present, it lies completely posterior to the greatest
width of the bone. In Terrapene, the anterolateral tips of the first
vertebral scute do not extend beyond the margin of the nuchal bone.
The fossil nuchals fall into the range of variation seen in Clemmys,
but cannot be assigned to any particular extant species. Dorsally
the fossil nuchals match the appearance of those of C. guttata and
marmorata with the anterolateral corners of the first vertebral scute
extending off the nuchal onto the first peripheral. In C. insculpta,
these corners do not approach the edge of the nuchal closely, whereas
the corners in C. muhlenbergi end at the edge as in Terrapene. Ven-
trally the nuchals of C. owyheensis possess a long nuchal scute un-
derlay as in C. insculpta and guttata.

The pygals of C. owyheensis are bifurcate posteriorly and have
posteriorly converging lateral sides. Only C. m. marmorata possesses
this condition; all other Clemmys have only a shallow nick. In
Chrysemys, pygals with deep notches have parallel or posteriorly
diverging sides.

The epiplastra from Hagerman have the deep gular overlay and
thick gular lip characteristic of Clemmys. The anterior edge of the
gular lip is smooth and bordered laterally by a distinct horn on each
side. Likewise, the fossil hyoplastra are from Clemmys, for they have
an anterior humeropectoral sulcus. On the hypoplastra of C. owy-
heensis, the abdominomarginal sulcus coincides with the hypoplas-
troperipheral suture, and a deep groove extends posteriorly from the
inguinal buttress along the abdominal and femoral scutes onto the
xiphiplastron. The condition of the hypoplastron is commonly ob-
served in Clemmys. Each xiphiplastron also possesses a deep dorsal
groove, and in addition the transition laterally between anal and
femoral scutes is smooth, i.e., no step is formed.

Discussion and Summary

The Hagerman chelonian fauna consists of two species, Chryse-
mys idahoensis and Clemmys owyheensis. No turtles occur in this
region today, although Stebbins (1954,192) lists an unconfirmed
report of a C. marmorata from Eagles Nest, Idaho, which is upstream
of the Hagerman locality. If this record is excluded, the closest

July 31, 1969 fossil chelorians in idaho 85

Clemmys population is C. rnarniorata in the True kee iind Carson
rivers of Nevada. It has been suggested that the Nevada poi)ulation
is introduced; however, two factors hint that it may be a natural
population. Although C owyheensis shares some characteristics with
the eastern species of Clemmys^ it appears to be closely related to
C. rnarniorata. Furthermore, the absence of inguinal scutes on the
fossil hypoplastra indicates that the 1 lagerman Clemmys are related
to the southern race of C. marmorala. Other aquatic animals, mol-
lusks (Taylor, 1966) and fish (Miller, 1965), show this relationship
between faunas of southern Idaho and central California. This has
led to the postulation of a different drainage pattern for the Snake
River during the Pliocene and most of the Pleistocene. The course
of the river was from southeastern Oregon across northwestern Ne-
vada to central California. Thus the Nevada population of C. mar-
morala may be a relict as its present distribution coincides with this
old drainage pattern.

The shell elements of the Hagerman turtles were obtained from
several localities representing a temporal span of about 100.000
years (Fig. 1). Clemmys occur throughout this entire temporal span.
In contrast, unquestionable remains of Chrysemys are clustered
around the Horse Quarry (Fig. 1). This is a strange occurrence, for
the buttress-hke inferior parietal process and broad alveolar surfaces
of C. idahoensis indicate that this taxon was molluscivorous. Taylor
(1966) reported the near absence of mollusks from this stratigraphic
position. Accordingly, if C. idahoensis were entirely molluscivorous
as has been suggested for the minnow, Mylocyprinus (Uyeno, 1961),
it should also be absent from the Lake Idaho fauna of that time and
would be expected to occur earlier in assocation with the abundant
mollusk fauna.

At the present, C idahoensis has been found in Kansas and Idaho.
This distribution is not matched by either of its suggested relatives,
rubriventris (Gilmore, 1933) or scripta. The geographic proximity of
C. scripta attests to the likelihood of C. idahoensis being a derivative
of the scripta lineage rather than the Atlantic coastal plain rubri-
ventris. The cranial characteristics of C. idahoensis also seem to be
more similar to scripta than rubriventris.

Acknowledgments

The collection of fossil turtles from Hagerman, Idaho, was made
by Dr. Hibbard and his students, who were supported by a grant
from the National Science Foundation, GB-1528. I wish to thank
Dr. C. W. Hibbard. University of Michigan Museum of Paleont-
ology (UMMP). for the opportunity to examine the Hagerman
material. Dr. N. Hotton, United States National Museum (USNM).
permitted me to examine the types of C. idahoensis. Skeletons of
the extant species of Clemmys were examined through the courtesy
of Dr. C. F. Walker. University of Michigan Museum of Zoology.
Drs. C. W. ilibbard and J. A. Peters have improved the niaiuiscript

The Great Basin Naturalist

86 G. R. zuG Vol. XXIX, No. 2

ELEVATION CHRYSEMYS CLEMMYS STRATIGRAPHIC

(ft.) MARKER

3400

50

3300

X* X Horse Quarry

X

50 X

3200

iBasal

3.48 million years
X
? X

3100 X

X

50 X

X
3000 X Peters Gulch Ash

50

2900

Figure 1. The stratigraphic distribution of Chrysemys idahoensis and Clem-
mys owyheensis in the vicinity of Hagerman, Idaho. The elevation is the height
of fossiliferous strata above sea level. X, specimen(s) in the University of Michi-
gan Museum of Paleontology; star, the type specimens of C. idahoensis (USNM
12059-60).

July 31, 1969 fossil chelorians in idaho 87

by their editorial comments. Dr. P. Bjork greatly aided the pre-
paration of the figure.

Literature Cited

Brattstrom, B. H. and a. Sturn. 1959. A new species of fossil turtle from

the Pliocene of Oregon, with notes on other fossil Clemmys from western

North America. Bull. So. Calif. Acad. Sci., 58: 65-71.
GiLMORE, C. W. 1933. A new species of extinct turtle from the Upper Plio-
cene of Idaho. Proc. U.S. Natl. Mus., 82 (9): 1-7.
HiBBARD, C. W. 1959. Late Cenozoic microtine rodents from Wyoming and

Idaho. Papers Mich. Acad. Sci., Arts, Lett., 44: 3-40.
HoLMAN, J. A. 1968. LIpper Pliocene snakes from Idaho. Copeia, 1968: 152-

158.
Miller, R. R. 1965. Quaternary freshwater fishes of North America. In: The

Quaternary of the United States. H. E. Wright, Jr. and D. G. Frey, eds.,

pp. 596-581. Princeton Univ. Press, Princeton, N. J.
Stebbins, R. C. 1954. Amphibian and Reptiles of Western North America.

i-xxii, 536 pp. McGraw-Hill Book Co., Inc., N. Y.
Taylor, D. W. 1966. Summary of North American Blancan nonmarine mol-

lusks. Malacologia, 4: 1-172.
Uyeno, T. 1961. Late Cenozoic cyprinid fishes from Idaho with notes on other

fossil minnows in North America. Papers Mich. Acad. Sci., Arts. Lett., 46:

329-344.

NOTES ON THE DISTRIBUTION AND BIONOMICS
OF SOME IDAHO CERAMBYCIDAE (COLEOPTERA)i

William F. Barr and Richard L. Penrose-

Distributional and bionomical information concerning some
species of Idaho Cerambycidae, especially those associated with range-
land shrubs, has been accumulated over the past several years pri-
marily during the course of field studies. Although fragmentary in
scope, the more pertinent of this information is presented at this
time for use by other workers. Representatives of each species
treated in this paper are deposited in the University of Idaho insect
collections.

Thanks is given to J. A. Chemsak and E. G. Linsley, University
of California, and J. N. Knull, Ohio State University, for their assis-
ance in the determination of some specimens.

Ammanus vittigera LeConte

Linsley (1962) recorded the known northern limits of the dis-
tributional range of this southwestern species as southern Utah. Two
specimens have been taken in southwestern Idaho. One was associated
with a stand of Atriplex canescens (Pursh) Nutt. four miles southeast
of Grandview, Owyhee County. June 20, 1959 (W. F. Barr), and the
other was collected from the foliage of A. confertifolia (Torr. &
Frem.) Wats, eight miles northwest of Walters Ferry, Canyon
County, June 15, 1961 (J. E. Henry).

Asemum caseyi Linsley

This species known previously from the Pacific Coast region is
recorded from Idaho for the first time on the basis of specimens col-
lected at Hot Springs, nine miles north of Mountain Home. Elmore
County, June 16, 1957 (W. F. Barr) and Pollock, Idaho County,
June 7, 1956 (R. W. Portman).

Asemum nitidum LeConte

This common Pacific Coast species has not been recorded pre-
viously from Idaho. Collection records from the state are: Craters
of the Moon National Mommient, Butte County, July 15, 1964
(D.S. Horning. Jr.), on Pinus flexilis James, Moscow, Latah County,
August 28, 1954 (W. F. Barr); and Troy, Latah County, August
30, 1954 (R. H. Abbott).

^Published with the approval of tlie Director of the Agricultural Experiment Station as research
paper no. 7+6.

^University of Idaho, Moscow, Idaho.

July il. 1969 NOTES OI- IDAHO CKHAMHVCIDAK 89

Atiniia dor soli s LeConte

A specimen from Wallace. July 9, 1925 (O. Houllenianii) and a
specimen labeled "west central Idaho" collected by W. E. Ferguson
in June 1963 represent new distributional records for the state.

Crossidius ater LeConte

Although this species has been reported from Idaho (Linsley and
Chemsak. 1961). its known distributional range within the state
now can be expanded. New collection sites are: six miles south of
Hailey, Blaine County, August 7, 1966 (M. A. Brusven) ; three
miles north and three miles northwest of Malta. Cassia Countv.
September 12. 1965 (E. J. Allen, W. F. Barr, R. L. Penrose);
Craters of the Moon National Monument, Butte County. September
6, 1964 (D. S. Horning, Jr.); 1.5 miles west of Dayton. Franklin
County, September 13. 1965 (W. F. Barr); 1.5 miles northeast of
Baker. Lemhi County, September 2. 1965 (R. L. Westcott); Haynes
Creek, 4.5 miles north of Tendoy, Lemhi County. August 28. 1962
(W. F. Barr); 19 miles north of Shoshone. Lincoln County. Septem-
ber 2. 1965 (R. L. Westcott); and 10 miles southwest of Midvale.
Washington County. September 12, 1948 (W. F. Barr).

Adults have been taken on flowers of Chrysothamnus nauseosus
(Pall.) Britt., C. viscidiflorus typicus (Hall & Clem.) Cron., C v.
latifolius (D. C. Eat.) Rydb., C. v. lanceolatus Nutt. and Guderrizia
sarothrae (Pursh) Britt. and Rusby in the state. In the Malta area
larvae and a teneral adult have been found in the roots of Artemisia
tridentata Nutt.

Crossidius discoideus blandi Casey

Idaho collection records of adults, pupae, and larvae are available
for this subspecies indicating that it ranges into the more southeast-
erly portions of the state. Collection sites are: two miles north of
Swan Lake. Bannock County, July 25. 1967 (R. L. Penrose) (larv^ae
and pupae); one mile east of Basin, Cassia County, June 15, 1967
(W. F. Barr, S. M. Hogue, R. L. Penrose) (larvae); three miles
northwest of Malta, Cassia County. September 11 and 12. 1965 (E.
J. Allen, W. F. Barr. R. L. Penrose) (adults); 2.5 miles north of
Dayton, Franklin County. September 13, 1965 (W. F. Barr. R. L.
Penrose) (adults); Register Rocks, Power County. January 28. 1967
(R. L.Penrose) (Larvae); 9.1 miles south of Rockland, Power Coun-
ty. July 25. 1967 (R. L. Penrose) (larvae and jiupae). .Adults have
been found mainly on the flowers of Gutierrezia sarothrae (Pursh)
Britt. and Rusbv and occasionally on the flowers of Chrysothamnus
viscidiflorus (Hook.) Nutt. Larvae and pupae have been found only
in the roots of G. sarothrae. Examination of this plant where it oc-
curs in southwestern and central Idaho has thus far failed to disclose
the presence of C. d. blandi.

The Great BelsIh Naturalist
90 BARR AND PENROSE Vol. XXIX. No. 2

Crossidius pulchellus LeConte

The occurrence of this species in Idaho is based on a collection
record from Riverdale, Franklin County (Linsley and Chemsak,
1961). Additional collections have been made from three miles west
of Dayton, Franklin County, September 13, 1965 (R. L. Penrose);
2.5 miles north of Dayton, Franklin County, September 13, 1965
(W. F. Barr); and Bannock Pass, Lemhi County, August 30, 1962
(W. F. Barr). Adults were taken on the flowers of Chrysothamnus
viscidiflorus (Hook.) Nutt, C. nauseosus (Pall.) Britt., and Gutier-
rezia sarothrae (Pursh) Britt. and Rusby.

Elatotrypes hoferi Fisher

Two small series of specimens were reared from dead wood of
Pinus flexilis James at the Craters of the Moon National Monument,
Butte County, Idaho, by D. S. Horning, Jr., during the summers of
1964 and 1965. These collections establish the occurrence of this
species in the state for the first time.

Evodinus vancouveri Casey

This species, not previously recorded from the state, has been
collected only at one northern Idaho location. Specimens were taken
at Laird Park, Latah County, May 14, 1962 (W. F. Barr and S. D.
Smith) and May 20, 1967 (R. L. Westcott). In both instances adults
were found on the flowers of Trillium ovatum Pursh growing in a
dense or partially open coniferous forest comprised mostly of Abies
grandis Lindl., Pseudotseuga menzesii (Murb.) Franco, and Pinus
monticola Dougl.

Mecas bicallosa Martin

This species is widely distributed in southern Idaho. It has been
taken in several different habitats present in foothill areas, in moun-
tain valleys, on a mountain pass, and on the Snake River Plains at
elevations that range from approximately 3100 feet to 8752 feet.
Collection records include: Galena Summit, Blaine County, July 22,
1961 (W. F. Barr, R. B. Hawkes), July 30, 1967 (S. M. Hogue); 5.5
miles southwest of Almo, Cassia County, June 11, 1962 (W. F. Barr);
Bear Creek Camp, Custer County, July 21, 1964 (R. L. Westcott);
Bonanza, Custer County, July 22, 1965 (W. F. Barr); Weston Can-
yon, Franklin County, July 11, 1964 (O. O. Fillmore); six miles
southwest of Sweet, Gem County, June 19, 1959 (J. E. Henry); ten
miles northeast of Richfield, Lincoln County, June 25, 1959 (W. F.
Barr); three miles south of Driggs, Teton County, July 11, 1965 (O.
O. Fillmore) ; and nine miles southwest of Rogerson, Twin Falls
County, June 18, 1961 (W. F. Barr). These collections have been
associated with Artemisia tridentata Nutt., A. t. var vesseyena
(Rydb.), A. tripartita Rydb. and A. arbuscula Nutt.

July 31, 1969 notes of idaho cerambycidae 91

Megacheuma brevipennis (LeConte)

This attractive species is known to have a rather broad but dis-
continuous distributional range in the western United States. How-
ever, it appears to occur most abundantly in the more northern por-
tions of the Great Basin and adjoining areas.

The host plants of this species in Idaho have been established as
Sarcobatus vermiculatus (Hook.) Torr.. Atriplex nuttalUi S. Wats.,
and A. confertifolia (Torr.) Wats. PTowever, this cerambycid has not
been found throughout the distributional ranges of these host plants
in the state. Collection records are as follows: three miles east of
Howe, Butte County, June 22, 1966 (W. F. Barr) (larvae in A. nut-
talUi), August 7. 1955 (W. F. Barr) (adult in pupal cell in A. nut-
talUi); ten miles north of Howe, Butte County, April 11. 1958 (W.
F. Barr) (larvae in A. nuttalUi), June 22, 1967 (W. F. Barr) (larvae
in A. confertifolia). August 20, 1966 (R. L. Penrose) larvae in A.
nuttalUi). August 31, 1960 (J. E. Henry) (adults); 18 miles north-
east of Howe. Butte County, June 14. 1955 (W. F. Barr) (larvae in
A. nuttalUi), July 14, 1956 (W. F. Barr) (pupae in A. nuttalli) ;
five miles north of Malta, Cassia County, July 23, 1957 (R. A. Mac-
kie) (adults); six miles southeast of Malta, Cassia County, May 31.
1958 (W. F. Barr) (larvae and pupae in S. vermiculatus) . September
6 and 15, 1955 (W. F. Barr and R. A. Mackie) (adults); seven miles
southeast of Malta, Cassia County, April 30, 1954 (W. F. Barr) (lar-
vae in A. nuttalUi) ; four miles east of Idahome, Cassia County. June
1, 1958 (W. F. Barr) (larvae and pupae in S. vermiculatus) , August
9, 1955 (W. F. Barr) (larvae and adults in S. vermiculatus); five
miles northwest of Oakley, Cassia County, June 2, 1958 (W. F. Barr)
(larvae in A. confertifolia), August 16, 1957 (W. F. Barr) (larvae
and adults in A. nuttalUi), October 15. 1956 (W. F. Barr) (larvae
in A. confertifolia) ; six miles south of Challis. Custer County, Sep-
tember 17, 1965 (W. F. Barr and R. L. Penrose) (larvae in A. con-
fertifolia); 12 miles west of Mountain Home, Elmore County. April

15 and August 11, 1953 (W. F. Barr) (larvae in A. nuttalUi). June

16 and November 16, 1955 (W. F. Barr) (larvae in A. nuttalUi); 12
miles northwest of Terreton, Jefferson County. July 14, 1956 (W.
F. Barr) (larvae and pupae in A. nuttalUi). August 30, 1962 (W. F.
Barr) (adults) ; 20 miles northwest of Terreton. Jefferson County,
March 27, 1967 (W. F. Barr) (larvae in A. confertifolia); one mile
north of Murphy. Owyhee County, March 25, 1955 (W. F. Barr)
(larvae in ^9. vermiculatus) ; nine miles northwest of Grandview.
Owyhee County, May 27, 1958 (W. F. Barr) (larvae in A. nuttalUi).
August 10, 1953 (W. F. Barr) (larvae and pupae in A. nuttalUi),
August 21, 1966 (E. J. Allen) (adults). September 14. 1955 (W. F.
Barr and A. R. Gittins) (adults).

Observations and rearings indicate that the larva of the clerid
beetle. Enoclerus acerbus Wolcott, preys on larvae and pupae of M.
brevipennis in the roots of the three host plant species.

The Great Basin Naturalist
92 BARR AND PENROSE Vol. XXIX, No. 2

Megacyllene robinae (Forst.)

This well-known species apparently has become established in
several areas of southern Idaho. Collection records are available from
Pocatello, Bannock County, September 28 and October 4, 1965; Good-
ing, Gooding County, in mid-September (E. Koster) ; and Idaho Falls,
Bonneville County, October 16, 1967. It has also been taken at Spo-
kane, Washington. August 5, 1963 (S. M. Hogue) where it was
found on apple wood.

Meriellum proteus (Kirby)

The expected occurrence of this species in the state is confirmed
by a single specimen from Moscow Mountain, Latah County, Aug-
ust 1, 1953 (R. H. Abbott).

Monochamus notatus morgani Hopping

A single Idaho collection record is now available for this sub-
species from Elk River, Clearwater County, August 30, 1963 (R.
Hoff). An adjacent state record is Swan Lake, Lake County, Mon-
tana, July 18, 1963 (M. M. Furniss) on Pinus monticola Dougl.

Neoclytus balteatus LeConte

This species has been taken at three widely separated localities
in southern Idaho. They are: Juniper Mountain, Owyhee County,
August 15, 1963 (M. M. Furniss); Smith Prairie, Elmore County,
August 14, 1963 (R. E. Miller); and St. Anthony, Fremont County,
July 25, 1963 (R. E. Miller). The altitudinal range of these locations
is from 4700 feet to 6800 feet.

It may be of interest to note that no species of Ceanothus, the
recorded host plant genus of A^. balteatus, occurs in the St. Anthony
area, thus suggesting a broader host range than is presently known
for this ceranbycid.

Prionus {Homaesthesis) integer LeConte

This species has been recorded previously from southcentral and
southeastern Idaho (Linsley, 1962). A collection from Parma, Can-
yon County, July 10, 1953, now establishes its distributional range
across the southern portion of the state. A specimen from Moscow,
Latah County, September 30, 1931, in northern Idaho must be re-
garded as suspected mislabeling.

Adults of P. integer are most commonly taken at lights. Larvae
have been associated with the roots of Artemisia tridentata Nutt. and
Chrysothamnus viscidi floras (Hook.) Nutt. On one occasion larvae
were found damaging newly planted bean seeds in a field near Bur-
ley, Cassia County, and in another instance, cutting underground

July 31. 1969 notes of idaho ckrambycidae 93

stems of potato plants in a field near American Falls, Power County.
In both cases the fields had been recently cleared for cultivation.

Rosalia funebris Motschusky

This attractive species, previously unreported from Idaho, is un-
common but widely distributed in the state. It has been collected at
Rocky Point. Benewah County, July 25, 1964 (R. W. Portman);
Sandpoint. Bonner County. July 26, 1932 (W. Virgin); Moscow.
Latah County, September 5, 1965 (IT. C. Manis). August 1951 (W.
F. Barr), June 1950 (H. C. Manis), June 23, 1961 (C. M. Ott), July
28, 1961 (H. C. Manis); Culdesac. Nez Perce County. July 28, 1934
(C. Wakeland); and Twin Falls, Twin Falls County. July 24. 1960
(R. L. Williamson) and August 18, 1953 (A. R. Gittins).

Specimens from Twin Falls have been reared from the wood of
box elder Acer negundo Linn, and some specimens from Moscow are
labeled "apple."

Semanotus juniperi (Fisher)

A significant extension of the distributional range of S. juniperi
is based on a collection from the west slope of Black Pine Mountains.
Cassia County, Idaho, June 10, 1954 (W. F. Barr). A single speci-
men was dug from the wood of Juniper us osteosperma (Torr.) Little.
This species was reported by Linsley (1964) as occuring in the
"mountains of southern Arizona."

Semanotus ligneus amplus (Casey)

Specimens from northern Idaho localities can be referred to
either S. I. amplus or S. I. basalis (Casey) on the basis of elytral
markings. I'he known host plants of the former subspecies do not
occur in this area whereas the host of the latter subspecies does. From
the standpoint of distribution as given by Linsley (1964), these Idaho
collections would best constitute an extension of the range of .V. /.
amplus.

Because of the distinct possibility of these two forms being synon-
ymous it is perhaps best to consider the Idaho material as S. I. am-
plus. The collection records are: seven miles east of Laird Park. I^tah
County, April 17. 1962 (S. D. Smith); Moscow Mountain. Latah
County, May 10. 1957 (G. N. Knopf); and St. Joe River. Shoshone
County, May 21. 1941 (O. Huellemann).

Semanotus ligneus conformis (Casey)
Dead specimens assignable to this subspecies were dug from dead
wood of Juniperus osteosperma (Torr.) Little five miles southwest of
Oakley. Cassia County, June 15, 1967 (R. L. Penrose). Its occurronce
in Idaho is not unexpected, inasmuch as Linsley (1964) recorded tins
subspecies from northern Utah.

The Great Basin Naturalist
94 BARR AND PENROSE Vol. XXIX. No. 2

Toxotus obtusus LeConte

This dimorphic species is known from several states and Canadian
provinces of the Pacific Northwest and northern Rocky Mountains
(Hopping, 1937). but has not been recorded from Idaho. Present
records for the state include: Targhee Pass. Fremont County, July
15. 1965 (R. L. Westcott); two miles southwest of Bannock Pass.
Lemhi County. July 23. 1965 (R. L. Westcott); and two miles south
of Troy. Latah County. June 11, 1965 (A. P. Gupta). The Bannock
Pass specimens were found crawling on the soil surface and on the
steins of Lupinus sp.

Typocerus bait eat us Horn

In Idaho this species appears to be confined to the broad areas
of the southcentral and southeastern portions of the state at elevations
ranging from 4800 to 6000 feet. Collections have been made at Elba-
Basin Pass. Cassia County. August 23. 1958 (W. F. Barr). Augvist
25, 1959 (J. E. Henry) and September 10, 1965 (W. F. Barr and R.
L. Penrose) ; five miles southwest of St. Anthony. Fremont County,
September 6, 1967 (W. F. Barr and J. M. Gillespie) and September
14, 1965 (W. F. Barr); four miles east of Menan. Jefferson County,
August 31, 1965 (R. L. Westcott); Register Rocks, Power County,
August 15, 1966 (R. L. Penrose); three miles west of Tetonia. Teton
County, August 11, 1966 (L. S. Hawkins); Rock Creek Ranger Sta-
tion, Twin Falls County, July 8, 1965; and Rogerson, Twin Falls
County, September 8. 1955 (R. A. Mackie). Most specimens were
taken on the flowers of Chrysothamnus viscidiflorus (Hook.) Nutt.
and a few on the flowers of C. nauseosus (Pall.) Britt.

A specimen from the Tetonia collection exhibits a considerable
reduction in elytra markings. The subbasal fascia is reduced to a pair
of small discal spots, the median fascia to a pair of transverse spots,
and the post median fascia is absent. Other specimens from this
collection exhibit a tendency towards this extreme condition.

Ulochaetes leoninus LeConte

The occurrence of this distinctive species in Idaho is based on
several collections from the following localities in the northern part
of the state: Flat Creek. Latah County, June 1952 (T. F. McGill);
south slope of Moscow Mountain, Latah County, August 22. 1964
(R. L. Westcott); and Troy, Latah County, August 6 and September
8. 1954 (R. H. Abbott). Linsley (1940) recorded it previously from
British Columbia. Washington. Oregon. California and Nevada.

Xylotrechus mormonus (LeConte)

Only four specimens of this northern species are in the Univer-
sity of Idaho collection. They were collected at the Craters of the
Moon National Monument, Butte County. July 11, 1964 (D. S. Horn-

July 31. 1969 notes of idaho cerambycidae 9'i

ing); Basin. Cassia County, Juno 14. 1953 (P. Ashlock); Murphy
Hot Springs. Owyhee County, June 20. 1965 (W. F. Barr) ; and
Donnelly. Valley County, July 18, 1961 (W. F. Barr). No host in-
formation is available for these Idaho specimens which constitute
the first records for the state.

Literature Cited

Hopping. R. 1937. The Lepturini of America North of Mexico. Part II. Nation-
al Museum of Canada, Bull. no. 85. pp. 1-42.

LiNsi.EY. E. G. 1940. A Revision of the North American Necydalini (Coleop-
tera, Cerambycidae). Annals of the Entomological Society of America. 33
(2): 269-281. ■

1962. The Cerambycidae of North America. Part II. Taxonomy and
Classification of the Parandrinae. Prioninae, Spondylinae. and Aseminae.
University of California Publications in Entomology, 19: 1-102.

. 1962. The Cerambycidae of North America. Part III. Taxonomy and

Classification of the Subfamily Cerambycinae tribes Opsimini through Mega-
derini. University of California Publications in Entomology, 20: 1-188.

. 1964. The Cerambycidae of North America. Part V. Taxonomy and

Classification of the Subfamily Cerambycinae, tribes Callichromini through
Ancylocerini. University of California Publications in Entomology. 22: 1-197.

LiNsi.EY, E. G. AND J. A. Chemsak. 1961. A Distributional and Taxonomic
Study of the Genus Crossidius (Coleoptera, Cerambycidae). Miscellaneous
Publications of the Entomological Society of America. 3 (2): 25-64.

OVERWINTERING PHYTOSEIID MIFES IN
CENTRAL UTAH APPLE ORCHARDS^

John W. Leethani and Clive D. Jorgensen-

Predaceous typhlodromids ■ have been observed overwintering as
adult females in fruit trees of northern temperate regions, but
specific studies have been rather limited and no taxonomic or
ecologic studies have been reported from Utah. Gilliatt (1935) and
Garman and Townsend (1938) suggested that some adult phytoseiids
overwintered under the bark of trees while others may seek shelter
on the ground. Gilliatt (1935) further reported that adult Seiulus
pomi Parrott (since divided into a number of Typhlodromus spp.)
overwintered on twigs in Nova Scotia, concealed in such places as
under rough bark, in old bud moth hibernacula. and in deserted oy-
stershell scales {Lepidosaphes ulmi Linn.). Herbert (1953) found
females of several species overwintering under bark scales of tree
trunks and larger branches, in the empty hibernacula of eye-spotted
bud moths (Spilonota ocellana D. and S.) . in empty oystershell scales,
and in empty codling moth {Carpocapsa pomoneUa Linn.) cocoons.
Chant (1959) found Typhlodromus bakeri (Garman) overwintering
in deep crevices around scars and wounds and in the splintered ends
of dying twigs. Rurrell and McCormick (1964) reported three species
{Typhlodromus longipilus Nesbitt. Typhlodromus rhenanus (Oude-
mans). and Typhlodromus occidentalis Nesbitt) hibernating under
loose bark of lower tree trunks and in debris around the bases of
trees.

In a study of the overwintering sites in Ontario peach orchards
large numbers of mites (mostly T. rhenanus) were found in bark
crevices and deep protected cracks of cankers (Putman. 1959). Some
were found in the splintered ends of broken or pruned branches, but
none were found in superficially rough areas of living bark on the
smaller limbs. Very few were found on twigs that were free of old
Lecanium scales although large numbers were taken from twigs pos-
sessing old scales.

Only one investigation of the movement of mites from leaves to
overwintering sites has been reported (Chant. 1959). He found no
mites on leaves that had yellowed, but reported increases in the pop-
ulations of Typhlodromus finlandicus Oudemans. Typhlodromus
tiliae Oudemans (:= pyri Scheuten of Chant. 1959), and Typhlod-
romus umbricatus Chant on the green leaves and twigs. Since mites
were not found in the soil and cover, he concluded that they abandon

'This study was supported, in part, by grant no. 12-14-100-3029 (33) from the U.S. Depart-
ment of Agriculture to Brigham Young (jniversity, and was completed by the senior author in
partial fulfillment of the requirements for a Master of Science degree at Brigham Young
University.

=John \y. Leetham is at the Dept. of Zool., Col. State Univ.. Ft. Collins, Col.; Clive D.
Jorgensen is at the Dept. of Zool. and Entomol., Brigham Young Univ., Provo. Utah.

•"The nomenclature proposed by (^hant (1965) was adopted for this study.

96

July 31.1969 ovkrwintf.ring phytoseiid mitks 97

the yellowing leaves and move to the hranches and trunk to over-
winter.

Mortality rates of overwintering phytoseiids have been investi-
gated only incidentally. Anderson and Morgan (1958) reported high
winter mortality rates for several unidentified species of phytoseiids
in British (>)lumbia. Their cak ulations were made by comparing
mite numbers on leaves in the fall and spring, the difference being
attributed to winter mortality. Using similar methods. Chant ( 1959)
found more than 90% mortality in overwintering T. tiliae in Eng-
land. Putman (1959) found mortality of Amblyseius faUacis (Gar-
man) in Ontario peach orchards to be "very high."

The purposes of this study were to identify the species of phyto-
seiids overwintering as adults in central Utah orchards, to determine
their overwintering sites, and estimate winter mortality rates and fall
movements of Typhlodromus mcgregori Chant and 7. occidentalis—
both of which are abundant in Utah apple orchards (Lee and Davis,
1968).

Methods

This study was made during three consecutive winters, 1965-
1968. in Utah County. Utah. Eleven derelict apple orchards (Jorgen-
sen. 1967) located throughout Utah County were qualitatively
sampled during the first winter to determine the species of mites
overwintering as adults and to locate their overwintering sites. Soil,
ground cover, twigs and spurs, and bark were sampled extensively
from each orchard. Berlese funnels were used to isolate mites from
the collected materials. Occasional samples of crotch refuse, aban-
doned bird nests, and mmrunified fruit still clinging to the trees were
collected. Twig-spur and bark samples were also collected from dere-
lict orchards of other tree fruit varieties. Frequency charts were then
developed to determine the primary overwintering sites of the most
prevalent phytoseiid species.

Movements of T. mcgregori and T. occidentalis from leaves to
their respective overwintering sites were investigated during October
and November, 1967. Two ap|)le orchards were used; a conunercial
orchard in Orem. Utah for T. occidentalis, and a derelict orchard
north of American Fork. Utah for 7'. mcgregori. Samples of 500
leaves were taken from five selected trees in each orchard once a
week. Samples of leaves falling from each tree were also taken to
check for mites falling to the ground with the leaves. Collections of
falling leaves were made by placing a drop cloth beneath the trees
and gently shaking the branches to dislodge the leaves. These data
were analyzed with a /-test to determine whether or not the mean
numbers of mites on dropping leaves were significantly smaller than
the numbers found on green leaves still adhering to the trees. Incid-
ental collections of twig-si^urs and bark were also taken from the
commercial orchard in Orem.

The specific location of overwintering sites was investigated dur-
ing the winters of 1966 to 1968. In 1966. about 3{)() twigs and spurs

The Great Btisin Naturalist
98 LEETHAM AND JORGENSEN Vol. XXIX, No. 2

from a derelict orchard in Mapleton, Utah were dissected to deter-
mine the specific overwintering sites of phytoseiids. Five samples of
500 twig-spurs were collected during the winters of 1966-67 and
1967-68 and each twig cut into three sections based on years of
growth as shown in Fig. 1. A year's growth was defined as the dis-
tance between two sets of terminal bud scars. Surface areas for the
three terminal sections were estimated from 25 arbitrarily selected
twig sections from each of the samples. The data were analyzed with
an analysis of variance (Ostle, 1963) to check for significant vari-
ation in the number of mites among the sections.

A study was attempted to determine the winter mortality rates of
T. mcgregori. The orchard (102 trees) was divided into five nearly
equal size blocks, and five trees randomly selected within each block.
One hundred twigs were taken from each tree once each month for
five months (December- April, 1967-68). The data were than anal-
yzed with an analysis of variance (Ostle, 1963) to detect possible
variations among the months.

Results

Nine species of physoseiids were found overwintering in central
Utah apple orchards. Amblyseius cucumeris Oudemans, T. mcgre-
gori, and T. occidentalis were the most numerous and most frequently
collected, while Typhlodromus caudiglans Schuster, Typhlodromus
columbiensis Chant, A. fallacis, Amblyseius floridanus (Muma),
Amblyseius ovatus (Garman), and Amblyseius rosellus Chant were
collected incidentally. A frequency table was developed to indicate
overwintering site preferences for the three major species among
the four habitats most frequently inhabited (Table 1). Typhlodro-
mus mcgregori was present in all twig-spur and bark samples al-
though in much smaller numbers on the bark. On the other hand,
T. occidentalis was most common in bark samples and A. cucumeris
most frequently collected from cover samples. All of the incidentally
collected species were taken from cover, except that T. caudiglans was
taken from twig-spurs and T. columbiensis from bark. Although T.
mcgregori was about the only species present in derelict pear, apri-
cot, peach, plum, cherry, and apple orchards, they were present in

Table 1. Frequency of occurrence for the three most common
phytoseiids among the habitats they most frequently inhabited.
Species

Frequency^
Twig-Spur Bark Cover Soil

T. mcgregori 100.0 100.0 66.7 28.6

T. occidentalis 18.2 40.0 22.2 0.0

A. cucumeris 0.0 20.0 88.9 71.4

''Frequency was computed as a percentage of all samples taken for
each habitat.

Jul V 3 1 . 1 969 OVERWINTERING PHYTOSEIID MITES

99

Section 1

One year's
growth

Section 2

Section 3

Section ^

two years, section 2 — area o o ved.s
section 4 — area 9-15 years old.

The Great Basin Naturalist
100 LEETHAM AND JORGENSEN Vol. XXIX. No. 2

almost all of them and were much more numerous in apple orchards
than any of the other tree fruits.

The numbers of mites collected from green and falling leaves
were contrasted in the fall of 1967. Typhlodromus mcgregori num-
bers on the falling leaves were not significantly smaller than those
on the green leaves still adhering to the tree {t ■=■ 1.40. ^(21) (.95)
= 2.08). In this analysis the green leaves for October and Novem-
ber were pooled. A similar analysis for T. occidentalis resulted in the
same conclusions, the numbers on the falling leaves were not signi-
ficantly smaller than those on the green leaves still attached to the
tree (t = 1.40, ^(13) (.95) = 2.16). Typhlodromus occidentalis mig-
rated from the leaves to their overwintering sites at about the same
time Tetranychus urticae Koch migrated to the trunk and cover in
1968, and since T. urticae usually migrates earlier than the data were
gathered for T. occidentalis in 1967, the few predators remaining on
the leaves may have been merely incidental and not indicative of the
behavior of most members in the population.

The few T. mcgregori observed on the twigs and spurs appeared
to prefer sites beneath the corky abscission layers of the fruit stalks.
The area of the twig that was 3 to 5 years old maintained the rough-
est portions of twigs, primarily because of the remaining corky ab-
scision scars of former leaf petioles and fruit stalks. The twigs lost
most of their corky material and became smoother after five years,
reducing the available overwintering sites. The terminal two years
of growth characteristically displayed dense pubescence, but few good
hibernating sites. Typhlodromus mcgregori moved from the leaves
into the abundant overwintering sites on the 3 to 5 year old portions
of the twigs in the fall.

Twigs were sampled during the 1967-68 winter to test the hypo-
thesis that T. mcgregori preferred the region from 3 to 5 years old for
hibernating (Table 2). An analysis of variance was used to deter-
mine if there were significant differences among the three sections
tabulated in Table 2 (Table 3). Since the variance was significant,
the section means were tested (?-test) to find which sections had
significantly higher nmnbers. The only ones which were determined
to be significantly different were sections 1 and 3 (? = 5.04, ^(8)
(.95) = 2.31). It appears that the collection on February 3 in sec-
tion 1 had a deciding influence on the significance tests.

Data for winter mortality rates of T. mcgregori were pooled each
month for all five blocks, since the numbers of mites were too small
to analyze the trees separately (Table 4). An analysis of variance
suggested that there was no significant winter mortality within the
orchard ( oc = .05). If there were significant mortality of the suan,-
mer populations during the winter, it would have been when they
were finding their hibernacula or in the early spring after they had
left there sites. When the mortality reported by Anderson and Mor-
gan (1958). Chant (1959) and Putman (1959) occurred is yet to be
determined.

July 31. 1969 ovKinviNTEiuNG phytoseiid mites

101

Table 2. Number of Typhlodromus nicirrcgori per cm- in each
twig section from a derelict orchard in Majjlelon. Utah.

Mites per

Sample

Section and

(sq cm)

Mites per

Date

Mites per Section

Surface Area

1800 sq cm"

Section 1

March 15, 1967

1800.30

1

1.00

February 3, 1968

1493.12

34

40.97

March 2. 1968

1452.84

4

4.94

March 23. 1968

1649.30

11

11.96

March 29, 1968

1776.74

9

9.09

Mean

13.59

Section 2

March 15, 1967

2689.82

26

17.45

February 3. 1968

2648.72

39

26.53

March 2, 1968

2669.98

24

16.22

March 23, 1968

2287.74

29

22.83

March 29. 1968

2105.94

13

11.11

Mean

18.83

Section 3

March 15. 1967

2860.44

5

2.34

February 3, 1968

2610.64

9

4.48

March 2, 1968

3754.24

10

4.78

March 23, 1968

3273.26

13

7.14

March 29, 1968

2373.26

7

5.11

Mean

4.77

''The number of mites in each sample is determined as the number
of mites per 1800 cm- of twig surface area.

Table 3. Summary of the analysis of variance of the data from
Table 2, based on mites per 1800 cm- of twig surface area.

MH

° s

<H-( C«

OJ o

^^ 5

o gJ

c

b '^

Source of

II

3 cr

2 ™
cr .

Variation

&^

C/DOC

CD fe

Mean

1

173.80

173.80

Twig sections

2

10.32

5.16

3.68*

Experimental error

12

16.84

1.40

Total

15

200.96

*Significant at cc = .05.

The Great Basin Naturalist
102 LEETHAM AND JORGENSEN Vol. XXIX, No. 2

Table 4. Numbers of Typhlodromus mcgregori collected from
twigs and spurs of a derelict apple orchard near American Fork,
Utah.

Mites per 500 Twigs
in each Block

Month

1

2

3

4

5

Total

December

8

11

4

1

12

36

January

7

22

7

2

7

45

February

8

9

5

1

4

27

March

14

5

3

4

1

27

April

13

15

11

0

8

47

Discussion and Conclusions

Mites of the genus Typhlodromus have been observed overwin-
tering as adult females in the northern temperate fruit orchards,
although there is some question of the location of their overwintering
sites. Different species seem to prefer different habitats: T. mcgre-
gori on the twigs and bark, T. occidentalis on bark, and A. cucumeris
in the cover (Table 1). These results supported the observations of
Anderson and Morgan (1958), Putman (1959), and Burrell and
McCormick (1964), who found that Typhlodromus species were
generally arboreal in winter and summer while Amblyseius re-
mained primarily in the ground cover. The only arboreal species
found in high numbers in the cover was T. mcgregori and it has been
found rather commonly in the cover during the summer also (Jor-
gensen, unpublished data). Amblyseius cucumeris was occasionally
collected from the bark, but it was only incidental and probably not
in its preferred overwintering site.

Chant (1959) found that nearly all T. finlandicus, T. tiliae, and
T. umbricatus moved to the twigs and branches before the leaves
fell in the autumn months. He reported an increase in the number
of mites on green leaves as other leaves began to turn yelloW'' but
mites were not found on yellow falling leaves or on the ground cover.
The number of T. mcgregori on green leaves in central Utah did not
increase as other leaves turned yellow, but there were many collected
from falling leaves and in the ground cover. It appears that substant-
ial numbers of T. mcgregori fell to the ground cover with dropping
leaves, although it is likely that the data from the yellowed falling
leaves would have been misrepresentative if the major mite move-
ment to twigs and bark had occured before the sampling began.

Typhlodromus occidentalis moved from the leaves to the bark at
about the same time as T. urticae. The lack of change in the munbers
of predators on the leaves during October and November is likely
due to the fact that the major movement had already occurred and

July 31. 1969 overwintering phytoseiid mites lOS

only incidental specimens remained — the fate of \vhi( li is probaljly
not indicative of the population as a whole.

Putman (1959) suggested that arboreal phytoseiids are guided in
seeking overwintering sites by two responses — thigmotactic and
chemotactic (seeking objects of animal origin). The thigmotactic
response was suggested when he found hibernating mites primarily
in protected crevices; and the chemotactic when large numbers were
found in strips of cardboard containing oriental fruit moth (Grapho-
litha molesta Busck) cocoons placed on the trees, as oj)posed to few-
mites in similar strips without the cocoons. The twig dissections in
this study confirm Putman's (1959) findings since overwintering 7.
mcgregori were found primarily in protected crevices and spent
insect egg cases. The thigmotactic response in locating overwintering
sites was supported by the predominance of mites on portions of the
twigs from 3 to 5 years old where crevices are more numerous.
Thigmotaxis seems to be more influential than chemotaxis in the
searching behavior for overwintering sites by T. mcgregori.

All preceeding studies on winter mortality of phytoseiids have
reported rather high mortality rates. Chant (1959) and Anderson
and Morgan (1958) both reported mortality rates over 90%; but
since both studies were based on comparisons of fall and spring mite
populations on green leaves, an error could have been introduced if
a large number of mites fall to the ground with the autumn leaf drop
and do not move back onto the tree in the spring. Chant (1959) re-
ported that almost all of the mites left the leaves before they began
to fall, so the reported high mortality was likely, but a more pre-
cise measurement is possible by periodically sampling the overwinter-
ing population. Typhlodromus mcgregori did not appear to suffer any
winter mortality between December and April, thus whatever mort-
ality occurred must have been before December or after April, and
these two periods may result in mortality as severe as those reported
by Chant (1959) and Anderson and Morgan (1958).

If predaceous mites are to be considered for use in integrated
control of phytophagous mites, their overwintering sites and mortal-
ity rates must be known. Their usefulness is determined, in part,
if high spring populations can be provided to counteract an early
spring build-up of the pest species. By knowing the principal over-
wintering sites a grower could manage his trees to provide abundant
sites for the predators and direct j)esticide treatments to avoid them.
Gilliatt (1935) and Chant (1959) reported that dormant sprays
reduced phytoseiid })opulations in addtion to the eggs of phytopha-
gous species. Since most of the T. mcgregori overwinter on the twigs,
they are extremely vulnerable to dormant sprays. On the other hand.
T. occidentalism which overwinter [)rimarily in bark of the trunk and
scaffold limbs, is less vulnerable since the deeper bark crevices ])ro-
vide better protection from pesticides. This is supjwrted by the fact
that T. mcgregori is seldom found in commercial orchards of cen-
tral Utah (Duke. Croft, and Jorgensen; 1968) where T. occidental is

The Great Basin Naturalist
104 LEETHAM AND JORGENSEN Vol. XXIX. No. 2

is frequently abundant. Based on these observations, it appears that
T. mcgregori would be difficult to manage in an integrated control
program since it is vulnerable to dormant sprays, which are usually
an integral part of an integrated control program in tree fruit or-
chards. The overwintering habits of T. occidentalis make it a more
promising predator in integrated orchards.

Literature Cited

Anderson. N. H. and C. V. G. Morgan. 1958. The role of Typhlodromus spp.
(Acarina: Phytoseiidae) in British Columbia apple orchards. Proc. Internatl.
Cong. Entomol. 10:659-663.

BuRRELL, R. W. AND W. J. McCoRMiCK. 1964. TyphlodroTTius and Amblyseius
(Acarina: Phytoseiidae) as predatore on orchard mites. Ann. Entomol. Soc.
Amer. 57:483-487.

Chant, D. A. 1959. Phytoseiid mites (Acarina: Phytoseiidae) Part I. Biono-
mics of seven species in southeastern England. Part II. A taxonomic review
of the family Phytoseiidae. with descriptions of 38 new species. Can. En-
tomol. 91, Suppl. 12. 64 pp.

. 1965. Genetic concepts in the family Phytoseiidae (Acarina: Mesostig-

mata). Can. Entomol. 97:351-374.

Duke. D. M.. B. A. Croft, and C. D. Jorgensen. 1969. Distribution and
searching behavior of Typhlodromus mcgregori Chant (Acarina: Phytoseii-
dae). Unpublished manuscript.

Garman, p. and S. F. Townsend. 1938. The European red mite and its con-
trol. Bull. Conn. Agric. Expt. Sta. 418. 34 pp.

GiLLiATT, F. C. 1935. Some predators of the European red mite, Paratetrany-
chus pilosus C. & F.. in Nova Scotia. Can. J. Res. 13: 19-38.

Herbert, H. J. 1953. Progress report on predaceous mite investigations in Nova
Scotia (Acarina: Phytoseiidae). Rep. Entomol. Soc. Ontario. 1952:27-29.

Jorgensen, C. D. 1967. Ecology of Utah apple mites: A statement of the prob-
lem. Proc. Utah Acad, of Sci., Arts, and Letters. 44:265-274.

Lee, M. S. and D. W. Davis. 1968. Life history and behavior of the predatorj'
mite Typhlodromus occidentalis in LItah. Ann. Entomol. Soc. Amer. 61:
251-155.

Ostle, B. 1963. Statistics in research. 2nd Ed. Iowa State LTniv. Press. Ames.
585 pp.

Putman, W. L. 1959. Hibernation sites of Phytoseiids (Acarina: Phytoseiidae)
in Ontario peach orchards. Can. Entomol. 91:735-741.

SPIDERS OF THE
NATIONAL REACTOR TESTING STATION"

Dorald M. AUred

Ecological investigations conducted by Rrigham Young Univer-
sity from June 1966 to September 1967 at the National Reactor
Testing Station in southeastern Idaho (Figs. 1. 2) were directed
specifically toward the ectoparasites of vertebrates (Allred. 1968).
Secondarily, however, some ground-dwelling, free-living arthropods
were collected in can pit-traps used for the capture of rej)tiles and
rodents. Considering the relatively little effort expended, an un-
usual variety of spiders was collected. These were kindly identified
by Dr. Willis J. Gertsch, American Museum of Natural History.
Their listing here should provide a basis for further studies of these
important arthropods in the Great Basin and other desert areas
of the western United States.

BLACKFOOT

Fig. 1. Location of the National Reactor Testing Station in relationship to some
cities in southeastern Idaho.

'BYU - AEG publication no. C00-135')i luider sponsorship of the .Monilc Knergy Commission.

105

106

D. M. ALLRED

The Great Basin Naturalist

Vol. XXIX, No. 2

LEGEND

PAVED flOADS

UNSURFACED ROADS

— ^— NRTS BOUNDARY

a NHTS INSTALLATIONS
•-Q BYU STUDY AREAS

Fig. 2. Principal study areas at the National Reactor Testing Station.

Annotated List of Species

Antrodiaetus montanus: m*; area 6; Oct.

Calilena restricta: 15m 71 5im; areas 1-12 (except 4, 5, 9), 18;

June-Oct. (mostly Aug.).
Callilepis eremellus. m f; area 1,19; June, July.
Castianeira descripta: f; area 3; Sept.
Ceratinella acerea: f; area 10; May.
Circurina grandis: Im; area 21.
C. par ma: 13m 2f; areas 6, 10, 11, 12, 23; April, Oct. (mostly),

Nov.
C. utahana: m; area 11; Nov.

July 31, 1969 spiders of n.r.t.s. 107

Circurina new species: 6m; areas 6, 10, 21 ; Oct., Nov.

Dictyna coloradensis: 2m 2f; area 10; July.

Drassodes robinsoni: 4m 2f Sim; areas 1, 2. 6, 7, 10; June. Aug..

Oct., Nov.
Drassyllus mannellus: 7m 3f lim; areas 1, 3, 6, 10. 18; May-July,

Sept.
Ebo mexicanus: 5f; areas 1,6, 7; June, July. Nov.
Enophognatha wyuta: f; area 10; June.
Euryopis scriptipes: m; area 2; July.

Gnaphosa brumalis: 9m lOf 2im; areas 3, 6, 8; June- Aug.
Gnaphosa sp.: 2m If 8im; areas 1, 2, 7, 8, 10, 11; April-Aug.
Haplodrassus eunus: 15m 14f llim; areas 1. 6, 7, 10. 11. 12, 21;

April, May. Oct., Nov.
Herpyllus sp.: im; area 8; Aug.
Latrodectus Hesperus: 13im; areas 8, 12; July-Sept.
Metaphidippus sp.: im; area 6; July.
Micaria foxi: m f; areas 1,3; June, Aug.
M. gosuita: f; area 12; Sept.

Micaria new species A: 3m 4f; areas 3, 12; June (mostly), Aug.
Micaria new species B: 2m areas 2, 12; June, Aug.
Misumenops sp.: im; area 10; July.
Pardosa new species: 2f; area 8; July.
Pellenes washonus: m; area 12; July.
Phidippus altanus: m; area 3; Aug.
P. formosus: m; lim; areas 3, 7; Aug.
Philodromus alascensis: lim; area 23.
Poecilochroa atomistica: f im; areas 2, 3; July, Aug.
Schizocosa avida: 46m 34f 154 im; areas 1, 2, 3. 6, 7, 8, 10, 11, 12,

18, 19, 21, 23; males were taken from April to July (mostly in

July), females from April to Aug., and immatures from April

to Nov. (mostly Aug., Oct., Nov.).
Steatoda albomaculata: f; area 3; Sept.
Tarentula kochi: 37m 19f 13im; areas 1, 3, 6, 7, 10, 11, 12, 23;

June, Aug.-Nov. (mostly Oct.).
Thanatus altimontis: 9m If; areas 1, 2, 3, 7, 12; June- Aug.
Xysticus coloradensis: 10m lim; areas 2, 5, 6, 7, 11, 12, 18; June,

July, Sept.-Nov.
X. knowltoni: 7m 5f lim; areas 1, 3. 10, 12; May-July, Nov.
X. montanensis: 3m 3f; areas 1, 3, 7, 10; April- June.
X. nigromaculatus: 2m; areas 7, 10; June.
X. qulosus: m; area 6; Oct.

Zelotes pullatus: 11m 8f 2im; areas 1. 2, 3, 7, 10, 11, 12; April-Oct.
Z. puritanus: 4f; areas 2, 3, 19; July, Aug.

•ni = adult male, f = adult female, im = immature or juvenile.

Summary

From June 1966 to September 1967 approximately 500 spiders
representing 42 species in 31 genera were collected in. can pit-traps
at the National Reactor Testing Station in Idaho. Four of the species

The Great Basin Naturalist
108 D. M. ALLRED Vol. XXIX, No. 2

are new to science. The greatest variety of species was found in
study area 10 where the vegetation analysis was 68% Artemisia,
7% Opuntia, 7% mixed grasses, and 5% miscellaneous forbs. The
fewest numbers of species were found in study areas 7 and 11,
typified predominantly by Chrysothamnus plants. Males of all
species of spiders combined were collected from April to November.
Greatest numbers were found in October. Females of all species
were taken from April to November, with slightly more in October
than in other months. Immature spiders also were found from April
to November, with greatest numbers occurring in August. October,
and November. The species that were the most widespread over the
station are Schizocosa avida and Calilena restricta. The most
abundant species was S. avida; Tarentula kochi was only half as
abundant as 5. avida, but twice as common as any other species.

Referenced

Allred, D. M. 1968. Ticks of the National Reactor Testing Station. Brigham
Young Univ. Sci. Bull., Biol. Ser. (in press).

SCIENTIFIC NOTES

INDIVIDUAL RODENT MOVEMENTS IN UPPER
SONORAN DESERT OF CENTRAL NEVADA

Most studies of distiiiues traveled In animals suffer from the impossibility
of being certain that an animal cannot travel farther than one imagines. Even
investigations on islands or in other physically limited sites are apt to be biased
by tlie limits themselves, and thus can be extended to larger areas only at peril
of underestimation. For this reason, it seems imprudent — and probably delusive —
to calculate "average" distances unless one can be sure that the greatest distances
moved b}' animals v\ill in fact be recoi'ded, or at least recognizes the inadequacies
accepted. The ma.ximum distances known to have been moved, however, are use-
ful in judging the adecjuacy of experimental design in investigating the signifi-
cance and actual validity of such ideas as home range, and in studies of habitat
utilization.

During a study of habitat selection by kangaroo mice {M icrodipodops megace-
phalus and M. pallidus) conducted in central Nevada in 1960 and 1961. all ro-
dents caught at surveyed points were marked by toe-clipping and released at the
traps, in an effort intended to reveal interactions of kangaroo mice with other
species.

Trapping was carried on at three places where Hall (Mammals of Nevada,
pp. 379-403,, 1946) had found the two species of kangaroo mice to be sympat-
ric. The dominant vegetation at all three may be referred to the Little Grease-
wood-Shadscalc (Sarcobatus baileyi-Atriplex confertifolia) association of Billings
(Butler Univ. Bot. Stud., 7: 89-123, 1945). At each area, small Sherman traps
baited with rolled oats were set within 0.5 yard (0.5 m) of positions 10.00 yards
(9.14 m) apart in grid patterns. The grids, their accuracy controlled by stakes
at the 20.00-yard (18.28-m) intersects, were extended as accumulating data
on kangaroo mice dictated. The areas, and the greatest distances between trap
positions on the grids there, were (i) Big Smoky Valley, 5,650 ft, 4.75 miles NE
San Antonio. Nye Co., 439 yards (401 m); (ii) Granite Springs Valley. 4,000
ft. 21 miles W and 2 miles N Lovelock, Pershing Co.. 241 yards (220 m); and
(iii) Penoyer Valley, 4,800 ft, 15 miles N Groom Baldy, Lincoln Co., 632 yards
(579 m). For each individual, the points of capture were expressed in rectilinear
coordinates (Ghiselin, Amer. Midland Natur., 79: 242-246, 1968). For each, the
distances between all sites of capture were calculated, and the greatest distance
for each individual was selected by a computer routine utilizing the Pythagorean
theorem. The results are set forth in Table 1. To indicate comparative sample
sizes, numbers of captures and of individuals involved are included.

The field work was aided hy grants from the Wisconsin Alunmi Research
Foundation and the National Science Foundation (Cj 14908. administered by W.
G. R€>eder). I am grateful to Dr. Reeder for his counsel. — Jon Ghiselin. Dept.
of Zoology, University of Wisconsin, Madison 53706. (present address: Biology
Dept.. Lycoming College. Williamsporl. Pa. 17701).

109

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SCIENTIFIC NOTES

AN ALBINO PACIFIC TREE FROG, HYLA REGILLA,
FROM DEATH VALLEY, CALIFORNIA

On 25 March. 1967, a partially albino 9 H. regilla (snout-vent, 35.0 mm)
was collected at Saratoga Springs, Death Valley National Monument, San Ber-
nardino Co.. California. Except for color, the frog appeared normal. Her behavior
at capture was normal and her ovaries contained well developed, pigmented
eggs. Four lung-worms (Rhabdias ranae Walton) were found, but there was no
evidence of pathological damage. No other abnormally colored frogs have been
found in over a year of collecting from this population. The specimen (A-540)
is on deposit in the Biology Museum, University of Nevada, Las Vegas.

The frog's dorsal coloration is broken by extensive non-pigmented areas,
but her eyes appear to be of normal color. The pigmented skin appears to be
lighter than normal for the population. Four melanophore counts taken on a
5x3 mm section of pigmented skin from near the front leg of the albino and
a normal $ H. regilla bear this out X normal = 4375/mm-, X albino =
3525 /mm-. P<;.001). The normal pattern, although interrupted, is evident on
the pigmented areas of the albino.

To my knowledge, only one other case of albinism has been reported for
Hyla regilla. Jameson and Myers (Herpetologica, 13:74, 1949) reported an albino
H. regilla from Oregon that developad from non-pigmented eggs. Hensley (Publ.
Mus. Michigan State Univ. Biol. Ser., 1: 135-59, 1959) reiterated the Oregon
record, but did not report any other instances. I have found no other reports
of albinism in H. regilla since 1959. However, Wright and Wright (Handbook
of frogs and toads of the U.S. and Canada, Comstock Publ .Co., New York.
1949) reported collecting a Hyla regilla in Las Vegas, Nevada, in 1925, that
was ". . . very yellow with very indistinct pattern." [sic]. Although the authors
made no comment, the frog may have bean an albino. In several hundred tree
frogs from the Las Vegas area, I have not seen a yellow one. Greenish-gray is
a common color, but the pattern is always distinct.

I wish to thank Dr. James E. Deacon for reviewing the manuscript and Dr.
Bert B. Babero for identification of the parasites. This study has been supported
in part by U. S. National Park Service contract no. 14-10-0434-0989 to Dr.
James E. Deacon. Fenton R. Kay, Department of Biological Sciences,

University of Nevada, Las Vegas, 89109.

Ill

'6-G-i2>^t^

The

Volume XXIX, No. 3
September 30, 1969

MUS. CO MP. ZOCL.
LIBRARY

MAR 11971

HARVARD
UNIVERSITY

Great Basin

nmmim

Published by
Brigham Young University

GREAT BASIN NATURALIST

Editor: Vasco M. Tanner, Department of Zoology and Entomology
Brigham Young University, Provo, Utah

Associate Editor: Stephen L. Wood, Department of Zoology and
Entomology, Brigham Young University, Provo, Utah

Members of the Editorial Board:

Perron L. Andersen (5), Zoology

Jay V. Beck (3) , Bacteriology

Robert W. Gardner ( 1 ) , Animal Science

Joseph R. Murdock (4) , Botany

WiLMER W. Tanner (2), Zoology, Chairman of the Board

Stanley L. Welsh (1), Botany

Ex officio Members:

A. Lester Allen, Dean, College of Biological and Agri-
cultural Sciences

Ernest L. Olson, Chairman, University Pubhcations, Uni-
versity Editor

The Great Basin Naturalist

Published at Phovo, Utah by
Brigham Young University

Volume XXIX September 30, 1969 No. 3

NEW SYNONYMY AND RECORDS OF PLATYPODIDAE
AND SCOLYTIDAE (COLEOPTERA)'

Stephen L. Wood-

During the past year the opportunity came for me to study the
types of bark and ambrosia beetles (Platypodidae and Scolytidae) in
three old, significant collections. The oldest of these is the collection
of Count C. (t. Mannerheim, presently housed in the Universitetets
Zoologiska Museum, Helsinki, Finland. It is noteworthy because it
contains the types of species taken in Russian America (mostly
Alaska) at an early date when few American species were known.
Most of the 17 species of concern here were previously correctly
known to specialists; noteable exceptions involve Bostrichus (now
Ips) tridens, Hulurgus (now Pseudohylesinus) sericeus, and a new
status for H. (now Dendroctonus) obesus.

The second collection considered here, in terms of age priority,
is that of Victor von Motschulsky presently housed at the Zoological
Institute of the USSR. Moscow. The 26 species of concern here
were described largely from Ceylon and. for the most part, have
been known to specialists only from the original descriptions and a
very few duplicate specimens evidently distributed by Motschulsky.
Previous workers were not successful in locating the types and have
either ignored the names or have considered the types lost or de-
stroyed. The policy adopted here in this article apparently is con-
sistent with the action of previous workers; that is, specific names
of these species were given their normal place in priority, but three
generic names, Anodius (1860) and Phloeotrogus (1863) (=Xyle-
borus Eichhoff, 1864), and Olonthogaster (1886) (=PhIoeosinus
Chapuis, 1869) clearly predate the priority of names now used for
the genera to which they apply. These names. Xyleborus and
Phloeosinus. have been applied consistently for a century to large
genera of almost world-wide distribution and it is not in the interest
of nomenclatorical stability that they be changed. .Mthough Anodius.
Phloeotrogus, and Olonthogaster have been hsted in catalogs in an

'The travel grant that made this report possible was provided by the National Sricnrc Founiln-
tion through the Entomological Society of America.

-Department of Zoology and Entomology, Brigham Young University. Provo. Utah; Scolytoidea
contribution No. H).

113

The Great Basin Naturalist
114 STEPHEN L. WOOD Vol. XXIX, No. 3

incerta sedis category, I feel they qualify as nomena oblita. The
case v^dth the monobasic Genyocerus (1858) is different. Strohmey-
er evidently deliberately named a homonym in the genus Diapus to
replace G. alhipennis, the type-species of Genyocerus. It apparently
has been clearly recognized that albipennis belongs to the species
group currently assigned to Diacavus Schedl. 1939. Contrary to
Schedl's (1959:515) statement, the type of albipennis is available
and the characters of Genyocerus are clearly definable as his place-
ment of albipennis in Diacavus indicates. Diacavus was established
recently and it involves a limited number of species (about 15) from
a restricted region (oriental). For these reasons I place Diacavus
Schedl in synonymy under the much older name Genyocerus Mot-
schulsky {New synonymy).

The third collection treated here is part of the Felicien Chapuis
material on which his 1869 (and 1873) Synopsis des Scoly tides was
based. This part of his material is currently housed in the Institut
Royal des Sciences Naturelles de Belgique. Brussells. Because a
more extensive treatment of the American material contained in
this collection is in preparation, only four items of immediate con-
cern are treated here.

My sincere appreciation is extended to the above named institu-
tions for their kindness in permitting me to examine their material;
and to Dr. Walter Hackman and Mr. Hans Silfverberg. at Helsinki.
Dr. A. N. Zhelochovtsev, at Moscow, and Dr. Roger Damoiseau, at
Brussells, for their kindness and cooperation.

In order to resolve some of the problems presented here and to
facilitate other work, six American species are described below as
new to science. These species are in the genera Pseudohyle sinus ( 1 ) ,
Phloeotribus (3) , and C hramesus (2).

Mannerheim Collection

Of the 17 species of Scolytidae named from Alaska by Manner-
heim, subsequent workers have correctly placed most of them. A
few have been misidentified. My comments on the collection follow.

Bostrichus affaber Mannerheim (1852)

This species has correctly been referred to Dryocoetes. The type
is a female.

Bostrichus cavifrons Mannerheim (1843)

Three specimens taken at Sitka by Holmberg are in the Manner-
heim drawer wdth the types, but none has been designated as the
type. In another drawer containing general Mannerheim Scolytidae
there are eight specimens designated "Spec, typ.," collected by D.
Eschscholtz or Blaschke. Evidently the type, collected by Eschscholtz,
is among the latter eight specimens. All are of Trupodendron linea-
tum (Olivier) with which this species has correctly been placed in
synonymy.

Sept. 30. 1969 new synonymy and rkcords 115

Bostrichus concinnus Mannerheim (1852)

This species has correctly been referred to the genus Ips. The
type is a female.

Bostrichus inter ruptus Mannerheim (1852)

This species has correctly been referred to the genus Ips. The
type is a female.

Bostrichus nitidulus Mannerheim (1843)

This species has correctly been referred to the genus Pityoph-
thorus. The type is a female.

Bostrichus semicastaneus Mannerheim (1852)

This species has correctly been made a junior synonym of Dry-
ocoetes autographus (Ratzeburg). The sex of the type was not
determined.

Bostrichus septentrionis Mannerheim (1843)

This species has been made a junior synonym of Dryocoetes auto-
graphus (Ratzeburg). A specimen bearing this name was not found
in the Mannerheim collection.

Bostrichus terminalis Mannerheim (1843)
This species belongs in the family Cisidae.

Bostrichus tridens Mannerheim (1852)

This species has correctly been referred to the genus Ips. The
type is a female exactly identical to the female type of Ips inter-
ruptus (Mannerheim) {new synonymy). The name interruptus
appears first on the page, but I exercise the right as first revisor in
this instance and select the name tridens for this species. The mor-
phological form having a greatly enlarged female frons, generally
known as Ips tridens, does not require a new name; it will be treated
in the near future in a detailed study by Dr. Gerald Lanier.

Cryphalus striatulus Mannerheim (1853)

A specimen bearing this name could not be found in the Manner-
heim collection. It evidently is the species currently known as
Trypophloeus nitidus Swaine.

Hylastes cristatus Mannerheim (1853)

A specimen bearing this name could not be found in the Manner-
heim collection. It has been treated as a junior synonym of Hylur-
gops subcostulatus (Mannerheim).

The Great Basin Naturalist
116 STEPHEN L. WOOD Vol. XXIX, No. 3

Hylastes subcostulatus Mannerheim (1853)

This species has correctly been transferred to the genus Hylur-
gops. There were no specimens under this name in the Mannerheim
material; however, it is easily recognized from the original descrip-
tion.

Hylurgus nigrinus Mannerheim (1852)

This species has correctly been referred to the genus Hylastes.
The type is a female.

Hylurgus obesus Mannerheim (1843)

This species has correctly been referred to Dendroctonus. While
my (Wood, 1963) revision of Dendroctonus was in press a delayed
comparative note on the type of D. rufipennis (Kirby) was received
indicating that obesus is a junior synonym. In my absence the cor-
rection was properly placed in the synonomy but the senior name
was not substituted elsewhere in the manuscript. Since then I have
re-examined the types of both obesus and rufipennis. There are
obvious synonyms, with the name rufipennis having priority {new
status).

Hylurgus pumilus Mannerheim (1843)

This species has correctly been referred to the genus Dolurgus.
The sex of the type was not determined.

Hylurgus rugipennis Mannerheim (1843)

This species has correctly been referred to the genus Hylurgops.
The specimen labeled "Type" in the Mannerheim type collection
was not collected by Eschscholtz and, therefore, probably is not
actually the type. Seven other specimens of this species in the
Mannerheim material are from Sitka, but were also taken by other
collectors.

Hylurgus sericeus Mannerheim (1843)

Under this name in the Mannerheim collection are two speci-
mens. The second specimen belongs to the genus Leperisinus and
is from Louisiana. The first is from Sitka, collected by Blaschke, and
is considered to be the type. This specimen is a female of the species
generally known as Pseudohylesinus grandis Swaine {new synony-
my) ; Swaine's name, therefore, must be placed in synonomy. Pseu-
dohylesinus sericeus of Swaine and Blackman is an unnamed species
(see below).

Polygraphus saginatus Mannerheim (1853)

This species has correctly been placed as a junior synonym of
P. rufipennis (Kirby). The only specimen of this species in the
Mannerheim material is a female without locality data, bearing the
generic and specific names, but no author or other identifying in-
formation. It could be the type, but it is not so designated.

Sept. 30, 1969 new synonymy and records 117

MOTSCHULSKY COLLECTION

The types of 26 species of Scolytidae and Platypodidae are in
the Motschulsky collection. Synonymy for nine of the species has
been suggested by previous workers, based on an examination of the
original descriptions or of a very few duplicate specimens ])resuma-
bly distributed by their author. The taxonomic j)lacement of the
1 7 remaining species has not been clear. It was noted that labels
were mostly written in (jemian and did not always corresj)ond with
the published type locality designated by Motschulsky. However,
after examining the entire Motschulsky collection presently avail-
able, I see no reason whatever to question the types as they are
presently designated. Five of them have been carefully remounted
on the same pin above the original microcard, and all bear a red
type label in addition to the original designation. My comments on
the alphabetically arranged Motschulsky species follow.

Anodius denticulatus Motschulsky (1860)

Six female specimens are mounted individually on six micro-
cards on one pin. The upper five are Xyleborus perforans Wol-
laston; the sixth specimen, placed in the lowest position on the pin.
is Xyleborus similis Ferrari. I here designate the uppermost speci-
men as the lectotype of denticulatus Motsch. {new synonymy).

Anodius distinctus Motschulsky (1866)

One female specimen. Comparative material was not at hand
for varification, but this species is Coccotrypes floridensis Schedl
which is the species I have recognized as C. carpophagus Hornung.
Schedl refers Hornung's name to a different species. C. pygmaeus
Eichhoff; I have not yet verified his synonymy. Regardless of how
the later problem is resolved Coccotrypes distinctus (Motsch.) is a
senior synonym of C. floridensis Schedl {new synonymy).

Anodius piceus Motschulsky (1863)

One female specimen. Fhis species is Xyleborus indicus Eichhoff.
The apices of the elytra are immersed in glue. The pronotuni is
subquadrate, the color is dark, the basal half of the elytral declivity
may be very slightly more convex than in many indicus, but it is
within the range of variation for this species. Xyleborus piceus
(Motsch.) has priority and must replace X. indicus Eichhoff {new
synonymy) .

Anodius tuberculatus Motschulsky (1863)

Two female specimens. 1 hey were compared directly to authen-
tic specimens of Xyleborus perforans Wollaston; there is no question
as to the synonymy. Wollaston's name has priority {n^w
synonymy) .

The Great Basin Naturalist
118 STEPHEN L. WOOD Vol. XXIX, No. 3

Eccoptopterus sexspinosus Motschulsky (1863)

Three females labeled "India occidental, Birma." The length of
these speciments is about 2.3 mm., and the second and third pairs of
teeth on the elytral declivity are smaller, with the second pair closer
together than in my specimens of E. spinosus {Olivier). In spite of
these differences, I feel the previously suggested snyonymy is correct.

Genyocerus adustipennis Motschulsky (1859)

One female specimen, from Japan. An examination of the litera-
ture treating the Platypodidae of Japan strongly suggests that this is
Crossotarsus niponicus Blandford. Synonymy should await varifica-
tion of this observation.

Genyocerus albipennis Motschulsky (1858)

One female speciman labeled "Aus Indien or (word illegible)."
It is about 2.5 mm. in length; the frons is elaborately ornamented by
slight impressions on lateral areas that extend dorsad to the upper
level of the eye, with tufts of coarse, long hair occupying the lateral
thirds from the upper level of the eye to two-thirds of the distance
toward the mandibles and projecting forward a distance equal to the
length of the head then curling back toward the mouth and touching
the mandibles; additional tufts arise on the mouthparts and curl up-
ward to the middle of the frons; median one-third to one-half of
frons glabrous; base of pronotum with about a dozen coarse pores.
This is a Diacavus species reminiscent of quadriporus Schedl, but
distinct. However, as indicated above Diacavus must be placed in
synonymy under the much older Genyocerus.

Hylesinus granulifer Motschulsky (1863)

Two male specimens mounted on one pin. These specimens
agree in all details with my male of H. despectus Walker, from
Tonkin.

Hylesinus sericeus Motschulsky (1866)

One female specimen. It belongs to the genus Cryphalus, and
probably is C. indicus Eichhoff. The size, sculpture and setae fit
indicus, but specimens were not at hand for comparison.

Hypoborus cinerotestaceus Motschulsky (1866)

One female specimen in very poor condition. It is about 1.3 mm.
in length, with erect, interstrial rows of stout setae, the length of
each less than half as great as distance between rows. It is similar to
paganus Eichhoff or pallidus Eichhoff, but specimens were not avail-
able for comparison.

Sept. 30. 1969 new synonymy and records 119

Hypoborus dorsalis Motschulsky (1866)

Two females and one male mounted on one card on one pin.
They are of Cryphalus indicus Eichhoff, or very near. The size,
sculpture and setae are as in indicus, however, specimens were not
available for direct comparison.

Hypoborus nebulosus Motschulsky (1866)

Two specimens, probably males, with the frons hidden. Also of
Cryphalus, probably indicus. The size, sculpture and setae are as in
indicus, however, specimens were not available for direct comparison.

Olonthogaster nitidicollis Motschulsky (1866)

One female specimen. This is a large Phloeosinus similar to
asper (Sampson). Specimens were not available for comparison.

Olonthogaster nitidifrons Motschulsky (1866)

One male specimen. This is a Phloeosinus species presently un-
known to me. It is about 2.5 to 2.7 mm. in length; decli vital inter-
striae 1 and 3 each bear three or four widely separated, sharply
pointed teeth.

Phloeotrogus attenuatus Motschulsky (1863)

One female specimen. This specimen is Xyleborus sordicauda
(Motsch.) (new synonymy) and is known to specialists by that
name. It agrees completely with my homotype.

Phloeotrogus bidentatus Motschulsky (1863)

One female specimen labeled "India occidental." This specimen
agrees with the species generally known to specialists as Xyleborus
bidentatus (Motsch.) My specimen was compared directly to the
type.

Phloeotrogus crassiusculus Motschulsky (1866)

Three females labeled "India occidental," mounted on individual
microcards on one pin. The specimens are somewhat greasy, making
the transition from shiny to apaque areas on the elytra difficult to
see. My homotype (2.8 mm.) is very slightly larger than the types,
but it is clearly Xyleborus semiopacus Eichhoff {new synonymy) .
Motschulsky's name has priority.

Phloeotrogus obliquecauda Motschulsky (1863)

Four females labeled "India occidental." This species is allied to
Xyleborus minor (Stebbing) and is of the same size and proportion;
however, the declivital sculpture and armature are much more
nearly like X. tachyderus (Zimmermann). It is a species presently
unknown to me.

The Great Basin Naturalist
120 STEPHEN L. WOOD Vol. XXIX. No. 3

Phloeotrogus sordicauda Motschulsky (1863)

Two female specimens. These specimens were compared to my
specimen and the characters agree completely. It is the sj)ecies
known generally among specialists as Xyleborus sordicauda
(Motsch.).

Phloiotrihus subquadratus Motschulsky (1866)

One specimen. This specimen superficially resembles the genus
Phloeotribus, but it belongs to the family Anobiidae.

Platypus caudatus Motschulsky (1863)

Six male specimens labeled "India oriental," all badly covered by
fungal growth, unquestionably are Platypus solidus Walker as was
previously suggested.

Platypus cordatus Motschulsky (1863)

Four female specimens labeled "India oriental." These speci-
mens agree with my female homotype except that the major prono-
tal pores are very slightly smaller. It is a junior synonym of Platy-
pus solidus Walker (new synonymy) .

Platypus luniger Motschulsky (1863)

Three males and three females labeled "India oriental." Notes
and diagrams indicates this species is Platypus caliculus Chapuis
{new synonymy).

Platypus rotundicauda Motschulsky (1863)

Two female specimens labeled "India oriental." At the present
time I am not familiar with the Platypus species to which this name
should be associated.

Tomicus adusticollis Motschulsky (1863)

One female labeled "Ceylon." This specimen is about 2.2 mm.
in length; my notes and diagram indicate it is Xyleborus pumilus
Eggers. A specimen was not available for direct comparison {new
synonymy) .

Tomicus quadrispinus Motschulsky (1863)

One female labeled "India or Birma." This specimen is about
2.5 mm. in length; it is either Xyleborus amphicranulus Eggers, a
small specimens of X. emarginatus Eichhoff or very closely related.
More specimens of these species are required to more fully under-
stand their ranges of variation.

Sept. 30. 1969 new synonymy and records 121

Chapuis Collection

Only a few items relating to the Chapuis (1869) tyjies are men-
tioned here, the remainder will he treated in a later publication.

Hylastes vastans Chapuis

The female holotype is labeled "Mexico, Dy." It was compared
directly to my homotype of Hylastes gracilis LeConte and is of the
same species. LeConte's name has priority {neu^ synonymy) .

Dendroctonus parallelocollis Chapuis

The type evidently is a female; it is labeled "Mexico. Dy." My
concept of this species (1963) was based on specimens supposedly
compared to this type by Hopkins, Eggers, and Schedl. All clearly
were in error. The type is identical to D. aztecus Wood {new
synonymy). The species for which the name parallelocollis has
been used in all references except Chapuis (1869) must now take the
name of the only available iunior synonym, D. approximatus
Hopkins.

Phloeotribus obliquus Chapuis

The Chapuis series consists of four specimens, two labeled as
types, and two as "ex-typus" identified by Chapuis. The first syn-
type is a female of P. transversus Chapuis (types compared direct-
ly); the second syntype and both of the other specimens (ex-typus)
are of the one species. I here designate the second syntype to be the
lectotype of P. obliquus Cha{)uis. There is no indication on either
specimen or in subsequently studied material to suggest a Mexican
(or even a Central American) origin for specimens of this species.

Phloeotribus sulcifrons Chapuis

This species has been widely reported from Central America.
The female type, however, from Colombia, is quite different. The
female frons is narrowly, rather strongly sulcate from the vertex to
the position normally occupied by a frontal fovea in this genus.
Pronotal and elytral characters also indicate that P. sulcifrons of
Blandford and other authors, from Central America, requires a new
name (see below).

Supplemental Notes

Liparthrum Wollaston (1854)

The original spelling of this generic name was Leiparthruni; in
1864, Wollaston changed it to Liparthrum which was used consis-
tently from then until Bright (1968) declared Wollaston's (1864)
action an unjustified emendation, evidently without seriously study-
ing the matter, and reverted to tlie 1854 spelling. Article 33a of the
International Code says a justifiable emendation "is the correction

The Great Basin Naturalist
122 STEPHEN L. WOOD Vol. XXIX. No. 3

of an incorrect original spelling and the name thus emended takes
the date and authorship of the original spelling." This generic name
was based on the Greek root leipo which transliterates into the Latin
lipo. VVollaston, in 1864. clearly indicated by a footnote that his
emendation was intentional; he evidently assumed his reason for
the change was obvious. The spelling Liparthrum, therefore, is
correct and should be continued in the interest of nomenclaturial
stability.

Phloeotribus Latreille (1802/3)

This generic name was originally presented as Phloiotribus
(Latreille, 1796), a nomen nudum; it was validated in 1802/3 by
the same author. Later, Erichson (1836), recognizing the same type
of problem in transliterating the Greek root word phloios into the
Latin phloeos as treated above, emended the name to Phloeotribus.
With one or two minor exceptions (including that of Motschulsky,
used above) the spelling Phloeotribus has been used consistently
from 1836 until an apparent attempt to revive the original spelling
by Barr (1969), presumably influenced by Bright. Erichson's
emendation was justifiable under article 33a of the International
Code and should be preserved.

Chaetophloeus phoradendri Wood, emendation

In the recent validation of this name (Wood. 1969:8), the spe-
cific designation inadvertently was spelled phorodendri. It should
be corrected to phoradendri to correspond with the generic name of
its host.

Pseudohylesinus pini, n. sp.

This is the species previously knovsoi as Pseudohylesinus sericeus
of Swaine, Blackman. etc., not Mannerheim. As indicated above,
the type of Mannerheim' s species is of a different species than was
treated by Swaine and Blackman.

This species is distinguished from sitchensis Swaine by the
smaller average size, by the more strongly arcuate sides of the pro-
notum. by the stouter scales on the female pronotum, and by the
host.

Female. — Length 2.6 mm. (para type 2.3-2.8 mm.) 2.2 times as
long as wide; color very dark brown with a mottled pattern of dark
and light scales.

A complete description of this species is given by Blackman
(1942:24) under the name P. sericeus.

Male. — Similar to female except slightly smaller, stouter; frons
broadly, more strongly impressed; pronotal scales somewhat stouter.

Type Locality. — Pacific Grove, California.

Type Material. — The female holotype, male allotype and three
para type were collected at the type locality on December 12. 1923,
from Monterrey pine; one male paratype is labeled "Carmel Cal.,
5-IV-25. F. 0. Ballon, Pinus radiata.''

The holotype. allotype, and paratypes are in my collection.

Sept. 30. 1969 new synonymy and records 123

Phloeotribus quercinus, n. sp.

This species is closely allied to pruni Wood, but it may be distin-
guished by the less strongly attenuate lateral extension of segments
of the antennal club, by the absence of submarginal crenulations
near the elytral bases, by the coarser pronotal punctures, and by
the host.

Female. — Length 2.0 mm. (paratypes 1.8-2.1 mm.), 1.9 times
as long as wide; color black.

Frons evenly convex above level of antennal insertion, trans-
versely impressed below, smooth epistomal margin slightly elevated;
premandibular lobe small, distinct; surface strongly reticulate, punc-
tures small, obscure, rather close; vestiture fine, hairlike, rather
sparse. Antennal club with segments 1 and 2 each very slightly less
than twice as wide as long.

Pronotum 0.82 times as long as wide; widest at base, sides
arcuate, converging to a slight constriction just before broadly round-
ed anterior margin; surface reticulate, devoid of all indications of
asperities, punctures coarse, moderately deep, separated by distances
up to diameter of a puncture; part of median line impunctate in
front of middle; vestiture very fine, long.

Elytra 1.35 times as long as wide, 2.0 times as long as pronotum;
sides almost straight and parallel on basal two-thirds, rather broadly
rounded behind; scutellum small, rounded, scutellar notch rather
deep; basal margins each armed by 14 rather narrow, high crenula-
tions. devoid of submarginal crenulations; striae weakly impressed
toward declivity, punctures moderately coarse, deep; interstriae
slightly wider than striae, devoid of punctures, moderately large,
rounded granules moderately confused. Declivity convex, steep;
striae moderately impressed; interstriae feebly convex, not elevated,
granules almost uniseriate. Vestiture consisting of fine, semirecum-
bent strial hair, and erect, confused interstrial bristles of uniform
length, fine at base, becoming stout and shorter toward declivit}^
each almost equal in length to width of an interspace.

Male. — Unknown.

Type Locality. — Sixteen km. (10 mi.) east of Pachuca, Hidalgo,
Mexico.

Type Materlal. — The female holotype and 22 female paratypes
were collected at the type locality on June 10. 1967. 2.600 m.. No. 5,
by S. L. Wood, in broken Quercus branches less than 2 cm. in
diameter.

The holotype and paratypes are in my collection.

Phloeotribus destructor^ n. sp.

This species is closely related to frontalis (Olivier), but it may
be distinguished by the absence of the tubercles on decli vital inter-
striae 1 to 4. by the much coarser pronotal punctures, with the inter-
spaces between them devoid of impressed points, and by the different
host and distribution.

The Great Basin Naturalist
124 STEPHEN L. WOOD Vol. XXIX, No. 3

Male. — Length 2.2 mm. (paratypes 1.8-2.5 mm.) 2.1 times as
long as wide; color dark brown.

Frons shallowly, broadly concave from epistomal margin almost
to level of upper margin of eyes; lateral margins armed by a pair of
moderately large, pointed tubercles at level of antennal insertion;
surface shining, subreticulate below, almost rugulose above, marginal
areas above tubercles with rather abundant, small, rounded granules;
vestiture of rather abundant, coarse, short hair. Segment 1 of an-
tennal club 2.6 times as wide as long.

Pronotum 0.81 times as long as wide; widest at base, sides rather
weakly, arcuately converging to a feeble constriction just behind
anterior margin, rather narrowly rounded in front; surface coarsely,
very closely, rather shallowly punctured, a few granules and fine
asperities in anterolateral areas, interspaces between punctures less
than half as wide as diameter of a puncture, devoid of impressed
points; vestiture of rather fine, moderately abundant, short hair.

Elytra 1.4 times as long as wide. 1.9 times as long as pronotum;
sides almost straight and parallel on basal two-thirds, rather narrow-
ly rounded behind; scutellar notch rather shallow, obtuse; basal
margins each armed by 12-14 coarse, overlapping crenulations;
striae moderately impressed, punctures deep, rather fine on basal
fourth, becoming coarser toward declivity; interstriae as wide as
striae, closely crenulate, crenulations confused, each about half as
wide as an interstriae, low except at base and much larger at base of
declivity. Declivity convex, moderately steep; striae narrowly im-
pressed, punctures rather small; interstriae 1 to 8 equally convex,
armed at base by crenulations that increase in size laterally, inter-
striae on declivital face not at all granulate, uniseriately. finely
punctured, except 9 joining costal margin, its elevation and course
tubercles continuing to apex. Vestiture of slender, interstrial bristles
of uniform length, slightly confused on most of disc, becoming
uniseriate toward and on declivity, on declivity each equal in length
to distance between rows.

Female. — Similar to male except frons convex, a narrow, trans-
verse impression immediately above epistoma, lateral tubercles
absent.

Type Locality. — Morelia, Mochoacan. Mexico.

Type Material. — The male holotype, female allotype, an,d 5
paratypes were taken in the gardens behind No. 25 Avenida Acue-
ducto at the type locality on June 14, 1965, in a peach tree. No. 57,
by S. L. Wood; 12 paratypes were collected 19 km. (12 mi.) east of
Carapan, Michoacan, on June 18, 1965, in wild cherry. No. 76, by
S. L. Wood.

The holotype, allotype, and paratypes are in my collection.

Phloeotribus furvus, n. sp.

As indicated above, this is the Central American species pre-
viously reported as Phloeotribus sulcifrons by Blandford (1897) and
others. From the type of sulcifrons it differs by the convex female

Sept. 30, 1969 new synonymy and records 125

frons. by the more closely punctured pronotum, and by the more
finely, more closely placed interstrial crenulations (obscurely three
to four ranked on each interstriae; only two to three ranked in
sulcifrons) .

Male. — Length 3.0 mm. (paratype 2.8-3.3 mm.), 1.6 times as
long as wide; mature color black.

Frons broadly, shallowly, subconcavely impressed from epistoma
to upper level of eyes, lateral margins weakly elevated, except more
strongly raised at level of antennal insertions, unarmed, a low.
transverse callus at level of antennal insertions; surface strongly
reticulate, with rather close, shallow punctures of moderately small
size. Antennal scape with a small tuft of rather short hair, segment
1 of club more than eight times as wide as long.

Pronotum 0.80 times as long as wide; basal margin obtusely ex-
tended posteriorly, with median fourth more acutely extended;
widest just in front of posterolateral angles, rather strongly, arcuate-
ly convergent, weakly constricted just before rather narrowly round-
ed anterior margin; surface very densely, rather coarsely punctured,
becoming somewhat granulate on anterior third; anterolateral angles
with two or three coarse asperities, a row of small asperities extend-
ing along anterior margin to opposite sides; vestiture of minute, dark
setae at least in lateral areas.

Elytra 1.03 times as long as wdde, 1.4 times as long as pronotum;
sides weakly arcuate to base of declivity, rather broadly rounded
behind; scutellar notch abrupt, deep, broad; basal margins each
armed by 19 low, overlapping crenulations, striae abruptly, narrow-
ly, deeply impressed, punctures small, deep; interstriae twice as wide
as striae, dense crenulations narrow, rather high, often obscurely
three-ranked, general surface evidently minutely punctured. De-
clivity convex, steep; interstriae as on disc but slightly narrower,
median tubercles larger, others largely absent, 9 moderately elevated
from middle of declivity anteriorly. Vestiture of minute, abundant,
confused, hairlike bristles, each about equal in length to one-third
width of an interstriae.

Female. — Similar to male except frons convex, with an in-
distinct central fovea, punctures larger, very shallow, antennal scape
not ornamented by hair; interstriae 9 less strongly elevated.

Type Locality. — Turrialba. Cartago Province, Costa Rica.

Type Material. — The male holotype, female allotype and 29
para types were collected at the type locality on March 9. 1964,
2,000 ft. elevation. No. 459, from a broken limb of an unidentified
tree, by S. L. Wood.

The holotype, allotype, and para types are in my collection.

Chramesus vitiosus, n. sp.

This species was confused with crenatus Wood until an attempt
was made to construct a key to include it. From crenatus it is distin-
guished by the shorter, erect interstrial bristles, by the absence of

The Great Basin Naturalist
126 STEPHEN L. WOOD Vol. XXIX, No. 3

male strial punctures, and by the greater distribution of pronotal
punctures.

Male. — Length 1.3 mm. (para types 1.2-1.5 mm.), 1.4 times as
long as wide; color very dark brown, vestiture pale.

Frons broadly, deeply concave from epistoma to well above eyes,
lateral margins acutely, strongly elevated, attaining a subserrate
summit at level of antennal insertion, lower tubercle predominating;
surface shining, finely reticulate, punctures not clearly evident;
vestiture fine, inconspicuous. Antennal club moderately large. 2.3
times as long as wide.

Pronotum 0.72 times as long as wide; widest on basal fom-th, sides
arcuately converging to slight constriction just before rather broadly
rounded anterior margin; surface subreticulate, rather closely asper-
ate, asperities largely replaced by fine punctures in postermedian
area, a few punctures occur almost to anterior margin in median
area; vestiture of moderately abundant, short, stout, pale bristles
over entire surface.

Elytra 0.91 times as long as wide, 1.4 times as long as pronotum;
basal margins each armed by about 12 partly contigious, low crenu-
lations; sides straight and parallel on less than basal half, broadly
rounded behind; striae distinctly impressed, glabrous, punctures ob-
solete; interstriae as wide as striae, almost smooth, small punctures
close, confused. Declivity convex, rather steep; a few strial punc-
tures obscurely present. Vestiture consisting of a ground cover of
closely set short scales, each scale about twice as long as wide; and
interstrial rows of erect, scalelike bristles, each bristle not more than
twice as long as ground scales, about half as long as distance between
rows or between bristles in a row.

Female. — Similar to male except frons weakly convex, foveate at
center, a weak, transverse impression iust above epistoma; punctures
on pronotum slightly coarser and more widely distributed; strial
punctures small, distinctly impressed; occasional very small inter-
strial granules evident; scales in interstrial ground cover only slight-
ly longer than wide.

Type Locality. — Eight km. (5 mi.) south of Rosamorada, Na-
yarit, Mexico.

Type Material. — The male holotype, female allotype, and 48
para types were collected at the type locality on July 14, 1965, 100 m.
elevation, from Inga paterno, by S. L. Wood. Additional paratypes
were taken in Nayarit as follows: 14 from 48 km. (30 mi.) north of
Rosamorada, July 15. 1965, 100 m.; 1 from I^s Corchos. July 10,
1965, 10m.; all from the same host and collector.

Chramesus minulus, n. sp.

This species is closely allied to acacicolens Wood, but it is distin-
guished by the smaller size, by the more slender form, by the much
shorter, erect, interstrial scales, and by the much stouter pronotal
scales.

Sept. 30, 1969 new synonymy and records 127

Female. — Length 1.25 mm. (paratype 1.3 mm.), 1.8 timies as
long as wide; color yellowish brown.

Frons weakly convex, somewhat flattened on upper half, a slight,
transverse impression just above epistoma; surface minutely rugu-
lose-reticulate, fine punctures indistinct; vestiture short, rather
sparse, stout. Antennal club rather small for this genus. 2.0 times
as long as wide.

Pronotum 0.80 times as long as wide; widest at base, sides rather
weakly arcuate and converging slightly on basal two-thirds, broadly
rounded in front; surface rather finely, closely granulate-punctate
(largely obscured by scales), becoming finely asperate toward antero-
lateral anglts; vestiture of short, oval, almost white scales.

Elytra 1.2 times as long as wide, 1.7 times as long as pronotum;
basal margins each armed by 13 crenulations; sides almost straight
and parallel on slightly more than basal half, rather broadly round-
ed behind; striae weakly impressed, punctures coarse, shar[)ly im-
pressed, rather shallow; interstriae about one and one-half times as
wide as striae, moderately convex, smooth, with minute, confused
punctures, each interstriae with a median row of very fine granules.
Declivity convex, moderately steep; essentially as on disc. Vestiture
consisting of a ground cover of short scales, each scale as long as
wide, slightly more than a third as wide as an interstriae; and
median interstrial rows of erect scales, each only slightly longer than
ground scales and mostly less than twice as long as wide.

Type Locality. — Le Ceiba, Honduras.

Type Material. — The female holotype and one female paratype
were collected at the type locality, at light, in 1949. by E. C. Becker,
the holotype on June 10, the paratype on May 29.

The holotype and paratype are in my collection.

References Cited

Blackman, M. W. 1942. Revision of the bark beetles belonging to the genus

Pseudohylesinus Swaine. U. S. Dept. Agric. Misc. Pub. 461, 32 p.
Bi-ANDFORD, W. F. H. 1895-1905. Fam. Scolytidae. Biol. Centr.-Amer., Coleopt.

4(6): 81 -298, pis. 4-9.
Barr. B. 1969. Sound production in Scolytidae (Coleoptera) with emphasis on

the genus Ips. Canadian Ent. 101:636-672.
Bright, D. E., Jr. 1968. Review of the genus Leiparthrum in North America.

with a description of one new species (Coleoptera: Scolytidae). Canadian

Ent. 100:636-639.
Chapuis, F. 1869. Synopsis des Scolytides, 61 p. (1873. Synopsis des Scolytides.

Mem. Soc. Roy. Sci. Liege, ser. 2, 3:213-269).
Erichson, W. F. 1836. Systematische Auseinandersetzung der Familie der

Borkenkafer (Bostrichidae). Arch. Naturg. 2(l):45-65.
Latreille, p. a. 1796. Phloiotribus. p. 50, in Precis des caracteres generiques

des insects, disposes dans un order naturel. 201 p.
. 1802/3. Phloiotribus, p. 204, in Histoire naturelle. generale et par-

ticuliere des crustaces et des insectes. vol. 3. 467 p.
Mannerheim, C. G. 1843. Xylophagi. p. 296-298, in Beitrag zur Kaferfauna

der Aleutischen Inseln, der Inscl Sitka und Neu-Californiens. Bull. Soc.

Imp. Nat. Mo-scou 16 (2): 175-31 4.
. 1852. (Species nos. 143-153, in) Zweiter Nachtrag zur Kaefer-Fauna

The Great Basin Naturalist
128 STEPHEN L. WOOD Vol. XXIX, No. 3

der nord-amerikanischen Laender des loissischen Reiches. Bull. Soc. Imp.
Nat. Moscou 25(1) :283-387.

1853. Bostrichina, p. 234-239, in Dritter Nachtrag zur Kaferfauna der

Nord-Amerikanischen Lander des Russichen Reiches. Bull. Soc. Imp. Nat.
Moscou 26:95-273.

MoTSCHULSKY, V. VON. 1858. Genyocerus albipennis, p. 68, in Entomologie
speciale. Insectes des Indes orientales. Etud. Ent. 7:20-122.

. 1859. {Genyocerus adustipennis) . Etud. Ent. 9:19 (From Blandford.

1894:141).

. 1863. (Platypodidae and Scolytidae, p. 509-517, in) Essai d'un cata-
logue des insectes de I'lle Cevlan. Bull. Soc. Imp. Nat. Moscou 36(1):
421-532.

1866. (Scolytidae). Bull. Soc. Imp. Nat. Moscou 39(1) :401-404.

ScHEDL, K. E. 1959. A check list of the Scolytidae and Platypodidae (Q)leop-

tera) of Cevlon with descriptions of new species and biological notes. Trans.

Roy. Ent. Soc. London 1 1 1 (15) :469-534.
WoLLASTON, T. V. 1854. Fam. 32. Toniicidae, p. 288-306, in Insecta Maderen-

sia; being an account of the insects of the islands of the Medeiran group.

634 p.
. 1864. Fam. 43. Toniicidae, p. 254-267, in Catalogue of the coleopterous

insects of the Canaries in the collections of the British Museum. 648 p.
Wood, S. L. 1963. A revision of the bark beetle genus Dentroctonus Erichson

(Coleoptera: Scolytidae). Gt. Basin Nat. 23:1-117.
. 1969. New records and species of Neotropical bark beetles (Scolytidae:

Coleoptera). Brigham Young Univ. Sci. Bull., Biol. Ser. 10(2): 1-46.

ZOOGEOGRAPIIIC AND SYSTEMATIC NOTES ON THE

LACE BUG TRIBE LITADEINI. WITH THE DESCRIPTION

OF TI IE NEW GENUS STRAGULOTINdlS

(IIEMIPTERA: TINGIDAE)

Richard C. Froeschner^

The tribe Litadeini was proposed for the genus Litadea by Drake
and Ruhoff (1965a. p. 18) and defined by the unusual modification
of the second tarsal segment: "Enormously swollen, nearly ovate
in outline, upper surface convex, beneath deeply concave, the con-
cavity filled with short, bristlelike hairs." Subsequently these two
authors (1965b, p. 247) transferred Holophygdon Kirkaldy (1908,
p. 364) from Fingini to Litadeini and described another genus,
Aeopelys^ as a member of Litadeini.

During preparation of a key to the genera of Tingidae of the
world, it was found that on the basis of the modified second tarsal
segment five more genera must be transferred from the tribe Tingini
to the Litadeini (new assignments here) : Aristobyrsa Drake and
Poor (1937, p. 164), Larotingis Drake (1960, p. 357). Oecharis
Drake and Ruhoff (1965b. p. 280). Ogygotingis Drake (1948, p.l49),
and Psilobyrsa Drake and Hambleton (1935, p. 148). In addition,
two species cataloged in the Tingini genus Pleseobyrsa Drake and
Poor have the modified tarsi of the Litadeini but do not fit well
into any of the other genera and so must have a new genus erected
for them.

The tribe Litaeini originally containing but one genus and species
from the Indian Ocean island of Rodriguez, now includes thirteen
species in nine genera and is almost Pan-Tropical in distribution.
It is known from Africa and South America, and from the islands
in the Indian (Rodreguez I.) and Pacific (Fiji and New Guinea)
Oceans.

In view of the several genera added to the tribe Litadeini, the
significant modifications of the second tarsal segment might profit-
ably be reviewed and slightly modified. The segment is very strongly
dilated at the base so that it is abruptly much wider than the first
segment. The ventral surface is flat or concave (apparently arching
inward during drying) and densely hairy with the hairs toward
the sides of the segment much lengthened. The shape is not always
broadly oval; in Oecharis it is elongate oval, tapering from the wide
base to the slender claw-bearing ajiex. The functional significance
of these modifications has not been investigated by observation of
the living insect in the field.

^Sniitlisonian Institution, AVashington, D. C.

129

130

RICHARD C. FROESCHNER

The Great Basin Naturalist

Vol. XXIX, No. 3

Stragulotingis, n. gen.
Fig. 1

Diagnosis: Within the tribe Litadeini, Stragulotingis is recogniz-
able by the broad, flat, horizontal paranota associated with the flat
discoidal areas.

Characters: Length 3.0-3.7 nun.; macropterus.

Head short, vertically deflexed, wdth 5 long, cephalic spines
(occipital and supraclypeal pairs plus a median one above base of

EHF

Fig. 1 Stragulotingis plicata (Champion)

Se|)t. 30. 1969 new genus stragulotingis 131

latter) ; eye about one-third as wide as Liiterot ular space; bucculae
short, not reaching apex of clypeus; labium extending onto apical
half of niesosternuni; antennae slender, cylindrical, all segments
with long, slender bristles, segment I nearly or quite as long as inter-
ocular width, about three times as long as II, III slightly longer
than I plus II. about one and a half times as long as IV.

Pronotum without inflated cysts; anterior margin convexly pro-
jecting almost to apex of head; median carina low, with a single
row of small cells; lateral carinae reaching to calli. weak, lower than
median carina, with cells obscure or absent; paranotum anterolater-
ally expanded obliquely, attaining a line across apex of head, lateral
margins concavely converging posteriorly, terminating just posterior
to humeral angles, anteriorly with long marginal bristles; posterior
})ronotal projection acute with apex narrowly rounded, extending to
basal third of discoidal area.

Elytra conjointly nearly flat, multiseriate subcostal area oblique;
outer margins, except basal half of sutural margin, with a row of
numerous close-set, long tapering bristles; discoidal area reaching al-
most to midlength of elytron; costal margins strongly convex basally.
extending distinctly cephalad of elytral articulation, thence straight
and nearly parallel; costal area broad, horizontal, with 5-7 irregular
rows of cells at widest point, surface transversely weakly depressed
at basal fourth and midlength; sutural areas narrowly overlapping,
leaving rounded apices widely separated; hypocostal lamina uniser-
iate, more or less remote from sides of abdomen.

Peritreme absent. Metapleural flange reduced, not expanded.
Sternal laminae widely separated, strongly diverging on mesoster-
num, more widely separated and strongly outbowed on metasternum
where their posterior apices are connected by a weak transverse car-
ina. Abdomen convex basoventrally.

Type of genus: Pleseobyrsa plicata Champion, here designated.

Geographical distribution: The genus occurs in Tropical America
from Costa Rica south to Brazil.

Comments: This genus admittedly is quite close to Aristobyrsa on
a number of significant features, such as the long hairs of the anten-
nae; the basal part of the costal area projecting convexly cephaled
of the elytral articulation; the wdde, anteriorly projecting paranotum;
and the broad, coarsely lacey, diverging elytra. But two elytral feat-
ures separate them. Aristobyrsa has the discoidal area very strongly,
tumidly elevated and tilted outward to overhang the subcostal area,
while Stragulotingis has the discoidal area low and flat. In addition,
Aristobyrsa has the subcostal area vertical, much narrowed, sub-
coriaceous, with the veins much thickened and the included two rows
of cells punctiform, while the subcosta on the two species of Strag-
ulotingis is four to six cells wide with the veins and cells developed
similarly to those of the discoidal area.

The j)refix of the new name is from the Latin "stragulus'' im-
plying a spreading out and refers to the wide paranota and costal
areas which expand anteriorly as well as laterally.

The Great Basin Naturalist
132 RICHARD C. FROESCHNER Vol. XXIX. No. 3

The two species included here were listed under the genus Plese-
obrysa in the Drake and Ruhoff (1965a) catalog.

List of species of Stragulotingis

atratarsis (Drake and Hambleton), n. comb.

Pleseobyrsa atratarsis Drake and Hambleton 1946b, p. 124. Peru
plicata (Champion), n. comb.

Leptobyrsa plicata Champion 1897, p. 26. Panama.

Pleseobyrsa parana Drake and Hambleton 1944a, p. 95 [synony-
my Drake and Ruhoff 1965a, p. 343]. Brazil.

Key to species of Stragulotingis

A. Tarsi black; size larger, length 3.6-3.7 mm. atratarsis (Drake
and Hambleton) .
Tarsi yellow to tan; size smaller, length 3.0-3.2 mm. plicata
(Champion)

References

Champion. G. C. 1897. Rhynchota. In Godman and Salvin, Biologia Centrali-
Americana, vol. 2, pp. 1-32.

Drake, C. J. 1948. New genera and species of Tingidae (Hemiptera). Proc.
Biol. Soc. Washington, vol. 61, pp. 149-156.

. 1960. Tingidae of New Guinea (Hemiptera). Pacific Insects, vol. 2,

pp. 339-380.

, AND Hambleton, E. J. 1935. New Brazilian Tingidae (Hemiptera).

Part I. Revista Ent., Rio de Janeiro, vol. 4, pp. 435-451.

. 1944. Four new American Tingitidae (Hemiptera). Proc. Ent. Soc.

Washington, vol. 46, pp. 94-96.

. 1946. Three new species and a new genus of American Tingidae

(Hemiptera). Ent. News, vol. 57, pp. 121-125.

, AND Poor, M. E. 1937.Concerning the genus Leptobyrsa Stal (Hemip-
tera). Proc. Biol. Soc. Washington, vol. 50, pp. 163-166.

, AND Ruhoff, F. A. 1965a. Lacebugs of the world, a catalog (Hemip-
tera: Tingidae). U. S. Nat. Mus. Bull. 243, pp. i-viii, 1-634, pis. 1-56.

1965b. Lacebugs from New Guinea, Borneo, Solomon, and other is-

lands of the South Pacific and Indian Ocean (Hemiptera: Tingidae). Pacific
Insects, vol. 7, pp. 243-290.
KiRKALDY, G. W. 1908. A catalogue of the Hemiptera of Fiji. Proc. Linnaean
Soc. New South Wales, vol. 33, pp. 345-391.

A NEW SPECIES OF OPPIA FROM COLORADO
(CRYPTOSTKiMATA; OPPIIDAE)

Tyler A. Woolleyi

A Student brought in a soil and humus sanij)le from a (commer-
cial?) earthworm bed that was teeming with enchytraeids and uropo-
dine mites. Associated with these organisms but not in such large
numbers, was a species of one of the larger Oppia. Specimens of this
species were compared with various species of this genus in the
author's collections and with species described in the literature, par-
ticularly the species of the Oppia complex delineated by Hammer
(1968). As a result of these comparisons, the species was determined
to be a new representative of Oppiidae and is described below.

Oppia color adensis, sp. n.

(Figs. 1-4)

Diagnosis. The new species resembles Oppia trichosa Hammer,
1958. and O. notata Hammer, 1958, as well as O. yodai Aoki, 1965,
but the sensillus in each of these last three species is pointed and
lanceolate rather than the blunt and slightly clavate type found in
O. coloradensis, sp. n. Aoki's species also has five genital setae com-
pared to four in O. coloradensis. The sensillus of the new sjiecies
resembles the type found in Amerioppia, but the latter genus lacks
interlamellar hairs, which are present in the new species. The ros-
trum of O. notata is pointed and has a transverse band with a small
tooth; O. trichosa shows a rounded rostrum, as does O. coloradensis.
In both O. notata and O. trichosa the interlamellar hairs are lacking,
while in O. coloradensis these hairs are prominent. The new species
also has similarity to O. simplex, Balogh, 1962, but the notogastral
hairs of the latter species are much longer and more robust than in
O. coloradensis; the notogastral hairs of O. yodai Aoki, 1965, are
about the same relative length as in the new species.

Compared to O. covarrubiasi Hammer, 1968. from New Zealand,
the new species has only four pairs of genital hairs (rather than
six), the interlamellar hairs, exobothridial hairs and notogastral
hairs of the new species are much longer.

The new species apparently lacks notogastral setae ta in all of
the specimens examined.

Description. Golden-brown in color, with darker, reddish-
brown margins; integument finely granlated on both prodorsum
and notogaster; prodorsum triangular in outline, rostrum rounded,
rostral hairs four-fifths as long as lamellar hairs, finely barbed;
lamellae or lamellar ridges absent, lamellar hairs a fifth again as
long as rostral hairs, finely barbed, decurved; dorsum between pedo-

'Department of Zoology, Colorado State University, Fort Colluis.

133

134

TYLER A. WOOLLEY

The Great Basin Naturalist

Vol. XXIX. No. 3

Fig. 1. Oppia color adensis from the dorsal aspect, legs omitted; A. enlarged

view of sensillus.
Fig. 2. O. coloradensis from the ventral aspect, legs partially shown.
Fig. 3. Prodorsum and part of nonogaster of O. coloradensis from the lateral

aspect, showing tubercles associated with bases of legs I-III and pedotecta I.
Fig. 4. Tibia and tarsus I of O. coloradensis.

tecta I with sculptured depressions or circles, interlamellar hairs
shorter than rostrals. more robust, barbed; exobothridial hairs about
as long as rostrals, but more definitely barbed; pseudostigmata small,
circular; sensillus clavate, with fine spines or barbs (terminal barbs
make the tip appear lanceolate, but the outline is clavate), curved
outward, upward (position varies in mounted specimens), surface
of head and two-thirds of pedicel barbed (Fig. lA); prodorsal surface
laterad of pseudostigmata finely tuberculated from pedotecta I to
posterior margin, small, rounded tubercles extending into concavity
of pedotecta I (Figs. 1,3).

Outline of notogaster nearly round, slightly narrowed posteriorly,
nine pairs of notogastral hairs present (setae ta appear to be missing

Sept. 30. 1969 new species of oppia 135

in all specimens observed), main dorsal setae robust, curved, barbed,
slightly longer than lamellar hairs; setae and fissures /«, im as in
Fig. 1, fissure im equidistant between the setae ti and ms.

Camerostome oval, ventral apodemata. and setae as seen in Fig.
2; apodemata IV arched from behind insertion of legs IV to join the
coxisternal apodeme anterior to genital aperture; genital aperture
about Yi as large as anal, genital covers nearly rectangular, each
with four setae, g:l-3 in nearly straight line nearer medial margin
than lateral, g:4 in posterolateral corner of cover; aggenital setae,
f issue iad and adanal setae as in Fig. 2; anal opening rounded, anal
covers each with two setae.

Legs monodactylous

Measurements. Length 468 /«,, prodorsum 162 /x. hysterosoma
324 ti; width 282 /t.

The type (a male), 10 male para types and 10 female paratypes
were collected from a dirt sample with enchytraeids and uropodines,
Fort Collins. Larimer County, Colorado. 1 December 1954. by T. A.
Woolley; two additional females were taken from Mount Meeker
Camp Ground, Boulder County, Colorado, 17 July 1952, by T. A.
Woolley. The type will be deposited in the U. S. National Museum.

DicussiON. Hammer (1968) admits the difficulty of differen-
tiating species of the "Oppia" complex, but she (1961, 1962. and
1968) evaluated and arranged species of this complex into a number
of genera in a way that should be helpful; at least it is a start toward
clarification of the complex. Admittedly, as Hammer indicates,
Oppia continues to be an extraordinarily complicated genus or cryp-
tically a series of genera, and the use of the genus Oppia {sensu
stricto Koch) for species that do not fit in allied genera is resorted to.
not as finally correct, but because the placement of such species in
this genus is as close as one can come within the present taxonomic
status of this group.

Hammer (1968) also delineates and provides a key for the genera
of the Oppia complex in both South America and New Zealand.
The North American representatives of this complex are little known
in comparison. Higgins and Woolley (1965) described Spinoppia
magniserrata as a new genus and species of this complex. Woolley
(1957) also redescribed Oppia minuta (Ewing) from this group.
Other representatives are known from North America, but no at-
tempt has yet been made to assess the numbers of species and evalu-
ate their placement in the current scheme of genera. The minute
size of the representatives and the immense variety of forms are
deterrants to a revision of this complex, even though such a revision
and evaluation is sorely needed.

Literature Cited

Balogh. J. 1962. Recherche sur la Faune Endogee de Madagascar VII

Naturaliste malgache 13:121-151.
AoKi, JuN-icHi. 1965. Oribatidens Thailands I. Nature and Life in Southeast

Asia IV: 129-193.

The Great Basin Naturalist
136 TYLER A. WOOLLEY Vol- XXIX, No. 3

Hammer, M. 1958. Investigations of the Oribatid Fauna of the Andes Moun-
tains I. The Argentine and Bolivia. Biologiske Skrifter Det Kongelige
Danske Videnskabemes Selskab 1 0 ( 1 ) : 1 - 1 29.

. 1961. Investigations on the Oribatid Fauna of the Andes Mountains

II. Peru. Biol. Skr. Dan. Vid. Selsk 10(1): 1-157.

. 1962. Investigations on the Oribatid Fauna of the Andes Mountains

III Chile. Biol. Skr. Dan. Vid. Selsk. 13 (2): 1-96.

1968. Investigations on the Oiibatid Fauna of New Zealand part III.

Biol. Skrifter det Kongelige Danske Videnskabemes Selskab. 16(2): 1-96.
HiGGiNs, H. G., AND T. A. WooLLEY. 1965. An Unusual New Genus of Mites

from Florida. Florida Entomologist 49(l):67-70.
WooLLEY, T. A. 1957. Redescriptions of Ewing's Oribatid Mites, V. — Families

Belbidae and Oppiidae. Entomological News 58(8) :21 1-221.

A NEW (tENUS and SPECIES OF ORIBATID MITE
(ACARI: LIACAHOIDEA. MEnUOPPIIDAE;'

Tylor A. Woolley-

About two years ago Dr. Henry Dybas very kindly sent me a
number of collection of mites for sorting. Among the many oribatid
specimens was one that slightly resembled Pyroppia lanceolata
Hammer, 1955. but was larger (930 /i x 750 /-. vs. 620-670 /,. .\ 400
ix), and with different sensilli and lamellae. After comparing this
mite with other known genera. I considered it to be a new genus and
a new species in the family Metrioppiidae. It is described below.

Metapyroppia, n. gen.

Diagnosis. With s])indleform sensillus. short lamellar cusps, an
incomplete translamella; Pyroppia lacks a translamella and the
sensillus is clavate-lanceolate. The large, straight, trochanteral setae
of Ceratoppia and Pyroppia are apparently lacking in this new genus.
The new genus differs from Paenoppia Woolley and Iliggins. 1965.
in the shorter lamellar cusps, the incomplete translamella and the
spindleform sensillus; superficially, it resembles, yet differs from
the oribatuloid Conoppia in the sensillus, the prodorsal hairs and six
pairs of genital setae.

Type-species, Metapyroppia doratosa Woolley, monobasic.

Metapyroppia doratosa n. sp.

(Figs. 1, 2)

Diagnosis. The new species differs from Pyroppia lanceolata
Hammer, 1955, in the barbed, spindleform sensillus, contrasting
with the clavate-lanceolate sensillus of P. lanceolata; it differs also
in the presence of transverse, prodorsal carina, in size and in the
lack of trochanteral setae. The prefix meta implies "near" Pyroppia.
and doratoSy a spear-like sensillus.

Description. Color dark reddish-brown, prodorsum broadly
triangular in outline; rostrum smooth, rostral hairs barbed, incurved,
about as long as lamellar hairs, inserted in slight notches at antero-
lateral margins of prodorsum (Fig. 1); a slight transverse carina
anterior of lamellar cus})s across surface of prodorsum; lamellae
narrow, extending from dorsosejugal suture to bases of rostral hairs,
attenuated anteriorly, lamellar cusps short, truncate, about as long as
each end of interrupted translamella; lamellar hairs stout, barbed,
nearly straight, about the same length as rostral hairs; translamella
interrupted medially, short medial bar each side about as long as
length of lamellar cusp; interlamellar hairs barbed, slightly longer

'Research supported by NSF Grant GB iSTl.

-Department of Zoology, Colorado State University. Fort Collins.

137

138

TYLER A. WOOLLEY

The Great Basin Naturalist

Vol. XXIX, No. 3

Fig. 1. Dorsum of Metapyroppia doratosa, legs omitted; hysterosoma broken in

type specimen.
Fig. 2. Venter of M. doratosa, legs omitted.

than lamellar hairs, inserted mediad of bases of lamellae; pseudo-
stigmata posterior to pedotecta I. slightly beneath hysterosomal
margin; sensillus spindleform, barbed; pedotecta I stout, angled
inward at anterior margin, forming lateral angles of broad triangular
prodorsum.

Hysterosoma glabrous, nearly round in outline (slightly broken
across dorsum of type specimen), with two humeral bristles posterior
to pseudostigmata at shoulders of hysterosoma. Other setae and fis-
sures as seen in Fig. 1.

Camerostome broadly oval, infracapitulum, ventral setae with
prominent alveoli, and apodemata as seen in Fig. 2; trochanteral
fossae II with slight tubercules on surface (like some other Lia-
caroidea); genital aperture nearly square, between insertions of
legs IV, each genital cover with six setae. g:l-g:4 in nearly straight
line near medial margin of cover, g:5 laterally displaced in pos-
terior half of cover, g:6 near posterior margin, but not as close to
medial margin as g:l-g:4; aggenital setae laterally placed at level
of posterior margin of genital opening; anal aperture pentagonal,
only slightly larger than genital aperture, each anal cover with two
setae, a:l in anterior half of cover, both anal setae about in middle
of width of cover; preanal piece large, trumpet-shaped; iad fissure
very narrow, almost indistinct, closely appressed to rim of anal
aperture anterior to level of a:l; three pairs of adanal setae. ada:3
between levels of a: 1 and a: 2 but closer to a: 1 and remote from anal
openings by widest width of cover; ada:2, ada:l posterior to anal
Oldening.

Sept. 30. 1969 new genus and species of mite 139

Legs heterotridactylous; middle (empodial) claw only slightl}-
larger than lateral (true) claws: stiff, straight seta of trochanter 11
(as found in Cerotoppia and Pyroppia) lacking.

Measurements. (As taken in broken type specimen) Length:
930 /t, prodorsum 180 /t. hysterosoma 750/i; width 726 /i.

Distribution. One female and type specimen was collected
from moss on log on Mt. Le Conte, Sevier County, Tennessee, August
1956, by H. Dybas. The type will be deposited in the U. S. National
Museum.

Discussion. This new genus and species represents yet another
example of oribatid mites in the family Metrioppiidae. The char-
acteristics of the sensilli. number of genital setae, tuberculous tro-
chanteral fossae II are characteristics that ally these mites with the
Liacaridae and Xenillidae in the Liacaroidea. It appears from re-
search completed previously and some currently in progress that
these families exhibit many characteristics in common. Delinea-
tions of the details of these comparative features and specifically
demonstrable relationships await the results of further studies now
in progress.

Literature Cited

Hammer, M. 1955. Alaskan Oribatids. Acta Arctica VII: 1-36.

WooLLEY, T. A.. AND H. G. HiGGiNs. 1965. A New Genus and Species of

Oribatid Mite from Colorado (Acari: Oribatei, Ceratoppiidae) . Great Basin

Naturalist 25(3-4) :59-62.

A NEW SPECIES OF LIACARUS FROM A WOOD RAT NEST
(ACARINA: CRYPTOSriGMATA, LIACARIDAE)

Harold G. Higgins^ and Tyler A. WooUey-

A number of different groups of mites are represented in the
fauna found in rodent nests. Many of the mites are predaceous
forms and the biotic relationships have not been explained fully.
Others are parasitic forms that may have temporarily moved off
their hosts (Daniel, 1969). Free-living oribatid mites are sometimes
found in nests (Woolley and Higgins, 1968), but have been reported
infrequently up to now. It is possible that these oribatids are natural
inhabitants of the soil adjacent to the nests of the rodents and have
migrated in, but they could also have been picked up from the soil
and carried from nest to nest in the fur of the animals.

In the spring of 1969 a collection of mites taken from the sleeping
nest of the Wood Rat {Neotoma fuscipes) was received from Mr.
Tom Ashley of El Cajon, California. Among these was an unde-
scribed species in the genus Liacarus as designated and summarized
in the article of Woolley (1968). The new species has the barbed
and spindleform sensillus typical of the genus, but specific differ-
ences separate it from known species.

Liacarus lectronus, sp. n.

(Figs. 1-3)

Diagnosis. The new species is most similar to Liacarus robustus
Ewing, 1918, but is smaller in size and has a much smaller mucro.
wider lamellae, pointed lamellar cusps and much longer interlamel
lar hairs. The trivial name comes from the Greek lektron implying
"bed" and refers to the sleeping nest of the rodent from which it
was taken.

Description. Color dark reddish-brown; prodorsum triangular,
about a sixth as long as notogaster; rostrum notched, tip rounded;
rostral hairs two-thirds as long as lamellar hairs, finely barbed, in-
serted in raised forward ridge of tutoria; lamellae broad, narrowed
anteriorly, with bidentate cusps, medial dens twice as large as lateral
dens; lamellar hairs about half as long as interlamellar hairs, insert-
ed in dorsal surface of lamellae just behind anterior notch of lamellar
cusp, finely barbed (Fig. 2); translamella with a prominent mucro
extending from base of lamellae to level of insertions of lamellar
hairs; interlamellar hairs twice as long as lamellar hairs, extended
forward from insertions to level of tips of rostral hairs, finely barbed,
inserted in medial margins of lamellae near dorsosejugal suture;
pseudostigmata covered by clear, narrow shoulder patches of noto-

'Granger High School. Salt Lake City, Utah.

-Department of Zoology. Colorado State University. Fort OUins; research supported in part hy
NIH-NIAID rO TOI-AIO0694-IO.

140

Sept. 30. 1969

NEW SPECIES OF LIACARUS

141

Fig. 1. Liacarus lectronus, from the dorsal aspect, legs omitted.
Fig. 2. L. lectronus, enlarged view of lamellae, mucro and lamellar hairs.
F'ig. 3. L. lectronus, from the ventral aspect; legs partially omitted, but show-
ing keeled femora I, II, III.

gaster, spiraled beneath surface; sensillus spindleform, finely barbed;
pedotecta I with rounded tip.

Notogaster oval in outline; narrowed, clear shoulder patches ex-
tended over sensillus, with two fine, humeral bristles; ten pairs of
fine notogastral setae visible in specimens (Fig. 1 ).

Camerostome oval, infarcapitulum diarthric; rutella, mentum,
ventral setae and apodemata as in Fig. 3; genital opening smaller
than anal; each genital cover with six setae. g:5 inserted more lat-
erally than other setae on cover; aggenital setae widely separated,
insertions closer to genital opening than to anal; anal opening nearly
square, each anal cover with two setae inserted nearer medial margin
than lateral; fissure iad and adanal setae as in Figure 3. ada:3 more
remote from opening than ada:2 or ada: 1.

Legs heterotridactylous; femora I. II. Ill each with fine pointed
keel.

Measurements. The type female measures 702 /i x 438 ^w.

Collection Data. Three specimens, two females and a male,
were collected from the sleeping nest of Neotoma juscipes, 20 March
1968, at Cuyanaca Reservoir. San Diego Co., California, by Tom
Ashley. The type will be deposited in the U. S. National Museum.
The paratypes will be retained by the authors.

Discussion. In the same collection with L. lectronus, sp. n.
from the Neotoma nest was another species that has been recorded
from Europe previously, but not from the United States.

The Great Basin Naturalist
142 H. G. HIGGINS AND T. A. WOOLLEY Vol. XXIX, No. 3

This unusual find consisted of four specimens of Cosmochthonius
reticulatus Grandjean, 1947. Although several species of Cosmoch-
thonius are reported from Europe, South America, and New Zealand,
to our knowledge this is the first time that this European form has
been reported from the United States.

Preliminary studies indicate that studies of rodent nests in the
future may disclose a source of desert forms of oribatids that is cur-
rently untapped.

Literature Cited

Daniel, M. 1969. Nests of small terricolous mammals as the environment of

nidicolous paiasites. Folia Parasitologica (Praha) 16:137-142.
EwiNG, H. E. 1918. The Beetly Mite Fauna of Mary's Peak, Coast Range,

Oregon. Entomol. News 29(3) :81-90.
WooLLEY, T. A. 1968. North American Liacaridae. W-Liacarus. J. Kansas

Entomol. Soc. 41 (3) : 350-366.
W00L1.EY, T. A.' AND H. G. HiGGiNs. 1968. A new genus and species of

oribatid from pack rat nests. Great Basin Nat. 28(3) : 144-146.

INVASION OF A "TRAPPRD-OU 1" SOUTI lERN NEVADA
HABITAT BY PER0(;NATIIUS LONGIMEMRIUS

Lester D. Flake' and Clive D. Jorgensen-

Abstracts a 6.3 ha grid was established at the United States Atomic Energy
Commission's Nevada Test Site and the small mammals trapped-out during the
summers of 1964 and 1965 to study invasion by Perognathus longimembris Coues.
Age analyses were made to detemiine the relationship between age and invasion.
The mean age of invading animals was lowest in mid- and late summer, but
varied widely with reproductive success. There was no statistically significant
difference in mean age between male and female invaders. Ratios of male to
female invaders varied directly with that of the natural population. Invasion
rates varied widely and were mainly influenced by population density outside
the grid.

Introduction

Small mammal invasion studies of "trapped-out" habitats are not
numerous; but rates of invasion, home site origin, relationship of
invasion behavior to population density, juvenile or adult classifica-
tion of invaders, and sex ratios of invaders have been examined for
several species by Andrzejewski (1963), Andrzejewski and Wroclo-
wek (1962), Blair (1940), Calhoun and Webb (1953), Stickel (1946)
and Webb (1965). These studies were made in non-arid environ-
ments and did not include any of the heteromyid rodents.

In this study a "trapped-out" habitat was created and Perog-
nathus longimembris Coues movement into the grid (invasion) ex-
amined. Special attention was given to age composition of invading
P. longimembris. The possibility that male and female invaders come
from different age groups was examined along with sex ratios and
rates of invasion. These analyses provide the basis for determining
how the population characteristics of P. longimembris affect its in-
vasion behavior.

Materials and Methods

The study was conducted at the United States Atomic Energy
Commission's Nevada lest Site, 70 miles northwest of Las Vegas,
Nevada. The study area was in relatively undisturbed Grayia-Lycium
community (Allred, Beck and Jorgensen, 1963) in the northeast por-
tion of Yucca Flat.

A 6.3 ha grid was established with 12 parallel transects placed
at 22.9 m intervals. Each transect contained 12 trapping stations
22.9 m apart. Two Young-type live traps were placed at each of the
144 trapping stations.

Small mammals were trapj)ed and removed daily from June 7
through Aug. 25, 1964, and from June 9 through Sept. 7. 1965. Small
mammals were removed and traps baited with small amounts of

'Natural Resource Ecology I.al)oratory, Colorado State University, Fort Collins, Colorado.
-Departiiipnt of Zoology and Entomology, Brighani Young University, Provo, Utah.

143

The Great Basin Naturalist
144 L. D. FLAKE AND C. D. JORGENSEN Vol. XXIX, No. 3

rolle doats prior to 9:00 a.m. each day. Animals were killed and
taken to the laboratory where age and sex were determined.

During Aug. 1965, three 460 m transects, 100 m apart, were es-
tablished three miles southwest of the grid to sample the untrapped
population for mean age. Fifty equally spaced Museum Special traps
were placed on each transect and operated for seven nights between
Aug. 15 and Sept. 7. All P. longimembris were taken to the laboratory
for age and sex determinations.

The tooth wear technique developed for Perognathus by Howard
Kaaz (University of California, Los Angeles, California, unpub-
lished data) was used for age determination. A key for this technique
and accompanying illustrations are included in Flake (1966).

Results

The number of P. longimembris trapped daily leveled off by the
sixth day after trapping began in 1964 and those trapped after that
were considered to be invaders. Numbers of P. longimembris trapped
per day for the first 10 days of trapping in 1964 were respectively:
39, 15, 32, 15, 22. 16, 4, 7, 4 and 2. In support of the above observa-
tion it was found that after the fifth day of trapping, almost all cap-
tures were within 68.7 m of the borders. Similar results were obtained
during the first six trapping days of 1965.

Mean ages of P. longimembris for the trapping-out periods and
subsequent invasion periods are presented in Table I. Table II gives
the mean age by sex and period of capture. Hereafter, the June 7-
11. June 13-30, July 1-31 and Aug. 1-25, 1964, periods; and the
June 9-13, June 15-30, July 1-31 and Aug. 15 - Sept. 7, 1965, periods
are referred to as the 1st. 2nd. 3rd and 4th periods for the respective
years. June 7-11, 1964, and June 9-13, 1965, are actually trapping-
out periods while all other trapping intervals represent, by definition,
invasion periods.

An analysis of variance of the age data, using the fixed effects
model (model I), disclosed statistically significant differences (P -
.05) in the main effect means for the years x periods interaction.
The years x sex. periods x sex, and years x periods x sex interactions
were not significant. Mean age comparisons within the years x per-
iods interaction were accomplished with Cramer's modification of
Duncan's multiple range test (Cramer, 1956). Use of the expression
"significant difference" throughout the papers refers to a statistically
significant difference at the 5 percent level.

Comparison of mean ages for 1964 and 1965 trapping periods
(Table I) yielded the following results. In 1964 and 1965 there was
no significant difference between means for the 1st and 2nd periods
within either year. For both years there was a significant difference
between the mean for the 1st period and the means for the 3rd and
4th periods. Similarly, there was a significant difference between the
mean for the 2nd period and the means for the 3rd and 4th periods
in both years. No significant difference was observed between the
means for 3rd and 4th periods within years. There was no significant

Sept. 30. 1969 perognathus longimembris 145

difference in 1st and 2nd period means between years, but there was
a significant difference in means for the 3rd and 4th periods between
years. In siunniary, the mean age of invaders decreased from early
to late sunnner in both 1964 and 1965 though the decrease was much
greater in 1965.

As noted previously, P. longimembris were trapped from an un-
tra])ped population three miles southwest of the grid between Aug.
15 and Sept. 7, 1965. The mean age (n = 88, x = 4.6 months,
s = 7.9 months was not significantly different from that of period
4, 1965. grid invaders (Table I).

The ratios of males to females for the trapping periods are given
in Table I. In 1964, there were 1.3 males per female captured during
period 1 (trapping-out period). Since reproduction was negligible in
1964 (French. Maza and Aschwanden. 1967), the sample ratio for
the 1st period may be used to estimate the sex ratio in the untrapped
population outside the grid. Among total invaders for 1964, 1.5 males
invaded per female. A chi-square test disclosed no significant dif-
ference in proportion of males to females between the period
1 estimate of the untrapped population and total invaders
(X- (1 df, .05) — .08. In 1965 among total invaders 0.8 males in-
vader per female. When the proportion of males among total
invaders was compared with that for P. longimembris trapped
three miles southwest of the grid from Aug. 15 to Sept. 7
(n = 88, 42 males, 46 females), no significant difference was found

X- '1 <lf, ,051— .01).

Invasion rates varied from one period to the next as well as be-
tween 1964 and 1965 (Table 1). One may question the high invasion
rate from Aug. 15 to Sept. 7. -965, since the grid was not trapped the
two previous weeks. The daily trapping records (Flake. 1966) show
that the largest numbers per day were observed during late August
and early September rather than mid-August. If there had been ex-
tensive invasion and settlement during the tw^o weeks when there
was no trapping, the highest trapping incidence would be expected
to occur early after trapping was resumed. These data suggest the
increase in rate was mainly due to invasion and not belated capture.

Discussion and Conclusion

Reproductive activity and its relationship to the environment is
an essential consideration in understanding age distribution and in-
vasion. Hall (1946) reported that P. longimembris have one litter
per year and noted pregnancies from early April through June. Duke
(1957) stated that Perognathus in Utah may have two litters each
year and that pregnancy is possible between early April and early
June. Speth. Pritchett. and Jorgensen (1968) found that Perognathus
parvus in southern Idaho may have two litters per year and suggested
that the number of litters depends on food availability. At the Nev-
ada Test Site. French. Maza and Aschwanden (1967) reported ex-
tremely low rainfall and poor plant growth in the winter and spring
of 1962-63 and 1963-64. They found essentially no P. longimembris

146

The Great Basin Naturalist
L. D. FLAKE AND C. D. JORGENSEN Vol. XXIX. No. 3

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The Great Basin Naturalist
148 L. D. FLAKE AND C. D. JORGENSEN Vol. XXIX. No. 3

reproduction in the summers of 1963 and 1964. When conditions im-
proved for plant growth in the winter and spring of 1964-65, they
noted rapid population growth the following summer. During this
period they found several individual P. longimembris living from 2
to 5 years in the field. Jorgensen (Brigham Young University. Provo,
Utah, personal communication) also noted that several P. longimem-
bris survived at least 3 years in the field at the Nevada Test Site.

The relatively high mean age of animals trapped in 1964 and
June of 1965 probably reflects extremely poor reproduction in the
summers of 1963 and 1964. Thus, a mean age of 22 months (June
of both years) is not unreasonable considering the extremely limited
addition of young to the population during and prior to trapping. The
difference in mean age of 3rd and 4th period invaders between years
is likely a reflection of poor reproduction in 1964 versus increased re-
production in 1965. The decrease in mean age through both summers
correlates well wdth appearance of young in the population and re-
production success.

There was no significant difference in the mean age of male and
female P. longimembris invaders (Table II) and no significant inter-
action involving sex; therefore, the sexes were treated as a single
group in age analysis.

As there was no significaant difference between age of animals
in the un trapped population (3 miles southwest of the grid) and 4th
period invaders for 1965, it could not be concluded that young an-
imals have a greater tendency to invade than adults. The mean age
of the untrapped population was extremely small x = 4.6 months,
s = 7.9 months), probably because of high reproductive success.
Hence, a preferential invasion by young P. longimembris probably
would not have been detectable.

Stickle (1946) observed that adult Peromyscus leucopus invading
a "trapped-out" grid consisted of 2 males per female while the nat-
ural population in the immediate area contained approximately equal
numbers of males and females. In the present study of P. longimem-
bris, males invading the study grid outnumbered females in 1964
and similar ratios were obtained within the grid's original population
(trapping out period) which was used to estimate the ratio in the un-
trapped population. Contrary to the 1964 results, females invaded
more frequently than males in 1965. In 1965, the sex ratio of P.
longimembris from the untrapped population 3 miles southwest of
the grid was compared with that in the study grid and ratios of males
to females for both samples were almost identical. These results in-
dicate the sex ratios of invading P. longimembris are approximately
the same as those in the natural population.

The rates of invasion are likely influenced by several population
characters. Stickle (1946) noted that Peromyscus with home ranges
overlapping the borders of a newly established void were first to in-
vade. Andrezejewski and Wroclowek (1962) concluded that the rates
of invasion of Apodemiis and Clethrionomys were determined by the
density of the adjacent population, and further Andrzejewski (1963)

Sept. 30. 1969 perognathus longimembris 149

stated that increased rates of invasion and settling during the mid-
sunnner were due primarily to emergence of young and subsequent
increase of density.

In this study, the highest invasion rate in 1964 occurred from
June 13 to 30 and was possibly the result of the rapid removal of
animals with home ranges overlapping or adjacent to the grid. Since
rej)roduction during 1964 was extremely poor, the invasion rate re-
mained low through the summer. In 1965 Uie low density in the
surrounding population (resulting from the 1964 low reproduction)
resulted in low invasion rates in June and July but the rate increased
greatly in August and September as the numerous young increased
the density of the surrounding population.

Acknowledgements

This study, No. COO- 1336-2, was supported in y)art by U. S.
Atomic Energy grant (AT(11-1)1336 to Brigham Young University,
Provo. Utah. The preparation of this manuscript was supported in
part by NUT Training Grant No. T01-ES00089 from Division of
Envioronmental Health Sciences, Bureau of State Services, U. S.
Public Health Service. The age determinations were made possible
with the aid of Howard Kaas at the University of California. Los
Angeles, who has developed the method of tooth wear age analysis
for Perognathus. Dr. Vincent Schultz (Washington State University,
Pullman, Washington) assisted in statistical analysis of the data.

Allred, D. M., D E. Beck, and C. D. Jorgensen. 1963. Biotic communities of

the Nevada Test Site. Brigham Young Univ., Sci. Bull.. Biol. Sen II (2): 1-52.
Andrezejewski, R. 1963. Processes of incoming, settlement and disappearance

of individuals and variations in the numbers of small rodents. Acta Theri-

ologica. 7(11): 169-213.
Andrezejewski, R. and H. Wroclowek. 1962. Settling by small rodents a

terrain in which catching out had been performed. Acta Theriologica 6(9) :

257-274.
Blair, W. F. 1940. A study of prairie deer mouse populations in southern

Michigan. Amer. Midi. Nat. 24:273-305.
Calhoun, J. B. and W. L. Webb. 1953. Induced emigrations among small

manmials. Science 1 17(3040) :358-360.
Cramer, C. Y. 1956. E.xtension of multiple range tests to group means with

unequal numbers of replications. Biometrics. 12(3) : 307-310.
Duke, K. L. 1957. Reproduction in Perognathus. .1. Mammal, 38(1):207-210.
Flake, L. D. 1966. Small mammal invasion of a trapped-out habitat. Master's

thesis. Brigham Young Univ.. Provo. Utah. 35 pp. (unpublished).
French, N. R., B. D. Maza and A. P. Asciiwanden. 1967. Life spans of

Dipodomys and Perognathus in the Mojave Desert. J. Mammal. 48(4):

537-548.
Hall. E. R. 1946. Mammals of Nevada. University of California Press. Berke-
ley. 710 pp.
Speth, R. L.. C. L. Pritchett, and C. D. Jorgensen. 1968. Reproductive

activity of Perognathus parvus. J. Mammal. 49(2):336-337.
Stickel, L. F. 1 946. The source of animals moving into a depopulated area.

J. Mammal. 27(4) : 301-307.
Webb. W. L. 1965. Small mammal populations on islands. Ecology 46(4):

479-488.

LIST OF THE INSECT TYPE SPECIMENS IN THE

ENTOMOLOGICAL COLLECTION OF IHE

BRIGHAM YOUNG UNIVERSITY, PROVO, UTAH No. III.

Vasco M. Tanner' and Don R. Hanis-

In 1935 and 1936 Lists of Insect types in the Entomological Col-
lection of Brigham Young University were published.' Since then
there has been added to the University Entomological Collection the
following type sjiecimens in the order Coleoptera. Lists of type mater-
ial in other orders will shortly be published. All type specimens are
segregated from the main research insect collection. These are avail-
able to competent students of the insects.

ORDER COLEOPTERA

FAMILY CICINDELIDAE

Amblycheila utahensis Tanner.

Great Basin Nat. XL (1-2): 47, 1951.

Type; Diamond Valley, Washington Co., Utah. (Richard

Schmutz)

Cicindela parowana platti Cazier.

Southern Calif. Acad. Sci., Bull. 35:161, 1937.

1 paratype; Bentons Crossing, Mono Co., Calif.; 12 Aug. 1936.

FAMILY CARABIDAE

Elaphrus torreyensis Tanner.

Great Basin Nat. VI. (4): 137, 1941.

Paratype; Torrey, Wayne Co., Utah, June 1938, (W. W. Tanner)

Brachinus pulchellus Blatch.

The Coleop. of Indiana, Bull. 1, Indiana Dept. Geol. & N. R. :161,

1910.

1 paratype; Posey Co., Ind., 4-11-01 (W. S. Blatchley).

FAMILY AMPHIZOIDAE

Amphizoa carinata Edwards.

Wasmann Journ. Biol., 8 (3): 326. 1950.

4 para types; Near Cameron Pass, Gould, Colo; 18-20 Aug. 1941,

(Vasco M. Tanner)

FAMILY OMOPHRONIDAE

Omophron {Honiophron) tanner i Chandler.
Great Basin Nat. 2(2): 100, 1941.

'Department of Zoology and Entomologj', Brigham Young University. Provo, Utah.
-Graduate student in Entomology, Brigham Young University.

■''Utah Academy of Sciences. Arts, and Letters (Vol. XII), p. 181-19,3; Utah .Academy of
Sciences, Arts, and Letters (Vol. XIII). p. 147-152.

150

Sept. 30. 1969 list of insect types 151

Holotype and allotype; Moab, Utah 1927 (Irvin Rasmussen).
36 paraty|)es; Moab. Utah, June 1927. (James Kartchner. Anson
Call, Jr.. Vasco M. Tanner, and Irvin Rasmussen).

Omophron {Homophron) tanneri proximum Chandler.
Great Basin Nat. 2(2): 102. 1941.

Holotype and allotype; Escalante River, Garfield Co., Utah,
Mouth of Calf Creek, 2 Aug. 1939 (U. P. Chandler).

FAMILY DYTISCIDAE

Coelambus solinarius Wallis.
Can. Ent.. 56:105
1 paratype; Baldur, Man.; 29-VII-22, (J. B. Wallis).

Hydroporus transpunctatus Chandler.
Great Basin Nat., 2(2): 103-104, 1941.

Holotype; Aspen Grove, Utah Co., Utah, 1 Aug. 1938 (H. P.
Chandler) .

Agabus colrmbus Leech.
Can. Ent., 70:125. 1938.
1 paratype; Churchill, Man. VII-7-1937; (W. J. Brown).

FAMILY MELYRIDAE

Eutrichopleurus concinnus Blais.

Tr. Amer. Ent. Soc. 66:321, 1940.

1 paratype; St. George, Washington Co., Utah, V-28-35 (E. C.

Van Dyke).

FAMILY MELOIDAE

Epicauta normalis Werner
Psyche, 50:65, 1944.

6 paratypes; Fillmore, Utah (D. E. Johnson); Sheep Creek, Du-
chesne Co., Utah; June 1926 (Vasco M. Tanner); Provo En-
virons, Utah, (Truman Swallow).

Acmaeodera parkeri Cazier. ^^AMILY BUPRESTIDAE
Wasmann Coir.. 4:22, 1940.
1 paratype; Globe, Arizona, 7 Sept. 1936 (F. H. Parker).

Acmaeodera ligulata Cazier.
Wasmann Coir., 4:20, 1940.

1 paratype; Graham Mts., Arizona, 21 June 1935 (D. K. Dun-
can).

Acmaeodera duboisi Cazier.

Bull. So. Cal. Ac. Aci., 37: 138, 1938.

2 paratypes; Death Valley, Furnace Creek, Co., California. 14
April 1938.

Santa B. Co.. California, 4 July 1938 (B. E. White) {Prosopis
juliflora glandulosa) .

The Great Basin Naturalist
152 V. M. TANNER AND D. R. HARRIS Vol. XXIX, No. 3

Acmaeodera holsteni White.
Pan. Pac. Ent. 15:71. 1939.

2 paratypes; Sunset Valley, Santa B. Co., California, 4 July 1938.
(B.'E. White), {Adenostoma fasiculatur) .

Acmaeodera adenostomae Cazier.

Bull. So. Cal. Ac. Sci., 37:127, 1938.

1 para type; Pasadena, California, 22 May 1925.

FAMILY TENEBRIONIDAE

Eschatomoxys tanneri Sorenson and Stones.
Great Basin Nat., 19 (2 and 3) : 63-66, 1959.
Holotype, allotype, 8 paratypes.

Crossing of the Father, Colorado River, Kane Co.. Utah, 5-8-1959.
(E. Sorenson and R. Stones).

Craniotus blaisdelli Tanner.

Great Basin Nat. 23 (3-4) : 167-170, 1963.

Type and 3 paratypes; Inyo Mts., California, 18 April 1949,

(Owen Bryant) ; Nevada Test Site, Nye Co., Nevada

Araeoschizus airmeti Tanner.

Great Basin Nat., 6(1-4) : 125-26, 1945.

Ty]3e and 9 paratypes; Nampa, Idaho, April 1944 (LeRoy Jack

Airmet) .

Eleodes lineata Blaisdell.

Trans. Am. Ent. Soc, 64:55, 1939.

2 paratvpes; Chiricahua Mts. Arizona, 5 Aug. 1933, (Owen
Bryant)'^Lot #417.

2 paratypes; (Owen Bryant). Lot #33-448, Chiricahua Mts. Ariz.,
23 June 1933. Elevation 6,000'.

Eleodes omissa pygmaea Blaisdell.

United States Nat. Mus. Bull. 63; 77. 1909.
5 plesiotypes; San Diego, California.

Eleodes leechi Tanner.

Geat Basin Nat, 21 (3): 63. 1961.

7 paratypes; 3-Moab, Utah June 1927 (Anson Call, Jr.; 2-Cor-
tez, Colo.: 2 May 1925; 1-Blanding, Utah (W. J. Gertsch) ; 1-La
Sal, Utah June 1927, (Irvin Rasmussen).

Eleodes inyoensis Tanner.

Great Basin Nat. 21 (3): 68, 1961.

23 paratypes; Saline Valley, Inyo Co.. California, 1-11 Nov. 1959

(B. H. Banta).

Eleodes dentipes sub sp. sordida Blais.
Can. Ent. 67:30. 1935.
1 paratype; Tulare Co., California.

Sept. 30. 1969 list of insect types 153

Eleodes novoi^errucula Boddy.

Pan. Pac. Ent.. 33(4): 195. 1957.

2 paratyjios; 1 Pullman, Washington. 27 May 1907, l-Coeur d'

Alene. Idaho. June (Wickham).

Eleodes ( ^ letahlapylis ) californica Blais .
Pan. Pac. Ent., 4(4) 1965. 1929.

1 para type; Pal mSprings, Riverside Co., California, 10 Mar.
1925 (E. C. Van Dyke).

Eleodes (Holeleodes) bryanti Blais.
Tr. Amer. Ent. Soc. 63: 137, 1937.

1 paratvpe; Graham Mts.. Ariz.. 12 Aug. 1933. (Owen Bryant),
Lot #294.

8 topotypes determined by Blaisdell; collected 10-25 Sept. 1937,
at 7400' and 9700' (Owen Bryant).

Lariversius tibialis Blais.

Pan. Pac. Ent. 23:59-62, 1947.

2 paratypes; Pyramid Lake, Nevada; 24 Aug. 1941 and 1 Sept.
1940; sand dunes (Ira LaRivers).

Alaephus nevadensis Tanner.

Brigham Young Univ. Sci. Bull; Biol. Ser. 6(1): 39, 1965.

Type and 4 paratypes; Nevada Test Site, Mercury, Nye Co.,

Nevada, 25 July 1961.

FAMILY SCARABAEIDAE

Scerica pruinosa Saylor.

J. Ent. &Zool. 27:2, 1935.

1 para type; July 1933; Colinga, Kings Co., California.

Diplotaxis brevicornis Cazier.

Great Basin Nat. 1:136-37, 1940.

1 paratype; Pyramid Peak, Dona Ana Co., New Mexico, 12 Aug.

1930 (F. R. Fesberg).

Diplotaxis impressijrons Cazier.
Great Basin Nat. 1 : 127. 1940.

1 paratype; Cedarville. Modoc Co., Calif., 20 May 1939 (P. C.
Ting, M. A. Cazier, J. A. Downes, T. Aitkens).

Coenonycha tingi Cazier.

So. Cahf. Acad. Sci.. Bull. 37:126, 1937.

2 paratypes; cT and 9. Nepa, Nepa Co., CaHf., 20 March 1937.
(H. R. Leech, M. Cazier), {Adenostoma fasciculatum H. and A.).

Coenonycha ampla Cazier.

Amer. Mus. Nov. 12 39:14, 1943.

1 d paratype; Coalings, Calif.. 20 Mar. 1940 (K. S. Hagen).

Coenonycha testacea Cazier.

So. Calif. Acad. Sci.. Bull. 36:127. 1937.

The Great Basin Naturalist
154 V. M. TANNER AND D. R. HARRIS Vol. XXIX. No. 3

2 paratv[)es: cf and $ , Clear Creek. Cuyama Canyon. Calif.,

7 Mar. 1937 (E. Ross. H. B. Leech, M. Cazier), (Erigonum).

Coenonycha hageni Cazier.

Anier. Mus. Nov. 1239:15. 1943.

2 paratypes; ? . seven miles West Coalinga, Fresno Co.. Calif.
20 Mar. 1940 (R. G. Dahl) {Erigonum fasciculatum Benth).
cf. Coalings, California. 14 May 1938 (M. Cazier), {Erigonum
fasciculatum Benth).

Coenonyclm barri Cazier.

Amer. Mus. Nov. 1239:19. 1943.

1 paratype; McKittrick, Calif., 21 Mar. 1940 (K. S. Hagen).

Phobetus palpalis Saylor.
J. Ent.&Zool. 28:1,1936.

1 paratype; Indio, Calif., 24 Mar. 1935 (F. R. Piatt).

FAMILY CHRYSOMELIDAE

Exema byersi Karren.

Univ.' Kansas Sci. Bull., 46:670-72.

2 paratypes: eight miles North Lawrence, Kansas, 8 July 1965,
(J. B. Karren) (taken on Gutierrez dracunculoides) .

Exema mormona Karren.

Univ. Kansas Sci. Bull.. 46:659-672.

3 paratypes; One Farr West, Utah (C. J. D. Brown), 2 Hobble
Creek Canyon, Uinta National Forest. Utah, 16 Aug. 1961, (S.L.
Wood. J. B. Karren) {Gutierreza sarothae) .

FAMILY ANTHRIBIDAE-PLATYSTOMIDAE

Brachytarsus annulatus Carr.
Can. Ent., 60:279. 1930.
1 paratype; Medicine Hat. Alberta. Canada, 1930 (F. S. Carr).

Brachytarsus beyeri Schaeffer.

Trans Amer. Ent. Soc. 32:277, 1906.

8 cot^^es; California.

FAMILY CURCULIONIDAE

Hyperodes inter punctatulus Dietz.

Amer. Ent. Soc. Trans. 16:36, 1881.

1 paratype; Cypress Mills. Texas (Schaupp).

Listronotus leechi Sleeper.

Pan. Pac. Ent. Vol. 31 (3): 156, 1955.

2 paratypes: Rawiings Lake. Lumby B.C. 3 Oct. 1946, (Hugh B.
Leech) .

Lepyrus oregonus tesselatus Van Dyke.
Pan. Pac. Ent.. 5:56, 1928.
1 paratype; Banff. Alta.. 13 June 1928 (Owen Bryant).

Sept. 30. 1969 list of insect types 155

Pantomorus peregrinus Buchanan.

U.S. Dept Agric. Misc. Pub. #341:14. 1939.

4 paratypps: 3-Gulfport, Mississippi. 3 Sept. 1937. (Gladney and

Padgett); 1 -Saucier, Mississippi, 3 Sept. 1937 (Dopson, Padgett,

Baker).

Pantomorus planitiatus Buchanan.

U.S. Dept. Agri. Misc. Pub. #341:36. 1939.

2 paratypes; Pueblo. Colo., (H. Soltau Collection).

Orchestes paUicornis var. pallidior Leng.

Rhynchophora of North ELastern America, Indianapolis, p. 281,

1916.

Typ>e and 4 paratypes; Nfld., July.

Pnndcleteius bryanti Tanner.
Great Basin Nat., 14: 76, 1954.

Holotype, allotype, 3 paratypes; Davis Mts., Texas, 9 May 1941.
Owen Bryant, 6,700'.

Dorytomus rubidus Tanner.

Great Basin Nat.. 1(1):23. 1939.

Type; St. George, Utah, 20 April 1934 (V. M. Tanner).

Lupinocolus blaisdelli Van Dyke.
Pan. Pac. Ent. 12:82. 1936.
1 paratype; Hope Valley, Eldorado Co.. California, July.

Endalus laticollis Blatchley.

Rhynchophora of N. E. America, Indianapolis. 1916.

1 cotype; Dunedin, Florida, 15 March 1913 (W .S. Blatchley).

3 topotypes from Leng Collection.

Endalus depressus Burke.

The Southwestern Nat. 6 (2): 65-72. 1961.

2 paratypes; Anderson Co., Texas, 31 Aug. 1958 (H. R. Burke).

TAxellus haldemani Burke.

Southwestern Nat., 8:167-168. fig. 6. 1963.

2 paratypes; College Station, Brazos Co., Texas, 5 and 13 May

1962 (H. R. Burke).

Onychylis essigi Tanner.

(Jreat Basin Nat.. 14:77-78. 1954.

4 paratyj)es: Saratoga Springs, Death Valley. California, 20
March 1944 and 23 April 1944, (J.N. Belkin).

Onychylis texanus Burke.

Coleop. Bull. 13:36-37, 1959.

2 paratypes; Anderson Co.. Texas. 31 Aug. 1958 (H. R. Burke).

Pscudoeucyllus boulderenis Tanner.

Great Basin Nat. 10:71-72, fig. 1. 1950.

Holotype; Boulder Dam, Arizona (V. M. Tanner).

The Great Basin Naturalist
156 V. M. TANNER AND D. R. HARRIS Vol. XXIX. No. 3

Eugnaniptus nigriventris Schaeffer.
Bull. Br. Inst. I. pp. 141-179. 1905.

1 cotype; Huach. Mts., Arizona, Aug.

Eugnamptus pallidas Schaeffer.

Journ. N.Y. Ent. Soc. XVI. p. 213, 1908.

2 cotypes: Huach. Mts.. Arizona.

Eupagoderes hryanti Van Dyke.
Pan Pac. Ent.. 27:31. 1951.

4 paratypes; Baboquivari Mts., Arizona. 20-24 Sept. 1933, (Owen
Bryant).

Eupagoderes simulans Van Dyke.

Pan. Pac. Ent., 10(4) : 176-177, 1934.

1 paratype; Allamore, Texas, 2 July 1930 (J. 0. Martin).

Eupagoderes californicus Ting.

Bull. So. Calif. Acad. Sci., 38:81-83. 1939.

1 paratype: Stove Pipe Wells, Death Valley, California, 13 April
1938 (J. J. DuBois).

Eupagoderes hardyi Tanner.

Great Basin Nat., 1:31-32, 1939.

Type: North Fork Provo Canyon, (D. Ehno Hardy) .

Eupagoderes utahensis Tanner.
Great Basin Nat. 1:31, 1939.
Type: St. George, Utah, (Vasco M. Tanner).

Tosastes columhianus Van Dyke.
Pan. Pac. Ent., 27:32, 1951.

2 paratypes; 1-Kamloops. B.C. 1 May 1932, (Owen Bryant).

Cimbocera buchanani Ting.

So. Cahf. Acad. Sci. Bull.,39:134, 1940.

1 paratype: San Marcial, Socorao Co.. New Mexico, 22 June

1921 (C. D. Duncan).

Cimbocera peter soni Tanner.

Great Basin Nat.. 2:29-30. figs. 1. 2, 1941.

9 holotype, d allotype, $ 11 paratypes, 3 cf paratypes. Virgin
River. Utah. (Weidt) ; 6 9 paratypes, St. George, Wash. Co.,
April, 1921. (Vasco M. Tanner); 1 paratype; Washington.
Wash. Co., Utah (Vasco M. Tanner) ; 1 9 paratype; Indianola.
Sanpete Co., April 1921 (Vasco M. Tanner).

Paracimbocera atra Van Dyke.
Pan. Pac. Ent, 14:2, 1938.

1 Topotype: Mt. Charleston, Less Canyon, Clark Co., Nevada, 24
May 1960 (W. Reeves, M. Cazier, P.C. Ting), determined by P.
Ting 1941.

Paracimbocera artemisiae Ting.

Bull. So. Calif. Acad. Sci., 39:39, 1940.

Sept. 30. 1969 list of insect types 157

1 paratype; Esmeralda Co., Nevada, 15 Oct. 1907 (F. W. Nunen-
macher).

Miloderes argenteus Van Dyke.
Pan. Pac. Ent.. 11:4. 1935.

1 d" paratvpe; Paonia. Delta Co., Colo.. 18 .June 1926 (E. C.
Van Dyke^

Miloderoides vandykei Tanner.
Great Basin Nat., 3:23-24, 1942.
Holotype, allotype, and 3 paratypes; Virgin River, Utah (Weidt).

2 paraty|)es; St. George, Utah (A. M. Woodbury).

Miloderoides niaculatus Van Dyke.
Pan. Pac. Ent. 12:72. 1936.
1 paratype; Parma. Idaho, 28 April 1932.

Cryptolepidus leechi Ting.

Bull. So. Calif. Acad. Sci., 39:147, 1940.

1 paratype; Baker, California (P. C. Ting, M. Cazier, H. Leech).

(Atriplex sp.).

Cryptolepidus rugicollis Ting.

Bull. Calif. Acad. Sci., 39:149, 1940.

1 9 paratype; 3 miles north of Lovelock, Nevada, 17 May 1938

{Atriplex sp.).

Cimbocera cazieri Van Dyke.
Pan. Pac. Ent. 12:73. 1936.

1 paratype, Baker, California. 27 March 1935 (M. Cazier).

Cryptolepidus aridus Tanner.

Brigham Young Univ. Sci. Bull., Biol. Ser. 8(2): 11-12. Figs. 5,
6, 7, 1966.

3 paratypes; 4 topotypes; Nevada Test Site, Nye Co.. Nev.

Miloderes mercuryensis Tanner.

Brigham Young Univ. Sci. Bull., Biol. Ser. 8(2): 16, 1966.

2 paratypes; Nevada Test Site. Nye Co., Nev.

Crocidema arizonica Van Dyke.
Pan. Pac. Ent, 27:33. 1951.

1 paratype; Santa Catalina Mts.. Arizona, 15 June 1933 (Owen
Bryant) .

Dyslobus alepidotus Ting.

So. Cahf. Acad. Sci. Bull. 36: 79, 1937.

2 paratypes, 2 topotypes; Sigmond Stern (jrove. San Francisco,
Cahf.

Dyslobus washatchensis Tanner.

Proc. Utah Acad. Sci., Arts, and Letters. 15:147-148. Figs. 1. 2.
3. 1938.

Holotype, alloty]ie, and 50 paratypes. Aspen Grove. Mt. Tim-
panogos. Utah Co.. Utah, July 1936 (Vasco M. Tanner).

The Great Basin Naturalist
158 V. M. TANNER AND D. R. HARRIS Vol. XXIX, No. 3

Omias albus Van Dyke.

Pan. Pac. Ent., 11:96, 1935.

2 paratypes; Lind, Washington, 3 June 1921 (M. C. Lane).

Cercopeus bolli Burke.

Southwestern Nat., 8:164-166, Fig. 5, 1963.

1 para type; 10 miles southwest of Elkhard, Texas. 15 June, 1961

(H. R. Burke).

Nemocestes longulus Van Dyke.
Pan. Pac. Ent, 12:26, 1936.

1 paratype; Saticoy, Ventura Co., California, June 1924 (S.
Flanders).

Nemocestes tuberculatus Van Dyke.
Pan. Pac. Ent. 12:28, 1936.
1 paratype; Muir Woods, Marin Co., California, 30 July 1908.

Nemocestes sordidus Van Dyke.
Pan. Pac. Ent., 12:36-27. 1936.
1 paratype; San Jose, California, 1 Aug. 1931.

Bagous lengi Tanner.

Great Basin Nat., 4: 13, fig. 3, 1943.

Type; 3 paratypes; Iowa 207; 2 paratypes; Lake Okoboji, Iowa.

14 July 1917 (L. L. Buchanan).

Bagous longirostrus Tanner.

Great Basin Nat., 4: 18, fig. 5, 1943.

1 piaratype; Buffalo, New York; 1 paratype; Michigan.

Bagous texanus Tanner

Great Basin Nat.. 4:20, fig. 6, 1943.

4 paratypes; Victoria, Texas (J. D. Mitchell) (Cyporus virens).

Bagous maculatus Blatchley.

Rhynch. of N. E. America. Indianapolis, 1916.

1 paratype; Dunedin, Florida, 19 March 1917 (W. S. Blatchley),

p. 232.

Bagous chandler! Tanner

Great Basin Nat. 4:26, fig. 8, 1943.

Type and 1 paratype; east side Utah Lake, Utah, 14 June 1941.

4,000' elevation (H. P. Chandler).

3 paratypes; Bear River. Utah.

Bagous sellatus LeConte.

Rhynch. of Amer., Proc. Am. Phil. Soc. SV, 1876.
1 homotype; St. Augustine, Florida, Liebeck Collection, Com-
pared by F. G. Werner, 1942.

Bagous obliquus LeConte.

Rhynch. of Amer.. Proc. Am. Phil. Soc. XV. 1876.

1 homotype; Enterprise, Florida, May 25, Museum Comparative

Zoology. Compared by F. G. Werner, 1942.

Sept. 30. 1969 list of insect types 159

Bagous americanus LeConte.

LeConte and Horn. Rhynch. 1876. p. 185.

1 honioty])e; N. J. Museum, Coni{)arative Zoology, Compared by

F. G. Werner. 1942.

Bogous cavijrons I^Conte.

LeConte and I lorn, Rhynch. 1876, p. 186.

1 homotype; Mass.. Blatchley Collection, Compared by F. G.

Werner.

Bagous magister LeConte.

Rhynch. of Amer.. Proc. Am. Phil. Soc. XV, 1876.

1 homotype; Good Haven. Mich. (W. G. Dietz). Compared by

F. G. Werner, 1942.

Bagous transversus LeConte.

Rhvnch. of Amer. Proc. Am. Phil. Soc. XV. :188. 1876.
1 homoty])e; Penn Yan, N. Y., July 19, 1925 (Babry). Com-
pared by F. G. Werner. 1942.

Bagous tingi Tanner

Great Basin Nat., 4:27, fig. 9, 1943.

Type, 1 paratype; Lake Pilarcitos, San Mateo, California. 27

Aug. 1929 (P. C. Ting, M. Cazier).

Bagous restrictus LeConte.

Rhynch. of Amer., Proc. Am. Phil. Soc. XV, 1876.

1 homotype. San Tomas. Brownsville, Texas. Vn-29 from Schaef-

fer Collection. Compared by F. G. Werner, 1942.

Bagous californicus LeConte.

Rhynch. of .\mer. Proc. Am. Phil. Soc. XV, :187, 1876.

1 homotype; Los Angeles Co., California. Compared by F. G.

Werner, 1942.

Bagous puritanus Blatchley.

Rhyn. of N. E. America, Indianapolis, 1916.

1 homotype, Drac. Mass. Blatchley Collection, compared by F. G.

Werner, 1942.

Bagous puslllus LeConte.

Rhynch. of Amer., Proc. Am. Phil. Soc. XV, 1876.

1 homotype. Mt. Lee. written with pen and ink, difficult to read.

Compared by F. G. Werner, 1942.

Bagous floridanus Tanner.

Great Basin Nat., 4:30, 1943.

3 paratypes; 2 - Enterprise, Florida. Liebeck Collection.

1 - Ithaca, New York, 3 June 1925 (M. D. Leonard).

Bagous dietzi Tanner.

Great Basin Nat., 14:74, 1954.

Type; Cyprus Mills. Texas (Schaupp).

1 paratype; Texas. Frederick (Blandnard Collection).

The Great Basin Naturalist
160 V. M. TANNER AND D. R. HARRIS Vol. XXIX, No. 3

Bagous angustus Tanner.

Great Basin Nat., 14:73, 1954.

Type; Newman Lake, Washington, 9 July 1927 (M. C. Lane).

Pnigodes tuberosus Tanner.

Great Basin Nat., 4:35-36. fig. 12, 1943.
Type; Iowa.

Pnigodes buchanani Tanner.

Great Basin Nat., 4:33-34. fig. 10. 1943.

5 paratypes; Wolfe Canyon, Texas, various dates.

1 paratype; Mexico, Texas, 17 May 1907.

1 paratype; Texas.

Tychius albidus Schaeffer.

Jour. N. Y. Ent. Soc. 16:219, 1908.

1 cotype; Death Valley. April 91 K on Prosopis julifiora.

Tychius suturalis Schaeffer.

Jour. N. Y. Ent. Soc. 16:218, 1908.

1 cotype; Nogales, Santa Cruz Co., Arizona (F. W. Nenen-

macher) .

Dinocleus bryanti Van Dyke.
Pan. Pac. Ent., 29:10f, 1953.

1 paratype; Selngman, Arizona. 3 August 1936 (Bryant).

Lixus perstriatus Chittnenden.

Proc. U. S. Nat. Mus. 77 (Art. 18): 5.

2 paratypes; Cortez, Colorado. 19 June 1925.

Cylindrocopturus eatoni Buchanan.

Proc. Ent. Soc. Wash., 43:180-181, 1940.

4 paratypes; 2cr and 2$; Big Springs. Lassen Nat'l Forest,
California, 2 May 1939 (C. B. Eaton) (ex. stems Ponderosa and
Jefferi Pine).

Cylindrocopturus furnissi Buchanan.
Proc. Ent. Soc. Wash., 42: 1 78, 1940.
4 paratypes; 2^" and 2 9 ; LaGrand, Washington (R. L. Furniss).

Eucactophagus weissi Barber.

Proc. Ent. Soc. Wash., 19:21, pi. 4, figs. 4, 4A, 4B, 1917.
1 paratype; Summit, New Jersey, from Chas. W. Leng Collec-
tion.

Eugnamptus nigriventris Schaeffer.
Bull. Br. Inst. 1:178. 1905.

1 cotype; Huachuca Mts.. Arizona, VII-II.

Eugnamptus pallidus Schaeffer.

Journ. N. Y. Ent. Soc. XVI, 1908:213.

2 paratypes; Husch. Mts., Ariz.

Kietana gressitti Tanner

Brigham Young Univ. Sci. Bull. Biol. Ser. 1969. 10(3) :8-9.

Sept. 30, 1969 list of insect types 161

1 paratype; Solomon Islands; Guadalcanal, (jold Ridge, March
21, 1955 (E. S. Brown).

Moluccohius marshalli Tanner

Creat Basin Nat. XX. 1960:23-26.

lype and 105 para types; U. S. F*itu Military Airfield, located
between Pitoeo and Sabatai, Morotai Island; Oct. and Dec. 1944
(Ernest Reimschiissel).

Atactophysis cor data Illlr.

Verb. Naturf. Ges. Basel XLV, 1934, 9.

1 honiotype; Solomon Islands, Bougainville, Is. Kokure. nr.
Crown Prince Ra. 900 M. June 10, 1956 (J. L. Gressitt).

Platyacus subalatus Hllr.

Wien. Ent. Zeit XXIX, 1910, p. 194.

2 homotypes; Solomon Islands, Bougainville, Is. Kokure, nr.
Crown Prince Ra. 900 M. June 10. 1956 (J. L. Gressitt).

Platyacus nigrocristatus Illlr.

Verb. Naturf. Ver. Basel XLV, 1934, p. 22.

2 bomo types; Solomon Islands, Bougainville, Simba Mission,

June 29, 1955 (J. L. Gressitt).

Platyacus websteri Hllr.

Stett. Ent. Zeit. LVIII, 1897, p. 270.

2 homotypes; Solomon Islands. Buka Is., Gagan. 40 m. June 15,

1956 (J. L. Gressitt).

Platyacus decoratus Fst.

Stett. Ent. Zeit LVIII, 1897, p. 272.

1 bomo type; Solomon Islands, New Georgia Group, N-Georgia,
Is. Munda, 1-30 m (J. L. Gressitt).

Trigonops minuta Tanner

Brigham Young Univ. Sci. Bull. Biol. Sci., 1969, 10(3):26-27.

2 paratypes; 1 - Cboiseul Is., Luti, Sept. 8:1, 1958 (F. G. Fene-
more); 1 - Ysabel Is., March, 1932 (R. A. Lever).

Trigonops marshalli Tanner

Brigham Young Univ. Sci. Bull. Biol. Ser., 1969, 10(3): 30-31.
Holotyjie and three paratypes; Santa Isabel Is., X-19-1960 (J.
Tubua).

Trigonops paravicinii Hllr.

Verb. Naturf. Ges. Basel XLX., 1934, p. 16.

1 homotype; Solomon Islands, New Georgia Island.

Trigonops planicollis Hllr.

Verb. Naturf. Ges. Basel XLX, 1934. p. 16.

1 homotype; Solomon Islands, Western Group, Kolombangara,

Oct. 2, 1954 (E. S. Brown).

Trigonops exophthalnius Hllr.

Wien. Ent. Zeit XXIX, 1910. p. 187.

The Great Basin Naturalist
162 V. M. TANNER AND D. R. HARRIS Vol. XXIX, No. 3

2 homotypes; Solomon Islands, Malaital (R. A. Lever and P. G.
Fenemore).

Trigonops platessa Hllr.

Verh. Naturf. Ges. Basel XLV, 1934, p. 19.

6 homotypes; Solomon Islands, Guadalcanal Isl. 1944-45 (D E.

Beck).

Trigonops granulosa Tanner

Brigham Young Univ. Sci. Bull. Biol. Ser., 1969, 10(3):35-36.
Holotype, allotype, and 4 para types; Solomon Islands, Florida
Island, April 1944 (H. P. Chandler).

Trigonops isabellae Tanner

Brigham Young Univ. Sci. Bull. Biol. Ser., 1969, 10(3):36-37.
8 para types; Solomon Islands, Guadalcanal Island, 194-4-45
(D E. Beck).

Trigonops carinithorax Hllr.

Verh. Naturf. Ges. Basel SLV, 1934, p. 20.

2 homotypes; Solomon Islands, Bougainville Is., Kieta, Oct. 10,

1927 (J. L. Fraggatt).

Trigonops bougainvillensis Tanner

Brigham Young Univ. Sci. Bull. Biol. Ser., 1969, 10(3):38-39.

5 paratypes; Solomon Islands, Bougainville Is., June 4, Boha
(J. L. Gressitt) ; Kohura, 690 M. June 8-18, 1956 (E. J. Ford, Jr.).

Trigonops guadalcanalensis Tanner

Brigham Young Univ. Sci. Bull., Biol. Ser., 1969, 10(3):39-40.
Holotype, allotype, and 18 paratypes; Solomon Island, Guadal-
canal Is., 1944-45 (D E. Beck and Ernest Reimschiissel).

Trigonops helleri Tanner

Brigham Young Univ. Sci. Bull., Biol. Ser., 1969, 10(3):40-41.

6 paratypes; Solomon Islands, New Georgia Group, N. Georgia
Island, July 15, 1959 (J. L. Gressitt).

Trigonops dilaticollis Gunther

Mitt. Deut. Ent. Ges., 8:3, 1937.

2 homotypes; Solomon Islands, S, Malaita Is., May 5-7, 1934
E. H. I^ver).

Trigonops notaticollis Hllr.

Wien. Ent. Zeit XXIX, 1910, p. 190.

3 homotypes; Solomon Islands, Sikaiana, March 23, 1936 (R. A.
Lever) .

Trigonops forticornis Hllr.

Wien Ent. Zeit XXIX, 1910, p. 187.

2 homotypes; Solomon Islands, Guadalcanal Is., Tenaru R.,

1945 (F. E. Bohart).

Trigonops gressitti Tanner

Brigham Young Univ. Sci. Bull., Biol. Ser., 1969, 10(3):44-45.

Sept. 30. 1969 list of insect types 163

4 paratypes; Solomon Islands, Bougainville Island. Boku and
Kohure, June 10, 1956 (J. L. Gressitt and E. J. Ford, Jr.)

Trigonops gloriosa Tanner

Brigham Young Univ. Sci. Bull. Biol. Ser., 1969, 10(3):45-46.

1 paratype; Solomon Islands, Bougainville Island. Kokure. nr.
Crown Prince Ra. 900 M., June 9, 1956 (J. L. Gressitt).

Lophothetes reimschiisseli Tanner.

Great Basin Nat. XX (1-2), 1960; 26-28.

Type and 39 paratypes; Admiralty Islands, Manus, Sept. 1944

(Ernest Reimschiissel ) .

Rhyncogonus debilis Van Dyke

Bernice P. Bishop Museum, Occasional Papers, XIII (11): 119,
1937.

2 paratypes; Rurutu Island. August 28, 1934, Austral Islands,
Mt. Manurova, 1,200 feet, (sweeping grasses and low herbage)
(E. C. Zimmerman).

Rhyncogonus griseus Van Dyke

Bernice P. Bishop Museum, Bull. 98:44; Marquesan Insects No.

2 homotypes; Nukuhiva, Marquesas Islands, September 20, 1920
(E. W. Quayle). Compared by Vasco M. Tanner.

Rhyncogonus excavatus Van Dyke

Bernice P. Bishop Museum, Occasional Papers, XIII (11): 116,
1937.

3 paratypes; Rurutu Island, September 2, 1934, 1000 feet elev.,
South Slope Mt. Teape, Austral Islands (Beating. Dryopleris) ;
(E. C. Zimmerman).

1 paratype; Rurutu Island, August 28, 1934, 1000 feet elev..
South Slope Mt. Manuseva, Austral Islands, on Piper (E. C.
Zimmerman) .

Rhyncogonus nigerrimus Van Dyke

Bernice P. Bishop Museum, Occasional Papers, XIII (11): 119,
1937.

1 paratype; Raivavae Island. Austral Islands (E. C. Zimmer-
man).

1 paratype; South Slope Mt. Muanui (beating shrubs), August
6, 1934.

1 paratype; Mt. Miro, 1,000-1.300 feet.

Rhyncoponus nodosus Van Dyke

Bernice P. Bishop Museum, Occasional Papers, XIII (11): 113,
1937.

2 paratypes; 1 9 Tubuai Island. August 20, 1934, S. W. Ridge,
Mt. Taita, 1,100 feet. Austral Islands (D. Anderson). 1 Tubuai
Island, August 21. 1934, Mt. Taita, 1,200 feet. Austral Islands
(beating) (E. C. Zimmerman).

The Great Basin Naturalist
1(34 V. M. TANNER AND D. R. HARRIS Vol. XXIX, No. 3

Holcolaceus viridulus Mshl.

Ann. Mag. Nat. Hist. (12), 6, p. 324.

5 cotypes, 2 d" , 3 9; South Africa, Natal, nr. 1 larding, Weza
plantations, March 1951, Wattle Res. Inst. (Damaging seedlings
of Pinus patula and Cryptomeri japonica) .

Orochlesis conspersa Zimmerman

Proc. Hawaiian Ento. Soc. 10(1): 165.
1 paratype; Solomon Is. (W. M. Mann).

Antliarrhianus verdcourti Mshl.

Ann. Mag. Nat. Hist. (12)8:300 figs.

8 cotypes, ^dd-, 5? 5; Kenya, Matthews Range. June 1954

(B. Verdicourt) on 9 cone of Encephalartos.

Blosyrus mazambicus Mshl.

Mem. Estud. Mus. Zool. Univ. Cambra (2) LXXXI, p. 3, 1935.
4 cotypes; Port E. Africa, Mozambique Is., 6, IX-1929 (Sir
G. A. K. Marshall).

Nematocerus perditor Mshl.

Ann. Mag. Nat. Hist. (12), 4, p. 839.

12 paratypes, Kenya, Eldoret, April, 1950 (R. H. Lee Pelley).

STUDIES ON RAPTOR MORTALII Y IN WESTERN IH AH'

David H. Ellis, Dwight G. Smith, and Joseph R. Murphy-

In past studies of predatory bird destruction. Hickey (1949)
found that 102 of his banded juvenile marsh hawks were shot during
their first year of life, while more recently Sprunt (1963) re{)orted
that 91 of the 118 Bald Eagles known to have been killed in 1962
had been shot. Although S]X)fford (1964) and others have called
attention to the high raptor mortality caused by illegal predator con-
trol programs, and Imler and Kalmbach (1955) have reviewed Bald
Eagle destruction in the years of the lx)unty laws, the extent of mor-
tality caused by indiscriminate shooting is unknown.

In this study we attempted to locate all raptors killed during a
two year period in Cedar Valley. Utah County, Utah. The study
was centered on a 12.1 mile stretch of gravel road paralleled by
single cross bar utility poles which were frequently utilized as
perches by predatory birds in the valley. The study area was located
within an intermontane valley comprising some 170 square miles,
and averaging 4,900 feet in elevation; dominant plants were big
sagebrush (Artemesia tridentata) and shadscale {Atriplex conferti-
folia). The land is used for dry farming and sheep range, and sup-
ports two small settlements. The valley is also a favorite rabbit
hunting locality and receives large concentrations of hunters, par-
ticularly on winter weekends.

Procedure

The initial survey of the study road took place on 29 April 1967.
and it was then subsequently surveyed at intervals of approximately
one to three months, until the fall of 1968 when the pole line was
dismantled. For sake of comparison, surveys were also made of the
same pole line following its divergence from the study road. Many
additional miles of roads and cliffs were also surveyed in an attempt
to locate dead birds.

Results and Discussion

Thirty dead raptors were found on the initial road survey; this
total included 14 Golden Eagles, 14 Buteo hawks, 1 Sparrow Hawk,
and 1 Great Horned Owl. By the end of the study period, a total of
38 dead raptors had been found along this road and an additional
ten in the remainder of Cedar Valley (see Table 1). Density of kills
on the study road averaged 3.14 birds per mile, and included an
extreme situation in which four dead Golden F^agles were found
under two adjacent poles. Those birds suffering the heaviest mor-

'This study was supported in part liy a prant from tlie National Audubon Society. Gratitude
is expressed to ihem, and also to the Department of Zoology and Entoniology, Brighani Young
University for transportation and facilities.

-department of Zoolopy and Kntomology, Brigliam Young ITniversity.

165

The Great Basin Naturalist
166 ELLIS, SMITH, AND MURPHY Vol. XXIX, No. 3

Table 1

Dead Raptors Found During Study Period

Cedar Valley, Utah

Golden Elagle {Aquila chrysaetos) 26

Bald Eagle {Haliaeetus leucocephalus) 2

Rough-legged Hawk (Buteo lagopus) 8

Ferruginous Hawk {Buteo regalis) 1

Red-tailed Hawk {Buteo jamaicensis) 2

Swainson's Hawk {Buteo swainsoni) 1

Buteo* 5

Cooper's Hawk {Accipiter cooperii) 1

Sparrow Hawk {Falco sparverius) 1

Great Horned Owl {Bubo virginianus) 1

Total 48

'Not identifiable to species owing to decomposition.

tality included the eagles (26 Golden Eagles and two Bald Eagles)
and the Buteos (seventeen dead birds of four different species).
During this same period only one dead raptor was found along the
segment of the pole line not paralleled by the road.

The approximate times of death were estimated from the degree
of decomposition at the time of discovery and the rate of decomposi-
tion observed in birds found shortly following death. Four time
periods were assigned from the study road data. It was estimated
that nineteen raptors, including ten Buteos and nine Golden Eagles,
had been killed before the winter of 1966-67. During the winter and
spring of 1966-67, fourteen raptors, including seven Golden Eagles
and five Buteos, were killed. Two more Golden Eagles were killed
during the summer of 1967, and an additional three during the
winter of 1967-68. The heaviest mortality occurred in the fall and
winter period, which also coincides with the periods of heaviest sport
hunting. Two final road surveys were taken after the dismantling
of the pole line in the fall of 1968, but no recently dead raptors were
found.

Twenty-three of the dead eagles were suitable for age analysis.
Fifteen were immature Golden Eagles, six were adult Golden Eagles,
and two were adult Bald Eagles. Both Hickey (1949) and Craighead
and Craighead (1956) similarly found that the highest raptor mor-
tality occurs among the iuveniles. and attribute this to the inex-
perience of these birds. We found that when driving along the study
road we could approach to within fifteen meters of juvenile eagles
perched on the pole, while the adult birds rarely displayed such
tolerance of human presence.

At least two known hunting methods were used to destroy the
birds. During the day some hunters were observed to drive back and
forth along the road for the express purpose of obtaining suitable
raptor targets, while at night spotlights were employed to locate
roosting birds. Evidently the large mmibers or raptors present on

Sept. 30. 1969 raptor mortality 167

certain winter days attract considerable attention from the s|)orts-
men. For example, on the morning of 21 December 1967 we found
eighteen Rough-legged Hawks, five Golden Eagles, two Ravens, three
Marsh Hawks and one Prairie Falcon perched on poles along this
twelve-mile segment of road.

An im]X)rtant aspect of this study concerns the pn)bable causes
of death of these predators. Of the 48 dead birds from the study
area, only two were found sufficiently close to the road to indicate
the possibility of their having been killed by vehicles. Poisoning was
also considered as a possible cause of death, since several 1080 bait
stations were located in Cedar Valley. Data furnished by U. S. Fish
and Wildlife Service personnel would tend to indicate that high
tolerance levels of predatory birds for this poison might eliminate it
as a factor; we feel that such evidence is inconclusive, however,
since no actual analyses of carcasses for 1080 or pesticide residues
were made. Such analyses should definitely be a part of future
studies of raptor mortality, in our area as well as elsewhere. In most
cases, there were clear indications that the birds had been deliberate-
ly shot. For example, a Ferruginous Hawk nested immediately next
to the study road and w^as photographed 8 April 1967 (Weston and
Ellis, 1968). On 19 April 1967 it was found dead v\dth holes in the
head and body, and its legs and tail had been removed. Bodies of
the raptors were almost always found intact and seldom scattered
by scavengers. These remains often exhibited shot holes in the
feathers and bones. The preponderance of the evidence available
appears to support the hypothesis that shooting was the cause of
death for a maiority of the raptors found.

In our opinion, this brief study highlights a major problem re-
lating to the future survival of raptorial birds; it also demonstrates
gross disregard of present federal and state laws protecting raptors
on the part of a segment of the public. Additionally, it emphasizes
the present danger of indiscriminate shooting to high concentrations
of raptors, and the possible extent of such mortality in relatively
small areas.

Literature Cited

Craighead, J. J., and F. C. Craighead, Jr. 1956. Hawks- Owls and Wildlife.
The Stackpole Co.. Hanisburg. Pa.

HiCKEY, J. J. 1949. Survival studies of banded birds. Unpublished Ph.D. Dis-
sertation. Univ. of Michigan. Ann Arbor.

Imler, Ralph H.. and E. R. K-IiI-mbach. 1955. The Bald Eagle and its eco-
nomic status. U. S. Fish and Wildlife Serv. Circ. 30.

Spofford. Walter R. 1964. The Golden Eagle in the Trans-Pecos and Ed-
wards Plateau of Texas. Nat. Aud. Soc. Cons. Report No. 1.

Sprunt. Alexander, IV. 1963. Bald eagles aren't producing enough vouiig.
Aud. Mag. 65:32-35.

Weston, J. B.. and D. H. Ellis. 1968. Ground nesting of the Ferruginous
Hawk in west-central Utah. Great Basin Nat. 28:111.

ac "&.

The

Volume XXIX, No. 4
December 31, 1969

Mus. coMP. zooa

LIBRARY

MAR 11971

HAaVARD

.SilY;

^^-^ — .^ HARVARD

Great Badn

pubushed by
Brioham Young University

GREAT BASIN NATURALIST

Editor: Vasco M. Tanner, Department of Zoology and Entomology
Brigham Yoimg University, Provo, Utah

Associate Editor: Stephen L. Wood, Department of Xoology and
Entomology, Brigham Young University, Provo, Utah

Members of the Editorial Board:

Ferron L. Andersen (5), Zoology

Jay V. Beck (3), Bacteriology

Robert W. Gardner ( 1 ) , Animal Science

Joseph R. Murdock (4), Botany

WiLMER W. Tanner (2), Zoology, Chairman of the Board

Stanley L. Welsh ( 1 ) , Botany

Ex officio Members:

A. Lester Allen, Dean, College of Biological and Agri-
cultural Sciences

Ernest L. Olson, Chairman, University PubHcations, Uni-
versity Editor

The Great Basin Naturalist

PUBLISHKD AT PrOVO, UtAH BY

Brigham Young University
Volume XXIX December 31, 1969 No. 4

QUALITATIVE BEHAVIOR OF A PUPFISH
(CYPRINODON ATRORUS) IN DIFFERINC, ENVIRONMENTS

Murray Itzkowitz and W. L. Minckley'

Introduction

Fishes have been used extensively by ethologists. and there are
many papers that have contributed greatly to. the understanding not
only of fish behavior, but of general ethological theory as well
(Aronson, 1949; Baerends and Baerends-Van Boon, 1950; Baerends.
et al. 1955; Morris. 1958; Barlow, 1961a, et seq.; Nelson. 1964; Liley.
1966; Simpson. 1968; Miller and Hall. 1968; many others). How-
ever, since certain, behaviors are modifiable (Hess. 1962; Marler and
Hamilton, 1966). it is surprising that there are so few studies that
compare a fish's behavior in differing environments. The present
report describes qualitative differences in behaviors elicited by a
pupfish, Cyprinodon atrorus Miller, in aquaria and in natural and
semi-natural habitats.

Materials and Methods

Cyprinodon atrorus inhabits saline marshes and lakes on the floor
of the Cuatro Cienegas basin, central Coahuila. northern Mexico
(Minckley and Itzkowitz, 1967; Miller, 1968; Minckley, 1969).
Such habitats are severely variable, warming and C(X>ling rapidly and
subject to radical changes in salinities, dissolved oxygen, and so on
(Minckley and Cole, 1968). and su{)port only the most resistant
biotic elements. C. atrorus is often the only fish species j^resent at a
locality, but another cyprinodontid. Lucanin interioris Hubbs and
Miller, and two poeciliids. Gambusia longispinis Minckley and G.
marshi Minckley and Craddock. are sometimes abundant. Observa-
tions on the behavior of C. atrorus have been made at nimierous
places in nature, most intensively by Minckley and Elena T. Arnold
in summer 1968. The activities observed were similar to those
observed in a semi-natural pond (described below).

The [)upfish used for study all originated from a stock caught
in December 1963 from marshes associated with Laguna de San

'Department of Zoology, Arizona State University, Tenipe 85J8I (Ilzko\vit/"s present ndilrcss:
Department of Zoology, University of Maryland, Colleee Park J074O). Ueseairli siipiMirtod in port
by N.S.K. Grants GB-2461 and GB-(i477X, to Minrkley.

169

170

The Great Basin Naturalist
ITZKOWITZ AND MINCKLEY Vol. XXIX, No. 4

Pablo. 13 kilometers south-southeast of the village of Cuatro Ciene-
gas de Carranza. Coahuila. Mexico. Seventeen individuals were
placed in an artificial pool in March 1964, and by August a large,
stabilized population had developed. It was studied intensively be-
tween 2 February and 30 April 1966. Breeding activities of this pup-
fish in nature begin in late January, intensify through early July,
then decline to continue throughout the summer at a lower level. A
second period of intense breeding activity often occurs in early
autumn. Vhe climate in Tempe. Arizona, is similar to that in the
Cuatro Cienegas area, and the fish also behave similarly. The semi-
natural pond was isolated from disturbances in a large patio at
Arizona State University. It was cement lined, but had a several-
year accumulation of flocculent calcareous sediment. Considerable
shade and organic input occurred from an overhanging mesquite
(Prosopis) . and city water continuously passed through the system.
Maximum dimensions of the irregularly-shaped pool were 6 by 8
meters (m). and maximum depth was about 1.25 m (Fig. 1).
Fishes were maintained without supplementary feeding. Observa-
tions were from a {)oint about two meters above the water, by naked-
eye and through use of binoculars. Other vertebrates in the pond
included poeciliid fishes, Xiphophorus couchianus (Girard) and
Flexipenis vittatus (Hubbs), and a box turtle, Terrapene coahuila
Schmidt and Owens. Xiphophorus were rare, Flexipenis built a large

Fig. 1. Sketch map of the semi-natural pond at Tempe, Arizona, where
Cyprinodon atrorus from the Cuatro Cienegas basin, Coahuila, Mexico, was studied.
Contours are approximate, depths are in cm, and the hatched area is in the position
of territories that were specifically observed.

Dec. 31. 19(S9 in-.ii w ion oi imim imi 171

|)()|)uhili(iii ill the List 1)1 the stiul\' iici-iod. and the lurtlc \va^ sci rclivo
during \hc diiy.

MoroK I' ATII.HNs

A behavioral sequence su( h as courtship consists of discrete
motor elements (e.^.. muscle contractions) that occur with other
such elements to produce synchroni/.ed patterns. Categorization i'^
subjective since an observer nuist interpret the beginning and end of
a given pattern, and evaluate variations. Major interpretive problems
arise in description of numbers or intensities of components, or in
the amount of time required for their completion. For pre.sent pur-
poses, motor j)atterns making uj) broader behavioral sequences were
weighted equally, and attemj)ts were made to describe and evaluate
variations. Where jiossible. names and definitions of patterns folhnv
Barlow^ (1961a), as denoted by quotation marks.

Contacting. — "As the female moves slowly over the bottom the male stays
beside her, and her head is usually just in front of his. Often they are actually
touching."

Tilting. — The female swims slowly within 0.5 centimeters (cm) of the bottom
and tilts her body downward at an angle of 30° to 4'5°. All unpaired fins are
spread. Barlow (1961a) considered this the beginning of nipping (see next), but
in C. alrorus nipping may be omitted following a tilt.

Nipping. — "From the tilted position, the female opens her mouth, presses
against the bottom, and noiTnally takes up a mouthful of the substrate. . . .
Then the body is dropped down against the bottom. When horizontal, the female
either immediately spits out the substi^ate. or swims forward a short distance,
stops, and then expels it." We include "halting" as described by Barlow (1961a)
as a terminal component of a nip.

Sidling. — "The male swims forward and laterally against the female. The
male's dorsal fin is folded or half open. The region supporting the anal fin of the
male is thrust against the posterior line of the abdomen of tlie female. The body
of the male is tipped out of the median plane only slightly, or not at all."

S-shaping. — "Seen from above, the body of the pupfish forms a gentle 'S'
.... At this time the male and female normally lie side by side on the bottom
in parallel S-shapes. The cuiA'ature is more pronounced in the male. The head
and anal region of the male are directed toward the female, and the anal fin of
the male is extended in her direction. The dorsal fin of the male is spread, and
sometimes is bent slightly toward the female. In the female the dorsal fin is
spread maximally. Moreover, her vent is pressed against the bottom and her
caudal fin beats rapidly, but with a very small amplitude."

Wrapping. — During S-shaping the male wraps his anal fin around the anal fin
of the female.

Jerking. — "While still S-shaped the head is jerked toward the side opposite
that to which it is already directed, tiius initiating a wave of contraction which
passes down the body reversing the direction of tlie S-shape. In the female one
egg [or more] is extruded by this flexure, and the male presumably emits sperm
at this moment."

Slow Rising. — Both the male and female slowly rise from the substrate after
a wrap. The female then moves away at high speed, and this is almost invariably
the termination of spawning by the pair.

Patrolling. — "The male in his territory swims straight ahead in spurts of
about 30 to 50 cm with the dorsal, anal, and i)ectoral fins folded back .... At
the end of each spurt the male stops abruptly by tluowing the colorful pectoral
fins forward together in a quick movement that catches the eye. Then the male

The Great Basin Naturalist
172 ITZKOWITZ AND MINCKLEY Vol. XXIX, No. 4

Stands momentarily with the median fins raised, and the pectoral fins beating
alternatly and rapidly. . . . Then he swims off again in a new direction. In this
manner the male pupfish continually crisscrosses his territory."

Chasing. — An aggressive fish swims at high speed, often with its mouth open,
toward a fleeing individual. If opportunity arises an aggressor will bite or butt
a fleeing fish.

Lateral Display. — A fish spreads all its fins and presents its flank to another
fish. The two fish often simultaneously present a lateral display in a parallel
position, with bodies held head high and at an angle of 20° to 45° to the substrate.

Tailbeat. — Through use of the caudal fin a fish appears to push water against
the side of another fish. Usually a lateral display is the beginning movement of
a tailbeat; a more complete description was given by Barlow (1961a).

Following. — This pattern appears to have three basic variations that depend on
the depth and speed of movement of a female and is subdivided for convenience
as follows: 1 ) below — a female swims at slow to moderate speed with all fins
extended. 10 cm or more above the substrate, and the male is either directly
2.5 to 3.0 cm below, or at a similar depth below, but behind, with his dorsal
and anal fins half extended to completely depressed; 2) above — similar to
"following below" except that the male is above a female, usually when the
female is less than 10 cm above the substrate; and 3) in line — similar to
"following above and below" except that the male is directly behind a female
at any depth, and the female usually is swimming faster than when the other
two patterns are utilized.

Looping. — There are five definable variations of looping. In all instances a
male swims in a circle, semi-circle, or "figure-eight." 1) Below — a female in
mid-water or near-surface, and stationary, often has a male directly below and
swimming in a circle with all fins extended; 2) in front — a female may be at
any depth, but usually at least 7.5 cm above the substrate with unpaired fins
spread, and the male makes a circular loop or figure-eight directly in front or
slightly in front and below; 3) to the side — similar to "looping in front" except
that a male is to the side of the female; 4) sidle — a male assumes a sidle position
(described above) then leaves a female to loop in front, then returns (most often
on the other side) to continue a spawning sequence; and 5) zig-zag — a female
stationary or moving slowly, with fins spread, is approached from a distance of
10 to 30 cm by a male performing continuous semi-circles (a male may also
swim awa>' making such movements, and in such instances the female may
follow the male). The looping pattern, with many variations, also is used by
territorial males in display against intruders.

Nuzzling. — This describes a motor pattern in which a male touches a
stationary female with the anterior part of his body. 1) one — A male is below
a stationary female and moves upward until the head and nape come into contact
with, or slightly posterior to her branchiostegal region, then moves slowly back
to the anal fin. around the side, and forward to a typical contact; 2) two —
similar to "nuzzling one" except that a male moves directly to contact position
without moving back to the anal fin; and 3) top — a male touches his lower head
and breast to the dorsum of a female's head, then moves laterally and down to
contact.

Courtship and Spawning in the Artificial Pond and in Nature

The male of Cyprinodon atrorus is a highly aggressive, vigorous-
ly territorial animal. The bright, irridescent blue body and contrast-
ing yellow-orange fins, plus bold patrolling within a limited area.
must make him conspicuous to other fish, and these features may
also serve as a display to attract females that are willing to spawn.
A female, when prepared to spawn, moves into a male's territory
and nips the substrate. If the male is inattentive, the female may nip

Dec. 31.1969 behavior oi- i'ipfimi 173

several times, until the male approaches. Upon siphtin^ a female,
the motor patterns <iiven above were scarcely evident in nature or
in the semi-natural pond. The male swam directly toward the female
at high speed, as if sh(> were an intruder, or sometimes used a zig-
zag loop in his approach. Moreover, most males innnediately as-
sumed a contact position, with no intermediary disj)lay. This was
followed by a tilt and nip or a lunging nij) by the female. The male
sidled, both S-shaped, he wrapped, and both fish slowly rose. The
female then swam from the territory at high speed, with the male
following behind (a pursuit indistinguishable in such instances from
a charge used when chasing an intruder). In totals of 116 observed
spawns in the pond and 50 in nature that ended in a slow rise, this
repertoire was invariable in sequence (Fig. 2).

One unusual series of observations were made of a pair "trapped"
in a small area by surrounding clum})s of filamentous algae. The
male was exceedingly active, looping in front almost continuously
then quickly contacting after the female nipped. The water was less
than 1.5 cm deep, so a slow rise rarely followed a spawn. Both lay
quietly for a second or so. then the female would dart forsvard with
the male following in line. Algal mats blocked the female's rush,
and she stoi)ped after 3 to 5 cm. The male inmiediately began nuz-
zling and attempting to contact. This resulted in a nip and another
spawn 12 of 33 times in a 31 -minute period. The other times the
female fled and dove into algae, followed by an active, looping
search by the male. The female would re-appear, fanning her ex-
tended fins "nervously," and would nip or remain stationary. In the
latter case she was immediately nuzzled and contacted, and nipped

PATROLS ^^-

APPROACHES -*^

TILTS AND NIPS-
LOOPS AND/OR
CONTACTS

^^"^^ CHASES

TILTS AND NIPS

S-SHAPES AND WRAPS

SLOW RISES

Fig. 2. Courtship and spawning sequences of Cyprinodon atrorus in natural
and semi-natural habitats.

The Great Basin NaturaUst
174 ITZKOWITZ AND MINCKLEY Vol. XXIX, No. 4

and spawned or again fled to hide. No other observations of nuz-
zhng have been made under field conditions, and no such hyper-
activity has otherwise been noted in C. atrorus under natural or
semi-natural conditions.

In any stage of a reproductive sequence a male would leave a
female to challenge an intruder. In the pond between February and
late April about 50 percent of the females that nipped the substrate
in a male's territory failed to spawn, and most times this resulted
from interruption by other fishes. No quantitative data are available
from observations in nature, but disturbances of a similar nature
were often seen. While some discrimination by a non-spawning male
between homo- and heterospecific interlopers was evident, no such
recognition was obvious when a male was interrupted during a
spawning sequence. Even intruding poeciliids were aggressively
driven from the area when a defender was spawning, as were young
C. atrorus that are oftentimes ignored by defending males. Since
looping also occurred when no interlopers were present, or when a
male was returning to a female after chasing, it may also serve to
pacify a "nervous" female or to block her retreat from the spawn-
ing area.

The motor patterns of tilting, jerking, following above, and
following below were not observed in the pond or in natiu-e.

Courtship and Spawning in Aquaria

Male and female Cyprinodon atrorus were first kept isolated for
a number of days in separate. 28-liter tanks (35 by 26 by 22 cm),
then a male was introduced into a female's tank. Upon introduction,
a male always initially sank to the bottom, or moved into a position
along a wall. The female also would sink to the substate, with all fins
extended. The female initiated activity, swimming to the male and
tilting and/or nii])])ing the substrate nearby. If a male was inatten-
tive a female would "mildly" butt or bite at his body. Males some-
times responded to such aggression with tailbeat. and if this occurred
the female always retreated. When a nip occurred the male usually
swam directly to a contact position. Typical behavior consisted of a
variable period of contact, followed by a sequentially-inconsistent
assemblage of motor patterns on the part of the male that eventually
terminated in a spawn (Fig. 3). On the average, in 668 observed
spawns, a sequence of contact^ tilt and/or nip-^ S-shape^
wrap^ ierk or slow rise predominated (Fig. 4).

Females under conditions of aquarium isolation often remained
receptive to a male after the initial spawn. Females wdth high sexual
tendencies spawned immediately after the pair was placed together,
then began swimming slowly through the tank, conveniently acces-
sible to the male. This is not the case in nature since a female im-
mediately leaves the male's territory after the spawn is consum-
mated. The presence and "receptivity" of a female stimulated hyper-
active sexual behavior on the part of a male, such as nuzzling, loop-
ing, and following. These dis])lays had little if any sequential cor-

Dec:. 31, 1909

liEIIAVIOU Ol I'l Pl-I.MI

173

REPRODUCTIVE DRIVE

FOLLOW ABOVE-

FOLLOW BELOW

FOLLOW IN LINE -^

Fig. 3. Relationships of courtship and spawning motors patterns of Cyprino-
don atrorus in laboratory aquaria.

relation to one another or to other patterns (contacting, nipping.
etc.) of a more terminal spawning sequence. However. 4-1 per-
cent of the courtship patterns (whatever they were) led directly to a
second spawn. 1 he behavior, notwithstanding its variations, must
therefore have succeeded in stimulating the female to continue re-
productive activities. When a female was not initially prepared to
spawn, a similar series of random patterns was produced by a male.
However, it was rare for a male to induce an initially unresponsive
female to spawn.

The male C. atrorus. when territorial, is always receptive to a
female, and the female controls spawning by presenting herself
within a territory and {)erforming the nij)ing pattern. The female
apparently requires little stimulation by a male (excepting, perhaps,
a short period of contacting, or some visual stimulation from male
defensive activities) when she is prepared to spawn. Random use
of courtship motor patterns api)ears to function in stimulation of a
female to continue the spawning activities, rather than as an
initiator. Rigid courtship sequences require reciprocal stimulation,
and randomness of behavior of male C. atrorus may result from a
lack of cues from the female.

Territori.ality in the Semi-n.\tur.\l Pond and in Natitre

Males of Cyprinodon atrorus begin to establish territories in late
January, and remain aggressive throughout the breeding season with
the intensity of behavior varying with intensities of rei)roductive
activities in the given habitat. Males began sporadic aggressive ac-

176

The Great Basin Naturalist
ITZKOWITZ AND MINCKLEY Vol. XXIX, No. 4

FOLLOW IN LINE-

59%

S-shape

20%

Contact

11%

Nip

28% Follow below

26% Nip

16% Contact

12% Follow in line

5% Follow above

3% Nuzzle

54% Follow below

26% Follow above

9% Contact

6% Nip

3% Loop

3% S-shape

38% Contact

35% Follow below

15% Nip
5% Follow above
5% Follow in line
3% S-shape

70% S-shape
21% Contact
5% Nip
3% Tilt

FOLLOW ABOVE -

FOLLOW BELOW-

48% Jerk
45% Contact
7% Slow rise

38% Contact

32% Nip

13% Follow below

5% Follow in line

5% Loop

4% Nuzzle

28% Follow in line
23% Nuzzle
18% Follow above
18% Loop

7% Nip

7% Contact

68% Nip
19% S-shape

8% Tilt

3% Follow above

1% Follow below

1% Nuzzle

100% Contact

Fig. 4. Quantitative relations among a motor pattern and those patterns ob-
served to follow it in Cyprinodon atrorus under laboratory aquarium conditions,
based on 668 observed spawnings.

tions in February in the artificial pond. Fights were not confined to
a particular area, and usually consisted of short charges with little or
no contacting. Some fighting was associated with feeding pits during
this period, an activity continued almost throughout the year by non-
reproductive males, females, and juveniles (Minckley and Arnold,
1969). Pit-building and defense is unrelated to reproductive activities
in C. atrorus, although it may have significance in the sexual be-
haviors of some other pupfish species (E. T. Arnold, pers. comm.).
In mid-February, several males established temporary areas in
shallows of the semi-natural pond, but these were not defended for
more than a few hours. They were 15 to 25 cm in diameter in water
not more than 5 cm deep. Boundaries were ill-defined and frequent-
ly changed more than five cm in a few minutes. Border fights be-
tween neighboring fish rarely occurred since territories were widely
disj)ersed and usually isolated. Permanent territories were formed
by late mid-February, and three, well-defined, adjacent ones were
selected for intensive observations. These were situated in a cove
that sloped gently to a maximum depth of eight cm. On the pond-
ward side was an abrupt slope leading to water more than 30 cm
deep. The territories were at depths between 2 and 8 cm (Fig. 1),
and occupied all available shallow water. The largest included about
2,100 square centimeters (cm-) and the smallest was 1.050 cm- in
size. The range in estimated sizes for territories in nature is 3,200 to
875 cm- (23 estimates), with a mean of about 1,950 cm-. Shapes of

Dec. 31.1969 behavioh oi' itpfimi 177

territories raniied from rouiul in open, sparsely po|)uliitecl area>. to
elongate and highly variable where heterogeneity of habitat, or the
presence of other nudes, influenied the l)oundaries. The tiiree males
in the pond vigorously defended tlieir respective areas until inid-
Marc li. Others tlien began to form peripheral territories, and wedges
defended by new males were progressively forced into and between
the three original ones. The three defended areas were markedly
compressed to about half their original sizes, and within a week the
fish had been displaced and disappeared.

Territorial defense by (\ atrorus consists of numerous lateral
displays, infrequent tailbeats against peripheral males, and charges
against non-territorial males and sometimes females and juveniles.
Actual contacts and severe fighting occur when territorial lx)undaries
of "matched" males (in size and coloration) are closely appressed,
especially w^hen the border meets in relatively deep water (>5 cm).
Under such conditions the males present sustained lateral display,
tailbeat. and biting, the last generally in the vicinity of the anal fin
and flanks. Contact fighting, although rare, usually begins near the
surface, and the two males spin about each other as they gradually
move toward the bottom. Fights are sustained for as long as 30
seconds, and a small cloud of sediment often obscures the terminal
phases of such conflict. Tinbergen (1953). Carranza and Winn
(1954), and Miller (1964) have noted that the ferocity of attack
by a defending male of fishes they studied {Gasterosteus aculeatus
Linnaeus. Fundulus notatus [ Rafinesque]. and Trichogaster trich-
opterus [Pallas], respectively) increased as tlie interloper more close-
ly approached the territorial mid-point. However, this type of dif-
ferential aggression did not obviously occur in C. atrorus. The de-
fending male appeared to attack all intruders (excepting young and
reproductively-oriented females) with equal aggressiveness, regard-
less of their distance from the center of this area.

Territoriality in Aquaria

Results from a number of experiments with Cyprinodon atrorus
in aquaria are similar, and only one is described here. A bare, sand-
bottomed. 160-liter tank was used to attemi)t to duplicate tlie terri-
torial sequences observed under field conditions. Seven adults of
generally equal size, 3 males and 4 females, were jilaced in the tank,
and they immediately formed a tightly-knit, stationary cluster (a
"pyramid") near the substrate. They remained in tliat position for
several minutes, then began exploratory swimming. When disturbed
they immediately returned to a pyramid cluster. By the second
day the fish began to seek individual refuge when disturbed. There
was sporadic aggression, not limited to any i>articular area. On the
third day one male defended a small area near the substrate for a
short time. At the end of the first week, one male rigorously defend-
ed the entire bottom of the tank, from the substrate to about eight cm
above, making frequent forays against fish positioned at higher
levels. When attacked, other fish fled without defensive display. At

The Great Basin Naturalist
1 78 ITZKOWITZ AND MINCKLEY Vol. XXIX, No. 4

the end of the second week another male began to defend a 10- by
38-cni area at one end of the tank, positioned about 8 cm above the
bottom and extending to the surface. This was defended against all
but the male who continued to patrol the substrate. The third male
did not establish an area, and was intimidated by both other males.
There was continual, mild, aggressive activity among the females,
and they often attacked the subservient male. None of the female
attacks was prolonged, but consisted merely of a short charge termin-
ating in a butt or bite. The hierarchy in males was maintained for
a month, when the experiment was terminated. These results are
similar to those obtained by Barlow (1961a) for Cyprinodon macu-
larius Baird and Girard. and have bee nduplicated in our laboratory
using a number of other pupfishes.

Discussion and Conclusions

The study of behavior of fishes in the wild is exceedingly diffi-
cult, and the abundance of variables existing under field conditions
makes it simple to overlook significant factors and to arrive at
spurious conclusions. In the laboratory, new factors are introduced
by creation of "standard conditions," and data acquired from fish in
aquaria may be equally misleading.

Behavior of Cyprinodon atrorus in natural and semi-natural habi-
tats lacked many components that were commonly present under
aquarium conditions. Nuzzling, for example, generally occurred
only when a stationary female was 10 or more cm above the sub-
strate, and when she remained for some period of time in a male's
territory. Since all territories in nature or in the pond were in water
shallower than eight cm (and usually less than five cm), physical
limitations precluded jierformance of the pattern. The absence of
following (especially above and below) under field conditions may
also be attributable to insufficient depths. In addition, females in the
pond did not remain in the territory after a spawn, but moved away
at high speed so that little time was allowed for the male to perform
complex display patterns (with the notable exception discussed on
page 00).

The lack of records of the jerk motor pattern in nature and in the
pond is confusing. Barlow (1961a) implied that this was correlated
with passage of gametes; since young were abundantly produced in
the semi-natural pond, a jerk is obviously not essential for successful
reproduction. The jerk motor pattern in aquaria always led
to a continuance of spawning (Fig. 3). and only a slow rise resulted
in its termination. However, on the one occasion when repeated
spawning by a pair of fish was observed under field conditions (p.
00), the female did not jerk, and repeatedly utilized a slow rise (or
a period of quiescence in shallow water) after spawning.

Tilting is difficult to observe from above, and may be physically
impossible for a female in shallow water, and these factors may
explain its almost-complete absence from the behavioral repertoire

Dec. 31, 1969 behavioh oi- itpiimi 179

ill tlio poiul. llij^lily motivated females in a(]uaria al\va\s tilled, hut
often omitted a nip in their pre-spawnin^ movements.

The ahbreviated. "to-the-point" spawning se(|uerK<' under natural
conditions. hi<rhly stereotyped and ahnosl invariable, may bo a re-
sult of the over-riding territorially of the male ( . atrorus. A male
will curtail spawning at any point to chase an intruder in his terri-
tory. When a pair is placed in a habitat devoid of other fishes the
barraging effect of stimuli from interlopers is removed and the pair
may receive and react to other facets of their imiTiediate surround-
ings. Sexuality becomes the only broad stimulus, and subtleties of
activities on the part of each fish create a diversity of reactions in
each. Variability in their behavioral repertoires, such as occurs in
aquaria with the appearance of several courtship patterns, may then
ensue. This is supported somewhat by the diverse behavior exhibited
by the pair of fish isolated under semi-natural conditions by algal
mats (p. 00).

Development of territorial behavior in the laboratory is very
similar to that occurring under field conditions. Sporadic fighting
j)recedes acquisition of temj)orary territories, and fixed territories
remain relatively permanent. The artificial confinement in a tank
allows (or forces) development of a hierarchy. If these occur in
nature they are highly com])lex and localized. Males unable to hold
or establish territories in nature remain in deep water and avoid
contact with aggressive individuals, and smaller males sometimes
defend stones or vertical banks in peripheral areas. These kinds of
behavior correspond generally to the activities of the non-dominant
males in aquarium situations.

Literature Cited

Aronson, L. R. 1949. An analysis of reproductive behavior of the mouth-
breeding cichlid fish, Tilapia macroccphala (Bleeker). Zoologica 34: 133-157.

Baerends. G. p.. and J. M. Baerends-Van Boon. 1950. An introduction to
the study of the ethology of cichlid fishes. Behav. Suppl. 1, 242 pp.

Baerends, G. P., B. Brouwer, and H. Tj. Waterboi.k. 1955. Ethological
studies on Lebistcs reticulatus (Peters): 1. An analysis of the male courtship
pattern. Behaviour 8: 249-334.

Barlow, G. W. 1961a. Social behavior of the desert pupfish. Cyprinodon
macularius, in the field and in the aquarium. Amer. Midi. Nat. 65: 339-358.

. 1961b. Ethology of the Asian teleost, Badis badis. 1. Locomotion, main-
tenance, aggregation and fright. Trans. 111. State Acad. Sci. 54: 175-188.

. 1962a. Ibid. 3. Aggressive behavior. Z. Tierpsychol. 19: 29-55.

. 1962b. Ibid. 4. Sexual behavior. Copeia 1962: 346-360.

1963. Ibid. 2. Motivation and signal value of the colour patterns. Anim.

Behav. 11: 97-105.

Carranza, J., and H. E. Winn. 1954. Reproductive behavior of the blackstripe
topminnow (Fundulus notalus). Copeia 1954: 273-278.

Hess, E. H. 1952. Temperature as a regulator of the attack-response of Ri-tta
splendens. Z. Tierpsychol. 9: 379-382.

LiLEY, N. R. 1966. Ethological isolating mechanisms in four sympatric species
of poeciliid fishes. Behav. Suppl. 13. 197 pp.

Marker, P. R., and W. J. Hamilton. 1966. Mechanisms of Animal Behav-
iour. John Wiley and Sons, New York.

Miller, R. J. 1964. Studies on the social behavior of the blue gourami.
Trichogaster trichoplerus (Pisces. Belontiidae). Copeia 1964: 469-496.

The Great Basin Naturalist
180 ITZKOWITZ AND MINCKLEY Vol. XXIX, No. 4

md D. D. Hall. 1968. A quantitative description and analysis of

courtship and reproductive behavior in the anabantoid fish Trichogaster leeri

(Bleeker). Behaviour 32: 85-149.
Miller, R. R. 1968. Two new fishes of the genus Cyprinodon from the Cuatro

Cienegas basin, Coahuila, Mexico. Occ. Pap. Mus. Zool., Univ. Mich. 659:

1-15.
MiNCKLEY. W. L. 1969. Environments of the Bolson of Cuatro Cienegas,

Coahuila, Me.xico, with special reference to the aquatic biota. Publ. Univ.

Tex. El Paso, Sci. Ser. 2: 1-65.
. and M. Itzkowitz. 1967. Ecology and effects of intestinal gas accumu-
lation in a pupfish (genus Cyprinodon) . Trans. Amer. Fish. Soc. 96: 216-218.
. and G. A. Cole. 1968. Preliminary limnologic information on waters

of the Cuatro Cienegas basin, Coahuila, Mexico. Southwest. Nat. 13: 421-431.
and E. T. Arnold. 1969. "Pit digging." a behavioral feeding adaptation

in pupfishes (genus Cyprinodon) . Trans. Ariz. Acad. Sci. 5: 254-257.
Morris, D. 1958. The reproductive behavior of the ten-spine stickleback

(Pygosieus pungitius L.). Behav. Suppl. 4, 154 pp.
Nelson, K. 1964. The temporal patterning of courtship behavior in the glandulo-

caudine fishes (Ostariophysi, Characidae). Behaviour 24: 90-145.
Simpson. M. J. A. 1968. The display of the Siamese fighting fish, BeUa

splendens. Anim. Behav. Monogr. 1: 1-73.
Tinbergen, N. 1953. Social Behavior in Animals, with Special Reference to

Vertebrates. Methuen and Co., Ltd., London.

DOES VIREO GILVUS SWAINSONII OCCUR IN UTAH?

Gary L. Worthen'

While attempting to subspecifically identify a series of forty-two
Warbling Vireos collected in south central, western Utah for a recent
work (Worthen, 1968). a problem of subspecific distribution be-
came apparent. Twomey (1942), Woodbury and Russell (1945),
Behle (1948), Woodbury, Cottam, and Sugden, (1949). Behle
(1955), A.O.U. Check-list Committee (1957). Behle. (1958). and
Behle (1960), state that Vireo gilvus swainsonii Baird occurs or may
occur in Utah. Most of these authors designate the subspecies as a
migrant but some record it as a summer resident in certain localized
areas of the state.

I examined a series of 200 birds in the collection of the Univer-
sity of Utah and can find little evidence that V .g. swainsonii occurs
in Utah. It should be pointed out, however, that I have not examined
any topotypes or any specimens positively identified as V. g. swain-
sonii. Characteristics were used that were given by the original de-
scriber (Oberholser, 1932) and by Sibley (1940).

Virtually all specimens in the University of Utah Museum that
have been identified as V. g. swainsonii are birds taken in August
and September in fresh fall plumage. Furthermore, six of the seven
birds taken by myself in August (Worthen. 1968) have character-
istics of V . g. swainsonii. There is a decided paucity of birds taken
in spring in the collection of the University of Utah referable to
^'swainsonii". These data present two possible hypotheses: that this
subspecies commonly migrates south through Utah but usually mi-
grates north by some other route, or that the observed relationship
is caused by something other than patterns of migration. If the lat-
ter is true the factors which seem to be responsible are those dealing
with molt and wear of plumage. Bent (1950:366) states that,

There is a partial postjuvenal molt, beginning early in Au-
gust, which involves the contour plumage, and the wing coverts,
but not the rest of the wings nor the tail. This produces a first
winter plumage which is practically indistinguishable from the
winter plumage of the adult, greener above and more buffy
white below than the previous plumage.

Dr. Dwight says that the nuptial plumage is acquired by
wear, but Ned Dearborn (1907) found March and April speci-
mens of the western race undergoing a scattered molt on the
head and breast.

It seems apparent that the fresh fall plumage of the resident sub-
species {V. g. leucopolius (Oberholser)) would more closely approx-
imate the characteristics of V. g. swainsonii than breeding birds of

'Museum ol Natural History, University ol Kansas, Lawrence, Kansas ()60+4.

181

The Great Basin Naturalist
182 GARY L. WORTHEN Vol. XXIX, No. 4

V. g. leucopolius in worn and faded plumage. It is, therefore, my
contention that most (if not all) of the ''swainsonn' in the collec-
tion of University of Utah are misidentified through confusion of
variation in seasonal plumage with subspecific variation and are, in
actuality, birds assignable to V. g. leucopolius in fresh fall plumage.

Literature Cited

American Ornithologists' Union. 1957. Check-list of North American

birds (5th Ed). Amer. Ornithologists' Union, Baltimore, Maryland. 691 p.
Behle. W. H. 1948. Birds observed in April along the Colorado River from

EQte to Lee's Ferry. Auk, 65:303-306.
. 1955. The birds of the Deep Creek Mountains of central western

Utah. Univ. Utah Biol. Ser., 11 (4): 1-34.
. 1958. The birds of the Raft River Mountains, northwestern Utah.

Univ. Utah Biol. Ser., 11 (6): 1-40.

1960. Problems of distribution and speciation in Utah birds. Utah

Acad. Proc, 37:13-36.
Bent, A. C. 1950. Life histories of North American wagtails, shrikes, vireos,

and their allies. Smithsonian Inst. U. S. Nat. Mus., Bull. 197. 411 p.
De.\rborn, N. 1907. Catalogue of a collection of birds from Guatemala. Field

Mus. Natur. Hist. Pub. 125 Ornithol. ser., 1:60-138.
Oberholser. H. C. 1932. Description of new birds from Oregon, chiefly from

the Warner Valley region. Sci. Pub. Cleveland Mus. Natur. Hist. 4:1-12.
Sibley, C. G. 1940. The Warbling Vireo of the Cape District of lower Cali-
fornia. Condor, 42:255-258.
Twomey. a. C. 1942. The birds of the L^inta Basin, LTtah. Ann. Carnegie Mus.

27:341-490.
Woodbury. A. M., C. Cott.\m, and J. W. Sugden. 1949. Annotated check-list

of the birds of Utah. Univ. Utah Biol. Ser., 11 (2): 1-40.
Woodbury, A. M. and H. N. Russell, Jr. 1945. Birds of the Navajo countrv.

Bull. Univ. Utah. 35(14): 1-160.
WoRTHEN. G. L. 1968. The taxonomy and distribution of the birds of the

southeastern Great Basin, Utah. Unpublished Masters Thesis, Univ. Utah

589 p.

LIST OF INSECT TYPE SPECIMENS IN THE

ENTOMOLOGICAE COLLECTION OF THE

BRIGHAM YOUNG UNIVERSITY. PROVO. UTAH. NO. IV.

Vasco M. Tanner' and Don R. Harris-
There is a tendency at present in the biological sciences to shy
away from systematics in favor of molecular biolof^ical studies. The
Linnaoan system of dealing with the multitudinous forms of aiumate
nature is being seriously questioned. Today, biologists are concerned
with the synthesis of the living environment at levels of organiza-
tion such as the "biota, community, population, organism, cell, and
molecule."' No doubt emphasis in this line of endeavor will prove
to be revealing as to the similarity and unity of organisms. As ad-
vancements are made in this new approach to understanding the
basic makeup of organisms, their similarity and life processes, there
will still be a need for recognizing the great diversity of life forms.
The causes that bring about diversity of organisms may be more
clearly known and manipulated, but the end product, the objective
entity, that which we now know as the species, will still exist.

Modern methods employed in taxonomy are su{)erior to those
of the Linnaean period and should, through additional refinements,
continue to be one way of categorizing and reporting on the product
of evolution. In systematizing plants and animals, workers have de-
scribed the diverse forms and preserved, for future studies, some
of the specimens used in the description of the new species. These
specimens have been variously designated as: type, holotype. allo-
type, cotype, paratype, topotype and other designations. Experience
has justified the preservation of specimens so designated by the
author in making a description. Without the object described, it is
well nigh impossible for biologists and research workers to make a
satisfactory determination of some species. It is, therefore, in the
interest of future research work with the insect group, that the
species representing types are listed in these reports.

Entomology is an important discipline since it is concerned \N'ith
the study of a large segment of animal life. Research in this field
in the western states may be aided by having authoritative collec-
tions and some type material available at a focal point such as at
this university.

ORDER ORTHOPTERA

FAMILY GRYLLACRIDIDAE

Ceuthophilus nevadansis Barnum.

Brigham Young Univ. Sci. Bui., Biol. Ser., 4(3):96-100. 1964.

^DeparUiient of ZiHjlnpy and Entomology, Brigham Young University, I'rnvo, I'tati.
-Graduate student in Kntomology. Brigham Young University.
3Raven, P. H., and II. W. Richard. I'Ki". S.vst. Zool. 16:1.

183

The Great Basin Naturalist
184 V. M. TANNER AND D. R. HARRIS Vol. XXIX, No. 4

Paratypes, 19 c?cf, 16 ? 9 ; Nevada Test Site, Nye Co., Nev.

(Andrew Bamum).
Ceuthophilus deserticola Bamum.

Brigham Young Univ. Sci. Bui., Biol. Ser., 4(3)100-103, 1964.
Paratypes, 7 cfd", 9 ? ?; Nevada Test Site, Nye Co., Nev.

(Andrew Barnum).

ORDER HEMIPTERA

FAMILY MIRIDAE

Chlamydatus becki Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3) :28, 1968.
Paratypes, 2 cfcT, 2 9 2; Nevada Test Site, Nye Co., Nev.
(H. H. Knight and J. M. Merino) .
Microphylellus symphoricarpi Knight.

Brigham Young Uni. Sci. Bui., Biol. Ser., 9(3) : 30, 1968.
Paratypes, 1 cT, 1 9 ; Nevada Test Site, Nye Co., Nev. (H. H.
Knight)
Plagiognathus salviae Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser.. 9(3) :30, 1968.
Paratypes, 4 d" cf , 4 9 9 ; Nevada Test Site, Nye Co., Nev. (D E.
Beck, H. H. Knight and J. M. Merino).
Merinocapsus ephedrae Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3) :34, 1968.
Partypes, 3 cT cT, 5 9 9 ; Nevada Test Site, Nye Co., Nev. (D E.
Beck, H. H. Knight) .
Beckocoris laticephalus Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9 (3): 36, 1968.
Paratypes, 5 cTcT, 5 9 9; Nevada Test Site, Nye Co., Nev.
(H. H. Knight. J. M. Merino).
Europiella punctipes Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3) :47, 1968.
Paratypes, 2 cT d', 1 9 ; Nevada Test Site, Nye Co., Nev. (H. H.
Knight, D E. Beck) .
Europiella lycii Knight.

Brigham Young Univ. Sci. BuL, Biol. Ser., 9(3) : 40. 1968.
Parat}Tpes, 1 cf , 3 9 9; Nevada Test Site. Nye Co., Nev. (D E.
Beck, H. H. Knight) .
Europiella grayiae Knight.

Brigham Young Univ. Sci. BuL, Biol. Ser., 9(3) :41. 1968.
Parat\T)es, 5 cf cT, 5 9 9 ; Nevada Test Site, Nye Co., Nev. (D E.
Beck. H. H. Knight and J. M. Merino).
Europiella stitti Knight.

Brigham Young Univ. Sci. Bui, Biol. Ser., 9(3): 46, 1968.
Paratype, 1 9 ; Gila Bend. Ariz.; 13 Feb. 1941 (Loyd L. Stitt).
Europiella nigrofemorata Knight.

Brigham Young Univ. Sci. Bui., Biol. Sen, 9((3):39. 1968.

Dec. 31, 1969 list of inskct types 185

Paratype, 1 cf ; Nevada Test Site, Nye Co.. Nev. (H. II. Riu^lit.
D E. Beck and J. M. Merino).

EuropicUa arizonac Knight.

Brigham Young Univ. Sci. Bui.. Biol. Ser.. 9(3): 46, 1968.
Paratype, 1 9 ; Aztec, Ariz., 3 Feb. 1941 (Loyd L. Stitt).

Europiella nigrirornis Knight.

Brigham Young Univ. Sci. Bui.. Biol. Ser., 9(3) :40. 1968.
Para types, 1 cf. 1 $ ; Nevada Test Site, Nye Co., Nev. (H. H.
Knight. DE. Beck).

Europiella nicholi Knight.

Brigham Young Univ. Sci. Bui.. Biol. Ser.. 9(3) :42, 1968.
Paratype, 1 9 ; Bincon Mts.. Ariz., 2 Sept. 1923, Ah. 3,300
(A. A. Nichol).
Europiella albipubescens Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3) :46, 1968.
Paratype, 1 d ; Nevada Test Site, Nye Co., Nev. (H. H. Knight.
D E. Beck and J. M. Merino) .

Europiella yampae Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3) :43, 1968.
Paratype. 1 d" ; Steamboat Sprs., Colo., 12 July 1964, 6700'

(H. H. Knight).

1 9. Steamboat Sprs., Colo., 11 July 1964, 6700' (H. H.

Knight) .
Europiella unipuncta Knight.

Brigham Young Univ. Sci. BuL, Biol. Ser., 9(3) :44, 1968.
Paratypes, 1 cT, 1 9 ; Nevada Test Site, Nye Co.. Nev. (J. M.

Merino, H. H. Knight) .

Europiella rubricornis Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3): 39, 1968.
Paratype, 1 d ; Richfield. Utah, 11 June 1930, (Light Trap);
(E. W. Davis).
Europiella rufiventris Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3) :42, 1968.

Paratypes, 1 cf , 1 9 ; Goldwater, Arizona, 6 April 1940 (Loyd
L. Stitt).

Psallus purshiae Knight.

Brigham Young Univ. Sci. BuL, Biol. Ser., 9(3) :48, 1968.
Paratype, 1 d" ; Nevada Test Site, Nye Co., Nev. (H. H. Knight,
J. M. Merino).

Psallus atriplicis Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser.. 9(3) :48. 1968.
Paratype. 1 d" ; Nevada Test Site, Nye Co., Nev. (H. H. Knight).

Phymatopsallus prosopidis Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser.. 9(3) :49, 1968.
Paratypes, 4 d' d", 4 9 9 ; Nevada Test Site, Nye Co.. Nev (D E.
Beck, H. H. Knight).

The Great Basin Naturalist
186 V. M. TANNER AND D. R. HARRIS Vol. XXIX, No. 4

Phymatopsallus rubesi Knight.

Brigham Young Univ. Sci. Bui, Biol. Ser., 9(3) :50, 1968.
Paratypes. 1 d. \ 9 ; Nevada Test Site, Nye Co., Nev. (J. M.
Merino).

Lepidopsallus arizonae Knight.

Brigham Young Univ. Sci. Bui, Biol. Ser., 9(3) :52, 1968.
Paratypes, 1 cf , 1 ? ; Mt. Lemon, Ariz., Santa Cat. Mts., 27 July
1917, 9000' (H.H. Knight).
Lepidopsallus nicholi Knight.

Brigham Young Univ. Sci. Bui, Biol. Ser., 9 (3): 52, 1968.
Paratype, 1 ? ; Empire Mts., 5000', Ariz., 20 May 1928 (A. A.
Nichol).
Lepidopsallus pini Knight.

Brigham Young Univ. Sci. Bui. Biol. Ser., 9(3) :53, 1968.
Paratype, 1 9 ; Ft. Garland. Colo., 10 Aug. 1925 (H. H. Knight)
(Pinus edulis).

Lepidopsallus longirostris Knight.

Brigham Young Univ. Sci. Bui, Biol. Ser., 9(3) :54, 1968.
Paratypes, 1 cf , 1 9 ; Pingree Pk., Colo., 20 Aug. 1925 (H. H.
Knight) .
Lepidopsallus monticola Knight.

Brigham Young Univ. Sci. Bui. Biol Ser., 9(3) :54. 1968.
Paratypes. 1 cT, 1 9; Veta Pass, Colo., 9 Aug. 1925 (H. H.
Knight) ( On Pinus).

Nevadocoris becki Knight.

Brigham Young Univ. Sci. Bui. Biol Ser.. 9(3) :59, 1968.
Paratypes. 3 cTcf-. 3 9 9; Nevada Test Site, Nye Co., Nev.
(H. H. Knight, D E. Beck and J. M. Merino) {On Tetra-
dymia).
Nevadocoris pallidus Knight.

Brigham Young Univ. Sci. Bui. Biol. Ser.. 9(3) :60, 1968.
Paratypes, 1 d", 1 9 ; Nevada Test Site, Nye Co., Nev. (D E.
Beck. H. H. Knight) (On Grayia spinosa).
Nevadocoris bullatus Knight.

Brigham Young Univ. Sci. Bui, Biol. Ser., 9(3): 60, 1968.
Paratypes, 2 cf cf ■ 2 9 9 ; Nevada Test Site, Nye Co.. Nev. (D E.
Beck, H. H. Knight) {On Chrysothamnus) .

Brachyceratocoris nevadensis Knight.

Brigham Young Univ. Sci. Bui. Biol Ser., 9(3) :61, 1868.

Paratypes. 2 cT cT, 2 9 9; Nevada Test Site, Nye Co., Nev. (H.
H. Knight).
Macrotylus salviae Knight.

Brigham Young Univ. Sci. Bui, Biol. Ser., 9(3): 62. 1968.

Paratypes, 2 cT cT. 2 9 9 ; Nevada Test Site, Nye Co., Nev. (D E.
Beck and H. H. Knght) .
Coquillettia luteiclava Knight.

Brigham Young Univ. Sci. Bui, Biol. Ser.. 9(3): 63. 1968.

Dec. 31, 1969 list of insi:ct types 187

Paratypes. 1 cT ; l9; Nevada Test Site. Nye Co.. Nev. (H. H.
Knight and J. M. Merino).
('o(juiUctti(i alhcl/a Knight.

Brighani Young Univ. Sci. Bui.. Biol. Sen, 9(3j:62, 1968.
Paratypes, \ d ; Nevada Test Site. Nye Co., Nev. (H. II. Knight
and J. M. Merino). On Eriogonum inflatum.

Dicyphus ribesi Knight.

Brigham Young Univ. Sci. Bui.. Biol. Ser.. 9(3): 70. 1968.
Paratypes, 26" d". 2 9 ? ; Nevada Test Site, Nye Co.. Nev. (Beck,
Marino and Knight).

Dicyphus hespcrus Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser.. 9(3): 73. 1968.
Paratypes, 1 d ; Moscow, Idaho. 18 Mar. 1934; 2560', (H. Ship-
man) (Mullen). 1 9, Genese, Ida.. 23 May 1936 (R. E.
Miller) .
Dicyphus stitti Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3): 74. 1968.
Paratype. 1 d ; Gila Bend. Ariz., 6 Oct. 1940 (Loyd L. Stilt).
On Martynia parviflora.

Macrolophus mimuli Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3): 75. 1968.
Paratypes, 1 d", 1 9; Huachuca Mts., 6000'. Ariz.. 14 June
1928 (A. A. Nicol). Breeding on Mimulus cardinalis.

Sixeonotus bebbiae Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3): 76, 1968.
Paratype, 1 cT ; Mohawk, Ariz., 23 Apr. 1940 (Loyd L. Stitt).

Sixeonotus dextratus Knight.

Brigham Young Univ. Sci. Bui, Biol. Ser., 9(3): 76. 1968.
Paratype, \ <S '-, Tucson, Ariz., Sabino Canyon. 23 July 1917 (II.
H. Knight).
Sixeonotus areolatus Knight.
Bui. Brook. Ent. Soc. 23:243.

Paratype, 1 cf; Rock Island, Tex., 3^4 m. Northwest; 20 June
1922 (Grace O. Wiley).

Largidea nevadensis Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3) :84, 1968.
Paratypes, 1 d', 1 9 ; Nevada Test Site. Nye Co.. Nev. (H. H.
Knight and J. M. Merino) . On Pinus monophylla.

Labops hirtus Knight.

Brigham Young Univ. Sci. Bui., Biol.Ser., 9(3) :88. 1968.
Paratype, 1 d ; Yellowstone Park, Wyo.. 20 July 1925 (A. A.
Nichol).
Slaterocoris longipennis Knight.

Brigham Young Univ. Sci. Bui.. Biol. Ser., 9(3) :90. 1968.
Paratypes, 2 d' cT, 2 9 9; Nevada Test Site. Nye Co., Nev. (H.
H. Knight).

The Great Basin Naturalist
188 V. M. TANNER AND D. R. HARRIS Vol. XXIX, No. 4

Slaterocoris rubrofemoratus Knight.

Brigham Young Univ. Sci. Bui., Biol. Sen, 9(3) :90, 1968.
Paratypes, 1 cf, 2 9 9 ; Nevada Test Site. Nye Co., Nev. (H. H.
Knight). On Artemisia tridentata.

Lopidea deserta Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3): 99, 1968.
Paratypes, ScT cT- 5 9 9 ; Nevada Test Site, Nye Co., Nev. (Beck,
Knight and Merino).
Lopidea colli Knight.

Bui. Brook. Ent. Soc, 29:11. 1934.

Paratype. LaSal. San Juan, Co., Utah. June, 1927 (Anson Call,
Jr.).
Lopidea becki Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3) :98, 1968.
Paratype. 1 fl; Nevada Test Site. Nye Co., Nev. (Beck, Knight,
and Merino). On Eriogonum umbellatum.

Ceratopidea daleae Knight.

Brigham Young Univ. Sci. Bui.. Biol. Ser., 9(3): 100. 1968.
Paratypes, 3 cf d", 3 9 9 ; Nevada Test Site. Nye Co., Nev. (H.
H. Knight) . On Dalea polyadenia.

Daleapidea daleae Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3): 101, 1968.

Paratypes, 1 cf , 1 9 ; Nevada Test Site, Nye Co., Nev. (Beck,
Knight and Merino).
Hesperocapus plagiatus Knight.

Brigham Young Univ. Sci. BuL, Biol. Ser., 9(3): 103, 1968.

Paratype. 1 9 ; Nevada Test Site, Nye Co., Nev. (H. H. Knight).

HesDerocapus artemisicola Knight.

Brigham Young Univ. Sci. BuL, Biol. Ser.. 9 (3): 102, 1968.
Paratype, 1 cT ; Hudson, Colo., 25 Aug. 1925 (H. H. Knight)
{Artem. f Hi folia).
Dichaetocoris nevadensis Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser.. 9(3): 103, 1968.
Paratypes. 2 cfd". 2 9 9; Nevada Test Site, Nye Co., Nev.
(Knight and Merino).

Dichaetocoris merinoi Knight.

Brigham Young Univ. Sci. BuL. Biol. Ser., 9(3): 111, 1968.
Paratype, 1 cT ; Nevada Test Site, Nye Co., Nev. (Knight and
Merino). On Piuus monophylla.

Dichaetocoris pinicola Knight.

Brigham Young Univ. Sci. BuL. Biol. Ser.. 9(3): 110. 1968.
Paratypes, 3 cfcf. 3 9 9; Nevada Test Site, Nye Co., Nev.
(Knight and Merino). On Pinus monophylla.

Dichaetocoris symphoricarpi Knight.

Brigham Young Univ. Sci. BuL, Biol. Ser.. 9(3): 112. 1968.
Paratypes, 5 d d ■ ^ 9 9; Nevada Test Site, Nye Co.. Nev.

Dec. 31, 1969 ltst nv inskct types 189

(Beck, Kniglit and Merino). I lost plant Syrnphoricarpos.

longifloris.
Dichacfororis junipcri Knight.

Brigham Young Univ., Sci. Bui.. Biol. Ser., 9(3): 113. 1968.
Paratypes. 3 cfcf. 3 9 9; Nevada Test Site. Nye Co.. Nev.

(Knight and Merino). On Juniperus ostrosperma.

Dichaetocoris utahensis Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3): 114. 1968.
Paratype, 1 9 ; Scipio, Utah. 29 June 1965 (H. H. Knight).

Dichaetocoris stanleyaea Knight.

Brigham Young Univ. Sci. Bui. Biol Ser.. 9(3): 115, 1968.
Paratypes. 2 d" cf ; Nevada Test Site. Nye Co., Nev. (Beck.
Knight and Merino). Host plant Stanleya pinnata.

Dichaetocoris coloradensis Knight.

Brigham Young Univ. Sci. Bui.. Biol. Ser.. 9(3): 116. 1968.
Paratypes, 1 cT, 1 9; Durango, Colo., 13 Aug. 1925 (H. H.
Knight).
Melanotichus shoshonea Knight.

Brigham Young Univ. Sci. Bui.. Biol. Ser., 9(3): 125, 1968.
Paratypes. 1 d" ; Yellowstone Nat. Park, Wyo., 8 Aug. 1937 (H.
H. Knight). 1 ?, Shoshone Nat. Forest. Wyo.. 7 Aug. 1927
(H. H. Knight).
Melanotrichus azteci Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3): 122, 1968.
Paratype, 1 9; Phoenix, Ariz.. 17 Mar. 1939 (Loyd L. Stitt).

Melanotrichus malvastri Knight.

Brigham Young Univ. Sci. Bui.. Biol. Ser., 9(3): 120, 1968.
Paratypes, 1 d', 1 9 ; Whittier, Cal., 18 April 1935 (E. L. Pad-
dock) .
Melanotrichus nevadensis Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser.. 9(3): 122, 1968.
Paratypes, 5d cT, 5 9 9; Nevada Test Site, Nye Co., Nev. (H.
II. Knight and J. M. Merino). On Artemisia tridentata.

Melanotrichus pollens Knight.

Brigham Young Univ. Sci. Bui.. Biol. Ser.. 9(3): 125, 1968.
Parat>-pes, 1 d, 1 9 ; Nevada Test Site, Nye Co., Nev. (D E.
Beck, H. H. Knight and J. M. Merino).

Melanotrichus eurotiae Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3): 128, 1968.
Paratypes, 5 d cT, 5 9 9; Nevada Test Site. Nye Co.. Nev. (D
E. Beck, H. H. Knight and J. M. Merino). Taken on Euratia
lanata.
Melanotrichus atriplicis Knight.

Brigham Young Univ. Sci. Bui.. Biol. Ser., 9(3): 125. 1968.
Paratype, 1 9 ; Nevada Test Site, Nye Co.. Nev. (11. II. Knight
and J. M. Merino). Taken on Atriplex canescens.

The Great Basin Naturalist
190 V. M. TANNER AND D. R. HARRIS Vol. XXIX, No. 4

Parthenicus furcatus Knight.

Brigham Young Univ. Sci. BuL, Biol. Ser., 9(3): 133, 1968.
Para type, 1 cT ; Nevada Test Site, Nye Co., Nev. (D E. Beck
and J. M. Merino). I'aken at incandescent light.
Parthenicus pinicola Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3): 140, 1968.
Paratypes. 1 ? ; Grand Canyon, Ariz.. 40 mi. South. 3 Aug. 1917
(H. H. Knight). 1 d'; Durango, Colo.. 13 Aug. 1925 (H. H.
Knight) .
Parthenicus merinoi Knight.

Brigham Young Univ. Sci. Bui.. Biol. Ser., 9(3): 135. 1968.
Paratype. 1 d" ; Nevada Test Site, Nye Co., Nev. (D E. Beck
and J. M. Merino).
Parthenicus atriplicis Knight.

Brigham Young Univ. Sci. Bui, Biol. Ser., 9 (3): 135, 1968.
Paratypes, 1 d". 1 ? ; Nevada Test Site, Nye Co.. Nev. (H. H.
Knight and J. M. Merino).
Parthenicus incurvus Knight.

Brigham Young Univ. Sci. Bui. Biol. Ser., 9(3): 150. 1968.
Paratype. 1 d" ; Nevada Test Site. Nye Co.. Nev. (D E. Beck
and J. M. Merino) .
Parthenicus becki Knight.

Brigham Young Univ. Sci. Bui, Biol. Ser.. 9(3): 149. 1968.
Paratypes, 1 d"- 1 ? ; Nevada Test Site, Nye Co., Nev. (H. H.
Knight).
Parthenicus conspersus Knight.

Brigham Young Univ. Sci. Bui, Biol Ser.. 9(3): 139, 1968.
Paratypes. 1 d". 1 9 ; Richfield. Utah, 8 Julv 1930 (Light trap).
(E. W. Davis).
Parthenicus cowaniae Knight. fl

Brigham Young Univ. Sci. Bui. Biol. Ser., 9(3): 148, 1968.
Paratypes. 1 d", 1 ? ; Grand Canyon. Ariz., 3 Aug. 1917 (H.
11 Knight) .
Parthenicus houtalouae Knight.

Brigham Young Univ. Sci. Bui. Biol. Ser., 9(3): 154. 1968.
Paratype, 1 9 ; Hauchuca Mts.. 6000'. Ariz., 14 June 1928 (A.
A. Nichol). On Bouteloua gracilis.

Ceratocapsus nevadensis Knight.

Brigham Young Univ. Sci. Bui, Biol. Ser., 9(3): 156. 1968.
Paratypes. 2 9 ; Nevada Test Site. Nye Co., Nev. (D E. Beck.
H. H. Knight and J. M. Merino). Taken on Sphaeralcea.

Pilophorus miscosetosus Knight.

Brigham Young Univ. Sci. Bui, Biol. Ser.. 9(3): 169. 1968.

Paratypes, 1 d" - 1 9 ; Nevada Test Site. Nye Co., Nev. (H. H.
Knight and J. M. Merino).
Pilophorus merinoi Knight.

Brigham Young Univ. Sci. Bui, Biol Ser., 9(3): 175, 1968.

Dec. 31, 1969 list oi- insect types 191

Paratypes, 1 ? ; Nevada Test Site, Nye Co.. Nev. ( I i. H. Knij^lit
and J. M. Merino).

Pilop/iorus i/taht^nsi^ Knight.

Bnghani Young Univ. Sci. Bui.. Biol. Ser.. 9(3): 175, 1968.
Paratypes. 1 d ■■, Gnd. June. Colo.. 28 July 1900; 1 9; Leeds.
Utah. 12 Oct. 1952 (E. W. Davis).

Pilophorus nevadensis Knight.

Brigham Young Univ. Sci. Bui.. Biol. Ser.. 9(3): 172. 1968.

Paratype, 1 cT ; Wells, 15 mi. East, Nev. 12 July, 1965 (H. 11.
Knight) .
Pilophorus hesperus Knight.

Brigham Young Univ. Sci. BuL, Biol. Ser., 9(3): 169. 1968.

Paratypes, 1 cf, 1 9 ; Estes Park. Colo.. 24 Aug. 1925 (II. H.
Knight).
Pilophorus diffusus Knight.

Brigham Young Univ. Sci. Bui.. Biol. Ser.. 9(3): 168. 1968.

1 Paratype; Pingree Park, Colo.. 20 Aug. 1923 (C. J. Drake).

Pilophorus dislocatus Knight.

Brigham Young Univ. Sci. Bui, Biol. Ser., 9(3): 171, 1968.
Paratype, 1 9 ; Huachuca Mts., Ariz., 29 July 1905 (L. L. Stitt).

Pilophorus salicis Knight.

Brigham Young Univ. Sci. BuL. Biol. Ser.. 9(3): 173. 1968.
Paratypes, 1 d"; Las Animas, Colo., 6 Aug. 1925 (H. H. Knight).
1 9 . Wray, Colo.. 4 Aug. 1925 (H. H. Knight).

Irbisia shulli Knight.

Bui. Brook. Ent. Soc, 36:75. 1941.

Paratypes, \ d ; Lenore, Ida., 7 May 1938, 1000' (W. E. Shull).

1 9, Lenore, Ida., 7 May 1938, 1000, (W. W. Shull. E.

Ritzheimer) .

Irbisia fuscipuhescens Knight.

Bui. Brook. Ent. Soc, 36:76, 1941.

Paratypes, W, 1 9; Tampico, Wash., 10 May 1925 (E. W.
Davis).
Irbisia elongate Knight.

Bui. Brook. Ent. Soc, 36:77, 1941.

Paratypes, 1 cT, 1 9; Wen. Mts., Wn., 9 July 1930 (F. P.
Dean).
Irbisia nigripes Knight.

Canadian Ent.. 57:94, 1925.

Autotype, Rosevere Creek, Raft River Mts., Utah, June 1928 (D
Elden Beck).

Pinalitus brevirostris Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3): 187. 1968.
Paratypes, 1 cf ; Trinidad, Colo.. Stonewall 8500', 7 Aug. 1925
(H. IT. Knight). 1 9; Trinidad, Colo., Stonewall 8500'. 8
Aug. 1925 (H. II. Knight).

The Great Basin Naturalist
192 V. M. TANNER AND D. R. HARRIS Vol. XXIX, No. 4

Pinalitus rubrotinctus Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3): 189, 1968.
Paratype, 1 9 ; Mt. Lemon, Ariz., Santa Cat. Mts., July 26,
1917,9000' (H.H. Knight).

Dichrooscytus flavescens Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3): 196, 1968.
Paratypes, 2 $ ; New^castel, Wyo., 19 Ang. 1927 (H. H. Knight).

1 cf ; Grand Can., Ariz., 6 Sept. 1931 (H. H. Knight); 1 9 ;

Nevada Test Site, Nye Co., Nev. (D E. Beck, H. H. Knight).

Dichrooscytus rufivenosus Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3): 196, 1968.
Paratypes, IcT. 1 ? ; Pagosa Sprs., Colo., 12 Aug. 1925 (H. H.
Knight) .
Dichrooscytus ruberellus Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3): 199, 1968.
Paratypes, 1 c^, 1 9 ; Pingree Pk:. Colo., 22 Aug. 1925 (H. H.
Knight) .
Phytocoris flavellus Knight.

Brigham Young Univ. Sci. Bui.. Biol. Ser., 9(3):234, 1968.
Paratypes, 1 d" , 1 9 ; Nevada Test Site, Nye Co., Nev. (D E.
Beck, H. H. Knight, and J. M. Merino).

Phytocoris albidosquams Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3):232, 1968.
Paratype, 1 cT ; Nevada Test Site, Nys Co., Nev. (D E. Beck).

Phytocoris juniperanus Knight.

Brigham Young Univ. Sci. Bui, Biol. Ser., 9(3)238, 1968.
Paratype, 1 d ; Nevada Test Site. Nye Co., Nev. (H. H. Knight
and J. M. Merino). On Juniper us osteosperma.

Phytocoris subcinctus Knight.

Brigham Young Univ. Sci. Bui.. Biol. Ser., 9(3): 254, 1968.
Paratype. 1 cf ; Richfield, Utah, 15 Aug. 1929 (E. W. Davis),
Light trap.

Phytocoris becki Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3): 214, 1968.
Paratypes. 5 cT cT, 3 9 9; Nevada Test Site, Nye Co.. Nev. (D

E. Beck, H. H. Knight, and J. M. Merino). Taken on Ephedra

nevadensis) .

Phytocoris tanneri Knight.

Brigham Young Univ. Sci. Bui.. Biol. Ser.. 9(3): 257, 1968.
Paratype, 1 d"; Richfield, Utah, 15 July 1929 (E. W. Davis).
Taken at light trap.

Phytocorois rostratus Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3): 253, 1968.
Paratype, 1 cT ; Nevada Test Site, Nye Co., Nev. H. H. Knight,

D E. Beck, and J. M. Merino). Swept from Chrysothamnus

nauseosus.

Dec. 31, 1969 list of insect types 193

Phytocoris tricintus Knight.

Brighani Young Univ. Sci. Bui.. Biol. Ser.. 9(3):256. 1968.

Paratype. 1 9; Tucson, Ariz.. 12 May 1929 (E. D. Ball).
P/iytocoris rnerinoi Knight.

Brigham Young Univ. Sci. Bui.. Biol. Ser., 9(3):227. 1968.

Paraty])es. 1 cT, 1 9 ; Nevada Test Site, Nye Co., Nev. (D E.
Beck, H. H. Knig'ht and J. M. Merino). Taken on Grayia
spinosa.
Phytocorius breviatus Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3):226, 1968.

Paratypes. 1 cT, 3 9 9; Nevada Test Site. Nye Co., Nev. (J. M.
Merino) . Taken at black light.
Phytocoris deserticola Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3):251, 1968.

Paratype. 1 9 ; Nevada Test Site. Nye Co.. Nev. (H. H. Knight).
Phytocoris nigrolineatus Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3):224, 1968.

Paratype, 1 d" ; Nevada Test Site, Nye Co., Nev. (H. H. Knight
and J. Merino). Taken on Solazaria mexicana.
Phytocoris reticulatus Knight.

Brigham Young Univ. Sci. Bui., Biol. Ser., 9(3): 21 7, 1968.

Paratypes, 1 cT, 1 9 ; Nevada Test Site, Nye Co., Nev. (Taken
from can pit-trap).
Sittocapsus franseriae Knight.

Entomological News, 53:156, 1942.

Paratype, Mohawk, Ariz., April 6, 1927 (Loyd L. Stitt).

Bolteria juniperi Knight.

Brigham Young Univer. Sci. Bui., Biol. Ser., 9(3):202, 1968.
Paratypes, 1 cT ; Nevada Test Site, Nye Co., Nev., 1 9 ; Scipio,
Utah, 29 June 1965 (H. H. Knight).

FAMILY SALDIDAE

Saldula varionis Drake and Hottes.

Great Basin Nat., 10(1-4) :57, 1950.

Paratypes, 7, 5 from Escalante Desert, Kane Co., Utah (D Elden

Beck); 1 from Gateway, Colo., Sept.-Oct. 1949 (F. C. Hottes

and C. Drake).

ORDER DIPTERA

FAMILY TIPULIDAE
Erioptera maria Alexander.

Amer. Midi. Nat., 39:20, 1948.

19 paratypes; Puffers Lake, Beaver Canyon, Utah; 20 July 1936
(D. Elmo Hardy).
Tipula edmundsi Alexander.

Amer. Midi Nat., 39:75, 1948.

The Great Basin Naturalist
194 V. M. TANNER AND D. R. HARRIS Vol. XXIX, No. 4

1 para type; Logan Canyon, Utah Experimental Station; 22 May

1938 (D. Elmo Hardy).

FAMILY BIBIONIDAE

Bibio knowltoni var. paltidus Hardy.

Utah Acad. Sci., Arts and Letters, 14:203, 1937.

Holotype d - Provo Environs. Utah; 8 May 1937 (D. Elmo

Hardy) .
Allotype 5 - Provo Environs. Utah; 8 May 1937 (D. Elmo

Hardy).
Para type <S and 2 - Provo Environs. Utah Co.. Utah (Lowell
Miller).
Bibio lobata Hardy.

Utah Acad. Sci.. Arts and Letters, 14:203. 1937.
Para type ? - Sierra Co.. California; W. M. Wheeler Collection.
Bibio paint eri James.

Amer. Mus. Noviates 832:2. 1936.

2 paratypes; Manhattan. Kansas; 17 April 1934 C. W. Sabrosky).
Bibio albipennis beameri Hardy.

Kans. Univ. Sci. Bui., 30:451, 1945.

8 paratypes; Douglas Co., Kansas; 5 November 1931 (R. H.
Beamer) .
Plecia americana Hardy.

Kans. Ent. Soc. Jour.. 13:15. 1940.

1 paratype; Pensacola. Florida; 30 April 1888 (A. S. Stevens).
Plecia nearetic Hardy.

Kans. Ent. Soc. Jour.. 13:20, 1940.

1 paratype cf - Morgan City, Louisiana; 17 April 1938 (John
Standish).
Bibo melanopilosus var. bisepta Hardy.

Utah Acad. Sci., Arts and Letters. 14:204, figs. 7. 8. 9. 10. 1937.
Holotype, allotype, and 44 paratypes; Provo Environs (Brigham
Young University Campus), Utah, 12 May 1936 (D. E.
Hardy) .
Bibio lucens Hardv.

Utah Acad. Sci.. Arts and Letters. 14:203-204. 1937.

3 paratvpes; 1 - Hennepin Co.. Minn.. Oak Grove; 13 Mav 1933

(A.^A. Nichol). 1 - Mer Blue, Ont.; 28 May 1927 (G. S.

Walley). 1 - Miners Bay, Ont.; 26 May 1929 (G. S. Walley).
Bibio knowltoni Hardy.

Utah Acad. Sci.. Arts and Letters. 14:202-203, figs. 5 and 6. 1937.
Allotype ? ; Sandy. Utah; 5 May 1931 (G. F. Knowlton) (On

Sophia sophia) .
Holotype d" ; Granger. Utah; 29 April 1931 (G. F. Knowlton)

{On Lepidium).
Paratype d" ; Sunset. Utah; 5 May 1931 (G. F. Knowlton) {On

Erodium cleutarium).

Doc. 31,1969 LIST OI" INSECT TYPES 195

Bibio sericata Hardy.

Utah Acad. Sci.. Arts and Letters, 14:207, 1937.
14 paratvpes; 6 - Sunrise, ele. 6318 ft. Mt. Rainier. Washington;
13 August 1931 (J. Wilcox). 8 - Headley. B.C.; 29 August
1923 (C. R. Garrett).
Bibio rnickeli 1 lardy.

Utah Acad. Sci., Arts and Letters. 14:205, figs. 11. 12. 1937.
47 jiaratypes; 27 - Prontenac, Minn.; 29 May 1930 (C. E.
Mickel). 18 - Goodhue Co.. Minn.; 29 May 1930 (Carl T.
Schmidt). 1 - Minneapolis. Minn.. Mississippi River near
"U;'; 18 May 1922 (Wm. E. Hoffmann). 1 - Ecology Col-
lection; Hennepin Co., Minn.; basswood-maple ass'n; 27 May

1922 (Wm. E. Hoffmann).

Bibio rufalipes Hardy.

Utah Acad. Sci., Arts and Letters. 14:207, 1937.

2 para types - Waco, Texas (Belfrage) Collection of C. V. Riley.

Bibio nigrifemoratus Hardy.

Utah Acad. Sci., Arts and Letters, 14:206. 1937.

6 paratypes; 2 - Monte. B.C.; 13 May 1936 (J. K. Jacob). 1 -
Rochchester, Washington; 30 March 1932 (Wm. W. Baker).
2 - Mecham. Oregon; Ele. 3680 ft.; 8 May 1927 (H. A.
Schullen). 1 - Kiger's Island, Oregon; 12 April 1930 (J.
Wilcox).
Bibio utahensis Hardy.

Utah Acad. Sci., Arts and Letters. 14:208-209, 1937.

Holotype and 33 paratypes.

Holotype and 16 paratypes - Provo Environs, Utah Co.. Utah
(Harry Thomas). 5 - South Fork, Provo Canyon, Utah; 8
May 1937 (D. E. and Agnes Hardy). 1 - Utah Lake. East
Side. Utah Co., Utah. 3 - Provo Environs. Utah; 10 May 1937
(M. L Killpack). 1 - Provo, Utah (A. Sutherland). 6 - Provo.
Utah (D. E. Hardy). 1 - Aweme, Man.. Canada; 30 May

1923 (R.M.White).

Bibio signatus Hardy.

Utah Acad. Sci., Arts and Letters. 14:208 .1937.

Holotype d' - Spanish Fork, Utah; 29 April 1936 (D. E. Hardy).

Bibio velcida Hardy.

Utah Acar. Sci., Arts and Letters, 14:209, 1937.

4 <S paratypes; 3 - Low Bush, Ontario, Lake Abitibi; 12 June

1925 (N. K. Bigelow). 1 - Fredricton, New Brunswick; 30

May 1931 (R. P. Gorham).

Philia oklahomensis Hardy.

Utah Acad. Sci., Arts and Letters, 14:211, figs. 19, 20, 21. 1937.
11 paratypes; 1 - J^cust Grove, Oklahoma; 5 May 1934 (E. E.

Ivy) (genitalia dissected). 2 - Paris. Texas; 9 April 1904 (C.

T. Brues). 3 - College Station, Texas; 27. 30 March 1932 (H.

J. Reinhard) (4642). 3 - Victoria, Texas; 13 March 1908 (in

The Great Basin Naturalist
196 V. M. TANNER AND D. R. HARRIS Vol. XXIX, No. 4

Spanish Moss) (J. D. Mitchell). 1 - Dallas, Texas; 25 March
1906 (on salix) (W. D. Pierce).

Philia arizonaensis Hardy.

Utah Acad. Sci., Arts and Letters, 14:209, 210, figs. 13, 14, 1937.
3 paratypes; Huachuca Mt., Arizona (Brooklyn Museum Coll.
1929, Catalog No. 162 (2) and 163).

Phila breviceps var. atelestes Hardy.

Utah Acad. Sci.. Arts and Letters, 14:210, 1937.
51 paratypes; Meach Lake, Quebec; 21 June 1916.

Philia stigmaterus var. nigra Hardy.

Utah Acad. Sci.. Arts and Letters. 14:212. 1937.

14 paratypes; 3 - Plummer, Minn.; 10 and 12 September 1931

(Donald Denning). 5 - Polk Co., Minn.; 5 September 1936

(D. G. Denning). Kittson Co.. Minn.; 20 June 1936 (D. G.

Denning). 1 - Lancaster, Minn., 26 August 7936 (D. G.

Denning) .

Philia jamesi Hardy.

Utah Acad Sci.\ Arts and Letters, 14:210, 211, figs. 15, 16, 17,
18. 1937.

31 paratypes; 4 - Pingree Park. Colorado; 19 August 1935 (M.
and H. James). 1 - Pingree Park, Colorado; 15 August 1934
(A. E. Pritchard). 11 - Pingree Park. Colorado; 14 August
1934 (C. W. Sabrosky). 6 - Masonville. Colorado, 5 and 6
September 1934. 9 - Marshall Pass. Colorado; 16 September
1917, Cornell University Expedition, ele. 10,250 ft. (R. C.
Shannon) .

FAMILY STRATIOMYIDAE

Ptilocera bergi James

Proc. U. S. Nat'l. Mus.. Vol. 98. No. 3228:202. 1948.

5 9 9 ; Solomon Islands, Guadalcanal; 1944 (D E. Beck
collection #419. 325, 367. 420, 587. 381). 1 5 - Solomon
Islands. Guadalcanal; 1944 (Ernest Reimschiissel). 3 cT cf -
Solomon Islands, Guadalcanal; 1944 (D E. Beck collection
#587, 367. 362).

FAMILY TABANIDAE

Chrysops dilatus Rowe and Knowlton.
Ohio Jour. Sci., 36:256, 1936.
1 paratype; St. George, Utah, #28 (head missing).

FAMILY THEREVIDAE

Zionea tanneri Hardy.

Ent. Soc. Amer. Ann.. 31:144", 1938.

Genotj-pe-Holotype; Zion Nat'l. Park, Utah (Vasco M. Tanner).

Dec. 31, 1969 list of insect types 197

FAMILY ASILIDAE

Mallophora [Mallophorina) pallida Johnson.

Great Basin Nat., 18:41, 1958.

Holoty])e, Allotype and 12 para types; HoUotype. Allotype and 1
paratype - Little Granite Mts., Dugway Pr. Grounds, Tooele
Co., Utah; 21 August 1955 (D. E. Johnson). 5 paratypes -
same except 19 August 1955. 1 paratype - same except 12
August 1955. 1 paratype - S.E. end of Cedar Mts., Tooele Co..
Utah; 28 August 1955 (D. E. Johnson). 2 paratypes - same
except 7 August 1957. 1 paratype - 9 miles east of Delta,
Millard Co.. Utah; 19 August 1957 (D. E. Johnson). 1
paratype - Cane Spring, Cedar Mts.. Dugway Pr. Grounds.
Tooele Co., Utah; 10 August 1955. (D. E. Johnson).
Erax tanneri Bromley.

N.Y. Ent. Soc. Jour., 44: 105, 1936.

Holotype - Buena Vista, Colo., ele. 7,800 ft.; August 1932 (C.
Lynn Hayward).

Erax knowltoni Bromley.

Utah Acad. Sci., Arts and Letters, 14: 104 ,1937.
Paratype; Milford, Utah; June.

Cyrtopogon albi fades Johnson.
Great Basin Nat, 3(1): 1, 1942.
Holotype, Allotype and 1 paratype; Holotype - Glacier Lake,

Timpanogos, Utah Co., Utah (Vasco M. Tanner). Allotype

and paratype - same (E. T. Vest).

Neoitamus hardy i Bromley.

Utah Acad. Sci., Arts and Lettters, 15:61, 1938.

Holotype and 1 paratype; Spanish Fork, Utah (D. Elmo Hardy).

FAMILY BOMBYLIIDAE

Conophorus painteri Priddy.

Kan. Ent. Soc. Jour. 31:12, 1958.

1 paratype; Bear Ears, Elk Ridge, Utah (Vasco M. Tanner).

Poecilanthrax marmoreus Johnson and Johnson.
Great Basin Nat., 17(1-2): 16, 1957.

4 paratypes; Zion Nat'l. Park, Utah (Vasco M. Tanner).

Poecilanthrax varius Painter and Hall.

Kans. Agr. Expt. Sta. Tech. Bui., 106:114. 1960.
3 paratypes; 1 - North Fork, Provo Canyon, Utah (D. Elmo
Hardy); 2 - Spanish Fork, Utah (D. Elmo Hardy).

Poecilanthrax tanbarkensis Painter and Hall.

Kans. Agr. Expt. Sta. Tech. Bui.. 106(2): 109-110. 1960.

5 paratypes; 3 - Tanbark Flat. L./\. Co., California; 2 - 12. 14.

July, 1956 (J. C. Hall). 1 - (.lendale, L.A. Co.. California;
29 July, 1956 (E. I. Schlinger).

The Great Basin Naturalist
198 V. M. TANNER AND D. R. HARRIS Vol. XXIX, No. 4

Exoprosopa arenicola Johnson and Johnson.

Great Basin Nat., 18(3-4): 71, 1958.

4 para types; 2 - Little Granite Mt., Tooele, Co., Utah, 24 August
1956 (D. E. Johnson). 1 - Little Granite Mt., Dugway Prov-
ing Grounds, Tooele Co., Utah, 12 August 1956 (D. E. John-
son). 1 - S.E. end of Cedar Mts., Tooele Co., Utah, 21 Aug-
ust 1956 (D. E.Johnson).
Exoprosopa utahensis Johnson and Johnson.

Great Basin Nat, 18(34): 81, 1958.

1 para type, Antelope Springs, Millard Co., Utah, 10 August
1943 (D. E. Johnson).
Exprosopa sharonae Johnson and Johnson.

Great Basin Nat., 18 (3-4): 78, 1958.

4 paratypes; 2 - Little Granite Mt., Tooele Co., Utah, 16 August
1958 (D. E. Johnson). 2 - Little Granite Mt., Dugway Prov-
ing Grounds, Tooele Co., Utah, 1 September 1955 (D. E.
Johnson) Field #39.
Poecilanthrax marginatus Johnson and Johnson.

Great Basin Nat, 17(1-2): 14, 1957.

Holotype and Allotype. Johnson's Pass, Tooele Co.. Utah, 4
September 1955 (D. E. Johnson).
Poecilanthrax ingens Johnson and Johnson.

Great Basin Nat.. 17(1-2): 13, 1957.

Holotype and Allotype. Ramsey Canyon, Huachuca Mts.,
Cochise Co., Arizona, 5 September -955 (F. G. Werner and
and G. D. Butler) .

Poecilanthrax sackenii monticola Johnson and Johnson.
Great Basin Nat.. 17(1-2) :23. 1957.
Holotype - Aspen Grove, Utah Co., Utah, 23 July 1955 (D. E.

Johnson.
Allotype - Emerald Lake. Mt. Timpanogos, Utah Co., Utah, 4

August 1955 (D. E. Johnson).

Poecilanthraax robusta Johnson and Johnson.
Great Basin Nat, 17(1-2) :20, 1957.

Holotype and Allotype. Walker's Pass, California, 29 September
1945 (D. E. Johnson).

Poecilanthrax fasciata Johnson and Johnson.
Great Basin Nat., 17(1-2) : 11. 1957.
Holotype - Crowley, Colorado. 2 September 1939 (M. T. James).

Poecilanthrax butleri Johnson and Johnson.
Great Basin Nat. 17(1-2): 11, 1957.
Holotype - Tucson. Arizona. 15 October 1955 (F. G. Werner)

Baccharis saraothroides) .
Allotype - Tucson Arizona, 14 October 1955 (G. D. Butler)
Baccharis.
Exoprospa sharonae Johnson and Johnson.
Great Basin Nat (1958), 18:78.

Dec. 31. 1969 list of inskct typks 199

llolotype - West Side LiUle (.ranite Mt.. Tooele (>o.. I tal). ^

September 1955 (D. E. Johnson).
Allotype - S.E. end of Cedar Mts.. Tooele Co.. ITtah. S Se[)teni-
ber 1955 (Sharon Johnson).
Exoprosopn arenicola Johnson and Johnson.
Great Basin Nat.. 18(3-4): 71, 1958.

Holotype and Allotyj)e - Westside Little Granite Mrs., Tooele
Co.. Utah, 27 July 1955 (D. E. Johnson).
Exoprosopa utahensis Johnson and Johnson.
Great Basin Nat.. 18(3-4): 81. 1958.
Holotype - Antelope Springs. Millard Co.. Utah. 10 August 1943

(D. E. Johnson).
Allotype - S.E. end of Cedar Mts.. Tooele Co.. Utah. 28 August
1955 (D. E. Johnson).
Exoprosopa abdominalis Johnson and Johnson.
Great Basin Nat. 19(1). 10, 1959.

Holotype and Allotype - Weldon, Kern Co.. California; 5 Aug-
us"t 1945 (D. E. Johnson).
Exoprosopa butleri Johnson and Johnson.
Great Basin Nat., 18 (3-4): 74, 1958.
Holotype 4 miles East Apache Junction. Arizona; 15 May 1954

(G.D.Butler).
Allotype - Sycamore Canyon, Atascosa Mts., Arizona, 22 May
1955 G. Butler) (Senecio longilobus).
Oestranthrax farinous Johnson and Maughan.
Great Basin Nat., 13: 18. 1953.

Holotype and 6 paratypes; Delta. Utah., 5 July (D. E. Johnson).
Allotype and 1 paratyi^e; Delta, Utah; 5 July 1943 (D. E. John-
son).
2 paratypes - Delta. Utah., 4 July (D. E. Johnson).
Conophorus sackenii Johnson and Maughan.
Great Basin Nat.. 13:22, 1953.

Holotype, allotype and 4 paratypes; Herlong, California, 14 May
1944 (D. E. Johnson).

Exepacmus johnsoni Copuillett.

Amer. Ent. Soc. Trans., 21 : 101, 1894.

1 Androty]ie cT. Johnson and Johnson. Inyokern. California. 28

April 1945 (D. E. Johuson).

Lordotus lutcelus Hall.

Calif. Univ. Pubs. Ent.. 10:26. 1954.

2 paratypes; Walkers Pass. California, 16 September 1945 (D.

E. Johnson).

Lordotus miscellus melanosus Johnson and Johnson.
Great Basin Nat.. 19:24. 1959.

Holotvpe, Allotype and 9 parat\i^es; Holotype. .\llotv]>e - Little
Granite Mt., Tooele Co., Utah; 14 Sept^ember 1957 (D. E.
Johnson). 3 paratypes - same except 11 September 1956. 3 -

The Great Basin Naturalist
200 V. M. TANNER AND D. R. HARRIS Vol. XXIX, No. 4

para types - same except 7 September 1956. 2 - Dog Area,

Dugway Pr. Grounds, Tooele Co., Utah, 12 September 1955

(D. E. Johnson) Field #44. 1 - same except 31 August 1955

Field #38.
Lordotus lutescens Johnson and Johnson.
Great Basin Nat., 19(1): 15. 1959.
6 para types; Las Cruces. New Mexico; 25 April 1954 (R. H.

Beamer) .
Lordotus cinqulatus lineatus Johnson and Johnson.
Great Basin Nat.. 19(1) :22. 1959.
Holotype, Allotype and 2 paratj^Des; Ten miles E. San Quintin,

Mexico. Baja California; 10 September 1955 (Paul H.

Arnaud) .
Lordotus perplexus Johnson and Johnson.
Great Basin Nat., 19(1): 16. 1959.
Holotype and Allotype; Rillite, Arizona; 20 March 1934 (A. J.

Basinger).
Lordotus apicula Coquillett.
Ent. Amer., 3:116, 1887.
Homotype and Plesiotype; Skull Valley. Tooele Co., Utah; 31

May 1956 (D. E. Johnson). One Pleisotype - Skull Valley,

Tooele Co.. Utah; 9 June 1957 (D. E. JohnsonV

Lordotus abdominalis Johnson and Johnson.
Great Basin Nat., 19(1): 10.

Holotype - Tucson. Arizona; 24 April 1954 (F. G. Warner).
Allotype - West of Standfield. Arizona; 11 April 1955 (Butler
and Werner).
Lordotus arizonensis Johnson and Johnson.
Great Basin Nat.. 19(1 ) : 19, 1959.

Holoty]:)e - Sabino Canyon, Santa Catalina Mts., Arizona; 17
October 1955 (G. D. Butler).

Lordotus cingulatus Johnson and Johnson.
Great Basin Nat.. 19(1 ):21. 1959.

5 paratypes; 17 miles north of Wolf Hole, Mohave Co.. Arizona;
9 September 1958.

Lordotus sororculus nigriventris Johnson and Johnson.

Great Basin Nat.. 1 9 ( 1 ): 1 7. 1 959.

Holotype. Allotype, and 8 paratypes; Holotype. Allotype and 1
para type - Skull Valley, Tooele Co.. Utah; 13 May 1956 (D.
E. Johnson). 1 para type - same except 3 IMay 1956. 1 para-
type - same except 18 May 1958. 2 paratypes - same except
2 June 1957. 1 paratype - Dugway Mts.. Juab Co.. Utah; 25
May 1958 (D. E. Johnson). 2 paratypes - Wig Mts.. Dugway
Pr. Grounds. Tooele Co., Utah; 23 April 1956 (D. E. John-
son).
Poecilanthrax johnsonorum Painter and Hall.

Kans. Agr. Expt. Sta. Tech. Bui.. 106:69. 1960.

Dec. 31. 1969 list of inskct types 201

Holotype and Allotype:; Little (iranite Mt.. Tooolo Co.. I'tah;
14 September 1957 (D. E. Johnson).

Poecilanthrax hyalinipcnnis Painter and Hall.
Kans. Agr. Expt. Sta. Tech. Bui.. 10b: ()4. 19(i().
4 paratypes. 3 - Little Granite Mt.. Tooele Co.. Utah; 14 Sep-
tember 1957. 1 - Little Granite Mt.. Dugway Pr. Grounds.
looele Co.. Utah; 15 September 1955 (D. E. Johnson) Field
#45.
Poecilanthrax apache Painter and Hall.

Kans. Agr. Expt. Sta. Tech. Bui.. 106:31. 1960.
2 paratypes; Tucson, Arizona; 15 October 1955 (F. G. Werner)
{Baccharis sarotJiroides) .

Poecilanthrax painteri Maughan.

Kans. Ent. Soc. Jour.. 8:56-57. 1935.

Allotype and 3 paratypes; Allotype and 1 paratype - Blacksmith

Fork Canyon, Cache Co., Utah. 20 June 1944 (D. E. Johnson;

2 paratjT^es - Lava Hot Springs, Idaho. 27 June 1948 (D. E.

Johnson).

Poecilanthrax alpha zionensis Johnson and Johnson.
Great Basin Nat.. 17 (1-2): 6. 1957.

Holotype - Zion National Park. Utah, 8 July 1932 (D. E. John-
son).
Lordotus cingulatus rufotibialis Johnson and Johnson.
Great Basin Nat, 19:23, 1959.

Holotype and Allotype; Phoenix, Arizona, 29 September 1940
(Parker).
Conophorus hiltoni Priddy.

Kans. Ent. Soc. Jour., 31:24, 1958.

Holotype. Allotype and 2 paratypes; Herlong, California, 14 May
1944 (D. E.Johnson).

Conophorus collini Priddy.

Kans. Ent. Soc. Jour., 31:31. 1958.

1 paratype; Oceano, S. Luis Obispo Co., California, 24 April 1951
(R. M. Bohart).

Conophorus painteri Priddy.

Kans. Ent. Soc. Jour., 31:12, 1958.

Holotype. Allotype and4 paratypes; American Fork Canyon.
Utah. 4 July 1953 (D. E. Johnson); 1 paratype - .^spen
Grove, Utah Co., Utah, 19 July (D. E. Johnson).

Amphicosmus arizonensis Johnson and Johnson.
Great Basin Nat.. 19:67. 1960.

Holotype and Allotype; Tucson Mts.. Arizona; 16 August 1955
(G. D. Butler).

DicranocUsta fasciata Johnson and Johnson.
Great Basin Nat., 19:71, 1960.
Holotype; Cedar Creek, Arizona; 15 miles West of Fort Apadie.

The Great Basin Naturalist
202 V. M. TANNER AND D. R. HARRIS Vol. XXIX, No. 4

21 June 1957 (G. Butler and F. Werner). Allotype; Provo, Utah Co.,
Utah. July 1933 (D. E. Johnson).
Villa utahensis Maughan.

Kans. Ent. Soc. Jour.. 8:51. 1935.

2 paratypes; Provo, Utah; 29 April 1934 (D. E. Johnson).

1 metatype; Provo environs, Utah (D. E. Johnson).

FAMILY DOLICHOPODIDAE

Dolichopus indianus Harmston and Knowlton.
Amer. Midi. Nat.. 36:672, 1946.

1 paratype; Indianapolis, Indiana; 7 March 1943 (F. C. Harms-
ton).
1 paratype; Indianapolis. Indiana; 7 March 1943 (F. C. Harm-
ston).
Medeterus arnaudi Harmston.

Great Basm Nat, 11 (1-2): 12, 1951.

cT paratype; Redwood City, California, San Mateo Co.; 4 June
1946 (PaulH. Amaud).
Aphrosylus ivirthi Harmston.

Great Basin Nat., 11 (1-2): 13, 1951.

1 paratype; Moss Beach. San Mateo Co.. California; 21 March

1948 (W. W. Wirth) (Intertidal rocks).

FAMILY PIPUNCULIDAE

Chalarus latifrons Hardy.

Kans. Univ. Sci. Bui., 29:33, 1943.

6 parat>^)es; 3 - Austin. Nevada; 12 August 1940 (R. H. Beamer,
D. E. Hardy). 1 - Cloudcroft, New Mexico; 27 June 1940 (R.
H. Beamer); 1 - Ruidoso. New Mexico; 26 June 1940 (R. H.
Beamer). 1 - Pacific, California; 9 August 1940 (D. E.
Hardy) .
Tomosvaryella tumida Hardy.

Kans. Ent. Soc. Jour., 13:112. 1940.

2 paratypes; Belle Glade, Florida; 18 July 1939 (R. H. Beamer).
Tomosvaryella xerophila Hardy.

Kans. Univ. Sci. Bui., 29:188, 1943.

2 paratypes; 1 - Cuervo. New Mexico; 23 June 1940 (L. C.
Kuitert). 1 - Rustler's Peak, Chiricahua Mts., Arizona; 5 July
1940 (D. Elmo Hardy).
Tomosvaryella tumida Hardy.

Kans. Ent. Soc. Jour.. 13:112. 1940.
1 paratype; Belle Glade, Florida; 18 July 1939 (R. H. Beamer).
Tomosvaryella lepidipes Hardy.

Kans. Univ. Sci. Bui. 29:166, 1943.

4 paratypes; 2 - Garnett. Kansas; 22 September 1940 (D. Elmo
Hardy). 1 - Maybell. Colorado; 18 August 1940 (R. H.

Dec. 31, 1969 list of inskct typks 203

Bonnier). 1 - Yosemite National Park. California; 1 .\ii^ust
1940 (D. Elmo Hardy).
TomosvarycUa heomcri Hardy.

Kans. Ent. vSoc. Jour., 13:107, 1940.

1 para type; Cherokee Co.. Kansas; 30 August 1939 (R. H.
Beamer) .
Tomosvaryella jloridensis Hardy.

Kans. Ent. Soc. .Tour.. 13:109, 1940.

1 cf paratype; Adel, Ga.; 11 August 1939 (J. D. Beamer).

Tomosvaryella wilburi Hardy.

Kans. Ent. Soc. Jour.. 12:22, 1939.

1 paratype, 8 metatypes; paratype - Manhattan, Kansas; 13

August 1937 (D. A. Wilbur). 3 metatypes - Tonganoxie.
Kansas; 6 May 1939 (D. Elmo Hardy). 4 metaypes -
Cherokee Co.. Kansas; 30 August 1939 (R. H. Beamer). 1
metatype - Lawrence. Kansas; 22 August 1939 (D. Elmo
Hardy).
Tomosvaryella coquilletti coquilletti (Kertesz).

Tomosvaryella proxima Cresson.

Amer. Ent. Soc. Trans., 36:318, 1911.

3 homotypes; Tonganoxie. Kansas; 6 May 1939 (R. H. Beamer)
by D. Elmo Hardy.

Tomosvaryella similis Hough.

Boston Soc. Nat. Hist. Proc. 29:84, 1899.

2 homotypes; Garnett, Kansas; 29 August 1939 (R. H. Beamer)

compared by D. Elmo Hardy.

Tomosvaryella contorta Hardy.

Kans. Ent. Soc. Jour., 12:18. 1939.

10 metatypes; 1 - Onyx, Cahfornia; 23 August 1940 (D. Elmo
Hardy). 1 - Onyx, California; 23 August 1940 (R. H.
Beamer). 1 - Kernville, California; 24 August 1940 (R. H.
Beamer). 3 - Tonganoxie, Kansas; 6 May 1939 (D. Elmo
Hardy). 2 - Garnett, Kansas; 29 August 1939 (J. D. Beamer).

Tomosvaryella subvirescens Loew.
BerHn Ent. Ztschr., 16:87. 1872.
7 homotypes; 1 - Toganoxie, Kansas; 6 May 1939 (D. Elmo

Hardy). 2 - Toganoxie, Kansas; 3 September 1939 (R. H.

Beamer). 4 - Douglas Co., Kansas; 23 August 1939 (R. H.

Beamer) compared by D. Elmo Hardy).

Tomosvaryella exilidens Hardy.

Kans. Ent. Soc. Bui., 29:162, 1943.

2 para types; 1 - Moriarty, New Mexico; 24 June 1940 (E. E.

Kenaga). 1 - Sunnyside Canyon, Huachuca Mts., Arizona;

9 July 194- (D. Elmo Hardy).

Tomosvaryella turgida Hardy.

Kans. Ent. Soc. Jour., 13:113, 1940.

The Great Basin Naturalist
204 V. M. TANNER AND D. R. HARRIS Vol. XXIX, No. 4

2 para types; Griffin, Georgia; 12 August 1939 (R. H. Beamer,
A. T. Hardy).
Tomosvaryella agnesea Hardy.

Kans. Ent. Soc. Jour.. 13:103, 1940.

2 para types and 4 meta types; Para types, 1 - Manhattan, Kansas,
15 June 1943 (D. A. Wilbur). 1 - Douglas Co., Kansas; 23

August 1938 (R. H. Beamer). Metatypes, 1 - Glasco, Kansas; 24
August 1940 (E. E. Kenaga). 1 - Arivaca, Arizona; 12 July
1940 (R. H. Beamer). 1 - Kernville, California; 24 July 1940
(R. H. Beamer). 1 - Onyx, California; 23 July 1940 (D.
Elmo Hardy).
Tomosvaryella dissimilis Hardy.

Kans. Univ. Sci. Bui.. 29:161. 1943.

2 paratypes; 1 cf - Chiricahua Mts.. Arizona; 4 July 1940 (D.

Elmo Hardy). 1 - Bishop, CaHfornia; 28 July 1940 (L. C.
Kuitert) .
Tomosvaryella aliena Hardy.

Kans. Ent. Soc. Jour., 20:146-153, 1947.

New name for propinqua Hardy.

Kans. Univ. Sci. Bui.. 29: 169. 1943.

5 paratypes; 1 - Goshen. Utah; 16 August 1940 (R. H. Beamer).
2 - Goshen. Utah; 16 August 1940 (L. C. Kuitert). 1 - Fruita,
Colorado; 4 September 1938 (D. Elmo and A. Hardy). 1 -
Lone Pine, California; 28 July 1940 (R. H. Beamer).

Tomosvaryella quadradentis Hardy.
Kans. Univ. Sci. Bui.. 29:172, 1943.

3 paratypes; 2 - Carson City, Nevada; August 1940 (D. Elmo

Hardy). 1 d - Silver Lake, Kansas; 24 August 1940 (L. J.
Lipovsky).
Tomosvaryella brevijuncta Hardy.
Kans. Univ. Sci. Bui., 29:155, 1943.

1 paratype; Palm City, California; 19 July 1940 (R. H. Beamer).

Dorylomorpha tridentata Hardy.

kans. Univ. Sci. Bui., 29:141. 1943.

2 paratypes; 1 - Lake Tahoe, California; 11 August 1940 (R. K,

Beamer). 1 - Lake Tahoe, California; 11 August 1940 (D.
Elmo Hardy).
Pipunculus nevadensis Hardy.

Kans. Univ. Sci. Bui.. 29: 106. 1943.

1 paratype; Fallon, Nevada; 12 August 1940 (E. E. Kenaga).

Pipunculus huachueanus Hardy.

Kans. Univ. Sci. Bui., 29:95, 1943.

2 paratypes; 1 - Sunnyside Canyon, Huachuca Mts., Arizona; 9

July 1940. (R. H. Beamer). 1 - Sunnyside Canyon, Huachuca
Mts., Arizona; 9 July 1940 (D. Elmo Hardy).

Pipunculus cinctus subtilis Hardy.
Kans. Univ. Sci. Bui., 29:84, 1943.

Dec. 31, 1969 list of inskct types 205

1 i)aratype; Pingree Park. Calorado; 11 July 1937 (C. L. John-

ston).
Pipunculus kansetisis Hardy.

Kans. Ent. Soc. Jour., 13:102, 1940.

2 paratypes; 1 - Tekio, North Dakota; 28 July 1937 (C. L.

Johnston). 1 - Douglas Co.. Kansas; 25 August 1939 (R. H.
Beamer) .

Pipunculus alhofascintus Hough.

Boston Soc. Nat. Hist. Proc. 29:85. 1899.

3 honiotvpes; 2 - Garnett, Kansas; 29 August 1939 (R. H.

Beamer). 1 - Clarksville. Tenn.; 15 July 1939 (D. Elma

Hardy) compared by D. Elmo Hardy.
Pipunculus cingulatus Loew.

Berlin Ent. Ztschr. (1865), 9:176, 1866.

2 homotypes; Tangonoxie, Kansas; 6 May 1939 (D. Elmo Hardy)

compared by D. Elmo Hardy.

FAMILY CHLOROPIDAE

Oscinella hesperia Sabrosky.
Canadian Ent., 72:219, 1940.

4 paratypes; Spanish Fork, Utah Co., Utah (D. Elmo Hardy).

ORDER HYMENOPTERA
FAMILY SCOLIIDAE

Campsomeris solomonis Krombein.

Nova Guinea (Zool.) No. 22:570. 1963.

2 paratypes; Solomon Islands, Guadalcanal; 1944 (Ernest Reim-
schiissel).

Scolia pulchripennis franclemonti Krombein
Nova Guinea (Zool.) No. 22:636, 1963.
1 paratype; Solomon Islands, Guadalcanal; 1944 (D E. Beck).

206

INDEX TO VOLUME XXIX

The new genera, species, and varieties in this volume appear in bold
type in this index.

A New Genus and Species of Ori-
batid Mite (Acari: Liacaroidea,
Metrioppiidae), p. 137.

A New Species of Liacarus from a
Wood Rat Nest (Acarina: Crypto-
stigmata, Liacaridae), p. 140.

A New Species of Oppia from Colo-
rado (Cryptostigmata; Oppiidae),
p. 133.

A Note on Geranium richardsonii
Fisch. & Trautv., p. 25.

Acmaeodera diffusa, p. 13.

Alexander, Charles P., Article by,
p. 1.

AUred, Dorald M., Articles by, pp.
20, 42, 105.

An Albino Pacific Tree Frog, Hyla
regilla, from Death Valley, Cali-
fornia, p. 111.

Andersen, Ferron L., Keith H.
Hoopes, and J. Carl Fox, Article
by, p. 35.

Barr, WiUiam F., Article by, p. 11.

Barr, William F., and Richard L.
Penrose, Article by, p. 88.

Bees of the Nevada Test Site, p. 20.

Chramesus minulus, p. 126.

Chramesus vitiosus, p. 125.

Cryptantha shackletteana, p. 28.

Cryptantha subgenus Oreocarya
(Greene) stat. nov., p. 30.

Does Vireo gilvus swainsonii Occur
in Utah?, p. 181.

Ellis, David H., Dwight G. Smith,
and Joseph R. Murphy, Article by,
p. 165.

Flake, Lester D., and Clive D. Jor-
gensen, Article by, p. 143.

Fossil Chelonians, Chrysemys and
Clemmys, from the Upper Plio-
cene of Idaho, p. 82.

Fox, Carl J., Ferron L. Andersen,
and Keith H. Hoopes, Article by,
p. 35.

Froeschner, Richard C, Article by,
p. 129.

Ghiselin, Jon, Article by, p. 109.

Gonomyia (Odiocera) persimplex, p.

9.

Harris, Don R., and Vasco M. Tan-
ner, Articles by, pp. 150, 183.

Higgins, Harold G., and Tyler A.
Woolley, Article by, p. 140.

Higgins, Larry C, Article by, p. 28.

Hoopes, Keith H., Ferron L. Ander-
sen, and J. Carl Fox, Article by,
p. 35.

Individual Rodent Movements in
Upper Sonoran Desert of Central
Nevada, p. 109.

Invasion of a "Trapped-Out" South-
ern Nevada Habitat by Perog-
nathus longimembris , p. 143.

Itzkowitz, Murray, and W. L. Minck-
ley, Article by, p. 169.

Jorgensen, Clive D., and John W.
Leetham, Article by, p. 96.

Jorgensen, Clive D., and Lester D.
Flake, Article by, p. 143.

Kay, Fenton R., Article by, p. 111.

Leetham, John W., and Clive D. Jor-
gensen, Article by, p. 96.

Lepidoptera of the Nevada Test Site,
p. 42.

Liacarus lectronus, p. 140.

List of the Insect Type Specimens
in the Entomological Collection of
the Brigham Young University,
Provo, Utah, No. Ill, p. 150.

List of the Insect Type Specimens
in the Entomological Collection of
the Brigham Young University,
Provo, Utah, No. IV, p. 183.

Metapyroppia doratosa, p. 137.

Metapyroppia, n. gen., p. 137.

Minckley, W. L., and Murray Itz-
kowitz, Article by, p. 169.

Morris, Ronald L., and Wilmer W.
Tanner, Article by, p. 45.

Murphy, Joseph R., David H. Ellis,
and Dwight G. Smith, Article by,
p. 165.

207

New Combinations and a New Spe-
cies of Perennial Cryptantha, p.
28.

New Records and Distributional
Notes for Reptiles of the Nevada
Test Site, p. 31.

New Synonymy and Records of
Platypodidae and Scolytidae (Cole-
optera), p. 113.

Notes on the Distribution and Bio-
nomics of Some Idaho Ceramby-
cidae (Coleoptera), p. 88.

Oppia coloradensis, p. 133.

Overwintering Phytoseiid Mites in
Central Utah Apple Orchards, p.
96.

Penrose, Richard L., and William F.
Barr, Article by, p. 88.

Phloeotribus destructor, p. 124.

Phloeotribus furvus, p. 124.

Phloeotribus quercinus, p. 123.

Pseudohylesinus pini, p. 122.

Qualitative Behavior of a Pupfish
(Cyprinodon atrorus) in Differing
Environments, p. 169.

Smith, Dwight G., David H. Ellis,
and Joseph R. Murphy, Article by,
p. 165.

Spiders of the National Reactor
Testing Station, p. 105.

Stragulotingis, n. gen., p. 130.

Studies on Raptor Mortality in West-
ern Utah, p. 165.

Tanner, Vasco M., and Don. R. Har-
ris, Articles by, pp. 150, 183.

Tanner, Wilmer W., Article by, p.
31.

Tanner, Wilmer W., and Ronald L.
Morris, Article by, p. 45.

The Buprestidae and Cleridae of the

Nevada Test Site (Coleoptera), p.
11.

The Ecology of the Western Spotted
Frog, Rana pretiosa pretiosa
Baird and Girard, A Life History
Study, p. 45.

The Efficacy of Haloxon and
Thiabendazole as Anthelmintics
Against Gastro-Intestinal Nema-
todes in Sheep, p. 35.

The Infection of Immature Aquatic
Insects by Larval Paragordius
(Nematomorpha)

Tipula ( Arctotipulaj denali, p. 2.

Tipula (Arctotipula) epios, p. 3.

Tipula (Arctotipula) mckinleyana,
p. 4.

Tipula (Eremotipula) eurystyla, p. 8.

Tipula (Platytipula) knowltoniana,
p. 5.

Tipula (Sinotipulaj denningi, p. 7.

Undescribed Species of Nearctic
Tipulidae (Dipteral. X, p. 1.

Van Cott, John W.. Article by. p. 25.

Wislouchiella planctonica Skvortz
(Chlorophyta, Volvocales), A New
Algal Record for Nevada, p. 43.

Wood, Stephen L., Article by. p. 113.

Woolley, Tyler A., Articles by, pp.
133, 137.

Woolley, Tyler A., and Harold G.
Higgins, Article by, p. 140.

Worthen, Gary L., Article by. p. 181

Zoogeographic and Systematic Notes
on the Lace Bug Tribe Litadeini,
with the Description of the New
Genus Stragulotingis (Hemiptera:
Tingidae). p. 129.

Zug, G. R., Article bv, p. 82.

III! Ill

3 2044 072 23

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